Cicadakiller

Tachysphex morosus (F. Smith)

Tachytes morosus F. Smith, 1858:18, . [Holotype: Indonesia: Sulawesi: no specific locality (OxFORD), examined. Transferred to Tachysphex by Bohart and Menke, 1976:275.].—Maindron, 1879:179 [nesting habits].

Tachysphex bengalensis.—Bingham, 1897:193 [revision; specimens from Burma, corrected by R. Turner, 1917b:198].—Williams, 1928:92 [Philippines, corrected to Tachysphex tinctipennis by Pulawski, 1975:311] [also authors who followed Williams's interpretation: Krombein, 1949:382 (in key), 393 (Mariana and Caroline Islands); Tsuneki, 1963:7 (Thailand; redescription); 1967:49 (Taiwan; regarded japonicus as a junior synonym of bengalensis); 1976:54 (Philippines; as bengalensis bengalensis); Iwata, 1964:369 (life history);Haneda, 1971:30 (Taiwan)].

Tachysphex tinctipennis Cameron, 1904:301, . [Holotype: , India: Assam: Khasia Hills (OxFORD), examined. New synonym.].—Pulawski, 1975:311 [interpretation of the species].—Bohart and Menke, 1976:277 [listed].—Tsuneki, 1977b:269 [Taiwan; as Tachysphex nigricolor tinctipennis, new status]; 1983:64 [Philippines; redescription; as full species], 68 [in key].

Tachysphex bituberculatus.—Weber, 1948:203 [Hawaii, corrected to Tachysphex bengalensis by Krombein, 1949:393].—Yoshimoto, 1960:331 [listed].

Tachysphex lihyuetanus Tsuneki, 1971:15, . [Holotype: , Taiwan: Nantou Prefecture: Lihyuetan (originally K. Tsuneki coll., now USNM), not examined. Synonymized with Tachysphex tinctipennis by Tsuneki, 1983:64.]

Tachysphex morosus.—Bohart and Menke, 1976:275 [listed].

Tachysphex tinctipennis titidzimaensis Tsuneki, 1984:4, , . [Holotype: , Japan: Ogasawara (= Bonin) Islands: Chichidzima: Mount Chuoh (K. Tsuneki personal collection, Mishima), not examined. Valid subspecies.]

DIAGNOSIS.—Tachysphex morosus is one of the many species in which the mesopleuron is shiny, with well-defined punctures; the setae are erect on the vertex and nearly so on the scutum and midfemoral venter (as in Figures 32, 33), inclined obliquely anterad on the propodeal dorsum; the gaster as well as legs are black; and the female tarsi are of the pompiliformis type. Species of this complex are difficult to distinguish, but morosus differs as follows:

1. From most species with the above characteristics, by an almost flat middle clypeal section in the female (bevel shorter than basomedian area) and the presence of a well-defined foretarsal rake in almost all males.

2. From nigricolor Dalla Torre (which occurs in China: Beijing; Korea, Japan, Ryukyu Islands, Taiwan), in having relatively long scutal setae, a distinctive male clypeus (Figures 15, 17), and a foretarsal rake present in almost all males; the all black or mesally dark reddish mandible helps in identification. Specifically, the setal length, anterolaterally on scutum, is about 0.3–0.4 × basal mandibular width; in the male, the apical portion of the clypeal lobe (including lip) is situated basically in the same plane as the remaining clypeus; the male forebasitarsus has two to five rake spines, of which at least one is longer than basitarsal width; and the apicoexternal spine of foretarsomere II is as long as foretarsomere III or longer. In nigricolor, the setal length, anterolaterally on scutum, is about 0.2 × basal mandibular width, an obvious difference when specimens are compared; in the male, the apical portion of the clypeal lobe (including lip) is bent toward the back of the head and separated from the remaining surface by an inflection; the male foretarsomere has no rake spines or at most one minute spine near base; and the apicoexternal spine of foretarsomere II is shorter than tarsomere III. The two species are largely allopatric, but occur together in Taiwan. According to Tsuneki (1983), the mandible is “usually wholly black” in morosus while “always medianly broadly reddish” in nigricolor, but in fact this difference is not very reliable. For example, the mandible is as dark in two paratypes of nigricolor yaeyamanus Tsuneki (CAS) as it is in most morosus. Two other characters mentioned by Tsuneki are also variable (microsculpture of mesopleural interspaces and propodeal ridges).

