Corresponding author: Søren Achim Nielsen (
Academic editor: Vladimir Blagoderov
In the context of a major monitoring program of
Three species are described as new to science based on both morphological and molecular data:
Following the outbreak of bluetongue disease in 2007–2008, a major entomological monitoring program of
Biting midges were collected in 2007–2008 in Sweden (
For documentation of the identification, the head, wings and the posterior abdominal segments were removed from the female individuals and slide mounted, and the remaining parts of the animal were processed for DNA analysis as described by
The holotypes and paratypes of the three new species are deposited in the collection of the Natural History Museum of Denmark, University of Copenhagen (ZMUC).
Female: Eyes bare and contiguous dorsally (Fig.
Wing length 1,641 ± 10 μm (Fig.
Two normal-sized spermathecae, ovoid, of almost equal size, with a short neck and a third rudimentary one (Fig.
Male: Unknown.
Named as a tribute to Boy Overgaard Nielsen, an outstanding Danish entomologist and current emeritus at Aarhus University.
The length divided by the width of the first flagellomere is significantly higher in
Female: Eyes bare and contiguous dorsally (Fig.
Wing length 1,339 ± 33 μm (Fig.
Spermathecae not observed.
Male: Unknown.
The species epithet refers to the name of the major Danish island Sjælland (Latin = Selandia; English = "Zealand" or more rarely "Sealand"), where the type series was collected.
Female: Eyes bare and contiguous dorsally (Fig.
Wing length 1,423 ± 39 μm (Fig.
Two functional spermathecae, ovoid, of almost equal size, with a short neck; a third rudimentary one present (Fig.
Male: Unknown.
The species epithet refers to the municipality Kalix in northern Sweden, in which the type series was collected.
1 | Wing with a spot in cell cu |
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– | Wing without a spot in cell cu |
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2 | Third segment of maxillary palp about as long as or longer than the second segment (P3/P2 ≥ 0.94). Wing with the dark hour-glass mark in cell r3 broadest above the longitudinal fold above vein M1 |
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– | Third segment of maxillary palp shorter than the second segment (P3/P2 ≤ 0.93). Wing with the dark hour-glass mark in cell r3 broadest above the longitudinal fold above vein M1 or broadest at or below the fold above M1 |
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3 | Wing darkened around the entire length of vein M1 |
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– | Wing darkened for some part of vein M1 |
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4 | Wing with two dark marks in cell m1. The dark hour-glass mark in cell r3 is broad and roughly square in outline. Third segment of maxillary palp longer than second segment (P3/P2 > 1.01); PR (palp ratio, i.e., length/width of third segment) < 2.8. Fronto-vertex/ocellus ratio < 0.78 | |
– | Wing with only one dark mark in cell m1. The dark hour-glass mark in cell r3 is not broad and square in outline. Third segment of maxillary palp at most as long as second segment (P3/P2 ≤ 1.0); PR > 3.0. Fronto-vertex/ocellus ratio > 0.87 |
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5 | Wing with small pale spots at the tip of veins M1, M2, and Cu1. AR (antennal ratio, i.e., length of flagellomeres 9–13 divided by the length of flagellomeres 1–8) > 1.08. Ratio length/width of first flagellomere < 1.6 |
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– | Wing without spots at the tip of veins M1, M2 and Cu1. AR < 1.08. Ratio length/width of first flagellomere > 1.7 |
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6 | Wing with the dark hour-glass mark in cell r3 broadest above the longitudinal fold above M1 or hour-glass mark with continuous outline and of equal width above and at the longitudinal fold above vein M1 |
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– | Wing with the dark hour-glass mark in cell r3 broadest at or below the longitudinal fold above vein M1 |
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7 | Wing with the dark hour-glass mark in r3 broadest above the longitudinal fold above vein M1. The dark areas in wings surround the entire length of the veins M1 and M2. Mandibular teeth 12–14, maxillary teeth 12–17. Number of antennal sensilla coeloconica 9–12 |
