A review of Skutzia Reiss, 1985, with the description of three new species (Diptera:Chironomidae:Chironominae)

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1 J. N. Am. Benthol. Soc., 2009, 28(1): Ó 2009 by The North American Benthological Society DOI: / Published online: 16 December 2008 A review of Skutzia Reiss, 1985, with the description of three new species (Diptera:Chironomidae:Chironominae) Luiz Carlos de Pinho 1 Laboratório de Entomologia Aquática, Departamento de Biologia da Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Av. Bandeirantes, 3900, Monte Alegre, CEP: Ribeirão Preto - São Paulo, Brazil Humberto Fonseca Mendes 2 AND Trond Andersen 3 The Natural History Collections, Bergen Museum, University of Bergen, Muséplass 3, NO-5020 Bergen, Norway Abstract. Skutzia epleri sp. n. from USA, S. inthanonensis sp. n. from Thailand, and S. quetzali sp. n. from Panama and Mexico are described and figured as male imagines, and S. gaianii Andersen is recorded from Trinidad and Tobago. The genus now consists of 6 species. In addition to the species mentioned above, S. inopinata Reiss from Canada and S. bahiensis Reiss from Brazil are included. Skutzia is placed in the subtribe Zavreliina of the tribe Tanytarsini, but because the immatures are not known, this placement must be regarded as tentative. The distribution of the genus, previously known only from the Nearctic and the Neotropical regions, is expanded to include the Oriental region, indicating a Beringian connection. An emended diagnosis and a key to the males of Skutzia are provided. Key words: Chironomidae, Tanytarsini, Skutzia, new species, Neotropics, Oriental region. The genus Skutzia was erected by Reiss (1985) based on S. inopinata from Vancouver Island, Canada, and S. bahiensis from Bahia State, northeastern Brazil. Later, Andersen (2000) described S. gaianii from Venezuela. The female and immatures of Skutzia are not known. Within Chironominae, the genus is placed in the subtribe Zavreliina of the tribe Tanytarsini, which also includes Afrozavrelia Harrison, 2004; Constempellina Brundin, 1947; Friederia Sæther and Andersen, 1998; Neostempellina Reiss, 1984; Seppia Ekrem and Sæther, 2000; Stempellina Thienemann et Bause in Bause, 1913; Stempellinella Brundin, 1947; Thienemanniola Kieffer, 1921; and Zavrelia Kieffer, However, the placement of Friederia, Seppia, and Skutzia is still tentative because the immatures are not known for these genera (Sæther and Andersen 1998, Ekrem and Sæther 2000, Sæther and Roque 2004, Ekrem 2007). Three species new to science are described: Skutzia epleri from North Carolina, USA; S. inthanonensis from 1 address: chironomidae@usp.br 2 To whom correspondence should be addressed. humberto.mendes@bm.uib.no 3 address: trond.andersen@zmb.uib.no 196 northern Thailand; and S. quetzali from Panama and southeastern Mexico. Thus, the known distribution of the genus is expanded to include the Oriental Region. Furthermore, S. gaianii Andersen is recorded from Trinidad and Tobago. An emended generic diagnosis, diagnoses for all included species, and a key to the males of Skutzia are provided. Material and Methods The specimens were mounted on slides in Canada balsam or Euparal following the procedure outlined by Pinder (1989). Morphological terminology and abbreviations follow Sæther (1980; see Table 1 for a list of the ratios used in the descriptions). Measurements are given as ranges, followed by the mean, when 4 specimens were measured, followed by the number of specimens measured in parentheses if fewer than initially stated. Pencil drawings were made using a Nikon t light microscope with drawing tube (Nikon Optiphot-2; Nikon Corporation, Tokyo, Japan). The drawings of the hypopygia were later inked, scanned, and assembled in Adobe Photoshop 7 t (Adobe Systems Inc., Berkeley, California). The remaining illustrations

