Pison argentatum Shuckard, 1838

Pulawski, Wojciech J., 2018, A Revision of the Wasp Genus Pison Jurine, 1808 of Australia and New Zealand, New Guinea, and the Pacific Islands (Hymenoptera: Crabronidae), Proceedings of the California Academy of Sciences 65, pp. 1-584 : 64-72

Pison argentatum Shuckard

Figures 80 -89.

Pison argentatum Shuckard, 1838:79 , ♀ (as argentatus , incorrect original termination). Holotype: ♀, Mauritius: no specific locality ( OXUM), examined. – As Pison argentatum : Le Guillou, 1842:320 ( Singapore); F. Smith, 1856:314 (in catalog of Hymenoptera in British Museum); Kohl, 1885:186 (in checklist of world Pison ); de Saussure, 1892:528 ( Madagascar, redescription); Bingham, 1897:220 (in revision of Indian subcontinent aculeates); Dalla Torre, 1897:710 (in catalog of world Hymenoptera ); Turner, 1911:371 (Island of Aldabra); Bordage, 1912:32 (Isle of Réunion: nesting habits, occasional cleptoparasitism using nests of Sceliphron hemipterum , now fuscum); Kohl in Bordage, 1912:86 (description of ♂); Perkins, 1912:727 (introduced into Hawaii); Turner, 1916b:594 (in key to Afrotropical Pison ), 619 (introduced to Hawaii); Bridwell, 1919b:123 (in key to Hawaiian Pison ); Williams, 1919:143 ( Philippines: Los Baños); Swezey, 1921:522 (Hawaiian Islands: Kauai); nec Maidl, 1924:234 (= Pison carinatum Turner , present correction), 1925:390 ( Indonesia: Sumatra); Guiglia, 1928:500 ( Somalia, almost certainly in error); Pagden, 1934:459 (Malay Peninsula; nests constructed of mud, prey: Lycosidae spiders; nest parasites: mutillid Smicromyrme decora (F. Smith) and bombyliid Petrorossia ceylonica Brunetti ; hyperparasite: Melittobia hawaiiensis Perkins ); Swezey, 1942:185 ( Guam, nesting habits); Arnold, 1945:3 (in key to Pison of Madagascar); Krombein, 1949:384 (in key to Sphecidae of Micronesia), 403 (diagnostic characters; Mariana and Caroline Islands), 1950:139 (island of Yap); Vesey-Fitzgerald, 1956b:362 ( Seychelles); Evans, 1957b:98 (description of larva); Yoshimoto, 1960:334 (Hawaiian Islands); Tsuneki, 1963:11 ( Thailand); Iwata, 1964b:374 (nesting habits in Thailand); Yoshimoto, 1965:291 (nesting habits); Baltazar, 1966:335 (in catalog of Hymenoptera of Philippines); Tsuneki, 1974:636 ( Thailand); R. Bohart and Menke, 1976:335 (in checklist of world Sphecidae ); Casolari and Casolari Moreno, 1980:114 (specimens in M. Spinola collection); Tsuneki, 1983a:89 ( Philippines; comparison with Pison ignavum ), 102 (in key to Pison of Philippines); Radović, 1985:65 (sting apparatus analyzed); Callan, 1990:20 ( New Caledonia: no specific locality); Madl, Matyot, and Schödl, 1996:832 ( Seychelles Islands); D. Baker, 1998:173 (origin and depository of type material); Kami and Miller, 1998:57 (in checklist of Samoan insects); Pulawski, 2003:797 (in checklist of Malagasy Sphecidae ); Starr, 2004:565 (nesting habits); Evenhuis, 2007:6 (in checklist of Hymenoptera of Fiji); Terayama and Nambu, 2009:5, 18 (in key to Trypoxylini of Japan); Haneda, 2011:46 ( Philippines: Palawan); Pagliano, 2011a:114 (specimens in coll. Spinola, Torino, are Pison sp. ); Jennings, Krogmann, and Burwell, 2013:32 (in checklist of Hymenoptera of New Caledonia); Madl, 2014a:976 (in catalog of Ampulicidae , Crabronidae , and Sphecidae of Madagascar, with synonymy and locality records). – As Pisonitus argentatus : F. Smith, 1869a:298 (new combination, in checklist of Pisonitus ), 1871a:366 (in catalog of Oriental Aculeata).

Pison sarawakense Cameron, 1903:163 , ♀. Lectotype: ♀, Malaysia: Borneo (Sarawak): no specific locality ( BMNH), present designation, examined. New synonym. – R. Bohart and Menke, 1976:336 (in checklist of world Sphecidae ).

