Research Article |
Corresponding author: Emma Wahlberg ( emma.wahlberg@nrm.se ) Academic editor: Martin Husemann
© 2020 Caroline Rhodén, Emma Wahlberg.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rhodén C, Wahlberg E (2020) The phylogeny of Empis and Rhamphomyia (Diptera, Empididae) investigated using UCEs including an over 150 years old museum specimen. Evolutionary Systematics 4(1): 21-33. https://doi.org/10.3897/evolsyst.4.49537
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The genera Empis Linneus, 1758 and Rhamphomyia Meigen, 1822 (Empidoidea, Empididae Latreille, 1809) are two large genera of flies commonly named dagger flies. They are widely distributed in the world with most species described from the Palearctic Region. Empis comprises about 810 described species and Rhamphomyia comprises about 610 described species, together they represent one third of the known species diversity in Empididae. Two recent studies on the phylogeny of the two genera using Sanger sequencing on a few genetic markers, did not support monophyly of them. In this study high throughput sequencing of target enriched molecular data of ultraconserved elements or UCEs was used to investigate the phylogenetic relationships of included representatives of the genera. This method has proven useful on old and dry museum specimens with high amounts of degraded DNA, which was also tested herein. For this purpose, a commercially synthesized bait kit has previously been developed for Diptera which this study was the first one to test. Three out of nine old and dry museum specimens were successfully sequenced, one with an age of at least 154 years. Higher DNA concentration yielded a greater number of reads. Analyses conducted in the study confirmed that both Empis and Rhamphomyia are non-monophyletic.
Entomology, high throughput sequencing, next generation sequencing, systematics, taxonomy, target enrichment, UCE
The family Empididae Latreille, in the superfamily Empidoidea Latreille, commonly known as dagger flies, is a family within Diptera consisting of around 3 051 known species in the world (
Previous studies have indicated that Empis and Rhamphomyia are non-monophyletic (
Former studies on the phylogeny of Empis and Rhamphomyia focused on either morphological or molecular data from traditional Sanger sequencing on only few genetic markers (
A total of 48 taxa, 23 of Empis and 21 of Rhamphomyia, representing twelve subgenera of Empis and nine of Rhamphomyia were sampled from the collections at the Swedish Museum of Natural History in Stockholm (
List of specimens including voucher number, collection data, geographic origin and accession number of COI barcode sequence in GenBank (if available). SMTP referring to Swedish Malaise Trap Project (
Taxon | Voucher | Locality and collector | Collection date | GenBank accession number (COI) |
---|---|---|---|---|
Chelipoda sp. | AH7C | NEW CALEDONIA: Province Sud, Platou de Dogny, source of Dogny River, about 500 m SE summit of Platou de Dogny, –21.613588; 165.883401. Leg. K. A. Johanson. | 25 Nov.–16 Dec. 2003 | MN868983 |
E. (Coptophlebia) albinervis Meigen, 1822 | AA9C | SWEDEN: Öland, Mörbylånga kommun, Gamla Skogsby (Kalkstad), 56.616700; 16.507617. Leg. SMTP. | 3–20 Jul. 2006 | – |
E. (Empis) bicuspidata Collin, 1927 | AB1C | SWEDEN: Torne lappmark, Kiruna kommun, Abisko nationalpark, Nuolja, 68.359492; 18.719197. Leg. SMTP. | 26 Jun.–15 Jul. 2006 | MN868986 |
