Entomology Publications Entomology

2008 A New Species of Blepharicera Macquart (Diptera: ) from Western North America Amanda J. Jacobson Iowa State University

Gregory W. Courtney Iowa State University, [email protected]

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This Article is brought to you for free and open access by the Entomology at Iowa State University Digital Repository. It has been accepted for inclusion in Entomology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. A New Species of Blepharicera Macquart (Diptera: Blephariceridae) from Western North America

Abstract During a review of the Blepharicera of western North America, we discovered a new species from several mid- sized rivers in southwestern Oregon and northwestern California. We hereby present descriptions of the larvae, pupae, and adults of B. kalmiopsis, new species. Diagnostic characters and a brief discussion of bionomics and distribution are also provided. Based on previous and ongoing studies, B. kalmiopsis clearly belongs to the B. micheneri Alexander species group and appears closely related to B. zionensis Alexander.

Keywords Blepharicera, Blephariceridae, net-winged , new species, Nearctic

Disciplines Biology | Entomology

Comments This article is from Proceedings of the Entomological Society of Washington 110 (2008): 978, doi: 10.4289/ 0013-8797-110.4.978. Posted with permission.

This article is available at Iowa State University Digital Repository: https://lib.dr.iastate.edu/ent_pubs/190 A New Species of Blepharicera Macquart (Diptera: Blephariceridae) from Western North America Author(s): Amanda J. Jacobson and Gregory W. Courtney Source: Proceedings of the Entomological Society of Washington, 110(4):978-987. 2008. Published By: Entomological Society of Washington DOI: http://dx.doi.org/10.4289/0013-8797-110.4.978 URL: http://www.bioone.org/doi/full/10.4289/0013-8797-110.4.978

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PROC. ENTOMOL. SOC. WASH. 110(4), 2008, pp. 978–987

A NEW SPECIES OF BLEPHARICERA MACQUART (DIPTERA: BLEPHARICERIDAE) FROM WESTERN NORTH AMERICA

AMANDA J. JACOBSON AND GREGORY W. COURTNEY

Department of Entomology, Iowa State University, 3222 Science II Bldg., Ames, IA 50011-3222, U.S.A. (e-mail: [email protected]); AJJ current address: Department of Entomology and Plant Pathology, University of Tennessee, 2431 Joe Johnson Dr., Suite 205, Knoxville, TN 37996-4560. E-mail: [email protected]

Abstract.—During a review of the Blepharicera of western North America, we discovered a new species from several mid-sized rivers in southwestern Oregon and northwestern California. We hereby present descriptions of the larvae, pupae, and adults of B. kalmiopsis, new species. Diagnostic characters and a brief discussion of bionomics and distribution are also provided. Based on previous and ongoing studies, B. kalmiopsis clearly belongs to the B. micheneri Alexander species group and appears closely related to B. zionensis Alexander. Key Words: Blepharicera, Blephariceridae, net-winged midges, new species, Nearctic

