Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest : Why inventory is a vital science

Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurqui de Moravia, San Jose Province, Costa Rica (hereafter referred to as Zurqui), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification. Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods. Comparison of species numbers from each of two permanent Malaise traps from Zurqui with those of single Malaise traps at each of Tapanti and Las Alturas, 40 and 180 km distant from Zurqui respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurqui did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase. Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurqui is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera. Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites. Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.Peer reviewe

Comprehensive inventory of true flies (Diptera) at a tropical site

Peer reviewe

Comprehensive inventory of true flies (Diptera) at a tropical site

Estimations of tropical insect diversity generally suffer from lack of known groups or faunas against which extrapolations can be made, and have seriously underestimated the diversity of some taxa. Here we report the intensive inventory of a four-hectare tropical cloud forest in Costa Rica for one year, which yielded 4332 species of Diptera, providing the first verifiable basis for diversity of a major group of insects at a single site in the tropics. In total 73 families were present, all of which were studied to the species level, providing potentially complete coverage of all families of the order likely to be present at the site. Even so, extrapolations based on our data indicate that with further sampling, the actual total for the site could be closer to 8000 species. Efforts to completely sample a site, although resource-intensive and time-consuming, are needed to better ground estimations of world biodiversity based on limited sampling

Cryptodacus obliquus Hendel

Cryptodacus obliquus Hendel Figs. 12, 13, 24 Specimens examined. PANAMA: Panamá: Parque Nacional Chagres, Altos de Pacora [9 ° 15 ' 28 "N 79 ° 21 ' 24 "W], Lote H 4, McPhail trap 571, 1 May 1998, C. A. Korytkowski, 1 adult without abdomen (USNM USNMENT 00214175). Comments. The single examined specimen from Panama is without its abdomen, except for syntergite 1 + 2, and is missing most of its setae. It has a diffuse moderate brown area on the gena, but otherwise is similar to the previously known specimens of C. obliquus from Peru and Bolivia.Published as part of Norrbom, Allen L. & Korytkowski, Cheslavo A., 2008, New Cryptodacus (Diptera: Tephritidae) from Panama, with a key to the known species, pp. 31-43 in Zootaxa 1773 on page 32, DOI: 10.5281/zenodo.18218

