JHR 97: 349-361 (2024) OX JOURNAL OF = *2eereewee openasces journal

doi: 10.3897/jhr.97.124215 RESEARCH ARTICLE () I Tymenopter a
e

https://jhr.pensoft.net The international Society of Hymenopteriss REGEARCH

A new European species of Mesocrina (Hymenoptera,
Braconidae, Alysiinae, Alysiini) with notes on the
biology and systematics of the genus

H. Charles J. Godfray', Cornelis van Achterberg*

| Department of Biology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK 2 Naturalis Biodi-
versity Center, RO. 9517, 2300 RA Leiden, Netherlands

Corresponding author: H. Charles J. Godfray (charles.godfray@biology.ox.ac.uk)

Academic editor: Jovana M. Jasso-Martinez | Received 29 March 2024 | Accepted 22 April 2024 | Published 30 April 2024
Attps://zoobank. org/5893 CA68-2DB5-4D61-8FD3-3D54424D55D8

Citation: Godfray HCJ, van Achterberg C (2024) A new European species of Mesocrina (Hymenoptera, Braconidae,
Alysiinae, Alysiini) with notes on the biology and systematics of the genus. Journal of Hymenoptera Research 97:
349-361. https://doi.org/10.3897/jhr.97.124215

Abstract

Mesocrina chandleri Godfray & van Achterberg, sp. nov., is described in the small Holarctic genus Meso-
crina Foerster, 1863, the second European species. The holotype was collected in England and further
specimens are recorded from Finland, France and the Netherlands. A key is provided to the Palaearctic
Mesocrina species. DNA sequence from the CO1 barcode locus was obtained and the new species is 10%
divergent from M. indagatrix (the other European species) and 5% divergent from an undetermined
North American species. We provide evidence that Mesocrina spp. parasitise cyclorrhaphan Diptera in
fungi (and that previous host records from phytophagous insects are incorrect) and that the taxon is not

part of the Dapsilarthra genus-group as often previously assumed.

Keywords
DNA barcode, Europe, hosts, Mesocrina chandleri, Palaearctic key

Introduction

The genus Mesocrina Foerster, 1863 was erected for M. indagatrix Foerster, 1863,
which for a long time was the sole member of the genus as understood today. In the

last quarter of the 20" century four further species were described from the East-
ern Palaearctic and India (van Achterberg 1983; Belokobylskij 1998) and two from

Copyright H. Charles J. Godfray & Cornelis van Achterberg. 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.

350 H. C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

North America (Wharton 1980). Females of the genus are distinctive as they have a
laterally compressed metasoma with dorsal carina on the third and distal tergites, but
both sexes can be identified to genus relatively easily in standard keys to the Alysiini
(Fischer 1971; Tobias 1986; Wharton 1997; Belokobylskij and Tobias 1998; Zhu et
al. 2017).

Mesocrina indagatrix, hitherto the only species known from Europe (K6nigs-
mann 1959; van Achterberg 1983, 2014), is a widespread but uncommon species
with relatively few specimens in major collections. An examination of the four Meso-
crina in the collection of the National Museum of Scotland found that in addition to
three M. indagatrix, there is a single female specimen that differs from M. indagatrix
in several aspects, notably it is larger in size and has a longer ovipositor. DNA se-
quence from the CO1 barcode locus was obtained from both M. indagatrix and the
aberrant specimen and were clearly distinct (10% divergence). The BOLD database
contained a specimen from Finland with identical sequence to the new taxon which
was borrowed and found to be a male. Four further specimens were then discovered
in recent collections from the Netherlands and France. The wasps differ from the
species known from the East Palaearctic and North America and here are described
as belonging to a new species.

Methods

Details of the holotype and five paratypes are given below in the type designation.
The holotype is deposited in the National Museum of Scotland, Edinburgh, the male
paratype in the Zoological Collections of the Finnish Museum of Natural History,
University of Helsinki, Finland, and the remaining paratypes in the entomological col-
lection of the Naturalis Biodiversity Center, Leiden.

The specimen of Mesocrina indagatrix from which DNA sequence was obtained
has the data: 2, Savernake Forest, Wiltshire, England, United Kingdom (51.402N,
1.694E; UK Grid Reference SU214671); Malaise trap, 22 August — 25 Septem-
ber 1991; (collector not recorded); (Sample ID: NR1040; BOLD process ID:
BRAAL476-19).

