Pison punctifrons Shuckard

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 : 373-380

Pison punctifrons Shuckard

Figures 908 -921.

Pison punctifrons Shuckard, 1838:77 , ♀. Holotype: ♀, “ India or St. Helena ”, actually Austro-Papuan Region: no specific locality (OXUM), examined. – F. Smith, 1856:313 (in catalog of Hymenoptera in British Museum), 1869a:290 (in checklist of Pison ); Kohl, 1885a:188 (in checklist of world Pison ); Cameron, 1889c:118 (in checklist of Oriental Pison ); Dalla Torre, 1897:712 (in catalog of world Hymenoptera ); R. Bohart and Menke, 1976:336 (in checklist of world Sphecidae ); D. Baker, 1998:173 (type origin and depository). Nec following authors (= Pison suspiciosum ): Bingham, 1897; Turner, 1916b; Maidl, 1925; Yasumatsu, 1935, 1936, 1937; Iwata, 1939; Yasumatsu, 1939; Krombein, 1949b, 1950; Yasumatsu, 1953; Iwata and Yoshikawa, 1961; Tsuneki, 1962; Iwata, 1964b; Tano, 1964; Tsuneki, 1964; Baltazar, 1966; Tsuneki, 1967; Haneda, 1968a, 1968b; Tsuneki, 1968c, 1970; Haneda, 1971; Tsuneki, 1971; Yamada, 1971; Haneda, 1972; Tano, 1972; Haneda, 1973; Murota, 1973a, 1973b; Tsuneki, 1974; Nambu, 1975; Tsuneki, 1976, 1977, 1982b, 1982c, 1982d, 1983a, 1983c, 1984a; Paik, 1985; Radović, 1985; Takahashi 1993; Miyatake, 1996; Wu and Zhou, 1996a; Porter, Stange, and Wang, 1999; Yamane, Ikudome, and Terayama, 1999; Lee and Shin, 2000; Krombein and Norden, 2001; Haneda, Nosaka, Tano, Kurokawa, and Murota, 2004, 2005; Hua, 2006; Haneda, Nozaka, Tano, Kurokawa, H. Murota, and T. Murota, 2007; Terayama and Nambu, 2009; Takahashi, 2010; Haneda, 2011; T. Li, and Q. Li, 2011; Kim, 2014.

Pison nitidum F. Smith, 1859:160 , ♀ (as nitidus , incorrect original termination). Lectotype: ♀, Indonesia: Maluku: Key (now Kai) Islands (OXUM), present designation, examined. New synonym.– F. Smith, 1863a:35 ( Indonesia), 1863b:135 (known from islands of Misool, Key, and Aru), 1869:291 (in checklist of Pison ), 1871:366 (in catalog of Oriental Aculeata); Maindron, 1879b:181 ( Indonesia: Maluku: Ternate, redescription, as nitidus ); Kohl, 1885:187 (in checklist of world Pison ); Dalla Torre, 1897:712 (in catalog of world Hymenoptera ); Cameron, 1905:62 (Java, as nitidus ); Turner, 1916b:627 ( Indonesia: Aru and Ké islands, as nitidus ); R. Bohart and Menke, 1976:336 (in checklist of world Sphecidae ).

Pison morosum F. Smith, 1865:85 , ♀ (as morosus , incorrect original termination), junior primary homonym of Pison morosum F. Smith, 1856 . Holotype or syntypes: ♀, New Guinea: no specific locality (BMNH). Synonymized with Pison constrictum by Turner, 1916b:627.

Pison collare Kohl, 1884:337 , ♀. Lectotype: ♀, Papua New Guinea: Eastern New Britain: Duke of York Island (NHMW), present designation, examined. New synonym. – Kohl, 1885:186 (in checklist of world Pison ); Froggatt, 1892:217 (in catalog of Australian Hymenoptera ); Dalla Torre, 1897:710 (in catalog of world Hymenoptera ); Turner, 1916b:627 (diagnostic characters); R. Bohart and Menke, 1976:335 (in checklist of world Sphecidae ); Dollfuss, 1989:11 (holotype in NHMW).