2 2009] AREVIEW OF SKUTZIA REISS 197 TABLE 1. List of the ratios used in the descriptions. Abbreviation Name AR Antennal ratio Length of ultimate antennal flagellomere divided by combined length of the remaining proximal flagellomeres BV Beinverhältnisse Combined length of femur (fe), tibia (ti), and basitarsus (ta 1 ) divided by combined length of the remaining tarsomeres (ta 2,ta 3,ta 4, and ta 5 ) HR Hypopygium ratio Length of gonocoxite divided by the length of gonostylus HV Hypopygium value Total length divided by the length of the gonostylus 3 10 LR Leg ratio Length of basitarsus (ta 1 ) divided by the length of tibia (ti) SV Schenkel-Scheine- Combined length of femur (fe) and tibia (ti) divided by the length of basitarsus (ta 1 ) Verhältnisse VR Venarum ratio Length of cubitus (Cu) divided by the length of media (M) were made using the software Adobe Illustrator CS2 t (Adobe Systems Inc., Berkeley, California), with scanned pencil drawings defined as templates. The type material is housed in Zoologische Staatssammlung, Munich, Germany (ZSM); National Museum of Natural History, Washington, DC, USA (NMNH); Museo del Instituto de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela, Maracay, Venezuela (MIZA); and the Natural History Collections, Bergen Museum, University of Bergen, Bergen, Norway (ZMBN). Skutzia Reiss Skutzia Reiss, 1985: 173; Cranston et al. (1989: 412). Type species. Skutzia inopinata Reiss, 1985: 174, by original designation. Other included species. Skutzia bahiensis Reiss, 1985: 175; S. epleri sp. n.; S. gaianii Andersen, 2000: 121; S. inthanonensis sp. n.; S. quetzali sp. n. Diagnostic characters The males of the genus may be distinguished from all other Tanytarsini on the broadly based superior volsella with long, slender, sickle-shaped distal part. Generic diagnosis Male. Small species, wing length mm. Thorax pale to yellowish brown; head, legs, and abdomen pale. Antenna. With 12 flagellomeres, fully plumed; groove beginning on flagellomere 4; sensilla chaetica present on flagellomeres 1 to 4, and 12; apical setae present. AR Head. Eye bare, with short, triangular dorsomedial extension. Frontal tubercles absent. Palpomeres normal; palpomere 3 with 1 7 lanceolate sensilla clavata subapically. Temporals few, inner and outer verticals and postorbitals present. Cibarial pump long, narrow, with concave dorsal margin. Thorax. Antepronotal lobes widely separated dorsally. Scutal tubercle absent. Antepronotals absent; 5 9 uniserial dorsocentrals; 8 20 long, biserial acrostichals; 1 prealar, supraalars absent. Scutellum with 2 4 setae. Wing. Membrane densely covered with setae. Costa not extended; R 2þ3 indistinct distally; R 4þ5 ending above or somewhat proximal to apex of M 3þ4 ; FCu slightly distal to RM; false vein above Cu and M 3þ4. Brachiolum with 1 seta, M and RM bare, remaining veins with setae. Anal area of wing not developed. Squama bare. Legs. Apex of foretibia with straight, slender spur; mid- and hind tibiae each with short, well-developed combs, without spurs. Sensilla chaetica and pulvilli absent. Hypopygium. Anal tergite bands short to well developed. Anal point moderately long, wide and broadly rounded distally, with or without distally fused anal crests. Superior volsella broad basally, with single inner seta; distal part long and slender, sickle shaped, with 2 3 long setae distally and 3 4 long lateral setae; digitus absent; microtrichia sometimes present in basal 2 3. Inferior volsella extending to about midpoint of gonostylus. Median volsella short to moderately long, straight; distally with clump of slender pointed lamellae. Gonostylus very short, with or without short thorn-like process distally. Female and immatures. Unknown. = Systematics and Biogeography The tribe Tanytarsini of the subfamily Chironominae includes 3 main groups of genera, the subtribe Zavreliina and the subtribe Tanytarsina, which can be divided into a Microspectra group and a Tanytarsus group. The most important character for separating the subtribes Zavreliina and Tanytarsina is that, in Zavreliina, the costa ends well proximal to apex of M 3þ4, whereas in Tanytarsina, the costa ends slightly proximal to, opposite, or distal to the apex of M 3þ4 (Sæther and Roque 2004). In Skutzia, the costa ends

