Editing Bee (section)

== Evolution ==
The immediate ancestors of bees were [[Aculeata|stinging wasp]]s in the family [[Crabronidae]], which were [[Predation|predators]] of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same [[evolution]]ary scenario may have occurred within the [[vespoidea|vespoid]] wasps, where the [[pollen wasp]]s evolved from predatory ancestors.<ref name="Cardinal danforth 2011"/>

Based on phylogenetic analysis, bees are thought to have originated during the [[Early Cretaceous]] (about 124 million years ago) on the supercontinent of [[Gondwana|West Gondwana]], just prior to its breakup into [[South America]] and [[Africa]]. The supercontinent is thought to have been a largely [[Deserts and xeric shrublands|xeric]] environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong [[Phylogenetic niche conservatism|niche conservatism]] among bees ever since their origins.<ref name=":1">{{Cite journal |last1=Almeida |first1=Eduardo A. B. |last2=Bossert |first2=Silas |last3=Danforth |first3=Bryan N. |last4=Porto |first4=Diego S. |last5=Freitas |first5=Felipe V. |last6=Davis |first6=Charles C. |last7=Murray |first7=Elizabeth A. |last8=Blaimer |first8=Bonnie B. |last9=Spasojevic |first9=Tamara |last10=Ströher |first10=Patrícia R. |last11=Orr |first11=Michael C. |last12=Packer |first12=Laurence |last13=Brady |first13=Seán G. |last14=Kuhlmann |first14=Michael |last15=Branstetter |first15=Michael G. |date=2023-08-21 |title=The evolutionary history of bees in time and space |journal=Current Biology |volume=33 |issue=16 |pages=3409–3422.e6 |doi=10.1016/j.cub.2023.07.005 |issn=0960-9822|doi-access=free |pmid=37506702 |bibcode=2023CBio...33E3409A }}</ref>

Genomic analysis indicates that despite only appearing much later in the fossil record, all modern bee families had already diverged from one another by the end of the Cretaceous. The [[Melittidae]], [[Apidae]], and [[Megachilidae]] had already evolved on the supercontinent prior to its fragmentation. Further divergences were facilitated by West Gondwana's breakup around 100 million years ago, leading to a deep Africa-South America split within both the Apidae and Megachilidae, the isolation of the Melittidae in Africa, and the origins of the [[Colletidae]], [[Andrenidae]] and [[Halictidae]] in South America. The rapid radiation of the South American bee families is thought to have followed the concurrent radiation of [[flowering plant]]s in the same region. Later in the Cretaceous (80 million years ago), colletid bees colonized [[Australia]] from [[South America]] (with an offshoot lineage evolving into the [[Stenotritidae]]), and by the end of the Cretaceous, South American bees had also colonized North America.<ref name=":1" /> The North American fossil taxon ''[[Cretotrigona]]'' belongs to a group that is no longer found in North America, suggesting that many bee lineages went extinct during the [[Cretaceous–Paleogene extinction event|Cretaceous-Paleogene extinction event]].<ref name=":1" />

Following the K-Pg extinction, surviving bee lineages continued to spread into the Northern Hemisphere, colonizing [[Europe]] from Africa by the [[Paleocene]], and then spreading east to [[Asia]]. This was facilitated by the warming climate around the same time, allowing bees to move to higher latitudes following the spread of tropical and subtropical habitats. By the [[Eocene]] (~45 mya) there was already considerable diversity among eusocial bee lineages.<ref name="pnas3054932">{{cite journal |last=Engel |first=Michael S. |year=2001 |title=Monophyly and Extensive Extinction of Advanced Eusocial Bees: Insights from an Unexpected Eocene Diversity |journal=PNAS |publisher=National Academy of Sciences |volume=98 |issue=4 |pages=1661–1664 |bibcode=2001PNAS...98.1661E |doi=10.1073/pnas.041600198 |jstor=3054932 |pmc=29313 |pmid=11172007 |doi-access=free}}</ref>{{efn|[[Triassic]] nests in a petrified forest in Arizona, implying that bees evolved much earlier,<ref name=Buchmann2012/> are now thought to be beetle borings.<ref>{{cite journal |last1=Lucas |first1=Spencer G. |last2=Minter |first2=Nicholas J. |last3=Hunt |first3=Adrian P. |title=Re-evaluation of alleged bees' nests from the Upper Triassic of Arizona |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |date=February 2010 |volume=286 |issue=3–4| pages=194–201 |doi=10.1016/j.palaeo.2010.01.010 |bibcode=2010PPP...286..194L }}</ref>}} A second extinction event among bees is thought to have occurred due to rapid climatic cooling around the [[Eocene–Oligocene extinction event|Eocene-Oligocene boundary]], leading to the extinction of some bee lineages such as the tribe [[Melikertini]]. Over the Paleogene and [[Neogene]], different bee lineages continued to spread all over the world, and the shifting habitats and connectedness of continents led to the isolation and evolution of many new bee tribes.<ref name=":1" />