Photographs were taken through a Leica M125C microscope with focus stacking
using the Leica Application Suite X (LAS X) image analysis software with final pro-
cessing in Photoshop. Ratio measurements were also made using the LAS X system.
Morphology terminology follows Wharton et al. (1997).

CO1 sequencing was carried out by the Biodiversity Institute of Ontario at the
University of Guelph with initial Sanger sequencing supplemented in the case of the
M. indagatrix specimen by “next generation” short read sequencing (Prosser et al.
2016). Our sequence data are publicly available at http://v4.boldsystems.org/ which
also provides full details of primers. Sequence analysis was carried out on the BOLD
platform and using the programme MEGA11 (Molecular Evolutionary Genetics Anal-
ysis version 11, Tamura et al. (2021)).

Mesocrina chandleri sp. nov. go!

Systematics

Mesocrina chandleri Godfray & van Achterberg, sp. nov.
https://zoobank.org/F38D6FA7-860D-4378-9 150-23 D8E1CF53C3
Eres iy.

Type material. Holotype, 9, south side of Haugh Wood, Herefordshire, England,
United Kingdom (52.021°N, 2.600°W; UK Grid Reference $0589360); 10 Octo-
ber 1998; swept, PJ. Chandler; National Museum of Scotland (Sample ID: NR980;
BOLD process ID: BRAAL477-19).

Paratype, 3, near Nurmijarvi, Uusimaa, Finland (60.523°N, 24.842°E; Finnish
Grid Reference 6711:381); 8 September 1994; Malaise Trap, M. Koponen; Finnish
Museum of Natural History (MZH) (Sample ID, MZH_GQ.22; BOLD process ID:
LEFIJ28469-22).

Paratype, 2, Lac de Remoray, Doubs, France (46.785°N, 6.254°E; French [Lam-
bert 93] Grid reference 948242, 6634560); 27 October 2021; Malaise Trap, H. Gens,
(RMNH’23”) Naturalis Biodiversity Center.

Paratypes 2. 9, Veluvia-Hamelakkers, Wageningen, Gelderland, The Nether-
lands (51.969°N, 5.681°E); 25-29" October 2022; caught at a skylight, D. Belgers,
RMNHW’23”; Naturalis Biodiversity Center.

Paratype, 2°, Wassenaar, Zuid-Holland, The Netherlands (52.142°N, 4.379°E;
Dutch [RDS] Grid reference 86.60, 462.12); 20 October 2023; on underside of Armil-
laria mellea (honey fungus), PH. Hoekstra, RMNH’23”; Naturalis Biodiversity Center.

Name. ‘The new species is named for Peter J. Chandler, the eminent Diptera ento-
mologist who collected the first specimen in 1998.

Description of female holotype. Holotype, 9, length of body 3.8 mm, of fore
wing 4.7 mm (Fig. 1).

Head. Antenna with 36 segments, 1.1 times length of fore wing and 1.5 times
body, densely clothed with anteriorly directed setae projecting at an angle of 30°; length
of third segment 0.7 times fourth segment, lengths of third, fourth and penultimate
segments 3.9, 2.7 and 2.4 times their widths, respectively; maxillary and labial palps
with 6 and 4 segments, respectively; length of maxillary palp ~ 2.6 times height of eye;
OOL 2.3 times greater than POL, POL 1.4 times posterior ocellus diameter; in dorsal
view head 2.0 times wider than maximum length, margin of temples posterior to eye
slightly convex; vertex shiny with scattered anteriorly-directed setae especially near oc-
ciput and margin of eye, a weak furrow runs from between posterior ocelli to occiput;
frons largely smooth and glabrous with small areas of rugosity and a small number of
setae; dorsal length of eyes 0.6 times maximum length of head, glabrous; face 1.8 times
wider than high medially, 0.6 times as wide as head, largely smooth with some slight
rugosity medially, with moderately dense setae apart from a narrow medial glabrous
band, the setae dorsally directed except at sides where they point laterally and latero-
ventrally; clypeus glabrous with sparse punctures, ventral margin very slightly convex,
epistomal suture distinct; malar space 0.15 times basal width of mandible; mandibles

352 H.C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

Figure |. Montage of photographs of the female holotype (Sample ID NR890) of M. chandleri a lateral
view b dorsal metasoma € whole insect d lateroventral head e face and mandible f dorsal head and meso-

soma. The length of the body (excluding antennae and ovipositor) is 3.8 mm.

