Home Field and Swamp: Animals and Their Habitats

        

Bees, Wasps, Sawflies and Ants (order Hymenoptera)

Taxonomic classification categories are valid according to the Integrated Taxonomic Information System (ITIS).   Information about Hymenoptera species history is based on Grimaldi and Engel (2005).  I got some great expert help with bee and wasp identification from John S. Ascher, who also reviewed the changes I made under his direction to make sure they were right.  Even then, there is some chance of error, and I claim full responsibility for it.

The general consensus is that bees (unlike wasps) are a monophyletic group, meaning that they and they alone come from a certain common ancestor: bees are more closely related than the various wasps, and are more closely related to some wasps than are other wasps.  Most bee species are solitary bees, i.e., females of those species are designed to reproduce.  The main behavioral distinction between these categories is that bees feed on honey which they create from gathered flower nectar, while wasps are predators, parasites or parasitoids (parasites that kill their host, usually another much larger animal that they live inside as larvae).

Web-spinning and Leaf-rolling Sawflies (family Pamphiliidae, superfamily Megalodontoidea, suborder Symphyta)

The Pamphiliidae family first appeared about 190 million years ago, during the Jurassic Period, 40 million years before Pangaea, the original continent, started to break up, according to Grimaldi and Engel (2005).

Leaf-rolling Sawfly, Mason Farm Biological Reserve, Orange County, NC, 6/7/06.  According to John S. Ascherit is an Onycholyda amplecta.  See BugGuide page. Leaf-rolling Sawfly (Onycholyda amplecta), Pettigrew State Park, Washington County, NC, 4/28/09

Horntails (family Siricidae, superfamily Siricoidea, suborder Symphyta)

This is also a very old family, which first appeared about the same time: 190 million years ago.

Horntail (Tremex columba), Penny's Bend, Durham County, NC, 6/4/06.  This was an especially large insect, at least 2 inches long. 

Argid Sawflies (family Argidae, superfamily Tenthredinoidea, suborder Symphyta)

         
Argid Sawfly (Arge genus. quidia or scapularus species), Mason Farm Biological Reserve, Orange County, NC, 5/21/09.  Genus ID and limitation of species ID possibilities thanks to Dave Smith. Sawfly (Arge humeralis), Ft. Fisher Basin Trail, New Hanover County, NC, 8/12/11 Same Arge humeralis          

Cimbicid Sawflies (family Cimbicidae, superfamily Tenthredinoidea, suborder Symphyta)

Not all sawflies are members of unusually ancient families.  The Cimbicidae first appeared about 60 million years ago.

Sawfly (Abia genus), Sandy Creek Park, Durham, NC, 4/1/10.  Not a bee, but an Abia!  This genus used to be known as Zaraea, apparenatly. Sawfly (Cimbex Americana), Indian Creek trail in Chatham County, NC, on 5/3/05.  Thanks to Prof. Norman F. Johnson for family ID.

Common Sawflies (family Tenthredinidae, superfamily Tenthredinoidea, suborder Symphyta)
         
Common Sawfly family member (Tenthredo genus, Tenthredinidae family).  ID thanks to Dave Smith. Willow sawfly (Nematus genus), Durham, NC, 4/22/14. ID thanks to Dave Smith.          

Sawfly Larvae
         
Sawfly larva, curled up on this chilly morning          

Bees and Sphecid Wasps (superfamily Apoidea, suborder Apocrita)

Many members of this group are tiny solitary bees much smaller than honeybees.  Solitary bees are those whose females are all genetically capable of reproducing.  They are frequently found together with Large Bee Flies (Bombylius major), which parasitize their larvae.  Social bees, those that live in colonies that each rely on a single queen bee for reproduction, include the Honeybees, Bumblebees, Stingless Bees (not pictured), and some Sweat Bees (which have small colonies and minimal size differences between queens and workers).

Bees may not be the most common pollinators, but they are the most significant economically because they are the most efficient, in part because of their special pollen-gathering hairs and "pollen baskets" which carry large amounts of pollen.

Performing formal identifications of solitary bees from photos is hard because they are so closely related, and therefore such identifications depend heavily on very small details, such as wing venation.   In some cases one can distinguish the long-tongued bees (the Megachilidae and the familiar Apidae) from the short-tongued bees (the Colletidae, Halictidae, Andrenidae, Strenotridae, and Melittidae) by the types of flowers that they visit.

