Leioproctus

{{Automatic taxobox

| image = Leioproctus-fulvescens.jpg

| image_caption = Leioproctus fulvescens, Otago, New Zealand

| taxon = Leioproctus

| authority = F. Smith, 1853

| type_species_authority = Smith, 1853 {{cite web |url=http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/Leioproctus_%28Leioproctus%29 |title=Subgenus Leioproctus (Leioproctus) Smith, 1853 |work=Australian Faunal Directory |publisher=Department of the Environment, Water, Heritage and the Arts |date=March 29, 2010 |accessdate=April 7, 2010}}

}}

Leioproctus, the hairy colletid bee,{{Cite book |last=Crowe |first=Andrew |title=Which New Zealand insect? with over 650 life-size photos of New Zealand insects |date=2002 |publisher=Penguin |isbn=978-0-14-100636-9 |location=Auckland, N.Z}} is a genus in the plaster bee family Colletidae. Its members are primarily found in Australasia and temperate South America,{{cite journal |author=Lawrence Packer |year=2006 |journal=Zootaxa |volume=1104 |pages=47–57 |title=A new Leioproctus with unique wing venation in males (Hymenoptera: Colletidae: Paracolletinae) with comments on unusual wing modifications in bees |doi=10.11646/zootaxa.1104.1.4 |url=http://www.mapress.com/zootaxa/2006f/zt01104p057.pdf }} and include the most common native bees in New Zealand.{{cite encyclopedia |url=http://www.teara.govt.nz/en/wasps-and-bees/4 |author=John Early |title=Wasps and bees - Native bees |encyclopedia=Te Ara: The Encyclopedia of New Zealand |date=March 1, 2009 |accessdate=April 7, 2010 |publisher=Ministry for Culture and Heritage / Te Manatū Taonga}}

File:Female Native bee on Haemodorum plant.jpg species in Western Australia]]

Description

Species within the genus Leioproctus are small, black, hairy bees ranging from 4 –16 mm. Most are less than 10mm, but the largest species Leioproctus Muelleri reach up to 16mm.{{Cite journal |last1=Batley |first1=Michael |last2=Popic |first2=Tony J. |date=2016-11-30 |title=An unusual new Leioproctus species (Hymenoptera: Colletidae) |url=https://journals.australian.museum/batley-2016-rec-aust-mus-686-263268/ |journal=Records of the Australian Museum |volume=68 |issue=6 |pages=263–268 |doi=10.3853/j.2201-4349.68.2016.1659}} The legs and thorax are covered in hairs ranging from black to red to yellow to white, although hair colour typically fades with age. The dorsal surface of the abdomen is mostly hairless showing the shiny black cuticle underneath. The clypeus and supraclypeus are typically hairy while the forehead is often mostly hairless, likely to avoid interference with the ocelli. Supraclypeus is almost completely flat and is densely punctured throughout.{{Cite journal |last=Hart |first=Ngaire Hiria |date=2007-11-01 |title=Industrious Native Bees: A Case Study in Whangarei |url=https://www.gefree.org.nz/assets/Uploads/Industrious-Native-Bees-A-Case-Study-.pdf |journal=The University of Auckland |via=gefree}}

Females can be distinguished from pollen clumps, carried externally on scopae.{{Cite journal |last=Hart |first=Ngaire Hiria |date=2007-11-01 |title=Industrious Native Bees: A Case Study in Whangarei |url=https://www.gefree.org.nz/assets/Uploads/Industrious-Native-Bees-A-Case-Study-.pdf |journal=The University of Auckland |via=gefree}}

Ecology

= General diet =

All their nutrients come directly or indirectly from pollen and nectar. Pollen is the main source of protein, required for growth, so it is consumed mostly during the larval stage, while nectar provides sugar, which provides the energy adults and larvae need throughout their life.{{Cite journal |last1=Villanueva |first1=M.T. Orzáez |last2=Marquina |first2=A. Díaz |last3=Serrano |first3=R. Bravo |last4=Abellán |first4=G. Blazquez |date=January 2002 |title=The importance of bee-collected pollen in the diet: a study of its composition |url=http://www.tandfonline.com/doi/full/10.1080/09637480220132832 |journal=International Journal of Food Sciences and Nutrition |language=en |volume=53 |issue=3 |pages=217–224 |doi=10.1080/09637480220132832 |pmid=11951585 |issn=0963-7486}}