The Blephariceridae (net-winged first ventral sucker) (Courtney 2000a). midges) are a small dipteran family of The suckers are important for locomo- approximately 320 described species in tion and provide a strong hold to rocks, 28 described genera; however, these bedrock, or other smooth surfaces (Fru- can be an important component of tiger 1998, 2002). Larvae are grazers stream ecosystems. These flies are usual- whose main diet is diatoms (Alverson et ly restricted to cool, clear, well-oxygen- al. 2001, Alverson and Courtney 2002). ated mountain streams, and many spe- Like larvae, blepharicerid pupae are cies are known to be sensitive highly adapted to fast-flowing streams, bioindicators (Lenat 1993, Courtney et being streamlined, somewhat dorsoven- al. 2007). Furthermore, with densities in trally flattened, and possessing 3–4 pairs some streams exceeding 1,000/m2 (Johns of ventrolateral adhesive discs that are 1996, Courtney unpublished) and bio- used to adhere the pupa to the substrata. mass at least temporally high in other Adult blepharicerids are delicate, long- streams (Anderson 1992), blepharicerids legged flies that superficially resemble can have a significant trophic impact. crane flies. Adults usually remain close Blepharicerids are noteworthy in their to the natal stream, where they often rest adaptations to life in and around water- on the underside of riparian vegetation, falls and torrential streams. Perhaps the overhanging logs, or shaded rocks most striking adaptations are in the (Courtney 2000b). larvae, which possess six ventral suckers Four genera of Blephariceridae occur and a specialized cephalic division (com- in the Nearctic Region (Hogue 1987, prising the fused head, thorax, and first Courtney 2000a). von Ro¨der is abdominal segment, and including the found in western North America and VOLUME 110, NUMBER 4 979 central and eastern Asia, comparative material originated from Osten Sacken in western North America the collections of C. L. Hogue, C. P. and Japan, Philorus Kellogg in western Alexander and the junior author. Exam- North America and central and eastern ined specimens were on loan from or are Asia, and Blepharicera Macquart in deposited with the following institutions western and eastern North America and (acronyms used throughout the text): throughout Eurasia. The latter genus ISIC, Iowa State Collection, Iowa currently contains four described species State University, Ames, Iowa; LACM, (B. jordani Kellogg, B. micheneri Alex- Natural History Museum of Los An- ander and B. zionensis Alexander [5 B. geles County, Los Angeles, California; micheneri species group] and B. osten- USNM, National Museum of Natural sackeni Kellogg) in western North Amer- History, Smithsonian Institution, Wa- ica (Hogue 1987). The distributions of shington, D.C. these species vary considerably but gen- Specimen preparation.—Most speci- erally are limited to mountainous areas mens were collected and preserved in from southwestern Canada to southern 70% or 95% ethyl alcohol (EtOH). California and Arizona (Hogue 1987, Morphological studies were based on Courtney unpublished). whole- preparations, pupal dissec- We hereby add a fifth species of tions, slide mounts, and scanning elec- Blepharicera to the fauna of western tron microscopy (SEM). Slide-mounted North America. Descriptions are provid- material was cleared in cedarwood oil ed for the larvae, pupae, and adults of B. and mounted in Canada balsam, follow- kalmiopsis new species. Diagnostic char- ing procedures described elsewhere acters and brief discussions of bionom- (Courtney 1990). Additional material ics, distribution, and taxonomic affinities was prepared by removing soft tissues are also provided. with dilute (approximately 10%) potas- The authors would like to dedicate this sium hydroxide (KOH). Specimens were paper to Jack Lattin, emeritus professor examined using an Olympus SZX-12 at Oregon State University. Jack’s en- dissecting microscope and a Nikon E- thusiasm, instruction, and mentorship 800 compound microscope, and draw- during GWC’s undergraduate years pro- ings were rendered with the aid of a vided not only a sound introduction to drawing tube on both systems. Material systematic entomology but the incentive for SEM examination was sonicated to consider a career in this field. It is briefly (5–10 s) in EtOH or an EtOH- perhaps befitting that the senior author KOH mixture and prepared by critical of this paper was under GWC’s mentor- point drying and gold-palladium coating ship as an undergraduate a few short in a sputter coater. Material was exam- years ago, ‘‘saw the light’’ of systematic ined with a JEOL JSM-5800LV SEM, entomology, and now is pursuing a which captured direct digital images. career in this field. Things have indeed Terminology.