John S. Ascher, Ph.D. provided identifications and background information for a large proportion of the bees and wasps pictured below, making a major contribution to the quality of this website.   If errors remain, however, they may represent mistakes I have made in my interpretation of the information he has offered.

Bee Phylogeny at Cornell University provides detailed descriptions of recent bee phylogeny research using morphological, DNA, and fossil data.

Sweat Bees (Halictidae family)

These are short-tongued bees, which these photos illustrate: they prefer composite flowers and "flat" individual ones.  They are called "sweat bees" because of their reputation for landing on people and drinking their sweat.

Some of these bees are social, living in small colonies, i.e., with less than a dozen members.  There is a lot of diversity in this respect, so they are of special interest to evolutionary biologists.

Sweat Bee (Augochlorini tribe), Eno River SP, Old Cole Mill Road access, 5/29/05.   This green bee is one of the most obvious members of this family; however, most Halictidae are not green, although they are less hairy than other bees. Sweat bee (Augochlorini tribe), Durham, NC, 7/1/08 Sweat bee (Augochlorini tribe), Durham, 9/28/06 Sweat bee (Augochlorini tribe), Indian Creek Trail, a Jordan Lake Game Land, Chatham County, NC, 7/7/06

 

         
Halictid bee (subgenus Dialictus, Lasioglossum genus, Halictini tribe, Halictinae subfamily), Indian Creek Wildlife Observation Trail, Chatham County, NC, 4/5/06.  This flower was about ⅓ inch across. Female Halictid bee (Lasioglossum genus), White Pines Nature Preserve, Chatham County, NC, 4/16/06.  This bee was about inch long.  The tiny fly was barely visible to me without the camera's magnifying lens. Halictid bee (Lasioglossum genus, Durham, 5/3/05.           

 

Sweat bee,  Moses Cone Memorial Park, Watauga County, NC, 8/8/06 Sweat bee,  Moses Cone Memorial Park, Watauga County, NC, 8/8/06 Sweat bee,  Moses Cone Memorial Park, Watauga County, NC, 8/8/06


Female Halictid Bees (Agapostemon virescens), Winston-Salem, Forsyth County, NC, 7/31/06.

Mining Bees (Andrenidae family)

These are short-tongued bees solitary bees, although they apparently build their nests in holes in the ground close to one another in the early spring when pollen is first available.  As is the case with all flower-visiting bees, they have long hairs over many parts of their bodies.

   
Andrena genus, Penny's Bend, Durham County, NC, 3/16/06.  This was a member of a very fast-moving cluster of bees near the ground on a wooded trail.  This bee seemed ill or injured, otherwise this photo would have been impossible to take!  This bee was about inch long.    

Plasterer Bees (Colletidae family)

   
Plasterer Bee (Colletes thoracicus), Durham, NC, 3/9/07.  This bee wandered onto the asphalt path, apparently unable to fly because of what seemed to be clay on its thorax. Two Plasterer Bees (Colletes thoracicus), Durham, NC, 3/18/09 Plasterer Bees (Colletes thoracicus), Johnston Mill Nature Preserve, Orange County, NC, 3/14/08.    Plasterer Bee (Colletes inaequalis, right) with nest, in right bottom of this photo.  Nags Head Woods, Kill Devil Hills, Dare County, NC, 2/18/11  

         
Plasterer bee, hovering near nest entrance.  She crawled in and out of it several times.  Durham, NC, 3/18/11 Plasterer bee entering nest          

 Mason Bees and Leafcutter Bees (Megachilidae family)

These are long-tongued bees, able to visit longer, skinnier flowers.  Members of the Megachile genus do the leaf-cutting, as shown in the first two photos.  Some of these bees are actually parasites and are therefore not either masons or leafcutters; however, since they share this characteristic with members of other families, this description is not included in the accepted common name for this family. 

Leafcutter Bees (Megachile genus)

There was a large group of such bees in the location I found those two, mostly hidden under a clump of wild grape vines, buzzing very loudly.  Note the yellow pollen pocket on the abdomen underside, a unique Megachilid characteristic.

