Review Article

Spider-Ant Associations: An Updated Review of Myrmecomorphy, Myrmecophily, and Myrmecophagy in Spiders

Table 2

Spider myrmecophages. *Categories (defined in text) include R: Reluctant myrmecophage; I: Indifferent acceptor; F: Facultative ant predator; O: Obligatory ant predator; Unk: cannot be determined from information about their biology presented in the literature (these are most likely R or I myrmecophages). Spider taxonomy according to Platnick [63]; ant taxonomy according to http://antbase.org/.

Spider myrmecophageCategory of myrmecophage*Notes on biologyReferences

Araneidae
Metepeira gosoga Chamberlin and IvieUnkAuthor suggests that spiders may feed on ants found only on cholla where spider is also found.[93]
Metepeira sp.UnkReported feeding on Crematogaster opuntiae Buren.[93]

Deinopidae
Deinopis sp.Probably IThrows web over ants passing below.[94]

Eresidae
Seothyra sp.FLives in silk lined burrows. Mouth of burrow covered by prey capture web. Captures mostly ants. Male spider runs on ground during day and is myrmecomorph and behavioral mimic of Camponotus sp. and mutillid wasps (dimorphic mimicry).[95]

Gnaphosidae
Callilepis nocturna (L.)May be FFeeds on Formica spp. and Lasius spp. Actively searches for ants and may enter nests to hunt workers. Approaches ant and bites on base of antenna. Antennae seem to act as stimulus to trigger attack.[9698]

Linyphiidae
Frontinella communis (Hentz)IOccasionally preys on ants.[99]

Oecobiidae
Oecobius annulipes LucasOMain food is Plagiolepis pygmaea (Latreille) but other ants (e.g., Lasius flavus (Fabricius)) accepted in lab. Bites at base of antenna. Swaths ant in silk and encircles it. Sometimes uses last pair of legs as well as spinnerets to direct silk over prey. Reduced chelicerae and enlarged gnathocoxae may be adaptations to myrmecophagic lifestyle.[100]
O. cellariorum (Dugès)OFeeds on Plagiolepis pygmaea (Latreille). Bites at base of antenna.[100]
O. templi O. P.-CambridgeO[100]

Oonopidae
Triaeris stenaspis Simon (publ. as T. patellaris)UnkReported attacking Cyphomyrmex costatus Mann.[101]

Oxyopidae
Oxyopes apollo BradyUnkEats ants.[102]
O. globifer SimonI/FAnts constitute large % of prey.[99, 102]
O. licenti SchenkelUnkEats ants.[102]
O. salticus HentzUnkEats ants.[102]
O. scalaris HentzIOccasionally eats ants.[99, 102]
O. sertatus L. KochUnkEats ants.[102]
Peucetia viridans (Hentz)UnkEats ants.[103]

Pholcidae
Crossopriza lyoni (Blackwall) (publ. as Crossopriza stridulans)UnkFeeds on fire ants, Solenopsis invicta Buren.[104]