E. (Platyptera) borealis Linnaeus, 1758 | AA8C | SWEDEN: Torne lappmark, Kiruna kommun, Abisko nationalpark, Nuolja, 68.359492; 18.719197. Leg. SMTP. | 11–17 Aug. 2005 | MN868968 |
E. (Leptempis) grisea Fallén, 1816 Old specimen | AE3C | SWEDEN: Skåne. Leg. C. H. Boheman. | 1852 or 1865 | – |
E. (Coptophlebia) hyalipennis Fallén, 1816 Old specimen | AE5C | SWEDEN: Skåne. Leg. C. H. Boheman. | 1852 or 1865 | – |
E. (Planempis) latro Frey, 1953 | AF4C | JAPAN: Hyōgo Prefecture, Arimafuji Park, 34.9194; 135.2356. Leg. Japan 2011 Exp. | 12 Apr. 2011 | MN868961 |
E. (Kritempis) livida Linnaeus, 1758 | AA3C | SWEDEN: Östergötland, Ödeshögs kommun, Omberg, bokskogsreservatet (beech forest), 58.297183; 14.634817. Leg. SMTP. | 5–19 Jul. 2005 | MN868964 |
E. (Anacrostichus) lucida Zetterstedt, 1838 | AA4C | SWEDEN: Torne lappmark, Kiruna kommun, Abisko nationalpark, Nuolja, 68.359492; 18.719197. Leg. SMTP. | 1–13 Jul. 2005 | MN868972 |
E. (Leptempis) nigricans Meigen, 1804 Old specimen | AD9C | SWEDEN: Skåne. Leg. P. F. Wahlberg. | 1846 | – |
E. (Lissempis) nigritarsis Meigen, 1804 | AD1C | SWEDEN: Öland, Mörbylånga, Gamla Skogsby (Kalkstad), mixed deciduous forest. Leg. M. & C. Jaschhof. | 1–25 May 2014 | MN868979 |
E. (Anacrostichus) nitida Meigen, 1804 Old specimen | AE1C | SWEDEN: Jämtland, Undersåker. Leg. O. Ringdahl. | 17 Jun. 1914 | – |
E. (Empis) nuntia Meigen, 1838 | AB5C | SWEDEN: Öland, Mörbylånga, Lilla Vickleby Lunds NR, old oak forest. Leg. M. & C. Jaschhof. | 1–27 May 2014 | MN868989 |
E. (Euempis) picipes Meigen, 1804 Old specimen | AE2C | SWEDEN:Södermanland, Stormossen. Leg. A. Orbe. | 19 Apr. 1991 | – |
E. (Empis) planetica Collin, 1927 | AA7C | SWEDEN: Uppland, Knivsta kommun, Rickebasta alsumpskog, western part, 59.734350; 17.720417. Leg. SMTP. | 18 Jun.–6 Jul. 2003 | MN868966 |
E. (Euempis) tessellata Fabricius, 1794 | AA6C | SWEDEN: Småland, Gränna kommun, Lönnemålen, 58.048917; 14.573033. Leg. SMTP. | 15 Jun.–1 Jul. 2005 | MN868980 |
E. (Xanthempis) trigramma Wiedemann, 1822 | AB2C | SWEDEN: Öland, Mörbylånga, Gamla Skogsby (Kalkstad), mixed deciduous forest. Leg. M. & C. Jaschhof. | 1–25 May 2014 | MN868960 |
E. (Xanthempis) univittata Loew, 1867 | AB3C | SWEDEN: Öland, Mörbylånga, Kalkstad NR, mixed deciduous forest. Leg. M. & C. Jaschhof. | 27 May–27 Jun. 2014 | MN868951 |
Empis sp. 1 | AD5C | RUSSIA: Chukota Autonomous Okrug, Chaunsky, Ajon Island, 70 m from base camp, 69.5840; 168.6955. Leg. P. Mortensen. | 11–24 Jul. 2015 | MN868973 |
Empis sp. 2 | AD7C | JAPAN: Ehime Prefecture, Matsuyama–shi, Ehime University Forest. Leg. Japan 2011 Exp. | 2 May 2011 | MN868957 |
Empis sp. 3 | AF2C | GREECE: East Macedonia, Paranesti by Nestos River, east bank of river north of road to Drama. Leg. P. Lindskog & B. Viklund. | 3–5 May 1995 | MN868970 |
Empis sp. 5 | AG9C | CHILE: Region de los Lagos, Isla Grande de Chiloé Rio Melilebú, along road between Tebuhueico and Hullinco, 5.4 km S crosspoint to Curaco, river, 100 m upstream bridge, –42.7181; –73.8965. Leg. K. A. Johanson. | 6 Jan. 2006 | MN868969 |
Empis sp. 6 | AH2C | NEW CALEDONIA: Province Sud, Platou de Dogny, source of Dogny River, about 500 m SE summit of Platou de Dogny, –21.613588; 165.883401. Leg. K. A. Johanson. | 25 Nov.–16 Dec. 2003 | |
Hilara cornicula | AG7C | SWEDEN: Uppland, Älvkarleby kommun, Älvkarleby kommun, Båtfors, between Milsten and Båtforstorpet, 60.46065; 17.317817. Leg. SMTP. | 27 Jun.–1 Jul. 2004 | MN868950 |
