111-475: Scaptotrigona is a genus of bees belonging to the family Apidae . The species of this genus are found in Central and South America. Species: Bees Apiformes (from Latin 'apis' ) Bees are winged insects closely related to wasps and ants , known for their roles in pollination and, in the case of the best-known bee species, the western honey bee , for producing honey . Bees are
222-401: A division of labour into reproductive and non-reproductive adults, plus overlapping generations. This division of labour creates specialized groups within eusocial societies which are called castes . In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered semisocial . The group is called eusocial if, in addition,
333-577: A monophyletic lineage within the superfamily Apoidea . They are currently considered a clade , called Anthophila . There are over 20,000 known species of bees in seven recognized biological families . Some species – including honey bees , bumblebees , and stingless bees – live socially in colonies while most species (>90%) – including mason bees , carpenter bees , leafcutter bees , and sweat bees – are solitary. Bees are found on every continent except Antarctica , in every habitat on
444-663: A bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless larva , a pupation stage during which the insect undergoes complete metamorphosis , followed by the emergence of a winged adult. The number of eggs laid by a female during her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social species. Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. Like
555-429: A compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when
666-642: A complex series of orogenic events assembled the eastern parts of Gondwana (eastern Africa, Arabian-Nubian Shield, Seychelles, Madagascar, India, Sri Lanka, East Antarctica, and Australia) c. 750 to 530 Ma . First, the Arabian-Nubian Shield collided with eastern Africa (in the Kenya-Tanzania region) in the East African Orogeny c. 750 to 620 Ma . Then Australia and East Antarctica were merged with
777-599: A continuous arc chain, the direction of subduction was different between the Australian-Tasmanian and New Zealand-Antarctica arc segments. Many terranes were accreted to Eurasia during Gondwana's existence, but the Cambrian or Precambrian origin of many of these terranes remains uncertain. For example, some Palaeozoic terranes and microcontinents that now make up Central Asia, often called the "Kazakh" and "Mongolian terranes", were progressively amalgamated into
888-466: A distinct set of genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees , mason bees, plasterer bees , squash bees , dwarf carpenter bees , leafcutter bees, alkali bees and digger bees . Most solitary bees are fossorial , digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollow reeds or twigs, or holes in wood . The female typically creates
999-746: A diversified assemblage of true insects. In Gondwana, in contrast, ice and, in Australia, volcanism decimated the Devonian flora to a low-diversity seed fern flora – the pteridophytes were increasingly replaced by the gymnosperms which were to dominate until the Mid-Cretaceous. Australia, however, was still located near the Equator during the Early Carboniferous, and during this period, temnospondyl and lepospondyl amphibians and
1110-472: A few million years, reached its peak at c. 200 Ma , and coincided with the Triassic–Jurassic extinction event . The reformed Gondwanan continent was not precisely the same as that which had existed before Pangaea formed; for example, most of Florida and southern Georgia and Alabama is underlain by rocks that were originally part of Gondwana, but this region stayed attached to North America when
1221-594: A gradual decline during the Triassic while ferns, though never dominant, managed to diversify. The brief period of icehouse conditions during the Triassic–Jurassic extinction event had a dramatic impact on dinosaurs but left plants largely unaffected. The Jurassic was mostly one of hot-house conditions and, while vertebrates managed to diversify in this environment, plants have left little evidence of such development, apart from Cheiroleidiacean conifers and Caytoniales and other groups of seed ferns. In terms of biomass,
SECTION 10
#17327719520381332-525: A larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees. Most other bees, including familiar insects such as carpenter bees , leafcutter bees and mason bees are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and worker bees for these species. Solitary bees typically produce neither honey nor beeswax . Bees collect pollen to feed their young, and have
1443-440: A length of 39 millimetres (1.54 in). Bees feed on nectar and pollen , the former primarily as an energy source and the latter primarily for protein and other nutrients. Most pollen is used as food for their larvae . Vertebrate predators of bees include primates and birds such as bee-eaters ; insect predators include beewolves and dragonflies . Bee pollination is important both ecologically and commercially , and
1554-729: A male is haploid (has only one copy of each gene), his daughters (which are diploid , with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton termed "supersisters", more closely related to their sisters than they would be to their own offspring. Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of