—Terms for structures come full circle, especially given that are based primarily on Courtney (2000a). Jack received his undergraduate degree Descriptive format.—Measurements at Iowa State University! are given in millimeters, as a mean followed by a range in parentheses. METHODS Descriptive format is based on, and Material.—Although some collections values are recorded according to, proce- of the new species date back to the mid- dures outlined in Courtney (2000b). 1980’s, most material was obtained more Abbreviations for label and locality data recently (e.g., 1999, 2007). Additional are Ck 5 Creek; Co 5 County; coll. 5 980 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON collected by; Fk 5 Fork; Hwy 5 Cranial sclerites yellow with dark brown Highway; M 5 Middle; mi 5 mile(s); outline or yellow frontoclypeal apotome Mtn 5 mountain; nr 5 near; S 5 South; with brown genae; ecdysial line with stem $ 5 Dollar. Abbreviations for life stages line, posterior margin of frontoclypeal are L 5 larvae; P 5 pupae; Pex 5 pupal apotome not extended to posterior cra- exuviae; A 5 adult. nial margin. Cephalic division, abdomen, and lateral lobes of uniform coloration, Blepharicera kalmiopsis Jacobson and light brown; color variation includes Courtney, new species lateral lobes with darker pigmentation. (Figs. 1–16) Lateral lobes with proleg and two addi- tional appendages; anterolateral lobes Diagnosis.—A medium-sized Blephar- larger than posterolateral lobes. Antero- icera. : Anterolateral lobes on lateral lobes on cephalic division minute, cephalic division minute, much smaller smaller than lobes on abdominal seg- than lobes on abdominal segments II– ments II–VI. Anal division rounded, with VI; lateral lobes with setose anterolateral round lateral lobes. Chaetotaxy: Cranial and posterolateral appendages; sensilla sclerites, cephalic division, and abdomi- absent on ventral portion of lateral nal segments with fusiform sensilla; nu- lobes. Pupa: Body outline ovoid; bran- merous setiform sensilla along frontal chial sclerite glabrous; apices of respira- margin of cephalic division; substernal tory lamellae separated medially. Adult sensilla digitiform, pale, < 40 in number; male: Eye division contiguous; ultimate dorsal secondary sensilla fusiform, antennal flagellomere 3.23 length of opaque, pale in color, largely confined penultimate; cerci triangular, with dor- to apex of lateral lobes, prolegs, and solateral ridge extending obliquely to anterolateral and posterolateral append- outer margin, dividing cerci into basal ages; ventral side of lateral lobes without two-thirds and distal third; gonostylus setae or sensilla; anal division with 8–10 truncate with two lobes; aedeagal rods prominent setiform sensilla marginally. equal in length; ventral parameres thick- Pupa (Figs. 6–8, 9, 10): Measure- ened, horseshoe-shaped, slightly longer ments, male (N 5 18) length 5.52 mm than aedeagal rods. Adult female: Eyes (5.00–5.80), width 3.39 mm (3.10–3.60); divided; callis oculi present but very female (N 5 20) length 6.50 mm (5.60– small; sternite VIII slightly bilobate, 7.40), width 3.81 mm (3.20–4.00). Body medial depression shallow; genital fork outline ovoid. Integument: Dorsal papil- broadly Y-shaped; hypogynial valves lae uniformly distributed on abdominal elongate, parallel, and rounded apically, segments; metatergite with papillae pre- inner margin slightly concave and outer sent medially, absent on lateral surface margin slightly convex. beyond abdominal segment I. Papillae Description.—Larva (Figs. 1–5): Mea- dark brown with small, well-developed surements, instar III (N 5 20) total spinules. Spinules directed posteriorly, length 5.09 mm (3.94–6.30), cranial cone-shaped, of variable size, uniformly width 0.46 mm (0.43–0.49), antennal distributed on papillae. Cuticle between segments 0.15 mm (0.09–0.18), 0.13 mm papillae glabrous and dark brown. Bran- (0.11–0.16), membrane 0.04 mm (0.03– chial sclerite without papillae. Alar and 0.05); instar IV (N 5 20) total length cephalic sclerites glabrous. Anal tergite 7.88 mm (5.30–9.30), cranial width wrinkled. Respiratory lamellae wider at 0.70 mm (0.66–0.75), antennal segments base and rounded apically; inner and 0.28 mm (0.18–0.33), 0.18 mm (0.13– outer margins curving medially; apices 0.22), membrane 0.05 mm (0.03–0.13). separated medially. VOLUME 110, NUMBER 4 981