Hilara flavipes | AG8C | SWEDEN: Ångermanland, Örnsköldsviks kommun, Skuleskogen, Långrå, 63.088717; 18.498383. Leg. SMTP. | 5–25 Jul. 2004 | MN868984 |
Phyllodromia melanocephala | AH5C | SWEDEN: Hälsingland, Hudiksvalls kommun, Stensjön–Lomtjärn, Stensjön, Marsh pine wood close to bog, 62.140333; 16.286100. Leg. SMTP. | 8–23 Apr. 2005 | – |
R. (Megacyttarus) anomalina Zetterstedt, 1838 | AC5C | SWEDEN: Ly, Sorsele kommun, Ammarnäs, Vindelfjällens naturreservat, Tjulträsklaspen. 65.966783; 16.060500. Leg. SMTP. | 28 Jun.–15 Jul. 2004 | – |
R. (Megacyttarus) crassirostris (Fallén, 1816) | AC2C | SWEDEN: Lycksele lappmark, Sorsele kommun, Ammarnäs, Vindelfjällens naturreservat, Tjulträsklaspen, 65.966783; 16.060500. Leg. SMTP. | 28 Jun.–15 Jul. 2004 | MN868975 |
R. (Pararhamphomyia) curvula Frey, 1913 | AC9C | SWEDEN: Lycksele lappmark, Sorsele kommun, Ammarnäs, Vindelfjällens naturreservat, Tjulträsklaspen, 65.966783; 16.060500. Leg. SMTP. | 28 Jun.–15 Jul. 2004 | |
R. (Lundstroemiella) dudai Oldenberg, 1927 | AB7C | SWEDEN: Ångermanland, Örnsköldsviks kommun, Skuleskogen, Långrå, 63.088717; 18.498383. Leg. SMTP. | 5–25 Aug. 2004 | MN868958 |
R. (Amydroneura) erythrophthalma Meigen, 1830 | AD3C | SWEDEN: Skåne, Ystad kommun, Sandhammaren, Järahusen, 55.403781; 14.199936. Leg. SMTP. | 26 Sep. 2005–10 Feb. 2006 | MN868965 |
R. (Pararhamphomyia) fascipennis Zetterstedt, 1838 | AB9C | SWEDEN: Värmland, Munkfors kommun, Ransäter, Ransberg Herrgård, 59.790442; 13.415169. Leg. SMTP. | 22 May–5 Jun. 2005 | MN868954 |
R. (Amydroneura) gibba (Fallén, 1816) Old specimen | AF1C | SWEDEN: Skåne. Leg. C. H. Boheman. | 1852 or 1865 | – |
R. (Lundstroemiella) hybotina Zetterstedt, 1838 | AD2C | SWEDEN: Hälsingland, Hudiksvalls kommun, Stensjön–Lomtjärn, Stensjön, 62.140333; 16.286100. Leg. SMTP. | 14–27 Jul. 2005 | MN868949 |
R. (Holoclera) nigripennis (Fabricius, 1794) | AC7C | SWEDEN: Öl. Mörbylånga kommun, Västerstads almlunds naturreservat, old elm forest. 56.427307; 16.421942. Leg. SMTP. | 15 May–9 Jul. 2014 | MN868962 |
R. (Rhamphomyia) nigrita Zetterstedt, 1838 | AD6C | RUSSIA: Chukota Autonomous Okrug, Chaunsky, Ajon Island, 70 m from base camp, 69.5840; 168.6955. Leg. P. Mortensen. | 11–25 Jul. 2015 | MN868988 |
R. (Collinaria) nitidula Zetterstedt, 1842 Old specimen | AE8C | SWEDEN: Torne lappmark, Kiruna kommun, Abisko nationalpark. Leg. O. Ringdahl. | 30 Jun. 1918 | – |
R. (Pararhamphomyia) pilifer Meigen, 1838 | AB8C | SWEDEN: Ångermanland, Örnsköldsviks kommun, Skuleskogen, Långrå, 63.088717; 18.498383. Leg. SMTP. | 5–25 Aug. 2004 | MN868990 |
R. (Rhamphomyia) plumipes (Meigen, 1804) Old Specimen | AE6C | SWEDEN: Lappland, Hemavan, Klippen. Leg. H. Bartsch. | 4 Jul. 1993 | – |
R. (Eorhamphomyia) spinipes (Fallén, 1816) Old specimen | AE7C | SWEDEN: Lappland. Leg. N. J. Andersson. | 1843 | – |
R. (Rhamphomyia) sulcata (Meigen, 1804) | AC8C | SWEDEN: Lycksele lappmark, Sorsele kommun, Ammarnäs, Vindelfjällens naturreservat, Tjulträsklaspen, 65.966783; 16.060500. Leg. SMTP. | 28 Jun.–15 Jul. 2004 | MN868982 |
R. (Rhamphomyia) trilineata Zetterstedt, 1859 | AB6C | SWEDEN: Ångermanland, Örnsköldsviks kommun, Skuleskogen, Långrå, 63.088717; 18.498383. Leg. SMTP. | 5–25 Aug. 2004 | MN868987 |
R. (Holoclera) umbripennis Meigen, 1822 | AC1C | SWEDEN: Torne lappmark, Kiruna kommun, Abisko nationalpark, Nuolja, 68.359492; 18.719197. Leg. SMTP. | 1–13 Aug. 2005 | MN868952 |
Rhamphomyia sp. 1 | AD4C | RUSSIA: Chukota Autonomous Okrug, Chaunsky, Ajon Island, 70 m from base camp, 69.5840; 168.6955. Leg. P. Mortensen. | 11–24 Jul. 2015 | MN868956 |