=== Fossils ===
The oldest non-compression bee fossil is ''[[Cretotrigona prisca]]'', a [[Apinae|corbiculate bee]] of [[Late Cretaceous]] age (~70 mya) found in [[New Jersey amber]].<ref name="Cardinal danforth 2011">{{cite journal |author1=Cardinal, Sophie |author2=Danforth, Bryan N. |year=2011 |title=The Antiquity and Evolutionary History of Social Behavior in Bees |journal=PLOS ONE |volume=6 |issue=6 |pages=e21086 |bibcode=2011PLoSO...621086C |doi=10.1371/journal.pone.0021086 |pmc=3113908 |pmid=21695157 |doi-access=free}}</ref> A fossil from the early Cretaceous (~100 mya), ''[[Melittosphex burmensis]]'', was initially considered "an extinct lineage of pollen-collecting Apoidea [[Cladistics|sister]] to the modern bees",<ref name="Poinar">{{cite journal |author1=Poinar, G. O. |author2=Danforth, B. N. |year=2006 |title=A fossil bee from Early Cretaceous Burmese amber |url=http://fossilinsects.net/pdfs/Poinar_Danforth_2006_MelittosphexBurmese.pdf |url-status=dead |journal=Science |volume=314 |issue=5799 |page=614 |doi=10.1126/science.1134103 |pmid=17068254 |s2cid=28047407 |archive-url=https://web.archive.org/web/20121204122518/http://fossilinsects.net/pdfs/Poinar_Danforth_2006_MelittosphexBurmese.pdf |archive-date=4 December 2012}}</ref> but subsequent research has rejected the claim that ''Melittosphex'' is a bee, or even a member of the superfamily [[Apoidea]] to which bees belong, instead treating the lineage as ''[[incertae sedis]]'' within the [[Aculeata]].<ref name="Allommation">{{Cite journal |last1=Rosa |first1=B. B. |last2=Melo |first2=G. A. R. |year=2021 |title=Apoid wasps (Hymenoptera: Apoidea) from mid-Cretaceous amber of northern Myanmar |journal=Cretaceous Research |volume=122 |pages=Article 104770 |bibcode=2021CrRes.12204770R |doi=10.1016/j.cretres.2021.104770 |issn=0195-6671 |s2cid=234071940}}</ref>

