Biodiversity Data Journal :
Taxonomic paper
|
Unveiling of a cryptic Dicranomyia (Idiopyga) from northern Finland using integrative approach (Diptera, Limoniidae)
Corresponding author:
Academic editor: Pierfilippo Cerretti
Received: 11 Nov 2014 | Accepted: 28 Nov 2014 | Published: 03 Dec 2014
© 2014 Jukka Salmela, Kari Kaunisto, Varpu Vahtera
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:
Salmela J, Kaunisto K, Vahtera V (2014) Unveiling of a cryptic Dicranomyia (Idiopyga) from northern Finland using integrative approach (Diptera, Limoniidae). Biodiversity Data Journal 2: e4238. https://doi.org/10.3897/BDJ.2.e4238
|
The subgenus Idiopyga Savchenko, 1987 is a northern hemisphere group of short-palped crane flies (Diptera, Limoniidae). In the current article we describe a new species, Dicranomyia (I.) boreobaltica Salmela sp.n., and redescribe the male and female post-abdomen of a closely related species, D. (I.) intricata Alexander. A standard DNA barcoding fragment of 5′ region of the cytochrome c oxidase I (COI) gene of the new species is presented, whilst the K2P minimum distances between the new species and 10 other species of the subgenus were found to range from 5.1 to 15.7 % (mean 11.2 %). Phylogenetic analyses (parsimony and maximum likelihood) based on COI sequences support the identity of the new species and its close relationship with D. (I.) intricata and D. (I.) esbeni (Nielsen). The new species is known from the northern Baltic area of Finland. The new species has been mostly collected from Baltic coastal meadows but an additional relict population is known from a calcareous rich fen that was estimated to have been at sea level circa 600-700 years ago. Dicranomyia (I.) intricata (syn. D. suecica Nielsen) is a Holarctic species, occurring in the north boreal and subarctic vegetation zones in Fennoscandia.
Crane flies, DNA barcoding, Baltic coastal meadows, mires
The subgenus Idiopyga (
Dicranomyia (I.) intricata was reported from Finland by Nieminen (
The morphological terminology used here mainly follows
Layer photos were taken using an Olympus E520 digital camera, attached to an Olympus SZX16 stereomicroscope. Digital photos were captured using the programmes Deep Focus 3.1 and Quick PHOTO CAMERA 2.3. Layer photos were finally combined with the program Combine ZP.
A 658 bp fragment of mitochondrial protein-encoding cytochrome c oxidase subunit I (COI) was sequenced from a total of 22 Dicranomyia specimens and one Metalimnobia specimen. Legs or 2–3 abdominal segments of the specimens were placed in 96% ethanol in a 96-well lysis microplate and dispatched to the Canadian Centre for DNA Barcoding, Biodiversity Institute of Ontario where DNA was extracted and sequenced using standard protocols and primers (
Dicranomyia (Idiopyga) and outgroup specimens (Dicranomyia (D.) didyma (Meigen), D. (Numantia) fusca (Meigen), Metalimnobia (M.) charlesi Salmela & Starý) used in DNA barcoding (COI). Co-ordinates are given in WGS84 decimal format.
species_sample ID | GenBank | year | country | locality | N | E |
Dicranomyia danica_JES-20110422 | KP064166 | 2009 | Czech Republic | Hrabetice | 48.788 | 16.426 |
Dicranomyia danica_JES-20110418 | KP064167 | 2009 | Czech Republic | Hrabetice | 48.788 | 16.426 |
Dicranomyia esbeni_JES-20120182 | KP064169 | 2005 | Finland | Oulunsalo | 65.039 | 24.818 |
Dicranomyia halterella_JES-20120184 | KP064171 | 2008 | Finland | Kankaanpää | 61.768 | 22.639 |
Dicranomyia halterella_JES-20110097 | KP064172 | 2009 | Finland | Enontekiö | 68.636 | 22.784 |
Dicranomyia intricata_JES-20120082 | KP064173 | 2009 | Finland | Kittilä | 67.639 | 25.427 |
Dicranomyia intricata_JES-20110082 | KP064174 | 2009 | Finland | Enontekiö | 68.636 | 22.538 |
Dicranomyia boreobaltica_JES-20120094 | KP064175 | 2005 | Finland | Oulunsalo | 64.906 | 25.376 |
Dicranomyia klefbecki_JES-20120377 | KP064176 | 2011 | Finland | Eckerö | 60.253 | 19.541 |
Dicranomyia lulensis_JES-20120081 | KP064177 | 2009 | Finland | Kemijärvi | 66.997 | 27.150 |
