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Research Article
Lochetica ramii sp. nov. – a new species of Lochetica Kriechbaumer, 1892 (Hymenoptera, Ichneumonidae, Phygadeuontinae) from Finland, with a key to world species
expand article infoJuuso Paappanen
‡ Unaffiliated, Kuopio, Finland
Open Access

Abstract

Two tentatively distinct morphological forms belonging to the Darwin wasp genus Lochetica Kriechbaumer, 1892 (Hymenoptera, Ichneumonidae) were found to occur in Finland, although only one species is known in Europe. The identity of the two forms were resolved by examining additional museum material, DNA barcoding and revising the relevant types. Both morphology and molecular results support the recognition of a new species, Lochetica ramii sp. nov., from Finnish specimens – in addition to Lochetica westoni (Bridgman, 1880) already known from Finland. New host associations are given for both species and their ecology is discussed. An identification key is given to the known species of Lochetica of the world to facilitate the recognition of the new species.

Key Words

Parasitoid wasps, Phygadeuontinae, north Europe, Palearctic, saproxylic, identification, key

Introduction

The Darwin wasp genus Lochetica Kriechbaumer, 1892 is a species-poor genus within the megadiverse Ichneumonidae, comprising only four species, all occurring in the Holarctic Region (Yu et al. 2012; Watanabe 2021). Information regarding the life history of the genus Lochetica only accounts for one species, Lochetica westoni (Bridgman, 1880), which has been associated with the crabronid genus Passaloecus Shuckard, 1837 (Barbey and Ferriere 1923; Jonaitis 1981; Kreisch 2000), which nest in above-ground structures such as beetle burrowings in (dead) wood, galls and plant stems (e.g. Blösch (2000)).

For the greater part of the 20th century, two species of Lochetica have been known: L. westoni (Bridgman, 1880) and Lochetica pimplaria (Thomson, 1888), until Horstmann (1972) established the synonymy between the two species. The initial placement of L. westoni in a separate genus (Cecidonomus Bridgman, 1880) likely maintained their separation for such a long period. Townes (1983) revised the genus and provided the first identification key to the genus describing two new species: Lochetica farta from Taiwan and Lochetica agonia from the western United States. Watanabe (2021) described Lochetica japonica from Japan, which increased the number of species to four. All species, except for L. westoni, are known by only a few specimens.

Two tentatively distinct morphological forms of Lochetica were recognised to occur in Finland. This prompted the present study with the aim to resolve their identities. Subsequently, a new species of Lochetica is here described from Finnish specimens and an identification key for all five known species of Lochetica is provided. New host associations are given and the ecology of the genus is discussed.

Methods

Specimen data and depositories

The studied material comprises 88 specimens of Lochetica, which are deposited in the following collections (later referred by the given abbreviations):

MZH Natural History Museum, Helsinki, Finland.

MZLU Lund University Biological Museum, Lund, Sweden.

NCM Norwich Castle Museum, Norwich, United Kingdom.

KPM-NK Kanagawa Prefectural Museum of Natural History, Odawara, Kanagawa, Japan.

RJK Secondary collection of Reijo Jussila, Kuhmo, Finland.

All specimens have been given ID labels by the associated museum. This facilitates later revision of the specimens mentioned in this article. In addition, all but three specimens are deposited in public museums.

Morphology

Morphological terms follow Broad et al. (2018). The following abbreviations are used in the text:

Tergite Metasomal tergite.

Temple ratio The width of the temples, in dorsal view, measured halfway between the posterior ocelli and the occipital carina, divided by the maximum width of the head (in dorsal view).

Ovi-tib ratio The length of the ovipositor projecting beyond the apex of the metasoma (in Lochetica, equal to the length of the ovipositor sheaths) divided by the length of the hind tibia (measured in outer lateral view).

POL:OOL The distance between the lateral ocelli divided by the distance between a lateral ocellus and the compound eye.

OD:OOL The diameter of the lateral ocellus divided by the distance between the lateral ocellus and the compound eye.

The specimens were examined with a common main objective type Olympus SZX16 stereo-microscope with a plan apochromatic objective, using magnifications 7×–115×. Measurements were done with an ocular micrometer, with one eye closed. The length of the body is usually impossible to measure since the body is rarely straight. The fore-wing length has been widely used as a proxy for body size. Here both measures are reported; however, the body length is not an absolute measurement of the specimen, but partly an evaluation on how long the specimen would be if straightened. This is not the ideal method, but was favoured instead of taking an absolute measurement of the specimen regardless of its posture.

Temple ratio and ovi-tib ratio were statistically compared between the species with the Welch t-test. These analyses, as well as the graphs, were done in R (v. 4.3.2) via the RStudio (v. 12.0-369) software. The package stats (v. 4.3.2) was used to perform the t-tests, the package effsize (0.8.1) was used to calculate the effect sizes and the package beeswarm (0.4.0) was used to produce the graphs.

All morphological examinations, measurements and identifications were done by the author in 2023. Photographs of the types of the Cecidonomus westoni, Phygadeuon pimplarius and L. japonica were examined.

Photographs

Figs 3, 6, 7 are focus stacks, composed of dozens of individual photographs, taken by the author with the Olympus OM-5 camera with an extended and reversed Schneider Kreuznach Componon S 50mm F/2.8 lens, using the in-camera pixel shift technology (tripod high-res mode). Individual photographs were stacked with the CombineZP (v. 1.0) software using the Quick Align and Do Stack –macros. The photographs of type material deposited in collections other than the MZH, have been taken with various equipment by the corresponding curators.

Molecular methods

Ten specimens of Lochetica were sequenced for the cytochrome oxidase subunit I (COI) marker (the standard barcode region sensu Hebert et al. (2003)). The tissue sample used for DNA extraction was a single leg, in most cases one of the middle pair, of each specimen. DNA extraction, PCR and sequencing were carried out by the Canadian Centre for DNA Barcoding, according to the prescribed protocols described in deWaard et al. (2008). The forward primer LepF1 (ATTCAACCAATCATAAAGATATTGG) and the reverse primer LepR1 (TAAACTTCTGGATGTCCAAAAAATCA) were used to amplify the selected marker. The Sanger sequencing protocol was used to obtain the sequence.