The [[Allodapini]] (within the Apidae) appeared around 53 Mya.<ref name="danforth2012">{{cite journal |last1=Danforth |first1=Bryan |last2=Cardinal |first2=Sophie |last3=Praz |first3=Christophe |last4=Almeida |first4=Eduardo |last5=Michez |first5=Denis |s2cid=28274420 |title=The Impact of Molecular Data on Our Understanding of Bee Phylogeny and Evolution |journal=Annual Review of Entomology |date=28 August 2012 |volume= 58 |pages=57–78 |doi=10.1146/annurev-ento-120811-153633 |pmid=22934982 }}</ref>
The Colletidae appear as fossils only from the late [[Oligocene]] (~25 Mya) to early [[Miocene]].<ref>{{cite journal |last1=Almeida |first1=Eduardo A. B. |last2=Pie |first2=Marcio R. |last3=Brady |first3=Sean G. |last4=Danforth |first4=Bryan N. |title=Biogeography and diversification of colletid bees (Hymenoptera: Colletidae): emerging patterns from the southern end of the world |journal=Journal of Biogeography |year=2012 |volume=39 |issue=3 |pages=526–544 |url=http://entomology.si.edu/staffpages/Brady/2012_AlmeidaPieBradyDanforth_jBiogeog.pdf |doi=10.1111/j.1365-2699.2011.02624.x |bibcode=2012JBiog..39..526A |s2cid=34626231 |url-status=live |archive-url=https://web.archive.org/web/20130921025319/http://entomology.si.edu/StaffPages/Brady/2012_AlmeidaPieBradyDanforth_jBiogeog.pdf |archive-date=21 September 2013}}</ref>
The Melittidae are known from ''Palaeomacropis eocenicus'' in the [[Ypresian|Early Eocene]].<ref>{{cite journal |last1=Michez |first1=Denis |last2=Nel |first2=Andre |last3=Menier |first3=Jean-Jacques |last4=Rasmont |first4=Pierre |title=The oldest fossil of a melittid bee (Hymenoptera: Apiformes) from the early Eocene of Oise (France) |journal=Zoological Journal of the Linnean Society |year=2007 |volume=150 |issue=4 |pages=701–709 |url=http://www.atlashymenoptera.net/biblio/194_Michez_et_al_2007_Oldest_Melittid_from_Oise_Palaeomacropis.pdf |doi=10.1111/j.1096-3642.2007.00307.x |url-status=live |archive-url=https://web.archive.org/web/20150923175837/http://www.atlashymenoptera.net/biblio/194_Michez_et_al_2007_Oldest_Melittid_from_Oise_Palaeomacropis.pdf |archive-date=23 September 2015|doi-access=free }}</ref>
The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the [[Eocene|Middle Eocene]].<ref>{{cite journal |last1=Sarzetti |first1=Laura C. |last2=Lanandeira |first2=Conrad C. |last3=Genise |first3=Jorge F. |title=A Leafcutter Bee Trace Fossil from the Middle Eocene of Patagonia, Argentina, and a Review of Megachilid (Hymenoptera) Ichnology |journal=Palaeontology |year=2008 |volume=51 |issue=4 |pages=933–994 |doi=10.1111/j.1475-4983.2008.00787.x |url=https://www.researchgate.net/publication/229562255 |bibcode=2008Palgy..51..933S |url-status=live |archive-url=https://web.archive.org/web/20150924123512/http://www.researchgate.net/profile/Conrad_Labandeira/publication/229562255_A_LEAFCUTTER_BEE_TRACE_FOSSIL_FROM_THE_MIDDLE_EOCENE_OF_PATAGONIA_ARGENTINA_AND_A_REVIEW_OF_MEGACHILID_%28HYMENOPTERA%29_ICHNOLOGY/links/53fe0d4b0cf23bb019bd1b59.pdf |archive-date=24 September 2015|hdl=11336/100644 |hdl-access=free }}</ref>
The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.<ref>{{cite journal |last1=Dewulf |first1=Alexandre |last2=De Meulemeester |first2=Thibaut |last3=Dehon |first3=Manuel |last4=Engel |first4=Michael S. |last5=Michez |first5=Denis |title=A new interpretation of the bee fossil Melitta willardi Cockerell (Hymenoptera, Melittidae) based on geometric morphometrics of the wing |journal=ZooKeys |year=2014 |issue=389 |pages=35–48 |doi=10.3897/zookeys.389.7076 |pmid=24715773 |pmc=3974431|doi-access=free |bibcode=2014ZooK..389...35D }}</ref>
The Halictidae first appear in the Early Eocene<ref>{{cite journal | last1=Engel | first1=M.S. | last2=Archibald | first2=S.B. | year=2003 | title=An Early Eocene bee (Hymenoptera: Halictidae) from Quilchena, British Columbia | doi=10.4039/n02-030| journal=The Canadian Entomologist | volume=135 | issue=1| pages=63–69| hdl=1808/16473 | s2cid=54053341 | url=https://kuscholarworks.ku.edu/bitstream/1808/16473/1/Engel_CE_135%281%2963.pdf |archive-url=https://web.archive.org/web/20170812132942/https://kuscholarworks.ku.edu/bitstream/1808/16473/1/Engel_CE_135%281%2963.pdf |archive-date=2017-08-12 |url-status=live | hdl-access=free }}</ref> with species<ref name="Engel1995">{{cite journal |last1=Engel |first1=M.S. |year=1995 |title=''Neocorynura electra'', a New Fossil Bee Species from Dominican Amber (Hymenoptera:Halictidae) |journal= Journal of the New York Entomological Society |volume=103 |pages=317–323 |jstor=25010174 |issue=3}}</ref><ref name="Engel2000">{{cite journal |last1=Engel |first1=M.S. |year=2000 |title=Classification of the bee tribe Augochlorini (Hymenoptera, Halictidae) |journal=Bulletin of the American Museum of Natural History |volume=250 |pages=1 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/1598/1/B250.pdf |url-status=live |archive-url=https://web.archive.org/web/20110110015144/http://digitallibrary.amnh.org/dspace/bitstream/2246/1598/1/B250.pdf |archive-date=10 January 2011|doi=10.1206/0003-0090(2000)250<0001:COTBTA>2.0.CO;2 |hdl=2246/1598 |s2cid=85810077 }}</ref> found in amber. The Stenotritidae are known from fossil brood cells of [[Pleistocene]] age.<ref>{{cite journal | last1=Houston | first1=T.F. | year=1987 | title=Fossil brood cells of stenotritid bees (Hymenoptera: Apoidea) from the Pleistocene of South Australia | url=http://eurekamag.com/research/001/840/001840568.php | journal=Transactions of the Royal Society of South Australia | volume=1111–2 | pages=93–97 | url-status=live | archive-url=https://web.archive.org/web/20150701102937/http://eurekamag.com/research/001/840/001840568.php | archive-date=1 July 2015}}</ref>

