Phylloicus aeneus (Hagen)

PRATHER, AYSHA L., 2003, Revision of the Neotropical caddisfly genus Phylloicus (Trichoptera: Calamoceratidae), Zootaxa 275 (1), pp. 1-214 : 21-26

Phylloicus aeneus (Hagen)

Figs. 1 , 6 , 15 , 16 , 118

Phylloicus aeneus ( Hagen, 1861:285) [Type locality: Mexico [Veracruz]; MCZ; female; in Macronema ]; 1864:804 [to Anisocentropus ]. — Ulmer 1905a:79 [to Phylloicus , redescription of male now in P. nigripennis ]. Ross 1952:34 [male]. — Flint 1967:17 [illustration of male now in P. nigripennis ]. — Denning et al. 1983:182 [redescription, illustration is probably of P. mexicanus ]. —Bowles & Flint 1997:58 [variation].

Phylloicus ornatus ( Banks, 1909:342) [Type locality: United States, Texas, Brownwood; MCZ; female; in Notiomyia ]. — Flint 1967:17 [to Phylloicus ]. — Holzenthal 1988:72 [distribution]. — Bowles & Flint 1997:53 [redescription, male, female, larva, pupa]. NEW SYNONYM.

Phylloicus centralus ( Navás, 1924:82) [Type locality: Costa Rica; MNHNP; male; in Macronema ]. — Holzenthal 1988:72 [to Phylloicus ]. NEW SYNONYM.

The identity of P. aeneus has been the subject of some confusion, largely due to the holotype being a female in very poor condition. Several authors have redescribed this species based on specimens mistakenly associated with the type female, and thus judged P. nigripennis , mexicanus , latus , and sagittosa to be conspecific with P. aeneus . Based on the evidence discussed below, mexicanus and nigripennis are valid species — with latus and sagittosa junior synonyms of the latter — and ornatus and centralus are synonyms of aeneus .

Hagen (1861) described Macronema aeneum from a female specimen labeled “Sallé, V.C.” [for Veracruz, Mexico, handwritten on a single green label]. Ross (1952) identified this type specimen in a paper on lectotypes of Hagen species in the Museum of Comparative Zoology. He did not refer to the specimen as a lectotype in the paper. A yellow label in Ross’s hand on the specimen names it lectotype, but there is nowhere in Hagen’s original description or Ross’s 1952 paper (except by implication in the paper’s title) any indication that it was selected from among the type series. This specimen is badly damaged; the abdomen, which has been cleared and placed in a microvial of glycerin, is in many pieces; only a portion of one forewing remains, and even this is faded, so that it is not possible to determine what color pattern it might once have exhibited, although the epithet “ aeneum ” translates as “coppery.” The holotype female of M. aeneum bears a tuft of dark stout setae at each posterolateral margin of tergite VIII.

The female holotype of Notiomyia ornata was collected in central Texas. It is in good condition and the wing pattern is clearly discernable. The pattern is very distinctive, and the species can be easily discriminated on the basis of wing pattern alone. The wings can be described as copper­colored. The tufts of setae on tergite VIII observed in P. aeneus are present also on the female holotype of N. ornata . I have seen these setal tufts only on females of P. aeneus , ornatus , mexicanus , and panamensis , new species. Specimens of P. mexicanus have other distinguishing characters that rule out conspecificity; of the remaining two species, only P. ornatus is known to be from Mexico. The terminalia of the type specimens of P. aeneus and ornatus are indistinguishable. Only one form of Phylloicus is found in Texas, and thus there is no doubt of the association of male and female. The males bear short coremata on abdominal segment IV, with very short, broad projections of the posterolateral tergal margin.

The male holotype of M. centrale has terminalia and modifications of abdominal segment IV identical with those observed in P. ornatus males. The wing pattern is faded, but what remains is consistent with the distinctive pattern of P. ornatus .