         
Leafcutter bee (Megachile genus) on an aster at the North Carolina Zoo, Asheboro, NC, 9/30/11 Leafcutter bee (Megachile genus), apparently in the process of leaf-cutting at the Eno River State Park, Durham County, NC, 8/17/08 in a power line cut. Same Megachile  genus leafcutter bee, further along in the process Megachilid bee (Megachile genus), Durham, NC, 8/19/07 Megachilid bee (Megachile genus), Penny's Bend Nature Preserve, Durham County, NC, 6/30/06 Female Megachilid bee (Megachile genus), Carolina Beach State Park, New Hanover County, NC, 10/13/06        

       
Megachilid bee (Megachile xylocopoides, family Megachilidae), Carolina Beach State Park, New Hanover County, NC, 10/19/05.  Unlike the other bees on this page, this one was large, perhaps an inch long.  Thanks to Eric Eaton for ID info. Megachile texana, Mason Farm Biological Reserve, Orange County, NC, 7/12/10.  ID thanks to John S. Ascher.        

Coelioxys genus

These are kleptoparasites or "cuckoo" bees, i.e., they lay their eggs in other Megachilidae family members' nests to be raised by unsuspecting adults.

Bee (Coelioxys mexicana), Buccaneer State Park, Waveland, Hancock County, MS, 10/15/10.  First expert-identified photo of this species on BugGuide, on this page. Female Megachilid bee (Coelioxys modesta), NC Botanical Garden, 7/2/05.  Thanks to Eric Eaton for ID.

Apidae family

These female Nomada genus bees are long-tongued bees.  According to Dr. Ascher, most Nomada species are kleptoparasites of genus Andrena, laying their eggs in Andrena nests, while other Nomada species are cleptoparasites of Agapostemon. 

Nomadinae subfamily

Nomada genus

 
Cuckoo Bee (Nomada ruficornis?), North Carolina Botanical Garden, Chapel Hill, Orange County, NC, 4/23/15 Cuckoo Bee (Nomada ruficornis), Penny's Bend Nature Preserve, Durham County, NC, 4/22/09.  ID thanks to John S. Ascher.  Full ID: BugGuide page.   Solitary bee (probably Nomada genus), Penny's Bend Nature Preserve, Durham County, NC, 4/20/07.   This bee was in frantic motion. Nomada species bee, Penny's Bend Nature Preserve, 5/7/07.

Triepeolus genus

         
Cuckoo bee (Triepeolus genus).  Tip of the tongue is visible.  Mason Farm Biological Reserve, Orange County, NC, 9/19/11 Same Cuckoo bee (Triepeolus genus).          

Xylcopinae subfamily

         
Carpenter Bee, Museum of Life and Science grounds, Durham, NC, 4/23/09  Carpenter Bee (Xylocopa virginica, Xylocopinae subfamily), Eno River SP, Orange County, NC, 4/2/06.   Valid taxa according the the ITIS.  This large bee is well-known to all, and dreaded by owners of wooden houses.  Thanks to Eric Eaton for ID. Small carpenter bee (Ceratina genus, Zadontomerus subgenus), on a Smooth Coneflower, Penny's Bend Nature Preserve, Durham County, NC, 6/30/06.  Little carpenter bee (Ceratina genus, Zadontomerus subgenus). ID thanks to John S. Ascher.
       

Apinae subfamily

We depend on honeybees for pollination of most fruit, vegetable and nut crops.  Because honeybees are dying in significant numbers, beekeepers predict disaster for humans, according to http://www.allheadlinenews.com, article by Ihuoma Ezeh.
European Honeybee (Apis mellifera), Raulston Arboretum, Raleigh, Wake County, NC, 3/17/06.  European Honeybee. Durham, NC, 10/21/09 European Honeybee, Duke Gardens, Durham, NC, 7/21/07.  The pollen of this moss rose was enough to cover several bees.
       
         



       
Long-horned bee (Eumelissodes subgenus, Melissodes genus, Eucerini tribe),  Boone, Watauga County, NC, 8/7/06 Long-horned bee (Eucerini tribe), Durham, 9/20/08        

Bumblebees (genus Bombus, tribe Bombini)
         
Bumble Bee on thistle, Flat River Impoundment, Durham County, NC, 8/5/11 Bumblebee on an aster, Weymouth Woods-Sandhills Nature Preserve, Southern Pines, Moore County, NC, 10/31/14          

Thread-waisted Wasps (family Sphecidae, superfamily Apoidea, suborder Apocrita)

The Sphecidae first appeared about 140 million years ago.