3 198 L. C. PINHO ET AL. [Volume 28 more or less opposite to the apex of M 3þ4, so based on this character alone, Skutzia should belong in Tanytarsina. Another important character is that the superior volsella lacks a digitus in Zavreliina, whereas it has a digitus in Tanytarsina. The superior volsella in Skutzia lacks a digitus, so based on this character, the genus possibly belongs in Zavreliina. In an attempt to establish the phylogenetic placement of the genus Nandeva Wiedenbrug, Reiss and Fittkau, 1998, Sæther and Roque (2004) did a parsimony analysis for all genera of Tanytarsini and found that the tribe Tanytarsini and the subtribe Zavreliina are monophyletic. In both cladograms presented, Skutzia is included in Zavreliina as the sister genus of Pontomyia Edwards, However, Sæther and Roque (2004) concluded that phylogeny of the subtribe will remain tentative until the females and immatures of more genera are known. With the 3 new species included, Skutzia shows a Beringian distribution pattern, i.e., track 4 in Sæther (2000) and Sæther and Ekrem (2003). This pattern of distribution, over the Beringian Strait Bridge that connected North America and East Asia, also is found in some orthoclad genera, such as Antillocladius Sæther, 1981 and Compterosmittia Sæther, 1981 (Mendes et al. 2004). Skutzia, Antillocladius, and Compterosmittia, also are recorded from the Caribbean. Biology The larvae of Skutzia are not known. However, they can be expected to construct transportable cases of sand grains, small wood, or plant remains, as seen in other larvae of Zavreliina species. The adults of S. epleri sp. n., S. gaianii, and S. inopinata have been captured close to coldwater streams, or streams and seeps in mountainous areas.900 m asl. Skutzia inthanonensis sp. n. also was collected in a mountainous area rich in streams and small rivers. Skutzia quetzali sp. n. is recorded from a tropical lowland rainforest with small lakes and intermittent streams. Skutzia bahiensis also has been recorded from a lowland area. Key to the males of Skutzia Reiss 1. Median volsella long, base.24 lm long Median volsella short, base,17 lm long Gonostylus with strong, blunt apical tooth; superior volsella with two pairs of setae medially, lacking microtrichia; base of median volsella ;25 lm long; AR Brazil (fig. 3 in Reiss 1985)... Skutzia bahiensis Reiss 2 0. Gonostylus with small, pointed apical thorn, apparently bearing a long, thin seta; superior volsella with row of 3 4 setae medially, with microtrichia on elevated ridge in basal 2 3; base of median volsella.30 lm; AR Trinidad and Tobago, Venezuela (figs 1 10 in Andersen 2000)... Skutzia gaianii Andersen 3. Superior volsella narrow apically; anal crest conspicuous, nearly V-shaped (Fig. 2D) Superior volsella wider apically; anal crest weak to absent (Figs 1D, 3D) Inferior volsella parallel sided, weakly curved; AR Canada (figs 1 2 in Reiss 1985)...Skutzia inopinata Reiss 4 0. Inferior volsella narrow medially, strongly curved; AR Thailand (Fig. 2A F)... Skutzia inthanonensis sp. n. 5. Anal point lm long, widest medially. Wing ;1.20 mm long; gonostylus lm long. USA (Fig. 1A F)... Skutzia epleri sp. n Anal point lm long, tapering from base. Wing mm long; gonostylus lm long. Panama, Mexico (Fig. 3A F)......Skutzia quetzali sp. n. Skutzia bahiensis Reiss Skutzia bahiensis Reiss, 1985: 175; Spies and Reiss (1996). Material examined. BRAZIL: Bahia, Itabuna, 1970, 10 male paratypes, JA Winder leg (ZSM). Diagnostic characters. The species is similar to S. gaianii Andersen in having a comparatively long median volsella. The 2 species can be separated because S. bahiensis has a strong, blunt tooth apically on gonostylus, whereas S. gaianii has a small, pointed thorn bearing a long, thin seta. The superior volsella has 2 pairs of setae medially and is lacking microtrichia, while S. gaianii has an elevated ridge with microtrichia and row of 3 4 setae in basal 2 3. The AR is , whereas S. gaianii has an AR of The species is described in detail by Reiss (1985). = =

4 2009] AREVIEW OF SKUTZIA REISS 199 FIG. 1. Skutzia epleri sp. n., male. A. Tentorium, stipes, and cibarial pump. B. Thorax. C. Wing. D. Anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus. E. Hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect. F. Median volsella.

5 200 L. C. PINHO ET AL. [Volume 28 FIG. 2. Skutzia inthanonensis sp. n., male. A. Tentorium, stipes, and cibarial pump. B. Thorax. C. Wing. D. Anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus. E. Hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect. F. Median volsella.

6 2009] AREVIEW OF SKUTZIA REISS 201 FIG. 3. Skutzia quetzali sp. n., male. A. Tentorium, stipes, and cibarial pump. B. Thorax. C. Wing. D. Anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus. E. Hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect. F. Median volsella.