3. From the female of sri (an endemic of Sri Lanka), by the long vertex setae (setal length about two midocellar diameters in morosus, one diameter in sri).

4. From formosanus Tsuneki (which is known from Taiwan and Ryukyu Islands), by subtle characters that probably intergrade. In morosus, the scutal setae are as described under 2 above and the mandible is black or dark reddish mesally. In formosanus, the scutal setae are insignificantly shorter (equal to 0.2–0.3 of basal mandibular width) and slightly more inclined posterad, and the mandible is yellowish red mesally (particularly on the anterior face). There are other differences according to Tsuneki (1983), but I have observed full intergradation: the tarsal apex (black or dark brown in morosus, reddish brown in formosanus), color and configuration of the female rake spines, and male forefemoral notch. The species overlap in Taiwan.

5. From fugax (Radoszkowski), which occurs in Africa, southern Europe, east to Tajikistan, in having a shorter female flagellum. For example, flagellomere IV is 3.5–3.6 as wide as long in morosus and 4.0–5.0 in fugax. There are no constant differences between males, although the flagellum averages longer in fugax. Males of fugax from southeastern Europe, however, are distinctive: length of flagellomere III is about 0.6–0.7 of IV and the apicoexternal spine of tarsomere II is markedly shorter than tarsomere III. In morosus and most fugax, length of flagellomere III is 0.9 of IV, and apicoexternal spine of foretarsomere II is as long as tarsomere IV or longer.

6. From helveticus Kohl (which occurs in the western Palearctic to Mongolia), by a combination of characters: scutal setae as under 2 above, mandible black or dark reddish mesally, mesopleural interspaces shiny, apical silvery fascia on terga I–III (I–IV in occasional specimens). In helveticus, scutal setae are shorter in the female of the nominotypical subspecies (0.2 × basal mandibular width) and longer in the male of helveticus aegyptiacus Morice (0.5 X), the mandible is yellowish red in helveticus aegyptiacus (Egypt), the mesopleural interspaces are microsculptured in the female and many males of helveticus helveticus and helveticus quadrifasciatus Pulawski, and the apical fascia is present on terga I–IV in helveticus quadrifasciatus Pulawski (Cyprus, Jordan, Tajikistan).

7. From angustatus Pulawski (which is known from Greece, Turkey, Iran, Transcaspia, and Mongolia), by the broad clypeal lobe of male and finely punctate sterna in both sexes. In the male, the distance between the orbit and corner of the clypeal lobe is more than the clypeal midline in morosus, but less than that in angustatus. Sternal punctures are markedly larger in angustatus than in morosus, a feature that is easily seen but difficult to describe or to quantify.

DESCRIPTION.—Scutal punctures well defined, about one diameter apart on disk, less than that near margins. Mesopleural punctures well defined, averaging about two to three diameters apart beneath scrobe (more than that in some specimens); interspaces shiny, also posteriorly. Episternal sulcus complete, but evanescent anteroventrally in many specimens. Propodeal dorsum rugose or irregularly ridged; side ridged. Hindcoxal dorsum: inner margin carinate basally, carina practically not expanded. Apical tarsomeres without spines on venter or lateral margins.

Setae (figures in parentheses refer to setal length expressed as fraction of basal mandibular width): erect on vertex (0.4–0.5) and between mandibular base and occipital carina (0.4), slightly inclined posterad on scutum mesally (0.3–0.4 anteriorly), suberect and inclined posterad on mesopleuron (0.4); inclined obliquely anterad on propodeal dorsum.

Body black, tarsal apex dark brown. Frontal vestiture silvery. Gastral segments I–III silvery fasciate apically (I–IV in a male from Kokmotte area, Sri Lanka). Wings moderately infumate.