Pison papuanum W. Schulz, 1905:217 . Substitute name for Pison morosum F. Smith, 1865 . New synonym. – Turner, 1917:326 (valid name for Pison constrictum ); R. Bohart and Menke, 1976:336 (in checklist of world Sphecidae ); Tsuneki, 1983:42 (in key to Pison of New Guinea).

Pison constrictum Turner, 1912:201 , ♂. Holotype by monotypy: ♂, Indonesia: Western Papua (= Indonesian New Guinea): Mimika River (BMNH), examined. Synonymized with Pison papuanum by Turner , 1917b:326. – Turner in Turner , Meade-Waldo , and Morley , 1915:7 ( Western Papua : Mimika River ; redescription); Turner , 1916b:627 (almost certainly the male of Pison morosus F. Smith, 1865 ).

Pison bismarckianum Tsuneki, 1982:41 , ♀, ♂. Holotype: ♀, Papua New Guinea: Bismarck Archipelago: New Britain: Yalom (ZMUC), examined. New synonym. – Tsuneki , 1983:42 (in key to Pison of New Guinea), 45 ( New Guinea; geographic variation).

Pison biroi Tsuneki, 1983b:46 , ♀. Holotype: ♀, Papua New Guinea: Morobe Province: Simbang (MTM), examined. New synonym.

Pison huonense Tsuneki, 1983b:48 , ♀. Holotype: ♀, Papua New Guinea: Morobe Province: Satterberg, correctly Sattelberg , (MTM), paratype examined. New synonym.

LECTOTYPE DESIGNATION.– F. Smith did not provide the number of specimens examined in his description of Pison nitidum , but at least one female is present in the Oxford University Museum. I have designated it as the lectotype of this species.

Kohl’s description of Pison collare does not contain the number of specimens studied. I have examined the only specimen present in the Naturhistorisches Museum , Wien, and have selected it as the lectotype of this species .

RECOGNITION. – Pison punctifrons is an all black species with three submarginal cells and erect setae on tergum I and sternum II. It shares with P. pandambai and P. suspiciosum the conspicuously coarse frontal punctures ( Fig. 911 ), some punctures equal to 0.4-0.6 × midocellar diameter (compared to about 0.1-0.2 × in the other Australian species) and with the first species the absence of the longitudinal carina between the propodeal spiracle and the gastropropodeal articulation and the hindcoxal dorsum with a conspicuous tooth at the base of the inner carina.

Pison punctifrons differs from the New Guinean P. pandambai in having the clypeal lamella acutely to slightly obliquely angulate in the female and acutely angulate in the male ( Figs. 908, 909, 910 ), rather than, respectively, truncate and obtusely tridentate or truncate. Unlike P. suspiciosum , the propodeum of P. punctifrons lacks the longitudinal carina separating the side from the dorsum and posterior surface (carina present in P. suspiciosum except rudimentary in some specimens), the hindcoxal dorsum has a prominent tooth at the base of the inner carina (tooth insignificant in P. suspiciosum ), and male sternum VIII has a prominent median process (conspicuously emarginate apically in P. suspiciosum ). Also, the apical depressions of tergum I is finely punctate (unsculptured mesally in most P. suspiciosum ), and the lateral setae of tergum II are appressed (in most P. suspiciosum suberect, about as long as the midocellar diameter).

INTERPRETATION OF PISON PUNCTIFRONS .– The holotype of P. punctifrons is labeled “ S. Helena ? India?”, as recorded in the original description. The reference to India probably prompted Bingham (1897) to include P. punctifrons in his revision of aculeates of British India, and Turner (1916b) to include it in his key to Indian Pison . Apparently neither of the two authors has seen the holotype (that I have received for study through the kindness of Dr. James E. Hogan). Unfortunately, both Bingham and Turner used the name P. punctifrons for a different although closely relat- ed species ( P. suspiciosum ), and they were followed by all subsequent authors. In fact, however, the holotype of P. punctifrons is conspecific with P. nitidum and its synonyms (but not P. suspiciosum ); as the oldest name, it becomes the valid name for them. It must have been collected somewhere in the Austro-Papuan Region (the species does not occur in India).

JUSTIFICATION OF NEW SYNONYMY.– The holotype of P. bismarckianum and the lectotype of P. collare are clearly conspecific with the lectotype of P. punctifrons , and I synonymize these names.

