Research Article |
Corresponding author: Marla D. Schwarzfeld ( marla.schwarzfeld@unbc.ca ) Academic editor: Gavin Broad
© 2014 Marla D. Schwarzfeld, Felix A. H. Sperling.
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:
Schwarzfeld MD, Sperling FAH (2014) Species delimitation using morphology, morphometrics, and molecules: definition of the Ophion scutellaris Thomson species group, with descriptions of six new species (Hymenoptera, Ichneumonidae). ZooKeys 462: 59-114. https://doi.org/10.3897/zookeys.462.8229
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The diverse genus Ophion is almost entirely undescribed in the Nearctic region. In this paper we define the Ophion scutellaris species group. This species group is well-supported by analysis of DNA (ITS2, COI, and 28S D2-D3) and morphology. It includes the Palearctic species O. scutellaris and the Nearctic species O. idoneus. An integrative analysis of DNA, geometric wing morphometrics, classical morphometrics and qualitative morphology indicates that this species group contains a minimum of seven species in North America, although the full diversity of the group has likely not been sampled. O. clave Schwarzfeld, sp. n., O. aureus Schwarzfeld, sp. n., O. brevipunctatus Schwarzfeld, sp. n., O. dombroskii Schwarzfeld, sp. n., O. keala Schwarzfeld, sp. n. and O. importunus Schwarzfeld, sp. n. are described, and a key to the known Nearctic species of the O. scutellaris group is provided.
Barcoding, geometric morphometrics, Ichneumonidae , integrative taxonomy, molecular taxonomy, Ophion , species delimitation
Ophion Fabricius is a genus of large nocturnal Ichneumonidae in the subfamily Ophioninae. Most species parasitize medium to large-sized Lepidoptera larvae, especially Noctuoidea (
While several species groups of Ophion have been proposed (summarized by
Ophion species are difficult to distinguish morphologically and have a great deal of intraspecific variability (
To avoid relying on a single mitochondrial gene, nuclear genes provide an additional data source to evaluate species boundaries (
Quantitative morphometric analyses have been shown to be at least as accurate as molecular analyses in the delimitation of morphologically challenging species, and possibly more so (
The use of multiple lines of evidence to circumscribe species is often referred to as “integrative taxonomy” (
In this paper, we define the O. scutellaris species group, the first Ophion species group to be circumscribed based on both molecular and morphological characters. We then revise the known Nearctic species within this group, including descriptions of six new species.
BMNH The Natural History Museum, London, UK
CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada
CUIC Cornell University, Ithaca, New York, USA
NFRC Northern Forest Research Centre, Edmonton, Alberta, Canada
RBCM Royal British Columbia Museum, Victoria, British Columbia, Canada
SEMC Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA
SFUC Simon Fraser University, Burnaby, British Columbia, Canada
UASM E. H. Strickland Entomology Collection, University of Alberta, Edmonton, Alberta, Canada
UBCZ Spencer Entomology Collection, University of British Columbia, Vancouver, British Columbia, Canada
This species group was first identified as part of a large-scale taxonomic and phylogenetic study of Ophion (Schwarzfeld et al., in prep.). From the sequencing results of several dozen Ophion morphospecies, a well-supported clade that included the Palearctic species O. scutellaris Thomson was identified (Schwarzfeld et al., in prep.). Morphological comparison showed that several of the characters that define O. scutellaris, according to
Most specimens were newly collected as part of a large-scale study of Canadian Ophion. Out of more than 4000 specimens that were collected from a range of habitats and localities across Canada (primarily from light-traps and Malaise traps), 196 were from the Ophion scutellaris species group. We also borrowed 662 specimens of Ophion from the following institutions: CUIC, UBCZ, SEMC, RBCM, CNC, NFRC, SFUC; these included 42 specimens from the species group. Five specimens of Ophion scutellaris were sequenced from British material provided by G. Broad (BMNH). We examined three of these, while for two specimens, we were sent legs for sequencing but have not seen the specimens.
We sequenced one mitochondrial and two nuclear genes for the O. scutellaris group: the cytochrome c oxidase I gene (COI), the internal transcribed spacer 2 (ITS2), and the D2-D3 variable region of 28S rRNA. We also sequenced these three genes for eight species of Ophion outside of the target species group; these were chosen to represent the diversity across Ophion, based on a large-scale molecular analysis of Ophion (Schwarzfeld and Sperling, in prep.). Finally we sequenced one individual of Enicospilus, another genus within Ophioninae, as an outgroup. DNA was extracted from a single hind leg using DNeasy Blood & Tissue Kits (Qiagen, Toronto, ON); the final elution volume was 150 μL. We conducted PCR in either 50 μL or 15 μL reactions. The 50 μL reactions contained 4–8 μL genomic DNA, 5 μL 10× PCR buffer (containing 15 mmol/μL MgCl2) (Promega, Madison, WI), 3 μL of 25 mmoles/μL MgCl2 (Promega), 1 μL of 10 mmoles/μL dNTP’s (Roche, Switzerland), 1 μL each of 5pmol/μL forward and reverse primers, 0.5 μL of 5 U/μL Taq polymerase (Fermentation Service Unit, University of Alberta) and 30.5–34.5 μL of autoclaved Millipore water. The 15 μL reactions used 4–8 μL DNA, 1.5 μL PCR buffer, 0.9 μL MgCl2, 0.3 μL each of dNTP’s, forward and reverse primers, 0.15 μL Taq and 3.55–7.55 μL water (all concentrations as above). All PCR products were purified using ExoSap-IT (USB Corporation, Cleveland, OH), and were sequenced using BigDye Terminator version 3.1 cycle sequencing kit (Applied Biosystems, Foster City, CA), followed by ethanol precipitation. Sequencing reactions were run on an ABI Prism 3730 DNA analyser. Sequences are deposited in NCBI GenBank, and Genbank accession numbers are listed in Appendix
To sequence COI, we used the primers lco hym (5’–CAA ATC ATA AAG ATA TTG G–3’) and hco out (5’–CCA GGT AAA ATT AAA ATA TAA ACT TC–3’) (
We analyzed ITS2 using the primers ITS2-F (5’-GGG TCG ATG AAG AAC GCA GC-3’) and ITS2-R (5’-ATA TGC TTA AAT TCA GCG GG-3’) which anneal to the flanking 28S and 5.8S genes (
We sequenced the D2-D3 region of 28S rDNA using the following primers: Forward: 5’-GCG AAC AAG TAC CGT GAG GG-3’; Reverse: 5’-TAG TTC ACC ATC TTT CGG GTC-3’ (
Alignments for all three gene regions have been deposited on TreeBASE, and can be accessed at: http://purl.org/phylo/treebase/phylows/study/TB2:S16341.
All molecular analyses were conducted using MEGA version 5 (
We analysed wing morphometrics for 118 specimens (71 female and 47 male; Appendix
Only the fore wings were used for the geometric morphometrics. A total of 23 landmarks were digitized in tpsDig version 2.16 (
Wing characters used in classical morphometric analysis and in species descriptions. DC = discocubital cell; Fen = fenestra; Ram = ramellus. Wing indices: AI = xy/wz; CI = ef/fh; ICI = wz/wx; SDI = fg/de; cu-a = ab/bc; red circles indicate the locations of the 23 landmarks used for geometric morphometrics.
Eighty-four specimens (52 females and 32 males; Appendix
List of abbreviations for morphological characters included in this study. Abbreviations marked with an asterisk were included in the classical morphometric analysis of the O. scutellaris species group.
Abbreviation | Character | Details |
---|---|---|
Head | ||
CAW | Clypeus apical width | A straight line across ventral margin of clypeus (Figure |
CH | Clypeus height | At centre of clypeus, from ventral margin to epistomal sulcus (Figure |
F1 | Flagellomere 1 length/width | In ventral view at midpoint of flagellomere |
F20 | Flagellomere 20 length/width | In ventral view at midpoint of flagellomere |
Flag* | Number of flagellomeres | For the morphometric analysis, three specimens of O. idoneus (one male and two females) had broken antennae. The number of flagellomeres for each of these specimens was estimated by averaging the number of flagellomeres for all remaining O. idoneus of the respective sex. One female specimen of an unknown species had broken antennae; since the analysis did not allow missing data, the number of flagellomeres was estimated by averaging the number of flagellomeres of all female specimens of a similar size (i.e. excluding O. aureus and O. keala). |
FH* | Face height | From the ventral margin of the clypeus to the bottom of the facial tubercle in frontal view (Figure |
FM* | Face maximum width | Widest part of the face between the maximum indentation of the eyes (Figure |
FW* | Face width | Between the inner eye margins at the level of the clypeal foveae (Figure |
GI | Genal inflection | Length of the hypostomal carina between base of mandible and intersection with genal carina ( |
HW* | Head width | Across the widest part of the eyes in frontal view (Figure |
IOD* | Interocellar distance | Shortest distance between the posterior ocelli in dorsal view |
MS* | Malar space | Shortest distance between the eye and the base of the mandible (Figure |
MW* | Mandible width | Measured at the base (Figure |
OC | Postocellar vertex | Minimum distance between occipital carina and posterior ocellus |
OL* | Ocellus length | Maximum length of the posterior ocellus |
OOD* | Ocellar-ocular distance | Shortest distance from the eye margin to the deepest part of the sulcus adjacent to the posterior ocellus |
Mesosoma | ||
SL* | Scutellum length | From the base to the apex of the scutellum |
SW* | Scutellum width | Across the base of the scutellum between the inner margins of the lateral scutellar carinae |
Metasoma | ||
M1A* | Metasomal segment 1 apical width | The width of the first metasomal segment measured in dorsal view at the apex (Figure |
M1B* | Metasomal segment 1 basal width | The width of the first metasomal segment measured in dorsal view at the base (Figure |
M1S* | Metasomal segment 1 spiracle width | The width of the first metasomal segment measured in dorsal view at the level of the spiracles (Figure |
Legs | ||
CL | Metacoxa length | Maximum length of metacoxa in lateral view |
CW | Metacoxa width | Maximum width of metacoxa in lateral view |
FL* | Metafemur length | Midpoint of the base of the hind femur to midpoint of the apex, in lateral view |
FW* | Metafemur width | Maximum lateral width of the hind femur |
MT1 | Metatarsomere 1 length | In lateral view at midpoint of tarsomere |
MT2 | Metatarsomere 2 length | In lateral view at midpoint of tarsomere |
MTS | Mesotibial spur ratio | Length of shorter spur/Length of longer spur |
Wings | ||
Wing L* | Length of the fore wing | For most specimens, this was measured in ImageJ from the photographs used in the wing geometric morphometric analysis; photographs were calibrated for size using the known size of the coverslip on the slide-mount; for those specimens not photographed for the wing analysis, wing length was measured using a calibrated ocular micrometer |
cu-a* | cu-a index of hind wing | Length of cu-a above Cu 1/length of cu-a below Cu 1 (=ab/bc in Figure |
AI | Alar index of fore wing | Length of 1m-cu between bulla and 2m-cu/length of 3rs-m (=xy/wz in Figure |
CI | Cubital index of fore wing | Length of Cu 1 between 1m-cu and Cu 1a/length of Cu 1b (=ef/fh in Figure |
ICI | Intercubital index of fore wing | Length of 3rs-m/length of M between 2m-cu and 3rs-m (=wz/wx in Figure |
SDI | Second discoidal index of fore wing | Length of Cu 1a between Cu 1b and 2m-cu/length of Cu 1 between Rs+M and 1m-cu (=fg/de in Figure |
The data underpinning the analyses reported in this paper are deposited at Canadensys (www.canadensys.net), and are available at https://doi.org/10.5886/y2bbsq5.
The O. scutellaris species group was well-supported by both molecular and morphological characters, and thus far includes seven Nearctic and potentially two Palearctic species (Figure
Statistics from the molecular analyses are found in Table
Summary of statistics from maximum likelihood (ML) and maximum parsimony (MP) analyses of COI, ITS2 and 28S genetic markers.
