FNZ 57 - Apoidea (Insecta: Hymenoptera) - History of research
Donovan, BJ 2007. Apoidea (Insecta: Hymenoptera). Fauna of New Zealand 57, 295 pages.
( ISSN 0111-5383 (print), ; no. 57. ISBN 978-0-478-09389-6 (print), ). Published 07 Sep 2007
History of research on New Zealand bees
Families. The history of bee classifications world-wide is outlined by Michener (2000). Latreille (1802) recognised 2 families which became the Andrenidae and the Apidae. Of relevance to the native bees of New Zealand, the genera Colletes, Hylaeus, and Andrena were assigned to the Andrenidae. Over the next nearly 200 years there were many changes, both in numbers of names for various categories of higher names, and their status and combinations. During about the first 100 years the number of families increased, with Ashmead (1899) naming 14, and Robertson (1904) naming 18. Lepeletier (1835, 1841), under the family Merilegides, assigned the genus Halictus to the tribe Andrenites, and used the name Colletides for a tribe for the genus Colletes; whereas Hylaeus was placed in the tribe Prosopites in the family Monomorphides among the parasitic bees. Ashmead (1899) assigned Paracolletes to the family Colletidae, Prosopis = Hylaeus to the Prosopidae, and Halictus to the Andrenidae. Robertson (1904) made one change by placing Halictus in the Halictidae.
In a major work, Michener (1944) established a classification of all bees which is followed by most workers today. The genera Paracolletes and Hylaeus were placed in the Colletidae, and the genus Lasioglossum was placed in the Halictidae. However, this system has not been universally followed, and as pointed out by Michener (2000), Warnke (1977) assigned the genera Colletes and Hylaeus to the subfamily Colletinae within the Andrenidae, and the genus Halictus to the subfamily Halictinae, again within the Andrenidae. Except for the superfamily name, this present revision follows the classifications of Michener (1944, 1965, 2000).
Native and adventive bees
Genera and species. The first record of a bee collected from New Zealand appears to be the name Andrena trichopus and a drawing of a female made in manuscript by White. Publication was in Butler (1874), where under Andrenidae the name Dasycolletes metallicus is followed beneath by “Andrena trichopus White Ms., tab. 7, f. 12.”, and under this, “Dasycolletes metallicus, Smith, Cat. Hymen. Ins. I., p. 15, n.1 (1853).” The indication is that the work by White was completed before that of Smith but not published. The insects described were collected during the voyage of H.M.S. Erebus and H.M.S. Terror, supplemented with other insects collected by Dr Sinclair, Mr Earl, and others. The Erebus and the Terror were in the Bay of Islands from 17 August to 23 November 1841, and after reaching Folkstone, England, on 4 September 1843, various authors were assigned to different zoological groups. White was responsible for the insects other than Lepidoptera, and most were published in White (1846). However, financial problems arose delaying publication of White’s final part of his contribution until it appeared under the authorship of Butler in 1874 (Andrews 1986, Crosby pers. comm.).
Dr Sinclair was in the Bay of Islands at the same time as the Erebus and Terror expedition, and subsequently sent plant and animal material to the British Museum (Andrews 1986). Mr Earl arrived at Wellington on 23 September 1842 and left in October 1844. Most of his collecting seems to have taken place in the south of the South Island, and on his return to England, insects and other organisms ended up in the British Museum (Andrews 1986). Among several other collectors mentioned by Andrews from the 1840s and 1850s was Lt Col Bolton, after whom Cockerell (1904a) named Leioproctus boltoni, from female and male specimens collected by him. Bolton toured New Zealand with Governor Grey in 1847, and was stationed in Wellington between 1848 and 1850. He was then based in Auckland between March 1850 and November 1853 and commanded the Royal Engineers: this is most likely when he did his insect collecting (Dugdale 1988). He embarked for England in November 1853, and his collection reached the British Museum early in 1854. According to Andrews (1986) there were also many other, lesser known collectors during this period. The last organised collecting expedition of the era was that of the Acheron, which reached Auckland on 7 November 1848, and visited Akaroa, Lyttelton, the east coast of Otago, Cook Strait, and Tasman Bay before departing for Sydney and England. While the ship was at Akaroa and Lyttelton, Frederick Strange collected in the area and as far afield as the Torlesse Range in Canterbury. Over 100 insects from collections made by David Lyall and a few collected by Frederick Strange passed to the British Museum.