Figs. 1–8. Scanning electron micrographs of larval and pupal Blepharicera kalmiopsis.1,Cephalic division, dorsal view. 2, Cephalic division, ventral view. 3, Proleg on abdominal segment III, dorsal view on right side. 4, Proleg on abdominal segment III, ventral view on left side.5,Analdivision,abdominal segments VI–X, dorsal view. 6–8, Pupal abdominal tergite microstructure.Scalebars5 1 mm(Fig.8),10 mm (Fig. 7), 100 mm (Fig. 6), 500 mm (Figs. 3–5), 1 mm (Figs. 1, 2). 982 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Adult male: Size: medium. Measure- Forecoxae light brown; other coxae pale. ments (N 5 11): Total length 6.02 mm Abdominal tergites light brown, sternites (5.30–6.60), wing length 6.96 mm (6.00– pale. Specimens from type locality gen- 7.50), width 2.12 mm (1.80–2.40). erally darker than those from other sites Head (Fig. 12): Structure: Eyes semi- which might be an artifact of age and dichoptic, interocular ridge present; in- fixation. terocular distance 0.07 mm; eye divided; Terminalia (Figs. 14–16): Abdominal callis oculi absent; dorsal division con- segment VIII greatly reduced, barely tiguous with ventral, smaller (0.333) visible dorsally. Epandrium simple, than ventral; dorsal ommatidia larger in emarginate posteromedially; setae more diameter; dorsal division with 15 rows of numerous laterally. Cerci triangular, ommatidia along mid-meridian. Clypeus with dorsolateral ridge extending obli- length/width 5 1.7. Proboscis about quely to outer margin, dividing cerci into 0.433 head width; mandibles absent; basal two-thirds and distal third; inter- palpi with 5 palpomeres, distal 4 segment cercal area consisting of a wide U-shaped proportions 1.0, 1.3, 1.2, 3.0. Antennal depression, setiform sensilla along inner flagellomeres barrel-shaped; ultimate fla- margin and apical third of cerci. Genital gellomere 3.23 length of penultimate capsule about as wide as long. Gonosty- flagellomere; scape and pedicel light lus and gonocoxite setose. Gonostylus brown with prominent, dark brown seti- small, truncated, less sclerotized distally form sensilla; flagellomeres setose and with 2 lobes, anterior and posterior. brown (Smith River); light brown (Ump- Gonocoxal lobe heavily sclerotized, dark qua River). Chaetotaxy: Setiform sensilla brown, simple, glabrous, dorsoventrally groups as follows (number per side): flattened, curved medially near apex, clypeals (10–24), parietals (9–18), occip- almost as long as gonostylus. Aedeagal itals (40–50), postgenals (< 22). rods equal in length; medial rod straight, Thorax and Appendages: Structure: lateral rods curve laterally at base and Tibial spurs 0–0–1, length 0.31 (0.28– apex away from medial rod. Ventral 0.34). Leg-segment proportions: foreleg— parameres slightly longer than aedeagal 33:30:18:7:5:3:4, midleg—36:30:14:7:5:4:4, rods, thickened, horseshoe-shaped, al- hindleg—39:35:14:4:3:2:3. Chaetotaxy: Tho- most encircling aedeagal rods. Dorsal rax glabrous except for few sparse setae paramere rectangular at apex. Dorsal and setae on posterior margin of anepis- carina prominent, rounded. Gonocoxal ternum (3–10); scutellum with setae apodeme and lateral parameral lobes grouped at posterolateral corner (N < well developed. Ejaculatory apodeme 13) and along margin; coxae with prom- longer in comparison to lateral param- inent setae. eral lobes. Coloration: Frons, face and clypeus Adult female: Size: medium. Measure- pruinose, light brown. Thorax brown. ments (N 5 10): Total length 7.10 mm

