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A new species of gigantodax enderlein (diptera: simuliidae) from the United States PDF

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Preview A new species of gigantodax enderlein (diptera: simuliidae) from the United States

PROC. ENTOMOL. SOC. WASH. 98(4), 1996, pp. 741-751 A NEW SPECIES OF GIGANTODAX ENDERLEIN (DIPTERA: SIMULIIDAE) FROM THE UNITED STATES J. K. MOULTON Department of Entomology, Forbes Building, Rm. 410, University of Arizona, Tucson, AZ 85721, U.S.A. Abstract.(cid:8212)The larva, pupa, and adults of Gigantodax adleri, new species, are described from small springs in eastern Arizona and western New Mexico. This species represents the first United States record for the genus and a northern range extension of more than 3000 km. The preimaginal stages are associated with those of two species, Simulium patziciaense Takaoka and Takahasi and Simulium puigi Vargas, Martinez Palacios, and Diaz Najera, not previously recorded north of Guatemala and southern Mexico, respec- tively. Gigantodax adleri belongs to the Gigantodax wrighti group of Wygodzinsky and Coscaron, and its life stages are superficially similar to those of several congeneric species. Diagnoses that differentiate the female, male, and pupa from those of all other species and a detailed description of the preimaginal habitat are provided. Key Words: North America, springs, aquatic insect, Nematocera Gigantodax Enderlein, with 67 currently around the anal proleg, and posteroventrally recognized species, is the third largest black directed struts underlying the base of the fly genus and the second largest in the dorsal arms of the anal sclerite (Wygodzin- southern hemisphere. Typically, it is placed sky and Coscar6n 1989). The pupae are within the Prosimuliini, which along with best known for their spectacular diversity in the Simuliini comprise the Simuliinae (Wy- gill structure, which alone will separate godzinsky and Coscar6n 1973, 1989, Cros- many species. Some species, however, pos- skey 1988, Takaoka et al. 1988, Vargas and sess more typical gills and are scarcely sep- Ramirez-Pérez 1989). Currie (1988), how- arable from those of other genera. The ever, redefined the tribal limits within the adults are easily recognized by the straight, Simuliinae and restricted the Prosimuliini to rather than sinuous, CuA, vein and the dis- include only the following genera: Prosi- tinctive terminalia. The terminalia, howev- mulium Roubaud, Helodon Enderlein, Uro- er, are indistinguishable from those of spe- simulium Contini, Levitinia Chubareva and cies in the genus Cnesia Enderlein, which Petrova, Gymnopais Stone, and Twinnia is its sister group (Wygodzinsky and Cos- Stone and Jamnback. As a result, Gigan- caron 1973, 1989). todax and the remaining genera within the The greatest diversity in this genus oc- Simuliinae were assigned to the Simuliini, curs in the Andes, but representatives of the a view that is also corroborated by DNA G. wrighti species-group (Wygodzinsky sequence data (J. K. Moulton, unpublished). and Coscar6n 1989) are known as far north Larvae of Gigantodax are the only ones as the Sierra Madre de Occidental of central in the family with a seventeen-toothed hy- Mexico (Dalmat 1955, Vargas and Diaz Na- postoma, a complete, well sclerotized ring jera 1957, Wygodzinsky and Coscar6n 742 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1989). Larval habitats range from small the two males were obtained from these springs and rivulets to medium-sized rivers, rearings. The majority of adults were col- and some species are found only in acidic lected over the course of three days using or alkaline watercourses (Dalmat 1955, an emergence trap placed directly above an Wygodzinsky and Coscar6n 1989). Females exposed portion of the spring. The adults are presumably ornithophilic based upon were placed into 95% ethanol immediately the large basal tooth of the tarsal claw upon their discovery in the net. Paratype (Shewell 1955). larvae, deposited in collections other than Wygodzinsky and Coscarén (1989) rec- the author(cid:8217)s, that were initially fixed in Car- ognized eight species groups within Gigan- noy(cid:8217)s solution were transferred to 80% eth- todax, the G. brophyi, G. cilicinus, G. cor- anol for permanent storage. Pupae were monsi, G. cortesi, G. igniculus, G. minor, G. preserved in 60% ethanol to prevent col- multifilis, and G. wrighti groups, which con- lapsing of the gills, and then transferred to tained 17, 11, 10, 4, 2, 4, 1, and 15 species, 80% ethanol for permanent storage. The respectively, at that time. Since their treatise, holotype and all pinned adults were dehy- three additional species have been described drated with absolute ethanol and dried us- (Takaoka et al. 1988, Vargas and Ramirez- ing hexamethyldisilazane, HMDS (Poly- Pérez 1989). Gigantodax biergi and G. wil- sciences, Inc.). The holotype and some lei, described from Costa Rica by Vargas paratypes are deposited in the National Mu- and Ramirez -Pérez (1989), belong to the G. seum of Natural History, Smithsonian In- wrighti group, and Gigantodax lazoi, de- stitution, Washington, DC (USNM). Addi- scribed from Ecuador by Takaoka et al. tional paratypes are deposited in the Cana- (1988), is assignable to the G. cormonsi dian National Collection, Ottawa (CNC), group. The Natural History Museum, London In this paper, all life stages, after the egg, (NHM), the American Museum of Natural are described for a new species of black fly History, New York (AMNH), and the au- in the G. wrighti species group that repre- thor(cid:8217)s personal collection (JKM). The fig- sents the northernmost record for the genus ures of the larval head capsule, pupa, and and a new generic record for the United male and female terminalia were drawn by States. Also provided are diagnoses that dif- Ralph Idema for the upcoming book on the ferentiate the pupa and adults from those of black flies of North America and were all other species in the genus and a detailed made available to me by D. M. Wood. The account of the preimaginal habitat. The de- remaining drawings were rendered by the scription below provides a name for this author with the aid of an MTI CCD72 im- species that will enable its inclusion in a aging system connected to an Olympus book on the black flies of North America SZH10 stereo microscope. Length and being prepared by P. H. Adler, D. C. Currie, width measurements were made using ei- and D. M. Wood. ther this imaging system or by an ocular micrometer in an Olympus SZH zoom ste- MATERIALS AND METHODS reo microscope. Terminalia, mouthparts, Life-stage descriptions largely follow and legs were cleared in 85% lactic acid to that of Adler and Currie (1986). Taxonomic facilitate visibility and interpretation. terms predominantly follow that of Peterson (1981). Representative larvae were pre- DESCRIPTION served in absolute ethanol or Carnoy(cid:8217)s fix- Gigantodax adleri Moulton, new species ative, while others were transported live to (Figs. 1-17) the laboratory and reared in water taken from the type locality and maintained at Larva (mature).(cid:8212)Length 6.9(cid:8212)7.2 mm (x 10°C. All but one of the pupae and one of = 7.0, n = 5). General color of larva pink- VOLUME 98, NUMBER 4 743 \ 4 nal sensillum 3.4 times longer than basal width. Labral fan with 24(cid:8212)31 primary rays (x = 28, n = 4). Maxillary palpus (Fig. 4) about 1.9 times longer than basal width. Mandible (Fig. 5) with 10(cid:8212)11 inner teeth and 9-10 marginal serrations. Hypostomal bridge slightly darker than remainder of un- dersurface of head capsule, with faintly per- ceptible median longitudinal, light-colored stripe. Hypostoma (Fig. 6) with 17 teeth, one median tooth and 8 others per side; me- dian tooth shorter than tallest lateral tooth; first tooth from median arising outwardly from near the base of median tooth; second tooth from median deeply set and barely surpassing dorsal wall; third tooth from me- dian about as tall as median tooth; fourth tooth from median tallest tooth, its apex sloped outwardly; fifth and sixth teeth from median subequal in size although arising along downward slope; seventh tooth from median smallest of outer group of 5 teeth; eighth tooth from median prominent, its apex directed slightly inwardly. Lateral margin of hypostoma with 5(cid:8212)6 hypostomal setae and 7-8 lateral serrations per side. : (cid:8220)Keelof Postgenal cleft nearly imperceptible, at I° dema most a small notch. Cervical sclerites par- Figs. 1-2. Gigantodax adleri. 