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FIGURE 11. A–B in The Frog-Biting Midges of Borneo—From Two to Eleven Species (Corethrellidae: Diptera)
No full textFIGURE 11. A–B. Thoraces and legs of female Corethrella, lateral view; slide-mounted. C. Portion of midfemur of female C. calathicola, at about distal third of femur, showing scales, dorsal view (with leg extended laterally from body).Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo—From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on page 34, DOI: 10.5281/zenodo.21086
Corethrella (Corethrella) nanoantennalis Borkent & Grafe, new species
No full textCorethrella (Corethrella) nanoantennalis Borkent & Grafe, new species (Figs. 3 A, 4 C, 6 C, 8 C, 9 C, 12 C, 13 C, 14 A) DIAGNOSIS: Male adult. Unknown. Female adult. Only extant species of Corethrella in the Palaearctic, Oriental, Oceanian and northern Australasian Regions with the combination of no pigmentation distal to a single distinct midlength wing band (no subapical band) (Fig. 12 C), flagellomeres 1–4 short, each notably shorter than flagellomere 5 (Fig. 8 C), with non-discrete darker pigmentation on the basal 0.5 of the midfemur and basal 0.3–0.4 of the hind femur, uniformly yellow or light brown hind tibia, and abdominal tergites each with an anterior darker band (Fig. 9 C). Only extant species in Borneo with the combination of flagellomeres 1–4 short (Fig. 8 C) and palpal segments 1–2 dark brown and contrasting with pale segments 3–4 (5 grading from pale to medium brown) (Fig. 6 C). DESCRIPTION: Male adult. Unknown. Female adult. Descriptive statistics: See Tables 1 –5. Head: Outline in anterior view somewhat circular (Fig. 6 C). Coronal suture elongate, extending ventrally past ommatidia. Two large setae on frons between ventromedial area of ommatidia. Antenna uniformly medium brown; pedicel without distinctive elongate, stout setae. Antennal flagellomeres as in Fig. 8 C; sensilla coeloconica on flagellomeres 1 –2, 9– 13, only flagellomere 1 with more than one; flagellomere 13 with well-developed apical bifurcation. Clypeus (Fig. 6 C) somewhat square. Mandible with very small, pointed teeth. Palpus (Fig. 6 C) with segments 1–2 dark brown, very base of segment 3 dark brown or pale, most or all of segment 3, segments 4 light brown, segment 5 grading from light to medium brown apically; segment 3 swollen apically. Thorax (Fig. 9 C): Scutum, scutellum mostly light to medium brown, contrasting with darker brown pleura; pale sclerites around base of wing. Posterior portion of dorsocentral row with 2 elongate setae situated somewhat lateral to one another. Prescutal suture elongate, thick, uninterrupted, extending to near dorsocentral row of setae. Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin not thick. Wing (Fig. 12 C): Apex of R 2 slightly distal to apex of M 1. Anterior margin with differently, discretely pigmented scales (indicating anterior margin of midlength band), with midlength band, with darker more basal scales restricted to vein C, Sc, slightly in more posterior veins, with basal scales on posterior margin of wing dark; veins (other than costa and wing margin) with well-developed scales. Halter medium brown, equal to scutellum. Legs (Fig. 9 C): Light brown, with about basal 0.5 of forefemur slightly darker, about basal 0.5 of midfemur darker, about basal 0.2–0.3 of hind femur darker; with darker pigmentation not discrete. With only slender setae, lacking scales. Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Empodia thick. Abdomen (Fig. 13 C): Tergites 2–7 pale with anterolateral corners more darkly pigmented, sternites light brown with sternites 2–6 with posterior margin pale. Segments 8–9 medium brown. Cercus pale. Immature stages. Unknown. DISTRIBUTION AND BIONOMICS: Corethrella nanoantennalis is known from female specimens collected in Ulu Temburong National Park in Brunei at 110 m and Lanjak Entimau, Sarawak, Malaysia at 80 m (Fig. 14 A). Habitats are found in steep terrain in mature mixed dipterocarp forest (as in Fig. 1 F). One female was collected with a frog-call trap broadcasting 4 kHz, the maximum frequency (Hz) attractive to any Corethrella species (Grafe et al. unpublished). The others were collected feeding on calling males of two species of frogs (Table 7). TAXONOMIC DISCUSSION: We have placed C. nanoantennalis in the drakensbergensis species group based on the presence of synapomorphies 21–23 and its plesiomorphic complete prescutal suture (character 28) (Fig. 19). It could equally form an earlier lineage anywhere between the nodes defined by Borkent (2008: fig. 141) by " 21–23 " and "26, 28– 30 ". Discovery of the male will help resolve its phylogenetic position. TYPES: Holotype, female adult on microscope slide, labeled " HOLOTYPE Corethrella nanoantennalis Borkent and Grafe ", "Sungai Mata Ikan, Ulu Temburong National Park, Brunei, 10 -VII- 2008; on M. nasuta, U. Grafe; 11 n 1 ", " 11 n 1 050109 " (CNCI). Paratypes on microscope slides: 4 Ƥ, Lanjak Entimau, Sarawak, Malaysia, 112 ° 4 'E, 1 ° 28 'N, 80 m, 24 -VI- 2008 (CNCI); 1 Ƥ, from previous locality but 18 -VI- 2008 (UBD). Paratypes on pins: 11 Ƥ, Lanjak Entimau, Sarawak, Malaysia, 112 ° 4 'E, 1 ° 28 'N, 80 m, 24 -VI- 2008 (9, CNCI; 2 UBD); 1 Ƥ, from previous locality but 18 -VI- 2008 (CNCI). DERIVATION OF SPECIFIC EPITHET: The name nanoantennalis (small, antenna) refers to the relatively short flagellomeres 1–3 of females of this species.Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 9-10, DOI: 10.5281/zenodo.21086