The first published descriptions of bees from New Zealand were by Frederick Smith (1853). The new genera relevant to New Zealand, Paracolletes, Leioproctus, Lamprocolletes, and Dasycolletes, were also described, to which were assigned the new species Leioproctus imitatus, Dasycolletes metallicus, and D. purpureus. A fourth species, Halictus sordidus, was also described. Smith (1854) described Prosopis laevigata from New Zealand. However, on the next page a species with the same name is recorded as being described from Russia in 1852. Subsequently, Smith’s species from New Zealand proved to be conspecific with Prosopis agilis Smith 1876.
Dasycolletes metallicus (= Leioproctus metallicus) occurs over much of New Zealand, so unfortunately no light is cast on possible collectors or collection areas. However, for L. imitatus and D. purpureus, W. F. Kirby, who was an assistant in the Zoology Department of the British Museum, names Churton as the collector (and in addition to New Zealand for both species, also gives Australia for L. imitatus), and for Prosopis laevigata and Halictus sordidus, Bolton is named as the collector and the locality is Auckland (Kirby 1881). The Reverend J. F. Churton arrived in New Zealand at Port Nicholson (Wellington) on 20 April 1840, but by 10 January 1841 was colonial chaplain in Auckland. He died in Auckland on 26 January 1853 (Blain 1999).
The next species of bee to be described as from New Zealand was Prosopis vicina Sichel (1867). The Novara expedition visited Auckland from 22 December 1858 to 8 January 1859. On board was geologist Ferdinand von Hochstetter: he left the Novara to make a rapid geological survey for the New Zealand government and collected extensively. However, this species was later found to be from Tasmania (Cameron 1902).
Hutton (1874) listed the bees from New Zealand, but incorrectly included Lamprocolletes obscurus, which Smith (1853) described from Van Diemen’s Land (= Tasmania), as well as repeating the incorrect record of Prosopis vicina.
The first descriptions for which locality data are given are those of Smith (1876) for “Hymenopterous Insects of New Zealand, collected by C. M. Wakefield, Esq., principally in the neighbourhood of Canterbury”. Prosopis agilis, P. relegatus, P. capitosus, Lamprocolletes fulvescens, and Halictus familiaris are from Canterbury, and Dasycolletes vestitus is from Wellington. All are placed in Andrenidae. Two years later, Smith (1878) described Dasycolletes hirtipes, which was collected by Prof. Hutton at Otago.
Hutton (1881) provided a brief description of Andrenidae, a key to the genera of bees in New Zealand, and a description of the genera and all species. Prosopis vicina was said to occur in both Auckland and Tasmania. Kirby (1884) also listed the bees from New Zealand, and pointed out that Lamprocolletes obscurus described by Smith (1853) is from Tasmania, not New Zealand. Cameron (1898) described P. sulcifrons and P. innocens from Greymouth, and listed P. agilis, Dasycolletes vestitus and D. hirtipes from the same locality, while Cameron (1900) described Halictus huttoni from Christchurch, and provided a key to this species and for H. sordidus and H. familiaris.
