Shore Fly

Hecamede (Hecamede) inermis Malloch

Hecamede inermis Malloch, 1933:12.—Cresson, 1948:23 [note].

Hecamede granifera.—Of authors [misidentification].—Mathis, 1989a:641 [Australasian/Oceanian catalog]: 1989b:8 [list of Diptera from the Pitcairn Group].

DESCRIPTION.—Small to medium-sized shore flies, length 1.30 to 3.00 mm.

Head: Fronto-orbits narrow, gray, or indistinct, with similar coloration as mesofrons; ocellar triangle densely microtomentose, mostly tan, occasionally mostly gray; mesofrons less densely microtomentose than ocellar triangle, coloration darker posteriorly with some mostly brown to black coloration, becoming yellowish orange to reddish orange anteriorly; proclinate fronto-orbital setae 1, inserted anterolaterad of reclinate seta. Antenna usually mostly yellow; flagellomere 1 usually darker, sometimes extensively so, especially dorsobasally near insertion of arista; arista with 4 dorsal branches. Face mostly whitish gray, frequently with some yellowish coloration, especially at sutures; facial setae 3 or 4, arising from bare, slightly elevated spots within densely microtomentose face; facial prominence bare, not much larger than shiny area at base of facial setae; gena mostly concolorous with face, bearing 1 larger seta and a few other black setae, these scattered; gena high, eye-to-cheek ratio averaging 0.50. Clypeus black, densely invested with gray microtomentum.

Thorax: Mesonotum usually mostly light to moderately brown, frequently with some faint mostly red or golden coloration, sometimes mostly gray, nearly always darker than pleural areas, if dark, becoming grayer, whitish to silvery gray, especially ventrad through notopleuron and dorsal half of anepisternum; presutural seta and usually prescutellar acrostichal setae well developed, some specimens with indistinct prescutellar setae; posterior notopleural seta inserted distinctly above level of anterior seta; pleural sclerites mostly gray to whitish gray, dorsal portion of anepisternum sometimes faintly to conspicuously tan; anepisternal setulae mostly on dorsal , occupying an area that is more or less triangular, with a line of setulae to ventral portion of pleuron where several more setulae are inserted. Coxae and trochanters lightly blackish gray to gray; middle and hind femora gray anteriorly, posterior surface blackish; fore femur lacking a row of setae along anteroventral margin, anterior surface mostly blackish, mostly gray posteriorly; tibiae mostly yellow, lightly invested with whitish gray microtomentum; fore basitarsus lacking black setae toward base of posteroventral surface; apical tarsomeres usually yellow, occasionally with apical tarsomere lightly brown. Wing with costal vein ratio averaging 0.35; M vein ratio averaging 0.60; last section of vein M shallowly arched posteriorly.

Abdomen: Male terminalia (Figures 34–36) as follows: 5th tergum with anterior to more lightly sclerotized and bearing numerous, evenly scattered spicules, from dorsal view with anterior margin broadly bifid, posterior to a sclerotized band bearing 8 to 11 setae along posterolateral margin, with 2 globular apodemes anterobasally; epandrium (Figure 34) a dorsal band, gradually enlarged posteriorly; surstylus (Figures 34, 35) in lateral view long and narrow, posterior margin more or less straight, anterior margin sinuous, anteroapical margin curved, apex bearing approximately 6–7 moderately longer, sharply pointed setulae; cercus (Figure 34) long and narrow, length extended nearly to apex of surstylus, bearing a conspicuously longer seta at posteroventral margin; gonite (Figure 36) roughly triangular in shape, with a narrow connection above aedeagus, ventral angle as a narrowed, short, parallel-sided, blunt process; aedeagal apodeme (Figure 36) allantoid in lateral view, slightly broader toward attachment with hypandrium; aedeagus (Figure 36) long and narrow, tapered gradually to apex, shaped like a pipe, with right angle curve at basal to ; hypandrium slightly emarginate anteriorly.