1665-480: A much older supercontinent, Rodinia , to amalgamate. One of those orogenic belts, the Mozambique Belt , formed 800 to 650 Ma and was originally interpreted as the suture between East (India, Madagascar, Antarctica, and Australia) and West Gondwana (Africa and South America). Three orogenies were recognised during the 1990s as a result of data sets compiled on behalf of oil and mining companies:
1776-518: A north-east motion about 90 million years ago. While subduction direction changed, it remained oblique (and not perpendicular) to the coast of South America, and the direction change affected several subduction zone -parallel faults including Atacama , Domeyko and Liquiñe-Ofqui . Insular India began to collide with Asia circa 70 Ma , forming the Indian subcontinent , since which more than 1,400 km (870 mi) of crust has been absorbed by
1887-639: A number of species related to those of the laurissilva of Valdivia, through the connection of the Antarctic flora . These include gymnosperms and the deciduous species of Nothofagus , as well as the New Zealand laurel, Corynocarpus laevigatus , and Laurelia novae-zelandiae . New Caledonia and New Zealand became separated from Australia by continental drift 85 million years ago. The islands still retain plants that originated in Gondwana and spread to
1998-544: A pattern that reflects the Jurassic break-up of Pangaea. The Cretaceous saw the arrival of the angiosperms , or flowering plants, a group that probably evolved in western Gondwana (South America–Africa). From there the angiosperms diversified in two stages: the monocots and magnoliids evolved in the Early Cretaceous, followed by the hammamelid dicots . By the Mid-Cretaceous, angiosperms constituted half of
2109-521: A peak in biodiversity – the end-Permian extinction was enormous and so was the radiation that followed. Two families of conifers, Podocarpaceae and Araucariaceae , dominated Gondwana in the Early Triassic, but Dicroidium , an extinct genus of fork-leaved seed ferns, dominated woodlands and forests of Gondwana during most of the Triassic. Conifers evolved and radiated during the period, with six of eight extant families already present before
2220-853: A series of events severally restricted the Proto-ACC: change to shallow marine conditions along the North Scotia Ridge; closure of the Fuegan Seaway, the deep sea that existed in Tierra del Fuego; and uplift of the Patagonian Cordillera. This, together with the reactivated Iceland plume , contributed to global warming. During the Miocene, the Drake Passage began to widen, and as water flow between South America and
2331-694: A single jaw from Australia. The closure of the Rheic Ocean and the formation of Pangaea in the Carboniferous resulted in the rerouting of ocean currents that initiated an Ice House period. As Gondwana began to rotate clockwise, Australia shifted south to more temperate latitudes. An ice cap initially covered most of southern Africa and South America but spread to eventually cover most of the supercontinent, save for northernmost Africa-South America and eastern Australia. Giant lycopod and horsetail forests continued to evolve in tropical Laurasia together with
SECTION 20
#17327719520382442-533: A single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as Halictus hesperus . Some species are eusocial in parts of their range and solitary in others, or have a mix of eusocial and solitary nests in the same population. The orchid bees (Apidae) include some primitively eusocial species with similar biology. Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning :
2553-512: Is Cretotrigona prisca , a corbiculate bee of Late Cretaceous age (~70 mya) found in New Jersey amber . A fossil from the early Cretaceous (~100 mya), Melittosphex burmensis , was initially considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees", 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
2664-476: Is believed that mites eat fungi that attack pollen, so the relationship in this case may be mutualistic . 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 paraphyletic . In their study, the placement of the monogeneric Heterogynaidae was uncertain. The small family Mellinidae
2775-818: Is now the Transantarctic Mountains ): the Antarctic Peninsula , Marie Byrd Land , Zealandia , and Thurston Island ; the Falkland Islands and Ellsworth–Whitmore Mountains (in Antarctica) were rotated 90° in opposite directions; and South America south of the Gastre Fault (often referred to as Patagonia ) was pushed westward. The history of the Africa-Antarctica break-up can be studied in great detail in
2886-576: Is now the southern Weddell Sea where initial break-up occurred during the Jurassic c. 180 to 160 Ma . Gondwana began to break up in the early Jurassic following the extensive and fast emplacement of the Karoo-Ferrar flood basalts c. 184 Ma . Before the Karoo plume initiated rifting between Africa and Antarctica , it separated a series of smaller continental blocks from Gondwana's southern, Proto-Pacific margin (along what
2997-541: Is presumed to have been linked to the subduction of cold oceanic lithosphere . During the mid to Late Cretaceous ( c. 90 million years ago ), the Andean orogeny changed significantly in character. Warmer and younger oceanic lithosphere is believed to have started to be subducted beneath South America around this time. Such kind of subduction is held responsible not only for the intense contractional deformation that different lithologies were subject to, but also