Most species show medium degrees of specialisation for native flowering plants and trees. However, they multible species in New Zealand have been recorded visiting kiwifruit flowers,{{Cite journal |last=Hart |first=Ngaire Hiria |date=2007-11-01 |title=Industrious Native Bees: A Case Study in Whangarei |url=https://www.gefree.org.nz/assets/Uploads/Industrious-Native-Bees-A-Case-Study-.pdf |journal=The University of Auckland |via=gefree}} and agricultural studies have caught them in clover pastures and orchards.{{Cite journal |last1=Malone |first1=Louise |last2=Aulsford |first2=James |last3=Howlett |first3=Brad |last4=Scott-Dupree |first4=Cynthia |last5=Bardol |first5=Nicolas |last6=Donovan |first6=Barry |date=January 2010 |title=Observations on bee species visiting white clover in New Zealand pastures |url=https://www.tandfonline.com/doi/full/10.3896/IBRA.1.49.3.09 |journal=Journal of Apicultural Research |language=en |volume=49 |issue=3 |pages=284–286 |doi=10.3896/IBRA.1.49.3.09 |bibcode=2010JApiR..49..284M |hdl=10214/2634 |issn=0021-8839|hdl-access=free }} This plasticity varies between species as some studies have shown to be more willing to feed from introduced species than others.{{Cite journal |last=Donovan |first=B. J. |date=1980 |title=Interactions between native and introduced bees in New Zealand. |url=https://www.jstor.org/stable/pdf/24052010.pdf |journal=New Zealand Journal of Ecology |volume=3 |pages=104–116|jstor=24052010 }}

= Foraging behaviour =

Leioproctus species show almost no territorial behaviours, and often nest and feed within close proximity of each other. They can show a range of distance and foraging time between individuals as well as species. Although a common theme to all species is that they typically nest close to a large food source (<10m) such as a large Leptospermum tree (Mānuka) which are common in New Zealand and east Australia.{{Cite journal |last=Batley |first=Michael |date=2020 |title=A TALE OF TWO SITES |url=https://resources.austplants.com.au/wp-content/uploads/Tale-of-Two-Sites.pdf |journal=The Cross Pollinator |issue=12}}

= Pollination =

Most species are somewhat specialised often to a particular genus or family, which is beneficial to plants as specialised pollinators increases the likelihood of pollen being spread to the correct species.{{Cite journal |last1=de Araújo |first1=L. D. A. |last2=Quirino |first2=Z. G. M. |last3=Machado |first3=I. C. |date=September 2014 |editor-last=Dafni |editor-first=A. |title=High specialisation in the pollination system of M andevilla tenuifolia (J.C. M ikan) W oodson ( A pocynaceae) drives the effectiveness of butterflies as pollinators |url=https://onlinelibrary.wiley.com/doi/10.1111/plb.12152 |journal=Plant Biology |language=en |volume=16 |issue=5 |pages=947–955 |doi=10.1111/plb.12152 |pmid=24628969 |bibcode=2014PlBio..16..947D |issn=1435-8603}}

The males are relatively short lived and have little impact on pollination, but the females intentionally collect pollen by accumulating pollen in the hairs around the head and thorax while feeding on nectar before moving it to the scopae. This morphology allows pollen to be more effectively spread between flowers than other potential pollinators such as other insects and birds.{{Cite journal |last1=Goulnik |first1=Jérémie |last2=Plantureux |first2=Sylvain |last3=Van Reeth |first3=Colin |last4=Baude |first4=Mathilde |last5=Mesbahi |first5=Geoffrey |last6=Michelot-Antalik |first6=Alice |date=December 2020 |title=Facial area and hairiness of pollinators visiting semi-natural grassland wild plants predict their facial pollen load |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/een.12913 |journal=Ecological Entomology |language=en |volume=45 |issue=6 |pages=1296–1306 |doi=10.1111/een.12913 |bibcode=2020EcoEn..45.1296G |issn=0307-6946}}

Leioproctus are particularly important pollinators for many endangered native plants, such as rare Proteas in Australia, many of which are strictly self-incompatible and rely on pollinators for survival. Multiple studies on Persoonia (geebungs) found that Leiproctus was the most common genus of pollinators at their study sites.{{Cite journal |last1=Bernhardt |first1=Peter |last2=Weston |first2=Peter |date=1996-07-01 |title=The pollination ecology of Persoonia (Proteaceae) in eastern Australia |url=http://plantnet.rbgsyd.nsw.gov.au/emuwebnswlive/objects/common/webmedia.php?irn=75863&reftable=ebibliography |journal=Telopea |volume=6 |issue=4 |pages=775–804 |doi=10.7751/telopea19963035|bibcode=1996Telop...6..775B }}{{Cite journal |last1=Emery |first1=Nathan J. |last2=Offord |first2=Catherine A. |date=2021-12-07 |title=Reproductive biology of the threatened and highly fragmented shrub Persoonia hirsuta (Proteaceae) |url=https://www.publish.csiro.au/BT/BT21068 |journal=Australian Journal of Botany |language=en |volume=70 |issue=1 |pages=56–62 |doi=10.1071/BT21068 |issn=1444-9862|doi-access=free }} Lots of these species live in dry woodlands on acidic sandy soils that are suited to Leioproctus as they are mining bees and can tolerate these harsh environments.