Corethrella (Corethrella) bipigmenta Borkent & Grafe, new species
No full text<i>Corethrella</i> (<i>Corethrella</i>) <i>bipigmenta</i> Borkent & Grafe, new species <p>(Figs. 2 C, 4D, 6D, 8D, 9D, 12D, 13D, 15B)</p> <p> DIAGNOSIS: <i>Male adult.</i> Unknown. <i>Female adult.</i> Only extant species of <i>Corethrella</i> in the Palaearctic, Oriental, Oceanian and northern Australasian Regions with the combination of no pigmentation distal to the single distinct midlength wing band (no subapical band) (Fig. 12D), flagellomeres 1–4 elongate, each similar in length to flagellomere 5 (Fig. 8 D), with non-discrete darker pigmentation on the basal 0.5 of the midfemur and basal 0.3–0.4 of the hind femur, uniformly yellow or light brown hind tibia (Fig. 9D), and abdominal tergites each with anterolateral darker pigmentation (Fig. 13D). Only extant species in Borneo with the combination of flagellomeres 1–4 elongate, each similar in length to flagellomere 5 (Fig. 8 D) and palpal segments 1 and 2 dark brown and contrasting with pale segments 3 and 4 (5 grading from pale to medium brown) (Fig. 6 D).</p> <p> DESCRIPTION: <i>Male adult</i>. Unknown. <i>Female adult.</i> Descriptive statistics: See Tables 1 –5. <b>Head</b>: Outline in anterior view somewhat circular (Fig. 6 D). Coronal suture elongate, extending ventrally past ommatidia. Two large setae on frons between ventromedial area of ommatidia (as in Fig. 16B). Antenna uniformly medium brown; pedicel with distinctive elongate, stout setae. Antennal flagellomeres as in Fig. 8 D; sensilla coeloconica on flagellomeres 1–2, 9–13, only flagellomere 1 with more than one; flagellomere 13 with well-developed apical bifurcation. Clypeus somewhat square. Mandible with moderately small, pointed teeth. Palpus (Fig. 6 D) with segments 1–2 dark brown, segments 3–5 light brown, segment 5 darker apically; segment 3 swollen apically. <b>Thorax</b> (Fig. 9D): Scutum, scutellum mostly light to medium brown, contrasting with darker brown pleura; pale sclerites around base of wing. Posterior portion of dorsocentral row with 2 elongate setae situated somewhat lateral to one another. Prescutal suture elongate, thick, uninterrupted, extending to near dorsocentral row of setae.</p> <p> Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin not thick. <b>Wing</b> (Fig. 12D): Apex of R2 slightly distal to apex of M1. Anterior margin with differently, discretely pigmented scales (indicating anterior margin of midlength band), with midlength band, with darker more basal scales restricted to vein C, Sc, slightly in more posterior veins, with basal scales on posterior margin of wing dark; veins (other than costa and wing margin) with well-developed scales. Halter medium brown, equal to or slightly lighter than scutellum. <b>Legs</b> (Fig. 9D): Light brown, with about basal 0.5 of forefemur slightly darker, about basal 0.5 of midfemur darker, about basal 0.3–0.4 of hind femur darker; with darker pigmentation not discrete. With only slender setae, lacking scales. Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Empodia thick. <b>Abdomen</b> (Fig. 13D): Tergites 2–7 pale or light brown with anterolateral corners more darkly pigmented, tergite 8 pale or light brown, sternites light brown with sternites 2–6 with anterior portions medium brown, posterior margins pale, sternites 7–8 medium brown. Cercus medium light brown.</p> <p> <i>Immature stages</i>. Unknown.</p> <p> DISTRIBUTION AND BIONOMICS: <i>Corethrella bipigmenta</i> is known only from steep terrain in the mature mixed dipterocarp forest (Figs. 1 E, F) of Ulu Temburong National Park, Brunei, at elevations of 50–120 meters (Fig. 15B). Three females in the type series were collected on 22-VII-2006 from a calling male <i>Megophrys nasuta</i> (Schlegel) at 8:30 PM (Fig. 2 C). A photograph by the second author (Fig. 2 C) shows numerous identical looking female <i>Corethrella</i> on and surrounding the frog: seven were on the legs, two were on the back, and at least 10 were on nearby substrate. Further specimens were collected from other <i>M. nasuta</i> and one other frog species (Table 7). This species has not yet been collected with a frog-call trap.</p> <p> TAXONOMIC DISCUSSION: The three type specimens of <i>C. bipigmenta</i> were noted by Borkent (2008) in the taxonomic discussion of <i>C. nippon</i> (but mistakenly stated as coming from Sungai Belalong). This species is very similar to <i>C. nanoantennalis</i> but differs in the relative lengths of flagellomeres 1–4.</p> <p> <i>Corethrella bipigmenta</i> is placed in the <i>drakensbergensis</i> species group for the same reason that <i>C. nanoantennalis</i> was placed there, along with the same caveats (Fig. 19). However, it appears that this species has synapomorphy 40 (pedicel with strong setae), which would place it elsewhere in the phylogeny of the Corethrellidae. We are inclined however, to question the validity of this synapomorphy – it is often difficult to interpret consistently in specimens and its distribution may have been misrepresented by Borkent (2008).