A New Species of Smicridea Mclachlan (Trichoptera:Hydropsychidae) from Venezuela and Its Role in Travertine Biogenesis

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A new species of Smicridea McLachlan (Trichoptera:Hydropsychidae) from Venezuela and its role in travertine biogenesis

HENRIQUE PAPROCKI1,RALPH W. H OLZENTHAL Department of Entomology, University of Minnesota, 1980 Folwell Avenue, St. Paul, Minnesota 55108 USA

CLAUDIA CRESSA Instituto de Zoologı´a Tropical, Universidad Central de Venezuela, 1041-A Caracas, Venezuela

Abstract. We collected an undescribed hydropsychid caddisﬂy, Smicridea (Smicridea) travertinera,n. sp., from 2 sites in Venezuela. One of the sites, Quebrada El Charo, ﬂowed over extensive calcareous formations of travertine, which were covered with retreats and capture nets of the new species. Smicridea travertinera was the most abundant aquatic insect colonizing travertine. We describe the adult male, the retreat and net, and gut contents. The retreat consisted of an aperture in the travertine with a capture net. Retreat-making behavior appears to cause both the biogenesis and erosion of the travertine formations. Key words: Smicridea, Hydropsychidae, Trichoptera, travertine, Venezuela, taxonomy, systematics, biogenesis, retreat-making behavior, gut contents.

Travertines are composed of calcareous sedi- where strong currents prevail. These caddisﬂies ments, chemically or biologically precipitated constructed cylindrical retreats with capture from waters of karstic, geothermal, or artesian nets on the travertine surface, which became im-

origin, which are supersaturated with CO2 portant substrates for CaCO3 deposition (Drys- (Drysdale 1999, Fig. 1A). In such conditions, dale 1998). Other important studies on traver-

CO2 rapidly outgasses to the atmosphere and tine biogenesis include Drysdale (2001) on the

CaCO3 precipitates out of solution as calcite or hydrochemistry of a travertine-depositing aragonite. This precipitate can cover all available stream in Australia, Freytet and Verrecchia inorganic and organic substrates, such as rocks, (1995) on Ca crystals associated with fungi in leaves, twigs, and also caddisfly cases and re- travertines, Pentecost et al. (1997) on the influ- treats (Fig. 1B–D). Deposition of travertines can ence of phototrophic microorganisms on trav- be highly influenced by living organisms. The ertine deposition, and Humphreys et al. (1995) biogenesis of travertines has been widely stud- on aquatic insect biogenesis of freshwater tufa ied, but mostly through the activity of diatoms dams. and bacteria (Pentecost et al. 1997). Few re- Among the caddisflies, members of the family searchers have studied the role of other aquatic Hydropsychidae have the greatest potential to biota in travertine formation. influence travertine deposition. Hydropsychid Aquatic insects can influence both deposition larvae construct fixed retreats with silken cap- and erosion of travertines (Thienemann 1933, ture nets that provide a suitable surface area for Drysdale 1999). Drysdale (1998) reported that CaCO3 precipitation. We describe a new species aquatic insect larvae play direct and indirect in the hydropsychid genus Smicridea, subgenus geomorphological roles in the deposition of Smicridea McLachlan, from Venezuela, the larvae travertines in the Barkly Karst, Australia. He of which inhabit travertine formations. We also demonstrated that species of Cheumatopsyche discuss aspects of the biology of this species and (Hydropsychidae) were the dominant members its influence on biogenesis of travertine forma- of the aquatic fauna on the surface of travertines tions. The genus Smicridea is widespread in the Neo- 1 Winner of the Hydrolab Student Award for the Best Poster Presentation in Basic or Applied Research tropical realm where 162 species occur, 16 of at the 50th Annual NABS Meeting, Pittsburgh, Penn- which have been recorded from Venezuela (Flint sylvania, 27 May–1 June 2002. E-mail address: et al. 1999). Known Smicridea larvae build typi- [email protected] cal hydropsychid retreats and capture nets

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(Wiggins 1996); none have been reported pre- TABLE 1. Physicochemical parameters of stream viously from travertine formations. Previous de- water at Quebrada El Toro and Quebrada El Charo, scriptions of larvae and pupae of Smicridea are Estado Lara, Venezuela, measured at 1800 h on 11 and found in Correa et al. (1981), Flint (1974, 1989), 12 June 2001, respectively. and Wiggins (1996). Adult taxonomic works in- Quebrada Quebrada clude Flint (1974) on the North and Central Parameter El Toro El Charo American species, Flint (1989) on the Chilean Њ fauna, and Blahnik (1995) on the fasciatella com- Air temperature ( C) 26.90 26.60 Water temperature plex. (ЊC) 22.50 24.30 pH 8.10 8.10