Analysis | Statistics | COI | ITS2 | 28S |
---|---|---|---|---|
ML | Log likelihood | -3545.81 | -4076.72 | -1429.26 |
MP | Tree Length | 579 | 467 | 75 |
No. MP trees | 8 | 10 | 10 | |
CI/RI | 0.55/0.80 | 0.90/0.93 | 0.88/0.78 |
Interspecific and intraspecific percent sequence divergence for three genetic markers within the Ophion scutellaris species group. Genetic distances calculated with the Kimura-2-parameter model are found above the diagonal, uncorrected p-distances are below the diagonal, and intraspecific distances are along the diagonal. Intraspecific distances and all 28S distances were identical using both models. aur = aureus; bre = brevipunctatus; cla = clave; dom = dombroskii; ido = idoneus; imp = importunus; kea = keala; scu A = scutellaris A; scu B = scutellaris B.
a. COI | |||||||||
---|---|---|---|---|---|---|---|---|---|
aur | bre | cla | dom | ido | imp | kea | scu A | scu B | |
aur | 0.4 | 2.0–2.4 | 2.9–3.7 | 2.1–2.4 | 9.5–10.1 | 9.1–9.6 | 9.3–10.0 | 9.0–10.0 | 8.8–8.9 |
bre | 1.9–2.4 | n/a | 3.0–3.4 | 1.0 | 8.3–8.5 | 7.7 | 8.3–8.7 | 8.8–9.3 | 8.5 |
cla | 2.8–3.6 | 3.0–3.3 | 0.1–0.3 | 3.2–3.5 | 9.1–9.7 | 9.0–9.3 | 9.2–9.8 | 9.0–9.6 | 8.7–9.0 |
dom | 2.1–2.4 | 1.0 | 3.1–3.4 | n/a | 9.5–9.7 | 8.8 | 9.2–8.8 | 9.4–9.8 | 8.8 |
ido | 8.9–9.4 | 7.9–8.0 | 8.6–9.0 | 8.9–9.0 | 0–0.3 | 2.3–2.4 | 4.6–5.4 | 6.5–7.6 | 6.7–7.0 |
imp | 8.6–9.0 | 7.3 | 8.4–8.7 | 8.3 | 2.2–2.4 | n/a | 4.8–5.3 | 6.7–7.6 | 7.0 |
kea | 8.7–9.3 | 7.9–8.2 | 8.6–9.2 | 8.6–8.9 | 4.4–5.2 | 4.6–5.0 | 0.1–0.7 | 4.6–5.9 | 5.4–5.9 |
scu A | 8.5–9.2 | 8.3–8.7 | 8.5–9.0 | 8.8–9.1 | 6.2–7.1 | 6.4–7.1 | 4.5–5.6 | 0 | 3.4–3.8 |
scu B | 8.3–8.4 | 8.0 | 8.1–8.4 | 8.3 | 6.4–6.7 | 6.7 | 5.2–5.6 | 3.3–3.6 | n/a |
b. ITS2 | ||||||||
---|---|---|---|---|---|---|---|---|
aur | bre | cla | dom | ido | imp | kea | scu | |
aur | 0 | 0.6 | 0.2–0.3 | 0.3 | 2.0–2.1 | 2.3–2.4 | 1.9–2.0 | 1.8–1.9 |
bre | 0.6 | n/a | 0.5 | 0.3 | 2.3 | 2.6 | 2.2 | 2.1 |
cla | 0.2–0.3 | 0.5 | 0 | 0.1 | 1.8 | 2.1 | 1.7 | 1.6–1.7 |
dom | 0.3 | 0.3 | 0.1 | n/a | 1.9 | 2.2 | 1.8 | 1.7 |
ido | 1.9–2.1 | 2.2–2.3 | 1.7–1.8 | 1.8 | 0 | 0.9 | 0.1 | 0.4–0.5 |
imp | 2.3–2.4 | 2.6 | 2.0–2.1 | 2.1 | 0.9 | n/a | 0.8 | 0.7–0.8 |
kea | 1.8–1.9 | 2.1 | 1.6 | 1.7 | 0.1 | 0.8 | 0 | 0.3–0.4 |
scu | 1.7–1.9 | 2.0–2.1 | 1.5–1.7 | 1.6–1.7 | 0.4–0.5 | 0.7–0.8 | 0.3–0.4 | 0.1 |
c. 28S; p-distances and K2P distances were identical, and are therefore only reported above the diagonal | ||||||||
---|---|---|---|---|---|---|---|---|
aur | bre | cla | dom | ido | imp | kea | scu | |
aur | n/a | 0.1 | 0.1 | 0.1 | 0.6 | 0.7 | 0.6 | 0.6–0.7 |
bre | 0.1 | n/a | 0 | 0 | 0.4 | 0.6 | 0.4 | 0.5–0.6 |
cla | n/a | 0 | 0.4 | 0.6 | 0.4 | 0.5–0.6 | ||
dom | n/a | 0.4 | 0.6 | 0.4 | 0.5–0.6 | |||
ido | n/a | 0.1 | 0 | 0–0.1 | ||||
imp | n/a | 0.1 | 0.2–0.3 | |||||
kea | 0 | 0–0.1 | ||||||
scu | n/a |
Monophyly of the O. scutellaris species group was strongly supported in almost all analyses, with bootstrap support ranging from 72% in the 28S ML analysis to 100% in the ITS2 MP analysis (Figure
Sequence divergence within the O. scutellaris group was highest in the COI dataset, with a maximum of 9.4% (K2P: 10.1%) sequence divergence within the species group, and 14.8% (K2P: 16.6%) within all included Ophion. While ITS2 had a higher sequence divergence overall (17.4%; K2P: 19.9%), the divergence within the O. scutellaris species group was only 2.6%. 28S was highly conserved, with a 2.8% (K2P: 2.9%) divergence within all Ophion, and 0.7% within the O. scutellaris group.
For the geometric morphometric (GM) analysis, approximately 60% of the variation in wing shape could be summarized in the first three principal components (PC). In the analysis of females, the variation was quite evenly represented by the first two principal components (PC1: 26.5%, PC2: 21.7%, PC3: 10.3%; Figure
In the classical morphometric (CM) analysis, the first three principal components represented 77.9% of the total variation in the female dataset (PC1: 52.0%, PC2: 18.9%, PC3: 7.0%; Figure
Summary of measurements of 13 morphometric variables for females of the O. scutellaris species group. Morphometric variables are defined in Table
Variable | cla (13) | aur (1) | bre (1) | dom (1) | kea (8) | imp (3) | ido (21) | A (1) | B (1) | C (1) |
---|---|---|---|---|---|---|---|---|---|---|
FW/FM | 0.83±0.02 | 0.83 | 0.79 | 0.84 | 0.81±0.02 | 0.82±0.01 | 0.86±0.01 | 0.88 | 0.87 | 0.83 |
0.81–0.85 | 0.78–0.84 | 0.82–0.83 | 0.84–0.89 | |||||||
FW/FH | 1.23±0.08 | 1.37 | 1.27 | 1.34 | 1.33±0.09 | 1.26±0.02 | 1.37±0.05 | 1.30 | 1.29 | 1.29 |
1.19–1.26 | 1.24–1.49 | 1.25–1.29 | 1.28–1.45 | |||||||
FW/HW | 0.52±0.02 | 0.51 | 0.47 | 0.53 | 0.48±0.01 | 0.50±0.00 | 0.53±0.01 | 0.54 | 0.54 | 0.51 |
0.50–0.54 | 0.46–0.50 | 0.49–0.50 | 0.51–0.55 | |||||||
MS/MW | 0.51±0.10 | 0.60 | 0.50 | 0.58 | 0.40±0.04 | 0.51±0.06 | 0.59±0.04 | 0.60 | 0.58 | 0.54 |
0.43–0.63 | 0.33–0.47 | 0.46–0.58 | 0.54–0.70 | |||||||
IOD/OL | 0.77±0.08 | 0.71 | 0.69 | 1.20 | 0.36±0.06 | 0.72±0.16 | 0.74±0.08 | 1.00 | 0.73 | 0.71 |
0.53–1.00 | 0.27–0.44 | 0.53–0.85 | 0.61–0.86 | |||||||
OOD/OL | 0.22±0.05 | 0.29 | 0.13 | 0.45 | 0.16±0.04 | 0.19±0.04 | 0.33±0.04 | 0.32 | 0.20 | 0.21 |
0.13–0.29 | 0.11–0.22 | 0.15–0.23 | 0.25–0.43 | |||||||
SL/SW | 1.43±0.17 | 1.18 | 1.48 | 1.76 | 1.63±0.10 | 1.58±0.03 | 1.52±0.09 | 1.39 | 1.52 | 1.45 |
1.23–1.58 | 1.47–1.76 | 1.55–1.60 | 1.37–1.69 | |||||||
M1B/M1S | 0.80±0.11 | 0.70 | 0.80 | 0.78 | 0.63±0.06 | 0.78±0.04 | 0.80±0.06 | 0.82 | 0.88 | 0.63 |
0.71–0.92 | 0.57–0.74 | 0.75–0.82 | 0.71–0.92 | |||||||
M1S/M1A | 0.61±0.07 | 0.69 | 0.60 | 0.64 | 0.76±0.06 | 0.66±0.02 | 0.67±0.06 | 0.64 | 0.53 | 0.70 |
0.58–0.67 | 0.57–0.74 | 0.64–0.68 | 0.59–0.79 | |||||||
FL/FW | 7.82±0.95 | 7.95 | 9.50 | 7.24 | 10.26±1.09 | 8.58±0.31 | 6.87±0.26 | 7.56 | 7.89 | 7.79 |
7.00–8.59 | 8.41–11.63 | 8.25–8.87 | 6.41–7.50 | |||||||
Flag | 55.08±5.82 | 65 | 67 | 51 | 70.63±1.30 | 55.00±1.73 | 54.05±1.93 | 52 | 59 | 59 |
52–60 | 69–73 | 54–57 | 51–57 | |||||||
cu-a | 0.69±0.11 | 0.88 | 0.66 | 0.83 | 1.08±0.13 | 0.72±0.09 | 0.83±0.12 | 0.62 | 0.68 | 0.64 |
0.60–0.84 | 0.89–1.26 | 0.66–0.83 | 0.68–1.13 | |||||||
Wing L | 11.83±2.76 | 16.69 | 14.04 | 10.80 | 19.06±0.45 | 12.92±0.38 | 10.13±0.64 | 10.70 | 12.86 | 12.70 |
10.49–12.44 | 18.64–19.73 | 12.57–13.33 | 8.98–11.48 |
Summary of measurements of 13 morphometric variables for males of the O. scutellaris species group. Morphometric variables are defined in Table
Variable | cla (3) | aur (9) | kea (1) | ido (17) | D (1) | E (1) |
---|---|---|---|---|---|---|
FW/FM | 0.85±0.02 | 0.84±0.01 | 0.86 | 0.89±0.02 | 0.83 | 0.91 |
0.83–0.87 | 0.82–0.89 | 0.85–0.91 | ||||
FW/FH | 1.22±0.04 | 1.33±0.05 | 1.41 | 1.40±0.03 | 1.23 | 1.39 |
1.18–1.26 | 1.25–1.41 | 1.32–1.45 | ||||
FW/HW | 0.50±0.02 | 0.52±0.01 | 0.50 | 0.53±0.01 | 0.50 | 0.55 |
0.49–0.52 | 0.50–0.54 | 0.51–0.56 | ||||
MS/MW | 0.48±0.06 | 0.74±0.05 | 0.39 | 0.65±0.08 | 0.58 | 0.70 |
0.43–0.54 | 0.65–0.80 | 0.50–0.82 | ||||
IOD/OL | 0.79±0.13 | 0.74±0.05 | 0.44 | 0.87±0.16 | 1.04 | 0.90 |
0.67–0.92 | 0.65–0.80 | 0.58–1.25 | ||||
OOD/OL | 0.23±0.03 | 0.30±0.02 | 0.44 | 0.48±0.14 | 0.33 | 0.40 |
0.20–0.27 | 0.26–0.33 | 0.35–0.92 | ||||
SL/SW | 1.54±0.12 | 1.23±0.08 | 1.74 | 1.44±0.12 | 1.68 | 1.44 |
1.43–1.67 | 1.13–1.38 | 1.21–1.68 | ||||
M1B/M1S | 0.79±0.04 | 0.64±0.08 | 0.63 | 0.77±0.07 | 0.79 | 0.75 |
0.76–0.84 | 0.54–0.78 | 0.61–0.87 | ||||
M1S/M1A | 0.65±0.06 | 0.72±0.04 | 0.79 | 0.73±0.07 | 0.67 | 0.80 |
0.61–0.71 | 0.67–0.78 | 0.62–0.90 | ||||
FL/FW | 7.08±0.09 | 8.72±0.54 | 9.03 | 6.95±0.43 | 7.13 | 7.14 |
7.00–7.18 | 8.11–9.63 | 6.31–7.85 | ||||
Flag | 54.67±1.53 | 66.78±1.56 | 65 | 54.69±1.74 | 54 | 52 |
53–56 | 65–69 | 51–57 | ||||
cu-a | 0.69±0.09 | 0.69±0.10 | 1.43 | 0.97±0.22 | 0.76 | 0.96 |
0.59–0.74 | 0.51–0.83 | 0.72–1.53 | ||||
Wing L | 10.78±0.11 | 15.55±0.94 | 16.23 | 9.45±0.56 | 9.91 | 8.91 |
10.65–10.85 | 14.37–17.07 | 8.42–10.74 |
Examination of specimens from the two strongly supported subgroups from the molecular analysis uncovered additional morphological characters distinguishing them. The most useful character to distinguish these subgroups can be found on the propodeum. All specimens in subgroup A have a weakly arched anterior transverse carina (Figure
Clypeus of subgroup A (a–c) and subgroup B (d–f) within the O. scutellaris species group. a, d Clypeus in frontal view; arrow indicates epistomal sulcus b, e Clypeus in frontal view, showing representative clypeal punctures c, f Clypeus in lateral view; arrow indicates the more convex clypeus in subgroup A, compared to subgroup B.