In addition to these ten sequences, all public COI sequences assorted to BINs with at least one specimen identified as Lochetica were downloaded from the BOLD database (downloaded 26 Nov 2023). These included only three sequences, all from Norwegian specimens.

The resulting combined data of thirteen sequences were aligned with the MUSCLE algorithm (Edgar 2004) within the MEGA XI software (Tamura et al. 2021). The following (default) settings were selected for the alignment, cluster method: UPGMA, gap open penalty: -400, gap extend penalty: 0, min. diag. length (lambda): 24. The alignment was manually examined; however, no further changes were made to the alignment, for example, no sites were removed.

A distance matrix (p-distances) was calculated in R (v. 4.3.2) via the RStudio software (v. 12.0-369), with the package ape (v. 5.7-1). Sites with missing data were deleted in pairwise calculations. The resulting matrix was transferred to Microsoft Excel 2013 (v. 15.0.5153.1000), where the distance measures were obtained. The matrix was also used to create a Neighbour-Joining tree with the same package (ape) in R.

Data sharing

Full data used in this study are available as Suppl. material 1: a spreadsheet file with specimen and morphological data, Suppl. material 2: R script (a simple text file) containing the code to produce the statistical analyses and graphs and Suppl. material 3: aligned sequences in fasta format. The sequences and their associated accession numbers are also available at the GenBank and the BOLD database via the doi: dx.doi.org/10.5883/DS-LOCH24

Results

Morphological comparison

Comparing the mean ovi-tib ratio between L. westoni and L. ramii sp. nov. (Fig. 1A) using the Welch t-test, the ratio is larger in L. westoni (mean 2.8) than in L. ramii sp. nov. (mean 2.1) (t = -22.8, DF = 71.8, p < 0.001; Cohen’s d = -5.3). There are no overlapping specimens with regards to this character, although the gap is rather small: max. for L. ramii sp. nov. is 2.4 and min. for L. westoni is 2.6). The L. japonica specimens overlap with L. ramii sp. nov., but not with L. westoni. The ovi-tib ratio of L. japonica was not statistically compared to other species due to its small sample size.

Figure 1. 

Swarmplots comparing the selected morphological ratios of examined females. A. Comparison of the relative length of the ovipositor. The data on Lochetica japonica Watanabe, 2021 is derived from the original description; B. Comparison of the temple ratio (see the Methods section for definition). Each point represents one specimen, the points are moved horizontally to avoid overlap.

Comparing the mean temple ratio between L. westoni and L. ramii sp. nov. (Fig. 1B) using the Welch t-test, the ratio is larger in L. ramii sp. nov. (mean 0.95) than in L. westoni (mean 0.88) (t = 9.7, DF = 64.6, p < 0.001; Cohen’s d = 2.3). There are numerous overlapping specimens with regards to this character (Fig. 1B).

COI comparison

All specimens yielded long sequences, varying in length from 621 bp to 658 bp. The COI sequences of seven specimens of L. ramii sp. nov. and six specimens of L. westoni were compared. No sequences for the other three species of Lochetica are available. The interspecific distances between the two species range from 7.12% to 8.05%, while the intraspecific distances within the specimens of L. ramii sp. nov. and L. westoni are 0.00–0.16% and 0.00–0.61%, respectively. The barcode gap is distinct (Fig. 2).

Figure 2. 

A simple Neighbour-Joining phylogenetic tree (p-distance) illustrating the barcode gap between the specimens. The scale bar represents 1% divergence. The ID after the species name refers to the sample ID in the BOLD-database (same as the museum (specimen) ID in Finnish material). The specimens marked with an asterisk (*) have not been examined.

Systematics

Lochetica Kriechbaumer, 1892

Lochetica Kriechbaumer, 1892: 340.

Type species

Phygadeuon pimplarius Thomson, 1888. Monotypic.

Diagnosis

Dense punctation and hairs across the body surface. The clypeus wide, weakly separated from the face, anterior edge of clypeus and mandibles covered with very dense short hairs. Temples rather wide. Sternaulus long. Propodeum with complete carination and pentagonal area superomedia. Spiracles of first tergite near the middle. The fore wing with a closed areolet, vein 3rs-m is weakly pigmented; vein 2m-cu long, straight, forming acute angle with vein CU, with one short bulla.

Lochetica westoni westoni (Bridgman, 1880)

Figs 3, 4

Cecidonomus Westoni Bridgman, 1880: 264. Lectotype designated by Horstmann (1972).

Phygadeuon pimplarius Thomson, 1884: 941. Lectotype designation of Townes published by Frilli (1973).

Material examined

Lectotype of Cecidonomus westoni Bridgman, 1880 (photographs examined) United Kingdom (presumably) • 1 ♀; “from Mr. Weston”; “Westoni”; “lektotypus Cecidonomus westoni ♀ Bridgm. Horstm. det. 1970”; “J. B. Bridgman coll. Norwich Museum 1895.40.1124”.

Lectotype of Phygadeuon pimplarius Thomson 1884 (photographs examined) Sweden • 1 ♀; “Öke å” [Skåne, Sjöbo, Övedskloster]; [55.684°N, 13.633°E]; “Lectotypus Phygadeuon pimplarius Thm. Tow. 58”; MZLU 5385:1.

Other material

Finland • 38 ♀♀, 5 ♂♂ [MZH]. Russia • 2 ♀♀ [MZH].

Diagnosis

The female is distinguished by the long epomia (Fig. 3A), long ovipositor (ovi-tib ratio 2.6–3.1 in examined specimens) (Fig. 4E), almost completely red legs (Fig. 4E), yellow tegulae (Fig. 3F), narrow temples (temple ratio 0.80 to 0.95 in examined material) and strong latero-median carinae of the first tergite (Fig. 3E). The male is distinguished by the yellow tegulae, medially white mandibles, long epomia and, to some extent, the strongly rugose sculpture of mesopleuron and metapleuron (Fig. 3C).

Figure 3. 