===Coevolution===
[[File:Amegilla on long tube of Acanthus ilicifolius flower.jpg|thumb|Long-tongued bees and long-tubed flowers [[Coevolution|coevolved]], like this ''[[Amegilla ]]'' species (Apidae) on ''[[Acanthus ilicifolius]]''.]]
{{further|Coevolution}}

The earliest animal-pollinated flowers were shallow, cup-shaped blooms [[pollination|pollinated]] by insects such as [[beetle]]s, so the [[Pollination syndrome|syndrome of insect pollination]] was well established before the first appearance of bees. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of [[coevolution]], flowers developed floral rewards<ref name="Armbruster ch. 3 Evol. Plant-pollinator relationships">{{cite book |author=Armbruster, W. Scott |author-link=Evolution and ecological implications of 'specialized' pollinator rewards |editor1-last=Patiny |editor1-first=Sébastien |title=Evolution of Plant-Pollinator Relationships |date=2012 |publisher=Cambridge University Press |pages=45–67 |chapter=3}}</ref> such as [[nectar]] and longer tubes, and bees developed longer tongues to extract the nectar.<ref name=Michener1974/> Bees also developed structures known as [[scopa (biology)|scopal hairs]] and [[pollen baskets]] to collect and carry pollen. The location and type differ among and between groups of bees. Most species have scopal hairs on their hind legs or on the underside of their abdomens. Some species in the family Apidae have [[pollen baskets]] on their hind legs, while very few lack these and instead collect pollen in their crops.<ref name=Michener2000>{{cite book |title=The Bees of the World |date=2000 |publisher=Johns Hopkins University Press |isbn=0-8018--6133-0 |pages=19–25 |last1=Michener |first1=Charles D.}}</ref> The appearance of these structures drove the [[adaptive radiation]] of the [[Flowering plant|angiosperms]], and, in turn, bees themselves.<ref name=Buchmann2012>{{cite book |author1=Buchmann, Stephen L. |author2=Nabhan, Gary Paul |title=The Forgotten Pollinators |url=https://books.google.com/books?id=YWTZs5fSqb8C&pg=PA41 |year=2012 |publisher=Island Press |isbn=978-1-59726-908-7 |pages=41–42 |url-status=live |archive-url=https://web.archive.org/web/20160527145146/https://books.google.com/books?id=YWTZs5fSqb8C&pg=PA41 |archive-date=27 May 2016}}</ref> Bees [[Coevolution|coevolved]] not only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs called [[acarinaria]] that appear to provide lodgings for mites; in return, it is believed that mites eat fungi that attack pollen, so the relationship in this case may be [[mutualism (biology)|mutualistic]].<ref name=":5">{{cite journal |last1=Biani |first1=Natalia B. |last2=Mueller |first2=Ulrich G.|last3=Wcislo |first3=William T. |title=Cleaner Mites: Sanitary Mutualism in the Miniature Ecosystem of Neotropical Bee Nests |journal=The American Naturalist |date=June 2009 |volume=173 |issue=6 |pages=841–847 |doi=10.1086/598497 |pmid=19371167|bibcode=2009ANat..173..841B |url=https://repositories.lib.utexas.edu/bitstream/2152/31261/1/CleanerMites.pdf |archive-url=https://web.archive.org/web/20180328155201/https://repositories.lib.utexas.edu/bitstream/2152/31261/1/CleanerMites.pdf |archive-date=2018-03-28 |url-status=live |hdl=2152/31261 |s2cid=4845087 |hdl-access=free }}</ref><ref name="Pavel B 2007">{{cite journal|last1=Klimov |first1=Pavel B. |last2=OConnor |first2=Barry M. |last3=Knowles |first3=L. Lacey |title=Museum Specimens And Phylogenies Elucidate Ecology's Role in Coevolutionary Associations Between Mites And Their Bee Hosts |journal=Evolution |date=June 2007 |volume=61 |issue=6 |pages=1368–1379 |doi=10.1111/j.1558-5646.2007.00119.x |pmid=17542846|url=https://deepblue.lib.umich.edu/bitstream/2027.42/74970/1/j.1558-5646.2007.00119.x.pdf |archive-url=https://web.archive.org/web/20190504194936/https://deepblue.lib.umich.edu/bitstream/2027.42/74970/1/j.1558-5646.2007.00119.x.pdf |archive-date=2019-05-04 |url-status=live |hdl=2027.42/74970 |s2cid=32318137 |doi-access=free }}</ref>