Six Nearctic species are described, and one redescribed, based on the integrated results of these analyses (see Taxonomic Part below). Because of the nature of the iterative process, a single path was not followed in determining the species boundaries for all species.
Two species (O. aureus Schwarzfeld, sp. n. and O. keala Schwarzfeld, sp. n.) were easily recognizable as distinct based on gross morphology. In particular, they are both considerably larger than the remaining species in the group. Ophion aureus is only known from the Peace River region of Alberta, and was recognized based on its golden-orange colour and distinctive propodeal carinae (Plate
There is a strong sex bias in the specimens available for each of these species. Ophion aureus was represented entirely by males, except for a single female specimen, while the opposite occurred in O. keala. They could therefore only be thoroughly assessed in the morphometric analyses of male or female specimens, respectively. Both species were successfully delimited by both GM and CM analyses, with essentially no overlap with other specimens (Figures
O. idoneus Viereck, the only previously described Nearctic species within this species group, was also initially recognized on the basis of morphology, in particular the shape of the propodeal carinae, the lack of a ramellus in the fore wing and the small size (Plate
A single female specimen of O. importunus sp. n. was first noted as being very similar to O. idoneus, but differing slightly in the shape of the propodeal carinae, along with being slightly larger and having a longer ramellus (Plate
O. clave Schwarzfeld, sp. n. was the only species other than O. idoneus that was represented by multiple male and female specimens, though female specimens were more common. It was first identified as a putative species based on the molecular analyses. It was recovered as part of subgroup B by all analyses, and formed a well-supported monophyletic clade in the analysis of COI (Figure
The morphometric analyses generally recovered both males and females of this species as distinct clusters compared to other species. For the analysis of females, this result was strongest in the plots of PC1 vs PC2 of both GM and CM, whereas all plots from the analysis of males recovered this species as a distinct group (Figures
O. brevipunctatus Schwarzfeld, sp. n. and O. dombroskii Schwarzfeld, sp. n. are each represented by singletons in this study. Ophion dombroskii differs from all other members of the scutellaris species group, and from all other known Nearctic Ophion, by its distinctive black face and thorax (Plate
Five specimens (3 female and 2 male) remain unassigned to any species. Based on the structure of the propodeal carinae, one of the female specimens can be assigned to subgroup A, while the other two are part of subgroup B. The former specimen (labeled as “A” in figures and tables) was initially identified as O. idoneus, but based on its divergent position in the morphometric analyses, particularly in the GM analysis, we re-examined this specimen, and concluded that it also differs from O. idoneus morphologically. The other two female specimens (“B” and “C”) cluster near or within O. clave in both CM and GM analyses. However they were sufficiently morphologically divergent from O. clave and from each other that we suspect this does not indicate conspecificity.
One of the two unplaced male specimens (“D”) can be assigned to subgroup B, and the other (“E”) is in subgroup A. According to both the GM and CM analyses, these clustered with species from their respective subgroup, i.e. “D” was near O. clave and “E” was near O. idoneus. However, they are sufficiently different morphologically from these species that this is probably an artifact of the data. It is possible that specimen “E” represents the unknown male of O. importunus, whereas we are almost certain that specimen “D” is an additional undescribed species.
The genus Ophion is often considered a particularly challenging taxonomic puzzle, due to high intraspecific and low interspecific morphological variation (
While 28S was useful in supporting the monophyly of the species group as a whole, and particularly of the two subgroups, it was too conserved to distinguish species within each subgroup. COI and ITS2 were both apparently effective at separating species, since most species, as defined through the integrated analysis, were recovered as well supported clades, with greater divergence between species than within species. However, few individuals over a limited geographical range were sequenced for the majority of species; further sequencing is needed to fully assess the ability of these markers to distinguish species across their range (
Despite the apparent utility of these molecular markers, morphology is an essential component of diagnosing species, since clusters of haplotypes are not necessarily equivalent to biological species (
Qualitative morphology, geometric morphometrics and classical morphometrics provide three additional semi-independent datasets with which to evaluate species diagnoses. Each species in this study is distinguishable morphologically, though again small sample sizes and limited geographical sampling mean the morphological variability within each species has probably not been fully sampled. Four species (O. clave, O. brevipunctatus, O. importunus, and O. idoneus) look superficially very similar, however each has qualitative morphological characters that are sufficient to separate them. In comparison, O. aureus, O. keala, and O. dombroskii are very morphologically distinctive species. Both O. brevipunctatus and O. dombroskii are described from single specimens. While some taxonomists argue that species should never be described from singletons (e.g.
Principal Components Analysis is a one-group method for data exploration, meaning it does not include a priori group designations (
Both geometric and classical morphometrics were effective for clustering species, although the clusters were often not widely separated. In general, superficially similar species clustered more closely together than they did to morphologically divergent species, indicating congruence between quantitative measures of shape and subjective, qualitative analysis. Specimens that did not cluster with other members of their putative species were flagged for further examination. In some cases these were misidentifications, while in others morphology and the alternate morphometric analysis supported their placement within a given species. Conversely, inclusion within a cluster did not guarantee that the specimen was a member of that species. These examples further support the advantage of including multiple lines of evidence for accurate species delimitation.
This is a first attempt at describing species within this newly-defined species group; as such, it should be considered a work in progress. This study was almost entirely limited to Canadian material, most species were from only a few localities, and all species except O. idoneus were represented by small numbers of individuals. As well, there are almost certainly additional undescribed species in the material available. All of this indicates that we have just begun to sample the true diversity within this species group. Nonetheless, we have shown that by using an iterative analysis of morphology, molecular analysis and morphometrics, we can delimit and describe species within a genus that is so morphologically challenging that no new species have been described in North America for more than a hundred years. Furthermore, molecular and morphological recognition of this species group will now allow more targeted specimen collection and museum research, supporting a global revision of the species group in its entirety.
Morphological terms were matched as closely as possible to the Hymenoptera Anatomy Ontology (
Wing characters used in this study are illustrated in Figure
The carinae of the propodeum are often considered too highly variable to be of much use in species delimitation (
Propodeum of Ophion demonstrating propodeal carinae and areas. Carinae: ATC = anterior transverse carina; PTC = posterior transverse carina; MLC = median longitudinal carinae; LLC = lateral longitudinal carina; PC = pleural carina; Areas: ASu = area superomedia; AP = area petiolaris; AD = area dentiparis; APE = area postero-externa; AJC = area juxta-coxalis.
Other important characters are the sculpture of the face and clypeus, punctation of the metapleuron, number of flagellomeres, and overall size and colour.
1a | Face and most of thorax black, central flagellomeres approximately quadrate (Plate |
O. dombroskii Schwarzfeld, sp. n. |
1b | Body entirely orange or reddish, central flagellomeres distinctly longer than wide | 2 |
2a | Anterior transverse carina weakly convex in centre (Figure |
3 |
2b | Anterior transverse carina strongly convex in centre (Figure |
5 |
3a | Fore wing length greater than 16 mm; antennae with at least 65 flagellomeres, more in female (Plate |
O. keala Schwarzfeld, sp. n. |
3b | Fore wing length less than 14 mm; antennae with less than 58 flagellomeres | 4 |
4a | Median longitudinal carinae of propodeum strongly convergent posterior to areola, usually arriving at apex of propodeum as a single carina (Figure |
O. idoneus Viereck |
4b | Median longitudinal carinae of propodeum represented by wrinkles posterior to posterior transverse carina, convergent but not fusing into a single carina (Figure |
O. importunus Schwarzfeld, sp. n. |
5a | Yellow-orange base colour, wing length greater than 14 mm, propodeal carinae generally reduced, greatly reduced in male, but apical part of lateral longitudinal carinae expanded as a flange (Figure |
O. aureus Schwarzfeld, sp. n. |
5b | Reddish-orange base colour, wing length 14 mm or less, propodeal carinae more developed, apical part of lateral longitudinal carinae strong but not expanded into a flange (Figure |
6 |
6a | Facial punctures strong and separated by less than their diameter; orbits yellowish, stemmaticum and mesopleural fovea concolourous with base colour (Plate |
O. clave Schwarzfeld, sp. n. |
6b | Facial punctures very shallow and separated by 2–3× their diameter; orbits lacking yellow, concolourous with reddish head, stemmaticum and mesopleural fovea darker than base colour (Plate |
O. brevipunctatus Schwarzfeld, sp. n. |
Diagnosis. Most species are reddish-coloured, lacking yellow markings (except for narrowly on orbits, tegula, mesepimeron and rarely mandibles); early-season (most collected April to June); midtibial spurs nearly equal in length; eyes separated from posterior ocellus by 0.27–1.25× ocellar diameter; facial punctures more widely separated in centre of face than on sides; scutellum strongly carinate (at least partially); propodeum short, posterior area abruptly sloping so that it is nearly vertical at apex, anterior transverse carina (at least in centre) and lateral part of posterior transverse carina (along area juxta-coxalis) strong, otherwise carinae variable, but median longitudinal carinae always convergent apical of area superomedia; petiolar spiracle distinctly anterior to membrane of first metasomal segment.