Pison ignavum Turner, 1908:511 , ♀, ♂. Lectotype: ♀, Australia: Queensland: Mackay (BMNH), present designation, examined. New synonym. – Turner, 1910:355 (as synonym of Pison argentatum ), 1916b:596 (in key to Australian Pison ), 601 (as Australian ssp. of Pison argentatum ; Fiji), 1919:338 ( Fiji, Queensland); Williams, 1932:152 (Marquesas Islands), 1945:440 ( New Caledonia, recognition characters), 1947:318 and 330 ( Fiji); Krombein, 1949b:385 (in key to Sphecidae of Micronesia), 404 ( Caroline Islands); Yasumatsu, 1953:140 (bibliographic references, geographic distribution); Fullaway, 1957:279 (in checklist of Hymenoptera of Fiji); Tsuneki, 1967:21 (Taiwan), 1971:19 (Taiwan); R. Bohart and Menke, 1976:336 (in checklist of world Sphecidae ); Tsuneki, 1976:95 ( Philippines); Evans, Matthews, and Hook, 1981:222 (nest structure); Tsuneki, 1982a:36 ( Bismarck Archipelago), 1983a:89 ( Philippines), 102 (in key to Pison of Philippines), 1983b:42 (in key to Pison of New Guinea), 45 (New Guinea); Cardale, 1985:260 (in catalog of Australian Sphecidae ); Radović, 1985:65 (sting apparatus analyzed); Callan, 1990:20 ( New Caledonia: no specific locality); Porter, Stange, and Wang, 1999:9 (in checklist of Sphecidae of Taiwan). – As Pison argentatum ignavum : Cheesman, 1928:177 (Marquesas and Society Islands); Perkins and Cheesman, 1928:6 (listed from Samoa), 28 ( Samoa); Kami and Miller, 1998:57 (in checklist of Samoan insects); Evenhuis, 2007:6 (in checklist of Hymenoptera of Fiji); Jennings, Krogmann, and Burwell, 2013:32 (in checklist of Hymenoptera of New Caledonia).

As Pison perplexum : Roth, 1885:321 (nest structure, prey, as perplexus), present correction.

SPECIES IDENTITY.– The identity of Pison argentatum was not firmly established in the XIXth and XXth centuries. Apparently, none of the previous authors examined the holotype, but I received it for study through the kindness of Dr. James E. Hogan. The holotype, preserved in the Westwood collection at the Oxford University Museum, United Kingdom, is a male lacking the head, prothorax, and forelegs. Nevertheless, it is certainly conspecific with the specimens from Australia, Bali, East Malaysia, Java, New Guinea, Philippines, Singapore, and Sri Lanka in the California Academy of Sciences collection. It shares with them the second recurrent vein reaching the second submarginal cell at the middle of its length, the fine scutal punctures, the tegula of the usual form, the well-defined ridges on the propodeal dorsum, the non-pedunculate gaster, and the all-black body. Its sternum VIII has a well-defined apical emargination.

LECTOTYPE DESIGNATIONS.– Cameron (1903) did not indicate the number of specimens examined in his description of Pison sarawakense . I have designated as the lectotype of this species the only female present under this name in The Natural History Museum, London. Similarly, Turner (1908) did not indicate the number of specimens examined in his description of Pison ignavum . Of the five females of this species present in The Natural History Museum, London under this name, I have designated one as the lectotype and the remaining ones as the paralectotypes.

RECOGNITION.– Pison argentatum has the head, thorax, propodeum, and gaster all black (legs partly ferruginous in some Australian specimens), the second recurrent vein received near the middle of the second submarginal cell, the pronotal collar concealed by setae, and the tibial spurs whitish. Also, the scutal flange is slightly projecting beyond the anterior margin of the axilla, the posterior scutal margin is slightly concave next to the apex of the flange, and the propodeal dorsum is ridged in the vast majority of specimens. Pison rufipes is similar, but in P. argentatum the erect setae of the upper frons are about as long as 0.5 × midocellar diameter, although a few sparse setae may be as long as midocellar diameter (1.0-1.5 × midocellar diameter in P. rufipes ). In the female of P. argentatum , the ocellocular distance equals 0.6-1.1 × hindocellar diameter (1.2-1.5 × in P. rufipes ), and the legs are black in the vast majority of specimens, but partly ferruginous in some (in rufipes the legs are mostly ferruginous, but exceptionally all black, as in P. argentatum ). The males are easily differentiated by the sculpture and pilosity of sternum VIII: in P. argentatum , it is unsculptured and asetose except near the hindmargin, whereas in P. rufipes it is punctate and setose (except basally); the leg color is as in the females. Also Pison prostratum resembles P. argentatum , but in that species the setae of the upper frons are appressed (erect, about as long as half width of the midocellus in P. argentatum ) and the ocellocular distance is smaller than the distance between the hindocelli (the ocellocular distance and the distance between the hindocelli are about equal in the female of P. argentatum ).

JUSTIFICATION OF NEW SYNONYMY.– Pison sarawakense Cameron, 1903 is certainly conspecific wih Pison argentatum Shuckard. The two names ae therefore synonyms.

