Phylloicus passulatus Prather 2003

Souza-Holanda, Paula Mayara De, Pes, Ana Maria & Hamada, Neusa, 2020, Immature stages of three species and new records of five species of Phylloicus Müller (Trichoptera, Calamoceratidae) in the northern region of Brazil, Zootaxa 4851 (1), pp. 111-136 : 126-134

Phylloicus passulatus Prather 2003

( Figs 12–18 , 23 )

— Prather 2003: 83 [Type locality: Venezuela, Amazonas, Puerto Ayacucho (40 km S) El Tobogán, Caño Coromoto; NMNH; ♂; ♀] .

Diagnosis. Larva: It is similar of those of P. abdominalis , P. amazonas , and P. cressae , by the pale brown to yellow color of the head and very evident muscle scars ( Huamantinco et al. 2005, Barrios et al. 2020); it differs from that of P. fenestratus which has a uniformly brown head with the color of the scars of the dorsal and lateral muscles not conspicuous. Similar to P. amazonas , the foretrochantin has shorter spines on the ventral margin, whereas in P. fenestratus the foretrochantin has strong and prominent spines on the ventral margin. It differs from known larvae of the other species by having a narrow base on each of the anterolateral processes of the pronotum and a pre-apical region with an undeveloped lobe; in contrast, the larva of P. amazonas has anterolateral processes of the pronotum that are without curvature, gradually tapering to pointed apices and in P. fenestratus each process has a broad base and a pre-apical region with a rounded lobe that is convex, in P. cressae the anterolateral extensions of the pronotum are elongate as a pair of strongly sclerotized hooks ( Barrios et al. 2020), and in P. obliquus the anterolateral margins are strongly projecting with apices hooked anteroventrad ( Cavalcante et al. 2020). The larva of P. passulatus also differs in the color of the legs which are uniformly light brown, but those of P. amazonas are light yellow, with a brown band on median region of each femur, tibia, and tarsus and the legs of P. fenestratus are uniformly dark brown.

Case: It is flat and with each side composed of 3 to 4 fragments of leaves of rectangular or round shape such as those of P. mexicanus , P. abdominalis , P. obliquus , and P. cressae ( Wiggins 1996, Huamantinco et al. 2005, Cavalcante et al. 2020, Barrios et al. 2020). It differs from the flat cases of P. amazonas and P. lituratus ( Rueda Martín 2013) which are composed of two oval fragments of leaves on each side; it also differs from the case of P. fenestratus , which is built with 3 or 4 fragments of rounded leaves or rectangular, but it is nearly circular in cross section, not flat.

Pupa: It differs by the number of setae on the median lobe of the labrum which has a row of 24–26 long setae; in contrast, the median lobe of the labrum of the pupa of P. amazonas has a row of 16–18 median setae, that of P. fenestratus has 20–22 long setae, P. abdominalis has 24 ( Huamantinco et al. 2005), and that of P. obliquus 24–28 long setae ( Cavalcante et al. 2020). The pupa of P. passulatus also differs by the number of setae on the anal processes, each having 1 long sub-basal and 3 short apical setae on each process; the pupae of P. amazonas has 30 long setae on each process and that of P. abdominalis has only 3 short apical setae on each process ( Huamantinco et al. 2005).

Additional description: Adult. Length of each forewing, male: M = 9.23 mm (SD = 0.29, n = 6); female: M = 9.13 mm (SD = 0.38; n = 6). General body color dark gray (pinned, Figs 12a, 12b ) and brown in alcohol; dorsal portion of head black; posterior dorsal warts and setae golden yellow; maxillary palps black and labial palps pale yellow. Antennae twice as long as folded forewings, black.

Male: Tergum III with short anterolateral lateral and posterolateral flaps ( Figs 13a, 13c ); tergum IV with pair of lateral coremata each with basal globose lobe and with long tubular posterior lobe (eversible membranous sac) and most-lateral lobes sclerotized ( Figs 13a, 13c ). Phallus everted, with phallotremal sclerite horseshoe-shaped, ventral sclerite short, tubular ( Fig 13d ); endotheca with rounded lateral lobes and single dorsomesal lobe ( Figs 13 e– 13g).