Description. Head: Eyes convergent to nearly parallel, not strongly bulging (FW/HW: 0.46–0.56); eyes moderately indented (FW/FM: 0.69–0.91); face with minute to medium sized punctures, variously separated, but always closer together on sides and under toruli than in centre; clypeus coriaceous, variously punctate, CH/CAW: 0.49–0.70; mandible with internal angle well–defined (Figure
Mesosoma: Epicnemial carina with pleurosternal angle obtuse (rarely 90°); propodeum short, posterior area abruptly sloping so that it is nearly vertical at apex, ATC strong and convex along ASu, usually entirely strong, but sometimes obsolete or absent laterally, lateral part of PTC (along AJC) strong, usually absent or obsolete along APE, obsolete to strong along ASu; MLC variously developed, but always convergent apical of ASu, LLC mostly absent except strong to absent along AJC; PC strong, rarely connected to spiracle by a very weak carina, but usually not connected.
Wings: Wing L: 8.4–19.7 mm; CI: 0.39–0.84, AI: 1.13–3.31, SDI: 1.03–1.31, ICI: 0.44–1.00, Rs sinuate, ramellus absent to somewhat long, fenestra not extending below prestigma, lacking glabrous area in discocubital cell along Rs+M.
Legs: CL/CW: 1.41–2.37; trochantellus dorsally shorter than width; FL/FW: 6.3–11.6; midtibial spurs nearly equal in length (MTS: 0.70–0.97, usually > 0.80).
Metasoma: First metasomal segment with spiracles distinctly anterior to membrane; first metasomal segment 1.3–1.6× as long as second; second metasomal segment 2.4–4.5× as long as high.
Colour: Uniformly reddish, without yellow markings except orbit (usually narrowly), tegula, mesepimeron and rarely mandibles; females with ovipositor sheath concolourous with apex of metasoma; one species with head and mesosoma predominately black.
Seasonality. Early-flying species, most dates of capture in Canada are May to mid-June. Unusual dates of captures are one specimen that was collected in July, and one collected in August. At least in the northern Nearctic region, most other early-flying species can be distinguished from this species group by their distinct yellow markings.
Biology. There is virtually nothing known about the biology or hosts of the Nearctic species. The one exception is that O. idoneus has been recorded from the noctuid, Sunira bicolorago (Guenée) (Crumb 1924). The Palearctic species, O. scutellaris Thomsen, is a parasitoid of noctuids that overwinter as feeding larvae and are fully grown by spring (
Remarks. The O. scutellaris species group can be further divided into two monophyletic subgroups (see text). One group (A) includes O. idoneus, O. scutellaris, O. keala and O. importunus. It can be recognized by the weakly arched anterior transverse carina of the propodeum (so the base of the area superomedia is only slightly convex; Figure
Only a small proportion of collected Ophion will generally be from the O. scutellaris species group. Within the group, O. idoneus is by far the most common Nearctic species.
Holotype: ♀ (MS2249, DNA3965, GenBank KF594539, KF615948) CAN: AB: Porcupine Hills, Skyline Rd; 49.972 -114.087; 29 v 2008; UV trap; J.Dombroskie, J.Walker (CNC)
Paratypes: 7 ♀♀, 1 ♂. CAN: AB: 2 ♀♀ (MS2244, DNA3980, GenBank KF594552, KF615947; MS2238, DNA3960, GenBank KF594534, KF615950) Same data as holotype (CNC); 1 ♂, 1 ♀ (MS2235, DNA3904, GenBank KF594480, 615945; MS2237) Same data as holotype except date 28 v 2008 (CNC); 1 ♀ (MS8647, DNA6515, GenBank KF594730, KF615943, KF616332) Porcupine Hills, Skyline Rd, 49.972 -114.087, 15 vi 2009, UV trap J.Dombroskie, B.Brunet (CNC); 1 ♀ (MS7801, DNA7327, GenBank KF594917, KF615946) 62 km WNW of Dixonville, Mixedwood retention patch in clearcut, Site 7, 56.685 -118.641, 26 v 2008, UV trap, B.Bodeux (CNC); 1 ♀ (MS7912, DNA7324, GenBank KF594914, KF615944) 62 km WNW of Dixonville, Mixedwood retention patch in clearcut, Site 4, 56.684 -118.644, 7 vi 2008, UV trap, B.Bodeux (CNC); 1 ♀ (MS7746) 54 km NW of Dixonville, Mixedwood forest, 56.86 -118.31, 11 vi 2007, UV trap, B.Bodeux (CNC).
The name for this species is derived from keala, the Hawaiian word for path, as this large and distinctive species presents a rare clear path within the morphologically homogeneous jungle that is Ophion. It is a noun in apposition - and is also the name of the species author’s daughter.
♀: Wing L: 18.6–19.7 mm, Flag: 69–73; ♂: Wing L: 16.2 mm, Flag: 65; Very large species, uniformly dark reddish-orange with interocellar area often darker; hind femur long and slender (FW/FH: 8.4–11.6), scutellum strongly carinate.
Head:Female: Eyes convergent in frontal view; stemmaticum weakly raised, sulci surrounding stemmaticum complete; IOD/OL: 0.27–0.44; OOD/OL: 0.11–0.22; occipital carina rounded, often very slightly wavy, with a very small peak in the centre; OC/OL: 0.42–0.68; temple receding, approximately equal to width of eye in lateral view; CH/CAW: 0.52–0.63, coriaceous with evenly distributed medium-sized punctures, separated by approximately 1–2× their diameter, punctures smaller basally; face weakly coriaceous, with small punctures separated by 1–2× their diameter, closer on sides than in centre; FW/FH: 1.24–1.49; antennae with 69–73 flagellomeres; F1: 3.56–4.50; F20: 1.54–2.24; MS/MW: 0.33–0.47; GI/MW: 0.46–0.69; Male: Same, except: Eyes more weakly convergent, OOD/OL: 0.44; OC/OL: 0.81; flagellomeres: 65.
Mesosoma: Mesoscutum polished, evenly punctate with minute punctures separated 1–2× their diameter; mesopleuron coriaceous, densely punctate with small to medium-sized relatively shallow punctures, separated by approximately their diameter; subpolished with minute punctures separated by 2–3× their diameter above mesopleural fovea; epicnemial carina with pleurosternal angle obtuse, distinctly so in male, rounded (see variation); SL/SW: 1.47–1.76, lateral carinae strong along almost entire length of scutellum, slightly weaker in male; metapleuron strongly coriaceous, punctures approximately equal to those of posterior corner of mesopleuron, equally or slightly less dense.
Propodeum: ATC strong, very slightly arched above ASu; PTC weak to absent along ASu, represented by small crests at MLC, obsolete to absent along AD, very strong along AJC where it is expanded as a flange; MLC obsolete, absent to very faintly represented by parallel or slightly convergent wrinkles along ASu, present as a series of stronger convergent wrinkles along AP; LLC weak to obsolete along AJC, otherwise absent; PC strong, connected to spiracle by a very weak to obsolete carina; spiracular area sloping, subpolished with minute shallow punctures separated by 1–2× their diameter; posterior area weakly wavy-wrinkled with minute punctures to PTC, then more distinctly wrinkled to apex.
Wings:Female: Wing L: 18.6–19.7 mm, CI: 0.54–0.70, AI: 1.42–2.01, SDI: 1.13–1.27, ICI: 0.61–0.93; wings slightly brownish with black veins, stigma reddish-brown, ramellus very short to somewhat long, fenestra restricted to area under stigma; Male: Same, except wing L: 162 mm.
Legs:Female: CL/CW: 1.94–2.37; FL/FW: 8.41–11.63, MT1/MT2: 2.06–2.33; MTS: 0.78–0.88 (♀); Male: Same, except CL/CW: 1.83; MTS: 0.91.
Metasoma: First metasomal segment often abruptly (sometimes gradually) expanded from petiole to postpetiole, spiracles sometimes raised on tubercles.
Colour: Reddish-orange; stemmaticum often slightly darker, mandibles usually slightly paler; orbit (especially posterior to eye), tegula and dorsal part of mesepimeron yellowish; notauli and margins of mesocutum slightly darker.
Propodeal carinae of species from the O. scutellaris species group. All images are of female specimens; male specimens have the same basic arrangement, but often with weaker carinae. a keala b idoneus (arrow indicates fused carinae) c importunus d clave e aureus (arrow indicates expanded, flange-like carina) f brevipunctatus g dombroskii.
One ♀ with all punctures slightly more sparsely distributed than in the remaining specimens; One ♀ with pleurosternal angle of epicnemial carina approximately 90° and somewhat sharp.
All collections have been from late May until mid-June.
Holotype: ♂ USA: Kansas: Douglas Co. ft. 900; May; U. of K., Lot 878, 8ub (SEMC)
Other material: 109 ♀♀, 83 ♂♂, 2 unknown: CAN: AB: 2 ♀♀ (MS12403, MS12405) 11 km NE of Lacombe, J.J. Collett Natural Area, N-facing slope, P. glauca, B. papyrifera, 835m, 52.553 -113.641, 18 vi 2009, UV, C.D. Bird (CNC); 4 ♀♀, 1 ♂ (MS11824–28) 5km NEE of Dunstable, George Lake Research Station, aspen forest, MT-7, 53.957 -114.130, 25-29 v 2007, Malaise, M. Schwarzfeld (UASM); 11 ♀♀, 6 ♂♂ (MS13317, MS13319–20, MS13322–26, MS13328–36) same data except date is 29 v-6vi 2007 (CNC); 2 ♀♀, 6 ♂♂ (MS12920, MS12922–28) 5km NEE of Dunstable, George Lake Research Station; Black spruce forest; MT-8, 53.957 -114.128, 29 v-6 vi 2007, Malaise, M. Schwarzfeld (UASM); 4 ♀♀ (MS12433–35, MS12444) 8 km NW of Winfield, Bird East Poplar Creek quarter, mixed woods, 900m, 53.01 -114.50, 12 vi 2010, UV, C.D. Bird (CNC); 1 ♂ (MS12335) same data except date is 15 v 2010 (UASM); 1 ♂ (MS11457) same data except date is 16 v 2009 (UASM); 1 ♀ (MS13925) same data except date is 21 v 2011 (UASM); 1 ♀ (MS2261) 8 km SE Sherwood Park, aspen forest, 53.478 -113.229, 12-15 v 2008, MV light, G.R. Pohl (UASM); 1 ♀ (MS7373) Bragg Creek, 50.917 -114.533, 15 viii 2007, at light, F. Sperling (CNC); 1 ♀ (MS4026), Calgary, Edgemont, 51.115 -114.142, 18 v 2010, light, T. Pike (CNC); 3 ♀♀ (MS2275, DNA3910, GenBank KF594486, KF615953; MS2280; MS2274) Edmonton, nr. Fulton Ravine, 53.545 -113.439, 15-16 v 2008, light, G. Anweiler (CNC); 1 ♂ (MS2284) same data except date is 17 v 2008 (UASM); 1 ♀ (MS9688, DNA3974, GenBank KF594547, KF615956) same data except date is 23 v 2009 (CNC); 2 ♀♀ (MS2286, MS2290) same data except date is 24 v 2008 (UASM); 1 ♀, 1 ♂ (MS5632, DNA3933, GenBank KF594507, KF615951; MS4301) same data except date is 26 v 2011 (CNC); 9 ♀♀ (MS13809, MS13811, MS13815, MS13820-22, MS13826-28) same data except date is 27 v 2011 (UASM); 3 ♀♀ (MS12169, MS12171–72) same