Ulmer redescribed P. aeneus based on several specimens from Morelos; these specimens appear to be conspecific with the type of P. nigripennis . Flint (1967) compared the female types of aeneus and nigripennis and concluded that they were conspecific; he illustrated a male labeled “ Mexico, Sallé,” stating that it was probably part of the original type series. I have examined this and another specimen labeled “ Mexico, Sallé,” and labeled by Flint in 1965, as Phylloicus aeneus male. The collection labels on these specimens are printed (possibly cut from a journal article) with “ Mexico ” on the upper label, and “Sallé” on the second label; these labels do not resemble those on the type of aeneus except in the collector’s name. These specimens are males of P. nigripennis ; the wings are dark and without pattern, no coremata are present on the abdomen, and they exhibit other characteristics consistent only with that species.

The redescription of P. aeneus by Denning et al. (1983) includes a mention of the abdominal coremata. Their illustrations were prepared from specimens collected in Sonora, which may have been either P. aeneus or mexicanus ; if the specimens depicted in the illustration had abdominal coremata, they must have been P. aeneus .

Given this evidence, the logical conclusion is that P. aeneus , centralus , and ornatus are conspecific.

Pinned specimens of P. aeneus are easily identified by the distinctive pattern of dark patches near the base of the forewing. As well, in P. aeneus and mexicanus , nigripennis , and panamensis , there is an oblique fold of the forewing’s apical third, toward the midline ( Fig. 118 ). The male genitalia of P. aeneus are most similar to those of P. mexicanus , nigripennis , and panamensis . Phylloicus aeneus may be distinguished from these three species by the presence of short, single­lobed coremata, covered by a short posterior tergal process, on abdominal segment IV.

Adult. Forewing length 9.6­12.3 mm, n = 201.

Head orange­brown. Maxillary palps black. Antenna twice forewing length; dark brown, with narrow patches of pale sensilla on anteromesal surface of each flagellomere. Dorsal pterothorax orange­brown, lateral margins black; ventrolateral thorax golden brown. Femora golden brown; foretibiae pale tan; mesotibiae white proximally, dark distally; metatibiae white proximally, dark distally; tarsi black. Metathoracic leg of male without posterior fringe. Tibial spur formula 2,4,3. Forewing apical third folded obliquely toward midline; chestnut brown; with two transverse bands; proximal band golden, not reaching either wing margin, 1/2 width of wing or less; distal band golden, extending from anterior to posterior wing margins, zig­zagging, with golden oval surrounding nygma; with single basal stripe, golden; with large dark patch anterobasally and golden semicircle posterobasally ( Figs. 105 , 118 ). Hind wing basal brush present in female, orange.

Male. Preterminalic abdominal terga with anteromesal notch. Corematic structures present. Tergum IV with expanded lateral flanges, lateral coremata; lateral coremata with basal globose lobes. Tergum V without sclerotized modifications ( Fig. 15F ). Sternum VII with short, acute anteromesal process. Sternum VIII similar to anterior sterna, sternum IX not elongate. Tergum IX with mesal ridge extending 2/3 length of segment; posterior margin smoothly rounded; posteromesal surface covered with fine pilosity ( Fig. 15B ); lateral ridge absent; dorsal pleural setae approximately 15, ventral pleural setae approximately 5 ( Fig. 15A ); sternum IX with paired mesolateral ridges; sternum IX ( Fig. 15C ). Preanal appendage approximately length of tergum X, of uniform diameter throughout length, setae long, but not filamentous or longer than appendage ( Fig. 15A, B ). Tergum X sagittate basally; basodorsal process short and bifid; basolateral processes absent; apex, in lateral view, acute, in dorsal view, notched, notch square; with 2 longitudinal rows of short setae mesally; with paired shallow processes on lateral surfaces ( Fig. 15A, B ). Harpago long, curving mesally and tapering apically; peglike setae few, apical ( Fig. 15A, C ). Phallic endotheca with paired basolateral lobes, basolateral lobes large and round; phallotremal sclerites average size, longest dimension less than diameter of phallobase; dorsal sclerite narrow, width less than height ( Fig. 15D, E ).