TYPE MATERIAL.—The holotype male of Hecamede inermis is labeled “Tahauku, HivaOa VII-10-[19]29 X[crossing out a letter]/sea shore/Marquesas Islands/Mumford & Adamson/Pacific Entomological Survey/Hecamede inermis 1226 [number in red ink] Type det. JRMALLOCH [species name and number handwritten, black submargin].” The holotype is pointed, is in good condition, and is deposited in the BBM.

OTHER SPECIMENS EXAMINED.—Australasian/Oceanian: MARQUESAS. Hiva Oa: Tahauku, 10 Dec 1929, E.P. Mumford, A.M. Adamson (2, paratypes; BBM, USNM). Nuku Hiva: Taiohae, 9–15 Jun 1987, W.N. Mathis (1, 3; USNM).

PITCAIRN GROUP. Henderson Island: Northwest Beach, 12–20 May 1987, W.N. Mathis (88, 11; USNM); North Beach, 17–20 May 1987, W.N. Mathis (56, 10; USNM). Oeno Atoll, 28 May 1987, W.N. Mathis (34, 9; USNM). Pitcairn Island: Adamstown, 22–26 May 1987, W.N. Mathis (7, 3; USNM); Down Rope, 25 May 1987, W.N. Mathis (9, 6; USNM).

WAKE ISLAND. 8 Jan 1923, E.H. Bryan, Jr. (1, 1; ANSP).

DISTRIBUTION (Figure 37).—Australasian/Oceanian: South Pacific from Wake Island and the Marquesas south to the Pitcairn Group (Henderson, Pitcairn, and Oeno islands).

NATURAL HISTORY.—On islands of the Pitcairn Group, this species was abundant on the beach, especially where debris had accumulated at the high tide mark (Mathis, 1989b). Adults were also attracted in large numbers to specimens of corals and study skins of birds that had been set out to dry on exposed rocks.

DIAGNOSIS.—This species is distinguished from congeners by the following combination of characters: proclinate fronto-orbital seta 1, inserted anterolaterad of reclinate seta; gena with 1 larger seta and several smaller black setae; fore femur lacking a distinct row of setae along anteroventral margin; fore basitarsus lacking black setae at base of posteroventral surface; mesonotum golden brown to brown; 5th tergum of male appearing hirsute, invested with numerous, long, microtrichia; and the structure of the male terminalia (Figures 34–36), especially the shape of the surstylus and its setulae.

NOTES ON NOMENCLATURE.—Previously (Mathis, 1989b:8), I commented that “The genus Hecamede needs revision before many of the available names can be validly applied. For the present, I am using H. granifera for this species in accordance with the recent catalog of Australasian and Oceanian Diptera (Mathis, 1989a). A junior synonym, H. inermis Malloch, was used previously for the common species of Hecamede that occurs in the South Pacific.”

Hecamede (Hecamede) granifera (Thomson)

Notiphila granifera Thomson, 1869:594.—Becker, 1896:271 [list].—Jones, 1906:176 [list].—Cresson, 1945:56 [footnote, as grandifera (sic)]; 1948:18 [suggested to be in the genus Hecamede].

Hecamede granifera.—Cogan and Wirth, 1977:325 [synonymy with H. persimilis, generic combination].—Mathis, 1989a:641 [Australasian/Oceanian catalog].

Hecamede lacteipennis Lamb, 1912:318.—Bezzi and Lamb, 1926:561.—Cresson, 1946b:248 [review].—Canzoneri and Meneghini, 1969:1483 [list, Senegal].—Cogan, 1980:657 [Afrotropical catalog]. [New synonym.]

Hecamede persimilis Hendel, 1913:99.—Cresson, 1925:229 [discussion]; 1945:56 [list, Indoaustralian]; 1948:23 [note on synonymy].—Bohart and Gressitt, 1951:84–85 [review, natural history].—Adachi, 1952:353 [synonymy].—Hardy, 1952:468 [synonymy].—Cogan and Wirth, 1977:325 [Oriental catalog].—Tenorio, 1980:275–276 [review, list, Hawaii, notes on natural history], 318–319 [figures of larva and puparium], 339 [description of larva and puparium].

Hecamede nivea de Meijere, 1916:61.—Cresson, 1948:23 [synonymy with H. persimilis].