Leg Segment Lengths Foreleg Midleg Hindleg femur 3.72 (3.45–3.90) 3.88 (3.50–4.15) 5.20 (4.60–5.63) tibia 3.37 (3.10–3.60) 3.33 (3.05–3.65) 4.60 (4.35–4.88) tarsus 1 1.91 (1.71–2.10) 1.51 (1.35–1.62) 1.84 (1.70–2.00) 2 0.82 (0.75–0.88) 0.80 (0.75–0.86) 0.58 (0.50–0.64) 3 0.55 (0.50–0.60) 0.56 (0.54–0.62) 0.42 (0.40–0.47) 4 0.37 (0.35–0.39) 0.39 (0.35–0.43) 0.30 (0.27–0.32) 5 0.45 (0.42–0.49) 0.46 (0.42–0.49) 0.41 (0.37–0.47) VOLUME 110, NUMBER 4 983

(6.30–8.30), wing length 8.40 mm (7.70– 9.00), width 2.30 mm (2.10–2.60). Head (Fig. 11): Structure: Eyes sub- holoptic, interocular ridge present, in- terocular distance 0.07 mm; eye divided; callis oculi present, minute, easily over- looked; dorsal division separated from ventral, subequal in size; dorsal division with 18 rows of ommatidia along mid- meridian. Clypeus length/width 5 1.4. Proboscis about 0.583 head width; palpi with 5 palpomeres, distal 4 segment proportions 1.0, 1.4, 1.2, 2.1. Antennal flagellomeres barrel-shaped; ultimate fla- gellomere elongate, 2.33 length of pen- ultimate flagellomere; scape and pedicel brown with prominent, dark brown seti- form sensilla; flagellomeres setose and brown. Chaetotaxy: Setiform groups as follows (number per side): clypeals (< 30), parietals (18–24), occipitals (26–33), postgenals (23–30). Thorax and appendages: Tibial spurs 0–0–2; spurs asymmetrical, one each long (0.35 mm) and short (0.13 mm). Leg-segment proportions: foreleg—36: 30:17:6:4:3:4, midleg—38:32:13:6:4:3:4, hindleg—39:35:14:4:3:2:3. Chaetotaxy: Thorax glabrous except for few sparse setae and setae on posterior margin of anepisternum (< 8); scutellum with setae grouped at posterolateral corner (N < Figs. 9–10. Pupae of Blepharicera kalmiopsis.9, 13) and along margin; coxae with prom- Pupa, dorsal view. 10, Pupa, frontal view. Scale bars inent setae. 5 1 mm. (abbreviations: al sc 5 alar sclerite; atI– Coloration: Same as for male. Parietal atVII 5 abdominal tergites I–VII; atA 5 anal sclerites pruinose. As for male, more tergite; br sc 5 branchial sclerite; mtt 5 metatergite). recently collected specimens are signifi- cantly darker.

Leg Segment Lengths Foreleg Midleg Hindleg femur 4.43 (4.05–4.80) 4.32 (4.00–4.65) 5.90 (5.00–6.40) tibia 3.75 (3.35–4.00) 3.59 (3.20–3.90) 5.22 (4.65–5.60) tarsus 1 2.03 (1.80–2.25) 1.46 (1.25–1.60) 2.12 (1.85–2.35) 2 0.78 (0.70–0.85) 0.72 (0.62–0.80) 0.61 (0.54–0.70) 3 0.51 (0.47–0.55) 0.47 (0.42–0.50) 0.43 (0.35–0.47) 4 0.31 (0.27–0.37) 0.31 (0.30–0.35) 0.32 (0.30–0.35) 5 0.49 (0.44–0.56) 0.48 (0.47–0.51) 0.49 (0.45–0.54) 984 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 11–16. Adults of Blepharicera kalmiopsis. 11, Female head and antennal apex, frontal view. 12, Male head and antennal apex, frontal view. 13, Female terminalia, ventral view. 14, Male terminalia, dorsal view. 15, Male terminalia (phallic structures), lateral view. 16, Male terminalia (phallic structures), dorsal view. Scale bars 5 0.10 mm (Figs. 13–16), 1 mm (Figs. 11, 12). VOLUME 110, NUMBER 4 985