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– | Wing with the dark hour-glass mark in cell r3 with continuous outline and equal widths above the longitudinal fold and at the fold above vein M1. The dark areas in wings do not surround veins M1 and M2. Mandibular teeth 16–20, maxillary teeth 19–21. Number of antennal sensilla coeloconica 12–19 |
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8 | Small species (wing length < 1400 µm). Wing markings are faint but sharply defined. Third segment of maxillary palp rhomboid. AR (antennal ratio, i.e., length of flagellomeres 9–13 divided by the length of flagellomeres 1–8) < 1.09. Head/proboscis ratio > 1.26. Number of antennal sensilla coeloconica 7–12 |
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– | Large species (wing length > 1600 µm). Wing with extensive and vaguely defined dark markings. Third segment of maxillary palp cigar-shaped. AR > 1.09. Head/proboscis ratio < 1.24. Number of antennal sensilla coeloconica 12–20 |
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9 | Wing with distinct dark markings; the dark hour-glass mark in the middle of cell r3 broadest above the longitudinal fold above the longitudinal fold above cell M1 |
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– | Wing with vaguely defined markings; the dark hour-glass mark in cell r3 broadest at or below the fold above M1 |
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10 | PR (palp ratio, i.e., length/width of third segment) < 3.5. Fronto-vertex/ocellus ratio > 1.2 |
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– | PR > 3.5. Fronto-vertex/ocellus ratio < 1.0 |
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11 | Small species (wing length < 1400 µm). Third segment of maxillary palp rhomboid. AR (antennal ratio, i.e., length of flagellomeres 9–13 divided by the length of flagellomeres 1–8) < 1.09. Head/proboscis ratio > 1.26. Number of antennal sensilla coeloconica 7–12 |
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– | Large species (wing length > 1600 µm). Third segment of maxillary palp cigar-shaped. AR > 1.09. Head/proboscis ratio < 1.24. Number of antennal sensilla coeloconica 12–20 |
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The problems in constructing reliable keys to adults of the European species of
Note that we are following
Relevant comparisons for separating the three new species from their morphologically most similar Scandinavian congeners are given in Table
The recent arrival of bluetongue virus in northern and western Europe (
The European fauna of
Revisionary taxonomy of European
The molecular data from the three species named in the present paper were analysed by
It is noteworthy that the three new species were collected at single locations or from a few locations in close proximity in spite of a very large sampling. These species are most likely more widely distributed, as are the majority of the well-known biting midge species, and what may look like a restricted geographical occurrence may be due to either a patchy distribution or a very short adult flying period.
We are very grateful for constructive comments from Dr. Art Borkent, Research Associate of the Royal British Columbia Museum, American Museum of Natural History, and Instituto Nacional de Biodiversidad as well as from Dr. Bruno Mathieu, Institut de Parasitologie et de Pathologie Tropicale, Faculté de Médecine de Strasbourg. The project was supported by the Danish Ministry of Food, Agriculture and Fisheries by grant no. 3304FVFP077 to MK.
SAN and MK conceived the study. SAN identified
Vertex. Scale = 50 µm.
Antennal pedicellus and first two flagellomeres. Scale = 50 µm.
Left palp, dorsal view. Scale = 50 µm.
Wing, brightfield photo. Scale = 500 µm.
Wing, darkfield photo. Scale = 500 µm.
Spermathecae. Scale = 50 µm
Vertex. Scale = 50 µm.
Antennal pedicellus and first three flagellomeres. Scale = 50 µm.
Left palp, dorsal view. Scale = 50 µm.
Wing, brightfield photo. Scale = 500 µm.
Wing, darkfield photo. Scale = 500 µm.
Vertex. Scale = 50 µm.
Antennal pedicellus and first three flagellomeres. Scale = 50 µm.
Left palp, dorsal view. Scale = 50 µm.
Wing, brightfield photo. Scale = 500 µm.
Wing, darkfield photo. Scale = 500 µm.
Spermathecae. Scale = 50 µm.