1, Larval head cap- tially enclosed anteriorly by arms of pos- sule (dorsal view). 2, Larval head capsule (ventral tocciput. Thoracic proleg with lateral plate view). subquadrate, moderately sclerotized. Per- ianal scales not developed. Ventral papillae ish beige to brown; intersegmental bands small, but distinct. Anal sclerite X-shaped, poorly defined. Head capsule (Figs. 1, 2) with heavily sclerotized, posteroventrally extremely variable in color, ranging from directed struts arising from base of anterior pale yellowish brown with distinct, dark- arms; posterior arms continuous as a well- brown headspots (pinkish beige larvae) to sclerotized, thin, brown ring (Fig. 7) sur- brown with headspots only slightly darker rounding posterior proleg. Posterior proleg brown than background and rather indis- with approximately 135 rows, each with tinct (brown larvae). Headspots subequal in 20-27 hooks. Anal papillae of three simple intensity and clearly comprised of multiple lobes. smaller spots; anterior posterolateral spot Pupa (Figs. 8(cid:8212)9).(cid:8212)Length of pupal body very small and concave; line over eyespots = 3.2-3.5 mm (x = 3.3, n = 5). Antenna thin, brown, and with small oval spot po- sheath of female extended slightly beyond sitioned anterior of midlength. Antenna posterior margin of head; antennal sheath of (Fig. 3) shorter than stem of labral fan, male extended about 2/3 distance to poste- somewhat pigmented, with apical article rior margin of head. Frontoclypeus of both darker than basal two; proportions of arti- sexes of normal shape, except for square cles (distal to proximal) 1.4:1.0:2.3; termi- protuberance medially, and with numerous 744 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ZZ Figs. 3-7. Gigantodax adleri. 3, Larval antenna. 4, Larval maxillary palpus. 5, Left larval mandible (ventral view). 6, Larval hypostoma. 7, Larval anal salerite (terminal view). regularly spaced, lightly verrucose platelets. with one short apical filament. Thorax with Frontal and clypeal trichomes in the form numerous, rather regularly spaced, slightly of strong setae. Gill about 2/5(cid:8212)1/2 length of verrucose platelets. Dorsocentral and dor- pupa, comprised of five, flimsy, pale gray, solateral trichomes setalike. Abdomen with corrugated, tubular filaments arising indi- platelets dorsally. Abdominal chaetotaxy as vidually from a short, common base; two follows: tergite I with 4 setae per side; ter- dorsal branches directed posteriorly, two gite II with 6(cid:8212)7 setae per side; tergites III median branches directed anterodorsally, and IV each with 4 retrorse hooks and 3 and a ventral branch directed posteriorly; setae per side, one setae anterior to fourth dorsal branches simple or with one terminal hook from midline and 2 setae laterally; ter- or subterminal filament; median branches gite V with 3 setae per side; tergites VI and slightly thicker than others, the dorsalmost VII with anterior row of spine combs and one with 4(cid:8212)5 and the other with 1(cid:8212)2 short, 1 seta per side; tergite VIII with anterior somewhat coiled filaments positioned just row of spine combs and 2-3 setae per side; basal to midlength; ventral branch usually tergite IX with pair of strong terminal VOLUME 98, NUMBER 4 745 SZe EePSe a me DEa e / celof idema Figs. 8-9. Gigantodax adleri. 8, Pupal thorax and gills (dorsal view). 9, Pupa and cocoon (lateral view). hooks and 2 setae per side. Sternite IV with with 3 long setae per side. Cocoon pale 2 setae per side, situated medially; sternite brown, covering abdomen and posterior V with 3 evenly spaced hook-like setae per portion of thorax, and often encrusted with side; sternite VI divided medially, each half small organic and inorganic particles. with 2 short, grapnel-like hooks; sternite Female.