Terminalia (Fig. 13): Sternite VIII M. Fk. Smith River along Hwy 197, slightly bilobate, medial depression shal- 41u489N 124u059W, 10 June 1999 [L]. All low. Sternite IX (genital fork) broadly Y- material collected by G.W. Courtney. shaped, heavily sclerotized at base. Hy- Etymology.—The specific epithet re- pogynial plate broad basally, narrowed fers to the Kalmiopsis Wilderness Area slightly at base of apical valves; individ- in the Klamath Mountains of southwest ual valves elongate and parallel, widest Oregon, from which several tributaries of at midpoint, intervalvular area narrowly the type locality and other streams with U-shaped. Accessory gland not directly B. kalmiopsis flow. The wilderness area visible in prepared specimens. Sper- is, in fact, named after Peter Kalmia, mathecae three in number; corpora Finnish student of Linnaeus. ovoid, slightly longer than wide, with Distribution.—Blepharicera kalmiopsis long necks; ducts long, basal half and is currently known only from three apex darkly pigmented. Chaetotaxy: medium-sized rivers in the Klamath Sternite VIII with 2 setiform sensilla Mountains of southwestern Oregon and laterally; hypogynial plate with numer- northwestern California. ous small setae; epiproct with 2 promi- Bionomics.—Although all known col- nent setiform sensilla apically. lection records of Blepharicera kalmiop- Type material.—Holotype [adult male, sis are from late May to mid June, the reared]: UNITED STATES: OREGON: larvae are presumably active throughout Curry Co: Pistol River nr. Glade Ck., May and possibly earlier. Most June 42u169N 124u199W, 13 June 1999, coll. collections consist of third- and fourth- G.W. Courtney; pinned, genitalia in instar larvae, with one first instar taken glycerin, deposited USNM. Allotype on 9 June 1999 from Pistol River. [adult female, reared]: same data as Additional first- and second-instar larvae holotype, deposited USNM. Paratypes: are present in a sample taken on 10 June same data as holotype: [20L, 10P 1999 from the Middle Fork Smith River; (EtOH), reared A: 1 male (pinned with however, due to sympatry and temporal genitalia in glycerin), 1 male (pinned), 2 overlap of B. kalmiopsis and B. jordani at males (EtOH), 1 female (EtOH)]; 9 June this site and the current difficulties in 1999 [20L, 4P, 1Pex]; 31 May 2007 [3 identifying early instar larvae, it is not males (pinned)]. Paratypes deposited in possible to determine the identity of ISIC, LACM, and USNM. these specimens. Blepharicera kalmiopsis Other material examined.—UNITED is sympatric with other Blepharicera at STATES: OREGON: Curry Co:Pistol all known collection sites. The species is River nr. Glade Ck., 42u169N 124u199W, temporally isolated from B. ostensackeni, 9 June 1999 [LPPex], 13 June 1999 [LP], which typically emerges later, but ap- 31 May 2007 [LPPexA]. Douglas Co:S. pears to overlap considerably with B. Umpqua River, 43u029N122u489W, 25 jordani. Other blepharicerids (e.g., Aga- June 1983 [PA]; S. Umpqua River < 7mi. thon comstocki (Kellogg), Bibiocephala above Tiller, 42u599N122u519W, 31 May grandis Osten Sacken, and/or Philorus 1986 [LP], 11 June 1999 [L]. Josephine Co: californicus Hogue) also occur at some Illinois River @ 8$ Mtn. Road, 42u149N sites but emerge much earlier than either 123u419W, 16 June 1996 [PPex], 10 June B. kalmiopsis or B. jordani. Historical 1999 [LPPex], 30 May 2007 [LPA]. records (Courtney unpublished) of the CALIFORNIA: Del Norte Co: M. Fk. Smith latter two species suggest that popula- River@PantherFlat,41u509N123u559W, tions of one species will predominate 19 June 1984 [A], 16 June 1996 [LPPexA], during a given year (e.g., in 1999, B. 10 June 1999 [L], 30 May 2007 [LPPexA]; jordani was much more abundant at both 986 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Middle Fork Smith River and Illinois known only from the Virgin River, Zion River, whereas the reverse was true in National Parkin southwestern Utah, has 2007). Although based on only two years a more restricted distribution. Although of data, the type locality is the only site future collections are more likely to yield where B. kalmiopsis is consistently the additional records for known species than most abundant species. the discovery of new species, our study Taxonomic comparison.