data except date is 31 v 2009 (UASM); 7 ♀♀ (MS13834–35, MS13842–43, MS13858, MS13862) same data except date is 6-8 vi 2011 (UASM); 5 ♀♀, 2 ♂♂ (MS13763–65, MS13767, MS13757, MS13774, MS13782) same data except date is 9-11 vi 2011 (UASM); 2 ♂♂ (MS78, MS80) same data except date is v 2007 (CNC); 2 ♀♀ (MS2879, MS2886) Edmonton, Edith Ravine, 53.510 -113.622, 12 v 2010, UV light, J. Acorn (UASM); 1 ♀ (MS2885) same data except date is 27 v 2010 (UASM); 1 ♀ (MS2234, DNA5552, GenBank KF594576, KF615958) Edmonton, inside building, 53.521 -113.521, J. Dombroskie (CNC); 3 ♂♂ (MS100, MS103, MS107) Edmonton, River Valley at U.Alberta, 53.529 -113.519, 28 v 2007, Sweep, M. Schwarzfeld (CNC); 1 ♀ (MS2770, DNA6551, GenBank KF594763, KF615949) EMEND site, 48 km NW of Dixonville; Decid. forest, uncut, 56.7525 -118.3282, 28 v-10 vi 2008, Malaise, 852-2, M. Schwarzfeld (CNC); 2 ♀♀ (MS12420, MS12423) Erskine, 5 Maple Close, backyard with aspen, 830m, 52.322 -112.883, 19 v 2010, UV, C.D. Bird (UASM); 1 ♀, 1 ♂ (MS9763–64) George Lake Research Site, 53.953 -114.120, 29 v 2007, Sweep, M. Schwarzfeld (UASM); 1 ♂ (MS39) George Lake Research Site, Aspen forest, 53.957 -114.125, 25 v 2007, Sweep, M. Schwarzfeld (UASM); 1 ♀ (MS3613) N. Wyndham-Carseland Provincial Park, 50.8366 -113.4347, 31 v 2008, light, T. Pike (UASM); 2 ♀♀ (MS5650, DNA3977, GenBank KF594549, KF615957; MS5651, DNA3950, GenBank KF594524, KF615952) Pigeon Lake, Itaska, 53.072 -114.072 3 vi 2008, UV light, F. Sperling (CNC); 1 ♀ Pigeon Lake, Itaska, May 28, 2006, F.A.H. Sperling coll.; 9 ♀♀ 14 ♂♂ (MS12460, MS12463, MS12472, MS12474–83, MS12485–93, MS12495) Rochon Sands Prov. Park, 15 km N Erskine, 720m, aspen, chokecherry, aspen, 52.46 -112.88, 2 vi 2010, UV, C.D. Bird (UASM); 1 ♂ (MS12197) Spruce Grove, 13 km South, 53.4 -113.9, 28 v-2 vi 1989, Malaise, A.T. Finnamore (CNC); 1 ♀ (MS12265) Summer Village of Gull Lake, 52.460 -113.947, 31 v 2009, UV trap, J.H. Acorn (UASM); 1 ♂ (MS13966) Wintering Hills West, 51.2552 -112.6261, 29 v 2011, net, J. Dupuis (UASM); BC: 1 ♂ (ENT008-002270) Robson, “?” v 1951, H.R. Foxlee, Ex. H.R. Foxlee Collection U.B.C. 1971 (RBCM); 1 ♂ (ENT008-005274) same data except date is 14 v 1954 (RBCM); 1 ♀ (ENT008-002250) same data except date is 15 v 1951 (RBCM); 1 ♂ (ENT008-002274) same data except date is 15 v 1954 (RBCM); 1 ♀, 1 ♂ (ENT008-002252, ENT008-002275) same data except date is 17 v 1954 (RBCM); 1 ♂ (ENT008-002230) same data except date is 29 v 53 (RBCM); 1 ♂ (ENT008-002239) same data except date is 3 v 51 (RBCM); 1 ♂ Robson, 8 v 1954, H.R. Foxlee (UBCZ); MB: 2 ♀♀ 10 mi. S. of Winnipeg, 1 vi 1973, coll. C. Starr, at UV light (SEMC); 2 ♀♀ same data except date is 30 v 73 (SEMC); 1 ♀ Brandon, 29 v 49, Light Trap (NFRC); ON: 4 ♀♀, 2 ♂♂ (MS2927, MS2932–34, MS2940, MS2943) Bells Corners, Monaghan Forest, 45.272 -75.808, 18 v 2010, Light, B.C. Schmidt (UASM); 1 ♀, 1 ♂ (MS8020, MS8039) Bells Corners, Stony Swamp, 45.295 -75.830, 25 v 2008, B.C. Schmidt (UASM); 1 ♂ (MS12164) Bells Corners, Stony Swamp, 45.295 -75.830, 3 v 2010, MV light, J. Dombroskie, B.C. Schmidt (UASM); 1 ♀, 1 unknown (MS10755, DNA5554, GenBank KF594578, KF615955; MS10754) Grenadier Is., St. Lawrence Islands Nat. Pk., Carya grove, 44.4 -75.9 10–21 vi 1994, YPT, Coll. CNC Hym Team (CNC); 1 ♀ (MS10710) same data except date is 24 v–9 vi 1994 (CNC); 7 ♂♂ (MS10713, DNA6552, GenBank KF594764, KF615960; MS10711; MS10716–17; MS10719–21) same data except trap is Malaise, date is 24 v–9 vi 1994 (CNC); 1 ♀ (MS10751) same data except trap is Malaise, date is 3–13 v 1994 (CNC); 1 ♀ (MS8007, DNA3976, GenBank KF594548, KF615961) Leeds Grenville Co., Long Mtn. 44.487 -76.008, 7 vi 2008, B.D. Schmidt (CNC); 2 ♀♀ (MS10807, MS10813) same data except date is 21 v 2009 (UASM); 5 ♀♀, 22 ♂♂ (MS10793, DNA5717, GenBank KF594635, KF615959; MS10794, DNA5552, GenBank KF594576, KF615958; MS10779, DNA5551, GenBank KF594575, KF615954; MS10768–78, MS10780–92) Ottawa, city garden, 45.356 -75.707, 5 v-5 vi 2008; Malaise, Coll. H. Goulet (CNC); 1 ♂ Waterloo Co., Cambridge, malaise, Skevington & Cannings (RBCM); SK: 1 ♀ S’toon, May 22, 1940, D.R. Foskett (RBCM); USA: MI: 1 ♀ Ag. Coll. Mich 5–23 93 22 (CUIC); 1 ♂ Ag. Coll. Mich 5-23 95 22 (CUIC); NY: 1 ♀, 1 ♂ Ithaca, v 23 1936 (CUIC); 1 unknown Orient, L.I. June 2, 1932, Roy Latham/ Roy Latham Collection (CUIC).
Smallest species within the species group; MLC fused immediately apically of ASu, therefore appears Y-shaped; ramellus usually absent or extremely short; Wing L: 8.4–11.5 mm, Flag: 51–57.
Head: Eyes weakly convergent in frontal view; stemmaticum very weakly raised, sulci not complete; IOD/OL: 0.58–1.25, OOD/OL: 0.25–0.43 (♀), 0.35–0.92 (♂); occipital carina rounded, often very slightly rippled or wavy, usually with a small peak at centre, OC/OL: 0.58–0.95; temple receding, approximately equal to eye width in lateral view; clypeus convex in lateral view and distinctly separated from face, coriaceous, with small regular shallow punctures only slightly larger than on face, separated by approximately their diameter, CH/CAW: 0.57–0.70; face subpolished to very weakly coriaceous, with small shallow punctures separated by approximately their diameter, slightly more dense on sides; FW/FH: 1.28–1.45; antennae with 51–57 flagellomeres, F1: 2.93–4.00; F20: 1.33–2.00; MS/MW: 0.50–0.82; GI/MW: 0.46–0.73.
Mesosoma: Mesoscutum subpolished with very small to minute regular punctures separated by approximately 2× their diameter; mesopleuron weakly coriaceous, strongly closely punctate, punctures separated by their diameter ore less, above mesopleural fovea subpolished with small punctures separated by 1–3× their diameter; metapleuron slightly more coriaceous, punctures similar in size to those of mesopleuron, only slightly less dense; epicnemial carina with pleurosternal angle obtuse; scutellum with lateral carina strong along most of length, SL/SW: 1.21–1.69.
Propodeum: ATC strong, weakly arched above ASu; PTC weak to obsolete along ASu, raised into small crests where intersects with MLC, absent along AD, strong along AJC; MLC obsolete and weakly convergent along ASu, strongly convergent just apical to ASu, so that the carinae fuse into one stronger carina for most of AP; LLC almost absent, short carina in AJC may be remnant of this carina; PC strong, not connected to spiracle; spiracular area slightly sloping, nearly horizontal, subpolished with shallow minute indistinct punctures, separated by 2–3× their diameter; posterior area wavy-wrinkled with very shallow indistinct punctures basally, more wrinkled apically.
Wings: Wing L: 8.4–11.5 mm, CI: 0.48–0.84, AI: 1.95–3.31, SDI: 1.03–1.29, ICI: 0.44–0.80; wings with veins dark brown, stigma light brown with apex whitish-gray, ramellus completely absent or represented by a minute vestigial stub, rarely somewhat long, fenestra not extending below prestigma.
Legs: CL/CW: 1.41–1.85, FL/FW: 6.31–7.85, MT1/MT2: 2.09–2.50; MTS: 0.71–0.89.
Metasoma: Expansion from petiole to postpetiole gradual.
Colour: Reddish-orange, mandibles, palps and legs slightly paler; orbits, tegula and dorsal part of mesepimeron yellowish.
All collection records are from May and June, with the exception of a single specimen collected on Aug 15 from Bragg Creek, Alberta.
Common and wide-spread early-season species. Most small dark reddish individuals, lacking yellow notauli, collected in May will be this species, at least in the northern Nearctic region. The original description (
Holotype: ♀ (MS12343, DNA6907, GenBank KF594814, KF615963, KF616346) CAN: AB: 8 km NW of Winfield, Bird East Poplar Creek quarter, mixed woods, 900m; 53.01 -114.5; 15 v 2010; UV; C.D.Bird (CNC).
Paratypes: 2 ♀♀. CAN: AB: 1 ♀ (MS13904) 8 km NW of Winfield, Bird East Poplar Creek quarter, mixed woods, 900m, 53.01 -114.5, 21 v 2011, UV, C.D.Bird (CNC). ON: 1 ♀ (MS12153) Bells Corners, Stony Swamp, 45.295 -75.83, 3 v 2010, MV light, J.Dombroskie, B.C.Schmidt (CNC).
This species is most similar to O. idoneus. Since idoneus is the Latin word for “suitable” or “proper”, the name for this species is derived from the Latin word importunus, meaning “unsuitable”.
Similar to O. idoneus, but can be recognized by the larger size, LMC convergent but not meeting (thus propodeum lacks Y-shaped carinae), and the long ramellus. Wing L: 12.6–13.3 mm, Flag: 54–57.
Head: Eyes weakly convergent in frontal view; stemmaticum weakly raised, sulci not complete; IOD/OL: 0.53–0.85, OOD: 0.15–0.23; occipital carina rounded, very slightly rippled or wavy, OC/OL: 0.80–1.00; temple receding, approximately equal to eye width in lateral view; clypeus convex, weakly coriaceous, with small regular punctures only slightly larger than on face, separated by approximately their diameter, CH/CAW: 50–0.60; face subpolished with very small punctures separated by slightly more than their diameter in the centre and slightly less on the sides; FW/FH: 1.25–1.29; antennae with 54–57 flagellomeres, F1: 3.43–3.69; F20: 1.54–1.83; MS/MS: 0.46–0.58; GI/MW: 0.46–0.54.
Mesosoma: Mesoscutum subpolished with very small regular punctures separated by 1–2× their diameter; mesopleuron and metapleuron coriaceous, strongly closely punctate, punctures separated by less than their diameter; area of mesopleuron above mesopleural fovea subpolished with minute punctures separated by 1–3× their diameter; epicnemial carina: pleurosternal angle obtuse, rounded; scutellum with lateral carina strong along most of length, SL/SW: 1.55–1.60; Propodeum: ATC strong, moderately arched above ASu; PTC present along AJC, otherwise absent; MLC very weak and weakly convergent along ASu, stronger at apex of ASu, obsolete and represented by strongly convergent wrinkles along AP; LLC represented by indistinct wrinkles along AJC, otherwise absent; PC strong, sometimes weakly connected to spiracle by an obsolete carina; spiracular area sloping, subpolished with small punctures separated by less than their diameter; posterior area wavy-wrinkled with very shallow punctures basally, more wrinkled apically.