Female. Abdominal terga I­III black. Preterminal abdominal terga with anteromesal notch. Sternum VII with short pointed anteromesal process. Tergum VIII with dense posterolateral brush of setae ( Fig. 16B ); sternum VIII cleft posteromesally, cleft not reaching anterior ridge; sternum VIII ( Fig. 16C ). Tergum IX with mesal ridge extending length of segment ( Fig. 16B ). Sternum IX anterior and posterior lobes darkly sclerotized and striate, with deep sublateral invaginations ( Fig. 16A ). Tergum X appendage longer than mesal lobe, base indistinct, apex triangular; mesal lobe lightly sclerotized; digitate lateral processes absent ( Fig. 16B ). Sternum X without setae in membrane ( Fig. 16A ). Vaginal apparatus anterior and posterior sclerites equal in length; anterior sclerite rounded anteriorly, posterolateral projections rounded; posterior sclerite ovoid; posterior end of spermatheca a sclerotized ovoid ( Fig. 16A ).

Material examined. BELIZE: Caya: Rió Frio, mouth of Rió Frio Cave , San Ignacio , 18.vi.1975, Resh — 2 males, 1 female ( UMSP) ; COSTA RICA: 1920, Serre — P. centralus holotype male ( MNHNP) ; Alajuela: 20 km S Upala , 1­20.xi.1990, Parker — 1 female ( EMUS) ; Río Toro, 3.0 km (road) SW Bajos del Toro , 10°12'14”N, 84°18'58"W, 1530 m, 3­4.ix.1990, Holzenthal, Blahnik, & Huisman — 1 male, 2 females ( NMNH) ; Reserva Forestal San Ramón, Río San Lorencito & tribs., 10°12'58”N, 84°36'25"W, 980 m, 1­4.x.1986, I & A Chacón — 1 male ( UMSP) ; 30.iii.­1.iv.1987, Holzenthal, Hamilton, & Heyn — 1 male ( UMSP) ; 1­4.v.1990, Holzenthal & Blahnik — 1 female ( UMSP) ; 28­ 30.vii.1990, Holzenthal, Blahnik, & Muñoz — 1 female ( UMSP) ; 6­10.ii.1991, Holzenthal, Muñoz, & Huisman — 2 males, 1 female ( UMSP) ; Cartago: Reserva Tapantí, Quebrada Palmitos & falls, ca. 9 km (road) NW tunnel, 09°43'12”N, 83°46'48"W, 1400 m, 24­25.iii.1991, Holzenthal, Muñoz, & Huisman — 1 male ( UMSP) ; Río Grande de Orosí , 09°41'10”N, 83°45'22"W, 1650 m, 23.i.1985, Janzen & Hallwachs — 1 male ( NMNH) ; Guanacaste: Cerro El Hache, W side, 310 m, 31.v.1986, Janzen & Hallwachs — 1 male, 1 female ( UMSP) ; Finca Montezuma , 3 km SE R. Naranjo, 17.xi.1991, Parker — 1 female ( EMUS) ; 20­31.i.1992, Parker — 1 female ( EMUS) ; v.1992, Parker — 1 female ( EMUS) ; 16­31.v.1993, Parker — 1 female ( EMUS) ; Maritza, confluence with Río Tempisquito , 10°57'25”N, 85°29'42"W, 580 m, 31­19.v.1994 — 1 male & pupal exuviae, 1 female ( UMSP) ; 14.ii.­1.v.1994 — 2 males & pupal exuviae, 2 females ( UMSP) ; Maritza, Rio Orosi , 10°59'28”N, 85°25'42"W, 675 m, 31.i.­27.v.1994 — 2 males & pupal exuviae, 1 female ( UMSP) ; Quebrada Garcia, Quebrada Garcia, 10.6 km ENE Quebrada Grande , 10°51'43”N, 85°25'41"W, 470 m, 8.iii.1986, Holzenthal & Fasth — 1 female ( UMSP) ; Río Los Ahogados, Río Los Ahogados, 11.3 km ENE Quebrada Grande , 10°51'54”N, 85°25'23"W, 470 m, 7.iii.1986, Holzenthal & Fasth — 1 male ( UMSP) ; Parque Nacional Guanacaste, Estacion Maritza, Río Tempisquito , 10°57'29”N, 85°29'49"E, 550 m, 29.i.1992, Sweeney — 1 male ( NMNH) ; Río Orosí, Estación Pitilla , 10°59'28”N, 85°25'41"W, 700 