Carpenter bee
Carpenter bees are species in the genus Xylocopa of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant material such as dead wood or bamboo. The main exceptions are species in the subgenus Proxylocopa, which dig nesting tunnels in suitable soil.
Linnaeus named the type species Apis violacea in the Systema Naturae, and Latreille described the genus Xylocopa, romanized, in his Natural history, general and particular of the crustaceans and insects, to which the type species was reassigned.
Characteristics
Many species in this enormous genus are difficult to tell apart; most species are all black, or primarily black with some yellow or white pubescence. Some differ only in subtle morphological features, such as details of the male genitalia. Males of some species differ confusingly from the females, being covered in greenish-yellow fur. The confusion of species arises particularly in the common names; in India, for example, the common name for any all-black species of Xylocopa is bhanvra, and reports and sightings of bhanvra or bhomora are commonly misattributed to a European species, Xylocopa violacea; however, this species is found only in the northern regions of Jammu and Kashmir and Punjab, and most reports of bhanvra, especially elsewhere in India, refer to any of roughly 15 other common black Xylocopa species in the region, such as X. nasalis, X. tenuiscapa, or X. tranquebarorum.Non-professionals commonly confuse carpenter bees with bumblebees; the simplest rule of thumb for telling them apart is that most carpenter bees have a shiny abdomen, whereas bumblebee abdomens are completely covered with dense hair. Males of some species of carpenter bees have a white or yellow face, unlike bumblebees, while females lack the bare corbicula of bumblebees; the hind leg is entirely hairy.
The wing venation is characteristic; the marginal cell in the front wing is narrow and elongated, and its apex bends away from the costa. The front wing has small stigma. When closed, the bee's short mandibles conceal the labrum. The clypeus is flat. Males of many species have much larger eyes than the females, which relates to their mating behavior.
In the United States, two eastern species, Xylocopa virginica and X. micans, occur. Three more species are primarily western in distribution, X. sonorina, X. tabaniformis orpifex, and X. californica. X. virginica is by far the more widely distributed species.
Behavior
As a subfamily, they nest in a wide range of host plants, but any one species may show definite adaptations or preferences for particular groups of plants. Carpenter bees are traditionally considered solitary bees, though some species have simple social nests in which mothers and daughters may cohabit. Examples of this type of social nesting can be seen in the species Xylocopa sulcatipes and Xylocopa nasalis. When females cohabit, a division of labor between them occurs sometimes. In this type of nesting, multiple females either share in the foraging and nest laying, or one female does all the foraging and nest laying, while the other females guard.Solitary species differ from social species. Solitary bees tend to be gregarious and often several nests of solitary bees are near each other. In solitary nesting, the founding bee forages, builds cells, lays the eggs, and guards. Normally, only one generation of bees lives in the nest. Xylocopa pubescens is one carpenter bee species that can have both social and solitary nests.
Carpenter bees make nests by tunneling into wood, bamboo, and similar hard plant material such as peduncles, usually dead. They vibrate their bodies as they rasp their mandibles against hardwood, each nest having a single entrance which may have many adjacent tunnels. The entrance is often a perfectly circular hole measuring about on the underside of a beam, bench, or tree limb. Carpenter bees do not eat wood; they discard the bits of wood, or reuse particles to build partitions between cells. The tunnel functions as a nursery for brood and storage for the pollen/nectar upon which the brood subsists. The provision masses of some species are among the most complex in shape of any group of bees; whereas most bees fill their brood cells with a soupy mass and others form simple spheroidal pollen masses, Xylocopa species form elongated and carefully sculpted masses that have several projections which keep the bulk of the mass from coming into contact with the cell walls, sometimes resembling an irregular caltrop. The eggs are very large relative to the size of the female, and are some of the largest eggs among all insects. Carpenter bees can be timber pests, and cause substantial damage to wood if infestations go undetected for several years.
Two very different mating systems appear to be common in carpenter bees, and often this can be determined simply by examining specimens of the males of any given species. Species in which the males have large eyes are characterized by a mating system where the males either search for females by patrolling, or by hovering and waiting for passing females, which they then pursue. In the other mating system, the males often have very small heads, but a large, hypertrophied glandular reservoir in the mesosoma releases pheromones into the airstream behind the male while it flies or hovers. The pheromone advertises the presence of the male to females.