Wings: Wing L: 12.6–13.3 mm, CI: 0.60–0.67, AI: 1.58–1.89, SDI: 1.06–1.22, ICI: 0.62–0.80, wings with veins black, stigma reddish-brown with apex whitish-gray, ramellus long, fenestra not extending below prestigma.
Legs: CL/CW: 1.61–2.00, FL/FW: 8.25–8.87, MT1/MT2: 2.19–2.26; MTS: 0.79–0.87.
Metasoma: Expansion from petiole to postpetiole somewhat abrupt.
Colour: Reddish-orange, orbits and sometimes mandibles yellow, tegula and dorsal part of mesepimeron yellowish, scutellum slightly paler than base colour.
All three collection records are in May.
Holotype: ♀ (MS12193, DNA6944, GenBank KF594831, KF615972, KF616352): CAN: AB: Spruce Grove, 13 km South, 53.4 -113.9, 28 v-2 vi 1989, Malaise, A.T.Finnamore (CNC)
Paratypes: 13 ♀♀, 3 ♂♂. CAN: AB: 2 ♀♀(MS12198–99) Same data as holotype (CNC); 1 ♂, 3 ♀♀ (MS13746, DNA7383, GenBank KF594963; MS13749, DNA7381, GenBank KF594961; MS13748; MS13752) Jenner Bridge, S. Jenner, riparian willow/sagebrush shrub, 50.844 -111.154, 2 vi 2010, UV trap, G.Anweiler (CNC); 1 ♀ (MS1689, DNA7391, GenBank KF594970) Jenner rodeo grounds, 50.842 -111.151, 07 vi 2007, Light, G.Anweiler (CNC); 1 ♂, 1 ♀ (MS71, DNA7382, GenBank KF594962; MS72) Jenner rodeo grounds, 50.842 -111.151, 09 v 2007, UV trap, M.Schwarzfeld (CNC); 1 ♀ (MS1434) Jenner rodeo grounds, 50.842 -111.151, 26 v 2007, UV trap, J.Dombroskie & G.Anweiler (CNC); 1 ♀ (MS12386) 17 km S of Stettler, Lowden Springs Conserv. Area, aspen/buckbrush/grassland, 822m, 52.154 -112.713; 26 v 2010; UV; C.D.Bird (CNC); 1 ♂ (MS11578) 17 km S of Stettler, Lowden Springs Conserv. Area, native prairie, 825m, 52.154 -112.712, 24 v 2009, UV, C.D.Bird (CNC). MB: 1 ♀ Brandon, 29.v.49, light trap (NFRC); ON: 2 ♀♀ (MS10758, MS10763) Leeds, Grenville Co. Long Mtn, 44.487 -76.008, 27 iv 2009, B.C.Schmidt (CNC); 1 ♀ Waterloo Co., Cambridge, malaise, 18-21.v.1992, Skevington & Cannings (RBCM)
“Clave” is the Spanish word for key, and refers to the arrangement of the propodeal carinae, which resemble an old-fashioned keyhole. It is also the fundamental rhythm in salsa music, which was undoubtedly playing as this species was being described. It is a noun in apposition.
ATC of propodeum U-shaped above ASu; stemmaticum raised; occipital carina rounded; face coarsely densely punctate (most punctures separated by less than their diameter), and punctures connected by strong microreticulation; Wing L: 10.5–12.4 mm; Flag: 52–60. This species is most similar to O. brevipunctatus, but can be distinguished by the sculpture of the face, the more extensive yellow orbits and the smaller size. This species is also similar to O. idoneus and O. importunus, but can be most easily distinguished by the strongly convex ATC.
Head: Eyes slightly convergent to nearly parallel in frontal view; stemmaticum raised, sulci surrounding stemmaticum complete or nearly so; IOD/OL: 0.53–1.00; OOD/OL: 0.13–0.29; occipital carina rounded, OC/OL: 0.67–0.92; temple receding, approximately equal to width of eye in lateral view; clypeus weakly convex in lateral view and very weakly separated from face, coriaceous with irregularly scattered coarse punctures, often interspersed with at least a few minute punctures, smaller and denser towards lateral and dorsal margins, less dense than on face (see variation), CH/CAW: 0.49–0.67; face evenly coarsely punctate, most punctures separated by less than their diameter and connected by strong microreticulation, smaller on sides than in centre (see variation); FW/FH: 1.18–1.26; antennae with 52–60 flagellomeres, F1: 3.00–4.17, F20: 1.36–1.83; MS/MW: 0.43–0.63; GI/MW: 0.36–0.67
Mesosoma:Female: Mesoscutum polished to subpolished (see variation), evenly, shallowly punctate with small to minute punctures; mesopleuron coriaceous with strong punctures separated by approximately their diameter, polished with sparse minute punctures above mesopleural fovea; epicnemial carina of females with pleurosternal angle approximately 90°, and more or less sharp, varying from slightly acute to slightly obtuse; SL/SW: 1.23–1.58; lateral carina strong along anterior third to two-thirds of scutellum; metapleuron coriaceous with strong punctures separated by approximately their diameter; Male: Same, except SL/SW: 1.43–1.67; pleurosternal angle of epicnemial carina rounded and slightly obtuse.
Propodeum:Female: ATC strong, strongly arched above ASu (so anterior margin of ASu is highly convex), occasionally weak to obsolete along AE; PTC present (occasionally obsolete) along ASu, usually produced as small crests where intersects with MLC, obsolete to absent between AD and APE, strong along AJC; MLC weak and slightly convergent along ASu, strong (occasionally weak) and distinctly convergent (but not meeting) along AP; LLC weak to strong along AJC, otherwise absent; PC strong, not connected to spiracle; spiracular area slightly to strongly sloping, closely punctate, subpolished to coriaceous; posterior area rugulopunctate basally, strongly wrinkled apically; Male: Same pattern but all carinae tending to be weaker.
Wings: Wing L: 10.5–12.4 mm, CI: 0.39–0.59, AI: 1.13–2.16, SDI: 1.16–1.30, ICI: 0.52–0.93; veins dark brown to black, stigma light brown; fore wing with ramellus absent or present, discocubital cell entirely trichiose except for well-defined fenestra under stigma (not extending below prestigma).
Legs:Female: CL/CW: 1.58–1.93; FL/FW: 7.00–8.59 (♀), 7.00–7.18 (♂), MT1/MT2: 2.01–2.26 (♀); MTS: 0.72–0.88; Male: Same, except MT1/MT2: 1.69–2.02.
Metasoma: Sides of petiole gently divergent from spiracles to apex in females, more abruptly expanded at spiracles in males.
Colour: Reddish-orange; palps, mandibles and/or scutellum sometimes slightly paler; orbits narrowly yellowish; tegula and mesepimeron yellowish.
There is some geographic variation in this species, with less morphological variation in specimens from within each locality than there is between localities.
2 ♀♀ have clypeal punctures more densely and regularly spaced than in remaining specimens; 2 ♀♀ have facial punctures slightly smaller and facial microreticulation slightly weaker that in the remaining specimens; 1 ♂, 1 ♀ (from same locality) with mesoscutum coriaceous.
This species has been collected from late April until early June.
Holotype ♀ (MS7632, DNA3970, GenBank KF615968) CAN: AB: Machesis Lk Forest Prov. Rec. Area, 32 km W Fort Vermilion, 318m, Jack pine forest, 58.325 -116.578, 10 vi 2008, UV trap, D.&S. Macaulay (CNC).
Paratypes 9 ♂♂ (MS2318, DNA3961, GenBank KF594535, KF645966; MS2324, DNA3911, GenBank KF594487, KF615965; MS2310, DNA3975, GenBank KF615969; MS2311, MS2313–16, MS2320) nr. Tangent Park Campgrnd, 23km S of Peace River, meadow in spruce/aspen, 56.092 -117.542; 7 v 2008, UV trap, D.Macaulay (CNC).
The name aureus is the Latin word for golden, referring to the golden-orange colour of this species.
Wing L: 14.3–17.1 mm, Flag: 65–69. Largest species in subgroup B; stemmaticum raised with sulci complete; reduced propodeal carinae with posterior area strongly wrinkled; more golden-orange (less reddish) than the other species in this species group.
Head: Eyes weakly convergent in frontal view; stemmaticum distinctly raised, sulci surrounding stemmaticum complete and deeply impressed; IOD/OL: 0.65–0.80, OOD/OL 0.26–0.33; occipital carina rounded, OC/OL: 0.73–1.00; temple receding, approximately equal to width of eye in lateral view; clypeus moderately convex in lateral view and weakly separated from face, slightly more convex in males, coriaceous, sparsely punctate in males, very sparsely punctate in female, with irregularly sized (minute to coarse) punctures, punctures denser basally and especially laterally, CH/CAW: 0.0.53–0.64; face coriaceous with small punctures separated by slightly more than their diameter, smaller and denser on orbits than in centre, FW/FH: 1.25–1.41; antennae with 65–69 flagellomeres, F1: 3.25–4.00; F20: 1.47–2.38; MS/MW: 0.52–0.76; GI/MW: 0.32–0.66.
Mesosoma: Mesoscutum densely evenly punctate, subpolished with minute punctures in males, weakly coriaceous with slightly larger punctures in female; mesopleuron coriaceous with strong punctures separated by approximately their diameter in female, punctures smaller and separated by approximately 2× their diameter in males, subpolished with smaller sparser punctures above mesopleural fovea, epicnemial carina with pleurosternal angle 90° to slightly obtuse, rounded (occasionally somewhat sharp); scutellum with lateral carina strong at base, present along basal third to half; SL/SW: 1.13–1.38; metapleuron coriaceous (strongly coriaceous in female), punctures smaller and sparser than on mesopleuron.
Propodeum:Female: ATC strong along ASu, strongly arched (so anterior margin of ASu is highly convex), otherwise absent; PTC present along ASu, raised into crests where intersects with MLC, absent along APE, extremely strong (raised as a flange) along AJC; MLC very weak and slightly convergent along ASu, stronger and nearly parallel along AP; LLC absent; two strong longitudinal wrinkles in AJC, one of which probably represents remnant of LLC; PC strong, not connected to spiracle; propodeum short, spiracular area sloping, punctate, strongly coriaceous, posterior area rugose; Male: similar, but with all carinae much less developed: ATC present as vestige in centre; PTC present only as slight crests where intersects with MLC and as a flange along AJC, but much less raised than in female; MLC convergent, weak to obsolete, often reduced to longitudinal wrinkles; punctures on spiracular area very shallow.
Wings: Wing L: 14.3–17.1 mm, CI: 0.41–0.60, AI: 1.57–2.36, SDI: 1.17–1.31, ICI: 0.70–1.00; veins brown to dark brown, stigma light brown with an apical white spot, fenestra mostly confined to area below stigma, in several specimens indistinctly extending below prestigma; trichiae more or less sparse along Rs+M; ramellus long
Legs:Female: CL/CW: 1.67, FL/FW: 7.95; MT1/MT2: 1.22; MTS: 0.86; Male: CL/CW: 1.69–2.08; FL/FW: 8.11–9.63; MT1/MT2: 1.22–2.19; MTS: 0.70–0.90.
Metasoma: Sides of petiole gently divergent from spiracles to apex, some males with more abrupt expansion at spiracles
Colour: Uniformly golden-orange, males less reddish than other members of the species group, female distinctly less reddish. Orbits narrowly and indistinctly yellow, more yellow posterior to eye; tegula and extreme dorsal part of mesepimeron yellowish.
This species had been collected in only two collection events, the males in mid-May and the one female in mid-June.
The name is derived from the Latin words brevis and punctatus, referring to the unusually shallow punctures of the face.