Rhamphomyia sp. 2 | AD8C | FRENCH GUIANA: Approuague–Kaw, Kaw Mountain, 4.5508; –52.1949. Leg. N. Jönsson. | 4–12 Feb. 2007 | MN868967 |
Rhamphomyia sp. 3 | AF7C | JAPAN: Ehime Prefecture, Tōon–shi, Saragamine Range Prefectural Park, 33.71598; 132.8943. Leg. Japan 2011 Exp. | 18 Apr. 2011 | MN868963 |
Rhamphomyia sp. 4 | AF8C | JAPAN: Ehime Prefecture, Tōon–shi, Saragamine Range Prefectural Park, 33.71598; 132.8943. Leg. Japan 2011 Exp. | 18 Apr. 2011 | MN868955 |
For DNA extraction the KingFisher™ Cell and Tissue DNA Kit (Thermo Scientific, USA) was used together with KingFisher™ Duo (Thermo Scientific, USA) extraction robot following the manufacturer’s protocols. For extractions of large specimens one leg was removed from the body, for medium sized specimens the abdomen was removed and for small ones the whole animal was used. Lysis was performed in 56 °C overnight. After extraction the body part was returned to the specimen. For pinned material only oneg was used, due to restrictions from the museum. All extracted material is kept in 80% ethanol as vouchers at the Swedish Museum of Natural History (
The DNA concentration and fragmentation in each extraction of the nine old and dry samples was measured using Qubit (Thermo Fischer Scientific, USA) and BioAnalyzer (Agilent, USA). The samples from newer specimens were measured and compared to the older samples. The newer samples were fragmented on a Covaris sonicator, at SciLife Lab (Solna), to the target fragment length of 500–600 bp. Libraries for each sample were constructed following a modified version of the
Demultiplexed reads were quality checked and filtered using the pre-processing tool fastp (
Bayesian inference was performed on the partitioned dataset using MRBAYES v3.2.6 (
Of the 48 specimens ten lack sufficient reads or target loci and were removed from the analyses. Five out of the nine old and dry samples were successfully aligned but only three of them, E. picipes (AE2C), E. hyalipennis (AE5C) and R. gibba (AF1C), with a satisfactory amount of data. In 35 of the 38 samples there was an increase in DNA concentration after library amplification. DNA concentration for the old and dry samples in the study range between 0.626 and 1.09 ng/µl before the library amplification. For newer material stored in ethanol the concentration range between 0.198 and 11.0 ng/µl. Following library amplification DNA concentration of the old samples range between 1.58 and 13.9 ng/µl and for the newer 0.612 and 46.8 ng/µl. The number of reads for each of the 38 specimens varies between 38 000 and 3 000 000. Seven out of the ten poorly sequenced samples have a lower number of reads than the rest, ranging from 24 to 28 000. However, three of them have reads ranging between 80 000 and 176 000. The measured DNA concentration before and after library construction is depicted for each taxon in Appendix
The phylogenetic analysis of the dataset generated trees with a total of 38 taxa; 36 ingroup taxa and two outgroup taxa (Figs
The majority rule tree of partitioned UCE data of 70% completeness, inferred with Bayesian inference in MRBAYES v3.2.6 with a burnin of 25%. Posterior probability values above 50% are depicted at nodes. Voucher numbers are provided in parenthesis for all taxa. The blue branches mark the Clade A, B and C which are clades of both genera forming monophyletic groups, the red boxes mark monophyletic clades.