</p> <p> TYPES: Holotype, female adult on microscope slide, labeled " HOLOTYPE <i>Corethrella bipigmenta</i> Borkent and Grafe ", "Tributary of Sungai Temburong in Temburong district of Brunei Darussalam, 115° 09'E, 4° 33'N, 22-VIII-2006, U. Grafe, CD2069" (CNCI). Paratypes on microscope slides: 2 Ƥ, labeled as for holotype (1, CNCI; 1 UBD). Paratypes on pins: 2 Ƥ, from Kuala Belalong, Ulu Temburong NP, Brunei, 50 m, 23-VII-2007 (CNCI); 4 Ƥ, from Sungai Esu, Ulu Temburong NP, Brunei, 60 m, 11-VII-2008 (CNCI); 2 Ƥ, from Sungai Baki, Ulu Temburong NP, Brunei, 120 m, 18-VII-2008 (1 CNCI; 1 UBD); 1 Ƥ, Sungai Mata Ikan, Ulu Temburong NP, Brunei, 110 m, 10-VII-2008, (UBD).</p> <p> DERIVATION OF SPECIFIC EPITHET: The name <i>bipigmenta</i> (two, pigment) refers to differently pigmented palpal segments 1–2 (dark) and segments 3–5 (lightly pigmented) of females of this species.</p>Published as part of <i>Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279</i> on pages 10-11, DOI: <a href="http://zenodo.org/record/210864">10.5281/zenodo.210864</a>
Corethrella (Corethrella) bicincta Borkent, Grafe
No full textCorethrella (Corethrella) bicincta Borkent, Grafe & Miyagi, new species (Figs. 5 D, 7 C, 8 J, 11 A, 12 J, 13 I, 15 B) DIAGNOSIS: Male adult. Unknown. Female adult. Only extant species of Corethrella in the Palaearctic, Oriental, Oceanian and northern Australasian Regions with the combination of a subapical band on the wing (Fig. 12 J), a uniformly brown abdomen (Fig. 13 I), and with the hind tibia with darker pigmentation at base and apex (Fig. 11 A). Only extant species in Borneo with the combination of a subapical band on the wing (Fig. 12 J) and a uniformly brown abdomen (Fig. 13 I). DESCRIPTION: Male adult. Unknown. Female adult. Descriptive statistics: See Tables 6 –11. Head: Outline in anterior view laterally somewhat elongate (Fig. 7 C). Coronal suture elongate, extending to midheight to ventrally past ommatidia. Two large setae on frons between ventromedial area of ommatidia. Antenna uniformly medium brown; pedicel with distinctive, more elongate, stout, dorsal or dorsolateral setae; flagellomeres as in Fig. 8 J; sensilla coeloconica on flagellomeres 1, (8), 9–13, flagellomeres 1 with several, 9–13 with 2 sensilla; flagellomere 13 with well-developed apical bifurcation. Clypeus (Fig. 7 C) wide. Mandible with moderately small, pointed teeth. Palpus (Fig. 7 C) brown; segment 3 swollen subapically. Thorax (Fig. 11 A): Uniformly medium to dark brown but scutum with anterolateral darker patch of darker pigmentation, pale sclerites around base of wing. Posterior portion of dorsocentral row with 2 elongate setae situated somewhat lateral to one another. Prescutal suture elongate, thick, interrupted by area of pale cuticle, extending nearly to dorsocentral row of setae. Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin thick. Wing (Fig. 12 J): Apex of R 2 equal to apex of M 1. Anterior margin with differently, discretely pigmented scales (indicating anterior margin of wing bands), basal band with dark scales on C, Sc, R, subbasal band with dark scales on R, M, A 1, midlength band entire, subapical band with dark scales but R 4 + 5 pale; veins (other than costa and wing margin) with well-developed scales. Halter as dark as scutellum. Legs (Fig. 11 A): Uniformly pigmented dark brown, except for poorly defined lighter pigmentation on apex of forefemur, base of foretibiae in some, in all apical 0.3 of hind femur pale, hind tibiae pale but with basal and apical bands of dark pigmentation. With only slender setae, lacking scales. Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Empodia slender. Abdomen (Fig. 13 I): Uniformly medium brown, with segments 8–9. Cercus darker brown. Immature stages. Unknown. DISTRIBUTION AND BIONOMICS: Corethrella bicincta is known from Brunei and eastern Sarawak, Malaysia (Fig. 15 B) at elevations of 30–1083 m. Habitats include mid-elevation peat swamps or a nearby river (Fig. 1 G) and steep terrain in lowland mature mixed dipterocarp forest. They were collected with frog-call traps and on a calling male of one species of frog (Table 7). TAXONOMIC DISCUSSION: The specimens from Brunei and Gunung Mulu National Park (low elevations) were smaller (wing length less than 1.3 mm) than those from the Bario highlands (wing length more than 1.4 mm). Corethrella bicincta may be the first Old World species of the quadrivittatus species group. It has all the synapomorphies of the lineage identified by characters 39–44 (Borkent 2008: fig. 141) although character 44 could not be scored (present only in the currently unknown male). Corethrella bicincta lacks, however, both the apomorphic state of characters 33 and 68 and must be considered as unplaced until other life stages and character states become available. TYPES: Holotype, female adult on microscope slide, labeled " HOLOTYPE Corethrella bicincta Borkent, Grafe and Miyagi", "Gunung Mulu National Park, Sarawak, Malaysia, Headquarters, 14 -XI- 2009; on Hylarana glandulosa, U. Grafe;" (CNCI). Paratypes on microscope slides: 1 Ƥ, labeled as for holotype (CNCI); 10 Ƥ, Bario, Sarawak, Malaysia, 1060 m, on several dates: 2 -VIII- 2007, 6,7-IX-2007, 29-VIII- 2008 (9, CNCI; 1, UBD). Paratypes on pins: 1 Ƥ, Pa Umor, 5 km E. Bario, Sarawak, Malaysia, 1083 m, 25 -XI- 2010 (CNCI); 1 Ƥ, 0.5 km E Pa Umor, 03° 44 ' N 115 ° 30 'E, Sarawak, Malaysia, 1083 m, 27 -XI- 2010 (CNCI); 11 Ƥ, Sungai Ingei, 8.2 km S. Melilas, Brunei, 50 m, VII- 2010 (10, CNCI; 1 UBD); 12 Ƥ, from previous locality but 10 -VII- 2010 (11, CNCI; 1 UBD); 5 Ƥ, Gunung Mulu National Park, Sarawak, Malaysia, Headquarters, 30 m, 14 -XI- 2009 (CNCI). DERIVATION OF SPECIFIC EPITHET: The name bicincta (two belts) refers to the presence of both a midlength and subapical band on the wing of females of this species.Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 16-17, DOI: 10.5281/zenodo.21086