Turner (1910) thought that his P. ignavum (described in 1908) was a junior synonym of P. argentatum , and later (1916b) treated it as the Australian subspecies of P. argentatum , whereas Williams (1932) and Krombein (1949) regarded them to be separate species (although Krombein called P. ignavum to be “uncomfortably close to argentatum ”). The differences between the two species were supposed to be the shape of the clypeal lamella in both sexes and the shape of male sternum VIII. The clypeal lamella of the female, however, varies from slightly, evenly arcuate to one with a well-defined median projection, with intermediates. Moreover, it is not correlated with the shape of the male clypeus. For example, the males from the Wonga Beach, Queensland have the clypeal lamella widely obtusely angulate, whereas in the females it has a minimal median projection, and all four males examined from Moorea, French Polynesia also have the clypeal lamella widely obtusely angulate, but in the females the clypeal lamella varies from slightly, evenly arcuate to one with a well-defined median projection, with intermediates. As these females were found the same day at the same locality, I think it unlikely they represent two species.

The male of P. argentatum was supposed to have the clypeal lamella widely obtusely angulate and the apical margin of sternum VIII with a well-defined, moderately deep emargination, whereas in P. ignavum the clypeal lamella was obtusely pointed and sternum VIII was shallowly emarginate apically. A male from Suva, Fiji, however, combines the clypeal lamella of P. argentatum with sternum VIII of P. ignavum , and in other males from that locality sternum VIII is intermediate. Most males from Australia have the clypeal lamella obtusely pointed, but the point is barely expressed in a specimen from 1 km W Eumungerie, New South Wales, suggesting full intergradation.

Based on the above observations, I treat P. ignavum as conspecific with P. argentatum , and a junior synonym of the latter name.

DESCRIPTION.– Frons dull minutely punctate, punctures shallow, about one diameter apart. Distance between antennal socket and orbit equal to socket diameter in female, slightly larger than that in male. Occipital carina higher than hypostomal carina. Gena narrow in dorsal view. Labrum emarginate. Anteromedian pronotal pit transversely elongate, about as long as 1.0-1.5 midocellar width. Scutum not foveate along flange, with short longitudinal ridges adjacent to posterior margin; scutal punctures fine, on disk averaging about two diameters apart; interspaces microsculptured, dull; scutal flange slightly projecting beyond anterior margin of axilla, posterior scutal margin slightly concave next to apex of flange. Scutellum in most specimens with crenulate sulcus adjacent to scutal margin, but sulcus practically absent in specimens from Magnetic Island, Queensland, and intermediate in some others. Tegula not enlarged. Mesopleural punctures minute, averaging about one diameter apart, more than one diameter apart in some Australian males, partly concealed by vestiture, interspaces unsculptured. Postspiracular carina present, 1.0-1.5 × as long as midocellar diameter. Metapleural sulcus costulate and sunken between dorsal and ventral metapleural pits; metapleural punctures microscopically small. Propodeum with irregular longitudinal carina separating side from dorsum and posterior surface and extending from gastral socket area toward spiracle; dorsum with well-defined, oblique ridges in most specimens, but ridges evanescent in some specimens; side ridged at least dorsally; posterior surface transversely ridged; entire propodeum minutely punctate between ridges. Second recurrent vein received near midlength of submarginal cell II. Hindcoxal dorsum with outer margin rounded, not carinate. Punctures of tergum I minute, averaging about one diameter apart anterior of apical depression. Sterna evenly punctate throughout, those of sternum II averaging about 1-2 diameters apart mesally.

Setae silvery, subappressed on upper frons, appressed on scutum and tergum I, forming patch of dorsolaterally oriented setae on each side of upper frons (between dorsal end of middle carina and midocellus), on lower gena suberect, straight (curved apically), shorter than midocellar diameter; completely concealing integument on clypeus and pronotal collar. Apical depressions of terga with silvery, setal fasciae.

Body black, mandible ferruginous mesally, legs partly ferruginous in some specimens from Australia.

♀.– Upper interocular distance equal to 0.82-0.88 × lower interocular distance; ocellocular distance equal to 0.6-1.1 × hindocellar diameter, distance between hindocelli equal to 0.8-1.1 × hindocellar diameter; eye height equal to 1.04-1.14 × distance between eye notches. Free margin of clypeal lamella slightly obtusely angulate to practically straight ( Fig. 80 ) in specimens from Seychelles, Eastern Malaysia, Philippines, Singapore, and Thailand, but varying from slightly, evenly arcuate to one with a well-defined median projection (with intermediates) in other areas ( Australia, island of Moorea). Dorsal length of flagellomere I 2.2-2.7 × apical width, of flagellomere IX 1.1-1.3 × apical width. Mandible: trimmal carina with small incision at about midlength. Length 5.8-8.5 mm; head width 1.8-2.4 mm.