In New Zealand, Leioproctus are the largest genus of bees and are the next most common after Apis and Bombus. Both these genera are more efficient than Leioproctus at pollinating but they prefer introduced species over natives, making Leioproctus and other native bees still vital to native forests.{{Cite journal |last1=Newstrom |first1=Linda |last2=Robertson |first2=Alastair |date=January 2005 |title=Progress in understanding pollination systems in New Zealand |url=http://www.tandfonline.com/doi/abs/10.1080/0028825X.2005.9512943 |journal=New Zealand Journal of Botany |language=en |volume=43 |issue=1 |pages=1–59 |doi=10.1080/0028825X.2005.9512943 |bibcode=2005NZJB...43....1N |issn=0028-825X}} Research has gone into using native bees for commercial pollination and found positive results and successful trials of nest removal and re-establishment.{{Cite journal |last1=Donovan |first1=Barry J. |last2=Howlett |first2=Bradley G. |last3=Walker |first3=Melanie K. |date=February 2010 |title=Relocation and establishment of nesting populations of the native bee Leioproctus huakiwi Donovan (Hymenoptera: Colletidae). |url=http://www.tandfonline.com/doi/abs/10.1080/00779962.2010.9722198 |journal=New Zealand Entomologist |language=en |volume=33 |issue=1 |pages=109–113 |doi=10.1080/00779962.2010.9722198 |bibcode=2010NZEnt..33..109D |issn=0077-9962}} Because they nest individually, they would be more resistant to spread of disease, like Varroa mites or AFB, which is of increasing interest in agriculture.Newstrom-Lloyd, L. E. (2013). Pollination in New Zealand. Ecosystem services in New Zealand—conditions and trends. Manaaki Whenua Press, Lincoln, 408-431.

https://oldwww.landcareresearch.co.nz/__data/assets/pdf_file/0008/77057/2_11_Newstrom.pdf

= Impact from introduced honeybees =

The effect introduced bees have had on native plants and bees is still unclear, but some ecologists have concerns about the effect being negative.

Introduced honeybees have been recorded visiting many of the species previously associated with native bees. For most native plant species this is atypical and unintentional of beekeepers, but due to the high demand for Mānuka honey, Leioproctus bees are now seeing high competition in some places. Because domestic honeybees live in mobile hives, beekeepers can move these hives into high resource areas and then remove them once the nectar and pollen are depleted, forcing the sessile leioproctus bees to travel far distances to find resources. Although Leiproctus species feed from species other than Mānuka, many still rely on this food source to support them over the summer, so this increased competition could reduce leioproctus populations which could then affect the other native plants they pollinate.Fale, G. (2023). Competition between native bees and honey bees on mānuka flowers in New Zealand (Doctoral dissertation, The University of Waikato). https://researchcommons.waikato.ac.nz/server/api/core/bitstreams/cc490f66-c119-4bf6-b8f4-994eb757b934/content

Habitat

Leioproctus are solitary mining bees. They can tolerate a high range of substrate provided there is sufficient food resources nearby. This can range from forest undergrowth to roadside ditches to sandy costal banks to even currently occupied termite nests in at least one Australian species.{{Cite journal |last1=Maynard |first1=Glynn |last2=Rao |first2=Sujaya |date=January 2010 |title=The solitary bee, Leioproctus (Leioproctus) nigrofulvus (Cockerell 1914) (Hymenoptera: Colletidae), in SE Australia: Unique termite-mound-nesting behavior and impacts of bushfires on local populations |url=http://www.bioone.org/doi/abs/10.3956/2009-20.1 |journal=The Pan-Pacific Entomologist |language=en |volume=86 |issue=1 |pages=14–19 |doi=10.3956/2009-20.1 |issn=0031-0603}} Leioproctus need warm temperatures (at least 15 °C) before leaving their nest, typically waiting until the nest entrance is bathed in sun. Because of this, most nests face towards the north, or if in flat ground, will be well exposed to maximise heat absorption.