Lochetica westoni (Bridgman, 1880), a non-type specimen MZH GP.113444 (except for C). A. Mesosoma of the ♀ (lateral view); B. Head of the ♀ (dorsal view); C. Mesopleuron and metapleuron of the non-type ♂ MZH GP.109783 (lateral view); D. Propodeum of the ♀ (oblique posterior view); E. The first tergite of the ♀ (lateral view); F. Tegula of the ♀ (oblique anterolateral view). Scale bars: 0.5 mm.

Figure 4. 

A–C. The Lectotype of Cecidonomus westoni Bridgman, 1880. Photo credit: Tony Irwin. A. Labels; B. Head and part of the mesosoma (dorsal view); C. Habitus (lateral view); D–E. Lectotype of Phygadeuon pimplarius Thomson, 1884. Photo credit: Biological Museum, Lund University (MZLU); D. Labels; E. Habitus (lateral view). Scale bars: 1 mm.

The extent of the red colouration on the metasoma in females varies considerably. In the Finnish specimens, usually only the first tergite is laterally slightly reddish; however, some specimens have extensively red first and second terga. A subspecies Lochetica westoni rufiventris Habermehl, 1919 from Algeria (not examined) has the metasoma completely red.

Distribution

Most European countries, Turkey, Algeria, Japan (Yu et al. 2012), Russia (Jonaitis 1981) and Georgia (Riedel et al. 2018).

Ecology

Both open and forested habitats are used, especially sites with dead wood or other suitable nesting sites for hosts.

The type series has been reared from “galls” (Bridgman 1880), which Morley (1907) later states were the galls of Cynips Kollari (= Andricus kollari (Hartig, 1843)). Passaloecus gracilis (Curtis, 1834) have been recorded to nest in galls of A. kollari (Lomholdt 1975). Barbey and Ferriere (1923) reared L. westoni from the nests of Passaloecus gracilis inside empty beetle burrowings in the bark of Pinus sylvestris L. They also state that L. westoni overwinters as a larva in a silky cocoon and postulated that the female must oviposit to the host while the nest is still under construction, since the ovipositor is not long enough to reach the host larvae behind the outer resin plug that encloses the nest. Jonaitis (1981) also reported Passaloecus gracilis nesting inside branches as a host and Kreisch (2000) reported the host Passaloecus insignis (Vander Linden, 1829).

New rearing records published in this study are the following: a male reared from Passaloecus monilicornis Dahlbom, 1842 (MZH GP.109784), a female reared from Passaloecus eremita Kohl, 1893 (MZH GP.109755) and a male reared from Passaloecus borealis Dahlbom, 1844 (MZH GP.109783). In all cases, no additional information how the host was confirmed were given and no host remains or cocoon were included in the pin. Furthermore, P. borealis is extremely difficult to separate from Passaloecus turionum; thus, the record could refer to either of the two species. In addition, one specimen was reared from an unidentified Passaloecus larva inside an artificial nest (an Angelica L. stem) on the wall of a building (MZH GP.109758).

In conclusion, L. westoni utilises most species of the genus Passaloecus (recorded from P. gracilis, P. monilicornis, P. insignis, P. eremita, P. borealis/turionum and Passaloecus sp.) nesting in (dead) wood (including buildings and other structures), galls and most likely also plant stems.

Lochetica westoni rufiventris Habermehl, 1919

Lochetica pimplaria f. rufiventris Habermehl, 1919: 110. Lectotype designated by Horstmann (1991).

Remarks

This taxon is known from Algeria and, according to Horstmann (1991), it is distinguished from L. w. westoni only by its completely red metasoma. Horstmann considered it a subspecies and, since I have not examined the type or any material from the Mediterranean, his view of the taxon is followed here.

Lochetica japonica Watanabe, 2021

Fig. 5

Lochetica japonica Watanabe, 2021: 110, figs 47A–H, fig. 62J, fig 65Q, fig. 66Q.

Material examined

Holotype (photographs examined) Japan • 1 ♀; Tochigi Pref., Nasushiobara City, Shiobara, Utou-sawa; 22–28 May 2008; T. Matsumura leg.; Malaise-trap; KPM-NK 81989.

Diagnosis

The female is distinguished by the long epomia (cf. Fig. 3A), weak latero-median carinae of the first tergite, clearly defined punctation of the body (especially the vertex (Fig. 5E) and the area externa of the propodeum (Fig. 5B)), pronotum with transverse crenulae, wide (developed) temples (Fig. 5E), a shallow transverse depression followed by a low elevation on the vertex (Fig. 5E), almost completely black legs (Fig. 5D) and mandibles (Fig. 5A). The male is unknown.

Figure 5. 

The holotype of Lochetica japonica Watanabe, 2021. Photo credits: Kyohei Watanabe. A. Head (oblique anteroventral view); B. Propodeum (oblique posterodorsal view); C. Labels; D. Habitus (lateral view); E. Head and mesoscutum (dorsal view).

Distribution

Japan (Watanabe 2021).

Ecology

Unknown.

Lochetica ramii sp. nov.

Figs 6, 7

Material examined

Holotype Finland • 1 ♀ (pinned); Sb [Savonia borealis], Kuopio, Mökinoja; “6966126:3529339” [62.7976°N, 27.575°E]; 18 Jun 2021; Juuso Paappanen leg.; “niitty, hirsirakennuksen seinä” [meadow, on a wall of a log barn]; COI sequence GP.113446; MZH GP.113446.