Hecamede femoralis Malloch, 1930:245.—Wirth, 1947:21 [list, Hawaii].—Cresson, 1948:23 [synonymy with H. persimilis].—Miller, 1950:111 [New Zealand catalog].—Adachi, 1952:353 [synonymy].—Harrison, 1959:219–220 [description, list, New Zealand].

Hecamede albicans.—Of authors [misidentification].—Bryan, 1934:433 [list, Kahoolawe].—Miyagi, 1977:7–8 [probably H. granifera].

DESCRIPTION.—Small to medium-sized shore flies, length 1.60 to 3.20 mm.

Head (Figures 19–24): Fronto-orbits and ocellar triangle densely microtomentose, silvery gray to tan with some golden coloration; mesofrons less densely microtomentose, coloration darker posteriorly with some mostly brown to black coloration, becoming yellowish orange to orange anteriorly; proclinate fronto-orbital setae 1, inserted anterolaterad of reclinate seta. Antenna mostly yellow; flagellomere 1 slightly darker, especially dorsobasally near insertion of arista; arista with 4 dorsal branches. Facial setae 3 or 4; facial prominence bare, not much larger than shiny area at base of facial setae. Gena bearing 1 large and several smaller black setae, these scattered; gena high, eye-to-cheek ratio averaging 0.45. Clypeus black with an investment of gray microtomentum.

Thorax (Figures 25–28): Mesonotum golden tan to whitish gray, usually darker than pleural areas, if tan, lateral margins becoming grayer, whitish to silvery gray, especially ventral through notopleuron and dorsal half of anepisternum; presutural seta and prescutellar acrostichal setae well developed; posterior notopleural seta inserted distinctly above level of anterior seta; pleural sclerites mostly gray to whitish gray, dorsal portion of anepisternum sometimes faintly tan; anepisternal setulae mostly on dorsal , occupying an area that is more or less triangular, with a line of setulae to ventral portion of pleuron where several more setulae are inserted. Coxae and trochanters lightly blackish gray to gray; middle and hind femora gray anteriorly, posterior surface blackish; fore femur bearing distinct row of setae along anteroventral margin, anterior surface mostly blackish, mostly gray posteriorly; tibiae, especially the fore tibia, mostly yellow but moderately densely invested with whitish gray microtomentum, appearing whitish gray, only femoral/tibial articulation mostly yellow; fore basitarsus with 2 or 3 black setae, these inserted toward base of posteroventral surface; apical 2 tarsomeres blackish brown. Wing with costal vein ratio averaging 0.38; M vein ratio averaging 0.55; last section of vein M shallowly arched posteriorly.

Abdomen: Male terminalia (Figures 29–32) as follows: 5th tergum with anterior to more lightly sclerotized and bearing numerous, evenly scattered spicules, from dorsal view with anterior margin broadly bifid, posterior to a sclerotized band bearing 8 to 11 setae along posterolateral margin, with 2 globular apodemes anterobasally; epandrium (Figure 29) a dorsal band, gradually enlarged posteriorly; surstylus (Figures 29–31) in lateral view long and narrow, posterior margin more or less straight, anterior margin sinuous, anteroapical margin curved, apex bearing approximately 6–7 moderately long, broad setulae; cercus (Figure 29) long and narrow, length extended nearly to apex of surstylus, bearing a conspicuously longer seta at posteroventral margin; gonite (Figure 32) roughly triangular in shape, with a narrow connection above aedeagus, ventral angle a narrowed, short, parallel-sided, blunt process; aedeagal apodeme (Figure 32) allantoid in lateral view, slightly broader toward attachment with hypandrium; aedeagus (Figure 32) long and narrow, tapered gradually to apex, shaped like a pipe, with right angle curve at basal to ; hypandrium slightly emarginate anteriorly.