Early in the 20th century, lists of bees from New Zealand and re-descriptions and keys to some species were published by workers such as Alfken (1903), Cameron (1903), Hutton (1904), Meade-Waldo (1923), and Meyer (1931). However, the period from 1904 to 1936 was dominated by T. D. A. Cockerell, who, during this period published 16 papers that dealt to some extent, and often in detail, with the endemic bees of New Zealand. Among the most important was the unification with the genus Paracolletes Smith of the genera Leioproctus, Dasycolletes, and Lamprocolletes Smith (Cockerell 1905a). He also described 7 new species of Paracolletes, 4 of which, P. boltoni (Cockerell 1904a), P. monticola and P. hudsoni (Cockerell 1925), and P. maritimus (Cockerell 1936), are still valid specific names. The new hyaline species Prosopis maoriana (Cockerell 1909) and Hylaeus hudsoni (Cockerell 1925) and the halictine Halictus smithii var. a (Cockerell 1916a) are not valid names. The last descriptions of New Zealand bees were those published 70 years ago by Cockerell (1936).
The next major work relevant to the taxonomy of the bees of New Zealand was that of Michener (1965), who placed all the species formerly assigned to Paracolletes in a redefined genus Leioproctus, and allocated some species to a subgenus Leioproctus, and others to a newly described subgenus Nesocolletes, which was restricted to New Zealand. Also, New Zealand species previously assigned to Prosopis were placed in Hylaeus, which has priority, and all species were assigned to the subgenus Prosopisteron. In addition, species previously placed in the genus Halictus were assigned to the genus Lasioglossum and the subgenus Austrevylaeus. These classificatory arrangements are maintained in the present work.
The Australian species Euryglossina proctotrypoides was first reported to be present in New Zealand by Callan (1979) based on observations made by B. J. Donovan, and Donovan (1983b) discovered that Hyleoides concinna, also from Australia, was established. This revision records for the first time that 3 more Australian species (Hylaeus asperithorax, H. perhumilis, and Lasioglossum cognatum) are also established in New Zealand. Also, Donovan, Burnip & McCarthy (2006), and this revision, record the discovery in January and February 2006 of the wool-carder bee Anthidium manicatum, originally from Europe, at Napier, HB and Nelson, NN respectively.
The alkali bee, Nomia melanderi Cockerell, and the lucerne leafcutting bee, Megachile rotundata (F.), were imported and established during the early 1970s for the pollination of lucerne seed crops, Medicago sativa L. (Donovan 1975a). Research concentrated on maximising bee propagation, and subsequent seed yield increases were very good, but the dissolution of the Department of Scientific and Industrial Research in mid-1992 resulted in the collapse of the Government-funded promotion of both species of bees. Husbandry of alkali bees is now minimal, and lucerne leafcutting bee numbers are too few to provide an economic return to the handful of bee owners.
The red clover mason bee Osmia coerulescens (L.) was field-released in 1996 as a potentially manageable pollinator of red clover seed crops (Purves et al. 1998). However, numbers are still small, and management methods are still being developed.
The history of the introduction of bumble bees to New Zealand has been outlined by Farr (1889), Hopkins (1914), and Macfarlane & Griffin (1990). According to Hopkins (1914), the objective of introducing bumble bees was to “bring about the fertilization and seeding of red clover (Trifolium pratense)”. At the time, however, the identity of the species imported was uncertain.
As outlined by Macfarlane & Griffin (1990), from 1883 to 1885, 1,490 bumble bee queens of at least 3 species were imported from England. Most queens died during transport, but in 1883, 2 live queens were liberated near Pareora, SC, and in 1884, 2 live queens were liberated at Matamata, WO. There was apparently no establishment near Pareora, and none at Matamata. In 1885, 93 live queens were released at 2 sites near Christchurch, MC, from which Gurr (1957) believed that B. terrestris and B. ruderatus established. According to Macfarlane and Griffin (1990), this resulted in the establishment of B. terrestris, B. ruderatus and perhaps B. subterraneus. In 1906, 3 species, including B. hortorum, were imported from England. A total of 180 queens was liberated at Lincoln College, MC, of which 81 were alive, and another 145 were liberated at Tai Tapu, MC, of which 62 were alive. It seems that B. hortorum established from these liberations.
