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Hecamede (Hecamede) albicans (Meigen)
Notiphila albicans Meigen, 1830:65,—Macquart, 1835:522 [review, list, France].
Hecamede albicans.—Haliday, 1839:224 [generic combination].—Walker, 1856:344 [list].—Rondani, 1856:131–132 [list, Italy].—Loew, 1860:13 [review, synonymy.—Schiner, 1863:245 [review].—Becker, 1896:121 [list]; 1905:189 [palearctic catalog]; 1926:18 [revision].—Jones, 1906:181 [list].—Ringdahl, 1921:70 [note]; 1941:17 [note].—Cresson, 1925:229 [lectotype designation, list]; 1942:107 [review]; 1946b:248 [review].—Wahlgren, 1927:329 [review].—Séguy, 1934:408 [review, list, France].—Duda, 1942:6 [note on distribution].—Caspers, 1951:156 [list, Bulgarian coast].—Ardö, 1957:132 [natural history, list, Sweden].—Frey, 1958a:53 [list, Canary Islands]; 1958b:38 [list, Cape Verde Islands].—Dahl, 1959:160 [review, list, Sweden].—Wirth, 1965:737 [nearctic catalog].—Papp, 1975:26 [review].—Cogan, 1976a:83 [list, England]; 1980:657 [Afrotropical catalog]; 1984:130 [palearctic catalog].—Norrbom, 1983:118 [natural history].—Canzoneri and Meneghini, 1983:231–235 [review, list, Italy].—Valdenberg, 1985:172–173 [natural history, immature stages].—Steinly, 1992:329–335 [natural history, immature stages].
Notiphila (Hecamede) albicans.—Walker, 1853:254 [review].
Psilopa (Clasiopa) globifera Boheman, 1853:204.—Walker, 1856:344 [synonymy with H. albicans].—Loew, 1860:13 [synonymy].
Notiphila globifera.—Zetterstedt, 1855:4760 [review]; 1860:6314–6315 [review].
Hecamede grisescens Becker, 1903:166; 1905:189 [palearctic catalog].—Cogan, 1984:131 [palearctic catalog]. [New synonym.]
DESCRIPTION.—Small to moderately small shore flies, length 1.60 to 2.70 mm.
Head (Figures 10–12): Fronto-orbits and ocellar triangle densely microtomentose, mostly tan but quite variable (gray to blackish gray); mesofrons, except for ocellar triangle, sparsely microtomentose, only anterior of mesofrons yellowish orange to slightly reddish orange; 1 proclinate fronto-orbital seta, inserted anterolaterad of reclinate seta. Antenna mostly yellowish orange to orange; flagellomere 1 usually with some brown coloration toward base, near insertion of arista; arista bearing 4 dorsal branches, rarely 3. Face whitish to silvery gray, bearing 3 large lateral setae and 1 or 2 smaller ones ventrad, each arising from subshiny, bare, shallow tubercle; facial prominence bare at apex, bare area usually large, subequal to width of pedicel. Gena bearing 1 large and several smaller black setae; gena high, eye-to-cheek ratio averaging 0.55. Clypeus black with light to dense investment of gray microtomentum.