m, 22­25.v.1990, Holzenthal & Blahnik — 1 female ( UMSP) ; Río San Josecito , 10°55'19”N, 85°28'12"W, 960 m, 3­4.iv.1987, Holzenthal, Hamilton, & Heyn — 1 male ( INBIO) ; Río Tempisquito, Maritza , 10°57'29”N, 85°29'49"W, 550 m, 19­ 20.vii.1987, Holzenthal, Morse, & Clausen — 1 female ( UMSP) ; Parque Nacional Rincón de la Vieja, Río Negro , 10°45'54”N, 85°18'47"W, 810 m, 3.iii.1986, Holzenthal & Fasth — 1 male, 3 females ( UMSP) ; Límon: Limon , 16 km W Guapiles, 400 m, ii.­iii.1989, Hanson — 1 male ( UMSP) ; Reserva Biológica Barbilla, trib. to Río Dantas, 13 (km) S Pacuarito , 09°59'42”N, 83°28'37"W, 0 m, 1.ii.1992, Holzenthal, Muñoz, & Kjer — 1 male, 1 female ( UMSP) ; Puntarenas: Río Bellavista trib., Las Alturas , road to quarry, 08°57'07”N, 82°50'53"W, 1480 m, 19.iii.1991, Holzenthal, Muñoz, & Huisman — 1 female ( UMSP) ; Río Bellavista, ca. 1.5 km NW Las Alturas , 08°57'04”N, 82°50'46"W, 1400 m, 15­17.vi.1986, Holzenthal, Heyn, & Armitage — 1 male ( UMSP) ; 8­9.iv.1987, Holzenthal, Hamilton, & Heyn — 2 males, 2 females ( INBIO) ; — 9 males, 9 females ( UMSP) ; 2­3.viii.1987, Holzenthal, Morse, & Clausen — 2 males, 1 female ( UMSP) ; 10­ 11.viii.1990, Holzenthal, Blahnik, & Muñoz — 1 male, 1 female ( UMSP) ; 16­17.iii.1991, Holzenthal, Muñoz, & Huisman — 7 males ( UMSP) ; Río Cotón, in Las Alturas , 08°56'17”N, 82°49'34"W, 1360 m, 12.viii.1990, Holzenthal, Blahnik, & Muñoz — 2 females ( UMSP) ; 18.iii.1991, Holzenthal, Muñoz, & Huisman — 1 male ( INBIO) ; Río Guineal, ca 1 km (air) E Finca Helechales , 09°04'34”N, 83°05'31"W, 840 m, 22.ii.1986, Holzenthal, Morse, & Fasth — 4 males, 2 females ( UMSP) ; 4.viii.1987, Holzenthal, Morse, & Clausen — 1 male ( UMSP) ; Río Singrí, ca 2 km (air) S Finca Helechales , 09°03'25”N, 83°04'55"W, 720 m, 21.ii.1986, Holzenthal, Morse, & Fasth — 1 male, 1 female ( UMSP) ; San Vito, 2 km S., Finca Las Cruces , 08°42'00”N, 83°00'00"W, 4000 m, 20.xi.1988, Solis — 1 male ( INBIO) ; Jardín Botanico R & C Wilson, unnamed trib., Sendro del Agua , 08°48'00”N, 82°57'36"W, 1180 m, 8.viii.1990, Holzenthal, Blahnik, & Muñoz — 1 female ( UMSP) ; Zona Protectora Las Tablas, Río Cotón, Sitio Cotón , 08°56'28”N, 82°47'13"W, 1460 m, 15.iv.1989, Holzenthal & Blahnik — 11 males, 10 females ( UMSP) ; San José: Motel Prado, San Isidro del General — 1 male ( UMSP) ; Pedregoso , 640 m, 21.ii., Rounds — 1 male ( MCZ) ; Río Parrita Chiquito , rt. 12, 6.5 km SW jct. rt. 2, 09°42'11”N, 83°58'12"W, 1990 m, 10.iv.1987, Holzenthal, Hamilton, & Heyn — 1 male ( INBIO) ; GUATEMALA: Las Mercedes , 914 m, 1879­1881, Champion — 1 male, 1 female ( BMNH) ; Panima, Vera Paz , 1879­1881, Champion — 1 female ( BMNH) ; Chiquimula: Padre Miguel , 19.viii.1965, Flint & Ortiz — 2 females ( NMNH) ; El Progreso: Est. de la Virgen , 11­12.viii.1965, Flint & Ortiz — 1 male, 2 females ( NMNH) ; Jutiapa: San Jerónimo , 1879­1881, Champion — 2 males, 3 females ( BMNH) ; Suchitepequez: Cuyotenango , 10­20.vi.1966, Flint & Ortiz — 1 male ( NMNH) ; MEXICO: 3048 m, 1871, Bilimek — 1 male ( NMW) ; Chiapas: 2.9 km S of Jitotol , 11.viii.1967, Flint — 2 males ( NMNH) ; 58 