Salticidae
Aelurillus aeruginosus (Simon), A. cognatus (O. P.-Cambridge), and A. kochi RoewerFPrefer ants over other prey. Innately recognize ants even if ants are not moving. Attack from front unless ant is passing (then switch to rear attack). Use different hunting behavior for ants than for other prey. If hungry, show no preference for ants over other prey.[105]
Aelurillus m-nigrum KulczyńskiFPrefers ants over other prey; 85% of diet in field consists of ants. Uses different hunting behaviors for ants than for other prey: lunges, attacks from front, bites, releases, bites again.[92]
Aelurillus spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than for other prey.[106]
Anasaitis canosa (Walckenaer) (publ. as Corythalia canosa or as Stoidis aurata)FPrefers ants over other prey. Uses different hunting behaviors for ants than for other prey: attacks from front, holds forelegs away from struggling ant. Also stilts body off ground.[107, 108]
Anasaitis spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than for other prey.[106]
Chalcotropis spp.FUse different hunting behaviors for ants than for other prey: some attack from rear, some head-on, then lunge, bite, release, and wait.[106, 109]
Chrysilla lauta ThorellFPrefers ants. Uses different hunting behaviors for ants than for other prey: attacks from rear, bites gaster (not appendages), retreats and waits, may lunge and strike several times. When ant quiescent, spider approaches, bites again, and carries it away.[110]
Chrysilla spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than for other prey.[106]
Cosmophasis sp.UnkFeeds on ants and is myrmecomorph.[59]
Euophyrs spp.FUse different hunting behaviors for ants than for other prey: some attack from rear, some attack head-on, then lunge, bite, release, and wait.[106]
Evarcha albaria (L. Koch)I/FRobs ants of their prey and of their brood (eggs and larvae) that workers carry (kleptoparasites).[111]
Habrocestum pulex (Hentz)Some F
Some I
Some individuals prefer ants over other prey; some prefer other prey over ants. Myrmecophagic individuals use different behaviors for ants than for other prey: lunge or leap onto petiole or thorax, bite, release, repeat (up to 6 times). Keep front legs off ground away from ant. Reported preying on Crematogaster spp.[112114]
Habrocestum spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than other prey.[106]
Hasarius adansoni (Audouin)Probably IWill feed on ants.[115]
Hentzia palmarum (Hentz) (publ. as Eris marginata)UnkReported feeding on workers of Myrmica sp.[113]
Icius sp.UnkReported feeding on small brown ants.[113]
Menemerus fulvus (L. Koch) (publ. as Menemerus confuses)I/FRobs ants of their prey and of their brood (eggs and larvae) that workers carry (kleptoparasites).[111]
Myrmarachne foenisex SimonFRegularly feeds on weaver ant (Oecophylla) larvae. Also mimics weaver ants.[59]
Natta horizontalis Karsch (publ. as Cyllobelus rufopictus)FPrefer ants. Uses different hunting behaviors for ants than for other prey: attacks from rear, bites gaster (not appendages), retreats, and waits, may lunge and strike several times. When ant quiescent, spider approaches, bites again, and carries it away.[110]
Natta spp.FSpecies in genus generally prefer ants. Use different hunting behaviors for ants than for other prey: attack from rear, bite gaster (not appendages), retreat and wait, may lunge and strike several times. When ant quiescent, spider approaches, bites again, and carries it away.[106, 110]
Phidippus johnsoni (Peckham and Peckham)IOccasionally eats ants.[99, 116]
Plexippus setipes KarschI/FRobs ants of their prey and of their brood (eggs and larvae) that workers carry (kleptoparasites).[111]
Siler cupreus Simon (publ. as Silerella vittata)F/OEats ants. Spider population increases in areas infested with Argentine ants, Linepithema humile (Mayr). Also robs worker ants of brood including eggs, larvae, and pupae being carried by workers (kleptoparasitism).[117120]
Siler semiglaucus (Simon)FPrefer ants. Uses different hunting behaviors for ants than for other prey; bites gaster (not appendages), retreats and waits, may lunge and strike several times. When ant quiescent, spider approaches, bites again, and carries it away.[110]
Siler spp.FUse different hunting behaviors for ants than for other prey: some attack from rear, some from head-on, lunge, bite, release and wait.[106, 109]
Tutelina formicaria (Emerton)FAlso myrmecomorph. Preys on red and black ants.[121]
Tutelina similis (Banks)FPreys primarily on ants and is also a myrmecomorph. Uses different hunting behaviors for ants than for other prey: bites quickly, releases, retreats, carries paralyzed prey to safe area.[99, 113]
Tutelina spp.FOther species of Tutelina found on mound of Pogonomyrmex salinus Olsen (publ. as P. owyheei) feeding on worker ants.[113]
Xenocytaea spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than other prey.[106]
Zenodorus durvillei (Walckenaer), Z. metallescens (L. Koch), and Z. orbiculatus (Keyserling)FPrefer ants over other prey. Feed on ants caught in other spider’s webs—but only if spiders can approach safely without getting caught. Ambush ants; hang upside down and lunge at ant while releasing dragline. Repeatedly bite larger ants. Do not hold onto injured ant.[106, 108]
Zenodorus spp.FSpecies in genus prefer ants over other prey. Use different hunting behaviors for ants than other prey.[106]