—Although confirms that new taxa may indeed exist the larvae and pupae of Blepharicera in certain areas. Furthermore, it is possi- kalmiopsis superficially resemble Ble- ble that certain widespread species (e.g., pharicera jordani, the following charac- B. jordani) and morphologically variable ters differentiate between the two: (1) B. species (e.g., B. ostensackeni) may repre- jordani has dark brown digitiform sen- sent species complexes, which may be silla on the ventral portion of the particularly good candidates for genetic abdominal lateral lobes (absent in B. and/or phylogeographic studies. kalmiopsis); (2) the anterior appendage In addition, phylogenetic studies are on the cephalic division (first abdominal needed to determine the relationships segment) is larger in B. jordani. The among western Nearctic Blepharicera larvae of other species in the B. micheneri and between eastern and western Nearc- group (B. micheneri and B. zionensis) tic representatives of the genus. A with anterior and posterior appendages preliminary phylogenetic analysis using on the lateral lobes are not sympatric morphological characteristics (Jacobson with B. kalmiopsis. The pupae of B. 2006) confirmed Zwick’s (1984) earlier kalmiopsis and B. jordani are most hypothesis that the western Nearctic readily distinguished by the respiratory fauna comprises two separate lineages, organs (lamellae meet medially in B. B. ostensackeni, which is more closely jordani). Adult males of B. kalmiopsis related to a largely Asian clade, and a are unique in the structure of the male monophyletic and exclusively Nearctic terminalia and can be easily differentiat- B. micheneri species group. Furthermore, ed by the shape of the cerci, gonostylus, the Jacobson study clearly placed B. and ventral parameres. Adult females of kalmiopsis within the B. micheneri species B. kalmiopsis can be distinguished from group, and suggested a close relationship B. jordani by the size of the callis oculi between B. kalmiopsis and B. zionensis. (small and easily overlooked in B. Details of this analysis are the focus of a kalmiopsis) and the shape of the antennal forthcoming paper. flagellomeres (barrel-shaped in B. kal- The current paper provides the first miopsis, elongate in B. jordani). report of a new species of western Nearctic blepharicerid in nearly 40 years DISCUSSION (Hogue 1970) and the area’s first new Blepharicera since 1959 (Alexander The blepharicerids of western North 1959)! Despite the lengthy gap between America have been intensively surveyed species descriptions and the general per- by Hogue (1966, 1973, 1987) and Court- ception that North American blephari- ney (1991 and unpublished data, 1982– cerids are relatively well known, it is clear present). As a result, it is apparent that that much new information on these B. kalmiopsis has one of the most limited interesting flies awaits discovery. Future distributions among western Blephari- areas of study should emphasize collec- cera, being confined to a few river systems tion efforts throughout the year, careful in southwestern Oregon and northwest- examination of all life stages (including ern California. Only B. zionensis, which is definitive association of life stages!), and VOLUME 110, NUMBER 4 987 comparative investigations using both ———. 2000a. A.1. Family Blephariceridae, morphological and genetic characters. pp. 7–30. In Papp, L. and B. Darvas, eds. Contri- butions to a Manual of Palaearctic Diptera. Appendix. Science Herald, Budapest. 604 pp. ACKNOWLEDGMENTS ———. 