km S Palenque , 17.v.1981, C & O Flint — 1 male ( NMNH) ; Dolores, Rt. 190, km 1190, 7.viii.1965, Flint & Ortiz — 1 male ( NMNH) ; Soconusco, 27 Jul , Purpus — 1 female ( ZMHU) ; Chihuahua: Arroyo Toro, Toro Basin , 1720 m, 23.vi.1987, Kondratieff & Baumann — 1 male ( NMNH) ; Riito, Hwy 16, 10 mi E of Yepachic, 28.vi.1987, Kondratieff & Baumann — 1 male ( NMNH) ; Durango: 10 mi. W El Salto , 10.vi.1964, McAlpine — 1 female ( NMNH) ; Jalisco: 11.2 mi. E Cuititian, 4 mi. E Durazno , 1128 m, 29­ 30.viii.1987, Bloomfield — 1 female ( NMNH) ; 2.3 mi. E Durazno, 1158 m, 6­8.vi.1989, Bloomfield — 1 male ( NMNH) ; Mórelos: Cuernavaca , Banks — 1 female ( MCZ) ; Nayarit: 0.6 mi. E Riitos, road to San Andreas , 9­11.iii.1987, Bloomfield — 1 male ( NMNH) ; Nuevo Leon: Municipio de Santiago, Stream at La Nogalera above Cola de Caballo , 27.v.1991, Contreras & Harris — 1 male, 2 females ( UMSP) ; Oaxaca: 8 km S Valle Nacional , 25.v.1981, C & O Flint — 1 male, 1 female ( NMNH) ; San Luis Potosí: Huichihuayan, Rt. 85, km 399, 7.viii.1966, Flint — 1 male, 1 female ( NMNH) ; Sancti Spíritus: Sótano de Tlamaya , 26.xi.1964, Raines & Bell — 2 males ( NMNH) ; Sinaloa: Portrerillos , 15 mi. W El Palmito, 1524 m, 8.vii.1964, McAlpine — 1 female ( CNC) ; 20.vii.1964, Mason — 1 female ( CNC) ; Sonora: small stream near Yecora , 20.viii.1986, Kondratieff — 1 female ( NMNH) ; Tamaulipas: Rio verge. 25 km SW Ciudad Victoria, 14­16.iii.1982, Gillaspy — 1 male ( NMNH) ; Veracruz: Hagen — P. aeneus holotype female ( MCZ) ; 1.6 mi. N Coscomatepec , 22.vii.1966, Flint & Ortiz — 1 male, 1 female ( NMNH) ; Cordoba , 4­23.viii. Aug 1965, Lau — 1 female ( NMNH) ; 10­20.viii.1966, Lau — 1 male ( NMNH) ; 1.ix.1966, Lau — 1 male ( NMNH) ; nr. Huatusco , 25­26.vii.1965, Flint & Ortiz — 1 male ( NMNH) ; Río Tacolapan , 25­ 26.vii.1966, Flint & Ortiz — 1 male, 1 female ( NMNH) ; Las Tuxtlas Biological Station, Las Tuxtlas area, nr. Balzapote , 31 km NE of Catemaco, 3­15.v.1981, C & O Flint — 1 male, 3 females ( NMNH) ; NICARAGUA: Estelí: Area Protegida Miraflor, Quebrada Grande , 13°13'20”N, 86°15'33"W, 1200 m, 19.vii.2000, Chamorro & Lacayo — 1 female ( UMSP) ; Jinotega: Peñas Blancas , 13°17'00”N, 85°33'00"W, 1300 m, 25.vii.1997, Maes & Hernández — 5 males ( NMNH) ; PANAMA: Chiriqui: Boquete , 6.vi.1940, Fairchild — 1 female ( MCZ) ; Bugaba , 244­457 m, 21.x.1881 ­ 11.iii.1883, Champion — 1 male ( BMNH) ; Fortuna Dam Site nr. Hornitos , 08°55'00”N, 82°16'00"W, 1050 m, 24.xi.­ 20.xii.1976, Wolda — 6 males ( NMNH) ; 12­18.x.1977, Wolda — 1 male, 1 female ( UMSP) ; 9­6.xii.1977, Wolda — 1 male, 2 females ( NMNH) ; 16­29.xi.1977, Wolda — 2 males, 1 female ( NMNH) ; 30­6.xii.1977, Wolda — 3 males ( NMNH) ; UNITED STATES: Texas: Brownwood, Banks — P. ornatus holotype female ( MCZ) ; Fort Davis, Davis Mountains , 1524 m, vii­viii.1928, Poling — 1 female ( CAS) ; Brewster County, Windows Creek , 5­10.iv.1993, Gelhaus — 1 female ( NMNH) ; Jeff Davis County, Barrilla Draw , 4 mi. E Ft. Davis, 11­12.iv.1993, C & O Flint — 1 male ( NMNH) ; Kerr County, Kerrville , 1.iv.1959, Wigmore — 1 female ( CNC) .