Maximum likelihood tree of partitioned UCE data of 70% completeness, inferred with IQTREE v1.6.10. Bootstrap values are depicted at nodes. Voucher numbers are provided in parenthesis for all taxa. The blue branches mark the Clade A and B which are clades of both genera forming monophyletic groups.
The application of target enrichment of UCEs on old and dry museum samples of Diptera was in general successful. Three of the old samples were sequenced E. (Euempis) picipes (AE2C), R. (Rhamphomyia) plumipes (AE6C) and R. (Amydroneura) gibba (AF1C) with sufficient data, but two specimens, E. (Leptempis) nigricans (AD9C) and E. (Leptempis) grisea (AE3C), had too little data coverage. When performing Sanger sequencing of COI none of the old and dry specimens were recovered. This was expected because old and dry samples usually have highly degraded DNA, and to conduct a successful PCR, preceding Sanger sequencing, it is necessary with sufficient good DNA quality (
The average DNA concentration in this study (2.26–12.94 ng/µl) were within the same range as in
The phylogenetic analyses adopting Bayesian inference inferred a tree with high support values (Fig.
The species within the two genera are morphologically quite similar, and the traditional characters used to distinguish the genera are the wing venation and mouthpart length. However, there are exceptions. For example, species in the Rhamphomyia subgenera Aclonempis and Vockerotempis,
The first-time application of the Diptera 2.7Kv1 probe kit (
Thanks to Rodrigo Esparza-Salas and Bodil Cronholm at the Molecular Systematics Laboratory (Swedish Museum of Natural History) for the valuable help with the laboratory work. Also, a big thank to SciLife Lab (Solna) for guidance and help during sample preparation. The major part of the specimens used in this study was provided by the Swedish Malaise Trap Project (SMTP). Additional specimens were collected by Niklas Apelqvist, Mathias and Catrin Jaschhof, Kjell Arne Johanson, Peter Mortensen, Pär Lindskog and Bert Viklund. Thanks to Yngve Brodin (curator) for the permission to use specimens from the Swedish Museum of Natural History collection. Thanks to Yngve Brodin and Mattias Forshage for valuable comments on the manuscript.
DNA concentration before and after library construction and number of reads for each taxon. Old referring to pinned museum specimens sampled before year 1993, new referring to specimens kept in ethanol and sampled after year 1995. Specimens below lines of average, min and max, refers to specimens excluded from the study.
Taxon | Voucher | Old/new sample | DNA conc. before-library (ng/µL) | DNA conc. after-library (ng/µL) | No of reads |
---|---|---|---|---|---|
Chelipoda sp. | AH7C | New | 0.198 | 4.16 | 138000 |
Empis albinervis | AA9C | New | 9.04 | 2.9 | 161000 |
E. bicuspidata | AB1C | New | 6.08 | 3.6 | 118000 |
E. borealis | AA8C | New | 1.6 | 3.78 | 134000 |
E. latro | AF4C | New | 1.07 | 1.11 | 40000 |
E. livida | AA3C | New | 0.526 | 4.22 | 96000 |
E. lucida | AA4C | New | 0.552 | 2.46 | 46000 |
E. nigritarsis | AD1C | New | 4.22 | 25.6 | 656000 |
E. nuntia | AB5C | New | 1.21 | 4.86 | 265000 |
E. picipes | AE2C | Old | 0.714 | 13.9 | 521000 |
E. planetica | AA7C | New | 0.61 | 2.22 | 38000 |