Anastrepha inaequalis Norrbom & Korytkowski, 2012, new species

<i>Anastrepha inaequalis</i>, new species <p>Figs. 24, 25, 47, 68, 83, 97, 98</p> <p> <b>Diagnosis.</b> <i>Anastrepha inaequalis</i> differs from most other species of <i>Anastrepha</i> by its pattern of dorsobasal denticles on the eversible membrane, with one medial denticle much larger than the others. In this regard it is similar to other species of the <i>megacantha</i> clade, but differs as indicated in the key.</p> <p> In the key of Steyskal (1977) it runs with difficulty to <i>A. parallela</i> (Wiedemann) or <i>A. kuhlmanni</i> Lima. It differs from both species in having paler setae, the large medial denticle on the eversible membrane, and the much shorter aculeus tip (0.08 mm long vs. 0.16 mm in <i>A. kuhlmanni</i> and 0.25–0.33 mm in <i>A. parallela</i>) with uneven margins (lateral margin evenly tapering in the other two species).</p> <p> <b>Description.</b> Mostly yellow to orange. Setae orange to dark orange.</p> <p>Head: Yellow to orange except brown ocellar tubercle. 3 frontal setae; 2 orbital setae, posterior seta well developed. Ocellar seta weak, slightly longer than ocellar tubercle. Facial carina, in profile, slightly concave on dorsal two-thirds. Antenna not extended to ventral facial margin. Palpus in lateral view dorsally curved, evenly setulose.</p> <p>Thorax: Mostly yellow to orange, with typical white areas poorly differentiated in holotype except dorsal margin of anepisternum. Subscutellum and mediotergite entirely orange. Mesonotum 3.63 mm long. Postpronotal lobe and scutellum entirely microtrichose; microtrichia of scutum, which is concave due to pin, not observable. Scutal setulae orange. Chaetotaxy typical for genus. Katepisternal seta weak, approximately half as long as anepisternal seta, but much weaker, orange.</p> <p>Legs: Entirely yellow to orange.</p> <p>Wing (Fig. 47): Length 8.08 mm, width 3.24 mm, ratio 2.49. Apex of vein R1 at 0.57 wing length, slightly proximal to level of anterior end of crossvein r–m. Cell c 1.32 times as long as pterostigma; pterostigma 5.91 times as long as wide. Vein R2+3 strongly sinuous. Crossvein r–m at 0.66 distance from bm–cu to dm–cu on vein M. Vein M strongly curved apically; cell r4+5 0.92 times as wide at apex as at level of dm–cu. Cell bcu with distal lobe moderately long, length of bcu 1.61 times as long as anterior margin, lobe 0.72 times as long as vein A1+Cu2. Wing pattern mostly orange and moderate brown. C-band mostly orange, paler in cell c, darker orange brown in pterostigma and with brownish streak in cell br. C-band and S-band broadly separated along veins R2+3 and R4+5 by hyaline band that extends from cell bm to costa, slightly narrowed along vein R2+3. Basal hyaline area in cell dm medium sized. Basal half of S-band mostly orange, posterodistal margin brown, very broadly in cell cu1, without incision in cell cu1, proximal margin narrowly brown in radial cells; distal section orange and moderate brown, medium width, at apex of vein R2+3 0.58 times width of cell r2+3, not extended to apex of vein M; hyaline area proximal to apex of band extending to vein R2+3. V-band complete, brown except proximal arm orange bordering most of dm-cu and in cell r4+5 except proximal margin and anterior proximal margin of distal arm, proximal arm separated from S-band, on posterior margin extended half distance to vein A1+Cu2; distal arm connected to proximal arm; cell r4+5 with small diffuse hyaline area between V-band and vein M.</p> <p>Abdomen: Mostly orange, without brown markings. Setulae orange brown.</p> <p> Female terminalia: Oviscape 4.91 mm long, 1.35 times as long as mesonotum, straight in lateral view; spiracle at basal 0.25. Eversible membrane (Fig. 68; lost in holotype after dissection) similar to <i>A. macracantha</i>, with 1 very large medial hooklike denticle and numerous small hooklike dorsobasal denticles, less than half as long as medial denticle, in suboval pattern. Aculeus (Fig. 83) slightly ventrally curved in lateral view, 4.60 mm long, 0.94 times as long as oviscape, in ventral view base strongly expanded, 0.15 mm wide; shaft rapidly tapered, 0.045 mm wide at midlength; tip (Figs. 97–98) 0.08 mm long, 0.02 times aculeus length, 0.04 mm wide, 2.0 times as long as wide, slightly tapered then slightly expanded subapically, nonserrate but with large notch on distal 0.30, 0.02 mm wide in lateral view, 0.50 times ventral width. Spermathecae elongate ovoid.</p> <p> <b>Distribution.</b> <i>Anastrepha inaequalis</i> is known only from Trinidad.</p> <p> <b>Biology.</b> The host plants and other aspects of the biology of this species are unknown.</p> <p> <b>Type data.</b> Holotype Ƥ (AMNH USNMENT 00104564), Trinidad: Arima Valley, 19 Apr 1950, W. Beebe.</p> <p> <b>Etymology.</b> This name is an adjective, from the Latin “in”, meaning not, and “aequalis”, meaning equal, in reference to the difference in size between the single larger medial hooklike denticle versus the other dorsobasal denticles on the eversible membrane.</p>Published as part of <i>Norrbom, Allen L. & Korytkowski, Cheslavo A., 2012, New species of Anastrepha (Diptera: Tephritidae), with a key for the species of the megacantha clade, pp. 510-552 in Zootaxa 3478</i> on page 529, DOI: <a href="http://zenodo.org/record/282325">10.5281/zenodo.282325</a&gt