Wing L: 14.0 mm; Flag: 67; ATC strongly arched above ASu; stemmaticum raised with sulci complete; facial punctures small, very shallow, widely separated but connected with strong microreticulation; temple strongly receding, 0.6× eye width (other species in this group with temple approximately equal to eye width); stemmaticum dark, no yellow on orbits.
Head: Eyes convergent in frontal view; stemmaticum raised, sulci complete; IOD/OL: 0.69, OOD/OL: 0.13; occipital carina rounded, OC/OL: 0.78; temple strongly receding, 0.6× as long as eye width in lateral view; CH/CAW 0.58× apical width, only slightly convex in lateral view, weakly separated from face; clypeal punctures irregularly sized, sparsely, irregularly distributed across coriaceous clypeus; punctures of face small, very shallow, separated by 2–3× their diameter, connected by strong microreticulation; FW/FH: 1.27; antennae with 67 flagellomeres; F1: 3.6; F20: 1.8; MS/MW: 0.5; GI/MW: 0.5.
Mesosoma: Mesoscutum subpolished, evenly punctate with minute punctures; mesopleuron coriaceous with strong punctures separated by approximately their diameter, varying to subpolished with smaller punctures anteriodorsally; subpolished with minute punctures above mesopleural fovea; epicnemial carina with pleurosternal angle slightly obtuse; SL/SW: 1.5, strongly carinate along the anterior third; metapleuron coriaceous with shallow medium-sized punctures, more sparsely distributed than on mesopleuron.
Propodeum: ATC strong, strongly arched along ASu (so anterior margin of ASu strongly convex); PTC obsolete in centre, strong at intersection with MLC and for a short distance along AD, otherwise obsolete along AD, very strong along AJC; MLC obsolete and slightly convergent along ASu, obsolete (represented by wrinkles) and strongly convergent along AP; LLC present along AJC, very weakly represented at intersection with ATC, otherwise absent; PC strong, not connected to spiracle; spiracular area sloping, coriaceous with small shallow punctures; posterior area weakly rugopunctate, becoming wrinkled apically.
Wings: Wing L: 14.0 mm, CI: 0.62, AI: 1.73, SDI: 1.29, ICI: 0.80, wing veins dark brown, stigma reddish-brown, fenestra restricted to area below stigma, trichiae slightly sparser below prestigma, ramellus long.
Legs: CL/CW: 2.0; FL/FW: 9.5, MT1/MT2: 2.1; MTS: 0.78.
Metasoma: Sides of petiole gradually expanding at spiracles.
Colour: Uniformly reddish-orange; stemmaticum distinctly darker, dark reddish-brown; palps and scutellum very slightly paler than base colour, tegula and extreme dorsal part of mesepimeron dark yellowish, mesopleural fovea slightly darker than base colour.
The one collection record is from May 13.
This species is named for Jason Dombroskie, who was kind enough to collect the only known specimen of this species on an otherwise rainy and utterly unsuccessful moth-collecting trip.
Wing L: 10.8 mm, Flag: 51; The head and thorax of this species are almost entirely black, making this species easily recognizable. It also has unusually short antennal segments, widely separated ocelli and a long, narrow scutellum.
Head: Eyes slightly convergent in frontal view; stemmaticum slightly raised, sulci surrounding stemmaticum complete; IOD/OL: 1.20, OOD/OL: 0.45; occipital carina rounded, OC/OL: 1.20; temple receding, approximately equal to width of eye in lateral view; clypeus 0.5× as high as apical width, coriaceous, punctures irregularly sized (coarse to very small) and sparsely, irregularly distributed, denser on sides, CH/CAW: 0.50; face with medium-sized punctures, approximately separated by their diameter and connected with strong microreticulation, smaller on sides of face; FW/FH: 1.34; antennae short, 51 flagellomeres, F1: 2.7, F20: 1.1; MS/MW: 0.6; GI/MW: 0.8.
Mesosoma: Mesoscutum coriaceous, evenly punctate with minute punctures, separated by several times their diameter; mesopleuron and metapleuron strongly coriaceous, evenly, coarsely punctate with punctures separated by their diameter or less; mesopleuron above mesopleural fovea subpolished, punctures minute, separated by 2-3× their diameter; epicnemial carina pleurosternal angle obtuse; scutellum with lateral carina strong along anterior half, SL/SW: 1.8.
Propodeum: ATC strong and strongly arched along ASu (so anterior margin of ASu strongly convex), weak to obsolete along AD; PTC mostly obsolete, represented by wrinkles, forming small crests where intersects with MLC, strong along AJC, especially strong at propodeal apophysis; MLC obsolete, faintly represented by wrinkles along ASu and even more indistinctly along AP; LLC present as a wrinkle along AJC, otherwise absent; PC strong, not connected to spiracle; spiracular area sloping, coriaceous with numerous small punctures; posterior area weakly wrinkled and punctate, more wrinkled apically.
Wings: Wing L=10.8 mm, CI: 0.49, AI: 1.39, SDI: 1.15, ICI: 0.83; wings with veins dark brown, stigma light brown, fenestra not extending below prestigma, ramellus short.
Legs: CL/CW: 1.5, FL/FW: 7.2, MT1/MT2: 3; MTS: 0.85.
Colour: Head: black; orbits, temple, vertex, mandibles, palps and clypeus except for extreme base reddish-orange; Mesosoma: black; mesocutum (except margins and base of notauli), scutellum, anterior margin and an indistinct area in the centre of mesopleuron, apical margin of propodeum, apical half of coxae and legs reddish-orange; mesepimeron reddish-orange ventrally and yellowish dorsally; metasoma reddish-orange; ovipositor sheath same colour as metasoma.
The one collection record is from May 20.
This species is unusual because of its extensive black markings. The unusually short antennae and black markings indicate that this species may be diurnally active (
We are grateful to Gavin Broad (BMNH) for loaning specimens and sending legs for sequencing, for sharing his knowledge of Ophion, and for his valuable feedback on an earlier version of this manuscript. We are sincerely thankful to everyone who helped collect the material included in this paper (John Acorn, Gary Anweiler, Charley Bird, Bryan Brunet, Jason Dombroskie, Julian Dupuis, Doug Macaulay, Boyd Mori, Ted Pike, and Chris Schmidt) as well as to the assistants and volunteers who helped in the lab and field (Heather Bird, Jessica Edwards, Jeremy Fitzpatrick, Huda Khan, Maria Khan, Kerilynn Mercier, Cheryl Tebby, and Crystal Tebby). We are appreciative of the curators and collection managers who lent specimens: Andy Bennett (CNC), Claudia Copley (RBCM), Jason Dombroskie (CUIC), Steve Halford (SFUC), Dave Langor and Greg Pohl (NFRC), Karen Needham (UBCZ), and especially Zachary Falin (SEMC) for loaning the type specimen of Ophion idoneus, along with additional material. A huge thank you to Rylee Isitt and Don Griffiths for their assistance with photographing specimens. Finally, we would like to thank Pascal Rousse and an anonymous reviewer for their helpful feedback on an earlier version of the manuscript. This work was supported by the the Sustainable Forest Management Network, ACA Grants in Biodiversity (supported by the Alberta Conservation Association), Natural Sciences and Engineering Research Council (NSERC) and Alberta Ingenuity (Alberta Innovates) student scholarships to MDS and an NSERC Discovery grant to FAHS.
List of all specimens included in molecular and morphometric analyses.
Accession |
Provenance |
Lat-long | Date | Coll. |
Species |
GenBank | |||
---|---|---|---|---|---|---|---|---|---|
Sex | COI | ITS2 | 28S | ||||||
O. scutellaris species group | |||||||||
DNA3970 | AB: Machesis Lake PRA | 58.325 -116.578 | 10 vi 2008 | DM | aureus C,G | F | KF615968 | ||
DNA3911 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | KF594487 | KF615965 | KF616311 |
DNA3961 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | KF594535 | KF615966 | |
DNA3975 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | KF615969 | ||
MS2313 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
MS2315 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
MS2316 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
MS2314 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
MS2311 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
MS2320 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | aureus C,G | M | |||
DNA3939 | ON: nr. Carp | 45.385 -76.008 | 13 v 2008 | BCS | brevipunctatus C,G | F | KF594513 | KF615967 | KF616314 |
DNA7381 | AB: Jenner | 50.844 -111.154 | 2 vi 2010 | GGA | clave C,G | M | KF594961 | ||
DNA7382 | AB: Jenner | 50.842 -111.151 | 09 v 2007 | MDS | clave C,G | M | KF594962 | KF615970 | |
DNA7383 | AB: Jenner | 50.844 -111.154 | 2 vi 2010 | GGA | clave C,G | F | KF594963 | ||
DNA7391 | AB: Jenner | 50.842 -111.151 | 07 vi 2007 | GGA | clave C,G | F | KF594970 | ||
MS72 | AB: Jenner | 50.842 -111.151 | 9 v 2007 | MDS | clave C,G | F | |||
MS1434 | AB: Jenner | 50.842 -111.151 | 26 v 2007 | JJD | clave C,G | F | |||
MS13752 | AB: Jenner | 50.844 -111.154 | 2 vi 2010 | GGA | clave C,G | F | |||
MS13748 | AB: Jenner | 50.844 -111.154 | 2 vi 2010 | GGA | clave C,G | F | |||
DNA6944 | AB: Spruce Grove | 53.4 -113.9 | 28 v–2 vi 1989 | ATF | clave C,G | F | KF594831 | KF615972 | KF616352 |
MS12198 | AB: Spruce Grove | 53.4 -113.9 | 28 v–2 vi 1989 | ATF | clave C,G | F | |||
MS12199 | AB: Spruce Grove | 53.4 -113.9 | 28 v–2 vi 1989 | ATF | clave C,G | F | |||
MS12386 | AB: Stettler Co. | 52.154 -112.713 | 26 v 2010 | CDB | clave C,G | F | |||
MS11578 | AB: Stettler Co. | 52.154 -112.713 | 24 v 2009 | CDB | clave C,G | M | |||
n/a | MB: Brandon | 49.83 -99.96 | 29 v 1949 | ? | clave C | F | |||
n/a | ON: Cambridge | 43.46 -80.52 | 18–21 v 1992 | S&C | clave C | F | |||
MS10758 | ON: Leeds | 44.487 -76.008 | 27 iv 2009 | BCS | clave C,G | F | |||
MS10763 | ON: Leeds | 44.487 -76.008 | 27 iv 2009 | BCS | clave C,G | F | |||
DNA6548 | SK: nr. Newton Lake | 49.301 -107.764 | 20 v 2011 | JJD | dombroskii C,G | F | KF594760 | KF615971 | KF616341 |