Thorax: Scutum and disc of scutellum mostly grayish tan, frequently with some faint golden luster; lateral margins of scutum, beginning slightly above level of notopleuron or as far ventrad as dorsal portion of anepisternum, becoming grayer; prescutellar acrostichal setae and presutural seta usually well developed; microtomentum of pleural sclerites with faint shiny luster; anepisternum bearing several setulae on dorsal towards posterodorsal angle. Legs, except for tibiae and tarsi, mostly gray; fore femur mostly black anteriorly, mostly gray posteriorly; middle and hind femora mostly gray anteriorly, posterior surface blackish; fore femur lacking distinct row of setae along anteroventral surface; tibiae yellow, usually with thin investment of mostly white microtomentum; fore basitarsus with 2 or 3 black setae inserted toward base at posteroventral surface; tarsomeres mostly yellow except for apical 1–2, which are dark 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 13, 14) 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; epandrium (Figure 13) a dorsal band, gradually enlarged posteriorly; surstylus (Figure 13) in lateral view L-shaped, longer arm extended ventrally, long and narrow, parallel sided, posterior and anterior margins slightly curved, apex bluntly rounded, bearing 4 or 5 setulae; cercus (Figure 13) long and narrow, length almost twice that of epandrium, bearing conspicuously longer setae at posteroventral margin; gonite (Figure 14) roughly triangular in shape, with a narrow connection above aedeagus, ventral angle as a narrow, short, parallel-sided, blunt process; aedeagal apodeme (Figure 14) L-shaped, with arms curved, better developed toward angle; aedeagus (Figure 14) 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 lectotype male of Notiphila albicans (designated by Cresson, 1925:229) is labeled “TYPE [red]/Marseille [France; handwritten]/albicans [handwritten] Coll. Winth. [black border along both longer sides of label].” The specimen is double mounted (pin in a rectangular block of cork), is in good condition, and is deposited in the NMW. There are also five paralectotypes in the NMW.
The lectotype male of Psilopa globifera, designated herein, is labeled “[Sweden] Sc. [Skåne, handwritten]/Bhn [Boheman, handwritten]/Notiph[ila]. globifera Boh.Scan. a Bohem. 52. [handwritten]/LECTOTYPE Psilopa globifera Boheman By W.N.Mathis [handwritten except for “LECTOTYPE” and “By”, a black submarginal border].” The lectotype is directly pinned, is in fair condition (considerable verdigris present), and is deposited in ZIL (Göteborg collection). There are also four paralectotypes (2, 2; ZIL) that bear the same first two labels as the lectotype.
The holotype male of Hecamede grisescens Becker is labeled “Alexandrien 44158 XI [handwritten]/Hecamede n.sp. [handwritten]/Holotypus [red]/Zool. Mus. Berlin.” The holotype is double mounted (minuten in a rectangular block of pith ?), is in fair condition (the left wing is missing, the right wing is twisted), and is deposited in the ZMHU.
OTHER SPECIMENS EXAMINED.—OLD WORLD. Palearctic: BULGARIA. Achtopol (seashore, 42°09′N, 27°52′E), 7–10 May–Aug 1941, 1958, 1987, M. Barták, H. Caspers, B. Pisarski (6, 8; BMNH, MBP, USNM); Aladza bei Varna, 16 Jul 1930, H. Zerny (12; NMW); Irakli (damp meadow, 42°47′N, 27°54′E), 16 Jul 1987, M. Barták (24, 10; MBP). Primorsko ad Burgas, 7 Jun 1959, R. Bielwawski (1; USNM).
CROATIA. Dalmatia, G. Strobl (1; UZMC); Korcula, Lombarda, 19 Apr 1930, H. Zerny (1; NMW); Split (on shore), 2 Sep 1986, F. Kaplan (3; USNM).
DENMARK. Hellebaek, (1; UZMC). Hornbaek, 31 May 1887 (2, 2; UZMC). Roenne, Jul 1883, H.J. Hansen, W. Lundbeck (4, 19; UZMC, WSU).
EGYPT. Fayoum, Lake Karun, 2–23 Nov 1945, R.L. Coe (25, 6; BMNH).
FRANCE. Marseille, Winth (1; ANSP).
GREAT BRITAIN. Cornwall: Lelant Towans, 27 May 1935 (4, 3; BMNH); Mayle, 22 Jul 1934, L. Parmenter (1, 1; BMNH); Padstow, 16 Mar–3 ct 1905, C.G. Lamb (2, 3; BMNH, USNM). Devon: Dartmoor, Wistman's Wood, 5 Oct 1959, A.C. Pont (1; BMNH). Dorset: Studland, 13 May–26 Oct 1912, 1961, Verbury (3, 2; BMNH). Kent: Deal, 3–18 Jul 1905, 1948, J.J.F.X. King (4, 1; BMNH); Sandwich Bay, 23 Jul 1957, E.A. Fonseca (9, 6; ZIL).
