.jpg){kind=logo}
Platyhelminthes
The phylum Platyhelminthes includes two classes of worms that involve ants: the cestodes (tapeworms) and the trematodes (flukes and blood flukes). These two groups have complex life-cycles, with mature stages that live as parasites in the digestive systems of fish or land vertebrates, and intermediate stages that infest secondary hosts, including ants.
Cestodes
All cestodes are parasites of vertebrates; many have complex life histories, including a stage in a definitive (main, vertebrate) host in which the adults grow and reproduce, often for years, and one or two intermediate stages in which the larvae develop in other hosts (typically arthropods or other vertebrates). The adults live in the digestive tracts of vertebrates, while the larvae often live in the bodies of other animals.
Commenting on cestode infections in Manica bradleyi, Prebus (2023) were uncertain of the details of parasite transmission, but speculated that workers collect bird feces containing cestode eggs while foraging and feed these to its larvae via trophallaxis, which has been observed in other ants (Horsfall, 1938). Development of infected individuals is likely disrupted by nutrient deficiency, potentially causing the morphological modifications observed in infected workers.
Known Ant Hosts
| Ant | Cestode Species | Locality | Source | Notes |
|---|---|---|---|---|
| Brachyponera chinensis | Raillietina kashiwarensis | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Brachyponera sennaarensis | Raillietina tetragona | Sudan | Mohammed et al., 1988; Laciny, 2021 | |
| Carebara | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest; as ''Pheidologeton'' sp. | |
| Crematogaster lineolata | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Forelius pruinosus | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Formica rufa | Raillietina friedbergeri | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Harpagoxenus sublaevis | Type 2 (Choanotaenia unicoronata?) | South Tyrol, Italy | Buschinger, 1973; Laciny, 2021 | |
| Lasius niger | Mesocestoides sp. | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Leptothorax | Choanotaenia crateriformis | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Leptothorax acervorum | Choanotaenia unicoronata | South Tyrol, Italy; Bavaria, Germany | Buschinger, 1973; Peru et al., 1990; Laciny, 2021 | |
| Manica bradleyi | Raillietina sp. | California, United States | Prebus et al., 2023 | |
| Monomorium | Cotugnia digonopora | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium floricola | Cotugnia digonopora | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium floricola | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium indicum | Cotugnia digonopora | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium indicum | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Monomorium indicum | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Myrmica rubra | Davaineidae (Raillietina urogalli??) | Scotland | Muir, 1954; Laciny, 2021 | |
| Myrmica rubra | Raillietina urogalli | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Myrmica scabrinodis | Davaineidae (Raillietina urogalli??) | Scotland | Muir, 1954; Laciny, 2021 | |
| Myrmica scabrinodis | Raillietina urogalli | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina australis | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina beveridgei | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina chiltoni | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina circumvalata | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina dromaius | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina mehrai | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina mitchelli | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina singhi | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole bergi | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole bicarinata | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole bicarinata | Raillietina georgiensis | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole bicarinata | Raillietina georgiensis | Georgia, United States | Muir, 1954; Laciny, 2021 | as ''Pheidole vinelandica'' |
| Pheidole bicarinata | Raillietina loeweni | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole bicarinata | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole dentata | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole fervida | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole jucunda fossulata | Cotugnia digonopora | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole jucunda fossulata | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole jucunda fossulata | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole nodus | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Pheidole soritis | Raillietina loeweni | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest; as Pheidole sitarches campestris | |
| Prenolepis | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Prenolepis | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tapinoma sessile | Mesocestoides sp. | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Temnothorax affinis | Anomotaenia brevis | Germany | Scharf et al., 2012; Laciny, 2021 | |
| Temnothorax affinis | Dilepididae (type 1(?)) | Germany | Buschinger, 1973; Laciny, 2021 | |
| Temnothorax aveli | Dilepididae (type 1(?)) | Germany | Buschinger, 1973; Laciny, 2021 | |
| Temnothorax lichtensteini | Choanotaenia crateriformis | Spain | Espadaler & Riasol, 1983; Laciny, 2021 | |
| Temnothorax nylanderi | Anomotaenia brevis | France | Gabrion et al., 1976; Laciny, 2021 | |
| Temnothorax nylanderi | Anomotaenia brevis | France (lab colony) | Trabalon et al., 2000; Laciny, 2021 | |
| Temnothorax nylanderi | Anomotaenia brevis | Germany | Scharf et al., 2012; Beros et al., 2015; Feldmeyer et al., 2016; Laciny, 2021 | |
| Temnothorax nylanderi | Dilepididae (type 1(?)) | Germany | Buschinger, 1973; Laciny, 2021 | |
| Temnothorax parvulus | Dilepididae (type 1(?)) | Germany | Buschinger, 1973; Laciny, 2021 | |
| Temnothorax rugatulus | Dilepididae unspecified (cf. Anomotaenia) | Arizona/New Mexico, United States | Heinze et al., 1998; Laciny, 2021 | |
| Tetramorium caespitum | Raillietina caucasica | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium caespitum | Raillietina circumvalata | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium caespitum | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium caespitum | Raillietina fedjuschini | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium caespitum | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium simillimum | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium simillimum | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium tortuosum | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium tortuosum | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium tsushimae | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest; as Tetramorium caespitum jacoti | |
| Tetramorium tsushimae | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest; as Tetramorium caespitum jacoti | |
| Tetramorium wroughtonii | Raillietina echinobothrida | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Tetramorium wroughtonii | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Trichomyrmex destructor | Cotugnia digonopora | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Trichomyrmex scabriceps | Raillietina tetragona | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest |
Trematodes
Trematodes are primarily internal parasites of molluscs and vertebrates. Most trematodes have a complex life cycle with at least two hosts. The primary host, where the flukes sexually reproduce, is a vertebrate. The intermediate host, in which asexual reproduction occurs, is usually a snail, but includes several species of ants as well.