TYPE MATERIAL. The holotype female of Notiphila granifera is labeled “Ins. Ross. [Insulae Rossii (Cocos or Keeling Islands), as noted by Cogan and Wirth (1977)]/Kinb/Typus/472 73 [“73” handwritten, pink]/Hecamede granifera (Thom.) det. B.H. Cogan 1976. [handwritten except for “det. B.H. Cogan 19”]/HOLOTYPE Notiphila granifera Thomson det. B.H. Cogan 1976. [handwritten except for “det. B.H. Cogan 19”, red border]/Naturhistoriska Riksmusoet Stockholm Loan no 523/91 [light green].” The holotype is pinned directly, is in poor condition (wings, right legs, most dorsal setae, and antennae except for scapes missing), and is deposited in the NRS.

The three syntype females of H. lacteipennis are labeled “Type [round label with a red border]/Mahe [Long Island (seaweed on beach), Jul 1908], ’08–9. Seychelles Exp [glued to a rectangular, larger label, the underside of which has “Seaweed” handwritten in pencil]/Seychelles Is. Prof. J.S. Gardiner. 1914–537/TYPE [blue label glued to the larger determination label] S. Hecamede lacteipennis n. sp. CGL [except for the “TYPE” label, handwritten]. The syntypes are double mounted on a single block of cork (pin mounted in a paper-covered, rectangular block of cork with the number “107” handwritten on top), are in good condition, and are in the BMNH. ASTOVE. 1907, H.P. Thomasset (4; BMNH). Lamb (1912:319) also reported syntypes of this species from Astove Island, 1907, H.P. Thomasset and from Coetivy, 24 Sep 1905.

The lectotype male of H. nivea, designated herein, is labeled “Edw. Jacobson Pulu Bahi.Sim Sum.4, 1913 [black submargin]/Hecamede nivea det de Meijere. Type [black submargin, species name and “type” handwritten]/Lectotype [red]/Lectotype Hecamede nivea de Meij. det. B.H. Cogan 1971 [all but “det. B.H. Cogan 197” handwritten].” The lectotype is double mounted (minuten in foam rectangular block), is in good condition (the abdomen has been removed and dissected; the parts are in an attached microvial), and is deposited in the ZMA. My study of this specimen reconfirms the synonymy of this species with H. granifera. Although this specimen bears Cogan's lectotype label, it was never so designated in the literature. Data on the lectotype are as follows: 3.17 mm in length (before the abdomen was removed), the M vein ratio is 0.62, and the costal vein ratio is 0.35.

The lectotype male of H. persimilis Hendel, designated herein, is labeled “Tainan Formosa H. Sauter 11.09 [Nov 1909]/Hecamede persimilis H[endel]. det. Hendel [species name and “H.” handwritten]/Coll. Hendel/LECTOTYPE Hecamede persimilis Hendel By W.N. Mathis 1991 [“LECTOTYPE” and “By” in red, name, designator, and year handwritten, black sub–border].” Mounted with the lectotype is a paralectotype female. Other paralectotypes are as follows: the type locality (2); Anping (2). The lectotype is double mounted (minuten in a plastic foam block), is in poor condition (the left wing and most of the forelegs are missing, the abdomen is greasy), and is deposited in the NMW. I selected this specimen as the lectotype as it is the only male that remains from the original syntype series in NMW.

The holotype female of H. femoralis is labeled “Nelson N[ew].Z[ealand]. 20.II.24 [20 Feb 1924, not Nov as stated in the original description] A. Tonnoir Beach [date and “Beach” handwritten]/Hecamede femoralis Type Det. J.R. Malloch [species name and “Type” handwritten, black submargin].” The holotype is double mounted (minuten wired to main pin), is in good condition (head partially greasy), and is deposited in the NZAC. Although originally deposited in the Cawthom Institute (Nelson, New Zealand), it is now at the NZAC.

OTHER SPECIMENS EXAMINED.—Afrotropical: ALDABRA. Grande Terre: Anse Mais, 17 Mar 1986, W.N. Mathis (6, 14; USNM); Anse Takamaka, 7 Apr 1986, W.N. Mathis (11, 11; USNM); Cinq Cases, 12–13 Mar 1986, W.N. Mathis (8, 21; USNM). Malabar: Passe Houareau, 14–18 Mar 1986, W.N. Mathis (27, 21; USNM). Picard: La Gigi, 19–24 Mar 1986, W.N. Mathis (28, 9; USNM); Settlement, 15–21 Mar 1986, W.N. Mathis (44, 15; USNM).