Methods O2 (mg/L) (% of saturation) 7.15 (83.0) 6.92 (84.3) Study area Conductivity (␮S/cm) 468.80 608.00 Ca (mg/L) 61.60 31.70 We collected the new Smicridea species from 2 Mg (mg/L) 4.73 7.02 streams in the northwestern Venezuelan state of Na (mg/L) 7.60 11.62 Lara within or near Parque Nacional Cueva de K (mg/L) 0.47 0.73 la Quebrada del Toro (see Gabaldoń 1987, 1992 Cl (mg/L) 14.10 13.00 for description of the park). The undisturbed SO4 (mg/L) 16.00 30.00 vegetation in the region consists of dense, hu- mid, pre-montane forest. The canopy shades most of the riverbed, and contains mostly Ery- fall occurs ϳ250 m upstream from the collecting thrina poeppigiana (Fabaceae:Papilionoideae), site, just downstream from a large spring that Roystonea regia (Arecaceae), Trichilia hirta (Meli- forms the headwaters. We visited the site in aceae), Tabebuia rosea (Bignoniaceae), and Bau- June, just prior to the rainy season (delayed in hinia spp. (Fabaceae:Caesalpinioideae). The ge- 2001). Stream flow was at bankfull, suggesting ology of the region is characterized by a number that Quebrada El Charo is a perennial stream. of Cenozoic faults, with limestone bedrock Physicochemical parameters of the stream water forming the main component. The rainy season for both sites are shown in Table 1. lasts ϳ7 to 8 mo (May to November) with mean annual precipitation ranging from 1100 to 2200 Specimen collection and preparation mm. Water dissolves the limestone to produce deep fissures and sinkholes, and percolates Adults were primarily attracted to an ultra- through fissures contributing to a network of violet light placed in front of a white bed sheet karst. Average annual minimum and maximum and adjacent to the river; additional adults were temperatures in the area are 18 and 24ЊC, re- netted during the day. Adults were collected in- spectively. dividually in potassium cyanide kill jars and We collected adult caddisflies at 2 sites, Que- subsequently pinned. Larvae and pupae were brada El Toro, within the National Park (lat removed from the travertine, and 2 travertine 10Њ49.581Ј N, long 69Њ07.990Ј W), and nearby sections (range 140–850 cm3) were removed Quebrada El Charo (lat 10Њ46.771Ј N, long from the upper part of a barrage for examina- 69Њ12.174Ј W). The Quebrada El Charo site con- tion of larval retreats in the laboratory. sists of a series of emerald-green pools alternat- Male and female genitalia were cleared in lac- ing with travertine barrages (Fig. 1A). We clas- tic acid for 10 to 30 min at 125ЊC, and then sified these travertines as meteogene travertines rinsed in distilled water; remaining soft tissues (Pentecost and Viles 1994). A 70-m high water- were removed from the genitalia with fine-

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FIG.1. Smicridea travertinera habitat and retreat. A.—Travertine formations at Quebrada El Charo, Estado Lara, Venezuela. B.—Retreat of S. travertinera (scale bar ϭ 2.7 mm). C.—Anterior opening of retreat showing peripheral lip (scale bar ϭ 0.8 mm). D.—Cross section of travertine matrix showing larval tunnels (scale bar ϭ 7 mm).