♂.– Upper interocular distance equal to 0.84-0.90 × lower interocular distance; ocellocular distance equal to 0.7-1.3 × hindocellar diameter, distance between hindocelli equal to 0.8-0.9 × hindocellar diameter; eye height equal to 1.04-1.10 × distance between eye notches. Free margin of clypeal lamella varying from obtusely angulate ( Fig. 81 ) to one with obtuse median point. Dorsal length of flagellomere I 1.6-1.9 × apical width, of flagellomere X 1.1-1.4 × apical width. Sternum VIII ventrally unsculptured except for setigerous punctures adjacent to apical margin, with well-defined, moderately deep apical emargination ( Fig. 82 ). Genitalia: Figs. 83, 84 . Length 5.6-8.0 mm; head width 1.7-2.1 mm.

NESTING HABITS.– The nesting habits of this species were observed by Bordage (1912) Cheesman (1928, as P. ignavum ), Pagden (1934), Swezey (1942), Williams (1945, as P. ignavum ), Iwata (1964b), Yoshimoto (1965), Evans, Matthews, and Hook (1981, as P. ignavum ), and Starr (2004), of which Bordage is the most detailed. Bordage (1912) indicated that P. argentatum was very common on the island of Réunion, while Cheesman (1928) reported that the species was very numerous on the Society and the Marquesas islands “where these wasps build cells of clay pellets, usually choosing a sheltered position on walls, inside buildings, or under the eaves, against flat surfaces under overhanging rocks, suspended in clusters from exposed roots, in the interstices between the cells of Sceliphron , or on the undersurfaces of leaves, etc.”. Williams (1945) found that in New Caledonia the species “constructs free cells of mud pellets and sometimes hangs them from rootlets exposed in the bank”. Evans, Matthews, and Hook (1981) described a nest plastered to the underside of a Banksia leave overhanging a stagnant pool near Brisbane, Queensland; the nest was 4.5 cm long, 3.1 cm wide, and 0.8 cm deep. The nests are commonly found on buildings and other human-made structures, in a variety of situations, usually in somewhat protected places. Yoshimoto (1965) describes nesting of Pison argentatum in the entrance wall of the Bishop Museum in Honolulu, Hawaii. Some nests are built inside old cells of other dauber wasps such as Sceliphron sp. or Eumenes sp. Nests are built out by adding individual pellets of mud; they are smooth on the inner side and rough on the outer side ( Fig. 85 ). An individual cell is about 10-11 mm long, about 7-9 mm in diameter, with walls about 0.5 mm thick. The cell is opened at the top prior to provisioning, and is closed by a circular operculum about 3 mm across when provisioning is completed and the egg laid. The nest may consist from one up to 24 cells (Iwata, 1964). Nests may be covered by a plastering, an additional mud layer making individual cells unrecognizable. When the substrate is an approximately flat surface, the cells tend to form a line, with new cells added serially at one end only; when the substrate departs clearly from the horizontal, new cells are added at the top. The female works on one cell at a time. Starr (2004) confirmed the previous observations.

Roth (1885) observed the nesting habits of what he called Pison spinolae and P. perplexum at Mackay, Queensland, and provided the following description: “The nests are exceedingly brittle, and are apparently formed of small particles of loose dry earth stuck together by some gummy fluid secreted by these wasps. They fill their nests exclusively with small spiders, and the larva makes itself a dull grey brittle shell in the cell”. The determination of Pison spinolae is certainly in error, as this species does not range as far north as Mackay. A specimen of Pison argentatum (as ignavum ) at the BMNH bears the following labels: “85/2”, “ Australia, pres[ented by] Henry Ling Roth, BMNH (E) 1885-2, and “see Roth, 1885, …habits of some Australian Hymenoptera … J. Linn. Soc., Zool. 18:321”. I believe this specimen is what Roth called Pison perplexum .

The prey consists of spiders of the genera Attus (now Salticus ), Salticidae and Sphasus (now Peucetia ), Oxyopidae (Bordage, 1912) , immature Pardosa , Lycosidae (Pagden, 1934) , and members of Araneidae , Lycosidae , Oxyopidae , and Salticidae (Starr, 2014) .

RECORDS.– ALDABRA: no specific locality ( Turner, 1911).

AMERICAN SAMOA: Tutuila Island: Fagatogo (3 ♀, BISH) , Leone-Aluau trail (1 ♀, BISH) , Pago Pago (1 ♂, BISH) , Tafuna (1 ♂, BISH, as Tapuna), Taputimu (2 ♀, 2 ♂, BISH) , no specific locality (1 ♀, 1 ♂, BISH) .