Distribution. Belize, Costa Rica, Guatemala, Mexico, Nicaragua, Panama, USA (Texas).

Figures

FIGURE 15. Phylloicus aeneus. Male (A­C, P. centralus holotype; D­F, UMSP000067471): A— lateral view; B—dorsal view; C—ventral view; D—phallus, lateral view; E—phallus, ventral view; F—segment IV, dorsal view.

FIGURE 16. Phylloicus aeneus. Female (P. ornatus holotype): A—sterna IX, X and vaginal apparatus, ventral view; B—terga VIII­X, dorsal view; C—sternum VIII, ventral view.

FIGURES 1­3. —1, Phylloicus aeneus (UMSP000034901). Head, lateral. —2, Phylloicus mexicanus (UMSP000041378). Head, dorsal; — 3, Phylloicus mexicanus (UMSP000041378) Pro­ and mesothorax, dorsal.

FIGURES 6­8. —6, Phylloicus aeneus (UMSP000000137). Forewing. —7, Phylloicus angustior (UMSP000027443). Forewing. —8, Phylloicus pulchrus (UMSP000066748). Forewing.

FIGURES 117-118. — 117. Phylloicus abdominalis (UMSP000000285). A— male habitus, lateral; B— antenna, scape and basal flagellomeres, anterior view. — 118. Phylloicus aeneus (composite of UMSP000068398, UMSP000068399, UMSP000068400). Habitus, dorsal.

FIGURES 105-108. — 105. Phylloicus aeneus (composite of UMSP000068398, UMSP000068399, UMSP000068400). Forewing. — 106. Phylloicus amazonas (composite of UMSP000005724-5, UMSP000068383-9). Forewing. — 107. Phylloicus bicarinatus (composite of UMSP000068393 and UMSP000068394). Forewing. — 108. Phylloicus cubanus (composite of UMSP000009806 and UMSP000009807). Forewing.