Ammophila pictipennis,  Raulston Arboretum, Wake County, NC, 9/23/05.  Ammophila pictipennis, Greenville, NC, 9/20/08 Thread-waisted wasp (Ammophila genus) and two small weevils on Common Sneezeweed, Flat River Impoundment, Durham County, NC, 7/17/11 Eremnophila aureonotata,  Durham, NC, 9/2/09 Eremnophila aureonotata, mating pair, Durham, 7/28/03.  These are Sphecidae according to the University of Michigan Museum of Zoology.  Thanks to Josh Rose for genus and species ID. Grass-carrying Wasp (Isodontia genus), Durham, NC, 7/7/08, trying to access nectar of Trumpet Vine (Campsis radicans) flower buds.


         
Thread-waisted wasp (Prionyx genus, Prionchini tribe, Sphecinae subfamily), Greenville, Pitt County, NC, 9/20/08 Thread-waisted Wasp (Prionyx genus?), perhaps resting.  Ft. Fisher Basin Trail, New Hanover County, NC, 5/24/11.          

Bee Wolves and Sand Wasps (family Crabronidae, superfamily Apoidea, suborder Apocrita)

Crabronid wasp (Philanthus gibbosus), NC Botanical Garden, Orange County, NC, 8/28/06.  This is a predator of halictid bees. Cicada Killer (Sphecius speciosus, Handlirschiina subgenus, Bembicini tribe, Bembicinae subfamily), Durham, NC, 8/1/08 Cicada Killer, Duke Gardens, Durham, NC, 7/1/08

Gall Wasps (family Cynipidae, superfamily Cynipoidea, infraorder Terebrantes, suborder Apocrita)

       
Wool Sower Gall (Callirhytis seminator, tribe Cynipini, subfamily Cynipinae), Durham, NC, 4/19/09        

Encyrtid Wasps (family Encyrtidae,superfamily Chalcoidea, suborder Apocrita)

         
Wasp.  About 1 mm long.  Family ID thanks to Ken Wolgemuth and  Ross Hill.  Ross is an expert in the field.  Third Fork Creek Trail, Durham, NC, 12/31/11.          

Ichneumonid Wasps (family Ichneumonidae, superfamily Ichneumonoidea, suborder Apocrita)

We don't usually think of wasps as being especially beautiful, but this family has more than its share of bright colors and striking patterns.  What these parasitoids do to their prey is not, however:  Charles Darwin cited them in particular in his remarks to botanist Asa Gray about how his faith in God's goodness was tested by the torment these animals inflicted on one another.

IDs of family of wasps in first row thanks to Prof. Norman F. Johnson.

Ichneumon wasp (Ephialtini tribe, Pimplinae subfamily), Durham, NC, 11/9/13. ID thanks to Bob Carlson. Ichneumonid wasp, Durham, 10/28/05 Ichneumonid Wasp, Ft. Fisher Recreational Area, New Hanover County, NC, 12/3/07 Ichneumonid wasp, Falls Lake State Park, Wake County, NC, 1/6/07 Ichneumonid Wasp (Cratichneumon variegatus), Durham, NC  9/16/06.  Taken at night.  ID thanks to Bob Carlson Ichneumonid wasp (Arotes genus, Acaenitinae subfamily), Durham, 9/12/05

 

Ichneumonid Wasp, Durham, 10/29/05 Ichneumonid Wasp (Labena grallator, Labenini tribe), Eno River State Park, 11/7/05.  Ichneumonid Wasp, Penny's Bend Nature Preserve, Durham County, NC, 4/5/06. ID thanks to Dr. Doug Yanega of the University of California at Riverside. Ichneumonid Wasp, Durham, 4/14/06.  ID thanks to Dr. Doug Yanega of the University of California at Riverside.  Photo taken at night. Giant ichneumonid wasp, subfamily Rhyssinae.  This wasp's ovipositor was easily six inches long.   Moses Cone Memorial Park, Watauga County, NC, 8/8/06

 

Durham, 10/1/05 Another view of the wasp on the left. Ichneumonid wasp

 

Ichneumonid Wasp, Eno River State Park, Orange County, NC, 10/9/06 Ichneumonid wasp, Penny's Bend Nature Preserve, Durham County, NC  4/20/06 Ichneumonid wasp, Johnston Mill Nature Preserve, Orange County, NC, 6/16/07 Ichneumonid wasp, Durham, NC, 4/8/08.  Photo taken at night.