GREECE. Crete: Chania. 11 Jul 1981, A. Freidberg (5, 1; USNM); Längs vägen Malia-Hersonissos, 19 May 1979, R. Danielsson (1, 3; ZIL); Paleochora, 9 Jul 1981, A. Freidberg (4, 11; USNM).
ISRAEL. Akhziv, 19 Aug 1982, A. Aadka (1; TAU); Akko, 16 May 1980, W.N. Mathis (3, 2; USNM); Haifa, 18 May–25 Nov 1971, 1980, J. Kugler, W.N. Mathis (6, 2; TAU, USNM); Herzliyya, 26 May–19 Nov 1980, 1981, A. Freidberg (1, 2; TAU); Kefar Rosh HaNiqra, 2 Jun 1981, A. Freidberg, W.N. Mathis (2, 4; TAU, USNM); Ma'agan Michael (beach), 17 May–22 Aug 1980, 1984, F. Kaplan, W.N. Mathis (24, 39; TAU, USNM); Michmoret, 3 Aug 1983, A. Freidberg (1; TAU); Nahariyya (beach), 8 Aug 1986, W.N. Mathis (11, 1; USNM); Taba, 27 Apr 1974, A. Freidberg (1, 1; TAU); Tel Aviv (dunes), 8 Apr 1981, A. Freidberg (1; TAU); Zaharani spill, 19 Jun 1982, I. Yarom (1; TAU); Ziqim, 29 Feb 1984, A. Freidberg (1, 1; TAU).
ITALY. Friuli-Venezia Giulia: Trieste, 24 Jul 1886, A. Handlirsch (2, 2; ANSP). Marches: Porto Civitanova, 10 Aug 1897, M. Bezzi (2, 2; USNM): San Benedetto (10 km N, damp valley, 42°58′N, 13°53′E), 3 Aug 1988, M. Barták (1; MBP). Sicily: Capo San Vito, 29 Aug 1982, A. Freidberg (3, 4; USNM); Schioedte (2; UZMC). Veneto: Venezia (seashore, 45°27′N, 12°18′E), 7 Aug 1988, M. Barták (2, 1; MBP).
LIBYA. Tripoli, Wheelers Field, 26 Apr 1951, R.W. Brown (1, 2; USNM).
PORTUGAL. Estrema Dura, So Martinho do Porto, 16 Sep 1964, O.W. Richards (1; BMNH); Lisbon (on dead fish), 21 Jul 1962, L. Horacsek (1; BMNH); Porto, 5 Aug 1962, J. Abraham, L. Horacsek (2, 1; BMNH); Porto, Leca da Palmeira (low vegetation near seashore), 7 Aug 1962, J.A. and L.H. (19, 8; BMNH); Porto, Boa Nova, Jun 1960, N.L.H. Krauss (1; USNM). Azores: So Miguel, So Populo, Ponta Delgada (7.5 km E), 12 Mar 1957, P. Brinck, R. Dahl (2, 2; ZIL).
ROMANIA. Lake Sinoe (edge, low salinity maritime flats near Histria), 18–20 Jun 1969, B.H. and M.C. Cogan, R. Vane-Wright (3, 3; BMNH); Olimp (near Constanta) 20 Aug 1975, A.N. Clements (5; BMNH).
SPAIN. Provence. Fréjus (Var), 9 Jun 1924, H. Zerny (, ; NMW). Andalusia, Algeciras, 12–20 May 1925, H. Zerny (3, 3; NMW). Almeria, Albufera, 23 Mar 1966, L. Lyneborg (2, 2 [sex ?]; UZMC). Granada, Torrehueva (E Motril), 14 Apr 1966, L. Lyneborg (1; UZMC). CANARY ISLANDS. Lanzarote, El Golfo, 4 Sep 1976, M. Baez (1; USNM).
