Illidops naso , Kotenko 2007

Illidops naso (Marshall, 1885) Figs 29–30; Tables 4–5 Apanteles naso Marshall, 1885: 203. Apanteles contortus Tobias, 1964: 224–225, fig. 45. Apanteles crantor Nixon, 1965: 183, figs 205, 209. Apanteles evander Nixon, 1965: 183, fig. 207. Apanteles coresia Nixon, 1973: 198. Other material examined KAZAKHSTAN – [Karaganda Region] • 1 ♂; Kyzylzhar, Monadyr [Street]; [49.98° N, 72.61° E]; [460 m]; 22 Jun. 1958; V.I. Tobias leg.; [V.I. Tobias det.]; paratype of Apanteles contortus Tobias, 1964; CNC, CNCHYM 00168. – [Akmola Region] • 1 ♀; Kokshetau, “Tersakkan W Akmol.”; 15 Jun. 1957; V.I. Tobias leg.; [V.I. Tobias det.]; paratype of Apanteles contortus Tobias, 1964; CNC, CNCHYM 00169. Species concept Our species concept is based on our examination of the female and male paratypes of the junior synonym A. contortus stored in the CNC and the information available in Marshall (1885), Tobias (1964), Tobias & Kotenko (1986), Nixon (1965, 1973, 1976), and Papp (1981, 1988) → *. Ecology / host information Host unknown. Distribution PAL: Afghanistan, Armenia, Azerbaijan, Bulgaria, Croatia, Finland, Georgia, Greece, Hungary, Iran, Kazakhstan, Korea, Kyrgyzstan, Macedonia, Moldova, Mongolia, Romania, Russia (KC, VOR), Serbia, Slovakia, Switzerland, Turkey, Turkmenistan, United Kingdom, Uzbekistan. Molecular data We observed specimens of six barcoding clusters which morphologically rather resemble Illidops naso: BOLDAEI3308 (Figs 2B, 31), BOLD:AEI9858 (Fig. 32), BOLD:AEK1062 (Fig. 33) BOLD:AEO8212 (Fig. 34), BOLD:AAD3865, and a single disparate sequence of a male A. contortus paratype. However, these clusters are far apart from each other with minimum p-distances ranging between 6.03% and 11.35%. The clusters are clearly separated, even though only few specimens are available for some. The Nearest Neighbors (NN) in the BOLD database are different clusters, not identified to species level (compare Table 4). Only BOLD:AAD3865 is quite close to its NN which is BOLD:AEA1257 with a minimum p-distance of 2.37%. However, we were unable to observe any specimens of the NN and the morphology observed by us so far is cryptic in any case, so we cannot draw any conclusions. Most NNs have no species-level identification associated and we do not currently have access to specimens associated with most of these clusters. Only BOLD:AAK4166, which includes representatives from Canada that are stored at the CNC, allowed for some morphological analysis and we found the characters of representatives of this cluster similar to I. cloelia, not the I. naso complex. In addition, the NN BIN of BOLD:AEI9858 includes representatives that we associated with I. cloelia. What we can conclude based on the molecular data available to us is that these clusters likely form a complex of morphologically cryptic species associated with the names I. naso (and its four junior synonyms), I. buteonis and I. dauricus. We attempted matching one of our clusters to a pre-existing name via DNA barcoding by sequencing a male paratype of A. contortus stored at the CNC and collected in Kazakhstan in Kyzylzhar, Karaganda. CNCHYM 00168 was included in the shotgun sequencing of historical DNA and was mapped to the COI sequence of ZSM-HYM-33161-C11. Karaganda is the neighboring province to Akmola, in which the holotype locality “ 6km NE from Ylektykol Lake” is located. However, the sequence of this specimen did not match any of our clusters. We did a BLAST search in BOLD and the sequence matched the publicly available sequence of specimen CGTURK-1496, part of BIN BOLD:AAH1044 from northeastern Turkey with 99.05% similarity. The specimen appears to be a male and we cannot draw any conclusions based on the photo available in the database. Remarks Holotype male, NHMUK. Country of type locality: United Kingdom. The type was examined by Fernandez-Triana et al. (2020). This species is morphologically very similar to I. buteonis and I. dauricus. We do not think that it is currently possible to reliably differentiate these species based on morphological characters. Firstly, because we observed that some characters used by previous authors are quite variable, even in specimens within a single barcoding cluster we associate morphologically with these species. Secondly, we found additional cryptic diversity via DNA barcoding with at least six different barcoding clusters matching the morphology of these three species, which appear in several places in a NJ analysis of Illidops (Fig. 62). We did not find sufficient characters to differentiate these molecular clusters morphologically (see notes on T2 shape and Fig. 5) or unambiguously match any of them to one of the previously described species. These species are likely part of a complex of morphologically cryptic species, hence, in this paper, we refer to them as part of the I. naso complex. In addition to that, the name Illidops naso currently has 4 junior synonyms associated with it (see Table 5). The holotype of I. naso is male, was not reared and no female paratypes are assigned or other female specimens from the same locality mentioned in the original description. The type locality according to the description is: “On the sand-hills at the mouth of the Exe, opposite Star-Cross, Devon.” (Marshall 1885: 203). The holotype of A. contortus is female and a rather long series of female and male specimens is mentioned in the original description, not reared, but partially collected from the same localities. The holotype of A. crantor Nixon, 1965 is female and in the original description only female specimens as material examined are indicated. The holotype of A. evander is female and a single female paratype is mentioned in the original description. The holotype of A. coresia Nixon, 1973 is female and the only specimen associated with this species in the original description. Apanteles coresia was described as part of the metacarpalis -group of Apanteles, not the butalidis -group. Nixon mentions that it is aberrant within this group due to its rather uniformly rugose propodeum. Nixon (1976) synonymized A. crantor under A. contortus, Papp (1981) synonymized A. evander and A. contortus under A. naso without providing details on his reasoning, and A. coresia was synonymized between 1981 and 1986, but we could not find by which author. The association between male and female specimens of I. naso were likely made based on the type series of A. contortus; however, matching female and male specimens of Illidops based on morphology is rather difficult due to the sexual dimorphism within this group observed by previous authors and ourselves. Tobias’ male and female specimens were partially collected from the same single localities, but that is by no means a guarantee that they belong to the same species. We have found 2–3 different BINs associated with the I. naso complex in a single Malaise trap collecting event in several independent cases. We conclude that there is no good way to associate male and females of a species of Illidops unless both specimens were reared from the same host individual (or, less ideal, at least the same host species), or via DNA barcoding. With this strong sexual dimorphism combined with morphological crypsis, we cannot judge whether all of these current synonyms were rightly synonymized, and they will need to be dealt with when attempting to resolve this complex. One of the sequenced specimens (ZSM-HYM-42448-E05, a member of BOLD:AEI9858) was reared by Tina Schulz from Aroga velocella (Zeller, 1839), a species of Gelechiidae.

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