First tabulation and analysis of natural enemies of snail-killing flies (Diptera: Sciomyzidae), their position in the fly / mollusc ecosystem, and implications for use of sciomyzids in biological control Author Williams, C. D. Biological and Environmental Sciences, Liverpool John Moores University, Liverpool, UK; Author Knutson, L. Author Gordh, G. Plant Protection & Quarantine, APHIS USDA, Raleigh, NC, USA; Author Mc Donnell, R. J. Department of Crop and Soil Science, Oregon State University, Corvallis, OR, USA; Author Kula, R. Systematic Entomology Laboratory, USDA-ARS, Smithsonian Institution, Washington, DC, USA; Author Gates, M. Systematic Entomology Laboratory, USDA-ARS, Smithsonian Institution, Washington, DC, USA; Author Buffington, M. Systematic Entomology Laboratory, USDA-ARS, Smithsonian Institution, Washington, DC, USA; Author Broad, G. Department of Entomology, Natural History Museum, London, UK & Biological and Environmental Sciences, Liverpool John Moores University, Liverpool, UK; text Journal of Natural History 2025 2025-01-30 59 5 - 8 331 396 https://doi.org/10.1080/00222933.2024.2443125 journal article 10.1080/00222933.2024.2443125 1464-5262 14798887

Table 4. Summary table of Sciomyzidae (tribes Sciomyzini and Tetanocerini), including their recorded parasitoids, predators, and pathogens; distribution; and phenological and behavioural groups. Taxon authorities for genera and species of natural enemies are given in Table 5, and those for Sciomyzidae are given in Table 6.

Sciomyzidae host/			Behavioural	Phenological
prey	Natural enemies	Distribution	group	group
Sciomyzini
Atrichomelina pubera Trichopria sp.		N (NT)	1	1
Mesoleptus declivus
Mesoleptus sp. (misidentified as Atractodes)
Aphaereta pallipes
Spalangia sp.
Urolepis rufipes
Crabro sp.
Colobaea americana	Orthizema n. sp.?
Theroscopus sp.
Eupteromalus sp.
Colobaea bifasciella	‘ Cryptine’	P	3	1
‘ Chalcid’
Colobaea punctata	‘ Chalcid’	P(O)	2	1
Cephalobaris eskelundi
Ditaeniella parallela	Mesoleptus declivus
Mesoleptus n. sp. 4
Mesoleptus n. sp. 5
Phygadeuon sp.
Pherbellia albovaria	Mesoleptus declivus	N	6	5a
Pherbellia cinerella	Phygadeuontinae sp.	P(O)	7	5a,b
Scathophagidae
Pherbellia dorsata	Mesoleptus sp. C	P	2	1
Phygadeuon sp. C
Pherbellia griseola	Mesoleptus declivus	H	2	1
Mesoleptus sp. B
Eriplanus sp. J
Pherbellia n. nana	Mesoleptus declivus	H	2	1
Eupteromalus? dubius
Pherbellia obscura	Mesoleptus n. sp. 5	H
Aphaereta sp.
Pherbellia obtusa	Andrenosoma atrum	P	2	1
Pherbellia	Araneus umbraticus	P
pallidiventris
Pherbellia quadrata	Mesoleptus declivus	N	2	1?
Pherbellia s.	Mesoleptus declivus	N	4	2
maculata
Pherbellia s.	Mesoleptus sp. C	P	4	2
schoenherri
Pherbellia seticoxa	Mesoleptus sp. B	N	2	1
Eupteromalus sp.
Trichomalopsis dubia
Pteromicra	Leucosticte tephrocotis	H	2	1
angustipennis
Pteromicra pectorosa Mesoleptus declivus		H	2	1
Sciomyza simplex	Unidentified Ichneumonidae	H	2	1
Sciomyza varia	Mesoleptus declivus	N	3	5a?
Theroscopus sp.
Wingless proctotrupid
Tetanocerini
Anticheta borealis	Mesoleptus sp.	N	5	1
Phygadeuon sp. A, B
Anticheta brevipennis Unidentified Ichneumonidae		P	5	5a
Anticheta	Mesoleptus sp. nr. declivus	N	5	5a
melanosoma
Mesoleptus sp. 3
Phygadeuon sp. 1 nr. trichops
prey	Natural enemies	Distribution	group	group
Phygadeuon sp. 2
Phygadeuon sp. 7
Phaenocarpa antichaetae
Phaenocarpa impugnata
Anticheta testacea	Phygadeuon sp.	N	5	1?
Coremacera	Dioctria lateralis	P	-	-
marginata
Dichetophora	Tolmerus atricapillus	P	7	5b
obliterata
Dictya floridensis	Trichopria popei	N	11	1
Dictya pictipes	Entomophthora sp.	N	11	1
Dictya steyskali	Unidentified Ichneumonidae	N	11	1
Nematoda sp.