Paratypes (37 ♀♀, 1 ♂) Finland • 1 ♀; “Finby” [Särkisalo], [Finnarv, Vedudden]; [60.1143°N, 22.9485°E]; 14 Jul 1950; R. Elfving leg.; MZH GP.109724 • 4 ♀; “Keuru” [Keuruu]; [62.257°N, 24.708°E]; [1928]; Hellén leg.; MZH GP.109731, MZH GP.109732, MZH GP.109733, MZH GP.109779 • 1 ♀; Aitolahti; [61.5481°N, 23.8825°E]; 13 Jul 1932; V. Saarinen leg.; MZH GP.109725 • 1 ♀; “H:fors” [Helsinki]; [60.1797°N, 24.9393°E]; Hellén leg.; MZH GP.109734 • 1 ♀; Helsinki; [60.1797°N, 24.9393°E]; 11 Sep 1965; V. J. Karvonen leg.; MZH GP.109727 • 1 ♀; Ilomantsi, Koitajoki; 62.93310°N, 31.43978°E; 18 Jun–8 Aug 2023; Juuso Paappanen leg.; ~ 2 years ago dead standing Picea abies; [Tereshkin trap]; MZH GQ.7565 • 7 ♀; Ilomantsi, Koitajoki; 62.93390°N, 31.44084°E; 18 Jun.–8 Aug. 2023; Juuso Paappanen leg.; dying ø 39 cm standing Picea abies; [Tereshkin trap]; MZH GQ.7566, MZH GQ.7567, MZH GQ.7568, MZH GQ.7569, MZH GQ.7570, MZH GQ.7571, MZH GQ.7572 • 1 ♀; Karjalohja; [60.2609°N, 23.7138°E]; 25 Jul 1969; J. Perkiömäki leg.; MZH GP.109729 • 1 ♀; Kb [Karelia borealis], Ilomantsi, Koitajoki, Palokangas; 6989825:3724830 [62.9419°N, 31.4305°E]; 13 Jun–11 Jul 2022; Seppo Karjalainen & Maarit Similä leg.; [prescribed burn of a Pinus sylvestris dominated forest]; MZH GP.109742 •1 ♀; Kb [Karelia borealis], Ilomantsi, Patvinsuon kansallispuisto, Raanisuo; 7003901:3694317 [63.0855°N, 30.8475°E]; 11 Jul–17 Aug 2022; Seppo Karjalainen & Maarit Similä leg.; COI sequence BOLD sample id GP.113447; MZH GP.113447 • 1 ♀; Kb [Karelia borealis], Joensuu, Kuhasalo; 62.58126°N, 29.73376°E; 14 Aug 2017; Juuso Paappanen leg.; [Picea abies dominated forest]; MZH GP.109738 • 1 ♀; Kb [Karelia borealis], Lieksa, Kuikkasuo; 7034335:3673955 [63.3683°N, 30.4779°E]; 18 Aug–17 Sep 2021; Sampsa Malmberg & Maarit Similä leg.; [prescribed burn of a Pinus sylvestris dominated forest]; MZH GP.109743 • 1 ♂; Kn [Ostrobottnia kajanensis], Ristijärvi, pappilan peltoaukea; 71571:35582 [64.5074°N, 28.2125°E]; em. 30 Apr. 2008; Reima Leinonen leg.; koivupölkkykasvatus [reared from trap nest (birch wood block)]; COI sequence BOLD sample id FICH-001702; RJK FICH-001702 • 1 ♀; Ok [Ostrobottnia kajanensis], Kuhmo, Elimyssalo, Risulampi; 7122708:3660871 [64.1659°N, 30.3083°E]; 29 Jul–26 Aug 2022; Sampsa Malmberg et al. leg.; [prescribed burn of a Pinus sylvestris dominated forest]; COI sequence BOLD sample id GP.113448; MZH GP.113448 • 1 ♀; Ok [Ostrobottnia kajanensis], Paltamo, Itkonpuro; 7147421:526944 [64.45127°N, 27.55991°E]; 9 Jul–27 Jul 2023; I. Immonen & M. Laaksonen leg.; [moist/shaded forest next to a stream]; MZH GQ.7563 • 1 ♀; Ok [Ostrobottnia kajanensis], Talaskangas, Sopenjoki; 7098580:503358 [64.01405°N, 27.06866°E]; 22 Jul–6 Aug 2021; I. Immonen leg.; [moist/shaded forest next to a stream]; MZH GQ.7564 • 1 ♀; Parikkala, [Laurila]; [61.56718°N, 29.51232°E]; 12 Jul 1945; Hellén leg.; MZH GP.109726 • 1 ♀; Sa [Savonia australis], Mäntyharju, Kousa; 68022:34674 [61.327°N, 26.3922°E]; 23 Aug 2019; Juuso Paappanen leg.; Pinus sylvestris dominated forest; MZH GP.109741 • 1 ♀; Sa [Savonia australis], Ruokolahti, [Inkilänsaari]; [61.32062°N, 28.75517°E]; 26 Jul 1948; W. Hellén leg.; MZH GP.109735 • 1 ♀; Sb [Savonia borealis], Savonranta; 69077:6026 [62.261°N, 28.9761°E]; 13 Jul–10 Aug 1996; P. Martikainen leg.; dead aspen; Window trap; MZH GP.109730 • 1 ♀; Sb [Savonia borealis], Vieremä, Luvejoki; 63.98762°N, 26.74378°E; 21 Jul–17 Aug 2022; Juuso Paappanen leg.; a large, ~ 2 years ago dead standing Picea abies (shaded); MZH GP.109739 • 1 ♀; Ta [Tavastia australis], Vanaja; [60.978°N, 24.562°E]; 1954; Erkki Valkeila leg.; e Passaloecus monilicornis; MZH GP.109736 • 2 ♀; same as preceding; 1955; MZH GP.109737, MZH GP.109780 • 1 ♀; Tb [Tavastia borealis], Jyväskylä, Kotalamminmäki; 69028:34311 [62.2248°N, 25.6763°E]; 4 Jul 2019; Juuso Paappanen leg.; dying Picea abies; COI sequence BOLD sample id GP:113445; MZH GP:113445 • 1 ♀; Tb [Tavastia borealis], Jyväskylä, Roninmäki; 69014:34326 [62.2125°N, 25.7056°E]; 5 Jul–18. Jul 2019; Juuso Paappanen leg.; recently dead standing Picea abies; MZH GP.109740 • 1 ♀; V [Regio aboënsis], Nousiainen, Tepastus; 6735:344 [60.6991°N, 24.1525°E]; 4 Jul–8 Jul 2006; Reijo Jussila leg.; RJK FICH-001012 • 1 ♀; V [Regio aboënsis], Turku, Paattinen; 6727:244 [60.5763°N, 22.3374°E]; 29 Oct 2006; Reijo Jussila leg.; RJK FICH-001013 • 1 ♀; Vihti, Siikajärvi; [60.2882°N, 24.5214°E]; 22 Aug 1965; V. J. Karvonen leg.; MZH GP.109728.