tipped forceps. Genitalia were examined in no wing pattern and no white spot on the head. watch glasses with glycerin under an Olympus௡ However, an additional series of S. riita speci- SZX 12 stereomicroscope (up to 216ϫ). Pencil mens examined from another Venezuelan local- drawings were made using a 10 ϫ10 mm ocular ity displayed a range of variation from highly grid via transfer to 2.5 ϫ 2.5 cm grid drawing patterned, as in S. travertinera, to indistinctly pat- paper. Drawings were then scanned, and final terned. illustrations of male genitalia were made using ௡ Adobe Illustrator 9.0 . Terminology for male Description genitalic structures follows Schmid (1998). Adults and larvae were associated using the Adult. Length of forewing 4–6 mm. Head metamorphotype method (Wiggins 1996). flattened dorsally, with round patch of white se- We assessed larval diet by removing the di- tae on mesodorsal surface. Antennae and thorax gestive tract from 4th instars (n ϭ 10). Digestive fuscous. Forewing color fuscous, with an apical tracks were placed in alcohol and shredded. Gut fringe of white setae (easily rubbed off), and contents were then examined using an Olym- with 2 transverse bands of white setae, the 1st pus௡ BH-2 compound microscope (up to 500ϫ). band occurring at the basal 3rd,andthe2nd band Pieces of capture nets also were examined mi- at the apical 3rd of the forewing. Legs fuscous, croscopically and photographed with a digital with tarsi of fore and midlegs silvery. Abdomen camera attached to the stereomicroscope. Digital fuscous. images were imported into Auto-Montage௡ Male genitalia (Figs 2–6). Segment IX roughly (Syncroscopy, Frederick, Maryland) and then triangular when viewed laterally, anterior mar- measured using the ‘‘Measure Length’’ com- gin slightly produced medially, posterior mar- mand. gin slightly sinuous, excavated basally; sternum Type material of the new species described IX narrow. Segment X subrectangular, broad ba- was deposited in the University of Minnesota sally, tapering to broad, slightly transverse trun- Insect Collection, St. Paul, Minnesota (UMSP), cate apices; lateral margin rounded, setose; ter- National Museum of Natural History, Smithson- gum X membranous at base, medially with pair ian Institution, Washington, DC (NMNH), and of membranous dorsolateral ridges bordering the Instituto de Zoologıá Agrıćola, Maracay, deeply excavated area; surfaces covered with Venezuela (IZAM). minute setae. Inferior appendage 2-segmented, tubularly elongate, basal segment straight, slightly inflated apically, apical segment short Smicridea travertinera new species with apex rounded, slightly curved medially. Figs 2–10 Phallus tubular, angled from base, slightly en- larged apically; apex membranous ventrally, Diagnosis with pair of dorsolateral plates; internal sclerite with apex curved downward, with dorsolateral Morphology of male genitalia of S. travertinera plate connecting the dorsal and ventral arms. is similar to S. (Smicridea) riita Flint 1981; how- Larva (Figs 7–10). Length 5–7 mm. Head ever, some diagnostic differences occur. In the dark brown, circular in dorsal view, flattened new species the phallus is straighter (Fig. 3) and dorsally, with distinct U-shaped marginal cari- the apical sclerites differ from those in S. riita. na. Frontoclypeal apotome broad anteriorly, al- In S. travertinera the apical phallic sclerites con- most reaching carina posteriorly, coronal suture sist of a pair of dorsolateral plates, including an short; ventral margin of head broadly notched internal sclerite with its apex curved downward posteromesally, ventral apotome triangular. No- and with the dorsolateral plate connecting the tum of each thoracic segment sclerotized; pro- dorsal and ventral arms (Figs 3, 5). In addition, notum dark, divided on midline; meso- and the inferior appendages are broader in S. trav- metanota entire, lighter in color. Foretrochantin ertinera (Figs 2, 6). The forewing has 2 transverse simple. Legs similar, although prothoracic leg bands of white setae and an apical fringe of somewhat shorter than meso- and metathoracic white setae. The mesodorsal surface of the head legs. Abdomen covered with flattened, scalelike contains a small patch of white setae. The ho- setae and long setae (patterned as in Fig. 8), lotype of S. riita is completely fuscous, showing without lateral fringe; ventral gills with bulbous

FIGS 2–6. Male genitalia of Smicridea travertinera, n. sp. 2.—Abdominal segments IX and X (lateral view). 3.—Phallus (lateral view). 4.—Abdominal segments IX and X (dorsal view). 5.—Phallus apex (dorsal view). 6.—Inferior appendage (ventral view). base bearing 1–4 filaments, extending from tho- setal tuft on anal proleg bearing 5 long mesal racic segment II to abdominal segment VI; ven- and 5 long lateral setae, setal length not exceed- trolateral gills with 3–5 filaments on abdominal ing 2ϫ length of anal proleg. segments I–VII; lateral gills reduced, extending Larval retreat (Fig. 1B–D). Length up to 7 from abdominal segments III–VII. Abdominal mm, consisting of an elongate-oval, dome- sternum VIII with trapezoidal sclerite bearing shaped amalgamation of travertine particles numerous setae; sternum IX with 2 triangular fixed to the travertine matrix (Fig. 1B); anterior sclerites bearing numerous setae; segment X opening circular, 893.3–1025 ␮m in diameter with conical membranous projection dorsally; (mean ϭ 923, n ϭ 10, Fig. 1C), with silken cap- anal prolegs with subrectangular lateral sclerite ture net covering opening, often with peripheral narrowing anteriorly and projecting dorsally; lip of travertine particles (Fig. 1C); small orifice