Anastrepha lutea Stone

Anastrepha lutea Stone Figs. 5 –6, 13– 14. Anastrepha lutea Stone 1942: 95 [description, female; Panama]; Steyskal 1977: 27 [in key]; Norrbom et al. 1999 a: 80 [catalog]. Anastrepha bellicauda Norrbom in Norrbom & Kim 1988: 168 [description, male]; Norrbom et al. 1999 a: 77 [catalog; Venezuela]; Norrbom et al. 1999 b: 321, 324; McPheron et al. 1999: 345. New synonymy. Diagnosis. Anastrepha lutea also belongs to the schausi species group. The sexes are dimorphic in wing pattern and abdominal markings and setation. The male (Fig. 14) is easily recognized from other Anastrepha species by its highly reduced wing pattern, which is mostly diffuse yellow, and by the pattern of brown markings and clusters of large setae on the abdomen. The female (Fig. 13) differs from other species of the schausi group in having the C- and S- bands broadly connected along vein R 4 + 5. It was included in the key of Steyskal (1977). Description. Mostly yellow to orange. Setae dark brown to black. Body length: female 9.0–11.0 mm, male 6.5 –8.0 mm. Head: Yellow to orange except ocellar tubercle brown. Face entirely microtrichose and without white or brown markings in either sex, ventral margin not expanded laterally, carina weak, in profile concave. 3–4 frontal setae. Usually 2 orbital setae (posterior seta absent on 1 side in 2 specimens and on both sides in 2 of 18 specimens). Ocellar seta weak, 1–2 times as long as ocellar tubercle. Antenna extended 0.60–0.75 distance to ventral facial margin. Arista short pubescent. Thorax (Figs. 13–14): Mostly yellow to orange, without brown markings; postpronotal lobe, scutellum except base of disc, scutal vittae and dorsal margin of anepisternum white; medial scutal vitta slender, slightly broadened and rounded posteriorly, extended laterally to or slightly beyond acrostichal seta. Mesonotum 2.90–3.75 mm long. Scutum, postpronotal lobe, notopleuron and scutellum entirely microtrichose. Scutal setulae mostly yellow, brownish laterally. Chaetotaxy as usual for genus, katepisternal seta paler and weaker than other setae but moderately long, 2 / 3 to as long as postocellar seta. Wing (Figs. 5–6): Length 6.7–8.5 mm, width 2.55–3.35 mm, ratio 2.47–2.73. Cell c 1.11–1.23 times as long as pterostigma. Apex of vein R 1 at 0.53–0.58 wing length. Vein R 2 + 3 nearly straight. Vein M weakly curved apically; cell r 4 + 5 0.96–1.14 times as wide at apex as at level of dm-cu. Crossvein r-m at 0.66–0.70 length of cell dm, ratio of second to third sections of vein M 1.96–2.31. Distal lobe of cell bcu moderately long, bcu 1.43–1.57 times as long as its anterior margin. Pattern strongly sexually dimorphic. In male (Fig. 6) mostly hyaline with diffuse, pale yellow basal area of varying extent, usually covering cells bc, c, pterostigma, bm, bcu, base of cu 1, and cells r 2 + 3, br, and dm to level of crossvein r-m or slightly beyond, br usually with hyaline area posterior to pterostigma; crossvein dm-cu sometimes narrowly bordered with faint yellow. In female (Fig. 5; Stone 1942, pl. 20 A) pattern of more typical Anastrepha type, mostly yellow to orange brown, posterior margin of base of S-band in cell cu 1, sometimes extending to vein R 4 + 5, distal part of S-band, and proximal arm of V-band posterior to vein M darker. C- and S-bands broadly connected along vein R 4 + 5, hyaline area distal to apex of vein R 1 with somewhat diffuse margins and usually rounded or irregular in shape, extending only to or slightly posterior to vein R 2 + 3. S-band extended basally into posterior 1 / 4 – 1 / 2 and distal margin of cell bm; distal section moderately broad, at apex of vein R 2 + 3 0.57–0.74 times width of cell r 2 + 3; narrowly separated from or just reaching apex of vein M. V-band separated from S-band, incomplete, distal arm absent at least posteriorly, if present anteriorly broad and diffuse, without hyaline area between it and vein M; proximal arm extended basally along posterior wing margin more than half distance from vein Cu 1 to vein A 1 +Cu 2 but not connected to base of S-band. In male microtrichose except cell bc, extreme base of br (proximal to crossvein h), alula, and sometimes anterior and/or posterior areas in bcu or very small basal area in cell cu 1. In female microtrichose except cell bc, base and posterior 1 / 4 – 2 / 3 of cell c, extreme base (proximal to crossvein h) and posterior margin of subapical hyaline area of br, bm except distal and usually posterior margin, small basal or anterobasal area in dm, anteriorly and posteriorly in cell bcu (broadly to narrowly microtrichose along medial fold), very small basal area in cu 1, small narrow anterobasal area in a 1, and most or all of alula. Male abdomen (Fig. 14): Syntergite 1 + 2 mostly yellow to orange fading to white posteriorly. Tergites 3 and 4 with dark brown band on basal half, sometimes weakly and narrowly divided medially; white posteriorly. Tergite 5 dark brown on lateral 1 / 3 – 2 / 5, separated by white area slightly to strongly tapered posteriorly. White areas of all tergites with dense silvery white microtrichia. Brown areas of tergite 5 nonmicrotrichose. Setae on lateral margins of tergites 3 and 4, lateral and apical margins and on brown areas of tergite 5 large and stout. Lateral surstylus in posterior view elongate triangular, medial margin convex to very slightly concave, lateral margin slightly concave subapically, apex acute. Proctiger without lateral fold separating sclerotized areas. Phallus 4.95–5.50 mm long; 1.41–1.90 times as long as mesonotum. Glans 0.60–0.65 mm long. Female abdomen (Fig. 13): Tergites yellow to orange with posterior margins white, in Panama females tergite 3 with pair of narrow, diffuse brown bands on lateral 1 / 4 – 1 / 3 on basal half; basal, lateral parts of tergites 4 and 5 also slightly darker. Tergites mostly microtrichose but without denser, silvery white microtrichia; tergites 3–5 with nonmicrotrichose band at midlength on lateral 1 / 3 – 2 / 5. Oviscape 3.20–3.85 mm long, 0.97– 1.19 times as long as mesonotum, entirely yellow to orange; spiracle at basal 0.28–0.32. Eversible membrane with about 35 long, slender, hook-like dorsobasal scales in triangular to semicircular pattern. Aculeus 2.95– 3.70 mm long; base 0.19–0.23 mm wide; shaft 0.085–0.105 mm wide at midlength; tip (Stone 1942, fig. 19 A) 0.21–0.27 mm long, 0.100– 0.105 mm wide, 2.10–2.57 times as long as wide, nonserrate, gradually tapered. Spermathecae elongate ovoid. Type data. A. lutea: Holotype Ψ (National Museum of Natural History, Washington, DC (USNM USNMENT 00212764), PANAMA: Panamá: El Cermeño [8 ° 44 'N 79 ° 51 'W], Fruit fly trap, 3 Oct 1939, J. Zetek 4553. A. bellicauda: Holotype ɗ (USNM USNMENT 00212763), PANAMA: El Cermeño, Fruit fly trap, Dec 1939 – Jan 1940, J. Zetek 4621. Other specimens examined. PANAMA: Panamá: El Cermeño, 5 Dec 1939, J. Zetek 4600, 1Ψ paratype (USNM USNMENT 00212765); El Cermeño, Fruit fly trap, 12 Nov 1940, J. Zetek 4701, 1Ψ (USNM USNMENT 00212766). VENEZUELA: Trujillo: La Chira, 9 ° 12 ' 54 "N 70 ° 51 ' 23 "W, 300 m., emerged 15 Jul 1995, reared ex fruit of "cusco", K. P. Katiyar & J. Oroño MFAKP-00913, 8 ɗ 8 Ψ (USNM USNMENT 00048639- 54). Biology. In Venezuela K. P. Katiyar and colleagues reared A. lutea from fruit of a plant identified only as “cusco”, probably a species of Sapotaceae (K. P. Katiyar, pers. comm.). Distribution. Anastrepha lutea is known only from Panama and Venezuela. Comments. Norrbom described A. bellicauda from a male, presuming that the conspecific female would have the C- and S-bands of the wing separated as in other species of the schausi group, but the reared series from Venezuela indicates that it is conspecific with A. lutea, previously known only from females.Published as part of Norrbom, Allen L. & Korytkowski, Cheslavo A., 2007, A new species, new synonymy, and taxonomic notes in the Anastrepha schausi group (Diptera: Tephritidae), pp. 47-55 in Zootaxa 1497 on pages 51-55, DOI: 10.5281/zenodo.17705