Male bees often are seen hovering near nests and will approach nearby animals. However, males are harmless, since they do not have a stinger. Female carpenter bees are capable of stinging, but they are docile and rarely sting unless caught in the hand or otherwise directly provoked.
Ecological significance
In several species, the females live alongside their own daughters or sisters, creating a small social group. They use wood bits to form partitions between the cells in the nest. A few species bore holes in wood dwellings, chewing out burrows with their robust mandibles. Since the tunnels are near the surface, structural damage is generally minor or superficial. However, carpenter bee nests are attractive to woodpeckers, which may do further damage by drilling into the wood to feed on the bees or larvae.Carpenter bees have short mouthparts and are important pollinators on some open-faced or shallow flowers; for some they even are obligate pollinators, for example the maypop and Orphium, which are not pollinated by any other insects. They also are important pollinators of flowers with various forms of lids, such as Salvia species and some members of the Fabaceae. However many carpenter bees "rob" nectar by slitting the sides of flowers with deep corollae. Xylocopa virginica is one example of a species with such nectar robbing behavior. With their short labia the bees cannot reach the nectar without piercing the long-tubed flowers; they miss contact with the anthers and perform no pollination. In some plants, this reduces fruit and seed production, while others have developed defense mechanisms against nectar robbing. When foraging for pollen from some species with tubular flowers however, the same species of carpenter bees still achieve pollination, if the anthers and stigmata are exposed together.
Many Old World carpenter bees have a special pouch-like structure on the inside of their first metasomal tergite called the acarinarium where certain mites reside as commensals or symbionts. The exact nature of the relationship is not fully understood, though in other bees that carry mites, they are beneficial, feeding either on fungi in the nest, or on other harmful mites.
Predators
Woodpeckers eat carpenter bees, being attracted to the noise of the bee larvae and drill holes along the tunnels to feed on them. Other species of birds also prey on these bees, such as shrikes and bee-eaters as well as some mammals such as ratels. Other predators include large mantises and predatory flies, particularly large robber-flies.Apart from outright predators, parasitoidal species of bee flies lay eggs in the entrance to the bee's nest and the fly maggots live off the bee larvae.
Species