FIGURE 6 in A new tree-frog genus and species from Ivory Coast, West Africa (Amphibia: Anura: Hyperoliidae)
No full textFIGURE 6. Pectoral girdle and vertebral column of adult male Morerella cyanophthalma sp. nov. Pectoral apparati of SMNS 11940 (a: drawing after clearing and staining) and MNHG 3121-36 (b: scan). Vertebral column (scan of MHNG 3121-36) in dorsal (c) and ventral (d) view. CL = clavicle, CO = coracoid, ECO = epicoracoid, OM = omnosternum, ST = sternum; scale bar = 1mm.Published as part of Rödel, Mark-Oliver, Kosuch, Joachim, Grafe, Ulmar, Boistel, Renaud, Assemian, Emmanuel, Kouamé, N'Goran G., Tohé, Blayda, Gourène, Germain, Perret, Jean-Luc, Henle, Klaus, Tafforeau, Paul & Pollet, Nicolas, 2009, A new tree-frog genus and species from Ivory Coast, West Africa (Amphibia: Anura: Hyperoliidae), pp. 23-45 in Zootaxa 2044 on page 36, DOI: 10.5281/zenodo.18639
Corethrella (Corethrella) gilva Borkent & Grafe, new species
No full textCorethrella (Corethrella) gilva Borkent & Grafe, new species (Figs. 5 E, 7 D, 8 K, 11 B, 12 K, 13 J, 16 A) DIAGNOSIS: Male adult. Unknown. Female adult. Only extant species of Corethrella in the Palaearctic, Oriental, Oceanian and northern Australasian Regions with the combination of the palpus and clypeus equally dark brown (Fig. 7 D), a single, midlength wing band (Fig. 12 K), tergites 2–6 uniformly brown (Fig. 13 J), midfemur either uniformly brown or lighter apically (non-discrete), and hind tibia with, at most, very light pigmentation at apex (Fig. 11 B). Only extant species in Borneo with the combination of the palpus and clypeus equally dark brown (Fig. 7 D), a single, midlength wing band (Fig. 12 K), hind tibia with, at most, very light pigmentation at apex (Fig. 11 B), and tergites 2–6 uniformly brown (Fig. 13 J). DESCRIPTION: Male adult. Unknown. Female adult. Descriptive statistics: See Tables 1 –5. Head: Outline in anterior view somewhat circular (Fig. 7 D). Coronal suture elongate, extending ventrally nearly past ommatidia. Two large setae on frons between ventromedial area of ommatidia. Antenna uniformly medium brown; pedicel with distinctive elongate, stout setae. Antennal flagellomeres as in Fig. 8 K; sensilla coeloconica on flagellomeres 1 –2, 8– 13, only flagellomere 1 with more than one; flagellomere 13 with well-developed apical bifurcation. Clypeus (Fig. 7 D) somewhat square. Mandible with very small, pointed teeth. Palpus (Fig. 7 D) medium brown; segment 3 swollen near midlength. Thorax (Fig. 11 B): Scutum, scutellum mostly light to medium brown, contrasting with darker brown pleura; pale sclerites around base of wing. Posterior portion of dorsocentral row with 2 elongate setae situated somewhat lateral to one another. Prescutal suture elongate, thick, uninterrupted, extending to near dorsocentral row of setae. Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin not thick. Wing (Fig. 12 K): Apex of R 2 equal or slightly distal to apex of M 1. Anterior margin with differently, discretely pigmented scales (indicating anterior margin of midlength band), with midlength band, with poorly defined basal band; veins (other than costa and wing margin) with well-developed scales. Halter medium brown, equal to scutellum. Legs (Fig. 11 B): Medium brown, with about apical 0.3 of midfemur, basal 0.5 of midtibia, apical 0.5 of hind femur, all of hind tibia pale other than slightly darker pigmentation on apical 0.2 of hind tibia; with darker pigmentation not discrete. With only slender setae, lacking scales. Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Empodia thick. Abdomen (Fig. 13 J): Tergites 1–6 pale, sternites 1–6 light brown, segments 7–9 medium brown. Cercus medium brown. Immature stages. Unknown. DISTRIBUTION AND BIONOMICS: Corethrella gilva is known from Brunei at 30–110 m (Fig. 16 A). Habitats include lowland peat swamps (including standing water and very slow moving streams) and steep terrain in mature mixed dipterocarp forest (Fig. 1 A, as in Fig. 1 F). Females have been collected with frog-call traps and on calling males of three species of frogs (Table 7). TAXONOMIC DISCUSSION: Borkent (2008: 237) noted three species attracted to the taped call of Hyla gratiosa (LeConte) in Brunei and C. gilva was one of them (included as a paratype here). We are uncertain of the phylogenetic placement of C. gilva because it is known only as female adults and there were few synapomorphies present to interpret. Corethrella gilva has synapomorphies 21–23 and lacks synapomorphy 28. It has a banded wing but plain brown abdomen. It may belong to the drakensbergensis group but, as presently understood, the loss of wing pigmentation occurred before loss of abdominal pigmentation pattern, which would conflict with what is present in C. gilva. In addition, C. gilva has synapomorphy 40 (pedicel with strong setae), which would place it higher up in the phylogeny of the Corethrellidae. The discovery of the male of this species would help in its placement. TYPES: Holotype, female adult on microscope slide, labeled " HOLOTYPE Corethrella gilva Borkent and Grafe ", " 12 km S. Liang, Brunei, 