AUSTRALIA: Australian Capital Territory: Canberra: Black Mountain (2 ♀, BMNH) . New South Wales: Blacktown (1 ♀, AMS) , 1 km W Eumungerie at 31°56.7ʹS 148°36.9ʹE (1 ♂, CAS) , Fairfield (4 ♀, AMS) , 0.5 km SE Lansdowne near Taree (1 ♂, AMS; 1 ♀, ANIC), Mount Tomah in Blue Mountains (1 ♀, AMS) , Quakers Hill (1 ♀, AMS) , Whiskers 7 km WNW Hoskinstown at 35°24ʹS 149°23ʹE (1 ♀, ANIC) . Northern Territory: Adelaide River crossing with Daly River Road at 13°29ʹS 131°04ʹE (1 ♂, NTM) , 48 mi. SW Daly River at 14°11ʹS 130°08ʹE (2 ♀, ANIC) , Darwin (1 ♀, NTM) , Gregory National Park at 16°06.6ʹS 130°25.7ʹE (3 ♀, ANIC; 2 ♀, CAS), Keep River National Park at 15°45ʹ30ʺS 129°06ʹ28ʺE (1 ♂, ANIC; 1 ♀, 1 ♂, CAS), McArthur River 2 km SSE Borroloola at 16°05ʹS 136°19ʹE (1 ♀, ANIC) , 16 km NE Mount Cahill at 12°50ʹS 132°51ʹE (1 ♀, NTM) , Muirella Park in Kakadu National Park at 12°51ʹS 132°45ʹE (1 ♀, ANIC) , Springvale 8 km W Katherine (1 ♀, ANIC) , Virginia 31 km SE Darwin Central Business District at 12°33ʹS 131°02ʹE (1 ♂, NTM) . Queensland: Almaden (1 ♀, AMS) , Annandale, a southwestern suburb of Townsville , at 19°19ʹS 146°47ʹE (7 ♀, 2 ♂, NTM) , Arcadia on Magnetic Island at 19°09ʹS 146°52ʹE (9 ♀, ANIC; 1 ♀, CAS; specimens reared from mud nest), Atherton at 17°17ʹS 145°29ʹE (1 ♀, ANIC) , 14 mi NW Ayr (1 ♀, CAS) , Balgal Beach 51 km NW Townsville at 19°02.5ʹS 146°25.2ʹE (2 ♀, CAS) , Brisbane (4 ♀, 10 ♂, QMB) , Brisbane: Blunder Creek (6 ♀, QMB) , Brisbane: Indooroopilly (5 ♀, 4 ♂, BMNH) , Bundaberg (3 ♀, 1 ♂, ANIC; 1 ♂, BMNH), Burnett River at Bundaberg (1 ♀, ANIC) , Cairns (1 ♂, BMNH; 3 ♀, CAS; 1 ♂, RMNH), Cairns at 16°49ʹ07ʺS 145°41ʹ13ʺE (3 ♂, AMNH) , Cairns District (2 ♀, 1 ♂, SAM) , Cape Hillsborough National Park (1 ♂, QMB) , Cockatoo Creek Crossing 17 km NW Heathlands at 11°39ʹS 142°27ʹE (1 ♀, ANIC) , Cooktown at 15°28.3ʹS 145°15.5ʹE (2 ♀, CAS) , Davies Creek National Park at 17°00.2ʹS 145°34.1ʹE (1 ♀, CAS) , 2 km N Davies Creek National Park at 16°58.5ʹS 145°32.7ʹE (1 ♀, CAS) , 18 km S Dingo Beach at 20°16.0ʹS 148°31.2ʹE (1 ♀, CAS) , Eurimbula (1 ♀, AMS) , 5 km S Gympie (1 ♂, QMB) , Homevale National Park at 21°26.9ʹS 148°32.4ʹE (4 ♀, 2 ♂, CAS) , Innes Park E Bundaberg (1 ♂, ANIC) , Kuranda (1 ♀, ANIC; 2 ♀, AMS; 2 ♀, BMNH; 8 ♀, 3 ♂, CAS; 1 ♀, SAM), Kurrimine Beach 30 km S Innisfail at 17°46.6ʹS 146°06.5ʹE (2 ♀, 1 ♂, CAS) , Mabuiag Island in Torres Straits (1 ♀, AMS) , Mackay (18 ♀, including lectotype and 4 paralectotypes of Pison ignavum , 4 ♂, BMNH), Mareeba (3 ♀, ANIC) , Mount Byron area in D’Abguilar Range (1 ♀, QMB) , 56 road km WNW Mount Carbine at 16°19.4ʹS 144°43.2ʹE (1 ♀, CAS) , Mount Webb National Park at 15°04ʹS 145°07ʹE (2 ♀, ANIC) , Mungumby Lodge near Helenvale (1 ♀, AMS) , Old Annandale near Townsville (6 ♀, 1 ♂, NTM) , Paluma Range National Park at 18°51.6ʹS 146°07.6ʹE (1 ♀, CAS) , Pinnacle Creek 27 km N Archer Crossing (2 ♀, 1 ♂, ANIC) , Port Douglas (2 ♀, RMNH; 1 ♀, 1 ♂, SAM), Rockhampton (1 ♀, CAS) , 2 km N Rokeby at 13°39ʹS 142°40ʹE (7 ♀, 2 ♂, ANIC) , 61 km S Rolleston at 24°59.7ʹS 148°27.8ʹE (1 ♂, CAS) , Rowes Bay near Townsville (1 ♀, NTM) , South Mission Beach at 17°56ʹ10ʺS 146°05ʹ41ʺE (1 ♀, AMNH) , Split Rock 14 km SE Laura at 15°39ʹS 144°31ʹE (1 ♀, ANIC) , Stewart River 5 km W Port Stewart (1 ♀, ANIC) , The Bend 3 km NW Coen at 13°56ʹS 143°12ʹE (2 ♀, ANIC) , Townsville (1 ♀, ANIC; 5 ♀, 4 ♂, SAM), Waverley Creek Rest Area 65 km N Marlborough at 22°26.3ʹS 149°28.5ʹE (1 ♀, CAS) , Wenlock River at Moreton (1 ♀, ANIC) , Wonga Beach 11 km NNE Mossman at 16°19.9ʹS 145°25.3ʹE (2 ♀, 2 ♂, CAS) , Crail Bay (1 ♀, RMNH) . Victoria: Gunbower (2 ♀, BMNH) , Melbourne (1 ♂, BMNH) , 23 mi. E Orbost (1 ♂, CAS) . Western Australia: Carson escarpment at 14°49ʹS 126°49ʹE (1 ♀, ANIC) , 4 km W King Cascade at 15°38ʹS 125°15ʹE (1 ♀, ANIC) .