(cid:8212)General body color bright or- VII divided medially, each half with 1 grap- ange with golden pile. Length: thorax 1.2(cid:8212) nel-like hook; each half of sternites VI and 1.4 (x = 1.3, n = 4) mm; wing 3.4(cid:8212)4.4 (x VII bears a pleural sclerite with! grapnel- = 4.0, n = 4) mm. Frons gray, at vertex like hook; sternite VIII glabrous; sternite LX about 3 times broader than at narrowest 746 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-13. Gigantodax adleri. 10, Female left maxillary palpus (dorsal view). 11, Female tarsal claw (lateral view). 12, Female terminalia (ventral view with left anal lobe and cercus removed). 13, Female right anal lobe and cercus (lateral view). point, with sparse golden pile. Frontal angle and at end of short neck. Median proximal 77°. Fronto-ocular triangle about as high as space of cibarium broadly U-shaped, lack- wide. Clypeus brown, about as long as ing setae or armature. Postpronotum light wide, with golden pile. Occiput gray, with brown, with golden pile. Proepisternum golden pile reaching posterior margin of light brown, with tuft of long golden setae. eye; postocular setae golden. Antenna with Scutum bright orange, with purple-brown brown pile; scape and pedicel orange; fla- border from lateral of midline to superalar gellum orange, with apical two flagellom- region; anterior portion of this border ex- eres slightly darker. Proboscis brown. Man- tended posteriorly as a pair of thin, longi- dible with 20 teeth along inner margin. La- tudinal stripes extended to scutellum; be- cinia with 18 retrorse teeth. Palpus brown, tween this pair of stripes is a median stripe with stout brown setae; ratio of palpomeres of similar color that also reaches posterior (Fig. 10) 1.2:1.0:1.6; sensory vesicle of pal- declivity; scutum clothed in decumbent pomere III elongate, about 0.6 times length golden pile, with posteriormost portion also of segment, with external opening rounded with long golden brown recumbent setae. VOLUME 98, NUMBER 4 747 Anepisternum and katepisternum yellowish and with numerous setae. Spermatheca orange; membranous regions pale brown. spherical, brown, moderately sclerotized, Region anterior of mesothoracic spiracle lacking noticeable sculpturing, and with purple brown. Mesepimeral tuft golden distinct neck at junction with spermathecal brown. Mesbasisternum yellowish orange, duct. Anal lobe subquadrate, with some- rimmed in purple brown. Scutellum beige, what prominent rounded lobe ventrally. with long, golden-brown setae, these being Cercus flat, rounded posteriorly, the scler- recumbent anteriorly and decumbent pos- otized apical portion about twice as tall as teriorly. Metanotum dull orange brown, an- wide. terior 1/3 shiny brass brown. Wing trans- Male.(cid:8212)Thorax length 1.1(cid:8212)1.2 (x eZ lucent Veins pale brown. Setae of stem vein n = 2) mm. Wing length 3.1(cid:8212)3.2 (x = 3.1, and all other veins dark brown. Costa (C) n = 2) mm. Identical to that of female ex- with simple setae proximal to and simple cept as noted. Clypeus reduced to a narrow setae and spinelike setae distal to about strip, with long brown pile. Sensory vesicle midpoint of subcosta (Sc). Sc haired ven- subglobular, with distinct neck, 0.4 times trally. R, with one or two rows of setae, length of palpomere. Scutum with dark spinules present but not numerous, begin- brown recumbent setae posteriorly. Procoxa ning apical to level of insertion of Sc with yellowish orange. Wing much as in female, C. Rs simple, without any trace of forking. except spines on C beginning near junction False vein forked apically. M, and M, near- of R, + Rs and spines of R, beginning at ly straight, reaching wing margin. CuA, and junction of C and Sc. Scutellum shiny brass CuA, straight, reaching wing margin. A, brown, with long dark-brown setae. Protar- straight, nearly reaching wing margin. A, sus brown. Probasitarsus 8.5 times as long strongly curved, not nearly reaching wing as wide. Meta basitarsus 5.3 times