Dicranomyia lulensis_JES-20110365 | KP064178 | 2007 | Finland | Enontekiö | 68.484 | 22.353 |
Dicranomyia lulensis_JES-20110142 | KP064179 | 2009 | Finland | Enontekiö | 68.660 | 22.638 |
Dicranomyia magnicauda_JES-20120119 | KP064180 | 2007 | Finland | Kittilä | 68.026 | 25.111 |
Dicranomyia magnicauda_JES-20120080 | KP064181 | 2009 | Finland | Kemijärvi | 66.997 | 27.150 |
Dicranomyia magnicauda_JES-20120122 | KP064182 | 2007 | Finland | Kittilä | 67.589 | 25.662 |
Dicranomyia murina_JES-20120042 | KP064183 | 2009 | Finland | Sodankylä | 68.087 | 26.109 |
Dicranomyia ponojensis_JES-20110117 | KP064184 | 2009 | Finland | Enontekiö | 68.636 | 22.538 |
Dicranomyia ponojensis_JES-20120086 | KP064185 | 2007 | Finland | Suomussalmi | 65.230 | 28.170 |
Dicranomyia stigmatica_JES-20120415 | KP064186 | 2006 | Finland | Ruovesi | 61.837 | 24.064 |
Dicranomyia stigmatica_JES-20110360 | KP064187 | 2007 | Finland | Kittilä | 67.593 | 25.308 |
Dicranomyia didyma_JES-20110098 | KP064168 | 2009 | Finland | Enontekiö | 68.636 | 22.784 |
Dicranomyia fusca_JES-20110237 | KP064170 | 2008 | Finland | Nurmes | 63.786 | 29.350 |
Metalimnobia charlesi_JES-20110381 | KP064165 | 2008 | Finland | Lieksa | 63.468 | 29.942 |
Parsimony approach
Since the number of studied taxa was only 23, no heuristic methods were needed for the parsimony analysis. This allowed us to explore all possible evolutionary hypotheses for our data via explicit enumeration (branch and bound) analysis in TNT (Tree Analysis using New Technology) version 1.1 (
Maximum likelihood approach
Maximum likelihood analysis was conducted with RAxML ver. 8.0.22 (
Dicranomyia (Idiopyga) intricata
Dicranomyia (Idiopyga) cf. intricata
Male. Head. Vertex dark brown, with short black setae. Rostrum light brown with a few short dark setae. Palpus 5-segmented; first palpomere very short, globular, 1.5 times wider than long; other palpomeres elongated, p2 length 140 µm, p3 100 µm, p4 100 µm and p5 120 µm. First palpomere with a long ventral seta, approximately 2 times longer than width of palpomere. Second and third palpomeres with 5 setae, arranged in the apical half of segments. Fourth palpomere bearing ca. 12 setae and p5 with 13-15 setae, most of these on the apices of the segments. Antennae 14-segmented, dark brown, segments bearing black setae mostly exceeding width of respective segment; setae straight on scape (ca. 10 setae) and pecidel (ca. 15 setae), straight or curved on flagellomeres (ca. 5 setae on each flagellomere). Scape cylindrical, length 200 µm, width 75 µm, pedicel wider apically than basally, length 115 µm, width 75 µm. Flagellomeres oval, longer than wide; f1 length 120 µm, width 65 µm, f2 length 8 µm, width 5 µm, f10 length 110 µm, width 40 µm. Thorax mainly dark brown. Prescutum dark brown, only small yellowish spots on hind lateral corners. Scutum dark brown with longitudinal yellow median line and yellow lateral spots near wing base. Mediotergite and anepisternum dark brown, mediotergite sometimes with narrow yellowish anterior margin. Laterotergite and anepimeron yellowish brown. Katepisternum bicolored: anterior half dark brown, posterior half yellowish brown. Fore coxa brown, mid and hind coxae yellowish brown. Femorae light brown or brown, tibiae and tarsi dark brown. Length of fore femora 4500 µm, tibia 5250 µm, t1 3500 µm, t2 1100 µm, t3 875 µm, t4 300 µm, t5 175 µm, claw 130 µm. Length of mid femora 5575 µm, tibia 5625 µm, t1 3200 µm, t2 1150 µm, t3 625 µm, t4 275 µm, t5 175 µm, claw 130 µm. Length of hind femora 5600 µm, tibia 5750 µm, t1 3050 µm, t2 1150, t3 650 µm, t4 275 µm, t5 178 µm, claw 130 µm. Halter grayish-brown. Wing clear, veins light brown - brown, pterostigma brown (Fig.
Dicranomyia (I.) boreobaltica Salmela sp.n. male hypopygium, gonocoxite and gonostylus. gx=gonocoxite, algx=appendage of ventromesal lobe of gonocoxite, lgx=ventromesal lobe of gonocoxite, dg=dorsal lobe of gonostylus, vg=ventral lobe of gonostylus.