FIGS 7–10. Larva of Smicridea travertinera, n. sp. 7.—Head and thorax (dorsal view). 8.—Lateral view. 9.— Head (ventral view). 10.—Abdominal segments VIII–X and abdominal prolegs (ventral view). at posterior end, 240.4–360.8 ␮m in diameter W, el. 425 m, 12.vi.2001, R. W. Holzenthal, R. J. (mean ϭ 302.3, n ϭ 10, Fig. 1B), often with ac- Blahnik, H. Paprocki, & C. Cressa–1 ( (UMSP) cessory opening internally at retreat bottom; (pinned). Paratypes: Same, 10 (,2&(UMSP) mesh size of net ranging from 30.8 ϫ 41.2 ␮m (pinned), 2 (,1& (NMNH) (pinned); Parque to 42.2 ϫ 43.8 ␮m (mean ϭ 38.6 ϫ 43.0 ␮m, n Nacional Cueva de la Quebrada del Toro, Que- ϭ 20). brada del Toro, 10Њ49.581Ј N, 69Њ07.990Ј W, el. 530 m, 11.vi.2001, R. W. Holzenthal, R. J. Blah- ( Material examined nik, H. Paprocki, & C. Cressa–4 (UMSP) (pinned), 2 (,1& (IZAM) (pinned). Other ma- Holotype: VENEZUELA, Falco´n: Quebrada terial studied [VENEZUELA], Falco´n: Quebra- El Charo at cataratas, 10Њ46.771Ј N, 69Њ12.174Ј da El Charo at cataratas, 10Њ46.771Ј N,

69Њ12.174Ј W, el. 425 m, 12.vi.2001, R. W. Hol- travertinera, like M. carolina, has a posterior zenthal, R. J. Blahnik, H. Paprocki, & C. Cressa– opening that allows water to flow through the 78 larvae (UMSP) (in alcohol). retreat, which aerates the gills and evacuates fe- ces. Wallace and Sherberger (1974) also inferred Etymology that the flat, carinate head of M. carolina serves to direct water flow through the different cham- A travertine builder in Spanish would be a bers of the retreat and to protect against pred- ‘‘travertinera’’, referring to the biogenesis of the ators. We think that the flat, carinate head of S. travertine formations at the type locality. travertinera has similar functions. Using scanning electron microscopy, Drys- Biology dale (1998) observed that the size of the CaCO3 particles around the retreat aperture of an Aus- In Quebrada El Charo, the surfaces of the tralian Cheumatopsyche were comparable to the travertine barrages were mostly covered by re- size of larval mouthparts, indicating intentional treats of S. travertinera (Fig. 1A). Other aquatic placement. The shape and structure of the ap- insects were found in lower densities, including erture of the retreats of S. travertinera (Fig. 1C)

Diptera (Chironomidae), Ephemeroptera (Bae- also suggest that CaCO3 fragments are inten- tidae), Coleoptera (Elmidae), and Trichoptera tionally positioned by larvae, especially evident (Helicopsychidae, Philopotamidae). in the lip built around the rim of the anterior The retreat of S. travertinera (Fig. 1B–D) differs opening (Fig. 1C). from all known Smicridea retreats. Previously Continual retreat- and net-building activities described Smicridea larvae construct typical hy- of larvae contribute both to the deposition and dropsychid retreats with capture nets (Flint erosion of travertines. Flint (1996) reported that 1974, Correa et al. 1981, Oliveira and Froehlich pits gouged by Hydropsyche incommoda Hagen 1996, Wiggins 1996). Smicridea travertinera re- larvae into wooden bridge pilings eventually retreats consist of a structure amalgamated to the sulted in a loss of structural integrity and col- travertine matrix, with a capture net at the an- lapse of the pilings. Over time, the intricate net- terior end and a small oriﬁce at the posterior work of tunnels and retreats of S. travertinera ap- end (Fig. 1B). Over time, retreats become cov- pears to become the major biotic component of ered with CaCO3 precipitate, forming an intri- the travertine matrix (Fig. 1D). The process of cate network of tunnels within the travertine travertine alteration by S. travertinera as well as matrix (Fig. 1D). erosion by larval tunnels indicates both an ac-