as long margin. Halter whitish, slightly tinged with as wide. Abdominal tergites and sternites brown, especially on capitulum. Legs yel- brown with anterior and posterior margins low brown, with golden-brown, simple se- paler brown; membranous regions gray. tae. Procoxa brown. Probasitarsus 13 times Terminalia (Figs. 14-17) weakly sclero- as long as wide. Mesocoxa and metacoxa tized. Gonocoxite about three times longer yellowish orange. Metafemur with apex than greatest width. Gonostylus about two- brown. Metatibia with base and apex thirds as long as basimere, with two apical brown. Metabasitarsus 7.5 times as long as spines. Ventral plate about as wide as long, wide. Calcipala rounded, extended to center with median carina, and characteristic hor- of second tarsomere. Pedisulcus a series of izontal bridge connecting anterior arms. shallow wrinkles along slightly concave Median sclerite forked, the tines about 1/3 surface of second tarsomere. Basal tooth of length of structure. Paramere weakly scler- claw (Fig. 11) triangular, about 1/4 as long otized distally, apex membraneous and as claw. Abdominal tergites piceous, orange armed with approximately 12 large spines brown, with golden pile center of tergites and several additional spines progressively pale brown, with posterior margin clothed smaller proximally. with golden pile interspersed with sparse, Types.(cid:8212)Holotype: 2 (pinned), USA: brown pile. Membranous areas gray. Ster- ARIZONA: Apache County, Government nites orange brown with golden pile. Ab- Spring, 50 m S of Forest Service Road 1120 dominal fringe beige, with long golden se- (33°59'34"N, 109°27'54"W), elev. 8500 ft. tae. Terminalia (Figs. 12(cid:8212)13) weakly scler- (2591 m), 21 May 1994, J. K. Moulton otized. Genital fork with stem expanded (USNM). Paratypes: same data as holotype, apically, arms expanded distally, and with except May 21-24 19948 larvae [4 Car- long angled, dorsomedially projected apo- noy(cid:8217)s (CNC), 4 absolute ethanol (JKM)], 1 demes. Ovipositor flaps broadly rounded pharate pupa (JKM), 7 pupae (2 each in 748 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LUE (cid:8220)pr ertb hatt NAN Figs. 14-17. Gigantodax adleri. 14, Male terminalia (ventral view with left gonocoxite and gonostylus removed). 15, Male ventral plate, median sclerite, and parameres (lateral view). 16, Male ventral plate, median sclerite, and parameres (terminal view). 17, Male right dististylus (dorsal view). USNM and JKM, 1 each in CNC, NHM, ton(cid:8212)1 larva (JKM). NEW MEXICO: Cat- and AMNH), and 4 pinned @s (1 each in ron County, Fanny Spring @ Rt. 159 (Bur- CNC and NHM, 2 in JKM); same data, ex- sum Rd.) 4.2 km E of Mogollon cept 10 October 1994(cid:8212)10 larvae [5 Car- (33°22'55"N, 108°45'59"W), elev. ca. 9000 noy(cid:8217)s (NHM), 5 absolute ethanol (JKM)]. ft (2744 m), 20 May 1995, J. K. Moulton(cid:8212) Unnamed rivulet-like spring ca. 400 m S of 7 larvae, 1 pupa (NHM); same data, except Government Spring, 18 May 1994, J. K. 5 June 1995(cid:8212)4 larvae (NHM). Moulton(cid:8212)6 larvae (Carnoy(cid:8217)s) and | pharate Etymology.(cid:8212)This remarkable species is 3 (in ethanol) (JKM). Small spring (seep) named in honor of Peter H. Adler, a dear ca. 100 m N of Government Spring, 18 May friend and mentor to whom I will be forever 1994, J. K. Moulton(cid:8212)4 larvae (JKM). Gra- grateful for introducing me to the intriguing ham County, Pinaleno Mountains, Twilight world of simuliids. Spring @ Rt. 366, mile-marker 127 Diagnosis.