Tsuneki (1983b) differentiated his new Papuan species P. biroi and P. huonense from P. bismarckianum (i.e., P. punctifrons ) first by differences in the punctation of the clypeus, tegula, and sternum II. According to my observations, these characters are individually variable and have no value in species discrimination. The length of flagellomere I, according to the original description, is 2.8 × apical width in the holotype of P. biroi and 3.3 × in that of P. huonense , but I found no difference in this character between the two specimens. The large clypeal punctures on Tsuneki illustration of P. huonense are grossly exaggerated, when compared with the original specimen. The shallow median furrow on the postocellar area and the absence of the midfrontal carina in the holotype of P. biroi are also found in other specimens and in my opinion have no diagnostic value. In addition, the holotype of P. biroi originates from Simbang where a specimen of P. punctifrons was also collected. For these reasons, I consider P. biroi and P. huonense as synonyms of P. punctifrons . As a matter of fact, Tsuneki (1983b) himself suspected this synonymy, as he wrote about P. biroi “the present species may be a variety of bismarckianum ” and about P. huonense “this species may be a variation of the preceding species or of bismarckianum ”.

DESCRIPTION.– Frons coarsely punctate, punctures less than one diameter apart, some of them equal to 0.4-0.6 × midocellar diameter ( Fig. 911 ); interspaces aciculate, slightly shining; middle supraantennal carina present or absent; integument swollen above each antennal socket (swelling ill defined in smallest specimens). Occipital carina slightly expanded, its height at midlength about 0.3 × of midocellar diameter. Labrum not emarginate. Pronotal collar slightly raised apicomesally, roundly angulate laterally, slightly concave between median elevation and lateral swelling. Anteromedian pronotal pit transversely elongate, about twice as long as midocellar diameter. Scutum not foveate along flange, at most with a few ill defined, short longitudinal ridges adjacent to posterior margin; scutal punctures conspicuous, irregularly spaced, varying from several diameters apart on disk to less than one diameter apart ( Fig. 914 ). Tegula enlarged. Mesopleural punctures conspicuous, varying from less than one diameter apart to several punctures about three diameters apart beneath scrobe. Postspiracular carina present, about 3.0 × as long as midocellar diameter. Metapleural sulcus not costulate between dorsal and ventral metapleural pits. Propodeum without longitudinal carina separating side from dorsum and posterior surface; dorsum with series of punctures or minute transverse grooves along midline, otherwise variously sculptured: either sparsely punctate (punctures more than one diameter apart, Fig. 915 ), or with interspaces merging into ridges in some specimens, or ridged, densely punctate between ridges (female from Claudie River, Queensland and another from Santa Ysabel Island); side ridged, punctate between ridges, but ridges largely evanescent in specimen from New Britain; posterior surface ridged. Forecoxal venter finely, closely punctate and with larger punctures many diameters apart. Posteroventral forefemoral surface finely punctate, some punctures up to about three diameters apart. Hindcoxal dorsum with inner carina produced into conspicuous tooth basally. Punctures of tergum I well defined, minute and several diameters apart mesally. Sternum II sparsely punctate except closely punctate adjacent to lateral margin, impunctate apicomesally.

Setae silvery, erect on head, thorax, propodeum, femoral venters, and tergum I, not concealing integument on clypeus; genal setae sinuous, longest setae of lower gena about 1.3 × basal mandibular width.

Body all black, mandible dark reddish preapically.

♀.– Upper interocular distance equal to 0.54 × lower interocular distance; ocellocular distance equal to 0.3-0.6 × hindocellar diameter, distance between hindocelli equal to 0.5-0.7 × hindocellar

0.56-0.62 × lower interocular distance; ocellocular distance equal to 0.5-1.0 × hindocellar diameter, distance between hindocelli equal to 0.7-0.8 × hindocellar diameter; eye height equal to 1.00-1.06 × distance between eye notches. Free margin of clypeal lamella acutely angulate ( Fig. 910 ). Dorsal length of flagellomere I 2.7 × apical width, of flagellomere X 1.1-1.2 × apical width. Tergum VII compressed laterally to form an obtuse carina apicomedially ( Fig. 917 ). Sternum VIII with rounded apical projection ( Fig. 918 ). Genitalia: Figs. 919, 920 . Length 9.2-10.8 mm; head width 2.9-3.4 mm.