.—Clypeus (Figures 14, 16): middle section almost flat, with no inflection between basomedian area and bevel; the latter shorter than basomedian area, not extending laterad to lobe corner; free margin of lobe arcuate to obtusely pointed mesally. Vertex width 1.5–1.6 × length. Dorsal length of flagellomere I 2.0–2.3 × apical width. Outer surface of foretibia with one or two spines. Tarsi of pompiliformis type, forebasitarsus with five to seven rake spines. Pygidial plate shiny, with punctures that concentrate mainly along margins. Length 8.5–10.5 mm.

.—Innermandibular margin with tooth. Clypeus (Figures 15, 17): bevel absent; lip of equal length mesally and laterally, its free margin straight, arcuate, sinuate, or insignificantly concave; distance between lip corners 0.7–0.9 × distance between corner and orbit; corners prominent. Vertex width 1.6–1.8 × length. Dorsal length of flagellomere I 1.1–1.4 × apical width. Forefemoral notch with microsculptured, asetose bottom (Figures 21, 22). Foretarsus with rake in vast majority of specimens (Figure 23): outer margin of forebasitarsus with three to five preapical spines, apical spine of tarsomere II as long as tarsomere III or longer (one male from Chiang Mai has two preapical spines on left basitarsus, but only one minuscule spine on right one; one male from Phoenix Islands has no preapical spines). Tergum VII with minute punctures that are several diameters apart basomedially. Sterna densely, evenly punctate and setose. Volsella: Figure 19. Penis valve: Figure 20. Length 5.5–8.5 mm.

COLLECTING PERIOD.—All months except April and December.

HABITAT.—The species occurs in all three ecological zones in Sri Lanka at altitudes ranging from near sea level to an elevation of 1880 m, in localities with an average annual rainfall ranging from 970 to 4900 mm (Figure 24).

GEOGRAPHIC DISTRIBUTION.—Widely distributed in the Oriental Region and in Oceania, ranging from Sri Lanka, Sumatra, and New Guinea in the south to Nepal, China (Sichuan), and Taiwan in the north, and to Micronesia, Fiji, and Hawaii in the east. Quite possibly it was introduced into the last three areas by commerce.

RECORDS.—BHUTAN: Wangdu (1, CAS).

CHINA: SICHUAN: Hanyuan [as Fulin] (1, USNM).

FIJI: VITI LEVU: Korotogo, 8 km E Sigatoka (13, 20, CAS).

HONG KONG: Hong Kong Island: Potfulan (1, CAS).

INDIA: ASSAM: Khasia Hills (1, OXFORD, holotype of tinctipennis). BIHAR: Tirhut [as Tirhoot] (Bingham, 1897). KARNATAKA: Mudigere area (2,, 1 CAS; 5, 2, ZMK). KERALA: Walayar forests (1, USNM). TAMIL NADU: Anamalai Hills, Cinchona (1, 1, CAS; 2, 1, FSAG; 3, 1, OSU); Nilgiri Hills, Naduvatam (1, OSU); Nilgiri Hills, Cherangode (1, USNM); Shevaroy Hills, Yercano (2, OSU).

INDONESIA: IRIAN JAYA: Kebar, W Manokwari in Vogelkop (1, BISH). MALUKU: Ambon Island: no specific locality (2, CAS). SULAWESI: Senkang (1, 3, CAS; 5, 12, RMNH), no specific locality (1, OXFORD, holotype of morosus). WEST SUMATRA: Padang (1, 1, RMNH).

LAOS: VIENTIANE PROVINCE: Ban Van Eu (2, BISH), GiSion Vill., de Tha Ngon (1, BISH; 1, CAS), Phou Khao Khouai (1, BISH). SAVANNAHKET PROVINCE: Savannahket (2 , BISH).

MALAYSIA: Kuala Lumpur (1, BISH).

MYANMAR (Burma): Tenasserim (Bingham, 1897).

NEPAL: Arun Valley, Sabhaya River below Tumlingtar (2 , BMNH; 1, CAS), Sysbrubens, 35 km N Trisuli (1, BISH).

PACIFIC OCEAN ISLANDS: CAROLINE ISLANDS: Peleliu Island: Palaus (Krombein, 1949). HAWAIIAN ISLANDS: Oahu: Hickam Field (1, CAS), Honolulu (Weber, 1948). MARIANA ISLANDS: Saipan, Susupe (2, 2, BISH). PHOENIX ISLANDS: Canton Atoll (1, CAS; 3, 1, USNM).