ASTOVE. Around coconut plantation, 5 Mar 1968, B.H. Cogan, A.M. Hutson ( undetermined number, 2; BMNH).

COSMOLEDO. Menai Island: Station, 26 Mar 1986, W.N. Mathis (18, 9; USNM).

KENYA. Gazi (60 km S Mombasa, Rt. A14), 5 May 1991, A. Freidberg, F. Kaplan (18, 14; USNM). Mombasa, Yadini Beach, 15 Jan 1983, A. Valdenberg (4, 1; TAU, USNM). Mombasa (45 km S), 5 May 1991, A. Freidberg, F. Kaplan (1; USNM). Ngomeni (150 km N Mombasa), 4 Dec 1989, A. Freidberg, F. Kaplan (14, 8; USNM).

MADAGASCAR. Antseranana: Nosy Be, Ambatoloaka (beach), 4–7 Apr 1991, A. Freidberg, F. Kaplan (20, 14, TAU, USNM); Nosy Komba, 6 Apr 1991, A. Freidberg, F. Kaplan (6, 3; TAU, USNM); Nosy Tanikely, 6 Apr 1991, A. Freidberg, F. Kaplan (5, 4; TAU, USNM); Ramena, 10 Apr 1991, A. Freidberg, F. Kaplan (7, 3; TAU, USNM); Sambava (beach), 14 Apr 1991, A. Freidberg, F. Kaplan (5, 3; Tau, USNM). Toliara: Fort Dauphin (beach), 18 Apr 1991, A. Freidberg, F. Kaplan (11, 13; TAU, USNM); Fort Dauphin, Taolanaro (beach), 18–23 Apr 1991, A. Freidberg, F. Kaplan (1, 2; TAU, USNM); Fort Dauphin, Libanona (beach), 20 Apr 1991, A. Freidberg, F. Kaplan (4, 2; TAU, USNM); Morondava, 28–31 Mar 1990, W.E. Steiner, C. Kremen, V. Razafimahatratra (120, 9; TAU, USNM).

MOZAMBIQUE. Laurenco Marques, Polana Beach, Jan 1956, B. Stuckenberg (3, 1; USNM).

SEYCHELLES. Mahé: Anse aux Pins, 2 Apr 1986, W.N. Mathis (1; USNM); Anse Takamaka, 7 Apr 1986, W.N. Mathis (10, 10; USNM); Beau Vallon Beach, 20 Feb–4 Apr 1965, 1986, W.N. Mathis, I.B. Nye, W.T. Tams (9, 20; BMNH, USNM); Bel Ombre, 5 Apr 1986, W.N. Mathis (8, 2; USNM); Port Launay, 8 Apr 1986, W.N. Mathis (1, 2; USNM).

Australasian/Oceanian: AUSTRALIA. Queensland: Cairns, 19–25 Apr 1957, W.W. Wirth (3, 2; USNM); Cairns, Ellis Beach, 28 Apr 1957, W.W. Wirth (7, 2; USNM); Green Island (on decaying organism on beach), 17 Apr 1980, G.F. Hevel, J.A. Fortin (18, 6; USNM); Port Douglas (sand beach), 24 Apr 1957, W.W. Wirth (8, 12; USNM).

BONIN ISLANDS. Chichi Jima: Omura (camp beach), 2–25 Apr 1958, F.M. Snyder (3, 2; USNM); Takayama Bay, 22 Apr 1958, F.M. Snyder (1; USNM). Haha Jima: Okimura, 26 May–9 Jun 1958, F.M. Snyder (4; USNM).

COOK ISLANDS. Aitutaki Island, 27 Oct 1945, D.G. Hall (1; USNM).

FEDERATED STATES OF MICRONESIA. Kusie: Mutunlik (22 m), 1 Apr 1953, J.F.G. Clarke (2; USNM); Yeta Cave, 9 Apr 1953, J.F.G. Clarke (1; USNM). Pingalep Atoll, 22 Jul 1949, R.P. Owen (1; USNM). Ponape Island, Kolonia, Aug 1956, M.R. Wheeler (1; USNM). Yap Islands: Gagil, Gachapar, 19 Jun 1957, C.W. Sabrosky (3, 2; USNM); Weloy, 19 Jun 1957, C.W. Sabrosky (2; USNM).