Dicranomyia (I.) boreobaltica Salmela sp.n., details of male hypopygium. vg=ventral lobe of gonostylus, dg=dorsal lobe of gonostylus, gx=gonocoxite, rm=rostral prolongation (rostrum) of ventral gonostyle, srm=subrostral prolongation of ventral gonostyle, iagx=inner appendage of gonocoxite, lgx=ventromesal lobe of gonocoxite, algx=appendage of ventromesal lobe of gonocoxite.
Female. In general, similar to male. Wing length 6.5 mm. Cerci short, ca. 240 µm in length. Infra-anal plate with a strong caudal peak (Fig.
Dicranomyia (I.) boreobaltica Salmela sp.n., female, details of post-abdomen.
Dicranomya (I.) intricata Alexander, male hypopygium, gonocoxite and gonostylus. gx=gonocoxite, algx=appendage of ventromesal lobe of gonocoxite, lgx=ventromesal lobe of gonocoxite, dg=dorsal lobe of gonostylus, vg=ventral lobe of gonostylus, lvg=ventrobasal lobe of ventral gonostyle.
Dicranomyia (I.) intricata Alexander, details of male hypopygium. vg=ventral lobe of gonostylus, dg=dorsal lobe of gonostylus, gx=gonocoxite, rm=rostral prolongation (rostrum) of ventral gonostyle, srm=subrostral prolongation of ventral gonostyle, iagx=inner appendage of gonocoxite, lgx=ventromesal lobe of gonocoxite, algx=appendage of ventromesal lobe of gonocoxite.
Dicranomyia (I.) intricata Alexander, female, details of postabdomen.
Phylogenetic trees of Dicranomyia (Idiopyga) species and three out-group species based on COI sequences (raw data available in Suppl. material
Brownish, small species, very close to D. (I.) intricata. Ventrobasal lobe of ventral gonostylus sinuous, apex oval. Inner appendage of gonocoxite apically rounded. Rostral prolongantion of ventral apically rather narrow and subrostral prolongation simple, not bilobed, bearing dark stout spines. Female infra-anal plate with strong caudal peak.
Boreo (borealis, Latin)= north, baltica (Latin)= referring to the Baltic Sea. The species is so far known from the northern Baltic area. The species name is deemed to be a latinized adjective in nominative singular.
European, only known from Finland. The species is hitherto known from five separate localities; four of these are shore meadows in Oulunsalo and Hailuoto island (see
The species is probably halophilous, occurring in Baltic coastal meadows characterised by vascular plants such as Phragmites australis, Lysimachia thyrsiflora, Eleocharis palustris, Carex halophila and C. paleacea (
Due to its apparent rarity, that is, small area of occupancy and extent of occurrence, the species could most likely be assessed as a threatened species according to IUCN criteria. Habitats of this species are highly endangered, usually small and isolated. There are a total of ca. 4200 ha of Baltic coastal meadows along the Finnish coast, and all such habitat types are red-listed (
Based on morphology and COI sequence divergence, the new species is very closely related to the Holarctic species D. (I.) intricata. As already stated in the title of this article, the new species is cryptic, meaning that it is hard to distinguish from its sister species by morphological characters. Strictly speaking, cryptic species may mean taxa that are morphologically indisguishable (
External characters, such as wing venation and body coloration, between D. (I.) boreobaltica Salmela sp.n. and D. (I.) intricata are practically identical. The most important differences in male and female post-abdomen between the species are summarized in Table
Summary of the most important postabdominal differences between Dicranomyia (I.) boreobaltica Salmela sp.n. and D. (I.) intricata Alexander.