Cladogram based on COI-data showing clusters of species of
Pairwise morphometric comparisons between three new species of
1 - Species; 2 - Flagellum, length (μm); 3 - Antennal ratio (AR: length of flagellomeres 9–13 divided by length of flagellomeres 1–8); 4 - First flagellomere, length/width; 5 - Maxillary palp, length (μm); 6 - Maxillary palp ratio PR (length/width of third palp segment); 7 - Maxillary palp ratio P3/P2 (length of third maxillary palp segment divided by length of second); 8 - Length of wing (μm); 9 - Spermatheca ratio S/R; 10 - Head/proboscis ratio; 11 - Mandibular teeth; 12 - Maxillary teeth; 13 - Ratio M/M; 14 - Fronto-vertex /ocellus; 15 - Antennal sensilla coeloconica.
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742± 26 | 1.09± 0,03 | 1.52± 0.07 | 255.6± 14.3 | 2.9± 0.2 | 0.84± 0.05 | 1626± 67 | 121± 0.08 | 1.19± 0.04 | 16.7± 1.2 | 19.5± 1.1 | 1.17± 0.08 | 1.2± 0.3 | 15.47± 1.97 |
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746± 44 | 1.03± 0.04 | 1.78± 0.07 | 245.1± 17.7 | 2.9± 0.3 | 1.04± 0.11 | 1641± 10 | 1.05± 0.3 | 1.29± 0.07 | 15.1± 1.2 | 17.1± 1.4 | 1.13± 0.10 | 1.5± 0.2 | 14.71± 1.26 |
Divergence of COI sequences = 16.5% | ||||||||||||||
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591± 45 | 1.04± 0.05 | 1.53± 0.15 | 193.6± 18.7 | 2.6± 0.2 | 1.11± 0.10 | 1291± 12 | 1.04± 0.02 | 1.32± 0.13 | 13.4± 1.4 | 15.9± 1.5 | 1.18± 0.20 | 0.3± 0.3 | 7.70± 0.67 |
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616± 10 | 1.12± 0.04 | 1.56± 0.11 | 216.8± 8.6 | 3.2± 0.3 | 0.96± 0.06 | 1339± 33 | ND | 1.16± 0.06 | 15.0± 1.0 | 19.6± 1.5 | 1.31± 0.12 | 1.5± 0.8 | 12.29± 0.95 |
Divergence of COI sequences = 16.2% | ||||||||||||||
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591± 45 | 1.04± 0.05 | 1.53± 0.15 | 193.6± 18.7 | 2.6± 0.2 | 1.11± 0.10 | 1291± 12 | 1.04± 0.02 | 1.32± 0.13 | 13.4± 1.4 | 15.9± 1.5 | 1.18± 0.20 | 0.3± 0.3 | 7.70± 0.67 |
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646± 14 | 1.13± 0.04 | 1.46± 0.06 | 212.4± 4.0 | 2.9± 0.2 | 0.87± 0.08 | 1423± 39 | 1.17± 0.08 | 1.29± 0.07 | 12.8± 0.6 | 14.9± 1.5 | 1.17± 0.12 | 1.2± 0.3 | 10.70± 0.82 |
Divergence of COI sequences = 15.6% | ||||||||||||||
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616± 10 | 1.12± 0.04 | 1.56± 0.11 | 216.8± 8.6 | 3.2± 0.3 | 0.96± 0.06 | 1339± 33 | ND | 1.16± 0.06 | 15.0± 1.0 | 19.6± 1.5 | 1.31± 0.12 | 1.5± 0.8 | 12.29± 0.95 |
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646± 14 | 1.13± 0.04 | 1.46± 0.06 | 212.4± 4.0 | 2.9± 0.2 | 0.87± 0.08 | 1423± 39 | 1.17± 0.08 | 1.29± 0.07 | 12.8± 0.6 | 14.9± 1.5 | 1.17± 0.12 | 1.2± 0.3 | 10.70± 0.82 |
Divergence of COI sequences = 5.9% |