5th instar larva ( Figs 14–17 ). Body length: 11.7 mm, 12.9 mm (n = 2). General color, head and thoracic sclerites brown except muscle scars light yellow; legs light brown, uniform ( Figs 14 f–14h); abdomen pale yellow ( Fig 14a ).

Head: Muscle scars ovoid and round, including three on middle and posterior regions of frontoclypial apotome ( Figs 15a, 15i ), numerous on genae. Genae rough in anterior half, except for smooth muscle scars. Ventral apotome triangular and short, not reaching posterior edge of head.

Thorax: Pronotum sclerotized, light brown, with light yellow muscle scars in posterolateral 2/3, anterolateral processes long (about 1/3 as long as pronotal sclerites), each bearing 3 long and 3 short setae on inner margin with subtruncate apex ( Figs 14b14d ); foretrochantins each as long as forecoxa and with curved and acute apex ( Figs 14d , 15f , arrow). Mesonotum with central pair of lightly sclerotized sclerites, each sa 1 with 1 short seta, sa 2 with 1 long seta and 2 short ones, sa 3 with prominent anterolateral sclerite having 8 long and 5 short setae ( Figs 14b,14c ). Metanotum membranous, each sa1 with 1 long seta, sa2 with 1 long seta and 2 short setae, sa3 consisting of an anterolateral sclerite with 3 long and 4 short setae. Legs light brown, uniform, with few long setae ( Figs 14 f–14h).

Abdomen: Membranous. Segment I with one dorsal hump and one pair of ventrolateral humps, well developed, ventrolateral humps each rounded and with sclerotized apical region ( Fig 14k ). Gills with 2–4 filaments ( Fig 17 ). Segments III–VII with lateral line composed of long, fine, pale setae; lateral portions of segment VIII with row of forked lamellae. Tergum VIII with 1 pair of thick dorsal setae, 3 pairs of small fine setae, and 1 pair of long setae. Tergite IX with 1 pair of thick dorsal setae, 3 pairs of small thin setae, and 1 pair of long lateral setae ( Fig 14j ). Anal prolegs each with sclerotized hook and 4 long and 3 short setae ( Fig 14i ).

Case ( Figs 16 a–16c). Length: M = 21.94 mm (SD = 3.19; n = 6). Flat and each side formed by 3 to 4 leaf fragments of rectangular or rounded shape; anterodorsal fragment forming small projection as protective flap for larva.

Pupa ( Figs 18 a–18d). Body length: M = 11.8 mm (SD = 0.5; n = 4).

Head: Antennae each with stout scape, cylindrical pedicel ( Fig 18a ). Labrum trilobed, basolateral lobes about 1/3 as long as median lobe and with 3 long setae on each lobe; median lobe with triangular edge and with row of 24–26 long setae (divided into two groups) in dorsal view ( Fig 18b ). Maxillary palps long and 5-articulated. Labial palps short and 3-articulated. Mandibles symmetrical, falciform and well developed, each with 2 long thick basolateral setae ( Fig 18b ).

Thorax: Pronotum narrow; mesonotum wide, about 2.5X as long as metanotum.

Abdomen: Lateral line of setae on each side extending from anterior edge of segments III to VIII, curving ventrad on sternum VIII and touching ventral midline.

Pairs of anterior hook plates on terga III–VIII each with 2 or 3 hooks curved backward, pair of posterior hook plates on terga I and V each with 5 hooks curved forward ( Fig 18e ). Tergite IX sub-square, largely membranous with small incision posteromesally, bearing prominent dark tergite with anteriorly oriented lateral projections, only primary setae present ( Figs 18 c–18e). Apex ending in pair of very long anal processes each with one long seta on basal 1/3 and 3 short setae on apicoventral 1/3; apex spatula-shaped without teeth ( Figs 18 c–18d).

Bionomy. Larvae of P. passulatus were collected in streams on leafpacks, leaves retained between stones or wood. Pupae were fixed on leaves deposited on the bottom or under stones in areas of running water. In Presidente Figueiredo municipality (AM) and in Amajarí and Bonfim municipalities (RR), larvae were collected in second- to fourth-order streams, all with rocky bottoms and fast-moving water. In the streams near the airport in the urban area of Manaus (AM) larvae were collected in a first-order stream with low water flow and with substrate of small stones and sand.