Xylocopa abbotti Xylocopa abbreviata Hurd & Moure, 1963Xylocopa acutipennis Smith, 1854Xylocopa adumbrata Lieftinck, 1957Xylocopa adusta Pérez, 1901Xylocopa aeneipennis Xylocopa aerata Xylocopa aestuans Xylocopa aethiopica Pérez, 1901Xylocopa africana Xylocopa albiceps Fabricius, 1804Xylocopa albifrons Lepeletier, 1841Xylocopa albinotum Matsumura, 1926Xylocopa alternata Pérez, 1901Xylocopa alticola Xylocopa amamensis Sonan, 1934Xylocopa amauroptera Pérez, 1901Xylocopa amazonica Enderlein, 1913Xylocopa amedaei Lepeletier, 1841Xylocopa amethystina Xylocopa andica Enderlein, 1913Xylocopa angulosa Maa, 1954Xylocopa anthophoroides Smith, 1874Xylocopa apicalis Smith, 1854Xylocopa appendiculata Smith, 1852Xylocopa artifex Smith, 1874Xylocopa aruana Ritsema, 1876Xylocopa assimilis Ritsema, 1880Xylocopa atamisquensis Lucia & Abrahamovich, 2010Xylocopa augusti Lepeletier, 1841Xylocopa auripennis Lepeletier, 1841Xylocopa aurorea Friese, 1922Xylocopa aurulenta Xylocopa bakeriana Xylocopa balteata Maa, 1943Xylocopa bambusae Schrottky, 1902Xylocopa bangkaensis Friese, 1903Xylocopa barbatella Cockerell, 1931Xylocopa bariwal Maidl, 1912Xylocopa basalis Smith, 1854Xylocopa bentoni Cockerell, 1919Xylocopa bequaerti Xylocopa bhowara Maa, 1938Xylocopa biangulata Vachal, 1899Xylocopa bicarinata Alfken, 1932Xylocopa bicristata Maa, 1954Xylocopa bilineata Friese, 1914Xylocopa bimaculata Friese, 1903Xylocopa binongkona van der Vecht, 1953Xylocopa bluethgeni Dusmet y Alonso, 1924Xylocopa bombiformis Smith, 1874Xylocopa bomboides Smith, 1879Xylocopa bombylans Xylocopa boops Maidl, 1912Xylocopa bouyssoui Vachal, 1898Xylocopa brasilianorum Xylocopa braunsi Dusmet y Alonso, 1924Xylocopa bruesi Cockerell, 1914Xylocopa bryorum Xylocopa buginesica Vecht, 1953Xylocopa buruana Lieftinck, 1956Xylocopa caerulea Xylocopa caffra Xylocopa calcarata Xylocopa calens Lepeletier, 1841Xylocopa californica Cresson, 1864Xylocopa caloptera Pérez, 1901Xylocopa canaria Xylocopa cantabrita Lepeletier, 1841Xylocopa capensis Spinola, 1838Xylocopa capitata Smith, 1854Xylocopa carbonaria Smith, 1854Xylocopa caribea Lepeletier, 1841Xylocopa caspari van der Vecht, 1953Xylocopa caviventris Maidl, 1912Xylocopa cearensis Ducke, 1911Xylocopa ceballosi Dusmet y Alonso, 1924Xylocopa celebensis Xylocopa chapini Xylocopa chinensis Friese, 1911Xylocopa chiyakensis Xylocopa chlorina Xylocopa chrysopoda Schrottky, 1902Xylocopa chrysoptera Latreille, 1809Xylocopa ciliata Burmeister, 1876Xylocopa citrina Friese, 1909Xylocopa clarionensis Hurd, 1958Xylocopa claripennis Friese, 1922Xylocopa cloti Vachal, 1898Xylocopa cockerelli Maa, 1943Xylocopa codinai Dusmet y Alonso, 1924Xylocopa colona Lepeletier, 1841Xylocopa columbiensis Pérez, 1901Xylocopa combinata Ritsema, 1876Xylocopa combusta Smith, 1854Xylocopa concolorata Maa, 1938Xylocopa conradsiana Friese, 1911Xylocopa coracina van der Vecht, 1953Xylocopa cornigera Friese, 1909Xylocopa coronata Smith, 1861Xylocopa cribrata Pérez, 1901Xylocopa cubaecola Lucas, 1857Xylocopa cuernosensis Xylocopa cyanea Smith, 1874Xylocopa cyanescens Brullé, 1832Xylocopa dalbertisi Lieftinck, 1957Xylocopa dapitanensis Xylocopa darwini Cockerell, 1926Xylocopa dejeanii Lepeletier, 1841Xylocopa dibongoana Hedicke, 1923Xylocopa dimidiata Latreille, 1809Xylocopa