Mesocrina chandleri sp. nov. 353

Figure 2. Montage of photographs of the male paratype (Sample ID MZH_GQ.22) of M. chandleri

a lateral metasoma b face € dorsal head and mesosoma d whole insect € wing f lateral head and meso-

soma. The length of the body (excluding antennae) is 3.3 mm.

354 H.C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

1.6 times longer than maximum width which is 1.3 times basal width, finely rugulose
and setose; three teeth with the central pointed and 0.3 times mandible length, the
others obtuse and slightly reflexed relative to the axis of the mandible, no sharp inci-
sions between teeth; head 1.3 times as wide as mesoscutum.

Mesosoma. Length of mesosoma 1.25 times its height; pronope absent; side of
pronotum glabrous and largely smooth with some crenulation antero-medially and
ventrally; mesopleuron smooth with scattered setae, mesopleural sulcus finely crenu-
late, precoxal sulcus completely absent; mesosternum with moderately dense posteri-
orly directed setae and a narrow punctate sulcus; metapleuron shallowly rugulose with
sparse setae; mesoscutum with a triangular area of dense setae and punctation on its
anterior surface extending somewhat dorso-medially, but dorsal surface largely shiny
with sparse setae, notauli punctate anteriorly but absent from dorsal surface, a medio-
posterior groove runs from 2/3 to posterior edge, margin adjoining pronotum with
longitudinal striae; scutellar sulcus 3.0 times as wide as long, with a strong medial cari-
nae and two pairs of weaker lateral carinae; scutellum smooth with a few setae; axillar
depression well developed and weakly crenulate; metanotum rugose with an indistinct
antero-medial carina that bifurcates posteriorly, not protruding in lateral view; propo-
deum rugulose anteriorly, medially and laterally, with a relatively smooth area latero-
medially, posterior region adjacent to insertion of first metasomal tergite incised and
raised with horizontal striae, lateral posterior margin with small protuberance, surface
with scattered sparse setae.

Wings. Fore wing: closed marginal, three submarginal, discal and subdiscal cells;
pterostigma sub-elliptical, about 4.5 times as long as maximally wide, r inserted at
about 0.4 times the length of the pterostigma, anterior ventral margin of pterostigma
before insertion convex, beyond insertion it narrows and merges gradually with 1-R/;
length of 7 0.6 times the width of pterostigma, approximately orthogonal to anterior
wing margin; ratio of 7; 3-SR: SRI = 1: 5.2: 9.3, SRI slightly sinuate ending near
wing apex; second submarginal cell narrows distally, 5-sided (i.e. m-cu postfurcal),
ratio 2-SR: 3-SR: r-m: 2-M = 1: 1.45: 0.46: 2.16, angle subtended by 2-SR and 3-SR
125°, 2-SR slightly sinuate; /-CUI very short; 3-CUJa 1.3 times longer than CU10;
CU1a concave down and extends nearly to wing margin. Hind wing: closed basal cell;
cu-a and 2-M present.

Legs. Hind coxa smooth; hind femur ~ 5 times longer than maximally wide,
densely clothed with short setae dorsally projecting at an angle of 30°, ventrally ap-
proximately half the width of the femur and projecting at 60°; hind tibia slender,
densely clothed with setae (projecting at 30°), 1.15 times longer than tarsi; apical tibial
spur inconspicuous, less than 0.2 times length of basitarsus; tarsi with similar setae to
tibia, tarsal segment length ratios 1 (basitarsus): 0.53: 0.42: 0.31: 0.38; tarsal claws and
arolium well developed and 0.75 times length of fifth tarsal segment; structure of fore
and mid legs similar though femur more slender and legs shorter, ratio of hind: mid:
fore femur 1: 0.90: 0.81.

Metasoma. Length of first tergite 1.7 times its apical width, the latter 1.7 times
its narrowest width near its base, a pair of dorsal carinae arise basally from the lat-

Mesocrina chandleri sp. nov. 355

eral carinae and reach the dorsum at about 4 and run close together in parallel to
about 2 where they lose their distinctiveness, posterior dorsal surface with longitu-
dinal sculpturing and sparsely scattered setae, a distinct dorsope present; metasoma
beyond first tergite strongly lateral compressed with a strong dorsal medial carina
extending from the third tergite to the end; second tergite smooth; scattered setae
on second and posterior tergites; ovipositor straight and projecting, its exposed
setose part 0.7 times the length of the hind tibia; ovipositor sheath with posterior
directed setae projecting at an angle of 60—80°, their length up to twice the width
of sheath; hypopygium slightly postero-ventrally produced, terminating at level of
the cerci.