VARIATION.– At an early stage of this study I thought that the females of P. punctifrons represent two species, one, more numerous, with no carina above each antennal socket, a well-defined tooth on the trimmal carina of the mandible just above the incision ( Fig. 908 ), and the scutum sparsely punctate, the other species with an oblique carina above each antennal socket ( Fig. 912 ), no tooth on the trimmal carina of the mandible ( Fig. 909 ), and the scutum densely punctate.The troubling facts were that they occurred sympatrically in several localities, and that no male was available for the second form. The subsequent study of the rich material from the Bishop Museum demonstrated, however, that intermediates exist between all these character states and that the two forms are just one species.

GEOGRAPHIC DISTRIBUTION (Fig. 921).– Northeastern Australia, Maluku islands, New Britain, New Guinea, Solomon Islands. Recorded from Java by Cameron (1905), but his identification is not certain.

RECORDS.– AUSTRALIA: Northern Territory: Black Point in Cobourg Peninsula at 11°09ʹS 132°09ʹE (1 ♀, ANIC) ; Darwin: eastern suburb Berrimah (1 ♀, NTM) . Queensland: 8 km NW Bald Hill in Ilwraith Range at 13°45ʹS 143°22ʹE (3 ♀, ANIC) , 11 km NW Bald Hill at 13°44ʹS 143°20ʹE (23 ♀, 1 ♂, ANIC) ,

4 km NE Batavia Downs at 12°39ʹS 142°42ʹE (2 ♀,

ANIC), 5 km S Batavia Downs at 12°41ʹS 142°41ʹE

(1 ♂, ANIC), near Brisbane Forest Park at 27°26.0ʹS

152°55.4ʹE (1 ♀, CAS), Claudie River 1 mi. W (1 ♀,

1 ♂, AMS ) and 5 mi. W Mount Lamond (1 ♀,

AMS), Cockatoo Creek Crossing 17 km NW Heathlands at 11°39ʹS 142°27ʹE (1 ♀, ANIC) , Coen at

13°57ʹS 143°12ʹE (1 ♀, ANIC; 3 ♀, CAS), Cordalba State Forest 27 km S Bundaberg (1 ♀, ANIC) ,

Dividing Range 15 km W Captain Billy Creek (1 ♀,

CA), Heathlands at 11°45ʹS 142°35ʹE (2 ♀, ANIC),

14 km ENE Heathlands at 11°41ʹS 142°42ʹE (3 ♀,

1 ♂, ANIC), 15 km ENE Heathlands at 11°41ʹS

142°42ʹE (3 ♀, ANIC), 12 km NE Heathlands at

11°43ʹS 142°41ʹE (2 ♀, ANIC), 12 km SSE Heath- FIGURE 921. Collecting localities of Pison punctifrons lands at 11°51ʹS 142°38ʹE (2 ♀, 3 ♂, ANIC), Iron Shuckard .

Range : no specific locality (1 ♂, ANIC) , Leo Creek Road 30 km NE Coen (5 ♀, 3 ♂, ANIC; 1 ♂, CAS), Lockerbie area (2 ♀, 1 ♂, ANIC) , Middle Claudie River in Iron Range (2 ♀, AMS) , Moreton at Wenlock River at Moreton (1 ♀, 1 ♂, ANIC) , Mount Lamond in Iron Range (1 ♀, AMS) , 3 km ENE Mount Tozer at 12°44ʹS 143°14ʹE (1 ♀, ANIC; 1 ♀, CAS), 9 km ENE Mount Tozer at 12°43ʹS 143°17ʹE (2 ♀, 1 ♂, ANIC) , 11 km ENE Mount Tozer at 12°43ʹE 143°18ʹE (3 ♀, 1 ♂, ANIC) , 3 km NE Mount Webb at 15°03ʹS 145°09ʹE (2 ♀, ANIC) , 13 km SE Weipa at 12°40ʹS 143°00ʹE (2 ♀, ANIC) .