In Quebrada El Charo, precipitation of CaCO3 tive and passive contribution by larvae toward crystals seems to foul S. travertinera nets (Fig. biogenesis of travertine formations (Drysdale 1C), and larvae constantly rebuild and clean 1999). nets, presumably to reduce clogging. We hy- Apparently, S. travertinera larvae have specific pothesize that the larvae also chew (i.e., hollow flow requirements, which are achieved at the out) the travertine matrix to construct the re- travertine barrage. If the upper border of the treat. Gut content analysis indicates that larvae travertine barrage is heavily colonized, the com- ingest diatoms and plant detritus as well large plexity of the travertine surface increases and amounts of CaCO3 crystals. We assume that the velocity of the water decreases, favoring an crystals in larval guts result both from retreat increase in precipitation of CaCO3 crystals. This building and net-cleaning activity. event appears to be cyclical, as increased micro- topographical complexity likely also increases Discussion CaCO3 precipitation, building the height of the barrage until flow is directed elsewhere. At this Similar to S. travertinera, another species of point it appears that larvae stop colonizing that Hydropsychidae, Macrostemum carolina (Banks), area. Water becomes redirected around the bar- also produces tunnel-like retreats. Wallace and rage and begins flowing rapidly over other sub- Sherberger (1974) described the retreat of M. strates, exposing new areas for colonization and carolina as a network of tunnels and burrows the building up of other barrages. This succes- gouged by the larvae into bark and surface sion of events probably causes travertine barrag- wood of submerged tree limbs. The retreat of S. es to grow taller and stream pools to deepen.

There is little published information on the Terrence Ehrman, Russell Drysdale and 2 anon- biology of tropical Smicridea. Oliveira and ymous reviewers for valuable suggestions on the Froehlich (1996) examined gut contents, mesh manuscript. This material is based upon work size, and flight periods of an undetermined Smi- supported by the NSF grants DEB-9971885 and cridea from southeastern Brazil. The average DEB-0117772. Additional support was provided mesh size of capture nets for the species in their by the University of Minnesota Insect Collection study was 60 ϫ 68 ␮m, almost 2ϫ larger than and the Dayton-Wilkie Fund, Bell Museum of S. travertinera. Gut contents of larvae in their Natural History. study were largely diatoms. Boon (1988) reported on the habitat and biology of 3 species of Literature Cited Smicridea from Jamaica. An undetermined spe- BLAHNIK, R. J. 1995. New species of Smicridea (subge- cies of Smicridea was the only one found inhab- nus Smicridea) from Costa Rica, with a revision of iting cascades with limestone substrate. Larvae the fasciatella complex (Trichoptera: Hydropyschi- of the Jamaican species appeared to burrow into dae). Journal of the North American Benthological the limestone substrate, but did not make the Society 14:84–107. extensive excavations or retreats we report for S. BOON, P. J. 1988. Notes on the distribution and biology travertinera. Mean mesh dimensions of the Ja- of Smicridea (Trichoptera: Hydropsychidae) in Ja- maican species were 47 ␮m ϫ 36 ␮m, and gut maica. Archiv fu¨ r Hydrobiologie 111:423–433. ´ contents consisted of fine detritus and diatoms. CORREA,M.,T.MACHADO, AND G. ROLDAN. 1981. Tax- onomıá y ecologıá del Orden Trichoptera en el We also found diatoms as one of the main com- Departamento de Antioquia en diferentes pisos ponents of the diet of S. travertinera, but guts of altitudinales. Actualidades Biolo´gicas 10:35–48. final instars were so clogged with CaCO3 crys- DRYSDALE, R. N. 1998. Aquatic insect larvae as geo- tals that all food appeared miniscule in com- morphic agents in travertine-building: a case parison with ingested crystals. However, we be- study from the Barkly Karst, Australia. Supple- lieve that diatoms are a major food item for S. ment Geografia Fı´sica Dinamica Quaterna´ria 4: travertinera. 53–59. Studies of travertine-associated macroinver- DRYSDALE, R. N. 1999. The sedimentological signifi- tebrates are extremely limited. Research on the cance of hydropsychid caddis-fly larvae (Order: ecology and behavior of macroinvertebrates in- Trichoptera) in a travertine-depositing stream: Louie Creek, northwest Queensland, Australia. habiting travertine formations is needed to un- Journal of Sedimentary Research 69:145–150. derstand how these organisms influence trav- DRYSDALE, R. N. 2001. Factors controlling the hydro- ertine geomorphology. Studies of microhabitat chemistry of Louie Creek, a travertine-depositing preference by aquatic insects, faunal composi- stream in the seasonally wet tropics of northern tion of travertines, and rates of deposition and Australia. Marine and Freshwater Research 52: erosion of travertines, with or without a mac- 793–804. roinvertebrate component, would greatly im- FLINT, O. S. 1974. Studies of Neotropical caddisflies, prove our understanding of environmental in- XVII: The genus Smicridea from North and Cen- fluences on travertine formations. tral America (Trichoptera: Hydropsychidae). Smithsonian Contributions to Zoology 167:1–65. FLINT, O. S. 1981. Studies of Neotropical caddisflies, Acknowledgements XXVIII: The Trichoptera of the Rıó Limoń Basin, Venezuela. Smithsonian Contributions to Zoology HP was supported by a PhD fellowship from 330:1–61. CAPES (Coordenaçaõ de Aperfeiçoamento de FLINT, O. S. 1989. Studies of Neotropical caddisflies, Pessoal de Nı´vel Superior), Brazilian Ministry of XXXIX: The genus Smicridea in the Chilean Sub- Education. We are grateful to INPARQUES for region (Trichoptera: Hydropsychidae). Smithson- assistance with collecting permits, to Degnis Ra- ian Contributions to Zoology 472:1–45. mı´rez, Director, Parque Nacional Cueva de la FLINT, O. S. 1996. Caddisflies do count: collapse of the Quebrada del Toro, and to park rangers Ce´sar S.R. 675 bridge over the Pocomoke River, Poco- moke City, Maryland. Bulletin of the North Adans, Alı´ Baldallo, and Delcis Reyes. We also American Benthological Society 13:376–383. thank Ce´zar Cordero for field assistance, Oliver FLINT,O.S.,R.W.HOLZENTHAL, AND S. C. HARRIS. S. Flint, Jr., for insightful comments and the loan 1999. Catalog of the Neotropical Caddisflies (In- of specimens, Kris Kuda for illustrating larvae, secta: Trichoptera). Ohio Biological Survey, Co- and Fernando Munõz-Quesada, Roger Blahnik, lumbus, Ohio.