SWEDEN. Halland: Laxvik, Jul 1985, P. Ardö (2; ZIL); Laxvik, Trönninge, Jun–Jul 1954, 1955, P. Ardö, R. Dahl (6, 7; ZIL); Eldsberga, Tönnersa, 1 Jul 1954, R. Dahl (1; ZIL). Skåne: Ängelholm, 5 Sep 1915 (1; ZIL); Bjärred (strand), 4 Oct 1973, H. Andersson (2; ZIL); Hallands Väderö, 18 Jul 1924 (1; ZIL); Kullaberg, 24 Aug 1983, H. Andersson (1; ZIL); Lerberget (strand), 13 Aug 1969, H. Andersson (4, 1; ZIL); Lomma, 11 Aug 1974, H. Andersson (8, 2; ZIL); Mölle, 24 Jun–8 Aug 1926, 1969, H. Andersson (12, 9; ZIL); Strandbaden, Tallbacken, 4 Aug 1975, H. Andersson (1; ZIL); Torekov, Jul 1860, C.D.E. Roth (5; ZIL).
TURKEY. Istanbul, 30 Aug 1965, B.H. Cogan (1; BMNH); Silivri, 3 Nov 1965, B.H. Cogan (2, 2; BMNH).
NEW WORLD. Nearctic: BERMUDA. Devonshire Parish: Devonshire Bay (32°18′N, 64°44′W), 29 May 1991, W.N. Mathis (16, 4; USNM). Hamilton Parish: Shelly Bay Beach (32°19′N, 64° 44′W), 31 May 1991, W.N. Mathis (2, 5; USNM). Paget Parish: Hungry Bay (beach, 32°17′N, 64°45′W), 2 Jun 1991, W.N. Mathis (3, 2; USNM). Pembroke Parish: Spanish Point (beach, 32°18′N, 64°49′W), 2 Jun 1991, W.N. Mathis (1, 1; USNM). Warwick Parish: Long Bay Beach (32°15′N, 64°48′W), 30 May 1991, W.N. Mathis (2, 1; USNM); Warwick Long Bay, 18 Nov 1987, D.J. Hilburn, N.E. Woodley (4, 3; USNM).
UNITED STATES. Connecticut: Fairfield Co., Westport, 14 Aug 1939, A.L. Melander (1, 5; USNM). New Haven Co., Anchor Beach, Milford-West Haven, 5–15 Aug 1978, B.A. Steinley (about 1000 and; USNM); Branford, 30 Jun 1952, A.H. Sturtevant (1, 2; USNM). Delaware: Sussex Co., Rehoboth, 25 Jun–4 Aug 1939, 1941, A.L. Melander (4, 4; ANSP, USNM). Maine: Cumberland Co., Chebeague Island, 3 Sep 1952, A.H. Sturtevant (5; USNM). Maryland: Calvert Co., Lusby, G.C. Steyskal (1; USNM); Port Republic, 26 Apr 1971, R.J. Gagné (2, 5; USNM). Massachusetts: Barnstable Co., Falmouth, 11 Aug 1979, A. Freidberg (2; USNM); Cape Cod (end), 6 Jun–3 Aug 1939, 1970, A.L. Melander, H. Weems (5, 16; USNM); Woods Hole, 28 Jun–24 Aug 1950, 1952, A.H. Sturtevant (4, 4; USNM). Dukes Co. Pasque Island, 15 Aug 1952, A.H. Sturtevant (1, 3; USNM); Vineyard Haven, 17 Aug 1954, A.H. Sturtevant (2; USNM). North Beach (reared from seaweed), 3 Sep 1955, M. Williams (1, 2; USNM). New Hampshire: Rockingham Co., Rye Beach, 31 Jul 1950, A.H. Sturtevant (1, 2; USNM). New Jersey: Cape May Co., Town Bank (ex. flowering Prunus serotina), 31 May 1983, A.L. Norrbom (1, 2; USNM). New York: Queens Co., New York City, Douglaston, 2 Sep 1954, A.H. Sturtevant (1, 1; USNM). Suffolk Co., East Quoque, 10 Aug 1957, L. Wilcox (1; USNM); Montauk, 23 Jul 1932, A.L. Melander (5, 4; ANSP, USNM); Orient, 30 Apr 1953, R. Latham (1; USNM); Riverhead, Baiting Hollow (north shore), 9 Sep 1961, P.H. Arnaud, Jr. (10, 12; AMNH). Rhode Island: Newport Co., Tiverton, 13 Aug 1952, A.H. Sturtevant (2; USNM). Virginia: Accomack Co., Assateague Island, 19 Aug 1969, G.C. Steyskal (1; USNM). Westmoreland Co., Westmoreland State Park, 9 Jun 1988, D. and W.N. Mathis (3, 3; USNM). Norfolk, 14 Aug 1969, G.C. Steyskal (7, 4; USNM).