===Phylogeny===
====External====
Molecular phylogeny was used by Debevic ''et al'', 2012, to demonstrate that the bees (Anthophila) arose from deep within the [[Crabronidae]] ''[[sensu lato]]'', which was thus rendered [[paraphyly|paraphyletic]]. In their study, the placement of the monogeneric [[Heterogyna|Heterogynaidae]] was uncertain. The small family [[Mellinidae]] was not included in this analysis.<ref>{{cite journal |last1=Debevec |first1=Andrew H. |last2=Cardinal |first2=Sophie |last3=Danforth |first3=Bryan N. |title=Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea |journal=Zoologica Scripta |date=2012 |volume=41 |issue=5 |pages=527–535 |doi=10.1111/j.1463-6409.2012.00549.x |s2cid=33533180 |url=http://www.danforthlab.entomology.cornell.edu/files/all/debevec_etal_2012.pdf |url-status=live |archive-url=https://web.archive.org/web/20150923212548/http://www.danforthlab.entomology.cornell.edu/files/all/debevec_etal_2012.pdf |archive-date=23 September 2015}}</ref>

Further studies by Sann ''et al.'', 2018, elevated the subfamilies (plus one tribe and one subtribe) of [[Crabronidae]] ''sensu lato'' to family status. They also recovered the placement of ''Heterogyna'' within [[Nyssonini]] and sunk [[Heterogynaidae]]. The newly erected family, [[Ammoplanidae]], formerly a subtribe of [[Pemphredonidae|Pemphredoninae]], was recovered as the most sister family to bees.<ref name=Sann2018>{{cite journal |first1=Manuela |last1=Sann  |first2=Oliver |last2=Niehuis |first3=Ralph S. |last3=Peters |first4=Christoph |last4=Mayer |first5=Alexey |last5=Kozlov |first6=Lars |last6=Podsiadlowski |first7=Sarah |last7=Bank |first8=Karen |last8=Meusemann |first9=Bernhard |last9=Misof |first10=Christoph |last10=Bleidorn |first11=Michael |last11=Ohl |display-authors=5 |date=2018 |title=Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. |journal=BMC Evolutionary Biology |volume= 18 |issue=71 |page=71 |doi=10.1186/s12862-018-1155-8 |doi-access=free |pmid=29776336 |pmc=5960199 |bibcode=2018BMCEE..18...71S }}</ref>