3108-444: Is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees. Bees may be solitary or may live in various types of communities. Eusociality appears to have originated from at least three independent origins in halictid bees. The most advanced of these are species with eusocial colonies; these are characterised by cooperative brood care and
3219-555: Is thought to have occurred due to rapid climatic cooling around the 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. The oldest non-compression bee fossil
3330-787: The Antarctic Peninsula increased, the renewed ACC resulted in cooler global climate. Since the Eocene, the northward movement of the Australian Plate has resulted in an arc-continent collision with the Philippine and Caroline plates and the uplift of the New Guinea Highlands . From the Oligocene to the late Miocene, the climate in Australia, dominated by warm and humid rainforests before this collision, began to alternate between open forest and rainforest before
3441-886: The Antongil Block in far eastern Madagascar, the Seychelles , and the Napier and Rayner Complexes in East Antarctica ). The Azania continent (much of central Madagascar , the Horn of Africa and parts of Yemen and Arabia) was an island in the Mozambique Ocean. The continent Australia/ Mawson was still separated from India, eastern Africa, and Kalahari by c. 600 Ma , when most of western Gondwana had already been amalgamated. By c. 550 Ma, India had reached its Gondwanan position, which initiated
Scaptotrigona - Misplaced Pages Continue
3552-1075: The Australian Plate are now separated by the Capricorn Plate and its diffuse boundaries. During the opening of the Indian Ocean, the Kerguelen hotspot first formed the Kerguelen Plateau on the Antarctic Plate c. 118 to 95 Ma and then the Ninety East Ridge on the Indian Plate at c. 100 Ma . The Kerguelen Plateau and the Broken Ridge , the southern end of the Ninety East Ridge, are now separated by
3663-844: The Campbell Plateau , Chatham Rise , Lord Howe Rise , Norfolk Ridge , and New Caledonia , from West Antarctica c. 84 Ma . The opening of the South Atlantic Ocean divided West Gondwana (South America and Africa), but there is considerable debate over the exact timing of this break-up. Rifting propagated from south to north along Triassic–Early Jurassic lineaments, but intra-continental rifts also began to develop within both continents in Jurassic–Cretaceous sedimentary basins, subdividing each continent into three sub-plates. Rifting began c. 190 Ma at Falkland latitudes, forcing Patagonia to move relative to
3774-602: The Central Atlantic opened . Antarctica, the centre of the supercontinent, shared boundaries with all other Gondwana continents and the fragmentation of Gondwana propagated clockwise around it. The break-up was the result of the eruption of the Karoo-Ferrar igneous province , one of the Earth's most extensive large igneous provinces (LIP) c. 200 to 170 Ma , but the oldest magnetic anomalies between South America, Africa, and Antarctica are found in what
3885-834: The Drake Passage and the deepening of the Tasman Gateway. The oldest oceanic crust in the Drake Passage, however, is 34 to 29 Ma -old which indicates that the spreading between the Antarctic and South American plates began near the Eocene/Oligocene boundary. Deep sea environments in Tierra del Fuego and the North Scotia Ridge during the Eocene and Oligocene indicate a "Proto-ACC" opened during this period. Later, 26 to 14 Ma ,
3996-541: The Early Cretaceous (about 124 million years ago) on the supercontinent of West Gondwana , just prior to its breakup into South America and Africa . The supercontinent is thought to have been a largely xeric environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong niche conservatism among bees ever since their origins. Genomic analysis indicates that despite only appearing much later in
4107-598: The East African Orogeny ( 650 to 800 Ma ) and Kuunga orogeny (including the Malagasy Orogeny in southern Madagascar) ( 550 Ma ), the collision between East Gondwana and East Africa in two steps, and the Brasiliano orogeny ( 660 to 530 Ma ), the successive collision between South American and African cratons . The last stages of Gondwanan assembly overlapped with
4218-558: The East African Orogeny , the collision of India and Madagascar with East Africa, and culminating in c. 600 to 530 Ma with the overlapping Brasiliano and Kuunga orogenies, the collision of South America with Africa, and the addition of Australia and Antarctica, respectively. Eventually, Gondwana became the largest piece of continental crust of the Palaeozoic Era, covering an area of some 100,000,000 km (39,000,000 sq mi), about one-fifth of
4329-757: The Great American Interchange . The break-up of Gondwana can be said to continue in eastern Africa at the Afar Triple Junction , which separates the Arabian , Nubian , and Somali plates, resulting in rifting in the Red Sea and East African Rift . In the Early Cenozoic , Australia was still connected to Antarctica c. 35–40° south of its current location and both continents were largely unglaciated. A rift between
4440-803: The Himalayan - Tibetan orogen. During the Cenozoic, the orogen resulted in the construction of the Tibetan Plateau between the Tethyan Himalayas in the south and the Kunlun and Qilian mountains in the north. Later, South America was connected to North America via the Isthmus of Panama , cutting off a circulation of warm water and thereby making the Arctic colder, as well as allowing