PAPUA NEW GUINEA: MADANG: Nagada Harbor, 8 km N Madang (3, CAS), Sapi Forest Reserve, 30 km W Madang, 5°12′S, 145°30′ (1, CAS). MOROBE: Wau (2, USNM).

PHILIPPINES (Tsuneki, 1983, if not indicated otherwise): CEBU: Argao. LUZON: Albay Province: Tabaco. Camarinessur Province: Baao and Bato. Laguna Province: Alaminos, Los Baños (1, RMNH; 1, BISH), Mount Makiling (1, USNM), Pagsanjan. La Union Province: Baguio, Naguilian, St. Fernando. Manila Province: Manila (1, USNM). Mountain Province: Bontoc. MINDANAO: Bukidnon, Davao, Zamboanga. NEGROS: Kalaukalau (8, BISH), Mambucal, Taytay beach. PALAWAN: Uring Uring: Brookes Point, 8°47′, 117°50′E (3, ZMK).

SINGAPORE: Singapore (1, 2, CAS).

SRI LANKA: AMPARAI DISTRICT: Ekgal Aru Reservoir Jungle (1, USNM, Biological Note 61076D; 5, 2, USNM), Ekgal Aru tank (1, USNM). ANURADHAPURA DISTRICT: Padaviya (1, USNM), Ritigala Natural Reserve (2, 1, USNM), Wildlife Sanctuary Bungalow, Hunuwilagama in Wilpattu National Park (1, USNM). BADULLA DISTRICT: Ella (2, USNM). COLOMBO DISTRICT: Gampaha Botanical Garden (2, USNM); Colombo, Museum Gardens (1, USNM), Labugarna Reservoir Jungle (1, USNM). GALLE DISTRICT: Kanneliya Jungle (1, USNM). HAMBANTOTA DISTRICT: Palatupana (1, USNM). KANDY DISTRICT: Hasalaka Circuit Bungalow (1, USNM); Kandy (5, 3, USNM); Kandy, Peak View Motel (1, CAS); Kandy, Udawattakele Sanctuary (1, CAS; 5, USNM); Peradeniya, Botanical Gardens (1, USNM), Thawalamtenne (3, USNM). KEGALLA DISTRICT: Kitulgala in Bandarakele Jungle (1, USNM); Kitulgala, Makande Mukalana (1, USNM). KURUNEGALA DISTRICT: Kurunegala, Athugala (Elephant Rock) (1, USNM), Kurunegala, Badegamuwa Jungle (1, USNM). MANNAR DISTRICT: 0.5 mi (0.8 km) NE Kokmotte in Wilpattu National Park (2, 1, USNM); Marichchukkaddi (1, COLOMBO); Silavaturai, Kondachchi (1, COLOMBO; 1, USNM). MONARA GALA DISTRICT: Angunakolapelessa (2, USNM), 13 mi (20.8 km) E Uda Walawe (1, USNM). NUWARA ELIYA DISTRICT: Hakgala Botanical Gardens (3, 1, USNM), Hakgala Natural Reserve (8, 16, USNM). RATNAPURA DISTRICT: Rajawaka (1, USNM); Weddagala in Sinharaja Forest (1, USNM, Biological Note 62176); Sinharaja Forest, 2 mi (3.2 km) S Weddagala (2, 3, USNM); Sinharaja Forest, 3 mi (4.8 km) S Weddaggala (1, USNM). TRINCOMALEE DISTRICT: Trincomalee, China Bay (1, 1, USNM); Trincomalee, China Bay Ridge Bungalow (1, CAS; 4, 1, USNM).

TAIWAN (Tsuneki, 1967, if not indicated otherwise): CHIAYI PREFECTURE: Chuchi. HUALIEN PREFECTURE: Liyuchih. ILAN PREFECTURE: Tsukeng. NANTOU PREFECTURE: Lihyuetan, Penpuchi. PINGTUNG PREFECTURE: Hengchun, Kentin, Suchungchi. TAIPEI PREFECTURE: Yangmingshan. TAITUNG PREFECTURE: Chulu, Taitung, Taoyeh. TAOYUAN PREFECTURE: Taoyuan (Haneda, 1971). Also (Tsuneki, 1977b): Takao and Taihorinsho.