7 202 L. C. PINHO ET AL. [Volume 28 TABLE 2. Lengths (in lm) and ratios of legs of Skutzia epleri sp. n., male (n ¼ 2). Abbreviations: fe ¼ femur, ti ¼ tibia, ta ¼ tarsus; for ratios see Table 1. Segment Ratio Leg fe ti ta 1 ta 2 ta 3 ta 4 ta 5 LR BV SV Foreleg Midleg Hind leg Distribution The species is known only from the type locality in Bahia State, Brazil, where the specimens were collected during a cocoa pollination project at the Centre for Cocoa Research (CEPLAC) in Itabuna (Reiss 1985). Itabuna is situated in the lowlands (,200 m asl) close to the coast. Skutzia epleri sp. n. Fig. 1A F Skutzia sp. 1 Epler, Type material. Holotype male, USA: North Carolina, Haywood County, Blue Ridge Parkway, Soco gap (MP 455.7), N, W, 1311 m asl, 06 June 1991, boggy shrub and wetland seep, CR Parker leg (NMNH). Paratype: 1 male, same data as holotype except 13 June 1991 (ZMBN). Diagnostic characters. The species is similar to S. quetzali sp. n. in having short median volsellae, superior volsella comparatively wide apically, and anal crest weak or absent. It can be separated from S. quetzali on its larger size, with a wing length of ;1.20 mm and a gonostylus length of lm, compared to a wing length of mm and a gonostylus length of lmins. quetzali. Furthermore, it has an anal point lm long that is widest medially, whereas S. quetzali has an lm long anal point that is tapering from base. The AR is , whereas S. quetzali has an AR of Etymology. Named after Dr John H. Epler, who made the specimens available for us and for his contribution to chironomid research. Male (n ¼ 2, except when otherwise stated). Total length mm. Wing length mm. Total length/wing length Wing length/length of profemur Color. Cleared specimens with head, abdomen, and legs pale; thorax pale brown. Head. AR , ultimate flagellomere lm long. Temporals 6 (1). Clypeus with setae. Cibarial pump, tentorium, and stipes as in Fig. 1A; tentorium lm long, lm wide; stipes 86 (1) lm long. Palp segment lengths (in lm): 22 24, 32 39, 93 99, 86 91, 150 (1). Third palpomere with 5 7 sensilla clavata subapically, longest lm. Thorax (Fig. 1B). Dorsocentrals 6 7, acrostichals 12 14, prealar 1. Scutellum with 4 setae. Wing (Fig. 1C). VR Brachiolum with 1 seta, R with 18 24, R 1 with 12 18, R 4þ5 with 22 27, Sc, R 2þ3, M, and RM bare, M 1þ2 with 40 43, M 3þ4 with 19 46, Cu with 9 11, Cu 1 with 10 12, PCu with 20 22, and An with setae. Cells r 4þ5 with ; setae; m proximally of RM with 12 18, including 8 10 on false vein; m 1þ2 with ; , including on false vein; m 3þ4 with ; cu and an together with setae. Legs. Spur of foretibia lm long, combs of midtibia lm long, combs of hind tibia lm long. Width at apex of foretibia lm, of midtibia lm, of hind tibia lm. Lengths (in lm) and ratios of legs as in Table 2. Hypopygium (Fig. 1D F). Tergite IX with median and apical setae of which 3 4 below anal point. Anal point lm long, lm wide at base. Crest very weak, nearly V-shaped; tergal bands distinct. Laterosternite IX with 1 2 setae. Phallapodeme lm long. Transverse sternapodeme lm long. Gonocoxite lm long. Superior volsella lm long,with1strongbasal, 4 moderately strong median, and 3 strong apical setae. Inferior volsella lm long, with microtrichia and strong subapical to apical setae. Median volsella (Fig. 1F) with 15- to 16-lm-long base, and 23- to 25-lm-long, slender, pointed apical lamellae. Gonostylus lm long. HR , HV Distribution and habitat This species is known only from the type locality in North Carolina, USA. The 2 specimens were collected in an area with boggy shrub land and wetland seeps at 1300 m asl on the Blue Ridge Highway connecting