4551-845: The Ordovician . This is the Cuyania or Precordillera terrane of the Famatinian orogeny in northwest Argentina which may have continued the line of the Appalachians southwards. Chilenia terrane accreted later against Cuyania. The collision of the Patagonian terrane with the southwestern Gondwanan occurred in the late Paleozoic. Subduction-related igneous rocks from beneath the North Patagonian Massif have been dated at 320–330 million years old, indicating that
Scaptotrigona - Misplaced Pages Continue
4662-565: The Paleogene (from around 66 to 23 million years ago (Ma)). Gondwana was not considered a supercontinent by the earliest definition, since the landmasses of Baltica , Laurentia , and Siberia were separated from it. To differentiate it from the Indian region of the same name (see § Name ), it is also commonly called Gondwanaland . Regions that were part of Gondwana shared floral and zoological elements that persist to
4773-481: The Pangaea supercontinent during the Carboniferous. Pangaea began to break up in the Mid-Jurassic when the Central Atlantic opened . In the western end of Pangaea, the collision between Gondwana and Laurasia closed the Rheic and Palaeo-Tethys oceans. The obliquity of this closure resulted in the docking of some northern terranes in the Marathon , Ouachita , Alleghanian , and Variscan orogenies, respectively. Southern terranes, such as Chortis and Oaxaca , on
4884-499: The Southeast Indian Ridge . Separation between Australia and East Antarctica began c. 132 Ma with seafloor spreading occurring c. 96 Ma . A shallow seaway developed over the South Tasman Rise during the Early Cenozoic and as oceanic crust started to separate the continents during the Eocene c. 35.5 Ma global ocean temperature dropped significantly. A dramatic shift from arc- to rift magmatism c. 100 Ma separated Zealandia , including New Zealand ,
4995-401: The Stenotritidae ), and by the end of the Cretaceous, South American bees had also colonized North America. 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 . Following the K-Pg extinction, surviving bee lineages continued to spread into
5106-442: The Uralian orogeny and Laurasia . Pangaea was finally amalgamated in the Late Carboniferous-Early Permian, but the oblique forces continued until Pangaea began to rift in the Triassic. In the eastern end, collisions occurred slightly later. The North China , South China , and Indochina blocks rifted from Gondwana during the middle Paleozoic and opened the Proto-Tethys Ocean . North China docked with Mongolia and Siberia during
5217-422: The Variscan orogeny close to the Carboniferous–Permian boundary. South-east Asia was made of Gondwanan and Cathaysian continental fragments that were assembled during the Mid-Palaeozoic and Cenozoic. This process can be divided into three phases of rifting along Gondwana's northern margin: first, in the Devonian, North and South China , together with Tarim and Quidam (north-western China) rifted, opening
5328-403: The uplift and erosion known to have occurred from the Late Cretaceous onward. Plate tectonic reorganisation since the mid-Cretaceous might also have been linked to the opening of the South Atlantic Ocean . Another change related to mid-Cretaceous plate tectonic rearrangement was the change of subduction direction of the oceanic lithosphere that went from having south-east motion to having
5439-446: The Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the IUCN Red List criteria. There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family Halictidae , or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have
5550-502: The Carboniferous–Permian, followed by South China. The Cimmerian blocks then rifted from Gondwana to form the Palaeo-Thethys and Neo-Tethys oceans in the Late Carboniferous, and docked with Asia during the Triassic and Jurassic. Western Pangaea began to rift while the eastern end was still being assembled. The formation of Pangaea and its mountains had a tremendous impact on global climate and sea levels, which resulted in glaciations and continent-wide sedimentation. In North America,
5661-405: The Early Cretaceous, and West Burma and Woyla during the Late Cretaceous. Gondwana's long, northern margin remained a mostly passive margin throughout the Palaeozoic. The Early Permian opening of the Neo-Tethys Ocean along this margin produced a long series of terranes, many of which were and still are being deformed in the Himalaya Orogeny . These terranes are, from Turkey to north-eastern India:
SECTION 50
#17327719520385772-403: The Earth's surface. It fused with Euramerica during the Carboniferous to form Pangea . It began to separate from northern Pangea ( Laurasia ) during the Triassic , and started to fragment during the Early Jurassic (around 180 million years ago). The final stages of break-up, involving the separation of Antarctica from South America (forming the Drake Passage ) and Australia, occurred during
5883-410: The Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale. The Halictidae first appear in the Early Eocene with species found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age. The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated by insects such as beetles , so the syndrome of insect pollination was well established before
5994-448: The Equator during this period and remained a lifeless and barren landscape. West Gondwana drifted north during the Devonian , bringing Gondwana and Laurasia close together. Global cooling contributed to the Late Devonian extinction (19% of marine families and 50% of genera went extinct) and glaciation occurred in South America. Before Pangaea had formed, terrestrial plants, such as pteridophytes , began to diversify rapidly resulting in