disconota Friese, 1914Xylocopa distinguenda Pérez, 1901Xylocopa ditypa Vachal, 1898Xylocopa diversipes Smith, 1861Xylocopa dolosa Vachal, 1899Xylocopa dormeyeri Xylocopa duala Strand, 1921Xylocopa electa Smith, 1874Xylocopa elegans Hurd & Moure, 1963Xylocopa erlangeri Enderlein, 1903Xylocopa erythrina Gribodo, 1894Xylocopa escalerai Dusmet y Alonso, 1924Xylocopa esica Cameron, 1902Xylocopa euchlora Pérez, 1901Xylocopa euxantha Cockerell, 1933Xylocopa eximia Pérez, 1901Xylocopa fabriciana Moure, 1960Xylocopa fallax Maidl, 1912Xylocopa fenestrata Xylocopa fervens Lepeletier, 1841Xylocopa fimbriata Fabricius, 1804Xylocopa flavicollis Xylocopa flavifrons Matsumura, 1912Xylocopa flavonigrescens Smith, 1854Xylocopa flavorufa Xylocopa forbesii W. F. Kirby, 1883Xylocopa forsiusi Dusmet y Alonso, 1924Xylocopa fortissima Cockerell, 1930Xylocopa fransseni van der Vecht, 1953Xylocopa friesiana Maa, 1939Xylocopa frontalis Xylocopa fuliginata Pérez, 1901Xylocopa fulva Friese, 1922Xylocopa funesta Maidl, 1912Xylocopa fuscata Smith, 1854Xylocopa gabonica Xylocopa gabrielae Engel, 2001Xylocopa ganglbaueri Maidl, 1912Xylocopa gaullei Vachal, 1898Xylocopa ghilianii Gribodo, 1891Xylocopa gracilis Dusmet y Alonso, 1923Xylocopa graueri Maidl, 1912Xylocopa gressitti Lieftinck, 1957Xylocopa gribodoi Magretti, 1892Xylocopa grisescens Lepeletier, 1841Xylocopa griswoldi Mérida, Hinojosa-Díaz, & Ayala, 2022Xylocopa grossa Xylocopa grubaueri Friese, 1903Xylocopa gualanensis Cockerell, 1912Xylocopa guatemalensis Cockerell, 1912Xylocopa guigliae Lieftinck, 1957Xylocopa haefligeri Friese, 1909Xylocopa haematospila Moure, 1951Xylocopa hafizii Maa, 1938Xylocopa hellenica Spinola, 1843Xylocopa hirsutissima Maidl, 1912Xylocopa hottentotta Smith, 1854Xylocopa hyalinipennis Friese, 1922Xylocopa ignescens Xylocopa imitator Smith, 1854Xylocopa incandescens Xylocopa incerta Pérez, 1901Xylocopa incompleta Ritsema, 1880Xylocopa inconspicua Maa, 1937Xylocopa inconstans Smith, 1874Xylocopa inquirenda Vachal, 1899Xylocopa insola Vachal, 1910Xylocopa insularis Smith, 1857Xylocopa io Vachal, 1898Xylocopa iranica Maa, 1954Xylocopa iridipennis Lepeletier, 1841Xylocopa iris Xylocopa isabelleae Hurd, 1959Xylocopa javana Friese, 1914Xylocopa kamerunensis Vachal, 1899Xylocopa karnyi Maidl, 1912Xylocopa kerri Xylocopa kuehni Friese, 1903Xylocopa lachnea Moure, 1951Xylocopa lanata Smith, 1854Xylocopa langi Xylocopa lateralis Say, 1837Xylocopa lateritia Smith, 1854Xylocopa laticepsXylocopa latipes Xylocopa lautipennis Xylocopa lehmanni Friese, 1903Xylocopa lepeletieri Enderlein, 1903Xylocopa leucocephala Ritsema, 1876Xylocopa leucothoracoides Maidl, 1912Xylocopa levequeae Maa, 1943Xylocopa lieftincki Leys, 2000Xylocopa lombokensis Maidl, 1912Xylocopa longespinosa Enderlein, 1903Xylocopa longula Friese, 1922Xylocopa loripes Smith, 1874Xylocopa lucbanensis Xylocopa lucida Smith, 1874Xylocopa lugubris Gerstäcker, 1857Xylocopa lundqvisti Lieftinck, 1957Xylocopa luteola Lepeletier, 1841Xylocopa macrops