COOK ISLANDS: Aitutaki Island: Amuri (1 ♂, BISH) . Rarotonga Island: Avarua (2 ♂, BISH) , Avatiu (3 ♂, BISH) , Titikaveka (1 ♂, BISH) , and no specific locality (4 ♀, BISH) .

EAST TIMOR: foot of Mundo Perdido near Ossu (1 ♀, SAM) .

FEDERATED STATES OF MICRONESIA (Krombein, 1949, 1950; Yasumatsu, 1953, or as indicated): Chuuk (as Truk) Atoll: Tonowas Island (as Dublon Island), Toloas, Toloas – Erin. Kusaie Island (now Kusrae): Lelu, Mwot – Utwe. Pohnpei Island (formerly Ponape): Kolonia, as Colonia (2 ♀, 2 ♂, BISH), Ronkiti, Ronkiti-One, Sokehs Peninsula (2 ♀, BISH, as Jokaj). Yap Island: Yaptown.

FIJI: Viti Levu: Colo-i-Suva (1 ♀, BISH, as Tholo-i-Suva), Koronivia (3 ♀, ANIC; 2 ♀, CAS), Lautoka (1 ♀, BISH), Nadi (6 ♀, 2 ♂, BISH, as Nandi), Nanduruloulou (1 ♀, BISH), Raki Raki (1 ♂, BISH), 10 km E Sigatoka (1 ♀, 1 ♂, CAS), Suva (5 ♀, BISH; 1 ♀, 1 ♂, BMNH; 9 ♀, 5 ♂, RMNH). Also from Williams, 1947: Bua Province: no specific locality, Korovou, Lautoka, Nandarivatu, Rewa Province: no specific locality, Vunidawa.

FRENCH POLYNESIA (Cheesman, 1928; Williams, 1932, or as indicated): Austral Islands: Rurutu Island: Moerai (4 ♀, 1 ♂, BISH), Tubuai Island: Mahu (3 ♀, 1 ♂, BISH), no specific locality (2 ♀, BISH). Huahine Island: Maeva (1 ♀, BISH). Marquesas Islands: Fatu-Hiva: Omoa; Hiva-oa: Atuona, Hanaiapa Valley (18 ♀, BISH); Nuku Hiva: Taichae (3 ♂, BISH), Taiohae, Tovii; Tahuata: Hanamiai Valley, Hanatetena Valley, Vaitahu Village. Moorea: Atitia (10 ♀, 4 ♂, CAS), Baie de Cook (2 ♀, BISH). Raiatea Island: Uturoa (2 ♂, BISH). Tahiti: Papeiti (Yasumatsu, 1953). Tuamotu Archipelago: Rangiroa Atoll: Avatoru (1 ♀, BISH),

HAWAIIAN ISLANDS: Kauai: Waimea (Swezey, 1921). Maui: Haleakala (1 ♀, BISH). Oahu: Honolulu (5 ♀, 1 ♂, BISH; 1 ♀, CAS), Kalihi (1 ♂, BISH), and no specific locality (6 ♀, 2 ♂, BISH).