GUAM. Point Oca, 6–14 Jun 1945, G.E. Bohart, J.L. Gressitt (1, 1; ANSP); Point Ritidian, 16 Apr–Jun 1936, 1945, E.H. Bryan, Jr., G.E. Bohart, J.L. Gressitt (4, 2; ANSP, USNM).

HAWAIIAN ISLANDS. Kaua'i: Anini Beach (about 4 mi [6.4 km] NE Hanalei), 7 Jul 1985, R.S. and V.L. Zack (6; WSU). Maui: Hana, 10 Jan 1973, J.R. Vockeroth (1; USNM); Kihei (rocky beach), 30 Jun 1985, R.S. and V.L. Zack (1, 2; WSU). Oahu: Bellows Field, 8 Jul 1952, A.L. Melander (8, 1; USNM); Hanauma Bay (crab hole), 28 Jun-3 Jul 1936, 1952, A.L. Melander (4, 2; ANSP, USNM); Kailua Beach, 26 Mar 1968, J.A. Tenorio (2, 5; USNM, WSU); Moanalua (salt lake), 7 Mar 1945, Y. Tahada (2 USNM); Mokuleia, 16 May 1952, A.L. Melander (1, 2; USNM); Waiamae (resting on crab holes on sandy beach), 8 Nov 1946, W.W. Wirth (1, 3; USNM); Waimanalo Beach (crab hole), 8 May-2 Oct 1936, 1946, 1952, 1966, A.L. Melander, J.R. Vockeroth, W.W. Wirth (9, 5 ANSP, USNM); Sep 9–10 1955, M.R. Wheeler (1; USNM). Johnston Island, 10 Jul 1958, H.I. Rainwater (1; USNM).

KIRIBATI. Gilbert Islands. Aronuka Atoll: 20 Nov 1964, B.D. Perkins (1 USNM). Butaritari Atoll: Butaritari Island, Jul 1957, N.L.H. Krauss (1, 1 USNM). Makin Atoll: B.D. Perkins (2, 1 USNM). Tarawa Atoll: Bairiki Island, Nov 1957, N.L.H. Krauss (1, 1 USNM); Banraeaba, Jul 1957, N.L.H. Krauss (1 USNM); 23–27 Jul 1945, D.G. Hall (2; USNM). Phoenix Islands: Kanton Island, 25 Apr 1940, D.B. Langford (1; BBM). Line Islands: Palmyra Atoll, Feb-Jul 1948, 1962, N.L.H. Krauss, M.R. Wheeler (1, 2; USNM).

MARSHALL ISLANDS. Enewetak Atoll: Enewetak Island, 13 May 1946, H. Townes (1, 3 USNM). Jaluit Atoll: Jabor Island, 27 Apr 1958, J.L. Gressitt (1; USNM). Japtan Island, Aug-Sep 1955, M.R. Wheeler (1 USNM). Kwajalein Atoll: 8 Aug 1944, D.G. Hall (2, 1 USNM). Kwajalein Atoll: Kwajalein Island, 17 Aug 1946 (1 USNM); Loi Island, 3 Sep 1944, H.S. Wallace (1; WSU). Likiep Atoll: Likiep Islet, 30 Aug 1946 (1; USNM). Majuro Atoll: Uliga Island, Aug-Sep 1955, M.R. Wheeler (1; USNM). Parry Island, Aug-Sep 1955, M.R. Wheeler (2, 1; USNM); 24 Dec 1950, Y. Oshiro (1; USNM).

MOLUCCAS. Bacan: Labuha (sweeping shoreline debris), 26 Sep 1985, A.H. Kirk-Spriggs (10, 14; NMWL).