Comparative diagnosis

The female differs from L. japonica by the vertex uniformly sloping from the ocelli to the occiput, with shallow, almost granulate sculpture (Fig. 6A) (with a shallow transversely orientated depression, low elevation and large, rather deep punctures in L. japonica (Fig. 5E)); shallow, transversely orientated surface sculpture of area externa of propodeum, punctures very weakly discernible (Fig. 6F) (clearly defined deep punctures in L. japonica (Fig. 5B)); the pronotal sulcus without transverse crenulae (present in L. japonica); predominantly red legs (almost completely black in L. japonica); black or dark brown tegulae (Fig. 6C) (yellowish-brown in L. japonica (Fig. 5E)); predominantly red mandibles (black with reddish tips in L. japonica).

Figure 6. 

Lochetica ramii sp. nov. holotype ♀ (except for D). A. Head (dorsal view); B. Head (anterior view); C. Mesoscutum and tegulae (dorsal view); D. Mesopleuron of the paratype ♀ MZH GP.113447 (lateral view); E. The first tergite; F. The propodeum (posterodorsal view). Scale bars: 0.5 mm.

The female differs from L. westoni by the short and weak latero-median carinae of the first tergite, not extending beyond the level of spiracles (Fig. 6E) (strong carinae, extending beyond spiracles in L. westoni (Fig. 3E)); shorter and weaker crenulae of the sternaulus (Fig. 6D) (longer and stronger in L. westoni Fig. 3A); shorter ovipositor (ovi-tib ratio 1.9–2.4) (2.6–3.1 in L. westoni); black or dark brown tegulae (Fig. 6C) (yellow in L. westoni (Fig. 3F)). The male differs from L. westoni primarily by the black tegulae, contrasting with the pale yellow humeral plates (Fig. 7D, cf. Fig. 6C) (yellow tegulae, similarly coloured to the humeral plates in L. westoni (cf. Fig. 3F)). The sculpture of the mesopleuron and metapleuron is somewhat weaker (Fig. 7C) (stronger in L. westoni (Fig. 3C). The trochanters and hind tarsi dorsally darker in L. ramii sp. nov.; however, both of the latter characters are rather subtle and possibly become obsolete when more specimens are revealed.

Figure 7. 

Lochetica ramii sp. nov. A. Holotype ♀ habitus (lateral view); B. Tip of the ovipositor of the holotype ♀ (lateral view); C. Mesopleuron and metapleuron of the ♂ paratype RJK FICH-001702 (lateral view); D. Paratype ♂ habitus RJK FICH-001702 (lateral view). Scale bars: 1 mm.

Both sexes are differentiated from L. agonia by the long epomia ventrad to pronotal sulcus (cf. Fig. 3A) (absent or short in L. agonia) and by the black or dark brown tegulae (Fig. 6C) (yellow (♀) or “ivory” (♂) in L. agonia). The females are also distinguished by the weak, but angulate nodus of the ovipositor (Fig. 7B) (very weak and rounded in L. agonia).

Female is differentiated from L. farta by the area petiolaris of the propodeum having rather weak transverse sculpture concentrated to the sides, usually without sculpture medially (Fig. 6F) (the area petiolaris with very dense transverse sculpture in L. farta); strong latero-median longitudinal carina separating the area postero-externa from the area petiolaris (weak in L. farta).

Description

Female (holotype and paratypes). Length. Body length 5.6 mm (4.6–6.3 mm (n = 36)) excluding the ovipositor, fore wing length 3.6 mm (3.2–4.3 mm (n = 37)).

Head. Punctation dense, shallow, individual punctures indistinctly delimited, coalescing, vertex more granulate than punctate (Fig. 6A). Surface moderately matt, covered in dense, short (shorter than diameter of ocellus), white hairs, except clypeus with slightly longer, light brown hairs; hairs very dense on anterior edge of clypeus and mandibles (Fig. 6B); antennae, including scapes, with short hairs only. First flagellar segment without plate sensillae, second with one, remaining with gradually increasing number, being almost absent from posterior side of flagellum. Eyes bare. Occipital carina complete, joining hypostomal carina 0.6× basal length of mandible away from mandibular base. Malar space 0.8× as long as the basal width of mandible. Face very wide, with low central prominence (Fig. 6B). In anterior view, head 0.9× wider than high; in dorsal view, the medial length of head from occipital carina to level of antennal sockets 0.5× maximum width of the head; temple ratio 0.95 (paratypes: 0.9–1.0), POL:OOL 1.1, OD:OOL 0.5 (see Methods section for definitions). Both antennae with 20 flagellar segments (paratypes (n = 35): 20–23), all segments more than 1.5× longer than wide, decreasing in length towards apex. Width of segments fairly uniform across length of flagellum, increasing only very slightly apically. First flagellar segment 5× as long as wide and 1.3× longer than scape. Scape 1.7× as long as wide. Penultimate (next to last) flagellar segment 1.5× as long as wide. Head predominantly black, mandibles medially red with proximal fourth and teeth black (paratypes: from completely red with only teeth black to about basal third of mandibles black). Maxillary and labial palpi light brown, gradually darker proximally. Anterior side of scapes red (paratypes: from predominantly black with red patches anteriorly to predominantly red with dark spot posteriorly). Anterior side of two proximal flagellar segments red (paratypes: 0 to 4 basal-most segments with red anterior faces). Pedicel with narrow red apical annulus.