Lepeletier, 1841Xylocopa madida Friese, 1925Xylocopa madurensis Friese, 1913Xylocopa maesoi Dusmet y Alonso, 1924Xylocopa magnifica Xylocopa maidli Maa, 1940Xylocopa maior Maidl, 1912Xylocopa marginella Lepeletier, 1841Xylocopa mastrucata Pérez, 1901Xylocopa maya Mérida, Hinojosa-Díaz, & Ayala, 2022Xylocopa mazarredoi Dusmet y Alonso, 1924Xylocopa mcgregori Cockerell, 1920Xylocopa mckeani Xylocopa meadewaldoi Hurd, 1959Xylocopa mendozana Enderlein, 1913Xylocopa merceti Dusmet y Alonso, 1924Xylocopa metallica Smith, 1874Xylocopa mexicanorum Cockerell, 1912Xylocopa meyeri Dusmet y Alonso, 1924Xylocopa micans Lepeletier, 1841Xylocopa micheneri Hurd, 1978Xylocopa mimetica Cockerell, 1915Xylocopa minor Maidl, 1912Xylocopa mirabilis Hurd & Moure, 1963Xylocopa mixta Radoszkowski, 1881Xylocopa modesta Smith, 1854Xylocopa mohnikei Cockerell, 1907Xylocopa montana Enderlein, 1903Xylocopa mordax Smith, 1874Xylocopa morotaiana Lieftinck, 1956Xylocopa muscaria Xylocopa myops Ritsema, 1876Xylocopa nasalis Westwood, 1842Xylocopa nasica Pérez, 1901Xylocopa nautlana Cockerell, 1904Xylocopa negligenda Maa, 1939Xylocopa nigrella Hurd, 1959Xylocopa nigrescens Friese, 1901Xylocopa nigricans Vachal, 1910Xylocopa nigricaula Xylocopa nigripes Friese, 1915Xylocopa nigrita Xylocopa nigrocaerulea Smith, 1874Xylocopa nigrocaudata Pérez, 1901Xylocopa nigrocincta Smith, 1854Xylocopa nigroclypeata Rayment, 1935Xylocopa nigroplagiata Ritsema, 1876Xylocopa nigrotarsata Maa, 1938Xylocopa nobilis Smith, 1859Xylocopa nogueirai Hurd & Moure, 1960Xylocopa nyassica Enderlein, 1903Xylocopa oblonga Smith, 1874Xylocopa obscurata Smith, 1854Xylocopa obscuritarsis Friese, 1922Xylocopa occipitalis Pérez, 1901Xylocopa ocellaris Pérez, 1901Xylocopa ocularis Pérez, 1901Xylocopa ogasawarensis Matsumura, 1932Xylocopa olivacea Xylocopa ordinaria Smith, 1874Xylocopa ornata Smith, 1874Xylocopa orthogonaspis Moure, 2003Xylocopa orthosiphonis Xylocopa pallidiscopa Hurd, 1961Xylocopa parvula Rayment, 1935Xylocopa perforator Smith, 1861Xylocopa perkinsi Cameron, 1901Xylocopa perpunctata Xylocopa peruana Pérez, 1901Xylocopa perversa Wiedemann, 1824Xylocopa pervirescens Cockerell, 1931Xylocopa phalothorax Lepeletier, 1841Xylocopa philippinensis Smith, 1854Xylocopa pilosa Friese, 1922Xylocopa plagioxantha Lieftinck, 1964Xylocopa praeusta Smith, 1854Xylocopa prashadi Maa, 1938Xylocopa preussi Enderlein, 1903Xylocopa provida Smith, 1863Xylocopa proximata Maa, 1938Xylocopa pseudoleucothorax Maidl, 1912Xylocopa pseudoviolacea Popov, 1947Xylocopa pubescens Spinola, 1838Xylocopa pulchra Smith, 1874Xylocopa punctifrons Cockerell, 1917Xylocopa punctigena Maa, 1938Xylocopa punctilabris Morawitz, 1894Xylocopa pusulata Vachal, 1910Xylocopa ramakrishnai Maa, 1938Xylocopa rejecta Vachal, 1910Xylocopa remota Maa, 1938Xylocopa rogenhoferi Friese, 1900Xylocopa romeroi Villamizar, Fernández, & Vivallo, 2020Xylocopa rotundiceps Smith, 1874Xylocopa ruficeps Friese, 1910Xylocopa ruficollis Hurd & Moure, 1963Xylocopa ruficornis