Colour. Head, mesosoma and metasoma dark brown except for yellow brown
parts: scape, pedicel, base of the third antennal segment, mandibles (apart from tooth
tips) and ventral part of gena, latero-ventral prothorax, tegulum, medio-ventral region
of the laterally-compressed metasoma; precoxal area of mesopleuron slightly lighter
than the remainder of the surface; palps and legs yellow, the mid and hind tarsi slightly
darker, areola contrastingly dark brown; wing venation and pterostigma dark brown,
wing membrane hyaline.

Variation amongst females. The French and Netherland female specimens gen-
erally match the holotype. The number of antennal segments were 34 and 37 (two
specimens without complete antenna); extent of setation on dorsal surface of mesoscu-
tum varies from comparatively well developed as shown in holotype (Fig. 1) to largely
restricted to a few remote setae as figured in the male paratype (Fig. 2). In addition,
the medio-longitudinal carina of propodeum is weakly developed in some specimens.

Description of male. Paratype, 3, length of body 3.3 mm, of fore wing 4.1 mm
(Fig. 2).

This specimen is somewhat damaged with both antennae truncated and some legs
missing. Similar to the female but differing in the following features.

Head. Posterior ocelli slightly closer together (OOL 3.4 times POL); face some-
what less setose.

Wings. Veins thicker than in female; second submarginal cell slightly shorter —
ratio of 7; 3-SR: SRI = 1: 4.3: 8.3, ratio 2-SR: 3-SR: r-m: 2-M = 1: 1.37: 0.38: 2.06.

Legs. Hind leg first tarsal segment slightly longer — tarsal segment length ratios = 1
(basitarsus): 0.50: 0.36: 0.26: 0.34.

Mesosoma. Anterior surface of mesonotum setose but setae extending less onto the
dorsal surface than in the female holotype; propodeum smoother.

Metasoma. The metasoma is not laterally compressed and does not have longitu-
dinal carinae on the posterior tergites; thus, having a “normal” Alysiini appearance.

Colour. Ventral margin of clypeus yellow; metasoma more uniformly brown.

Molecular analysis. Sequence data from the mitochondrial CO1 gene (the stand-
ard barcode locus) were obtained from the M. chandleri holotype (607 base pairs)
and a British specimen of M. indagatrix (550 b.p.). A male Finnish specimen with
identical gene sequence to the holotype was found in the BOLD database. No further
Old World sequences were present in BOLD, but it did contain seven closely-related

356 H.C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

0.023

Mesocrina 0.021 ———————_| A

0.021

0.028

sp. Nov.

indagatrix

C
0.033

0,015 40
0.022
Figure 3. Maximum-likelihood tree (CO1 gene, Tamura-Nei model) with branch lengths of the 10
Mesocrina sequences in the BOLD database. The height of the terminal triangular wedges represents
sample number and their horizontal width the genetic variation within the species (where no variation the
wedge is a vertical line). Letters represent putative and undetermined Canadian (A, B, D) or Californian

(C) species of Mesocrina.

North American sequences, some identified as Mesocrina and others that appeared
from images in BOLD to belong to the genus. A maximum likelihood phylogenetic
tree (CO1 gene, Tamura-Nei model; default MEGA settings) of the 10 sequences was
created (Fig. 3), and they segregated into six putative species or BINs (Barcode Index
Numbers; BIN codes as of February 2024).

Mesocrina chandleri (BIN: BOLD:ADX0117) shows a 6.5% divergence from its
nearest relative, a Californian BIN (BOLD: AFO2946) represented by a single speci-
men. The two together with a Canadian BIN (3 specimens, BOLD:ACD3275) form
a clade separated by 6.1% from a clade containing M. indagatrix (BOLD:AED2809)
and two Canadian BINs (BOLD:AAU8494 & ACL6319). M. indagatrix shows a
11.6% separation from M. chandleri and 6.0% from the nearest Canadian BIN. The
genetic data thus strongly supports the distinctiveness of the new species.