6105-400: The Equator on landmasses then limited to Laurasia and, in Gondwana, to Australia. In the late Silurian, two distinctive lineages, zosterophylls and rhyniophytes , had colonised the tropics. The former evolved into the lycopods that were to dominate the Gondwanan vegetation over a long period, whilst the latter evolved into horsetails and gymnosperms . Most of Gondwana was located far from
6216-446: The Jurassic flora was dominated by conifer families and other gymnosperms that had evolved during the Triassic. The Pteridophytes that had dominated during the Palaeozoic were now marginalised, except for ferns. In contrast to Laurentia, very few insect fossils have been found in Gondwana, to a considerable extent because of widespread deserts and volcanism. While plants had a cosmopolitan distribution, dinosaurs evolved and diversified in
6327-406: The Kuunga orogeny (also known as the Pinjarra orogeny). Meanwhile, on the other side of the newly forming Africa, Kalahari collided with Congo and Rio de la Plata which closed the Adamastor Ocean . c. 540–530 Ma, the closure of the Mozambique Ocean brought India next to Australia–East Antarctica, and both North and South China were in proximity to Australia. As the rest of Gondwana formed,
6438-422: The Late Permian, many known from South Africa and Australia. Beetles and cockroaches remained minor elements in this fauna. Tetrapod fossils from the Early Permian have only been found in Laurasia but they became common in Gondwana later during the Permian. The arrival of the therapsids resulted in the first plant-vertebrate-insect ecosystem. During the Mid- to Late Triassic, hot-house conditions coincided with
6549-405: The Miocene, a warm and humid climate developed with pockets of rainforests in central Australia, but before the end of the period, colder and drier climate severely reduced this rainforest. A brief period of increased rainfall in the Pliocene was followed by drier climate which favoured grassland. Since then, the fluctuation between wet interglacial periods and dry glacial periods has developed into
6660-443: The Neoproterozoic to Palaeozoic phase of the Terra Australis Orogen , a series of terranes were rafted from the proto-Andean margin when the Iapteus Ocean opened, to be added back to Gondwana during the closure of that ocean. During the Paleozoic, some blocks which helped to form parts of the Southern Cone of South America, include a piece transferred from Laurentia when the west edge of Gondwana scraped against southeast Laurentia in
6771-413: 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. A second extinction event among bees
SECTION 60
#17327719520386882-441: The Palaeo-Tethys behind them. These terranes accreted to Asia during Late Devonian and Permian. Second, in the Late Carboniferous to Early Permian, Cimmerian terranes opened Meso-Tethys Ocean; Sibumasu and Qiangtang were added to south-east Asia during Late Permian and Early Jurassic. Third, in the Late Triassic to Late Jurassic, Lhasa , West Burma , Woyla terranes opened the Neo-Tethys Ocean; Lhasa collided with Asia during
6993-456: The South Atlantic (Brazil and Cameroon ) dating to around 120 million years ago , suggesting that some form of land connection still existed between Africa and South America as recently as the early Aptian . The first phases of Andean orogeny in the Jurassic and Early Cretaceous were characterised by extensional tectonics , rifting , the development of back-arc basins and the emplacement of large batholiths . This development
7104-465: The Southern Hemisphere, has a "Gondwanan distribution" and is often described as an archaic, or relict , lineage. The distributions in the Proteaceae is, nevertheless, the result of both Gondwanan rafting and later oceanic dispersal. During the Silurian, Gondwana extended from the Equator (Australia) to the South Pole (North Africa and South America) whilst Laurasia was located on the Equator opposite to Australia. A short-lived Late Ordovician glaciation
7215-408: The Taurides in southern Turkey; the Lesser Caucasus Terrane in Georgia; the Sanand, Alborz, and Lut terranes in Iran; the Mangysglak or Kopetdag Terrane in the Caspian Sea; the Afghan Terrane; the Karakorum Terrane in northern Pakistan; and the Lhasa and Qiangtang terranes in Tibet. The Permian–Triassic widening of the Neo-Tethys pushed all these terranes across the Equator and over to Eurasia. During
7326-488: The appearance of being social. Large groups of solitary bee nests are called aggregations , to distinguish them from colonies . In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when multiple females use that same entrance regularly. The life cycle of
7437-550: The base of the Absaroka sequence coincides with the Alleghanian and Ouachita orogenies and are indicative of a large-scale change in the mode of deposition far away from the Pangaean orogenies. Ultimately, these changes contributed to the Permian–Triassic extinction event and left large deposits of hydrocarbons, coal, evaporite, and metals. The breakup of Pangaea began with the Central Atlantic magmatic province (CAMP) between South America, Africa, North America, and Europe. CAMP covered more than seven million square kilometres over
7548-789: The bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae. English names, where available, are given in parentheses. Melittidae (inc. Dasypodainae , Meganomiinae ) at least 50 Mya [REDACTED] Apidae (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya [REDACTED] Megachilidae (mason, leafcutter bees) ≈50 Mya [REDACTED] Andrenidae (mining bees) ≈34 Mya [REDACTED] Halictidae (sweat bees) ≈50 Mya [REDACTED] Colletidae (plasterer bees) ≈25 Mya [REDACTED] Stenotritidae (large Australian bees) ≈2 Mya [REDACTED] Bees differ from closely related groups such as wasps by having branched or plume-like setae (hairs), combs on