INDIA (Bingham, 1897): Uttar-Pradesh (as North West Provinces, no specific localities).

INDONESIA: Bali: 9.6 km NW Ubud (1 ♂, CAS). Halmahera: between Payahe and Gita Woda (1 ♀, RMNH), Tayawi near Payahe (1 ♀, RMNH). Java: Bogor (1 ♀, CAS; 2 ♀, RMNH), Jakarta (8 ♀, 3 ♂, RMNH, as Batavia), Kaliwates (1 ♀, RMNH), Malang (1 ♂, CAS; 2 ♀, RMNH), Samarang (1 ♀, RMNH), Simpang (1 ♀, RMNH), Sindanglaja (12 ♀, RMNH). Maluku: Island of Ambon: Waai (18 ♀, BISH; 2 ♂, CAS), no specific locality (2 ♀, CAS; 12 ♀, RMNH). Seram: Hatumete 15 km NNE Tehoru at 3°17ʹS 129°39ʹE (2 ♂, RMNH). Sulawesi: Bogani Nani Wartabone National Park at 0°34ʹN 123°54ʹE (1 ♂, RMNH, as Dumoga Bone National Park). Sumatra: Bukittingi (Maidl, 1925, as Fort de Kock), Medan Island (1 ♀, RMNH), Palembang (1 ♀, CAS), Sinabang on Simeulue Island (2 ♀, RMNH), northeast Sumatra: no specific locality (1 ♀, RMNH). Western Papua (= Indonesian New Guinea): Bernhard Camp at Taritatu River (1 ♀, 1 ♂, RMNH, as Idenburg River), Hol Maffen 22 km E Sarmi (1 ♀, BISH), Jayapura (8 ♀, 3 ♂, RMHN, mostly as Hollandia), Sentani at 2°40ʹS 140°30ʹE (1 ♀, RMNH). West Timor: Bipolo: Pariti Forest at 10°01ʹS 123°49ʹE (1 ♂, NTM).

JAPAN: Ogasawara (= Bonin) Islands: Anijima Island in Chichijima Group: Southwest Bay (1 ♀, 1 ♂, BISH), Chichijima Island: Omura and no specific locality (2 ♂, BISH), Tatsumi Wan (1 ♂, BISH). Ryukyu Islands: Mount Omoto on Ishigaki Island (Evans, 1957).

KIRIBATI: Gilbert Islands: Banreaba Island (1 ♀, BISH) , Tarawa Atoll: Bairiki Island (1 ♀, 12 ♂, BISH) and Teaoraeke (1 ♀, BISH) .

MADAGASCAR: no specific locality (1 ♀, MHNG) .

MALAYSIA WEST: Perak: Parit Buntar (Pagden, 1934). Sabah: Forest Camp 19 km N Kalabakan (1 ♀, BISH), Forest Camp 9.8 km SW Tenom (1 ♀, BISH), Kennedy Bay (1 ♀, CAS), Koh Bersatu Estate 115 km W Sandakan at 5°42ʹN 117°09ʹE (Starr, 2004), Kota Kinabalu (1 ♀, as Jesselton, 1 ♂ CAS,), Singkor (2 ♀, BISH), Tuaran (2 ♀, 2 ♂, CAS), Ulu Dusun mile 30 on Labuk Road (1 ♂, CAS). Selangor: Dusun Tua in Hulu Langat area (3 ♀, RMNH, as Ulu Langat), Kuala Lumpur (Pagden, 1934), Seri Kembangan (Pagden, 1934, as Serdang).

MALAYSIA EAST: Sabah: Kalabakan River 48 km W Tawau (1 ♀, BISH), Singkor (1 ♂, BISH) .

MARIANA ISLANDS: Guam (Swezey, 1942; Krombein, 1949, or as indicated): Merizo, Nimitz Beach (1 ♀, BISH), Piti, Pago, Talofofo. Saipan Island: As Mahetog (1 ♂, BISH), no specific locality (5 ♂, BISH).

MARSHALL ISLANDS (Yasumatsu, 1935 or as indicated): Ine Island: Arno Atoll (1 ♀, BISH), Jaluit Atoll, Wotje Atoll.

MAURITIUS: no specific locality (Shuckard, 1838).

MYANMAR (as Burma, including Tenasserim): no specific locality (Bingham, 1897), but Rangoon and Kyaikkami (as Amherst) according to Turner, 1916).

NEW CALEDONIA: no specific locality (Callan, 1990; Jennings, Krogmann, and Burwell, 2013). Grande Terre: Hienghène (1 ♀, BISH), La Coulée (1 ♂, BISH), La Foa (1 ♂, BISH), Mouriance Pass (1 ♂, BISH), Nakale River (1 ♀, BISH), Nouméa (1 ♀, 8 ♂, BISH), 25 mi NW Nouméa (Williams, 1945), Ouégoa (1 ♀, UCD), Touho (1 ♀, UCD), Saint Louis (Williams, 1945), NE Yiambi (1 ♀, BISH).