Comments. Additional description of the male adult was included because Prather (2003), failed to evert the phallic endotheca fully, and did not illustrate tergum IV with its pair of lateral coremata each with basal globose lobes and with a long tubular posterior lobe with eversible membranous sac.

Material examined. BRAZIL, Amazonas, Barcelos, Base da Serra do Aracá, igarapé da Cobra no acampamento (#B01), 00°52’24.78”N 63°27’18.97”W, 136 m a.s.l., 02.viii.2009, A.M.P. Santos, N. Ferreira Jr., F. Salles leg., Malaise trap, 1♂, 1♀ (alcohol), 1 larva (alcohol); igarapé do Jabuti (#B07), 00°52’47.78”N 63°28’27.05”W, 125 m a.s.l., 26.vii.2009, emergence 07.viii.2009, A.M.P. Santos leg., 1♂ pupal exuviae and case (alcohol), also 26.vii.2009, emergence 15.x.2009, A.M.P. Santos leg., 1♂ pupal exuviae and case (alcohol); (#B08), igarapé da Cobra, 00°52’34.21”N 63°27’03.60”W, 157 m a.s.l., 25.vii.2009, emergence 18.ix.2009, A.M.P.Santos leg., 1♀ (pinned) and pupal exuviae and case (alcohol); Parque Estadual da Serra do Aracá , (#S07), 00°54’13.68” N 63°26’01.18” W, 1112 m a.s.l., 30.vii.2009, emergence 15.viii.2009, A.M.P. Santos leg., 1♂ (pinned) and pupal exuviae and case (alcohol); igarapé do Reginel (afluente da direita do rio Negro Px. a Barcelos) (#21), 01°02’30.98”S 62°50’09.20”W, 35 m a.s.l., 10.viii.2009, emercence 25.ix.2009, A.M.O. Pes leg., 2♀ (pinned). Manaus, igarapé na estrada do Turismo, mata do Aeroporto 03º02’29.95”S 60º04’12.53”W, 30.vi.2008, J.O. da Silva leg., 1♂ (alcohol). Presidente Figueiredo, igarapé Canoas (Cachoeira), 01º49’51”S 061º04’15”W, 07.iii.2002, A.M.O. Pes leg., 1 pupa (♀) (alcohol); 04.viii–05.ix.2008, A.M.O. Pes leg., 1♂ (alcohol) and 1♂ pupal exuviae and case (alcohol), 1♂, 1♀ (pinned), 1 pupa (♀) (alcohol), 1 pupa (♀) (alcohol); 04.viii.2008, emergence 10.viii.2008, A.M.O. Pes leg., 1♀, pupal exuviae, and case (alcohol); 04.viii.2008, emergence 02-ix.2008, A.M.O. Pes leg., 1♀, pupal exuviae, and case (alcohol); 04.viii.2008, emergence 04-ix.2008, A.M.O.Pes leg. 1♂ (alcohol); 04.viii.2008, emergence 09.ix.2008, A.M.O. Pes leg., 1♀ (pinned), pupal exuviae and case (alcohol); 04.viii.2008, A.M.O. Pes leg., light trap, 1♂ (alcohol); 12.viii.2014, light trap, A.M.O. Pes leg., 3♂ (alcohol) and 1♂ (pinned); 12.viii.2014, emergence 02.ix.2014, A.M.O. Pes leg. 1♀ (pinned), pupal exuviae and case (alcohol); igarapé da Caverna do Maroagoa , km 6 AM 240 ”, 01º04’20.33”S 059º58’54.24”W, 28.ix.2000, emergence 03.x.2000, A.M.O. Pes, J.O. da Silva leg., 1♀, pupal & larval exuviae, case (alcohol); 3–10.vii.2002, N. Hamada, M. Pepinelli, R. Kikuchi leg., 1♀, pupal & larval exuviae, case (alcohol); 12.viii.2014, A.M.O. Pes leg., 2 larvae, case (alcohol); 04.viii.2018, emergence 20.xi. 2008, A.M.O. Pes leg., 1♀, pupal & larval exuviae, case (alcohol); Reserva Biológica do Uatumã, igarapé 3, 01º48’54”S 059º15’93”W, 03.ii.2008, J.L.D. Albino leg., 2 larvae, 1 pupa (♂), larval exuviae, case (alcohol). Rio Preto da Eva , ZF3—Fazenda Esteio, igarapé, acampamento do Colosso ( PDBFF [ Programa de Dinâmica Biológica de Fragmentos ]), 02º25’01”S 59º52’15”W, 16.iv.2002, J.L. Nessimian leg., Pennsylvania trap, 1♂ (alcohol); Escola Adventista Agroindustrial, 02º41’41.3”S 59º44’06.05”W, 13.iv.2008, C.A.S. Azevedo, M. Pepinelli, U.G. Neiss leg., Pennsylvania trap, 1♂ (alcohol). Roraima: Bonfim, 02º00’57.8”N 60º00’56.6”W, 14.ix.2010, emergence 20.x.2010, A.M.O. Pes leg., 2♂, larval exuviae, case (alcohol). Amajarí, Serra do Tepequém , igarapé # R10 , 03°47’32.5”N 061°45’12.7”W, 505 m a.s.l., 03.vii.2016, emergence 11.vii.2016, A.M.O. Pes, G. Amora, J.O. da Silva, G. Dantas leg., Malaise trap, 1♂, 2♀ (alcohol) .