       
Ichneumonid wasp, Pimplinae subfamily, Camp Taylor, Marin County, CA, 8/4/07,  ID thanks to Ross Hill. Ichneumon wasp, Weymouth Woods-Sandhills Nature Preserve, Southern Pines, Moore County, NC, 10/23/12        

Brachonid Wasps (family Brachonidae, superfamily Ichneumonoidea, suborder Apocrita)

 
Braconid wasp, Mason Farm Biological Reserve, Orange County, NC, 9/23/07.  Braconid wasp (Atanycolus genus), Cypress Gardens, Berkeley County, SC, 10/12/07

Pelecinid Wasps (family Pelecinidae, superfamily Pelecinoidea, suborder Apocrita)

Pelecinid Wasp, Pelecinus polyturator, Penny's Bend Nature Preserve, Durham County, NC, 8/13/06 Pelecinid Wasp (Pelecinus polyturator), Mason Farm Biological Reserve, Orange County, NC, 9/29/06

Cuckoo Wasps (family Chrysididae, superfamily Bethyloidea, suborder Apocrita)

These wasps first appeared about 140 million years ago, during the early Cretaceous Period.

Cuckoo Wasp (possibly genus Holopyga), Duke Gardens, 9/17/05.  Chrysis seems to be most likely, however.  Prof. Johnson OKed this family ID. Cuckoo Wasp, Ft. Fisher Aquarium (outside), New Hanover County, NC, 6/26/08

Spider Wasps (family Pompilidae, superfamily Vespoidea, suborder Apocrita)

The Pompilidae evolved about 105 million years ago.

Spider wasp (Psorthaspis mariae), Falls Lake Dam area, 9/11/10.  It successfully mimicked a velvet ant in its movements.  It actually looks more like the checkered beetle on the left. Spider wasp, very active but with a wing problem.  Third Fork Creek Trail, Durham, NC, 7/18/11 Spider Wasp (Arachnospila genus), Johnston's Mill, Orange County, NC, 2/3/06 Spider Wasp in action, Eno River SP, Old Cole Mill Road access, Orange County, NC, 4/28/06.  This wasp dragged this spider at least five feet while I watched. Spider wasp attempting to drag a spider across pavement, Durham, NC, 5/13/08

Vespid Wasps (family Vespidae, superfamily Vespoidea, suborder Apocrita)

The Vespidae family evolved about 140 million years ago.

   
Possibly a European Hornet (Vespa crabro), Johnston Mill Nature Preserve, Orange County, NC, 6/10/06 Yellow Jackets (Vespula vulgaris?) mating, Durham, NC, 11/20/08.  Female is on left. Male Yellow Jacket.  Note how the male's abdominal markings differ from those of the female:  the male is mostly yellow, while the female is mostly black.    

     
Paper wasp (Polistes genus) on Pinkweed.  Durham, NC, 9/5/09. Paper wasp (Polistes genus), Durham, NC, 4/30/10 Paper wasp (Polistes genus), Southpoint Swamp, Durham, NC, 10/4/10      

         
Mason Wasp (Euodynerus schwarzi), Flat River Impoundment, Durham, NC, 8/15/10 Mason Wasps (Monobia quadridens), Eno River State Park, Orange County, NC, 5/18/11 Potter Wasp (Eumenes fraternus), Durham, 9/28/09.  This wasp is apparently mimicked by the Sphiximorpha willistoni flower fly. Potter Wasp, Penny's Bend Nature Preserve, Durham County, NC, 4/20/07.           

Tiphiid Wasps (family Tiphiidae, superfamily Vespoidea, suborder Apocrita)

Most of these are parasitoids, i.e., they lay their eggs within host animals.

   
Tiphiid wasps (Myzinum genus), Carolina Beach State Park, New Hanover County, NC, 6/23/07, struggling to get into one little hole in the sand.    