Mesosoma. Mesoscutum and scutellum moderately densely and shallowly punctate, similarly to head. Notauli not crenulate, weakly impressed, but long, extending to middle of mesoscutum. Posterior half of mesoscutum obscured by pin in the holotype. Scuto-scutellar groove smooth, without crenulae. Pronotum with same superficial punctures as rest of body, with stronger rugosity posteriorly and medially with larger shiny interstices between sculpture (about 2× wider than the punctures); pronotal sulcus without transverse crenulae; epomia not particularly strong, long, extending near posterior margin of pronotum ventrad to pronotal sulcus. Mesopleuron covered with denser, stronger, coalescing punctures, especially dorsad to sternaulus coalescing to form rugose areas (Fig. 6D); Dorsal part ventrad to subtegular ridge, with larger and distinctly shinier interstices between small punctures, interstices about 2× diameter of punctures. Sternaulus long, almost reaching posterior margin of mesopleuron, crenulae rather weak, longest distinctly weaker and only slightly longer than crenulae on mesopleural furrow (Fig. 6D) (paratypes: from short and weak to as highly raised and slightly longer than longest crenulae on mesopleural furrow). Epicnemial carina and posterior transverse carina of mesosternum complete. Mediosternal groove rather deep, not widened posteriorly with uniformly strong crenulae across its length. Metapleuron strongly rugosepunctate, juxtacoxal carina and metapleural carina distinct and complete (Fig. 6D). Sculpture of propodeum shallow, with indistinct transverse rugosity, individual punctures difficult to discern (Fig. 6F) (paratypes: rugosity variable, but never distinctly punctate); densely hairy. Area petiolaris differs in sculpture from rest of propodeum: shiny and sparsely scattered with longer setae, laterally with faint transverse sculpture, which is absent medially (Fig. 6F). Propodeal carination strong and complete; area superomedia slightly longer than wide (as long as wide in a few paratypes), pentagonal with very short, weak, parallel lateral carinae (paratypes: nearly absent lateral carinae in some specimens and less parallel in others); spiracles circular; apophyses not evident, carina somewhat enlarged in this position. Mesosoma 0.7× length of metasoma. Mesosoma (excl. legs and wings) black, including posterior corner of pronotum and tegulae (paratypes: black in most, but dark brown in a few and reddish-brown in one reared and evidently teneral specimen); humeral plates contrastingly pale yellow (Fig. 6C).

Legs. Hind femora 3.8× as long as wide. Lengths of tarsal segments in mm from first to fifth: 0.53, 0.27, 0.18, 0.10, 0.02. Claws moderately curved, simple, without teeth, basal lobe or pecten, few longer hairs present at base of claw. Arolium rather small, slightly more than half length of claw. Middle and hind tibiae with two reddish tibial spurs each, subequal in length and very slightly bent at apex. Coxae mainly black, fore and middle coxae with reddish spot on inner side (paratypes: from mostly red with dark patches to black with only small red patches on the inner side of middle coxae). Trochanters black with suffused red colouration, fore trochanters more red than black. Trochantelli, femorae and tibiae red, tibiae with darker outer surface, only weakly in fore leg. Tarsi with mixture of red and dark, with segments darker on outer surface and apically with narrow, indistinct reddish annuli.

Wings. The fore wing vein 3rs-m present, weakly pigmented, areolet symmetrical, pentagonal; vein 2m-cu rather long, straight, with one short bulla, forming acute angle with vein CU; vein RS straight; radius anterior (RA) slightly longer than pterostigma; pterostigma 2.8× as long as wide; vein 1cu-a slightly postfurcal relative to M&RS. Nervellus of hind wing intercepted at lower fifth (paratypes: from one fifth to one fourth below middle). Wing veins, including the pterostigma, uniformly dark brown, excluding narrow zone at the base of the wings with pale yellow veins; humeral plates pale yellow.

Metasoma. Punctation generally more well defined than on rest of body. First metasomal tergite with strong punctation (shiny interstices smaller than diameter of punctures), posterior part with weakly punctured area, interstices several times diameter of punctures. Dorso-lateral carinae sharp, complete, with spiracle situated just ventrad to carina (Fig. 6E). Glymmae absent. Latero-median carinae present as faint striae not extending up to level of spiracles (Fig. 6E) (paratypes: varying in length from virtually absent to extending up to level of spiracles as faint striae). Second and third terga with very strong, dense, coalescing punctures, posterior margins of terga with punctures slightly smaller and more superficial. Rest of terga somewhat densely punctulate with shiny interstices distinctly larger than punctures. Metasoma black, except for narrow, translucently brownish posterior margins of first and second terga (paratypes: first tergite from completely black to black with indistinctly delimited red patches laterally; second from completely black to black with anterior fourth red. Terga 1–3 with posterior margins from completely black to black with narrow reddish-brown translucent zones); covered with dense, short, light brown hairs. Thyridia of second metasomal tergite small, shiny, oval and red (black in some paratypes). Unsclerotised parts of metasomal sterna more or less brown, except for yellowish sclerite of first sternite. Sclerotised part of first metasomal sternite extends half way up length of sternite and up to level of spiracles on first tergite, suture between it and first tergite straight. First tergite 3.7× wider posteriorly than anteriorly, 1.6× longer than its maximum (posterior) width (paratypes: 1.1–1.7× its width (n = 32)). Second tergite 0.8× as long as wide.

Ovipositor. Dorsal valve with weak, but angulate nodus, tip beyond nodus gradually tapering, sharp (Fig. 7B). Lower valves with about five visible teeth (apical-most too small to distinguish), widely spaced, slanted, extending further towards apex ventrally than dorsally. Ovipositor sheaths covered with rather dense black hairs, about as long as diameter of sheaths. Portion of ovipositor projecting beyond metasomal apex 2.2× length of hind tibia, 0.5× length of body and 0.7× length of fore wing (paratypes: 1.9–2.4× length of hind tibia, 0.4–0.6× body and 0.6–0.9× fore wing (n = 36 for all).

Male (paratype). Length. Body length 4.9 mm, fore wing length 3.5 mm.

Head. Mandibles whitish-yellow with black teeth, reddish colouration next to mandibular teeth and mandibular base. Palpi pale reddish-yellow. Malar space 0.8× as long as basal width of mandible. Temple ratio 1.0, POL:OOL 1.3, OD:OOL 0.5. Anterior halves of scapes white, antennae otherwise black, with 21 flagellar segments, conversely to female, apical flagellomeres gradually narrower apically (Fig. 7D). Scape 1.8×, first flagellar segment 1.8× and penultimate flagellar segment 1.4× as long as wide. First flagellar segment 1.2× as long as scape; segments 8 to 12 with tyloidea, very narrow, strongly raised, more so apically, with acute apical end, almost as long as segments, except in 8th segment with medially situated tyloid about one fourth of length of segment.