8 2009] AREVIEW OF SKUTZIA REISS 203 TABLE 3. Lengths (in lm) and ratios of legs of Skutzia inthanonensis sp. n., male (n ¼ 5 7, except when otherwise stated). Abbreviations: fe ¼ femur, ti ¼ tibia, ta ¼ tarsus; for ratios see Table 1. indicates missing data caused by loss of tarsi when slidemounts were made. Segment Ratio Leg fe ti ta 1 ta 2 ta 3 ta 4 ta 5 LR BV SV Foreleg , , (1) Midleg , , (1) 101 (1) 83 (1) 55 (1) 27 (1) Hind leg , , (1) 203 (1) 166 (1) 111 (1) 55 (1) Shenandoah National Park and Great Smoky Mountains National Park. Skutzia gaianii Andersen Skutzia gaianii Andersen, 2000: 121. Material examined. VENEZUELA: Aragua, Parque Nacional Henri Pittier, hairpin bend ( curva regresiva ) on road to Choroni, 1200 m asl, 23 September 1999, male holotype, at light, T Andersen leg (MIZA); same as previous, 2 male paratypes (ZMBN). TRINI- DAD AND TOBAGO: Trinidad Island, Northern Range, Aripo Main Cave, 24 February 1991, 2 males, J Darlington leg (ZSM). Diagnostic characters. This species is similar to S. bahiensis Reiss in having a comparatively long median volsella. The 2 species can be separated because S. gaianii has a small, pointed thorn bearing a long, thin seta apically on gonostylus, whereas S. bahiensis has a strong, blunt tooth. The superior volsella has an elevated ridge with microtrichia and row of 3 4 setae in basal 2 3 whereas S. bahiensis has 2 pairs of setae medially and lacks microtrichia. The AR is , whereas S. bahiensis has an AR of = The species is described in detail by Andersen (2000). Distribution and habitat This species was originally described from Henri Pittier National Park in northern Venezuela (Andersen 2000). The present record from Trinidad and Tobago expands the distribution to the Caribbean. The specimens from Venezuela were collected at a light close to a small, fast-flowing river at 1200 m asl on the southern slopes of the coastal mountains near Maracay. The river has sandy and silty substratum with stones and boulders. The specimens from Trinidad were collected in the Northern Range, close to El Cerro del Aripo, the highest mountain in Trinidad, with an altitude of 940 m asl. Skutzia inopinata Reiss Skutzia inopinata Reiss, 1985: 174; Oliver et al. (1990). Material examined. CANADA: British Columbia, Vancouver Island, Lake Cowichan, Skutz Falls, 23 July 1979, 2 male paratypes, sweepnet, IM Smith leg (ZSM). Diagnostic characters. The species is similar to S. inthanonensis sp. n. in having short median volsellae, superior volsella comparatively narrow in apical 1 3, and anal crest V-shaped and conspicuous. It can be separated from S. inthanonensis in having a weakly curved, digitiform inferior volsella, whereas S. inthanonensis has a more strongly curved inferior volsella that is narrow medially. The AR is , whereas S. inthanonensis has an AR of The species is described in detail by Reiss (1985). Distribution and habitat This species is endemic to British Columbia (Scudder 1996). According to Reiss (1985), the type specimens were netted in the vegetation along Skutz Creek, a small, cold tributary to Cowichan River. Skutzia inthanonensis sp. n. Fig. 2A F Type material. Holotype male, THAILAND: Chiang Mai Province, Doi Inthanon National Park, 26 June 03 July 1990, H Malicky leg (ZSM). Paratypes: 6 males, same as holotype (ZSM and ZMBN). Diagnostic characters. The species is similar to S. inopinata Reiss in having short median volsellae, superior volsella comparatively narrow in apical 1 3, and anal crest V-shaped and conspicuous. It can be separated from S. inopinata in having a curved inferior volsella that is narrow medially, whereas S. inopinata has a weakly curved, digitiform inferior volsella. The AR is , whereas S. inopinata has an AR of / /