The best studied example is Dicrocoelium dendriticum, the lancet liver fluke. Adults of Dicrocoelium dendriticum are found in the bile ducts of sheep, cattle, pigs, deer, cottontail rabbits, and woodchucks, while earlier life stages inhabit snails and ants. Fluke eggs are eliminated in the feces of their vertebrate hosts and are ingested by terrestrial snails. Cercaria (the free-swimming larval stage) develop in the snails, are secreted in the snail’s mucus or slime ball, and are ingested by ants. Once within the ant the parasites develop in two areas of the ant’s body, the brain and the abdomen. Those in the brain cause the ant to disperse to the tips of grass blades or other vegetation at dusk, lock their mandibles to the vegetation, and remain there until dawn (this behaviour is called summiting behavior). In this position they are most likely to be eaten by herbivores (the fluke's primary host) in the early morning. This behaviour will be repeated every night until the ant is eaten by a vertebrate host in which the parasite can fully mature and complete its life cycle.
Formica aserva worker infected with the lancet liver fluke, Dicrocoelium dendriticum. (A) False-coloured 3D volume rendering in sagittal section. (B) Micro-CT-based virtual sagittal section of an ant gaster harbouring 6 encysted metacercariae. (C) Micro-CT-based virtual sagittal section of an ant gaster harbouring 98 encysted metacercariae. (D) False-coloured 3D volume rendering of an infected ant gaster in sagittal section, showing scar-like marks (arrow) on the crop surface. Abbreviations: br, brain; cr, crop; emc, encysted metacercaria; nmc, non-encysted metacercaria. From Martin-Vega et al., 2018, Fig. 2.
Diagrammatic representation of life cycle stages of Dicrocoelium dendriticum. A Final host: chiefly sheep and cattle; humans are incidental (“accidental hosts”). 1 Sexually mature liver fluke (see under III). 1 a Egg with a fully formed miracidium. B Intermediate host: land snails (Helicella, Zebrina). 2 Miracidium which has emerged from a snail. 3 a The youngest sprorocyst. b The oldest sporocyst stage (mother sporocyst). c Daughter sporocyst. d A single cercaria (the so-called Cercaria vitrina). e Shell of Helicella ericetorum. f Shell of Zebrina dendrita. C 2. Intermediate hosts: ants. 4 a Slime balls laid by the snail on grass stem. 4 b An isolated (single) slime ball laid by the snail. 5 a An ant eating slime balls. 5 b Mature metacercariae from ants. (I) A miracidium emerging. (II) “Cercaria vitrina”. (III) A sexually mature liver fluke. From Piekarski, 1963.
Known Ant Hosts
| Ant | Cestode Species | Locality | Source | Notes |
|---|---|---|---|---|
| Camponotus compressiscapus | Dicrocoelium hospes | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Camponotus herculeanus | Brachylecithum mosquensis | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Camponotus pennsylvanicus | Brachylecithum mosquensis | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Cataglyphis aenescens | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Cataglyphis aenescens | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | |
| Cataglyphis albicans | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | as ''Cataglyphis albicans armenia'' |
| Formica aserva | Dicrocoelium dendriticum | Canada (Alberta) | van Paridon et al., 2017; Martin-Vega et al., 2018; Criscione et al., 2020 | |
| Formica cunicularia | Dicrocoelium dendriticum | Germany | Hohorst & Graefe, 1961 | |
| Formica cunicularia | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | |
| Formica gagates | Dicrocoelium chinensis | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Formica gagates | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Formica lugubris | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Formica mesasiatica | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Formica picea | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Formica polyctena | Dicrocoelium dendriticum | Denmark | Botnevik et al., 2016; de Bekker et al., 2018 | |
| Formica pratensis | Dicrocoelium dendriticum | de Bekker et al., 2018 | ||
| Formica pratensis | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode secondary; indirect transmission; transmission outside nest | |
| Formica rufa | Dicrocoelium dendriticum | de Bekker et al., 2018 | ||
| Formica rufa | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | Arakelian et al. (1997) note this infection is questionable |
| Formica rufibarbis | Dicrocoelium dendriticum | de Bekker et al., 2018 | ||
| Formica rufibarbis | Dicrocoelium dendriticum | Germany | Hohorst & Graefe, 1961 | |
| Formica rufibarbis | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | |
| Formica transkaucasica | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | |
| Lasius | Dicrocoelium dendriticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest | |
| Proformica pilosiscapa | Dicrocoelium lanceatum | Armenia | Arakelian et al., 1997 | |
| Technomyrmex albipes | Eurytrema pancreaticum | Quevillon, 2018 | encounter mode primary; indirect transmission; transmission outside nest |
References
- Horsfall, M.W. 1938. Observations on the life history of Raillietina echinobothrida and of R. tetragona (Cestoda). J. Parasitol. Res. 24, 409–421 (doi:10.2307/3272117).
- Prebus, M., Georgiev, B.B., van de Kamp, T., Hamann, E., Baker, I., Rabeling, C. 2023. The rediscovery of the putative ant social parasite Manica parasitica syn. nov. (Hymenoptera: Formicidae) reveals an unexpected endoparasite syndrome. Biology Letters, 19, 20230399 (doi:10.1098/rsbl.2023.0399).
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||