DISTRIBUTION (Figure 15).—Holarctic along coasts of the northern Atlantic Ocean, including the Mediterranean Sea. Palearctic: Azores, Bulgaria, Canary Islands, Croatia, Denmark, mark, Egypt, France (type locality of senior synonym), Great Britain, Greece, Ireland, Israel, Italy, Libya, Portugual, Romania, Spain, Sweden, Turkey, (also reported from Cape Verde Island, Madeira). Nearctic: Bermuda, United States (CT, DE, MA, ME, MD, NH, NJ, NY, RI, VA).
NATURAL HISTORY.—In Ireland, Haliday (1839:224) first noted that this species occurs “On sandy coasts, especially on fresh marine rejectamenta; runs fast and does not fly readily.” Ardö (1957:184) found this species wholly in the “marine shore dune ecosystem” where it was baited to and trapped on decaying herring. Ardö reported two generations at higher latitudes, such as in Sweden, but on beaches of the eastern Mediterranean (Israel), adults are fairly common throughout the year (February to November), suggesting several generations per year. Dahl (1959:160), from studies also done in Sweden, suggested that this species prefers halophilous habitats but also has xerophilous tendencies and a preference for habitats that have soil substrates with microflora.
In addition to summarizing past work on the natural history of this species, Norrbom (1983:118) succeeded in rearing over 200 adults from larvae that were feeding on dead horseshoe crabs that had washed ashore in Delaware Bay near Town Bank, New Jersey (USA). He observed larvae mostly between the gills and on other moist membranous surfaces but found that pupariation occurred in the sand and on the sides of the rearing jar as well as within the crab. Norrbom also described and illustrated the third-instar larva and the puparium and noted that one puparium was parasitized by the pteromalid, Urolepsis rufipes (Ashmead).
Steinly (1992) recently confirmed Norrbom's work and added many details about the life cycle, especially how quality and quantity of nutrient resources affect size, from both field and in vitro rearings. His field work was conducted in Connecticut, where the species is multivoltine and can complete a life cycle in less than two weeks.
DIAGNOSIS.—This species is distinguished from similar congeners by the following combination of characters: proclinate fronto-orbital setae 1; gena with several black setae; fore femur lacking a distinct row of setae along anteroventral surface; tibiae mostly yellow, investment of mostly white microtomentum thin; fore basitarsus with 2 or 3 black setae inserted toward base of posteroventral surface; and structures of the male terminalia (see description and Figures 13, 14), especially the shape of the surstylus and gonite.
- bibliographic citation
- Mathis, Wayne Neilsen. 1993. "Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A Revision of the Shore-Fly Genus Hecamede Haliday." Smithsonian Contributions to Zoology. 1-46. https://doi.org/10.5479/si.00810282.541