9 204 L. C. PINHO ET AL. [Volume 28 TABLE 4. Lengths (in lm) and ratios of legs of Skutzia quetzali sp. n., male (n ¼ 2 3, except when otherwise stated). Abbreviations: fe ¼ femur, ti ¼ tibia, ta ¼ tarsus; for ratios see Table 1. indicates missing data caused by loss of tarsi when slidemounts were made. Segment Ratio Leg fe ti ta 1 ta 2 ta 3 ta 4 ta 5 LR BV SV Foreleg Midleg (1) 83 (1) 55 (1) 37 (1) (1) 5.55 (1) 3.36 (1) Hind leg (1) 157 (1) 129 (1) 83 (1) (1) 2.89 (1) 2.82 (1) Etymology. Named after the type locality, Doi Inthanon National Park in northern Thailand. Male (n ¼ 5 7, except when otherwise stated). Total length , 1.84 mm. Wing length , 1.16 mm. Total length/wing length , Wing length/length of profemur (3). Color. Cleared specimens with head, abdomen, and legs pale; thorax pale brown. Head. AR , 0.67; ultimate flagellomere , 243 lm long. Temporals 7 (1). Clypeus with 11 17, 14 setae. Cibarial pump, tentorium, and stipes as in Fig. 2A; tentorium , 85 lm long, 16 26, 18 lm wide; stipes 91 (1) lm long. Palp segment lengths (in lm): 18 27, 23; 20 30, 25; 52 75, 65; 89 95, 93 (4); (3). Third palpomere with 3 4, 3 sensilla clavata subapically; longest 11 16, 14 (4) lm long. Thorax (Fig. 2B). Dorsocentrals 6 8, 8; acrostichals 16 18, 17, biserial; prealar 1. Scutellum with 2 4, 4 setae. Wing (Fig. 2C). VR , Brachiolum with 1 seta; R with 19 22, 20; R 1 with 13 14, 14; R 4þ5 with 20 46, 32; Sc with 0 11, 5; R 2þ3, M, and RM bare; M 1þ2 with 31 38, 34; M 3þ4 with 14 20, 18; Cu with 8 10, 9; Cu 1 with 11 14, 12; PCu with 24 36, 30; and An with 12 26, 18 setae. Cells r 4þ5 with ; , 240 setae; m proximally of RM with 15 39, 24, including 14 29, 19 on false vein; m 1þ2 with ; , 280, including 47 63, 55 on false vein; m 3þ4 with ; , 120; and cu and an together with ; , 125 setae. Legs. Spur of foretibia 7 14, 9 lm long; combs of midtibia 7 11, 10 lm long; combs of hind tibia 11 14, 12 lm long. Width at apex of foretibia 29 36, 31 lm; of midtibia 25 32, 30 lm; of hind tibia 32 36, 34 lm. Lengths (in lm) and ratios of legs as in Table 3. Hypopygium (Fig. 2D F). Tergite IX with 9 13, 10 median and 12 17, 14 apical setae of which 3 4, 3 below anal point. Anal point 18 25, 22 lm long, 11 16, 14 lm wide at base. Crest short, nearly V-shaped; tergal bands distinct. Laterosternite IX with 1 2, 2 setae. Phallapodeme 66 75, 72 lm long. Transverse sternapodeme 20 27, 26 lm long. Gonocoxite 75 82, 79 lm long. Superior volsella 30 39, 37 lm long, with 1 strong basal, 3 4, 4 moderately strong median, and 2 strong apical setae. Inferior volsella 45 54, 47 lm long, with microtrichia and 10 13, 11 strong subapical to apical setae. Median volsella (Fig. 2F) with 9 14, 10 lm long base and 23 27, 25 lm long, slender, pointed, apical lamellae. Gonostylus 37 42, 41 lm long. HR , 1.15; HV , Distribution and habitat This species is recorded only from Doi Inthanon in northern Thailand. Doi Inthanon is the highest mountain in Thailand (2565 m asl). At altitudes.1000 m, annual rainfall is.2500 mm, and at the summit, the temperature can drop as low as 88C. At high altitudes, dominant plant species belong to families found in areas with a temperate rather than a tropical climate, and several species are considered to be relicts from periods with a cooler climate. Skutzia quetzali sp. n. Fig. 3A F Type material. Holotype male, MEXICO: Campeche State, Calakmul, Ejido Nuevo Becan, El Chorro, N, W, 130 m asl, 30 April 1997, light trap, A Contreras-Ramos et al. leg (ZMBN). Paratypes: 1 male, MEXICO: Campeche, Calakmul, 1 km NE Hormiguero, Aguada Pequeña, N, W, 300 m asl, May 1997, Malaise trap, A Contreras-Ramos et al. leg (ZMBN). 1 male, PANAMA: 09 August 1983, W Nentwig leg (ZSM). 1 male, same data as previous, except 27 December 1983 (ZSM). Diagnostic characters. This species is similar to S. epleri sp. n. in having short median volsellae, superior volsella comparatively wide apically, and anal crest weak or absent. It can be separated from S. epleri on its smaller size, with a wing length of mm and a gonostylus length of lm, compared to a wing length of ;1.20 mm and a gonostylus length of lm ins. epleri. Furthermore, it has an anal point 11 18