2000b. Revision of the net-winged midges Original drawings were rendered by of the genus Blepharicera Macquart (Diptera: G.W. Courtney (Figs. 9, 10), J. Dorn Blephariceridae) of eastern North America. (Figs. 13–16), and L. Rogers (Figs. 11, Memoirs of the Entomological Society of 12). Assistance with scanning electron Washington 23: 1–99. Courtney, G. W., R. W. Merritt, K. W. Cummins, microscopy was courtesy of R. Brown M. B. Berg, D. W. Webb, and B. A. Foote. and the Microscopy and NanoImaging 2007. Ecological and distributional data for Facility at Iowa State University. Valu- larval aquatic Diptera, pp. 747–771. In Merritt, able comparative material was provided R. W., K. W. Cummins, and M. B. Berg, eds. by B.V. Brown, Natural History Museum An Introduction to the Aquatic of North America. Fourth Edition. Kendall / of Los Angeles County, Los Angeles, Hunt Publishing Co., Dubuque, Iowa. 1158 pp. California, and W.N. Mathis, National Frutiger, A. 1998. Walking on suckers—new Museum of Natural History, Smithso- insights into the locomotory behavior of larval nian Institution, Washington, D.C. net-winged midges (Diptera: Blephariceridae). Funding for various aspects of this work Journal of the North American Benthological was from the National Science Founda- Society 17: 104–120. ———. 2002. The function of the suckers of larval tion (Grants DEB-9407153 and DEB- net-winged midges (Diptera: Blephariceridae). 9796275). This journal paper of the Iowa Freshwater Biology 47: 293–302. Agriculture and Home Economics Exper- Hogue, C. L. 1966. The California species of iment Station, Project No. 6693, was Philorus: , early stages and descrip- supported by Hatch Act and State of tions of two new species (Diptera: Blepharocer- Iowa funds. idae). Contributions in Science, Natural Histo- ry Museum of Los Angeles County 99: 1–22. ———. 1970. Description of a new species of net- LITERATURE CITED winged from the Great Basin, with a key to the North American species of the genus Diop- Alexander, C. P. 1959. Undescribed species of topsis. Contributions in Science, Natural History nematocerous Diptera. Part VI. Bulletin of Museum of Los Angeles County 178: 1–10. the Brooklyn Entomological Society 54: 37–43. Alverson, A. J., G. W. Courtney, and M. R. ———. 1973. The net-winged midges or Blephar- Luttenton. 2001. Niche overlap of sympatric iceridae of California. Bulletin of the Califor- Blepharicera larvae (Diptera: Blephariceridae) nia Insect Survey 15: 1–83. from the southern Appalachian Mountains. ———. 1987. Blephariceridae. In Griffiths, G. C. D. Journal of the North American Benthological ed. Flies of the Nearctic Region. Stuttgart: E. Society 20: 564–581. Schweizerbart’sche Verlagsbuchhandlung, 2(4). Alverson, A. J. and G. W. Courtney. 2002. Jacobson, A. J. 2006. Phylogenetic analysis of Temporal patterns of diatom ingestion by Nearctic Blepharicera Macquart (Diptera:Ble- larval net-winged midges (Diptera: Blephari- phariceridae). M.Sc. Thesis, Iowa State Uni- ceridae: Blepharicera). Freshwater Biology 47: versity, Ames. 122 pp. 2087–2097. Johns, J. A. 1996. The net-winged midges (Diptera: Anderson, N. H. 1992. Influence of disturbance on Blephariceridae) of the southeastern United insect communities in Pacific Northwest States: phenology and ecology. M.Sc. Thesis, streams. Hydrobiologia 248: 79–82. Clemson University, Clemson. 80 pp. Courtney, G. W. 1990. Cuticular morphology of Lenat, D. R. 1993. A biotic index for the southeast- larval mountain midges (Diptera: Deuteroph- ern United States: derivation and list of lebiidae): implications for the phylogenetic tolerance values, with criteria for assigning relationships of . Canadian Jour- water quality ratings. Journal of the North nal of Zoology 68: 556–578. American Benthological Society 12: 279–290. ———. 1991. Life history patterns of Nearctic Zwick, P. 1984. Phylogenetic system and distribu- mountain midges (Diptera: Deuterophlebii- tion of the genus Blepharicera (Diptera: dae). Journal of the North American Bentho- Blephariceridae). 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