E. tessellata | AA6C | New | 0.502 | 4.04 | 81000 |
E. trigramma | AB2C | New | 1.12 | 3.24 | 109000 |
E. univittata | AB3C | New | 0.892 | 3.82 | 150000 |
Empis sp. 1 | AD5C | New | 0.894 | 43.6 | 1000000 |
Empis sp. 2 | AD7C | New | 1.09 | 35.0 | 1000000 |
Empis sp. 3 | AF2C | New | 0.978 | 4.7 | 443000 |
Empis sp. 5 | AG9C | New | 0.562 | 2.42 | 41000 |
Empis sp. 6 | AH2C | New | 1.12 | 6.14 | 657000 |
Hilara cornicula | AG7C | New | 0.516 | 5.1 | 347000 |
H. flavipes | AG8C | New | 0.486 | 5.82 | 294000 |
Phyllodromia melanocephala | AH5C | New | 0.602 | 7.54 | 441000 |
Rhamphomyia curvula | AC9C | New | 4.02 | 18.5 | 647000 |
R. dudai | AB7C | New | 11.0 | 1.4 | 57000 |
R. erytropthalma | AD3C | New | 9.42 | 18.9 | 1000000 |
R. fascipennis | AB9C | New | 5.02 | 2.3 | 66000 |
R. gibba | AF1C | Old | 0.626 | 1.58 | 66000 |
R. hybotina | AD2C | New | 1.48 | 30.0 | 1000000 |
R. nigripennis | AC7C | New | 1.62 | 46.8 | 3000000 |
R. nigrita | AD6C | New | 0.944 | 46.4 | 2000000 |
R. plumipes | AE6C | Old | 1.09 | 6.02 | 197000 |
R. sulcata | AC8C | New | 0.868 | 41.0 | 1000000 |
R. trilineata | AB6C | New | 2.58 | 2.72 | 142000 |
R. umbripennis | AC1C | New | 0.768 | 1.29 | 54000 |
Rhamphomyia sp. 1 | AD4C | New | 1.52 | 30.8 | 2000000 |
Rhamphomyia sp. 2 | AD8C | New | 6.62 | 39.4 | 2000000 |
Rhamphomyia sp. 3 | AF7C | New | 2.7 | 7.96 | 721000 |
Rhamphomyia sp. 4 | AF8C | New | 1.08 | 6.58 | 507000 |
Average | 2.26 | 12.94 | 558763.2 | ||
Min | 0.198 | 1.11 | 38000 | ||
Max | 11.0 | 46.8 | 3000000 | ||
E. grisea | AE3C | Old | 0.688 | 3.62 | 110000 |
E. hyalipennis | AE5C | Old | 0.642 | 1.11 | 22000 |
E. nigricans | AD9C | Old | 0.518 | 2.72 | 176000 |
E. nitida | AE1C | Old | 0.506 | 0.946 | 80000 |
Empis sp. | AF3C | New | 0.522 | 0.612 | 27000 |
R. anomalina | AC5C | New | 0.362 | 0.818 | 19000 |
R. crassirostris | AC2C | New | 0.534 | 0.584 | 19000 |
R. nitidula | AE8C | Old | 0.71 | 0.332 | 24000 |
R. pilifer | AB8C | New | 1.78 | 0.138 | 24 |
R. spinipes | AE7C | Old | 0.672 | 0.194 | 28000 |
Table of partitions and best substitution models generated by Partition Finder v2.1.1.
Subset | Best model | Number of sites | Partitions |
---|---|---|---|
1 | GTR+I+G | 750 | uce_1872_core, uce_1165_core, uce_1022_right, uce_1022_left |
2 | GTR+I+G | 176 | uce_344_left, uce_1022_core |
3 | GTR+I+G | 376 | uce_1165_left, uce_212_right |
4 | GTR+I+G | 270 | uce_1165_right |
5 | GTR+G | 415 | uce_589_right, uce_1361_left, uce_1361_core, uce_3370_right |
6 | GTR+I+G | 462 | uce_2884_left, uce_1361_right |
7 | GTR+I+G | 177 | uce_1872_left, uce_2884_right |
8 | GTR+G | 216 | uce_1872_right, uce_830_core |
9 | GTR+I+G | 794 | uce_830_left, uce_212_left, uce_212_core, uce_589_core, uce_2156_left |
10 | GTR+G | 50 | uce_2156_core |
11 | GTR+G | 205 | uce_3999_right, uce_2156_right |
12 | GTR+I+G | 99 | uce_2884_core |
13 | GTR+I+G | 50 | uce_3078_left |
14 | GTR+I+G | 705 | uce_3078_core, uce_344_right |
15 | GTR | 142 | uce_3078_right |
16 | GTR+I+G | 265 | uce_3370_left, uce_3370_core |
17 | GTR | 80 | uce_344_core |
18 | GTR+G | 157 | uce_3999_left |
19 | GTR+I+G | 652 | uce_3999_core |
20 | GTR+I+G | 600 | uce_830_right, uce_589_left |
21 | GTR+I+G | 753 | uce_716_left, uce_716_right, uce_715_left, uce_715_core, uce_716_core, uce_715_right |