7659-617: The best known insects. Their colonies are established by swarms , consisting of a queen and several thousand workers. There are 29 subspecies of one of these species, Apis mellifera , native to Europe, the Middle East, and Africa. Africanized bees are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive. Stingless bees are also highly eusocial . They practise mass provisioning , with complex nest architecture and perennial colonies also established via swarming. Many bumblebees are eusocial, similar to
7770-414: The colonisation of Gondwana. The Baragwanathia Flora, found only in the Yea Beds of Victoria, Australia, occurs in two strata separated by 1,700 m (5,600 ft) or 30 Ma; the upper assemblage is more diverse and includes Baragwanathia, the first primitive herbaceous lycopod to evolve from the zosterophylls. During the Devonian, giant club mosses replaced the Baragwanathia Flora, introducing
7881-492: The continent Kazakhstania in the late Silurian . Whether these blocks originated on the shores of Gondwana is not known. In the Early Palaeozoic, the Armorican terrane , which today form large parts of France, was part of either Peri-Gondwana or core Gondwana; the Rheic Ocean closed in front of it and the Palaeo-Tethys Ocean opened behind it. Precambrian rocks from the Iberian Peninsula suggest that it, too, formed part of core Gondwana before its detachment as an orocline in
7992-424: The continent became the arid or semiarid landscape it is today. The adjective "Gondwanan" is in common use in biogeography when referring to patterns of distribution of living organisms, typically when the organisms are restricted to two or more of the now-discontinuous regions that were once part of Gondwana, including the Antarctic flora . For example, the plant family Proteaceae , known from all continents in
8103-401: The decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980. Human beekeeping or apiculture ( meliponiculture for stingless bees) has been practised for millennia, since at least
8214-833: The development of rifts systems on both continents, including the Central African Rift System and the Central African Shear Zone which lasted until c. 85 Ma . At Brazilian latitudes spreading is more difficult to assess because of the lack of palaeo-magnetic data, but rifting occurred in Nigeria at the Benue Trough c. 118 Ma . North of the Equator the rifting began after 120.4 Ma and continued until c. 100 to 96 Ma . Dinosaur footprints representing identical species assemblages are known from opposite sides of
8325-556: The end of it. Bennettitales and Pentoxylales , two now extinct orders of gymnospermous plants, evolved in the Late Triassic and became important in the Jurassic and Cretaceous. It is possible that gymnosperm biodiversity surpassed later angiosperm biodiversity and that the evolution of angiosperms began during the Triassic but, if so, in Laurasia rather than in Gondwana. Two Gondwanan classes, lycophytes and sphenophytes , saw
8436-946: The eruption of the Deccan basalts , whose eruption site may survive as the Réunion hotspot . The Seychelles and the Maldives are now separated by the Central Indian Ridge . During the initial break-up in the Early Jurassic a marine transgression swept over the Horn of Africa covering Triassic planation surfaces with sandstone , limestone , shale , marls and evaporites . East Gondwana, comprising Antarctica, Madagascar, India, and Australia, began to separate from Africa. East Gondwana then began to break up c. 132.5 to 96 Ma when India moved northwest from Australia-Antarctica. The Indian Plate and
8547-487: The eusocial Vespidae such as hornets in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year. In 2011, the International Union for Conservation of Nature set up
8658-563: The evolution of Voltziales , one of the few plant orders to survive the end-Permian extinction (57% of marine families and 83% of genera went extinct) and which came to dominate in the Late Permian and from whom true conifers evolved. Tall lycopods and horsetails dominated the wetlands of Gondwana in the Early Permian. Insects co-evolved with glossopterids across Gondwana and diversified with more than 200 species in 21 orders by
8769-480: The family Crabronidae , which were 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 evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps evolved from predatory ancestors. Based on phylogenetic analysis, bees are thought to have originated during
8880-530: The females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family Andrenidae ) are stingless. While solitary, females each make individual nests. Some species, such as the European mason bee Hoplitis anthocopoides , and the Dawson's Burrowing bee , Amegilla dawsoni, are gregarious, preferring to make nests near others of the same species, and giving
8991-564: The first amniote reptilians evolved, all closely related to the Laurasian fauna, but spreading ice eventually drove these animals away from Gondwana entirely. The Gondwana ice sheet melted, and sea levels dropped during the Permian and Triassic global warming. During this period, the extinct glossopterids colonised Gondwana and reached peak diversity in the Late Permian when coal-forming forests covered much of Gondwana. The period also saw