Discussion

The Palaearctic species of Mesocrina Foerster can be distinguished by the following key
which is based on Belokobylskij’s (1998) key (in Russian).

1 Precoxal sulcus distinctly crenulate; posterior surface of propodeum sloping
vertically; first metasomal tergite about as long as its apical width; mesoscu-
tum, pronotum and mesosternum brownish yellow; propleuron brownish;
[vein r of fore wing much longer than wide; vein 1-SR short]; India, N.
China (Jilin, Fujian, Hainan, Yunnan)....... M. dalhousiensis Sharma, 1978

7 Precoxal sulcus smooth or absent; propodeum gradually sloping posteriorly;
first metasomal tergite 1.2—2.0 times as long as its apical width; at most the
pronotum ventrally, and the propleuron, brownish yellow ........... ee eeeeeeee 2

Mesocrina chandleri sp. nov. D7

2 Setose part of ovipositor sheath about as long as first tergite, 0.3—0.5 times
as long as hind tibia (and shorter than apical height of metasoma and 0.13-—
0.14 times as long as fore wing); anterior half of middle lobe of mesoscutum
largely glabrous and without distinct punctures; Palaearctic & NE Oriental;
TL OAgA se taeisccea tener cheek vo Secam cel chcaat eaag aa cobmnchi ts M. indagatrix Foerster, 1863
- Setose part of ovipositor sheath 1.5—2.0 times as long as first tergite, 0.7—0.9
times as long as hind tibia (and about equal to apical height of metasoma and
about 0.2 times as long as fore wing); anterior half of middle lobe of mesos-
cutum: inore:or less: setose:and-with’some Pune tutes: ....6..22<¢.-e-2eseeveesPasnece! 3
3 Third antennal segment about 1.2 times longer than fourth segment; third
tooth of mandible much smaller than first tooth; pterostigma robust and
strongly sclerotised, about 5.0 times wider than length of vein r; [clypeus
blackish}; Kast Palacatctio®®: i290. tence. M. lesovik Belokobylskij, 1998
= Third antennal segment 1.5—1.7 times longer than fourth segment; third
tooth of mandible similar to first tooth or larger; pterostigma more slender
and less sclerotised, about twice times wider than length of vein r............... 4,
2 Mandible less robust, with first tooth of mandible not protruding and similar
to third tooth; vein 2-SR of fore wing distinctly sinuate; antenna with 31-32
segments; propodeum without sculpture medially; East Palaearctic...............
eee UM Reece, Oe see ie eee M. licho Belokobylskij, 1998
= Mandible more robust, with first tooth of mandible rather protruding and
sometimes larger than third tooth; vein 2-SR of fore wing straight or slightly
sinuate; antenna with 34—40 segments; propodeum more rugulose or with
(eebabavteantcra ble) lk gor eat he Seer Reinet te ae pars i ie ve eh eA 5
5 Vein r of fore wing 0.8 times maximum width of pterostigma and narrow;
first metasomal tergite 1.1 times as long as apically wide, twice as wide pos-
teriorly as minimum width; propodeum with distinctly developed medio-
longitudinal carina; clypeus largely yellow; second submarginal cell of fore
wing robust and vein SR1 about 2.2 times as long as vein 3-SR; [middle lobe
of mesoscutum only anteriorly setose (confined to anterior edge)]; East Palae-
AT CLIC saa, seed cclonmat «caret g Rbrenduaesindd eeBubeccanoe din taster M. leshii Belokobylskij, 1998
— Vein r of fore wing thickened and 0.3—0.6 times maximum width of pterostig-
ma; first metasomal tergite 1.7 times as long as apically wide, 1.7 times as wide
posteriorly as minimum width; propodeum at most with weakly developed
medio-longitudinal carina; clypeus dark brown or yellowish near ventral rim;
second submarginal cell of fore wing less robust and vein SR1 1.7—2.0 times as
long as vein 3-SR; [linear medio-posterior depression deep and comparatively
long; females with 34—37 antennal segments; setose area of middle lobe of mes-
oscutum variable]; West Palaearctic; Figs 1, 2.0.0.0... M. chandleri sp. nov.

In Europe only MZ. indagatrix (Fig. 4) and the new species are currently known.
Females of M. chandleri are easily distinguished by their longer ovipositor. Males are
harder to tell apart, but M. indagatrix is somewhat smaller and the anterior of the mes-
oscutum middle-lobe is smoother and bears fewer setae.