Distribution. Brazil (AM, RR new records) and Venezuela ( Fig 23 ).

Figures

FIGURE 12. Phylloicus passulatus Prather 2003, female. 12a, dorsal habitus; 12b, left lateral habitus (scale bar = 2 mm).

FIGURE 13. Phylloicus passulatus Prather 2003, male. 13a–13c, abdomen, dorsal, ventral, and right lateral, respectively (scale bar = 1 mm); 13d, 13e, phallus, left lateral and ventral, respectively; 13f, 13g, phallus apex, ventral and caudal, respectively.

FIGURE 14. Phylloicus passulatus Prather 2003, larva. 14a, right lateral habitus (scale = 2 mm); 14b, head, thorax, and abdominal segment I, dorsal (scale = 1 mm); 14c, thorax, dorsal (scale = 1 mm); 14e, pronotal plates, dorsal (scale = 0.5 mm); 14d, right foretrochantin (arrow) and right pronotal plate, right lateral (scale = 0.5 mm); 14f–14h, right fore-, mid-, hind legs, respectively, right lateral (scale = 0.5 mm); 14i, 14j, abdominal segment IX, lateral sclerite, and anal claw, right lateral and dorsal, respectively; 14k, abdominal segment I, ventrolateral humps, ventral.

FIGURE 15. Phylloicus passulatus Prather 2003, larva. 15a–15c, head, dorsal, ventral, and right lateral, respectively (scale = 0.5 mm); 15d, labrum, dorsal (scale = 0.2 mm); 15e–15g, left and right mandibles, dorsal, and right mandible, mesal, respectively; 15h, mentum and ventral apotome, ventral (scale bar = 0.2 mm); 15i, frontoclypeal apotome and labrum, dorsal; 15j, right foretrochantin (arrow) and forecoxa, right lateral view.

FIGURE 16. Phylloicus passulatus Prather 2003, larval case. 16a–16c, ventral, right lateral and anterior, respectively (scale bar = 2 mm).

FIGURE 17. Phylloicus passulatus Prather 2003, larval gill diagram with positions of gills and number of gill filaments on abdominal segments I–VIII.

FIGURE 18. Phylloicus passulatus Prather 2003, pupal exuviae. 18a, dorsal habitus (scale bar = 2 mm); 18b, head, anterodorsal view; 18c, 18d, abdominal segments VIII and IX, left = dorsal, right = ventral, respectively; 18e, hook plates of abdominal segments I–IX, dorsal (a = anterior and p = posterior position).

FIGURES 19–23. Distribution map of Phylloicus species. 19, P. amazonas Prather 2003; 20, P. dumasi Santos & Nessimian 2010; 21, P. elektoros Prather 2003; 22, P. fenestratus Flint 1974; 23, P. passulatus Prather 2003, (The states of the northern region of Brazil are highlighted in green).