Mesosoma. Generally rougher surface sculpture than in female: mesopleuron medially strongly rugose; sternaulus with longer, stronger crenulae than in female, longest crenulae as strong as and longer than crenulae on mesopleural furrow (Fig. 7C); metapleuron and propodeum rugose, no punctures discernible. Propodeal carination is strong, stronger than in female. Tegulae black, contrasting with pale yellow humeral plate. Pterostigmae of the fore wings dark brown. Coxae and hind trochanters mainly black with suffused red colouration, especially evident in fore coxae, about half red, half black; femorae, tibiae, trochantelli, fore- and mid-trochanters and tarsi, except fifth tarsal segment, red; hind tibiae and tarsi dorsally suffused with blackish (Fig. 7D). Hind femora 4.2× as long as wide; lengths of hind tarsal segments from first to last in mm: 0.59, 0.30, 0.18, 0.15, 0.14.

Metasoma. Latero-median carinae of first tergite long and rather strong, extending beyond level of spiracles. First tergite 1.5× and second tergite 0.7× as long as wide. Genitalia somewhat obscured. Parameres slightly narrowed apically and slightly downcurved, ventral margin slightly concave in lateral view, small parallel-sided gap between two parameres ventrally. Aedeagus not visible.

Etymology

L. ramii sp. nov. is my first species description. Thus, I take the opportunity to name the species in the honour of my grandfather Raimo “Rami” Konga (1936–2009) who had a profound impact on the development of my love for nature, which eventually led to the discovery of this species. The specific epithet is to be treated as a noun in the genitive case.

Distribution

Finland. Mainly in the Palearctic boreal forest biome sensu Olson et al. (2001). In addition, few specimens originate from the northernmost margin of the temperate broadleaf and mixed forest biome.

Ecology

The specimens have been collected from both open biotopes (Fig. 8A) and coniferous forests (Fig. 8B). More details are available for each specimen in the Material Examined section. Although both L. ramii sp. nov. and L. westoni occur frequently in the same habitats (and locations), it seems that L. ramii sp. nov. is more common in shaded, usually Picea abies L. dominated, forests (such as the one in Fig. 8B), while the opposite seems to be the case in the more exposed habitats. However, more data are needed for reliable comparisons.

Figure 8. 

Habitats of Lochetica ramii sp. nov. A. The type locality Mökinoja: an old log barn on the edge of a mesic meadow (the meadow is outside of the frame). B. Luvejoki, the habitat of paratype MZH GP.109739 (about 300 m from the exact spot).

Three examined specimens (MZH GP.109737, MZH GP.109736, MZH GP.109780) have been reared from Passaloecus monilicornis Dahlbom, 1842 (Crabronidae). The pins do not include the host remains and the labels do not state how the host was confirmed; however, considering the strong preference of L. westoni to parasitise Passaloecus, these rearing records are in line with expectations. The holotype was collected together with Passaloecus monilicornis and Passaloecus eremita on the same wall of a log barn (Fig. 8A).

There are not enough data to postulate whether a smaller subset of Passaloecus species is utilised as hosts (opposed to L. westoni, which seems to utilise most species) or whether additional genera of cavity-nesting aculeates are used.

Discussion

The support for the species status of L. ramii sp. nov

Several morphological characters and an ample 7% divergence in COI sequences readily separate the new species from L. westoni. However, no sequences were available for the remaining three species. Colour characters have been partly emphasised to differentiate the new species from the morphologically closest species L. japonica. Colour is known to vary geographically between distant populations on many insects, including the Ichneumonidae. Thereby, it is possible that the differences observed are not diagnostic, which could even imply the synonymy between L. ramii sp. nov. and L. japonica; however, the differences in head structure and surface sculpture of the head and mesosoma, in combination with the differences in colour adequately distinguish the new species from L. japonica. Despite several attempts, accessing the types of L. agonia and L. farta in the collection of the American Entomological Institute failed. The original descriptions are, however, sufficient for the new species to be distinguished from the two by at least two morphological characters. Furthermore, the two species occur in very different regions (western United States and Taiwan) from L. ramii sp. nov. (Finland).

The new species is described from a rather large number of specimens. However, as more specimens from a wider geographical area are revealed, the morphological and molecular gap between L. ramii sp. nov. and other species of the genus will no doubt become smaller. For example, there were no overlapping specimens of L. ramii sp nov. and L. westoni regarding their ovi-tib ratio, but the difference is so small that it is likely that overlapping specimens are likely to surface in the future.

The males of Lochetica

The males of the genus are rarely collected: three of the five species are only known from females and the number of females to males in the studied material is 38 ♀♀:1 ♂ in L. ramii sp. nov. and 40 ♀♀:5 ♂♂ in L. westoni. The sole male of L. ramii sp. nov. and three of the five males of L. westoni were reared. In addition, the reared type series of Cecidonomus westoni consisted of 9 ♀♀ and 15 ♂♂ (Bridgman 1880), which demonstrates the different sex ratio opposed to the studied museum material. On the basis of these data, it seems evident that the males are difficult to find other than via rearing. Female-biased sex ratios are not uncommon in Hymenoptera (Quicke 1997 and references therein); however, since rearing seems to produce males and females in a “normal” ratio, the most likely explanation is that the males of Lochetica are short-lived and thereby rarely collected other than via rearing. Another contributing factor could be that the males of Phygadeuontinae are usually more difficult to identify and, thus, are more likely to end up in the unsorted parts of collections than the females.

Future work

The distribution of L. ramii sp. nov. in Finland is rather widespread, extending from 60° to 64° in latitude. Thus, it is almost certain that L. ramii sp. nov. is also present in the neighbouring countries, but possibly also more widely in Europe and Asia. It is possible it has been previously confused with L. westoni, as it has been in Finland.

There are probably numerous species of Lochetica yet to be described, even in well-studied regions of the world. Future studies should attempt to examine large amounts of museum material from a wide geographic area – something the scope of this study did not allow. There is also an evident scarcity of information regarding most of the described species of Lochetica as well: the hosts and ecology is unknown in three out of five species, the males are unknown in two species and two species are only known from one or two specimens.