358 H.C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

Figure 4. Montage of drawings (by C.v.A. using a camera lucida) of male and female Mesocrina indagatrix.
Scale bar: 1.0 mm.

Mesocrina chandleri sp. nov. 359

Females of both European species of Mesocrina and an American species (Wharton
1980) have been captured on gilled fungi, and it has been hypothesised that the later-
ally flattened female metasoma allows it to be inserted between closely packed fungal
gills (Wharton 1980). We are unaware of any substantiated host rearing records but as
all Alysiinae attack Diptera we presume Mesocrina spp. parasitise fly larvae feeding in
fungal fruiting bodies.

K6nigsmann (1959) quotes two non-fungi-associated host records for M. indaga-
trix which he referred to as Pseudomesocrina venatrix Marshall, later synonymised by
van Achterberg (1983). The first a male from a Pegomya sp. (Anthomyiidae) leaf mining
Rumex sp. in the UK (Morley 1933). The first author and his students have reared 1880
parasitoid specimens from Pegomya spp. feeding on Rumex in the UK of which the only
Alysiini were Adelurola florimela Haliday which made up 35% of the rearings. As males
lack the distinctive compressed metasoma, and as Morley says the male only differs
from the female in antennal segment number (which would be a curious observation
to make about a highly sexually dimorphic Mesocrina), we suspect Morley’s record
is a misidentification, possibly of A. florimela. The second is Nanna (=Amaurosoma)
armillata and/or Nanna flavipes (Scathophagidae) in developing flower heads of grasses,
Phleum spp. (Poaceae), in Sweden (Borg 1959). We suspect this is a misidentification
of Synelix semirugosa Haliday (Braconidae, Alysiinae) which attacks these hosts (King
et al. 1935; Telenga 1935) and also has a laterally compressed metasoma. Our working
hypothesis thus remains that Mesocrina spp. are parasitoids of Diptera in fungi.

The UK, Netherland and French specimens of the new species were all captured in
October while the more northerly record from Finland is from September. ‘The three
specimens of M. indagatrix in the NMS collection were also collected in the autumn.
The abundance of fungal fruiting bodies peaks at this time of year, and so the phenol-
ogy and putative biology are consistent. Mesocrina chandleri is a relatively large and
distinctive species (for an Alysiini) and given how widespread it is we are surprised
it has not been noted before, especially as some of the recent records come from sites
that have been regularly surveyed for Braconidae. We speculate that the species may be
undergoing a current expansion of range and/or abundance.

There are currently eight described species of Mesocrina, six from the Palaearc-
tic and two from the Nearctic (Wharton 1980; van Achterberg 1983; Belokobylskij
1998). However, DNA COI sequences of the two European species cluster closely
with four North American taxa that we believe are Mesocrina (that may or may not
include the two described by Wharton) suggesting greater diversity in the Nearctic
than currently recognised.

The genus Mesocrina was described by Foerster (1863 [1862]) though van Achter-
berg (1983) treated it as a subgenus of Dapsilarthra. Later Papp (1991 [1990]) raised
it to genus level which was followed by other authors (van Achterberg 2014). Other
Dapsilarthra genus-group braconids attack phytophagous cyclorrhaphan Diptera and
they are thought to be the sister group of the Dacnusini which have the same biol-
ogy (Griffiths 1964). Van Achterberg (1983) noted that the association of Mesocrina
spp. with fungi was atypical for the genus group, and Wharton (1980) suggested an

360 H. C.J. Godfray & C. van Achterberg / Journal of Hymenoptera Research 97: 349-361 (2024)

affinity with Alysia which is supported by our (relatively limited) DNA sequence data.
Stronger evidence comes from a recent phylogenomic study of the Braconidae that
included single species of Alysia, Mesocrina and Dapsilarthra: the first two genera are
closely associated, while Dapsilarthra is distantly related and is indeed placed near the
root of the Dacnusini (Jasso-Martinez et al. 2022).

Acknowledgements

We are grateful to Sergey Belokobylskij for supplying photographs of the holotype of
Mesocrina leshii, to Mark Shaw for access to the National Museum of Scotland’s col-
lection, to Marko Mutanen and Juho Paukkunen for loan of the male paratype, and to
Paul Hebert & Jayme Sones (University of Guelph) for DNA sequencing and informa-
tion from the BOLD database.

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