15 -V- 2007; on Hylarana glandulosa; U. Grafe; 34 - 1 / 1, DNA extracted", " 34 - 1 / 1 231007 " (CNCI). Paratypes on microscope slides: 1 Ƥ, from type locality, but 10 -V- 2007 (CNCI); 1 Ƥ, from type locality but 15 -V- 2007 (CNCI); 1 Ƥ, from type locality but 23 -V- 2007 (CNCI); 1 Ƥ, from type locality but 9 -VI- 2007 (UBD); 7 Ƥ, from type locality but 11 -XII- 2007; 1 Ƥ, from type locality but 28 -V- 2007 (UBD); 1 Ƥ, 15 km S. Liang, Brunei, 40 m, 9 -V- 2007 (CNCI); 1 Ƥ, from previous locality but 22 -V- 2007 (CNCI); 1 Ƥ, lower Sungai Apan, 0.9 km E. headquarters, Ulu Temburong National Park, Brunei, 110 m, 9 -II- 2010 (CNCI); 1 Ƥ, tributary of Sungai Belalong, Temburong, Brunei, 110 m, 115 ° 09'E, 4 ° 33 'N, 7 -VIII- 2006 (CNCI). Paratypes on pins: 3 Ƥ, lower Sungai Apan, Ulu Temburong NP, Brunei, 110 m, 5 -II- 2010 (2 CNCI; 1 UBD); 1 Ƥ, from previous locality but 13 -VII- 2008 (CNCI); 2 Ƥ, Temburong, Ulu Temburong NP, Brunei, 7 -VII- 2008 (CNCI); 1 Ƥ, 12 km S. Liang, Brunei, 30 m, 11 -VI- 2007 (CNCI); 1 Ƥ, from previous locality but 11 -VI- 2011 (CNCI); 1 Ƥ, from previous locality but 28 -V- 2007 (CNCI); 2 Ƥ, from previous locality but 4 -VI- 2007 (CNCI; UBD); 1 Ƥ, from previous locality but 23 -V- 2007 (CNCI); 1 Ƥ, from previous locality but 9 -VI- 2007 (CNCI); 4 Ƥ, from previous locality but 15 -V- 2007 (CNCI); 1 Ƥ, from previous locality but 4 -XI- 2010 (CNCI); 1 Ƥ, 12.5 km S. Liang, Brunei, 33 m, 21 -II- 2009 (CNCI); 1 Ƥ, from previous locality but 3 -I- 2009 (CNCI); 1 Ƥ, 15 km S. Liang, Brunei, 40 m, 5 -VI- 2007 (CNCI); 2 Ƥ, from previous locality but 12 -V- 2007 (CNCI); 1 Ƥ, from previous locality but 22 -V- 2007 (CNCI); 1 Ƥ, 17 km N. Labi, Brunei, 40 m, 14 -V- 2007 (CNCI); 1 Ƥ, Sungai Ingei, 8.2 km S. Melilas, Brunei, 50 m, VII- 2010 (CNCI); 1 Ƥ, from previous locality but 10 -VII- 2010 (CNCI). DERIVATION OF SPECIFIC EPITHET: The name gilva (yellow) refers to the lightly pigmented hind tibiae of females of this species.Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 17-18, DOI: 10.5281/zenodo.21086
Corethrella (Corethrella) lutea Borkent & Grafe, new species
No full textCorethrella (Corethrella) lutea Borkent & Grafe, new species (Figs. 2 B, 4 B, 6 B, 8 B, 9 B, 12 B, 13 B, 15 A) DIAGNOSIS: Male adult. Unknown. Female adult. Only extant species of Corethrella in the Palaearctic, Oriental, Oceanian and northern Australasian Regions with the combination of a plain wing (Fig. 12 B), flagellomeres 1–4 relatively short (Fig. 8 B), scutum a lighter brown, contrasting with darker, mottled, pleura (Fig. 9 B) and midfemur relatively thick (Fig. 9 B) (compared to that of C. nippon Miyagi). Only extant species in Borneo with the combination of a plain wing (Fig. 12 B) and a thorax with patterned pigmentation (especially the pleura) (Fig. 9 B). DESCRIPTION: Male adult. Unknown. Female adult. Descriptive statistics: See Tables 1 –5. Head: Outline in anterior view somewhat circular (Fig. 6 B). Coronal suture elongate, extending ventrally past ommatidia. Two large setae on frons between ventromedial area of ommatidia. Antenna uniformly medium brown; pedicel without distinctive elongate, stout setae. Antennal flagellomeres as in Fig. 8 B; sensilla coeloconica on flagellomeres 1 –2, 9– 13, flagellomeres 1 with several, (10), 11–12 with 2 sensilla; flagellomere 13 with well-developed apical bifurcation. Clypeus (Fig. 6 B) somewhat square. Mandible with moderately sized, pointed teeth. Palpus (Fig. 6 B) brown; segment 3 swollen apically. Thorax (Fig. 9 B): Scutum mostly light to medium brown, contrasting with darker brown pleura; scutellum, mediotergite, most of pleura dark brown, katepisternum with pale area at midheight, posterior portion of anepimeron paler, pale sclerites around base of wing. Posterior portion of dorsocentral row with 2 elongate setae situated somewhat lateral to one another. Prescutal suture elongate, thick, uninterrupted, extending to near dorsocentral row of setae. Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin not thick. Wing (Fig. 12 B): Apex of R 2 slightly distal to apex of M 1. Plain, without pattern of pigmented veins and/or scales; veins (other than costa and wing margin) with slender scales. Halter light to medium brown, equal to or somewhat lighter than scutellum. Legs (Fig. 9 B): Light brown, forecoxa lighter than mid-, hind coxae, with about basal 0.4 of hind femur dark brown. With only slender setae, lacking scales. Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Empodia thick. Abdomen (Fig. 13 B): Pale or light yellow, tergites 2–6 with anterolateral light brown pigmentation, sternites medium brown, 3–5 somewhat darker anteriorly, segment 7 light to dark brown, segments 8–9 dark brown. Cercus light brown or pale. Immature stages. Unknown (but see below). DISTRIBUTION AND BIONOMICS: Corethrella lutea is known from Brunei and Sarawak, Malaysia from 30–200 m (Fig. 15 A). Habitats include lowland peat swamps (including standing water and very slow moving streams) and steep terrain in mature mixed dipterocarp forest (Figs. 1 A, D, E). They are relatively common in frog-call traps in Brunei (Grafe et al. 2008, as morphotype 2) and have been collected feeding on calling males of three species of frogs (Table 7). TAXONOMIC DISCUSSION: The seven females noted by Borkent (2008) in the taxonomic discussion of C. nippon, collected from a tributary of Sungai Belalong, Temburong