{{clade| style=font-size:85%;line-height:85%
|label1=Apoidea
|1={{clade
   |1=[[Ampulicidae]] (Cockroach wasps) [[File:Emerald Cockroach Wasp.JPG|70px]]
   |2={{clade
       |1=[[Astatidae]] [[File:Astata boopsDSC05135.JPG|70px]]
       |2={{clade
            |1={{clade
                  |1=[[Bembicinae|Bembicidae]] [[File:Bembix sp2.jpg|70px]]
                  |2={{clade
                     |1=[[Sphecidae]] (''[[sensu stricto]]'') [[File:Sceliphron spirifex TZ edit1.jpg|70px]]
                     |2={{clade 
                        |1={{clade
                           |1=[[Crabroninae|Crabronidae]] (''[[sensu stricto]]'') [[File:Ectemnius.lapidarius.-.lindsey.jpg|70px]]
                           |2=[[Mellinidae]] [[File:Mellinus arvensis - Flickr - gailhampshire (1).jpg|70px]]
                  }} }} }} }}
           |2={{clade
               |1={{clade
                  |1=[[Pemphredonidae]] [[File:Pemphredon ? (Crabronidae) - Zwerggrabwespe (11462528464).jpg|70px]]
                  |2=[[Philanthidae]] [[File:P. gibbosus57306787w.jpg|70px]]}}
               |2={{clade
                  |1=[[Psenidae]] [[File:Aphid Wasp - Flickr - treegrow (2).jpg|70px]]
                  |2={{clade 
                     |1={{clade
                        |1=[[Ammoplanidae]] 
                        |2='''Anthophila (bees)''' [[File:Abeille butineuse et son pollen.JPG|70px]] 
                  }} }} }} }} }}
         }}
      }}
   }}

====Internal====
This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae.<ref name=Hedtke2013>{{cite journal|last1=Hedtke|first1=Shannon M.|last2=Patiny|first2=Sébastien|last3=Danforth|first3=Bryan M.|title=The bee tree of life: a supermatrix approach to apoid phylogeny and biogeography|journal=BMC Evolutionary Biology|date=2013|volume=13|issue=138|pages=138|doi=10.1186/1471-2148-13-138|pmid=23822725|pmc=3706286 |doi-access=free |bibcode=2013BMCEE..13..138H }}</ref> English names, where available, are given in parentheses.

{{clade| style=font-size:85%;line-height:85%
|label1=[[Anthophila]] (bees)
|1={{clade
   |1={{clade
      |1=[[Melittidae]] (inc. [[Dasypodainae]], [[Meganomiinae]]) at least 50 Mya [[File:Macropis sp 01.jpg|70px]]
      |2={{clade
         |label1={{nowrap|long-tongued bees}}
         |1={{clade
            |1=[[Apidae]] (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya [[File:Apis mellifera flying2.jpg|70px]]
            |2=[[Megachilidae]] (mason, leafcutter bees) ≈50 Mya [[File:Leafcutter bee (Megachile sp.) collecting leaves (7519316920).jpg|70px]]
            }}
         |label2={{nowrap|short-tongued bees}}
         |2={{clade
            |1=[[Andrenidae]] (mining bees) ≈34 Mya [[File:Thomas Bresson - Hyménoptère sur une fleur de pissenlit (by).jpg|70px]]
            |2={{clade
               |1=[[Halictidae]] (sweat bees) ≈50 Mya [[File:Iridescent.green.sweat.bee1.jpg|70px]]
               |2={{clade
                  |1=[[Colletidae]] (plasterer bees) ≈25 Mya [[File:Colletes cunicularius m1.JPG|70px]]
                  |2=[[Stenotritidae]] (large Australian bees) ≈2 Mya [[File:Stenotritus pubescens, f, side, australia 2014-07-05-12.18.33 ZS PMax.jpg|70px]]
                  }}
               }}
            }}
         }}
      }}
   }}
}}