Cryptodacus tau Foote

Cryptodacus tau (Foote) Figs. 2, 3, 17, 29, 30 Specimens examined. COSTA RICA: Guanacaste: Río Naranjo, 3 km SE [10 ° 40 'N 85 ° 4 'W], 22–30 Mar 1992, F. D. Parker, 1 Ψ (USU USNMENT 00216547). MEXICO: Jalisco: Careyes, 12 Feb– 19 Mar 1997, F. D. Parker, 1 Ψ (USNM USNMENT 00213633). Comments. This species previously has been reported from Mexico (Sonora, Sinaloa, Morelos) and Guatemala (Foote 1978, Norrbom 1994). The examined specimens include the first specimen known from Costa Rica along with an additional record from Mexico (Jalisco).Published as part of Norrbom, Allen L. & Korytkowski, Cheslavo A., 2008, New Cryptodacus (Diptera: Tephritidae) from Panama, with a key to the known species, pp. 31-43 in Zootaxa 1773 on page 32, DOI: 10.5281/zenodo.18218

Cryptodacus ornatus Norrbom

<i>Cryptodacus ornatus</i> Norrbom <p>Figs. 14, 22, 26</p> <p> <b>Specimens examined.</b> ECUADOR: Napo: Reserva Etnica Waorani, Onkone Gare Camp, 1 km. S, Transect Ent., 0°39'10"S 76°26'W, Transect 5, Station 9, 220 m., insecticidal fogging, terra firme forest, 9 Oct 1994, T. L. Erwin et al. Project MAXUS lot 918, 1Ψ (USNM USNMENT00054001).</p> <p> <b>Comments.</b> This species previously was known from Colombia and Brazil (Amazonas) (Norrbom 1994).</p>Published as part of <i>Norrbom, Allen L. & Korytkowski, Cheslavo A., 2008, New Cryptodacus (Diptera: Tephritidae) from Panama, with a key to the known species, pp. 31-43 in Zootaxa 1773</i> on page 32, DOI: <a href="http://zenodo.org/record/182184">10.5281/zenodo.182184</a&gt