PALAU REPUBLIC: Koror (2 ♀, 1 ♂, BISH; 1 ♀, CAS) .

PAPUA NEW GUINEA: Madang Province: Brahman Catholic Mission at 6°45ʹS 145°23ʹE (1 ♂, CAS), Erima (1 ♂, MTM, determined as ignavum by Tsuneki), Nagada Harbor 8 km N Madang at 5°09ʹS 145°48ʹE (24 ♀, 24 ♂, CAS), Nobonob Hill 7 km N Madang at 5°10ʹS 145°45ʹE (1 ♂, CAS). Morobe Province: Lae (1 ♀, BISH). National Capital District: Boroko – a southern suburb of Port Moresby (1 ♂, BISH), Port Moresby (13 ♀, 8 ♂, CAS), Waigani, a suburb of Port Moresby (2 ♀, UCD). New Britain: Rabaul (1 ♀, BISH). New Ireland: Lavongai Island: Banatam (Tsuneki, 1982). Western Province: Daru Island (1 ♀, BISH).

PHILIPPINES: Cebu: (Tsuneki, 1983a): Argao, Cantabaco Mactan Island near Cebu. Leyte: (Tsuneki, 1983a; Starr, 2004): Baas at 10°22ʹN 124°45ʹE, Basey, near Baybay at 10°45ʹN 124°47ʹE, Palo (2 ♀, BISH), Tacloban. Luzon: (Turner, 1916; Tsuneki, 1983a; Balthasar, 1966, or as indicated), Bacoor, Baguio, Baso, Bontoc, Cavite, Ifugao (1 ♀, BISH), Los Baños (1 ♂, CAS), Manila, Muñoz (1 ♂, CAS), Naga City, Naguilian, Pagsanjan, Tabaco, Tanay in Rizal Province (1 ♂, BISH). Mindanao: (Tsuneki, 1983a, Starr, 2004): Busco at 8°16ʹN 124°58ʹE, Cagayan de Oro, Davao, Malaybalay, Zamboanga. Mindoro: San Jose (2 ♀, CAS). Negros: Taytay beach (Tsuneki, 1983 a). Palawan: (Starr, 2004, Haneda, 2011, or as indicated): near Aborlan at 9°26ʹN 118°33ʹE, Iwahig, Puerto Princesa, 3 km NE Tinabog (1 ♀, BISH). Samar: Basey (Tsuneki, 1983a). Tawi Tawi: Tarawakan (Tsuneki, 1976).

RÉUNION: no specific locality (de Saussure, 1892; Bordage, 1912; Kohl in Bordage, 1912).

SAMOA: Upolu Island: Apia (2 ♂, BISH), Mulinu’u (1 ♂, BISH) .

SEYCHELLES: (Vesey-Fitzgerald, 1956; Madl, Matyot, and Schödl, 1996): Mahé Island: Anse Bougainville (1 ♀, RMNH), Baie Lazare (1 ♀, NHMW), Glacis Village (3 ♀, RMNH), northeast point (1 ♀, RMNH), Port Glaud (1 ♀, NHMW), Val dʹEndor. Praslin Island: near Pasquière River. Silhouette Island: La Passe (2 ♀, NHMW).

SINGAPORE: Singapore (10 ♀, 2 ♂, CAS) .

SOLOMON ISLANDS: Ghizo Island: Gizo (1 ♂, BISH) . Russell Islands: Pavuvu (1 ♀, CAS) .

SOUTH KOREA: Chojusan (Yasumatsu, 1939, a Japanese name, current name unknown).

SRI LANKA: Western Province: Colombo (1 ♂, CAS) .

TAIWAN: Pingtung County: Fangliao (Tsuneki, 1967), Paoli (Tsuneki, 1971).

THAILAND: Bangkok: Bangkok (1 ♂, BISH). Chieng Mai: Chieng Mai (Tsuneki, 1963). Nakhon Nayok: Ban Na (1 ♀, BISH). Pathum Thani: Rangsit Rice Experimental Station (Iwata, 1964). Saraburi: Saraburi (Tsuneki, 1963). Songkhla: Songkhla (2 ♀, CAS). Wang Saphung: Loei (1 ♀, CAS).

TONGA: Tongatapu Island: Nukualofa (2 ♀, BISH) . Vavau Island: Neiafu (2 ♀, 1 ♂, BISH) .

Figures

FIGURES 80–85. Pison argentatum Shuckard. (80) Female clypeus and mandibles; (81) Male clypeus and mandibles; male: (82) Sternum VIII (ventral surface); (83) Genitalia in dorsal view; (84) Genitalia in lateral view; (85) Nest of Pison argentatum Shuckard.