Scytodidae
Scytodes sp.UnkFeeds on fire ants, Solenopsis invicta Buren.[104]

Theridiidae
Achaearanea spp.UnkFeed on “carpenter ants.” Ants become entangled in gum footed sticky thread attached to substrate. Movement of ant causes thread to snap and ant is lifted off ground.[93]
Argyrodes sp.UnkReported feeding on Pogonomyrmex rugosus Emery.[93]
Asagena fulva (Keyserling) (publ. as Steatoda fulva) and A. pulcher (Keyserling) (publ. as S. pulcher)UnkFeed on Pogonomyrmex badius (Latreille) and P. subnitidus Emery. When ant workers captured in webs, major workers (patrollers) may attempt to free them but become caught in webs themselves.[93, 122]
Cryptachaea riparia (Blackwall) (publ. as Theridion saxatile and as Acaeoraneae riparia)FCaptures ants with above-ground web that has sticky threads attached to substrate. Webs built in areas of high ant activity or traffic. Greater than 88% of diet made up of ants (mostly Formica spp.). Ant gets tangled in sticky silk, struggling causes line to snap, ant is suspended, spider responds to vibrations, bites ant several times in legs and antennae while wrapping in silk, cuts paralyzed ant, and carries it to sand-covered tube retreat.[123, 124]
Dipoena punctisparsa YaginumaUnkFeeds on small ants in genus Lasius.[125]
Enoplognatha ovata (Clerck) (publ. as Theridion lineatum or T. lineamentum)UnkFeeds on Pogonomyrmex barbatus (Smith). Builds webs in grass near colony. Ants crawling up into grass or passing below get entangled.[126]
Euryopis californica BanksI/FReported feeding on Pogonomyrmex rugosus Emery.[93]
Euryopis coki LeviI/FPreys on Pogonomyrmex salinus Olsen (publ. as P. owyheei). Spider captures ant on the mound by trapping ant against ground with sticky silk. Bites on leg. Ant swings off ground on thread. When paralyzed, spider drags it away using a web sling attached to the ant and to the spinnerets.[127]
Euryopis episinoides (Walckenaer) (publ. as E. acuminata)I/FFeeds on ants. Attacks Crematogaster ants and transports each attached to spinnerets.[128]
Euryopis formosa BanksI/FCaptures and carries workers of Pogonomyrmex salinus Olsen. Carries ant across ground. One attack described: spider bit gaster, released ant, moved to front and waited, reapproached paralyzed ant, climbed onto ant and began dragging across ant nest using web sling.[129]
Euryopis funebris (Hentz)F/OReported feeding on Camponotus castaneus (Latreille). Throws adhesive silk over ant passing by on tree trunk and fastens it to tree. Encircles ant, throwing silk. Bites leg. Cuts paralyzed ant free and carries it to crack or crevice or drops on line to feed.[130, 131]
Euryopis scriptipes BanksI/FFeeds on ants.[132]
Euryopis texana BanksI/FFemale reported preying upon moving line of small ants.[133]
Other Euryopis spp.I/FPrey on ants. Throw adhesive silk over ants and fasten to trees.[131133]
Latrodectus corallinus AbalosUnk[93, 134]
Latrodectus hesperus Chamberlin and IvieProbably IFeeds on Pogonomyrmex rugosus Emery. Builds web on colony mound over foraging trail. Spider throws silk on ant that gets caught in gum threads. Spider approaches ant from above, bites posterior femur, retreats, returns after ant paralyzed, and pulls ant to retreat or to hidden part of web. Also feeds on other species of ants.[93]
Latrodectus mactans (Fabricius)I/F75% of prey in cotton fields in Texas made up of fire ants, Solenopsis invicta Buren. Also reported feeding on Pogonomyrmex badius (Latreille) and P. barbatus.[89, 126, 135]
Latrodectus mirabilis (Holmberg)UnkFeeds on Acromyrmex spp. and Camponotus spp. Builds webs over colony entrances.[93, 134]
Latrodectus pallidus O. P.-CambridgeFPrimary prey are ants. Feeds on Monomorium semirufus (nomen dubium, but probably Messor semirufus (André)). Females build webs over foraging trails. Capture ants from above with trip line attached to substrate and pull prey into retreat. Spiders can also descend to ground and catch ants running on trails.[136138]
L. quartus AbalosUnkFeeds on Acromyrmex spp. and Camponotus spp. Builds webs over colony entrances.[93, 134]
Latrodectus revivensis ShulovUnkRemains of Messor sp. found in webs.[136]
Latrodectus tredecimguttatus (Rossi)UnkRemains of Messor sp. found in webs.[136, 137]
Latrodectus spp.UnkMembers of genus may generally be myrmecophages. Reported feeding on Monomorium sp. and Messor semirufus (André). [136138]
Parasteatoda tepidariorum (C. L. Koch) (publ. as Achaearanea tepidariorum)UnkFeeds on fire ants, Solenopsis invicta Buren.[107]
Phycosoma mustelinum (Simon) (publ. as Dipoena mustelina)UnkCaptures various species of ants of wide range of sizes.[125]
Steatoda albomaculata (De Geer)IFeeds on ants; ant remains found in webs.[139]
Steatoda fulva (Keyserling)I/FReported building webs near nest entrance of colonies of Pogonomyrmex badius (Latreille).[122]
S. triangulosa (Walckenaer)IFeeds on fire ants, Solenopsis invicta Buren.[104]
Yaginumena castrata (Bösenberg and Strand) (publ. as Dipoena castrata)UnkMostly feeds upon Camponotus sp. and Lasius sp. and most individual spiders feed upon single type of prey. The larger the spider, the larger the ant it can attack.[125]