district, Brunei, are paratypes of C. lutea here. Six females collected from the headquarters of Gunung Mulu National Park, Sarawak, Malaysia (113 ° 55 'E, 4 ° 23 'N) on 14 -XI- 2009 with a frog-call trap differed from other C. lutea in having only a single sensilla coeloconicum on each of flagellomeres 10–12 and relatively shorter flagellomeres 2–3 (only one slide-mounted). For the present we consider these as variants of C. lutea. The females of C. lutea are very similar to those of C. nippon but the latter have relatively thicker midfemora. Although the male of C. lutea is unknown, and therefore we do not know if this species shares the only synapomorphy for the group (unequal male midclaws), we consider it a member of the drakensbergensis species group (Fig. 19). It has synapomorphy 25 but lacks 26 and 27 (hence, an unresolved lineage along with C. nippon and C. harrisoni Freeman (Borkent, 2008)). TYPES: Holotype, female adult on microscope slide, labeled " HOLOTYPE Corethrella lutea Borkent and Grafe ", " 12 km S. Liang, Brunei, 30 m, 10 -V- 2007; on Hylarana glandulosa; U. Grafe; 18 - 2 / 3 ", " 18 - 2 / 3 261107 " (CNCI). Paratypes on microscope slides: 1 Ƥ labeled as for holotype but 18 - 2 /2, 231007 (CNCI); 3 Ƥ from type locality but 21 -IV-2007, 30-IV-2007, 15-V- 2007 (CNCI); 1 Ƥ, 15 km S. Liang, Brunei, 40 m, 9 -V- 2007 (CNCI); 1 Ƥ, 17 km N. Labi, Brunei, 40 m, 6 -IV- 2009 (UBD); 1 Ƥ, from previous locality but 8 -IV- 2009 (CNCI); 7 Ƥ, tributary of Sungai Belalong, Temburong district, Brunei, 110 m, 115 ° 09'E, 4 ° 33 'N, 50 m, 7 -VIII- 2006 (6 CNCI; 1 UBD); 1 Ƥ, Gunung Mulu NP, Sarawak, Malaysia, 30 m, 14 -XI- 2008 (CNCI); 5 Ƥ, Matang, Sarawak, Malaysia, 200 m, 4 -IX- 2008 (CNCI). Paratypes on pins: 1 Ƥ, Kuala Belalong Field Studies Centre, Brunei, 50 m, 17 -XI- 2010 (CNCI); 1 Ƥ, Belalong Münd (estuary), Ulu Temburong NP, Brunei, 4 -VII- 2008 (CNCI); 1 Ƥ, lower Sungai Apan, Ulu Temburong NP, Brunei, 110 m, 8 -VII- 2008 (CNCI); 1 Ƥ, from previous locality but 13 -VII- 2008 (CNCI); 8 Ƥ, 12.5 km S. Liang, Brunei, 33 m, 28 -I- 2011 (6, CNCI; 2 UBD); 12 Ƥ, from previous locality but 2 -IV- 2009 (CNCI); 6 Ƥ, 20 km N. Labi, Luagan Lalak Forestry Rec. Site, Brunei, 6 -XI- 2010 (CNCI); 2 Ƥ, Sungai Ingei, 8.2 km S. Melilas, Brunei, 50 m, 9 -VII- 2010 (CNCI); 11 Ƥ, from previous locality but 12 -VII- 2010 (CNCI); 4 Ƥ, from previous locality but VII- 2010 (CNCI); 2 Ƥ, from previous locality but 10 -VII- 2010 (CNCI); 5 Ƥ, Gunung Mulu NP, Sarawak, Malaysia, 30 m, 14 -XI- 2008 (CNCI); 3 Ƥ, from previous locality but 26 -XI- 2009 (CNCI). DERIVATION OF SPECIFIC EPITHET: The name lutea (yellow) refers to the overall yellowish appearance of the females of this species.Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 8-9, DOI: 10.5281/zenodo.21086
Corethrella (Corethrella) calathicola Edwards
No full textCorethrella (Corethrella) calathicola Edwards (Figs. 5 B, 7 A, F, H, 8 H, 10 C, 11 C, 12 H, 13 G, 18) Corethrella calathicola Edwards 1930: 529. Type locality: Singapore, Malaysia. Lectotype male (BMNH). Mogi and Yong 1992: 180; Borkent 2008: 145. DIAGNOSIS: Male and female adults: Only extant species of Corethrella in the Oriental, Australasian, and Oceanic Regions with the combination of a pale or light brown palpus (contrasting with dark brown clypeus) (Fig. 7 A), patterned wing (with a midlength band, darker where R 1 joins the costa) (Fig. 12 H), uniformly dark brown midfemur, hind tibia with dark pigmentation at base and apex (Fig. 10 C), and with abdominal tergites 2–6 uniformly brown (Fig. 13 G). Only extant species in Borneo with the combination of well-developed scales on midfemur (Fig. 11 C); also only species with palpus entirely pale and contrasting with a dark clypeus (Fig. 7 A) and with distinct dark pigmentation at both base and apex of hind tibia (Fig. 10 C). DESCRIPTION: This species was redescribed by Borkent (2008), but see Taxonomic Discussion below. DISTRIBUTION AND BIONOMICS: Corethrella calathicola is known from Singapore, Malaysia, Indonesia and Brunei (Fig. 18) at altitudes ranging from 1–1083 m. In Borneo, it has been recorded only from Brunei, Sarawak, Malaysia and eastern Kalimantan, Indonesia. This is the only species of Corethrella recorded from and restricted to species of Nepenthes. Its biology has been summarized by Borkent (2008). Habitats include lowland and mid-elevation peat swamp in Brunei (Fig. 1 A). Previously recorded from N. ampullaria (Jack), there were earlier records of an unnamed Corethrella species from N. bicalcarata (Hook). Here we confirm that C. calathicola occurs in N. bicalcarata in Brunei as well as in N. veitchii (Hook) in the Bario highlands of Sarawak. We have found it commonly in N. ampullaria in Brunei as well (Fig. 1 B). Immatures of C. calathicola were found in these three species of Nepenthes only when they were on or near the ground (both authors, pers. obs.; Miyagi pers. comm.). We reared nine males and five females from 14 larvae collected from N. ampullaria and N. bicalcarata. Aside from rearings, we examined female adults collected with frog-call traps in Brunei and Lanjak Entimau, Sarawak, as well as feeding on a calling male of one species of frog (Table 7). We examined the specimens reared from N. ampullaria at Matang, Sarawak, Malaysia noted by Miyagi et al. (2009). TAXONOMIC DISCUSSION: Borkent (2008) described the pigmentation of the male abdomen