9102-605: 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 such as nectar and longer tubes, and bees developed longer tongues to extract the nectar. Bees also developed structures known as 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
9213-522: The first trees, and by the Late Devonian this first forest was accompanied by the progymnosperms , including the first large trees Archaeopteris . The Late Devonian extinction probably also resulted in osteolepiform fishes evolving into the amphibian tetrapods , the earliest land vertebrates, in Greenland and Russia. The only traces of this evolution in Gondwana are amphibian footprints and
9324-526: The flora in northeastern Australia. There is, however, no obvious connection between this spectacular angiosperm radiation and any known extinction event nor with vertebrate/insect evolution. Insect orders associated with pollination, such as beetles , flies , butterflies and moths , and wasps, bees, and ants , radiated continuously from the Permian-Triassic, long before the arrival of the angiosperms. Well-preserved insect fossils have been found in
9435-470: The forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates. Bees have the following characteristics: The largest species of bee is thought to be Wallace's giant bee Megachile pluto , whose females can attain a length of 39 millimetres (1.54 in). The smallest species may be dwarf stingless bees in
9546-519: 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,
9657-636: The fracture zones and magnetic anomalies flanking the Southwest Indian Ridge . The Madagascar block and the Mascarene Plateau , stretching from the Seychelles to Réunion , were broken off India, causing Madagascar and Insular India to be separate landmasses : elements of this break-up nearly coincide with the Cretaceous–Paleogene extinction event . The India–Madagascar–Seychelles separations appear to coincide with
9768-464: The group consists of a mother (the queen ) and her daughters ( workers ). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many paper wasps ; when the castes are morphologically discrete, the system is considered highly eusocial. True honey bees (genus Apis , of which eight species are currently recognized) are highly eusocial, and are among
9879-719: 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 plants 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
9990-861: The lake deposits of the Santana Formation in Brazil, the Koonwarra Lake fauna in Australia, and the Orapa diamond mine in Botswana. Dinosaurs continued to prosper but, as the angiosperm diversified, conifers, bennettitaleans and pentoxylaleans disappeared from Gondwana c. 115 Ma together with the specialised herbivorous ornithischians , whilst generalist browsers, such as several families of sauropodomorph Saurischia , prevailed. The Cretaceous–Paleogene extinction event killed off all dinosaurs except birds, but plant evolution in Gondwana
10101-661: The lineage as incertae sedis within the Aculeata . The Allodapini (within the Apidae) appeared around 53 Mya. The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene . The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene . The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene . The Andrenidae are known from
10212-501: The multiple (at least nine) evolutions of eusociality within Hymenoptera . Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each other's genes. But, monogamy (queens mating singly)
10323-565: The necessary adaptations to do this. However, certain wasp species such as pollen wasps have similar behaviours, and a few species of bee scavenge from carcases to feed their offspring. Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to
10434-701: The opening of the Iapetus Ocean between Laurentia and western Gondwana. During this interval, the Cambrian explosion occurred. Laurentia was docked against the western shores of a united Gondwana for a brief period near the Precambrian/Cambrian boundary, forming the short-lived and still disputed supercontinent Pannotia . The Mozambique Ocean separated the Congo – Tanzania – Bangweulu Block of central Africa from Neoproterozoic India (India,
10545-600: The other hand, remained largely unaffected by the collision along the southern shores of Laurentia. Some Peri-Gondwanan terranes, such as Yucatán and Florida , were buffered from collisions by major promontories. Other terranes, such as Carolina and Meguma , were directly involved in the collision. The final collision resulted in the Variscan- Appalachian Mountains , stretching from present-day Mexico to southern Europe. Meanwhile, Baltica collided with Siberia and Kazakhstania which resulted in
10656-473: The other members of Hymenoptera bees are haplodiploid ; the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no diapause stage. Gondwana Gondwana ( / ɡ ɒ n d ˈ w ɑː n ə / )
10767-547: The planet that contains insect-pollinated flowering plants . The most common bees in the Northern Hemisphere are the Halictidae , or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than 2 millimetres (0.08 in) long, to the leafcutter bee Megachile pluto , the largest species of bee, whose females can attain
10878-561: The present arid regime. Australia has thus experienced various climate changes over a 15-million-year period with a gradual decrease in precipitation. The Tasman Gateway between Australia and Antarctica began to open c. 40 to 30 Ma . Palaeontological evidence indicates the Antarctic Circumpolar Current (ACC) was established in the Late Oligocene c. 23 Ma with the full opening of