Dictya texensis	Ectemnius paucimaculatus	N (NT)	11	1
Elgiva cucularia	Mesoleptus sp. (misidentified as	P	11	2
	Atractodes sp.)
Hemiteles sp.?
Elgiva solicita	Trichogramma julianoi	H	11	2
Trichogramma sp.
Trichopria atrichomelinae
Trichopria popei
Theroscopus sp. G
Unidentified Ichneumonidae
Ethiolimnia lindneri	Dasyproctus bipunctatus	AF	-	-
Ilione albiseta	Mesoleptus sp. 1	P	11	4
Unidentified Hymenoptera
Gordiacea
Limnia unguicornis	Empis tessalata	P	2/8?	5b
Neolimnia tranquilla	Spilomicrus barnesi	SA	12c	6
Pherbina coryleti	Mesoleptus sp. E	P	2	4
	Phygadeuon leucostigmus
Unidentified Ichneumonidae
Tolmerus atricapillus
Protodictya chilensis	Tetragnatha sternalis	NT	11	6
Renocera pallida	Empis tessalata	P	14	1?
Renocera striata	Unidentified Ichneumonidae	H	14	1
Sepedomerus	Pheidole megacephala	NT(N)	11	6
macropus
	Anisoptera naiads
Zygoptera naiads
Mesovelia mulsanti
Sepedon aenescens	Trichogramma japonicum	O(P)	11	6(2)
	Trichogramma sp.
Trichogrammatoidea bactrae
Trichogrammatoidea nana
	Phygadeuon yonedai
	Eupteromalus sp.
Trichomalopsis oryzae
Hirsutella citriformis
Sepedon ‘ angularis ’	Trichogramma kalkae	-	-	-
	Trichogrammatoidea simmondsi
Trichogramma pinneyi
Sepedon armipes	Mesoleptus declivus	N(NT)	11	2
	Mesoleptus sp. A
	Theroscopus sp. F
Urolepis rufipes
Ectemnius paucimaculatus
Sepedon ferruginosa	Trichogramma sp.	O	11	6
	Eupteromalus sp.
Sepedon fuscipennis	Trichogramma julianoi	N	11	2
prey	Natural enemies	Distribution	group	group
Trichogramma sp.
Trichogramma sp. nr. californicum
Trichogramma sp. nr. semblidis
Trichogramma semblidis
Trichopria atrichomelinae
Trichopria popei
Mesoleptus declivus
Mesoleptus sp. A
Theroscopus sp. A, B
Unidentified Ichneumonidae
Bembix spinolae
Tropisternus sp.
Notonecta sp.
Sepedon h. hispanica Phygadeuon trichops		P	4	6
Sepedon pacifica	Phygadeuon sp.	N	11	2?6
Sepedon plumbella	Salticidae	O-OC-A	11	6
Sepedon sphegea	Trichogramma japonicum	P(O)	11	2
Trichogramma semblidis
Mesoleptus ripicola
Mesoleptus sp.
Phygadeuon cinctorius (presumable
	misidentification)
Sepedon spinipes	Unidentified Trichogrammatidae	P	11	1?2
Sepedonea telson	Unidentified Ichneumonidae	-	-	-
Sepedon tenuicornis	Theroscopus sp.	N	11?	6?
Sepedonea telson	Unidentified Ichneumonidae	NT	11	6
Sepedonea trichotypa Spilomicrus n. sp.		NT	11	6
Sepedonea annulata	Trichopria n. sp.	NT	11	6
Sepedonea trichotypa Spilomicrus sp.		NT	11	6
Tetanocera annae	Unidentified Ichneumonidae	N	11	5
Tetanocera arrogans	Mesoleptus sp. D	P	4	1
Mesoleptus sp. 1
Mesoleptus incessor
Tetanocera	Mesoleptu laticinctus	H	11	1
ferruginea
Mesoleptus sp.
Theroscopus sp. A
	Theroscopus sp. B
	Theroscopus sp. C
Phygadeuon elegans
Phygadeuon sp.
	Phygadeuon pumilis
	Theroscopus sp. A, B, C
‘ Cryptine’
Wingless proctotrupid diapriid
	Hybomitra schineri
Tetanocera	Theroscopus sp. A	H	2	1
fuscinervis
Theroscopus sp. B
	Unidentified Ichneumonidae
Tetanocera	Nematoda sp.	N	11	4
obtusifibula
Tetanocera plebeja	Trichopria popei	H	9(8)	1
	Mesoleptus (Atractodes sp. – misidentified)
	Theroscopus sp. K
Theroscopus sp.
Unidentified Ichneumonidae
Tetanocera	Mesoleptus sp. F	N	4	1
rotundicornis
Unidentified Ichneumonidae – 2 spp.
prey	Natural enemies	Distribution	group	group
Tetanocera silvatica	Empis opaca	H	2	1?
Tetanocera vicina	Bembix amoena	N	11	4
Trypetoptera	Dioctria lateralis	P	7	5b
punctulata