as having "segments 1 –7, 9 light to medium brown, segment 8 pale". Some of our fresh slide-mounted material matches this description but at least some fresh material in alcohol or pinned (from Brunei, some from Bario, Sarawak) had tergites 2–7 pale and contrasting with a dark tergite 1 and darker sternites. Upon treatment with KOH, some specimens with lighter tergites had tergites and sternites appeared uniformly medium brown but in others, the lighter tergites were discernible. We are puzzled as to why there was this difference between alcohol and slidemounted material when this is not the case with other Corethrella specimens studied by Borkent (2008). The abdomens of all females were entirely medium to dark brown, so that there is a clear sexual dimorphism in pigmentation of the abdomen in at least some members of C. calathicola. More study and specimens are needed to further interpret this variation. There was significant variation in the relative sizes of flagellomeres 1–3 in the female C. calathicola we examined. In specimens from Brunei they were relatively short (Fig. 7 F), compared to those from Matang, Sarawak (Fig. 7 H) and other sites recorded and illustrated by Borkent (2008, fig. 30 H). Borkent (2008) designated the lectotype for C. calathicola and listed the paralectotypes known to him. There are an additional two larvae that should be considered paralectotype specimens on separate slides in the BMNH. Both are from Singapore and taken from a "pitcher plant" (Zoe Adams, pers. comm.). Corethrella calathicola belongs to the appendiculata species group, a group that otherwise includes only New World species occupying treeholes and is the sister species of the Neotropical C. melanica Lane and Aitken (Borkent 2008). MATERIAL EXAMINED: In addition to the material listed by Borkent (2008), we examined the following on microscope slides: 1 3, 12 km S. Liang, Brunei, 30 m, 9 -VIII- 2009, from Nepenthes ampullaria (CNCI); 1 Ƥ, from previous locality but 18 -V- 2007 (CNCI); 2 Ƥ, 15 km S. Liang, Brunei, 40 m, 12 -V- 2007 (1, CNCI; 1, UBD); 2 Ƥ, from previous locality but 5 -VI- 2207 (CNCI); 3 Ƥ from previous locality but 22 -V- 2207 (CNCI); 2 Ƥ, 17 km S. Liang, Brunei, 40 m, 6 -IV- 2009 (1, CNCI; 1, UBD); 2 3, Bario, Sarawak, Malaysia, 1060 m, 6 -IX- 2009, reared from Nepenthes veitchii (CNCI); 1 3, from previous locality but 17 -IX- 2006 (CNCI); 1 3, Pa Umor, 5 km E. Bario, 03° 44.118 ' N 115 ° 30.404 'E, 1083 m, Sarawak, Malaysia, 17 -IX- 2006, reared from 'pitcher plant' (CNCI); 2 3, 2 Ƥ, Matang, Sarawak, Malaysia, 200 m, 28 -VIII- 2006, reared from 'pitcher plant' (1 3, 2 Ƥ, CNCI; 1 3, UBD). Pinned specimens: 1 Ƥ, 12.5 km S. Liang, Brunei, 33 m, 10 -I- 2009 (CNCI); 2 3, 12 km S. Liang, Brunei, 30 m, 9 - VIII- 2009 (CNCI); 1 3, 2 Ƥ, previous locality but 23 -VII- 2009 (2 Ƥ, CNCI; 1 3, UBD); 1 3, 1 Ƥ, previous locality but 20 -VII- 2009 (CNCI); 2 3, from previous locality but 30 -VII- 2009 (1, CNCI; 1 UBD); 2 Ƥ, from previous locality but 9 -VIII- 2009 (UBD); 1 Ƥ, from previous locality but 15 -V- 2007 (CNCI); 2 3, from previous locality but 25 -VII- 2009 (CNCI); 4 Ƥ, 15 km S. Liang, Brunei, 40 m, 22 -V- 2007 (CNCI); 1 Ƥ, 17 km S. Liang, Brunei, 40 m, 17 -V- 2007 (CNCI); 1 Ƥ, Lanjak Entimau, Sarawak, Malaysia, 80 m, 28 -II- 2011 (SMKM).Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 14-15, DOI: 10.5281/zenodo.21086
Corethrella
No full textKey to the species of Corethrella of Borneo This key will successfully identify material in either alcohol or slide-mounted specimens. Material in good condition, retaining their setae and scales, are significantly easier to identify than rubbed specimens. Nevertheless, pigmentation patterns, although not as clear, are yet present in the underlying cuticle of damaged material. Identification of the antennal sensilla coeloconica requires slide mounting but we have added other features to the key so they are not vital to identification. Readers should be aware that most of our material came from Brunei or nearby localities in Sarawak. It is very likely that undescribed species are yet to be found in Borneo, especially from elsewhere on the island. As such, slide mounting of at least representative material will increase the likelihood of accurate identification of the species. The key below is based primarily on females because the males of only two of the 11 species are known, namely those of C. tigrina and C. calathicola. However, our use of colour patterns, likely shared by males and females, suggests that the key will work with both sexes. The pigmentation of the abdomen is described below based on cleared, slide-mounted specimens. At times, material in alcohol can be confusing because of underlying pigmentation of body tissues; in such instances, special care should be taken to study only the tergites and/or sternites. Rubbed, pinned specimens of C. gilva, which otherwise have a midlength wing band (Fig. 12 K), may appear to have no wing pigmentation. Such specimens would run to couple 2, but the combination of a scutum lighter than the dark pleura and pale abdominal tergites 2–6 (Fig. 5 E) will correctly identify these. 1. Wing plain, without discrete pattern of pigmented scales (Figs. 12 B, E, F, I)....................................... 2 - Wing with at least some pattern of pigmented scales (anterior margin always with at least one distinct and discrete band of dark scales) (Figs. 12 A, C, D, G, H, J, K)....................................................................... 