ENT008-002252 | BC: Robson | 49.34 -117.70 | 17 v 1954 | HRF | idoneus C | F | |||
ENT008-002250 | BC: Robson | 49.34 -117.70 | 15 v 1951 | HRF | idoneus C | F | |||
ENT008-005274 | BC: Robson | 49.34 -117.70 | 14 v 1954 | HRF | idoneus C | M | |||
ENT008-002230 | BC: Robson | 49.34 -117.70 | 29 v 1953 | HRF | idoneus C | M | |||
DNA6551 | AB: 48 km NW Dixonville | 56.753 -118.328 | 28 v–10 vi 2008 | MDS | idoneus C,G | F | KF594763 | KF615949 | |
MS7373 | AB: Bragg Cr. | 50.917 -114.533 | 15 viii 2007 | FAHS | idoneus C,G | F | |||
MS4026 | AB: Calgary | 51.115 -114.142 | 18 v 2010 | TP | idoneus C,G | F | |||
DNA3908 | AB: Edmonton | 53.521 -113.521 | 20 v 2008 | JJD | idoneus C,G | F | KF594484 | KF615962 | |
MS100 | AB: Edmonton | 53.527 -113.519 | 28 v 2007 | MDS | idoneus C,G | M | |||
MS107 | AB: Edmonton | 53.527 -113.519 | 28 v 2007 | MDS | idoneus C,G | M | |||
MS103 | AB: Edmonton | 53.527 -113.519 | 28 v 2007 | MDS | idoneus G | M | |||
DNA3933 | AB: Edmonton | 53.545 -113.439 | 15 vi 2008 | GGA | idoneus C,G | M | KF594507 | KF615951 | |
DNA3910 | AB: Edmonton | 53.545 -113.439 | 15–16 v 2008 | GGA | idoneus C,G | F | KF594486 | KF615953 | |
MS2274 | AB: Edmonton | 53.545 -113.439 | 15–16 v 2008 | GGA | idoneus G | F | |||
MS2280 | AB: Edmonton | 53.545 -113.439 | 15–16 v 2008 | GGA | idoneus G | F | |||
MS2284 | AB: Edmonton | 53.545 -113.439 | 17 v 2008 | GGA | idoneus G | M | |||
DNA3974 | AB: Edmonton | 53.545 -113.439 | 23 v 2009 | GGA | idoneus C,G | F | KF594547 | KF615956 | |
MS2286 | AB: Edmonton | 53.545 -113.439 | 24 v 2008 | GGA | idoneus G | F | |||
MS12169 | AB: Edmonton | 53.545 -113.439 | 31 v 2009 | GGA | idoneus G | F | |||
MS12171 | AB: Edmonton | 53.545 -113.439 | 31 v 2009 | GGA | idoneus G | F | |||
MS12172 | AB: Edmonton | 53.545 -113.439 | 31 v 2009 | GGA | idoneus G | F | |||
MS78 | AB: Edmonton | 53.545 -113.439 | v 2007 | GGA | idoneus C,G | M | |||
MS80 | AB: Edmonton | 53.545 -113.439 | v 2007 | GGA | idoneus G | M | |||
MS12420 | AB: Erskine | 52.322 -112.883 | 19 v 2010 | CDB | idoneus C,G | F | |||
MS9764 | AB: George L | 53.953 -114.120 | 29 v 2007 | MDS | idoneus G | F | |||
MS9763 | AB: George L | 53.953 -114.120 | 29 v 2007 | MDS | idoneus G | M | |||
MS39 | AB: George L | 53.957 -114.125 | 25 v 2007 | MDS | idoneus G | M | |||
MS11824 | AB: George L | 53.957 -114.125 | 25–29 v 2007 | MDS | idoneus G | F | |||
MS11825 | AB: George L | 53.957 -114.125 | 25–29 v 2007 | MDS | idoneus G | F | |||
MS11826 | AB: George L | 53.957 -114.125 | 25–29 v 2007 | MDS | idoneus G | F | |||
MS11827 | AB: George L | 53.957 -114.125 | 25–29 v 2007 | MDS | idoneus G | F | |||
MS11828 | AB: George L | 53.957 -114.125 | 25–29 v 2007 | MDS | idoneus G | M | |||
MS13334 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus C,G | M | |||
MS13333 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus C,G | M | |||
MS13319 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS13330 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS13331 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS13332 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS13336 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus C,G | F | |||
MS13320 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13322 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13324 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13325 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13326 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13328 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13329 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13335 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS13317 | AB: George L | 53.957 -114.125 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS12920 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12922 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12924 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12926 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12927 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12928 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | M | |||
MS12923 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
MS12925 | AB: George L | 53.957 -114.128 | 29 v–6 vi 2007 | MDS | idoneus G | F | |||
n/a | AB: Pigeon Lake | 53.072 -114.072 | 38 v 2006 | FAHS | idoneus C,G | M | |||
DNA3977 | AB: Pigeon Lake | 53.072 -114.072 | 3 vi 2008 | FAHS | idoneus C,G | F | KF594549 | KF615957 | |
DNA3950 | AB: Pigeon Lake | 53.072 -114.072 | 3 vi 2008 | FAHS | idoneus C,G | F | KF594524 | KF615952 | |
MS12475 | AB: Rochon Sands PP | 52.46 -112.88 | 2 vi 2010 | CDB | idoneus C,G | F | |||
MS12491 | AB: Rochon Sands PP | 52.46 -112.88 | 2 vi 2010 | CDB | idoneus C,G | M | |||
MS2261 | AB: Sherwood Park | 53.478 -113.229 | 12–15 v 2008 | GRP | idoneus G | F | |||
MS12197 | AB: Spruce Grove | 53.4 -113.9 | 28 v–2 vi 1989 | ATF | idoneus C,G | M | |||
MS12444 | AB: Winfield | 53.01 -114.50 | 12 vi 2010 | CDB | idoneus C,G | F | |||
MS12434 | AB: Winfield | 53.01 -114.50 | 12 vi 2010 | CDB | idoneus G | F | |||
MS12433 | AB: Winfield | 53.01 -114.50 | 12 vi 2010 | CDB | idoneus G | F | |||
MS12435 | AB: Winfield | 53.01 -114.50 | 12 vi 2010 | CDB | idoneus G | F | |||
MS12335 | AB: Winfield | 53.01 -114.50 | 15 v 2010 | CDB | idoneus C,G | M | |||
MS11457 | AB: Winfield | 53.01 -114.50 | 16 v 2009 | CDB | idoneus G | M | |||
MB: Brandon | 49.83 -99.96 | 29 v 1949 | ? | idoneus C | F | ||||
MB: Winnipeg | 49.74 -97.13 | 1 vi 1973 | CS | idoneus C | F | ||||
MS2933 | ON: Bells Corners | 45.272 -75.808 | 18 v 2010 | BCS | idoneus C,G | F | |||
MS2927 | ON: Bells Corners | 45.272 -75.808 | 18 v 2010 | BCS | idoneus C,G | M | |||
MS2932 | ON: Bells Corners | 45.272 -75.808 | 18 v 2010 | BCS | idoneus C,G | M | |||
DNA5554 | ON: Grenadier Is. | 44.4 -75.9 | 10–21 vi 1994 | CNC | idoneus | ? | KF594578 | KF615955 | KF616320 |
DNA6552 | ON: Grenadier Is. | 44.4 -75.9 | 24 v–9 vi 1994 | CNC | idoneus C,G | M | KF594764 | KF615960 | |
DNA3976 | ON: Leeds | 44.487 -76.008 | 7 vi 2008 | BCS | idoneus C,G | F | KF594548 | KF615961 | |
MS10780 | ON: Ottawa | 45.356 -75.707 | 5 v–5 vi 2008 | HG | idoneus C,G | M | |||
DNA5551 | ON: Ottawa | 45.356 -75.707 | 5 v–5 vi 2008 | HG | idoneus C,G | M | KF594575 | KF615954 | |
DNA5552 | ON: Ottawa | 45.356 -75.707 | 5 v–5 vi 2008 | HG | idoneus C,G | F | KF594576 | KF615958 | |
DNA5717 | ON: Ottawa | 45.356 -75.707 | 5 v–5 vi 2008 | HG | idoneus C,G | F | KF594635 | KF615959 | |
n/a | SK: Saskatoon | 52.1 -106.6 | 22 v 1940 | DRF | idoneus C | F | |||
DNA6907 | AB: 8 km NW Winfield | 53.01 -114.50 | 15 v 2010 | CDB | importunus C,G | F | KF594814 | KF615963 | KF616346 |
MS13904 | AB: 8 km NW Winfield | 53.01 -114.50 | 15 v 2010 | CDB | importunus C,G | F | |||
MS12153 | ON: Bells Corners | 45.295 -75.830 | 3 v 2010 | BCS | importunus C,G | F | |||
MS7746 | AB: 54 km NW Dixonville | 56.86 -118.31 | 11 vi 2007 | BBB | keala C,G | F | |||
DNA7324 | AB: 62 km WNW Dixonville | 56.684 -118.644 | 7 vi 2008 | BBB | keala C,G | F | KF594914 | KF615944 | |
DNA7327 | AB: 62 km WNW Dixonville | 56.685 -118.641 | 26 v 2008 | BBB | keala C,G | F | KF594917 | KF615946 | |
DNA3904 | AB: Porcupine Hills | 49.972 -114.087 | 28 v 2008 | JJD | keala C,G | M | KF594480 | KF615945 | KF616307 |
DNA3960 | AB: Porcupine Hills | 49.972 -114.087 | 28 v 2008 | JJD | keala C,G | F | KF594534 | KF615950 | |
DNA3965 | AB: Porcupine Hills | 49.972 -114.087 | 29 v 2008 | JJD | keala C,G | F | KF594539 | KF615948 | |
DNA3980 | AB: Porcupine Hills | 49.972 -114.087 | 29 v 2008 | JJD | keala C,G | F | KF594552 | KF615947 | |
DNA6515 | AB: Porcupine Hills | 49.972 -114.087 | 15 vi 2009 | JJD | keala C,G | F | KF594730 | KF615943 | KF616332 |
MS2237 | AB: Porcupine Hills | 49.972 -114.087 | 28 v 2008 | JJD | keala C,G | F | |||
MS11464 | AB: Winfield | 53.01 -114.50 | 16 v 2009 | CDB | A C,G | F | |||
MS4024 | AB: Calgary | 51.115 -114.142 | 18 v 2010 | TP | B C,G | F | |||
MS4310 | AB: Beaver Mines Lake | 49.371 -114.295 | 16 vii 2009 | TP | C C,G | F | |||
MS2322 | AB: nr. Tangent Park | 56.092 -117.542 | 7 v 2008 | DM | D C,G | M | |||
MS11823 | AB: George Lake | 53.957 -114.125 | 25–29 v 2007 | MDS | E C,G | M | |||
n/a | UK | not listed | not listed | scutellaris | EU378720 | ||||
DNA7524 | UK | ? | ? | GRB | scutellaris A | KF595042 | KM982696 | ||
DNA6914 | UK: Pitstone Commons | 51.81 -0.593 | 22 iv–7 v 2011 | GRB | scutellaris A | KF594819 | KF615964 | KF616305 | |
DNA7515 | UK: Radnage | 51.659 -0.858 | 3 v 2008 | AMG | scutellaris A | KF594993 | |||
DNA7558 | UK: Radnage | 51.659 -0.858 | 21 vii 2011 | AMG | scutellaris A | KF595030 | |||
DNA7519 | UK: Westcott | 51.848 -0.962 | 20 iii 2009 | DW | scutellaris B | KF594995 | |||
Not O. scutellaris species group | |||||||||
DNA6501 | AB: 6 km N Guy | 55.607 -117.161 | 22 vii–16 viii | BAM | Ophion sp. | KF594717 | KF616256 | KF616362 | |
DNA6913 | UK: Aldbury | 51.799 -0.601 | 9 v 2011 | GRB | minutus | KF594818 | KF616300 | KF616360 | |
DNA3902 | AB: Kootenay Plains PRA | 52.064 -116.422 | 14 vii 2007 | MDS | Ophion sp. | KF594478 | KF615984 | KF616308 | |
DNA6550 | SK: nr. Newton Lake | 49.301 -107.764 | 20 v 2011 | JJD | Ophion sp. | KF594762 | KF616150 | KF616343 | |
DNA5559 | BC: Vancouver | 49.2 -123.2 | 3–17 ix 1997 | IK | Ophion sp. | KF594583 | KF615978 | KF616371 | |
DNA5742 | BC: Glacier NP | 51.26 -117.57 | 8 vii 2010 | MDS | Ophion sp. | KF594660 | KF615911 | KF616337 | |
DNA3957 | AB: 21 km S Whitelaw | 55.928 -117.995 | 29 vi 2008 | DM | Ophion sp. | KF594531 | KF615930 | KF616381 | |
DNA5566 | MB: Winnipegosis | 51.651 -99.945 | 16 vi 2007 | HG | Ophion sp. | KF594589 | KF616242 | KF616321 | |
DNA3901 | AB: Kootenay Plains PRA | 52.064 -116.422 | 14 vii 2007 | MDS | Enicospilus sp. | KF594477 | KF594477 |