INDONESIA: Ambon: Waai (1 ♀, RMNH . Halmahera: between Payahe and Gila Woda (4 ♀, RMNH) . Maluku: Kai island (1 ♀, OXUM, lectotype of Pison nitidum ), Seram (formerly Ceram): Hatumete 15 km NNE Tehoru at 3°17ʹS 129°39ʹE (2 ♀, RMNH) and 9 km E Wahai (2 ♀, RMNH) , 11 km E Wahai (2 ♀, RMNH) , and 21 km E Wahai (1 ♀, RMNH) , Ternate ( Maindron , 1879) . Western Papua (= Indonesian New Guinea): Araucaria camp (1 ♀, 1 ♂, RMNH) , Bokondini (2 ♀, BISH) , Bomberi in Vogelkop Peninsula (2 ♀, BISH) , 11 km S Bupul at 7°39ʹS 140°53ʹE (5 ♀, 2 ♂, RMNH) , Eramboe 80 km NE from Merauke (1 ♀, BISH) , Genjam 40 km W Jayapura (2 ♀, BISH, as Hollandia), Ifar (1 ♀, RMNH) , Japen Island: Dawai River SSE Samberbaba (1 ♀, BISH) , Mimika River (1 ♀, BMNH, holotype of Pison constrictum ), Misool: no specific locality (1 ♀, BMNH; 1 ♂, RMNH), Nabire (1 ♀, BISH) , Paniai Lakes (as Wisselmeren): Enarotadi (2 ♀, BISH) , Rattan Camp (2 ♀, RMNH) , Waris S Jayapura (1 ♀, BISH) . Yapen Island: Mount Baduri (1 ♀, BMNH) .

PAPUA NEW GUINEA: Bougainville Province: bush E Buin (1 ♀, CAS) , Kukugai (1 ♀, BISH) , Simba Mission (1 ♀, BISH) . Central Province: Brown River (2 ♀, BISH) , Cape Rodney (1 ♀, BISH) , Ei Creek 30 km N Sogeri (2 ♀, RMNH) , SE Mamai E Port Glasgow (1 ♀, 1 ♂, BISH) , Roku (1 ♀, ANIC) . East Sepik Province: May River (1 ♀, BISH) . Eastern Highlands Province: Aiyra (2 ♀, BISH) , 22 km SE Okapa (4 ♀, BISH) . Gulf Province: Murua River (1 ♀, BISH) . Jiwaka Province: Tsenga in upper Jimi Valley . Madang Province: Baiteta 12 km NW Alexishafen at 5°00ʹS 145°45ʹE (2 ♀, CAS) , Duru 15 km SW Madang at 5°20ʹS 145°43ʹE (1 ♀, CAS) , Gogol River 12 km SW Madang at 5°20ʹS 145°42ʹE (6 ♀, CAS) , Karkar Island at 4°35ʹS 145°55ʹE (1 ♀, RMNH) , Madang (Tsuneki, 1983, as Friedrich-Wilhelmshafen ), Nagada Harbor 8 km N Madang at 5°09ʹS 145°48ʹE (1 ♀, CAS) , Nobonob Hill 7 km NW Madang at 5°10ʹS 145°45ʹE (2 ♀, CAS) , Saidor: Sibo in Finisterre Range (1 ♀, BISH) , Sapi Forest Reserve 30 km W Madang at 5°12ʹS 145°30ʹE (1 ♂, CAS) , Simbai at 5º17ʹS 144º26ʹE (1 ♀, CAS) , Tapo Creek 26 km SW Madang at 5°24ʹS 145°38ʹE (4 ♀, CAS) , Wanuma (1 ♀, BISH) . Manus Province: Rossum 6 km SE Lorengau (1 ♀, BISH) . Milne Bay Province: Milne Bay (1 ♀, BISH) , Woodland (= Murua) Island : Kulumadau Hill (1 ♀, BISH) . Morobe Province: Bulldog Road (1 ♀, BISH) , Bulolo River (3 ♀, BISH) , 10 km S Derim at 6°13ʹS 147°06ʹE (4 ♀, RMNH) , Finschhafen (7 ♀, BISH) , 14.4 km W Lae (1 ♀, BISH) , Lake Trist (1 ♀, BISH) , Mindik (1 ♀, BISH) , Mount Missim at 7°15ʹS 146°48ʹE (3 ♀, BISH) , Simbang (1 ♀, MTM) , Umboi Island : 8 km WNW Lab Lab (1 ♀, BISH) , Wau (31 ♀, BISH; 1 ♀, RMNH), Wau: Mount Kaindi at 2,100 m (1 ♀, RMNH) and at 2,350 m (3 ♀, BISH) . National Capital District: Laloki (1 ♀, BISH) . New Britain: Bainings in Gazelle Peninsula : Saint Paul (5 ♀, BISH) , Duke of York Islands : no specific locality (1 ♀, NHMW, lectotype of Pison collare ) ,