THAILAND: BANGKOK: Bangkok (1, OSU). CHIANG MAI Chiang Mai: (2, 16, CAS), Huai Tung Tao, 10 km N Chiang Mai (1, CAS). KANCHANABURI: Kanchanaburi (1, 3 CAS), Lam Ta Pen River bank, 5 km NW Lat Ya (7, CAS). LOEI: Loei (6, 2, CAS). RAYONG: Ko Samet Island (1, 2,CAS).

Tachysphex morosus (F. Smith)

Maindron (1879) was the first author to report the nesting habits of morosus, but his identification of the species is not certain.

NEST CONSTRUCTION.—We found this wasp, 9.2 mm long, nesting at 1515 on 10 June 1976, in Ekgal Aru Sanctuary Jungle, Amparai District, in the Dry Zone. The nest was in a flat, bare sandy spot along a jeep trail through the jungle. The wasp was closing the burrow entrance by kicking sand backward behind her. We disturbed the wasp, after which she flew around for a few seconds, and then alighted at the entrance, at which time we collected her.

We noted another female morosus, 9.4 mm long, excavating a nest in the rain forest at 1100 on 21 June 1976, near Weddagala, Sinharaja Jungle, Ratnapura District. The nest was located on a slight slope of a logging road made from a mixture of sand and fine gravel, a nesting medium that made it impossible to trace the exact nest architecture. The wasp was filling the burrow entrance when we returned to the nesting site at 1500, at which time we collected her and dug up the nest.

NEST STRUCTURE.—The burrow of the Ekgal Aru nest went downward into sandy loam at an angle of 20° for 3 cm, and ended in a horizontal cell, 12 mm long and 8 mm wide, containing four prey, the innermost bearing the wasp egg.

In Weddagala a grass stem probe penetrated the burrow at a slight angle for 6 cm. We found a completed cell 3 cm from the burrow axis; it contained three grasshoppers, one with an egg. A cell at the end of the burrow held three grasshoppers, one bearing the wasp egg. A third completed cell was on the other side of the burrow about 3 cm from the entrance; it contained four prey, one bearing a wasp egg. It is likely that this third cell was the oldest of these three, because the egg hatched that evening. A fourth cell was on the left side of the burrow, and about 3 cm from the entrance; it contained two prey, one with an egg, and a tiny dipterous maggot.

We continued digging and found three older cells that may have been made by this wasp. Considering the stage of larval development in each of these cells, it is probable that this was a separate nest. The first cell was 6 cm from the burrow entrance and about 8 cm from the burrow axis; it held two prey and a wasp larva half grown. A second cell was 17 cm from the end of the burrow and at the same depth; it contained four prey and a half grown wasp larva. The third cell was 5 cm from the second, and contained a nearly mature wasp larva and the remains of two prey.

PREY.—The grasshopper nymphs in the nest at Ekgal Aru consisted of two males and a female of Stenocatantops splendens (Thunberg) (Acrididae, Catantopinae), 8.5–9.5 mm long, and a female of a species of Chorotypus (Eumastacidae), 6.0 mm long.

The 21 nymphal prey in the Weddagala cells belonged to a single species of Mesambria (Acrididae, Catantopinae), and were 6–11 mm long. Both sexes were preyed upon, and a single cell contained one or more specimens of both sexes. Some of the prey had recovered partially from the wasp sting. They were able to get up on their legs, but could not walk.

IMMATURE STAGES.—Two eggs were 1.6–1.9 mm long and 0.4 mm wide. The egg was attached on the sternum of a prey behind either the left or right forecoxa and extended transversely to the right or left, respectively. A newly hatched larva at Weddagala was feeding through the intersegmental membrane behind the left forecoxa.

NEST ASSOCIATES.—We saw no adult parasitic miltogrammine flies (Sarcophagidae) around the nesting site. The tiny maggot that we recovered from one cell was too small to have been that of a miltogrammine, and was probably that of a commensal species of Phoridae.