Thomisidae
Amyciaea albomaculata (O. P.-Cambridge)OMyrmecomorph of Oecophylla smaragdina (Fabricius) (publ. as O. virescens). Adult spiders with eye spots on abdomen. Juvs. yellow and mimic other species of yellow ants (transformational mimics). Spider waits near foraging trail of ant, attacks from behind, bites back of body, drags paralyzed ant to edge of vegetation, drops down to feed.[140]
Aphantochilus rogersi O. P.-Cambridge (publ. as Cryptoceroides cryptocerophagum)OAlso a myrmecomorph of Cephalotes pusillus (Klug) (publ. as Zacryptocerus pusillus). Attacks from behind. Holds dead ant as “protective shield.” Females oviposit near ant nest and defend egg sacs against worker ants.[141143]
Aphantochilus spp.UnkFeed on cephalotine ants.[57, 141143]
Bucranium spp.UnkFeed on cephalotine ants. Hold dead ants as protective shield against attacks from other ants.[57, 141143]
Mecaphesa californica (Banks) (publ. as Misumenops californicus)UnkFeeds on Pogonomyrmex rugosus in vegetation near ant nests.[93]
Mecaphesa coloradensis (Gertsch) (publ. as Misumenops coloradensis)UnkFeeds on alate females of Pogonomyrmex maricopa Wheeler and P. desertorum Wheeler after they have removed their wings and while resting on bushes waiting for temperatures to drop in order to dig new nest chambers.[144]
Mecaphesa lepida (Thorell) (publ. as Misumenops lepidus)IOccasionally feeds on ants.[99]
Misumenops argenteus (Rinaldi)Probably I17% of prey are ants; mostly ants that get caught in trichomes of plant Trichogoniopsis adenantha (OC), where spider spends most of its time.[145]
Runcinioides argenteus Mello-Leitão (publ. as Misumenops argenteus)UnkIncludes ants in diet.[146]
Saccodomus formivorus RainbowMay be F or OBuilds a basket-like web that appears to attract wandering Iridomyrmex ants. Spider also uses behavioral tactics-tapping ant with its own legs before attacking.[4, 147]
Thomisus onustus WalckenaerI42.8% of diet consists of ants.[147]
Tmarus stoltzmanni KeyserlingOFeeds exclusively on ants; but only those without stings such as dolichoderine and formicine ants. Uses frontal attacks. May have sensory structures on 1st or 2nd pair of legs to detect chemical or tactile cues from ants.[148]
Other Tmarus sp. (from Australia)UnkIncludes ants in diet.[148, 149]
Xysticus californicus KeyserlingUnkAttacks harvester ants in California (cites unpubl. work of Snelling).[148, 149]
X. loeffleri RoewerRAnts comprise only a minor part of diet.[150]
Other Xysticus spp.I/F30–35% of diet of some spp. of Xysticus comprised of ants. One spider seen preying on Pogonomyrmex salinus Olsen. Spider seen on back of ant where it rode around, biting ant until paralyzed. Spider bit at base of petiole.[129, 150]