(cid:8212)Female: The following suite (32°40'47"N, 109°51'22"W), elev. ca. 7000 of characters distinguishes this species from ft. (2134 m), 15 March 1995, J. K. Moul- all others in the genus: (1) scutum bright VOLUME 98, NUMBER 4 TA9 orange, bordered with purple brown, and ft., 3477 m). Government Spring percolates clothed with golden-yellow pile; (2) setae at a modest rate through stones and herba- of stem vein, legs, and scutellum dark gold- ceous vegetation. The streambed is approx- en brown; (3) mandible with 20 teeth along imately 4 m in width, but the spring di- inner margin; (4) lacinia of maxilla with 18 verges such that any particular flow is only teeth; (5) calcipala extended to 1/2 length a few cm wide. The spring was once the of second tarsomere; and (6) basal tooth of water source for a Civilian Conservation claw small, triangular, about 1/5(cid:8212)1/4 as Corps (CCC) trout hatchery (Peter Taylor(cid:8212) long as claw. Male: The bright orange, pur- United States Forest Service, Apache-Sit- ple-brown bordered scutum clothed with greaves Division, personal communication). golden pile, R, with spines, and dark brown Remnants of concrete pipes through which stem vein and scutellar setae separate this the spring is still partially diverted and six species from all other species in the genus. rectangular, trough-like, earthen holding Pupa: The five, flimsy, pale gray, tubelike tanks are clearly evident. Two other simu- filaments of the respiratory organ distin- lids, Simulium hechti Vargas and Diaz Na- guishes this species from all others except jera and S. patziciaense Takaoka and Tak- G. nasutus Wygodzinsky and Coscaron. ahasi, belonging to the Simulium hunteri The gill of G. adleri differs from that of G. species-group were also found in the type nasutus in the following regards: (1) fila- locality. Simulium patziciaense was previ- ments less stout, especially basally, not aris- ously known only from Guatemala (Taka- ing from a somewhat swollen gill base, (2) oka and Takahasi 1982). The dominant or- filaments directed more anterolaterally than ganism by far in the type locality was an anteriorly, and (3) second filament from unidentified species of amphipod. dorsal and ventralmost filaments not strong- The remaining two apparently unnamed ly curved ventrally and dorsally, respective- springs near the type locality supported ly, and pressed against the anterolateral por- much smaller populations of both simultids tion of body. The protuberance of the fron- and amphipods. One is located approxi- toclypeus of G. adleri further distinguishes mately 400 m south of the type locality and it from G. nasutus and possibly all other is rivulet-like, with water flowing over large species. Larva: Larvae in this genus are not flat rocks and between large rocks and boul- sufficiently characterized to make accurate ders. Located approximately 100 m north of species-level diagnoses without the pres- Government Spring, the other spring is ence of a respiratory histoblast. A prelimi- slow-flowing and has a mud bottom with nary analysis of larvae of several species few rocks and sparse vegetation. Associated indicates that those of many species are with this new species in the rivulet was separable using nothing more than body Prosimulium travisi Stone. No other simu- coloration and pigment distribution if fixed liids were present in the mud-bottomed in Carnoy(cid:8217)s. spring. The only other significant popula- Remarks.(cid:8212)The preimaginal stages occur tion of this species was found near the or- in springs approximately 0.1(cid:8212)1.0 m wide, a igin of Fanny Spring near the historic min- few cm deep, and less than 75 m in length. ing town of Mogollon, New Mexico. This The temperature at the source of these spring is similar to the type locality in that springs is about 8(cid:8212)-10°C, although during it is well shaded and has watercress as the the day the temperature downstream may dominant herbaceous plant. Associated with reach 15°C. The type locality, Government this species in Fanny Spring were S. patz- Spring, is the largest of three closely situ- iciaense, S. piperi Dyar and Shannon, and ated springs issuing from the western ridge S. puigi Vargas, Martinez Palacios, and of Lee Valley, just south of the town of Diaz Najera. Simulium puigi was previously Greer in the shadow of Baldy Peak (11,403 known only from Chiapas, Oaxaca, Vera- 750 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cruz, and Guatemala (Vargas et al. 1946, G. fulvescens (Blanchard) and G. rufescens Takaoka and Takahasi 1982). (Edwards); and C. S. Chaboo (AMNH), for Larvae seem to prefer flat surfaces of the loan of