Scoliid Wasps (genus Scolia, family Scoliidae, superfamily Scolioidea, suborder Apocrita)

 
Scoliid wasp (Scolia nobilitata) on Common Sneezeweed.  ID thanks to Brian Bockhahn. Wasp (Scolia nobilitata), Durham, NC< 7/12/08.  NOTE: The ITIS does not list the species nobilitata. Wasp (Scolia dubia), Greenville, Pitt County, NC, 9/20/08 Wasp (Campsomeris plumipes), Fayetteville, NC, 7/16/08.  This was a big wasp, about an inch long.   Photo by Kathryn Cox. Wasp (Campsomeris genus), Weymouth Woods-Sandhills Nature Preserve, Southern Pines, Moore County, NC, 10/31/14  

 Velvet Ants (family Mutillidae, superfamily Scolioidea, suborder Apocrita)

These are parasitoids, i.e., females parasitize bee larvae by laying eggs on them.  They are generally considered to be wasps because of their behavior but their superfamily name implies that they aren't very distant relatives of ants!  The Cowkiller is named for its nasty sting, one informally hypothesized to be bad enough to kill a cow.

The Mutillidae family evolved relatively recently, about 45-70 million years ago.

   
Female Velvet Ant (Dasymutilla quadriguttata), White Pines Nature Preserve, Chatham County, NC, 4/16/06.  ID thanks to George Waldren Cowkiller (Dasymutilla occidentalis), Mason Farm Biological Reserve, Orange County, NC, 9/1/07.  See other Hymenopterans (bees, wasps and ants). Perhaps a Cowkiller, a velvet ant which is normally red and black.  It was hunched over, trying to hide. NOTE: the two small white spots which appear to be on this insect's back are actually on the ground: it has a typically skinny wasp waist. Carolina Beach State Park, 9/18/07 Velvet ant (Pseudomethoca simillima) #1. investigating the object on the left. Velvet Ant (Timulla genus?).  The extra abdominal ring is an artifact of photography.  Third Fork Trail, Durham, NC, 7/14/11    

Ants (family Formicidae, superfamily Scolioidea, suborder Apocrita)

Ants communicate with one another using pheromones, organic compounds which range from simple hydrocarbons called unbranched alkanes to somewhat more complex alkenes, each containing an oxygen atom.

The Formicidae family evolved about 140 million years ago.

Slavemaking Ants (Formica subintegra, Formicinae subfamily)

Social Parasitism in Ants

         
Slavemaking ant (Formica subintegra), carrying Formica subsericea ant captive.  ID thanks to James C. Trager.  One of a fast-moving colony.          

Small ants (Nylanderia Faisonensis, subfamily Formicinae)

Leaving a rather flat watering can on its side and turning it over every few days can produce surprising revelations of what normally goes on underground.   Ant larvae are unusual in that their limbs and wings are still developing (and somewhat visible), and the pupa stage in this species seems to be a brown-colored later part of the larva stage.  These ants quickly returned underground afterwards.

         
You can see some brown pupae; the one on the right has adult features showing through.  However, you can see developing legs on the white larvae. Winged ants, also walking on larvae.          

Yellow Ants (Acanthomyops interjectus, tribe Lasiini, subfamily Formicinae)

Winged Yellow Ants, Durham, 5/31/06.  The queens are brown, while the workers are yellow. These winged Yellow Ants came up out of the ground, crawled up plants in this meadow and flew into the air.  Southern Village, Orange County, NC, 11/13/07 Yellow Ant, with a red parasite egg on its right hind leg Winged ant being attacked by a tiny wasp?  Carolina Beach State Park, New Hanover County, NC, 3/9/10

Carpenter Ants (Camponotus genus, Camponotini tribe, Formicinae subfamily)

         
Black Carpenter Ant (Camponotus pennsylvanicus) tending aphids, Durham, NC, 4/25/08.  This ant was actually horizontal in the photo: rotation in the thumbnail is due to a glitch in the software.           

 Unknown Ants
   
Johnston's Mill, Orange County, NC, 2/3/06.  This ant was wandering around rocks in New Hope Creek. Worker ant subduing a winged termite, Occoneechee Mountain Natural Area, Orange County, NC, 4/9/06 Worker ant carrying a larva.    

Mystery Bee and Wasps
Mystery wasp, Durham, 6/28/06.  Maybe a braconid wasp.

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2006-2011 Dorothy E. Pugh, except where noted on this page.

REFERENCES

Genus Coelixys.  Retrieved on December 12, 2010 from http://bugguide.net/node/view/7605

Moisset, Beatriz (2010).  Personal communication.

Deslippe, R. Social Parasitism in Ants (2010). Nature Education Knowledge 1(8):27.  Retrieved 3 August 2011 from http://www.nature.com/scitable/knowledge/library/social-parasitism-in-ants-13256421