NEW ZEALAND. North Island. Auckland: Cape Reinga Beach, 20 Jan 1966, J.M. Riches (1; NZAC); Clarks Beach (weeds on beach), 2 Feb 1953, K.A.J. Wise (1 NZAC); Green Bay (on Asteraceae), 6 Apr 1975, B.Z. Holloway (1; NZAC); Kare Kare (kelp on beach), 28 May-6 Apr 1975, B.A. Holloway, G.Kuschel (5, 3; 4 adults reared from puparia; NZAC); Ruapuke (beach W of Hamilton, mussels on rocks), 27 Feb 1971, J. Gillespie (1, 1 NZAC); Wattle Bay (plants on mud flats or on dead stingray), 23–24 Feb 1977, B.A. Holloway, G. Kuschel (1, 2 NZAC). South Island. Canterbury: Napenape Scenic Reserve (near mouth of Hurunumi River, freshwater pond near ocean), 17 Nov 1977, E. Schlinger (1 NZAC). North Chatham Island: Ocean Mail point (on flowers of Sonchus grandifolius Kirk), 20 Jan 1976, R.P. Macfarlane (1; NZAC); Wharekauri (on beach, 1on flower of Cotula coronopifolia L.), 16 Jan 1976, R.P. Macfarlane (1, 2 NZAC). Whale Bay Road, 9 Dec 1970, J. Gillespie (1; NZAC).

NORTHERN MARIANA ISLANDS. Pagan: Songsofig, 27 May 1940, Yasu. et Yosi. (1 USNM).

PALAU. Babelthuap Island: Melekeiok (beach), 23 May 1957, C.W. Sabrosky (5, 3 USNM); Ngerehelong, 6 May 1957, C.W. Sabrosky (5, 2 USNM); Ngiwal (beach), 9 May 1957, C.W. Sabrosky (1; USNM). Ngaiangl Atoll, 9 May 1957, C.W. Sabrosky (1; USNM). Nguruukdabel Island, Ngaremediu, 24 May 1957, C.W. Sabrosky (14, 3 USNM). Ulebsehel Island, 21 May 1957, C.W. Sabrosky (4, 2 USNM).

PAPUA NEW GUINEA. Admiralty Islands: Los Negros, 27 Mar 1945, D.G. Hall (21, 25; USNM). Bismark Archipelago: Hermit Island, 26 Jun 1962 (13, 8 USNM, UZMC); Lavongai, Banatam, 24 Mar 1962 (1; USNM). Manus Lorengau, 14 Jun 1962 (8; UZMC). Papua New Guinea. Central Province: Boerg (seashore), 29 Nov 1982, J.W. Ismay (1, 1 USNM); Gabagaba (W Kwikila), 1 Jan 1983, J.W. Ismay (8, 4; USNM); LeaLea (saltpan margin), 6 Oct 1985, J.W. Ismay (1; USNM); Loloata Island (seashore), 31 Mar 1985, J.W. Ismay (2, 1 USNM); Yule Island (rocky shore, Strandline), 1 Jan 1983, J.W. Ismay (2, 3 USNM). West New Britain Province: Dami (beach, near Kimbe), 29 Feb 1983, J.W. Ismay (6, 2 USNM). West Sepik Province: Wutung (seashore), 12 Nov 1985, J.W. Ismay (3, 3 USNM).

SOLOMON ISLANDS. Guadalcanal: Koli Point, Oct 1944, J. Laffoon (2, 3 USNM). Guadalcanal and Florida Islands, 1 Mar 1945, J.R. Stuntz (1, 1 USNM).

WAKE ISLAND. Peale Island, Nov 1957, N.L.H. Krauss (1, 1 USNM). Oriental: CHAGOS ARCHIPELAGO. Diego Garcia: Baracholis Section (on dead fish), 25 Apr 1971, A. Hutson (1, 2 BMNH); East Point, 7 May 1971, A. Hutson (10, 6 BMNH).

Oriental: CHINA. Hong Kong: Lantau Island, Trappist Monastary to Silver Mine Bay (over hills), 24 Jul 1964, W.J. Woss (1 BBM).

INDIA. Maharashtra: Bombay, Juhu Beach, 7 Mar 1985, K.A. Spencer (1 NMWL).

MALAYSIA. Penang: Georgetown (dirty sand ocean beach and wave swept rocks), 2–17 Jan 1990, J.R. Vockeroth (27, 11; CNC). Sabah: Kota Kinabalu (17 km S), 9 Aug 1983, G.F. Hevel, W.E. Steiner (5, 12 USNM).