a paratype pupa of G. nasutus. rocks and the undersides of watercress This research was supported by D. R. Mad- leaves as substrates. Pupae were difficult to dison and the NSF Research Training find at the type locality. Undoubtedly, the Group Grant in the Analysis of Biological final instar larva moves into cracks and cer- Diversification (DIR-9113362) awarded to vices of rocks or into filamentous algae to the University of Arizona. spin its flimsy cocoon, which becomes fur- ther obscured by the accumulation of de- LITERATURE CITED bris. Of the two field-collected pupae, the one found at the type locality was in a Adler, P H. and D. C. Currie. 1986. Taxonomic res- olution of three new species near Simulium ver- clump of algae, and the one from Fanny num MacQuart (Diptera: Simuliidae). Canadian Spring was on the underside of a watercress Entomologist 118: 1207-1220. leaf. All laboratory-reared pupae were re- Crosskey, R. W. 1988. An annotated checklist of the moved from crevices formed by parts of the world black flies (Diptera: Simuliidae), pp. 425(cid:8212) stone bubbler assembly. Based upon the 520. In Kim, K. C., and R. W. Merritt, eds., Black flies: Ecology, Population Management, and An- presence of numerous larvae of different notated World List. The Pennsylvania State Uni- size classes in both May and October, this versity, University Park. species is presumably multivoltine. It is not Currie, D. C. 1988. Morphology and systematics of known whether eggs or larvae overwinter. primitive Simuliidae (Diptera: Culicomorpha). The habitats described are potentially re- Ph.D. thesis. University of Alberta, Edmonton, fugia, since they are inhabited by two spe- Alberta. Dalmat, H. T. 1955. The black flies (Diptera: Simu- cies, S. patziciaense and S. puigi, previous- liidae) of Guatemala and their role as vectors of ly known only from southern Mexico to onchocerciasis. Smithsonian Miscellaneous Col- Guatemala, and by G. adleri, whose closest lections 125: 1(cid:8212)425. relatives occur from southern Mexico to Peterson, B. V. 1981. Simuliidae, pp. 355-391. In northern South America. On the other hand, McAlpine, J. FE B., V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, these disparate distributions may be nothing coords., Manual of Nearctic Diptera, Vol. 1. Ag- more than an artifact resulting from a lack riculture Canada Research Branch, Ottawa. of faunistic research between these regions. Shewell, G. E. 1955. Identity of the black fly that Regardless, it is likely that additional spe- attacks ducklings and goslings in Canada (Dip- cies of Gigantodax occur in similar mon- tera: Simuliidae). Canadian Entomologist 87: taine habitats between southern Arizona 733-738. Takaoka, H. and H. Takahasi. 1982. A new species of and central Mexico. black fly (Diptera: Simuliidae) from upland areas of Guatemala. Journal of Medical Entomology 19: ACKNOWLEDGMENTS 63-67. I thank C. A. Olson (University of Ari- Takaoka, H., H. Hirai, and I. Tada. 1988. Description zona), for providing the emergence trap of a new species of Gigantodax (Diptera: Simu- liidae) from the Andean area of Ecuador. Journal used to obtain most of the imaginal mate- of Medical Entomology 25: 541-544. rial; D. G. Mead (University of Arizona), Vargas, L. and A. Diaz Najera. 1957. Simulidos Mex- for accompanying me on one of the expe- icanos. Revista del Instituto de Salubridad y En- ditions to the type locality; D. R. Maddison fermedades Tropicales 17: 143-399, + figures. (University of Arizona) and two anony- Vargas, L., A. Martinez Palacios, and A. Diaz Najera. mous reviewers for reviewing an earlier 1946. Simulidos de Mexico. Datos sobre siste- matica y morfologia. Descripcidn de dos nuevos draft of this manuscript; D. M. Wood subgéneros y especies. Revista del Instituto de (CNC(cid:8212)retired), for permitting me to use Salubridad y Enfermedades Tropicales 7: 101(cid:8212) the figures prepared by R. Idema; J. M. 192, 25 plates. Cumming (CNC), for the loan of adults of Vargas, M. V. and J. Ramirez-Pérez. 1989. Giganto-

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