D. (I.) boreobaltica | D. (I.) intricata |
apex of iagx simple, not furcated (Fig. |
apex of iagx bifurcated (Fig. |
apex of lgx angular (Fig. |
apex of lxg beak-like (Fig. |
stalk of lvg rather wide, apex oval (Fig. |
stalk of lvg tapering apically, apex spherical (Fig. |
apex of rm rounded, rather narrow (Fig. |
apex of rm pointed, rather wide (Fig. |
srm simple, not bilobed (Fig. |
srm bilobed (Fig. |
caudal margin of female infra-anal plate as in Fig. |
caudal margin of female infra-anal plate as in Fig. |
Standard 5′ region (658 bp) of the cytochrome c oxidase I (COI) sequence of Dicranomyia (I.) boreobaltica Salmela sp.n. BOLD Sample ID JES-20120094, holotype specimen:
TACCTTATACTTTATTTTTGGAGCTTGAGCAGGAATAGTGGGAACTTCATTAAGTATTATTATTCGAGCAGAATTAGGACACCCAGGTGCATTAATTGGAGACGACCAGATTTATAATGTGGTAGTTACTGCCCATGCTTTTATTATAATTTTCTTTATAGTTATACCAATTATAATTGGAGGATTCGGTAATTGATTAGTTCCTTTAATATTAGGAGCCCCAGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGAATACTTCCCCCTTCTTTAACTTTATTATTAGCTAGAAGCATAGTTGAAAACGGGGCAGGAACTGGCTGAACAGTATACCCTCCCCTTTCTTCTGGAATTGCCCATTCAGGGGCTTCTGTAGATTTAGCTATTTTTTCTCTTCACCTAGCAGGTATTTCTTCTATTTTAGGAGCTGTTAATTTTATTACAACTGTTATTAATATACGTTCAGCAGGAATTTCATTTGATCGAATACCATTATTTGTTTGATCAGTAGTAATTACTGCTATTTTATTGCTTTTATCACTTCCTGTTTTAGCCGGAGCTATTACAATATTATTAACAGATCGAAACTTAAATACTTCATTTTTTGATCCCGCAGGTGGAGGAGACCCTATTTTATATCAGCATTTATTT
Based on K2P (
Dicranomyia intricata
Limonia (Dicranomyia) suecica
Limonia (Dicranomyia) suecica
Dicranomyia (Idiopyga) intricata
The holotype specimen of D. (I.) intricata (Fig.
Male hypopygium. 9th tergite and proctiger as in Fig.
Female postabdomen. Cerci and hypogynial valves, see Fig.
Holarctic. Known from Canada (Alberta, Northwest territories, British Columbia), Sweden (North Sweden, Abisko,
The original description of D. (I.) intricata (
Dicranomyia (I.) intricata is red-listed in Finland (NT,
See Dicranomyia (I.) boreobaltica Salmela sp.n.
Standard 5′ region (658 bp) of the cytochrome c oxidase I (COI) gene of Dicranomyia (I.) intricata (BOLD Sample IDs JES-20120082 and JES-20110082, identical specimens):
TACCTTATACTTTATTTTTGGAGCTTGAGCAGGAATAGTAGGAACTTCACTAAGTATTATTATTCGAGCAGAATTAGGACACCCAGGAGCATTAATTGGAGATGACCAAATTTATAATGTAGTAGTTACTGCCCATGCTTTTATTATAATTTTTTTTATAGTTATACCAATTATAATTGGTGGATTCGGTAATTGATTAGTTCCTTTAATATTAGGAGCCCCAGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGAATACTTCCCCCTTCTTTAACCTTATTATTAGCTAGAAGTATAGTTGAAAACGGGGCAGGAACTGGTTGAACAGTTTACCCTCCCCTTTCTTCTGGAATTGCTCATTCAGGAGCTTCTGTAGACTTAGCTATTTTTTCTCTTCATTTAGCAGGTATTTCTTCTATTTTAGGAGCTGTTAACTTTATTACAACTGTTATTAATATACGTTCAGCAGGAATTTCATTCGACCGAATACCATTATTTGTTTGATCAGTAGTAATTACTGCTATTCTATTACTCTTATCACTCCCTGTTTTAGCTGGAGCTATTACAATATTATTAACAGATCGAAACTTAAACACTTCATTTTTTGACCCTGCAGGTGGAGGAGATCCTATTTTATACCAACACTTATTT
The phylogenetic tree (length 622 steps) resulting from the parsimony analysis is shown in Fig.
Barcode sequences were obtained at the Canadian Centre for DNA Barcoding based in the Biodiversity Institute of Ontario at the University of Guelph. Their work was supported by funding from the Government of Canada through Genome Canada and the Ontario Genomics Institute in support of the International Barcode of Life Project. Finnish Barcode of Life (FinBOL), led by Dr. Marko Mutanen (Oulu) is also thanked for a help in the barcoding process. JS would like to thank Mr. Teemu Nieminen for his trust and co-operation during his MSc project. Mr. Matti Mäkilä (Rovaniemi) is thanked for his in help in field work and painstaking sorting of Malaise traps samples. Dr. Tapani Sallantaus (Helsinki) kindly gave us his data on water quality from Kusiaiskorpi. KMK was financially supported by Emil Aaltonen foundation. English text was improved by Pete Boardman (Shrewsbury). We also thank referees for their constructive comments.
JS wrote the majority of the manuscript and took some layer photos. VV performed the phylogenetic analyses. KMK took most of the layer photos.
Non-type material of Dicranomyia (I.) intricata Alexander, males, permanently slide-mounted by C.P. Alexander, deposited in USNM (USA, Washington). Digital photos of the slides.
COI 5' standard DNA barcoding fragment