5 2 (1). Wing yellow (Fig. 4 B); scutum yellow (Fig. 4 B); katepisternum with at least some pale cuticle, contrasting with dark posterior anepisternum (Fig. 9 B); base of hind femur dark brown, strongly contrasting with yellow base of midfemur (Fig. 9 B)... lutea (Brunei, Malaysia (Sarawak)) - Wing brown (Figs. 4 E, F, 5 C); scutum brown; pleura (including katepisternum and posterior anepisternum) uniformly dark brown (scutum and pleura not equal in some) (Figs. 4 E, 10 A, D); bases of mid- and hind femora equally pigmented, medium to dark brown (Figs. 4 E, 10 A, D)........................................................................... 3 3 (2). Scutum more lightly pigmented than pleura (Fig. 10 A); abdominal tergites and sternites uniformly pigmented or at least each of abdominal tergites 2–5 more lightly pigmented posteriorly (Fig. 13 E)....................................... mitra (Brunei, Malaysia (Sarawak, Sabah)) - Scutum as dark brown as pleura (Figs. 4 E, 5 C, 10 D); abdominal tergites uniformly brown (Figs. 4 E, 13 H)............... 4 4 (3). Clypeus with about 23 setae; wing length 1.30 mm (known from one specimen).............................. brunnea (Malaysia (Sarawak)) - Clypeus with 2–5 setae; wing length of Bornean specimens less than 1.07 mm.............................. pauciseta (Brunei, Malaysia (Sarawak), Papua New Guinea) 5 (1). Wing with a distinct subapical band (Figs. 12 A, J)............................................................ 6 - Wing with single midlength band represented by at least dark pigmentation on anterior margin of wing (Fig. 12 C, D, G, H, K); with, at most, slightly darker pigmentation at very apex of veins R 2, R 3, and R 4 + 5 (Figs. 12 H).......................... 7 6 (5). Thorax uniformly brown (alcohol specimens with scutum with three narrow longitudinal vittae) (Fig. 11 A); basal 0.7 of forefemur, all of midfemur and about basal half of hind femur similarly and uniformly brown (pigmentation not discrete) (Fig. 11 A); abdomen uniformly brown (Fig. 13 I)................................................................ bicincta (Brunei, Malaysia (Sarawak)) Thorax with patterned pigmentation (Fig. 9 A); forefemur with basal, midlength and apical darker pigmentation, mid- and hind femur with narrow basal and apical bands of dark pigmentation (pigmentation discrete) (Fig. 9 A); at least each of abdominal tergites 2–6 more lightly pigmented posteriorly (Fig. 13 A)................................................ tigrina (Brunei, Malaysia (Sarawak)) 7 (5). Hind tibia with distinct basal and apical bands of dark pigmentation (Fig. 10 C); palpus pale, contrasting with dark brown clypeus (Fig. 7 A); midfemur with well-developed scales (Fig. 11 C); wing with midlength band somewhat scattered, with scales on R 4 + 5 pale (so that the dark scales on C, R 1 and R 2 + 3 appear as a separate anterior darker spot), and with posterior portion of band situated separately at apex of CuA 2 (Fig. 12 H)............................................. calathicola (Singapore, Malaysia (Pahang, Sarawak), Indonesia (East Kalimantan), Brunei) - Hind tibia with, at most, some darker pigmentation at very base and at apex (Fig. 10 B); palpus entirely medium brown or with segments 1 and 2 brown and contrasting with pale segments 3–5 (base of segment 3 may be brown in some) (Figs. 6 C, D, F, 7 D); midfemur without scales; wing with midlength band forming a single anterior to posterior band of uniform pigmentation (Figs. 12 C, D, G, K).................................................................................... 8 8 (7). Palpus with segments 1 and 2 (in some the very base of 3) dark brown, contrasting with light brown or pale segments 3 and 4 (5 grading apically to medium brown) (Figs. 6 C, D); at least each of abdominal tergites 2–6 with darker pigmentation restricted to lateral margins, sternites 2–6 more lightly pigmented posteriorly (Figs 13 C, D).......................... 9 - Palpus with all segments uniformly medium brown (Figs. 6 F, 7 D); abdominal tergites 2–6 uniformly pigmented, sternites 2–6 uniformly pigmented (in some with tergites lighter than sternites) (Figs. 13 F, J)................................... 10 9 (8). Flagellomeres 1–4 short, each notably shorter than flagellomere 5 (Fig. 8 C)............................ nanoantennalis (Brunei, Malaysia (Sarawak)) - Flagellomeres 1–4 elongate, similar in length to flagellomere 5 (Fig. 8 D)................................. bipigmenta (Brunei) 10 (8). Hind tibia with dark pigment apically (Fig. 10 B); 0–1 clypeal setae; flagellomeres 1, (sometimes 8), 9–13 with sensilla coeloconica and 9–12 (sometimes 13) with 2 sensilla coeloconica (1 with a few sensilla coeloconica).................. unizona (Brunei, Malaysia (Sarawak)) - Hind tibia with, at most, lightly pigmented apex (Fig. 11 B); with 13–25 clypeal setae; flagellomeres 1, 9– 13 with sensilla coeloconica and only flagellomere 1 with more than one sensilla coeloconicum.................................... gilva (Brunei)Published as part of Borkent, Art & Grafe, Ulmar, 2012, The Frog-Biting Midges of Borneo — From Two to Eleven Species (Corethrellidae: Diptera), pp. 1-45 in Zootaxa 3279 on pages 3-4, DOI: 10.5281/zenodo.21086
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