Zodariidae
Diores spp.Probably F or OFeed on ants.[151]
Habronestes bradleyi (O. P.-Cambridge)OSpider also myrmecomorph. Waves front legs around when hunting ants. When legs are amputated, spider has difficult time locating prey (Iridomyrmex purpureus (Smith)).[152154]
Lachesana insensibilis JocquéIPolyphagous but will eat ants smaller than themselves. Uses different hunting behaviors for ants than for other prey: bites, releases, re-approaches, bites again.[155]
Lachesana tarabaevi Zonstein and Ovtchinnikov FPreys mostly on harvester ants in genus Messor and on isopods.[156]
Pax islamita (Simon)IPolyphagous but will eat ants smaller than themselves. Uses different hunting behaviors for ants than for other prey: bites, releases, re-approaches, bites again.[155]
Trygetus sexoculatus (O. P.-Cambridge)OParalysis latency longer for male and juvenile attacks than for female attacks.[157]
Trygetus spp.OParalysis latency longer for male and juvenile attacks than for female attacks.[155, 157]
Zodariellum asiaticum (Tyschchenko)OSpecializes on formicine ants. Attacks other kinds of ants readily but there is shorter paralysis latency for formicine ants suggesting biochemical specificity of venom for certain kinds of ants.[155]
Zodariellum spp.Probably all OFeed on ants.[155]
Zodarion cyrenaicum DenisOShows cooperative foraging behavior. But some individuals steal prey from others (kleptoparasitism). Paralysis latency longer for male and juvenile attacks than for female attacks.[157159]
Zodarion frenatum (Simon)OFeeds on Cataglyphis bicolor (Fabricius). Locates nests at night (maybe via odor cues?). Sometimes builds retreats near nest. Digs open closed nest entrances, which triggers ants to come out and repair. Spider sometimes enters nest. Bites ant’s legs and carries paralyzed ant away from nest. Also kills ants in morning when they emerge from nest.[158, 160, 161]
Zodarion germanicum (C. L. Koch)OMyrmecomorph as well as myrmecophage. Waves 1st legs as antennal illusion. Holds dead ant in chelicerae and presents dead ant to approaching live ant while “antennating” live ant with its own forelegs. Presumably presenting both odor and tactile cues to living ant to deceive it and avoid attack. Attacks Cataglyphis bicolor (Fabricius).[162, 163]
Zodarion jozefienae BosmansOFemales and juveniles actively hunt ants. Mature males are kleptoparasites on females’ prey (spend energy on mate searching, not prey capture). Sexual size dimorphism (females larger).[161, 164, 165]
Zodarion lutipes (O. P.-Cambridge)OParalysis latency longer for male and juvenile attacks than for female attacks.[157]
Zodarion nitidum (Audouin)OParalysis latency longer for male and juvenile attacks than for female attacks.[157]
Zodarion rubidum SimonOMyrmecomorph as well as myrmecophage. Waves 1st legs as antennal illusion. Holds dead ant in chelicerae and presents dead ant to approaching live ant while “antennating” live ant with its own forelegs. Presumably presenting both odor and tactile cues to living ant to deceive it and avoid attack.[163, 166168]
Zodarion spp.OAll species obligate myrmecophages. Species also imperfect myrmecomorphs. Documented hunting various species. Do not survive well on non-ant diet. Seem to be behaviorally adapted to hunt ants and seem to have evolved nutritional limitations (non-ant prey do not provide required nutrients). Attack from rear, bite legs, retreat, may repeat, re-approach, pick up, and carry away paralyzed ants. Move front legs while hunting. Have femoral organ that may secrete chemical involved in prey capture.[49, 98, 151, 157, 158, 160, 161, 166, 168174]