They prefer dry ground and will often dig horizontal tunnels in banks and cliffs. Their nests are not lined with water repellent coatings, as seen in other bee nests, but nest architecture shows they can also make sink traps, potentially to prevent water from entering cells, suggesting they can tolerate wet environments.{{Cite journal |last1=Rozen |first1=Jerome G. |last2=Houston |first2=Terry F. |date=2022-06-28 |title=Descriptions of the Mature Larvae of Three Australian Ground-Nesting Bees(Hymenoptera: Colletidae: Diphaglossinae and Neopasiphaeinae) |url=https://bioone.org/journals/american-museum-novitates/volume-2022/issue-3989/3989.1/Descriptions-of-the-Mature-Larvae-of-Three-Australian-Ground-Nesting/10.1206/3989.1.full |journal=American Museum Novitates |issue=3989 |doi=10.1206/3989.1 |issn=0003-0082}} This tolerance to wet conditions has been observed in multiple studies where as long as it was reasonably warm, bees would be seen, no matter the rainfall.

Life cycle

Leioproctus are solitary, meaning all every female is fertile and creates their own nest for their young.

Leioproctus overwinters as diapausing prepupae in cells before they pupate and begin appearing in spring or early summer. Males typically appear first closely followed by females. Immediately they mate and the females begin nest mining while the males continue breeding. The males soon die while the females finish their nests and start producing cells. Larvae require a lot of pollen and nectar to grow and survive to adulthood, and the females have a lot of environmental pressures, so most only make 10-30 cells in their 6-8 week adult lifetime, before perishing. After the eggs are laid, the larvae hatch around 72 hours later and consume all stored food over the next 10 days, then over the next few weeks become prepupae and diapause until the following spring.{{Cite journal |last=Hart |first=Ngaire Hiria |date=2007-11-01 |title=Industrious Native Bees: A Case Study in Whangarei |url=https://www.gefree.org.nz/assets/Uploads/Industrious-Native-Bees-A-Case-Study-.pdf |journal=The University of Auckland |via=gefree}}

New Zealand

File:Inaturalist leioproctus distribution in NZ.png

= Diet =

Native New Zealand Leioproctus species primarily forages on the flowers of native species in the Myrtaceae family, such as Pōhutukawa (Metrosideros excelsa), Kānuka (Kunzea ericoides) and Mānuka (Leptospermum scoparium). But some species also show plasticity for introduced species, like certain fruit trees and pasture plants.

= Biogeography =

At least 18 species of Leioproctus can be found across New Zealand.

Australia

File:Inaturalist leioproctus distribution in Au.png

= Diet =

Native Australian Leioproctus species forage on the flowers much wider range of native species such as Persoonia (geebungs) in Family Proteaceae and many legumes such as Daviesia and Acacia,{{Cite journal |last=Bernhardt |first=Peter |date=1987 |title=A Comparison of the Diversity, Density, and Foraging Behavior of Bees and Wasps on Australian Acacia |url=https://www.jstor.org/stable/2399260 |journal=Annals of the Missouri Botanical Garden |volume=74 |issue=1 |pages=42–50 |doi=10.2307/2399260 |jstor=2399260 |bibcode=1987AnMBG..74...42B |issn=0026-6493}} in Family Fabaceae. Mānuka and other Myrtaceae family flowering trees are also visited by some species.

= Biogeography =

More than 80 species of Leioproctus can be found across Australia.{{Cite web |author1= |date=2024 |title=GBIF Occurrence Download |url=https://www.gbif.org/occurrence/download/0021240-241007104925546 |access-date=2024-10-18 |website=www.gbif.org |language=en |doi=10.15468/dl.tdxbnb}}

South America

File: Sa leioproctus.png

= Diet =

Research and information on Leioproctus species found in South America is seriously lacking. In New Zealand and Australia Leioproctus feed primarily on species within Family Proteaceae, Fabaceae and Myrtaceae. Between these three families there are over 3,000 species in South America, so Leioproctus feeding behaviour is likely similar to New Zealand and Australian species, although this cannot currently be confirmed.{{Cite web |date=2024 |title=Observations |url=https://www.inaturalist.org/observations?place_id=97389&subview=map&taxon_id=47122&view=species |access-date=18 October 2024 |website=iNaturalist}}

= Biogeography =

More than 60 species of Leioproctus can be found across South America.{{Cite web |author1= |date=2024 |title=GBIF Occurrence Download |url=https://www.gbif.org/occurrence/download/0021136-241007104925546 |access-date=2024-10-18 |website=www.gbif.org |language=en |doi=10.15468/dl.qbrrfc}}