10989-530: The present day. The continent of Gondwana was named by the Austrian scientist Eduard Suess , after the region in central India of the same name , which is derived from Sanskrit for "forest of the Gonds ". The name had been previously used in a geological context, first by H. B. Medlicott in 1872, from which the Gondwana sedimentary sequences ( Permian - Triassic ) are also described. Some scientists prefer
11100-760: The remaining Gondwana c. 570 to 530 Ma in the Kuunga Orogeny. The later Malagasy orogeny at about 550–515 Mya affected Madagascar, eastern East Africa and southern India. In it, Neoproterozoic India collided with the already combined Azania and Congo–Tanzania–Bangweulu Block, suturing along the Mozambique Belt. The 18,000 km-long (11,000 mi) Terra Australis Orogen developed along Gondwana's western, southern, and eastern margins. Proto-Gondwanan Cambrian arc belts from this margin have been found in eastern Australia, Tasmania, New Zealand, and Antarctica. Though these belts formed
11211-665: The still static remainder of South America and Africa, and this westward movement lasted until the Early Cretaceous 126.7 Ma . From there rifting propagated northward during the Late Jurassic c. 150 Ma or Early Cretaceous c. 140 Ma most likely forcing dextral movements between sub-plates on either side. South of the Walvis Ridge and Rio Grande Rise the Paraná and Etendeka magmatics resulted in further ocean-floor spreading c. 130 to 135 Ma and
11322-399: The subduction process initiated in the early Carboniferous. This was relatively short-lived (lasting about 20 million years), and initial contact of the two landmasses occurred in the mid-Carboniferous, with broader collision during the early Permian. In the Devonian, an island arc named Chaitenia accreted to Patagonia in what is now south-central Chile. Gondwana and Laurasia formed
11433-525: The term "Gondwanaland" for the supercontinent to make a clear distinction between the region and the supercontinent. The assembly of Gondwana was a protracted process during the Neoproterozoic and Paleozoic , which remains incompletely understood because of the lack of paleo-magnetic data. Several orogenies , collectively known as the Pan-African orogeny , caused the continental fragments of
11544-492: The times of Ancient Egypt and Ancient Greece . Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the Northern Hemisphere where beekeeping is far more common. In Mesoamerica , the Mayans have practiced large-scale intensive meliponiculture since pre-Columbian times. The immediate ancestors of bees were stinging wasps in
11655-604: The tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length. According to inclusive fitness theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when Cost < Relatedness * Benefit . The requirements for eusociality are more easily fulfilled by haplodiploid species such as bees because of their unusual relatedness structure. In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because
11766-636: The two developed but remained an embayment until the Eocene-Oligocene boundary when the Circumpolar Current developed and the glaciation of Antarctica began. Australia was warm and wet during the Palaeocene and dominated by rainforest. The opening of the Tasman Gateway at the Eocene-Oligocene boundary ( 33 Ma ) resulted in abrupt cooling but the Oligocene became a period of high rainfall with swamps in southeast Australia. During
11877-501: 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. The appearance of these structures drove the adaptive radiation of the angiosperms , and, in turn, bees themselves. Bees 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
11988-499: Was a large landmass, sometimes referred to as a supercontinent . The remnants of Gondwana make up around two-thirds of today's continental area, including South America , Africa , Antarctica , Australia , Zealandia , Arabia , and the Indian Subcontinent . Gondwana was formed by the accretion of several cratons (large stable blocks of the Earth's crust), beginning c. 800 to 650 Ma with
12099-483: Was followed by a Silurian Hot House period. The End-Ordovician extinction , which resulted in 27% of marine invertebrate families and 57% of genera going extinct, occurred during this shift from Ice House to Hot House. By the end of the Ordovician, Cooksonia , a slender, ground-covering plant, became the first known vascular plant to establish itself on land. This first colonisation occurred exclusively around
12210-540: Was hardly affected. Gondwanatheria is an extinct group of non- therian mammals with a Gondwanan distribution (South America, Africa, Madagascar, India, Zealandia and Antarctica) during the Late Cretaceous and Palaeogene. Xenarthra and Afrotheria , two placental clades, are of Gondwanan origin and probably began to evolve separately c. 105 Ma when Africa and South America separated. The laurel forests of Australia, New Caledonia, and New Zealand have
12321-895: Was not included in this analysis. 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 Pemphredoninae , was recovered as the most sister family to bees. Ampulicidae (Cockroach wasps) [REDACTED] Astatidae [REDACTED] Bembicidae [REDACTED] Sphecidae ( sensu stricto ) [REDACTED] Crabronidae ( sensu stricto ) [REDACTED] Mellinidae [REDACTED] Pemphredonidae [REDACTED] Philanthidae [REDACTED] Psenidae [REDACTED] Ammoplanidae Anthophila (bees) [REDACTED] This cladogram of
#37962