Fabricius, 1804Xylocopa rufidorsum Enderlein, 1913Xylocopa rufipes Smith, 1852Xylocopa rufitarsis Lepeletier, 1841Xylocopa rutilans Lieftinck, 1957Xylocopa samarensis Xylocopa sarawatica Engel, 2017Xylocopa schoana Enderlein, 1903Xylocopa scioensis Gribodo, 1884Xylocopa senex Friese, 1909Xylocopa senior Vachal, 1899Xylocopa shelfordi Cameron, 1902Xylocopa sicheli Vachal, 1898Xylocopa signata Morawitz, 1875Xylocopa similis Smith, 1874Xylocopa simillima Smith, 1854Xylocopa sinensis Xylocopa sinensis Smith, 1854Xylocopa smithii Ritsema, 1876Xylocopa sogdiana Popov & Ponomareva, 1961Xylocopa somalica Magretti, 1895Xylocopa sonorina Smith, 1874Xylocopa sphinx Vachal, 1899Xylocopa splendidula Lepeletier, 1841Xylocopa stadelmanni Vachal, 1899Xylocopa stanleyi Xylocopa steindachneri Maidl, 1912Xylocopa strandi Dusmet y Alonso, 1924Xylocopa subcombusta Xylocopa subcyanea Pérez, 1901Xylocopa subjuncta Vachal, 1898Xylocopa subvirescens Cresson, 1879Xylocopa subvolatilis Xylocopa subzonata Moure, 1949Xylocopa sulcatipes Maa, 1970Xylocopa sulcifrons Pérez, 1901Xylocopa suspecta Moure & Camargo, 1988Xylocopa suspiciosa Vachal, 1899Xylocopa sycophanta Pérez, 1901Xylocopa tabaniformis Smith, 1854Xylocopa tacanensis Moure, 1949Xylocopa tambelanensis Xylocopa tanganyikae Strand, 1911Xylocopa tayabanica Cockerell, 1930Xylocopa tegulata Friese, 1911Xylocopa tenkeana Cockerell, 1933Xylocopa tenuata Smith, 1874Xylocopa tenuiscapa Westwood, 1840Xylocopa teredo Guilding, 1825Xylocopa tesselata Maa, 1970Xylocopa thoracica Friese, 1903Xylocopa togoensis Enderlein, 1903Xylocopa torrida Xylocopa tranquebarica Xylocopa tranquebarorum Xylocopa transitoria Pérez, 1901Xylocopa tricolor Ritsema, 1876Xylocopa trifasciata Gribodo, 1891Xylocopa trochanterica Vachal, 1910Xylocopa truxali Hurd & Moure, 1963Xylocopa tumida Friese, 1903Xylocopa tumorifera Lieftinck, 1957Xylocopa turanica Morawitz, 1875Xylocopa uclesiensis Pérez, 1901Xylocopa unicolor Smith, 1861Xylocopa ustulata Smith, 1854Xylocopa vachali Pérez, 1901Xylocopa valga Gerstäcker, 1872Xylocopa varentzowi Morawitz, 1895Xylocopa varians Smith, 1874Xylocopa varipes Smith, 1854Xylocopa velutina Lieftinck, 1957Xylocopa vestita Hurd & Moure, 1963Xylocopa villosa Friese, 1909Xylocopa violacea Xylocopa virginica Xylocopa viridigastra Lepeletier, 1841Xylocopa viridis Smith, 1854Xylocopa vittata Enderlein, 1903Xylocopa vogtiana Enderlein, 1913Xylocopa volatilis Smith, 1861Xylocopa vulpina Alfken, 1930Xylocopa waterhousei Leys, 2000Xylocopa watmoughi Eardley, 1983Xylocopa wellmani Cockerell, 1906Xylocopa wilmattae Cockerell, 1912Xylocopa xanti Mocsáry, 1883Xylocopa yunnanensis Wu, 1982Subgenus ProxylocopaXylocopa altaica Xylocopa andarabana Hedicke, 1938Xylocopa hedickeiXylocopa mongolicus Xylocopa nitidiventris Smith, 1878Xylocopa nix Xylocopa olivieri Lepeletier, 1841Xylocopa parviceps Morawitz, 1895Xylocopa przewalskyi Morawitz, 1886Xylocopa rufa Friese, 1901Xylocopa versicolor Alfken, 1930Xylocopa wuiXylocopa zonata Alfken, 1930