FREYTET,P.,AND E. VERRECCHIA. 1995. Discovery of PENTECOST, A., S. BAYARI, AND C. YESERTENER. 1997. Ca oxalate crystals associated with fungi in moss Phototrophic microorganisms of the Pamukkale travertines (bryoherms, freshwater heterogeneous travertine, Turkey—their distribution and influ- stromatolites). Geomicrobiology Journal 13:117– ence on travertine deposition. Geomicrobiology 127. Journal 14:269–283. GABALDO´ N, M. 1987. Venezuela un paı´s sembrado de PENTECOST,A.,AND H. A. VILES. 1994. A review and parques. Instituto Nacional de Parques (IN- assessment of travertine classification. Geógra- PARQUES). Sede Principal de INPARQUES, Lado phie Physique et Quaternaire 48:305–314. Sur del Museo de Tansporte, Distribuidor Santa SCHMID, F. 1998. The insects and arachnids of Canada. Part 7. Genera of the Trichoptera of Canada and Cecilia, Caracas, Venezuela. adjoining or adjacent United States. NRC Re- GABALDO´ N, M. 1992. Parques Nacionales de Venezue- search Press, Ottawa, Ontario. la. Serie Parques Nacionales y Conservacioń Am- THIENEMANN, A. 1933. Mu¨ ckenlarven bilden Gestein. biental. Instituto Nacional de Parques, Caracas, Natur und Museum, Senckenbergische Naturfor- Venezuela. schende Gesellschaft 63:370–378. HUMPHREYS, W. F., S. M. AWRAMIK, AND M. H. P. JEBB. WALLACE,J.B.,AND F. F. S HERBERGER. 1974. The larval 1995. Freshwater biogenic tufa dams in Madang retreat and feeding net of Macronema carolina Province, Papua New Guinea. Journal of the Royal Banks (Trichoptera: Hydropsychidae). Hydrobio- Society of Western Australia 78:43–54. logia 45:177–184. OLIVEIRA,L.G.,AND C. G. FROEHLICH. 1996. Natural WIGGINS, G. B. 1996. Larvae of the North American history of three Hydropsychidae (Trichoptera, In- caddisfly genera (Trichoptera). 2nd edition. Uni- secta) in a ‘‘Cerrado’’ stream from northeastern versity of Toronto Press, Toronto, Ontario. Saõ Paulo, Brazil. Revista Brasileira de Zoologia Received: 16 October 2002 13:755–762. Accepted: 12 June 2003

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