Illugi on upper Warangoi River in Gazelle Peninsula (1 ♀, BISH) , Jacquinot Bay (1 ♂, CAS) , Linga Linga plain in Willaumez Peninsula (1 ♀, BISH) , upper Warangoi River (1 ♀, BISH) , Yalom (2 ♀, ZMUC, CAS, holotype and paratype of Pison bismarckianum ) . Oro (= Northern) Province: Kokoda , 2000 feet (1 ♀, BMNH) , Kokoda-Pitoki (1 ♀, BISH) , Mount Suckling , 500 m (3 ♀, BISH) , Popondetta (1 ♀, BISH) . Southern Highlands Province: 8 km W Mendi (1 ♀, BISH) , Mount Giluve at 2,550 m (5 ♀, BISH) . Western Province: Kiunga (1 ♀, BISH) , Oriomo (1 ♀, BISH) . West Sepik Province: junction of Green and Sepik Rivers (1 ♀, BISH) , Torricelli Mountains (1 ♀, SAM) .

SOLOMON ISLANDS: Choiseul Island: Kitipi River (3 ♀, BISH) , Kolombangara River (1 ♀, BISH) , Malangona (8 ♀, BISH) . Fauro Island: Toumoa (2 ♀, BISH) . Guadalcanal: Doma-Ruanu’u (1 ♀, RMNH) , Guadalcanal and Florida (now Nggela) Islands (1 ♀, CAS) , Honiara (8 ♀, 1 ♂, BISH) , 35 km E Honiara (2 ♀, BISH) , Lunga River , bridge (1 ♀, BISH) , Mount Jonapau in Suta-Gold Ridge (1 ♀, BISH) , Paripao (1 ♀, BISH) , Tenaru River (1 ♀, BISH; 2 ♀, 1 ♂, CAS), locality illegible (1 ♂, RMNH) . Malaita Island: Dala (29 ♀, 1 ♂, BISH) , Su’u (1 ♀, RMNH) . New Georgia Islands: Gizo Island (1 ♀, AMS; 4 ♀, BISH), Kolombangara Island : Pepele (8 ♀, BISH) , New Georgia Island: Munda (2 ♀, BISH) , Ringgi Cove (1 ♀, BISH) , and Vella Lavella Island: Dobeli jungle (1 ♂, RMNH) , Pusisama (1 ♀, BISH) , and Ulo Crater (3 ♀, BISH) . Nggela Islands (formerly Florida Islands): Gairava (1 ♀, BISH) , Haleta (9 ♀, BISH) . Russell Islands: Pavuvu (2 ♀, BISH; 1 ♀, CAS), Pepesala (5 ♀, BISH) . San Cristobal Island: Bweinaniawarikiapu (1 ♀, BISH) , Kira-Kira (1 ♀, BISH) , Wugiroga 1 ♀, BISH ). Santa Ysabel: Sukapisu (1 ♀, BISH) , SE Tatamba (1 ♀, BISH) , no specific locality (1 ♀, BISH) . Tulagi Island : no specific locality (2 ♀, BISH; 7 ♀, 1 ♂, RMNH) .

Figures

FIGURES 908-913. Pison punctifrons Shuckard. (908) Clypeus and mandibles of female without supraantennal carina; (909) Clypeus and mandibles of female with supraantennal carina; (910) Male clypeus and mandibles; (911) Upper frons of female in frontal view showing supraantennal carina; (912) Female frons in profile showing supraantennal carina; (913) Female vertex showing positions of ocelli.

FIGURES 914-917. Pison punctifrons Shuckard. (914) Female tegula and adjacent scutum; (915) Propodel dorsum of female; (916) Female tergum I; (917) Male tergum VII.

FIGURES 918-920. Pison punctifrons Shuckard, male. (918) Sternum VIII (ventral surface); (919) Genitalia in dorsal view; (920) Genitalia in lateral view.