Identification key to the species of Lochetica of the world

The keys include all known species of Lochetica; however, no specimens of L. agonia and L. farta have been examined and their characters are based on Townes (1983). Townes (1970) and Horstmann (1978) can be used for the identification of the genus.

♀♀

1 The epomia short or absent ventrad to the pronotal sulcus. The ovipositor with a rounded or absent nodus. The legs red and the tegulae yellow Lochetica agonia Townes, 1983
The epomia long ventrad to the pronotal sulcus, extending as a weak ruga nearly to the posterior edge of the pronotum (Fig. 3A). The ovipositor with a very small but angulate nodus (Fig. 7B). The colour of the legs and tegulae similar only in L. westoni 2
2 The area petiolaris of the propodeum very densely transversely sculptured. The latero-median longitudinal carina of the propodeum, separating the area postero-externa from the area petiolaris, is weak. The coxae, trochanters and tegulae dark Lochetica farta Townes, 1983
The area petiolaris weakly to moderately transversely sculptured, the sculpture concentrated on the sides (Figs 3D, 7B). The latero-median longitudinal carina of the propodeum, separating the area postero-externa from the area petiolaris, is strong (Figs 3D, 7B). Colouration variable 3
3 The latero-median carinae of the first tergite strong, extending at least to the level of spiracles (Fig. 3E). In dorsal view, the head narrowed behind the eyes (Fig. 3B). The sternaulus with strong crenulae, stronger and longer than the crenulae on the mesopleural furrow (Fig. 3C). The ovipositor about long as the body (ovi-tib ratio more than 2.5). The legs predominantly red (coxae and tarsi sometimes with dark colouration) and the tegulae yellow (Fig. 3F) Lochetica westoni (Bridgman, 1880)
The latero-median carinae of the first tergite weak or absent (Fig. 6E). The head slightly narrowed to widened behind the eyes (dorsal view). The sternaulus with weaker crenulae, not stronger than the ones on the mesopleural furrow (Fig. 6D). The ovipositor about as long as the metasoma or shorter (ovi-tib ratio less than 2.5) (Fig. 7A). Either legs or tegulae mainly black 4
4 The mesosoma with shallower, less defined punctures, especially the area externa of the propodeum with punctures weakly, or not, discernible amongst the weak transversely orientated sculpture (Fig. 6F). The vertex uniformly sloping from the ocelli to the occiput; punctation weak and shallow, sculpture almost granulate (Fig. 6A). The pronotal sulcus without transverse crenulae. The tegulae black or dark brown (Fig. 6C). The femorae and tibiae predominantly red (Fig. 7A). The mandibles reddish with black base and teeth (Fig. 6B) Lochetica ramii sp. nov.
The mesosoma with more clearly defined, deep punctation, especially the area externa of the propodeum with dense, clearly defined punctures (Fig. 5B). The vertex with a shallow transverse depression followed by a low elevation; punctures large and rather deep (Fig. 5E). The pronotal sulcus with transverse crenulae. The tegulae light brownish-yellow (Fig. 5E). The legs, including femorae and tibiae predominantly black or dark brown with some suffused reddish colouration (Fig. 5D). The mandibles mainly black (Fig. 5A) Lochetica japonica Watanabe, 2021

♂♂

This key is very tentative. It is based on a very small sample and the males of L. japonica and L. farta are unknown. The characters in parentheses are of uncertain diagnostic value.

1 The epomia short or absent ventrad to the pronotal sulcus. Tegulae “ivory”. The mandibles mostly red. Nearctic Lochetica agonia Townes, 1983
The epomia long ventrad to the pronotal sulcus, extending near the posterior margin of the pronotum, but rather weakly indicated. Tegulae yellow or black. The mandibles mainly white, with blackish or reddish colouration proximally and distally. Palearctic 2
2 The tegulae yellow, similar in colour to the adjacent humeral plate (Fig. 3C, cf. Fig. 3F). (mesopleuron and metapleuron with stronger rugosity (Fig. 3C). Trochanters and hind tarsi generally red) Lochetica westoni (Bridgman, 1880)
3 The tegulae black, contrasting with the pale yellow humeral plate (Fig. 7C, cf. Fig. 6C). (mesopleuron and metapleuron with weaker rugosity (Fig. 7C). The trochanters and hind tarsi darkened dorsally) Lochetica ramii sp. nov.

Acknowledgements

I thank all who gave access to specimens: Juho Paukkunen (Museum of Natural History, Helsinki) loaned the bulk of the material used in this study, Kyohei Watanabe (Kanagawa Prefectural Museum of Natural History) photographed the holotype of Lochetica japonica, David Waterhouse and Dr. Tony Irwin (Norwich Museum) photographed the lectotype of Cecidonomus westoni, Rune Bygebjerg and Christoffer Fägerström (Lund University Biological Museum) photographed the lectotype of Phygadeuon pimplarius, Gergely Várkonyi gave access to the previously barcoded Finnish specimens, including the sole male specimen of L. ramii. Many specimens also originated from recent species inventories conducted as part of the Beetles-LIFE and other projects carried out by Metsähallitus (Sampsa Malmberg, Mervi Laaksonen, Eerikki Rundgren). Marko Mutanen gave much needed comments, corrections and suggestions regarding the DNA barcoding sections of this paper and Ika Österblad sent me a crucial piece of literature.

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Supplementary materials

Supplementary material 1 

R script for statistical analyses and graphs

Author: Juuso Paappanen

Data type: R

Explanation note: The script file containing all the code to replicate the statistical analyses and recreate the graphs used in the study. The file is a simple text file, which can be opened in R or in any text editor (e.g. notepad).

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (4.39 kb)
Supplementary material 2 

Aligned sequences

Author: Juuso Paappanen

Data type: fas

Explanation note: The aligned COI sequences in .fasta format used in the study.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (8.85 kb)
Supplementary material 3 

Specimen and morphological data

Author: Juuso Paappanen

Data type: xlsx

Explanation note: A spreadsheet file (MS Excel) containing the data of specimens used in the study. The data includes specimen id, species name, collecting location and date, collector and selected morphological characters.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (38.78 kb)
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