10 2009] AREVIEW OF SKUTZIA REISS 205 lm long that is tapering from base, whereas S. epleri has an anal point lm long that is widest medially. AR is , whereas S. epleri has an AR of Etymology. Named after the bird Resplendent Quetzal (Pharomachrus mocinno de la Llave), which is also distributed from southern Mexico to western Panama. The name is a noun in apposition. Male (n ¼ 3, except when otherwise stated). Total length mm. Wing length lm. Total length/wing length Wing length/length of profemur Color. Cleared specimens with head, abdomen, and legs pale; thorax pale brown. Head. AR , ultimate flagellomere lm long. Temporals 7 8 (2), including 2 inner verticals, 2 3 outer verticals, and 2 4 postorbitals. Clypeus with setae. Cibarial pump, tentorium, and stipes as in Fig. 3A; tentorium lm long, lm wide; stipes 86 (1) lm long. Palp segment lengths (in lm): 18 19, 22 25, 50 59, 64 77, (2). Third palpomere with 3 sensilla clavata subapically, longest lm. Thorax (Fig. 3B). Dorsocentrals 6 9, acrostichals 14 20, prealar 1. Scutellum with 4 setae. Wing (Fig. 3C). VR Brachiolum with 1 seta, R with 14 25, R 1 with 7 14; R 4þ5 with 22 41, Sc with 8 16, R 2þ3, M, and RM bare, M 1þ2 with 21 34, M 3þ4 with 13 46, Cu with 8 15, Cu 1 with 10 12, PCu with 24 33, and An with setae. Cells r 4þ5 with ; setae; m proximally of RM with 24 36, including on false vein; m 1þ2 with ; , including on false vein; m 3þ4 with ; ; and cu and an together with ; setae. Legs. Spur of foretibia 7 8 lm long, combs of midtibia 7 11 lm long, combs of hind tibia 7 11 lm long. Width at apex of foretibia lm, of midtibia lm, of hind tibia lm. Lengths (in lm) and ratios of legs as in Table 4. Hypopygium (Fig 3D F). Tergite IX with 9 12 median and 8 12 apical setae of which 2 4 below anal point. Anal point lm long, lm wide at base. Crest apparently absent; tergal bands distinct. Laterosternite IX with 2 3 setae. Phallapodeme lm long. Transverse sternapodeme lm long. Gonocoxite lm long. Superior volsella lm long, with 1 strong basal, 4 moderately strong median, and 3 strong apical setae. Inferior volsella lm long, with microtrichia and 7 10 strong subapical to apical setae. Median volsella (Fig. 3F) with 9- to 11-lmlong base and 23- to 25-lm-long, slender, pointed apical lamellae. Gonostylus lm long. HR , HV Distribution and habitat This species is recorded from Panama and from the Yucatan Peninsula in southeastern Mexico. The specimens from Mexico were taken in Calakmul Biosphere Reserve, an area with wet tropical lowland forest. At the sampling sites there were small lakes (Hormiguero) and a small, clear-water permanent stream (El Chorro). Acknowledgements We are indebted to Marion Kotrba, Zoologische Staatssammlung, Munich, for the loan of the type material of S. bahiensis and S. inopinata and the specimens from Thailand, Panama, and Trinidad and Tobago; to John H. Epler for providing the material from the Smoky Mountains, USA; and to Dr Atilano Contreras-Ramos for the material from Mexico. LCdP received funding from Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) (05/ ), within the Biota/FAPESP The Biodiversity Virtual Institute Program ( while completing this paper. Literature Cited ANDERSEN, T A new species of Skutzia Reiss, 1985 (Chironomidae: Chironominae: Tanytarsini) from Henri Pittier National Park, Venezuela. Boletín de Entomología Venezolana 15: BAUSE, E Die Metamorphose der Gattung Tanytarsus und einige verwandter Tendipedidenarten. Ein Beitrag zur Systematik der Tendipediden. Archiv für Hydrobiologie und Planktonkunde, Supplement 2: BRUNDIN, L Zur Kenntnis der schwedischen Chironomiden. Arkiv för Zoologi 39A:1 95. CRANSTON, P. S., M. E. DILLON, L.C.V.PINDER, AND F. REISS The adult males of Chironominae (Diptera: Chironomidae) of the Holarctic region keys and diagnoses. Pages in T. Wiederholm (editor). Chironomidae of the Holarctic region. Keys and diagnoses. Part 3. Adult males. Entomologica Scandinavica, Supplement 11. EDWARDS, F. W On marine Chironomidae (Diptera); with descriptions of a new genus and four new species from Samoa. Proceedings of the Zoological Society of London 51: EKREM, T A taxonomic revision of the genus Stempellinella (Diptera: Chironomidae). Journal of Natural History 41: EKREM, T.,AND O. A. SÆTHER Seppia, a new Afrotropical tanytarsine genus (Chironomidae: Chironominae). Pages in O. Hoffrichter (editor). Late 20 th century