PHILIPPINES. Balabac Dalawan Bay, 12 Oct 1961 (1 UZMC). Calicoan, 27 Jul 1945, F.F. Bibby (11, 1; USNM). Jul 1945, F.F. Bibby (12, 13 USNM).

RYUKYU ISLANDS (JAPAN). Okinawa: Chizuka, Jul-Sep, G.E. Bohart (13, 3 USNM); Okinawa (rotting fish on shore), 17–20 Aug-23 Sep 1945, 1980, P.S. Cranston, W.D. Field (7, 10; BMNH, USNM).

SRI LANKA. West Province: Ratmalana, Colombo (9 mi S), 7–13 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (5, 2 ZIL). Northwest Province: Kadaimparu, Negombo (15 mi [24 km] N), 31 Jan 1962, H. Andersson, P. Brinck, L. Cederholm (1; ZIL).

TAIWAN. Takao, 3 May 1907, H. Sauter (5, 1; ANSP, USNM).

THAILAND. Bangphra, Cholburi (light trap), Oct 1962, J.E. Scanlon (1; USNM).

Palearctic: JAPAN. Honshu: Osaka, Hamadera Koen, 29 Aug 1954, P.H. Arnaud, Jr. (56, 31; CAS, USNM, WSU). Okitsu, Jun 1913, F. Muir (1; BBM); Okitsu (in cage), 27 Oct 1914, F. Muir (1 BBM); Noma Tanegashima, Jun 1958, N.L.H. Krauss (1; BBM). Amami Group: Tokuno Shima, Hetono, 23 Jul 1963, C.M. Yoshimoto (20, 1, 1 abdomen missing; BBM).

DISTRIBUTION (Figure 33).—Old World along coasts of the Indian and Pacific oceans. Afrotropical: Kenya, Madagascar, Mozambique, islands in the Indian Ocean (Aldabra, Astove, Cosmoledo, Seychelles). Australasian/Oceanian: Australia (QLD), Belau, Bonin Islands, Cook Islands, Federated States of Micronesia, Hawaiian Islands (Maui, Oahu), Kiribati, Marshall Islands, Moluccas, New Zealand, Northern Mariana Islands, Papua New Guinea (Admiralty Islands, Bismark Archipelago, Papua New Guinea), Solomon Islands, Wake Island. Oriental: Chagos Archipelago (Diego Garcia), China, India, Indonesia (Irian Jaya), Malaysia, Philippines, Ryukyu Islands, Sri Lanka, Taiwan, Thailand. Palearctic: Japan (Honshu).

NATURAL HISTORY.—Bohart and Gressitt (1952) found adults of this species (as H. persimilis) to be abundant on beaches of the island of Guam. Specimens occurred on moist sand, low-growing beach vegetation, and dead fish. They also discovered large numbers of maggots in damp, foul-smelling sand beneath a human carcass. Only rarely did they find adults on vegetation in a coconut grove, about a mile from sea.

Tenorio (1980:276), working on Hawaii, successfully reared this species (as H. persimilis) from larvae and puparia that had been collected and maintained on a medium of moist seaweed. He also reported an abundance of adults on rotting fish and crabs on the sand.

DIAGNOSIS.—This species is distinguished from all others of the albicans group by the distinct row of short and slightly stout setae along the anteroventral margin of the fore femur.

NOTES ON VARIATION.—Some variation in the coloration of this species appears to have a geographic component. In the Pacific, especially the north Pacific (Japan), the dorsum of the thorax is light brown, distinctly contrasted with the gray pleural areas. In the Indian Ocean (Madagascar, the Seychelles, India), the dorsal and pleural areas are essentially concolorous, gray to silvery gray. Setae also seem to be more robust in the Japanese specimens, perhaps best seen in the comblike row of setae along the anteroventral surface of the fore femur. Otherwise, however, the specimens are very similar, and the structures of the male genitalia are virtually identical. Because the variation is gradual, apparently clinal, I have interpreted it to be intraspecific.