11 206 L. C. PINHO ET AL. [Volume 28 research on Chironomidae: an anthology from the 13 th International Symposium on Chironomidae. Shaker Verlag, Aachen, Germany. EPLER, J. H Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. A guide to the taxonomy of the midges of the southeastern United States, including Florida. Special Publication SJ2001-SP13. North Carolina Department of Environment and Natural Resources, Raleigh, North Carolina, and St. Johns River Water Management District, Palatka, Florida. (Available from: esb.enr.state.nc.us/bauwww/chironomid.htm) HARRISON, A. D A contribution to the taxonomy of Tanytarsini (Diptera: Chironomidae) of sub-saharan Africa, with a description of a new genus (Afrozavrelia) and five new species from other genera. Annals of the Eastern Cape Museums 3(2002):1 15. KIEFFER, J. J Un nouveau genre de Chironomidae (Dipt.). Bulletin de la Société Entomologique de France 1919: KIEFFER, J. J Chironomides nouveaux ou peu connus de la région paléarctique. Bulletin de la Société d histoire naturelle de Metz 29: MENDES, H. F., T. ANDERSEN, AND O. A. SÆTHER A review of Antillocladius Sæther, 1981; Compterosmittia Sæther, 1981 and Litocladius new genus (Chironomidae, Orthocladiinae). Zootaxa 594:1 82. OLIVER, D. R., M. E. DILLON, AND P. S. CRANSTON A catalog of Nearctic Chironomidae. Research Branch, Agriculture Canada Publication 1857/B. Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Canada. PINDER, L. C. V The adult males of Chironomidae (Diptera) of the Holarctic region introduction. Pages 5 9 in T. Wiederholm (editor). Chironomidae of the Holarctic region. Keys and diagnoses. Part 3. Adult males. Entomologica Scandinavica, Supplement 34. REISS, F Neostempellina thienemanni n. gen. n. sp., eine europäische Chironomide mit gehäusetragenden Larven (Diptera, Insecta). Spixiana 7: REISS, F Die panamerikanisch verbreitete Tanytarsini Gattung Skutzia gen. nov. Spixiana 11: SÆTHER, O. A Glossary of chironomid morphology terminology (Diptera: Chironomidae). Entomologica Scandinavica, Supplement 14:1 51. SÆTHER, O. A Orthocladiinae (Diptera: Chironomidae) from the British West Indies, with descriptions of Antillocladius n. gen., Lipurometriocnemus n. gen., Compterosmittia n. gen. and Diplosmittia n. gen. Entomologica Scandinavica, Supplement 16:1 46. SÆTHER, O. A Zoogeographical patterns in Chironomidae (Diptera). Verhandlungen der Internationalen Vereinigung für theoretische und angewandte Limnologie 27: SÆTHER, O. A., AND T. ANDERSEN Friederia, a new Afrotropical tanytarsine genus (Diptera: Chironomidae). Entomologica Scandinavica 29: SÆTHER, O. A., AND T. EKREM Biogeography of Afrotropical Chironomidae (Diptera), with special reference to Gondwanaland. Cimbebasia 19: SÆTHER, O. A., AND F. O. ROQUE New Neotropical species of Nandeva (Diptera: Chironomidae), with a phylogeny of the Tanytarsini. Tijdschrift voor Entomologie 147: SCUDDER, G. G. E Terrestrial and freshwater invertebrates of British Columbia: priorities for inventory and descriptive research. Research Branch, British Columbia Ministry of Forests, and Wildlife Branch, British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia, Working Paper 9: SPIES, M., AND F. REISS Catalog and bibliography of Neotropical and Mexican Chironomidae (Insecta, Diptera). Spixiana, Supplement 22: WIEDENBRUG, S., F. REISS, AND E. J. FITTKAU Nandeva, gen. nov., a new genus of Chironomini (Insecta, Diptera, Chironomidae). Spixiana 21: Received: 24 April 2008 Accepted: 3 November 2008