In the first few years following the establishment of bumble bees, their progeny spread over the country with remarkable rapidity. Whole nests, and queens, were sent from Canterbury to various parts of the North Island (Hopkins 1914). Gurr (1957) surveyed the South Island, and found that 4 species were extant: B. terrestris and B. ruderatus were present throughout the island, whereas B. subterraneus and B. hortorum appeared to be confined to the eastern side of the main divide south of Balcairn, NC. During the following decades bumble bees were moved to several new areas (Gurr 1972, Macfarlane & Gurr 1995). Macfarlane & Gurr (1995) reported on their observations on the identity of over 28,000 bumble bees on over 300 species of flowers throughout New Zealand, and their examination of 707 specimens from 17 insect collections. The same 4 species were present as reported by Gurr (1957), but the range of B. hortorum had greatly increased.
Honey bees were first successfully imported to New Zealand from England when 2 straw hives were taken to Mangungu in Hokianga, ND, North Island, on 18 March 1839, by Miss Mary Anna Bumby (Barrett 1996). Because the bees were from England, they would have been the so-called English ‘brown bee’, A. m. mellifera. This subspecies is also found on the European continent north of the Italian Alps, where it is often called the black bee. The first bees to reach the South Island were imported from Australia to Nelson, NN on 18 April 1842 by Dr Imlay. According to Barrett (1996), honey bees were first established in Australia when hives from England were landed at Port Jackson on 9 March 1822. Other importations to Australia, all from England, were made between then and 1842, so the first bees that were imported to the South Island were also A. m. mellifera. Barrett (1996) relates reports of a number of subsequent importations of bees to New Zealand from many areas of the world. Some introductions were of colonies, but later importations were of queens with and without attendant workers. One of the most important was the introduction of 2 colonies of the yellow Italian subspecies A. m. ligustica from Los Angeles County, California, 1 to Coromandel, CL and the other to Christchurch, MC, in September 1880. Also, more recently, some feral colonies may have reached New Zealand in shipping containers. Yanke (2005) detailed the legal importation in 1988 and 1989 of small amounts of Italian-type semen from Western Australia, and in 2004, 2 importations of semen of the Carniolan bee, A. m. carnica, from Germany and Austria.
Soon after their introduction, honey bees began to thrive, due to the temperate climate, the absence of most natural enemies, and the abundance of both native and introduced nectar- and pollen-bearing flowers. Cotton (1848) reported that a 10-fold increase in hives could be expected, and a swarm and its offspring during 4 years yielded 983 kg of honey. Swarms that escaped from managed hives colonised the environment, so that soon honey bees could be expected almost throughout the vegetated areas of the country. Today, many North Island colonies both wild and managed, are being killed by the mite Varroa destructor Anderson & Trueman, and the discovery of the mite in beehives at Nelson NN in June 2006 means that many South Island colonies will die.
Biology of native bees
Meyer (1931), with his sketch of a nest of Prosopis laevigata, was the first to illustrate a nest of a New Zealand bee. The nest had been sent to him from Nelson by E. S. Gourlay. The first substantive description of the biology of an endemic New Zealand bee was that of Raymond Hansen in Rayment (1935). Mr Hansen outlined the biology and nest structure of a species identified by Tarlton Rayment as Paracolletes boltoni (but see ‘Remarks’ in the treatment of Leioproctus paahaumaa). Specimens of both adults and immatures from Orini, Taupiri, WO, were sent by Mr Hansen to Tarlton Rayment in Australia, who illustrated larvae and pupae and parts thereof, and pollen grains from the larval food. Tarlton Rayment also includes a description by Mr E. S. Gourlay of the activity of Paracolletes metallicus at a nest site in sand dunes at Tahuna(nui) near Nelson, NN.
In 1967 I completed an MSc thesis on the bionomics of Leioproctus boltoni at Island Bay Road, Auckland, AK (Donovan unpublished). Subsequent study showed that several closely related species of bees were involved. Since then I have published a number of papers on both native and introduced bees, either alone or with co-authors. Primary amongst these, which present most details of the biology of endemic bees, are Donovan (1980) on the interactions of native and introduced bees, Donovan (1983a) on the comparative biogeography of the Apoidea of New Zealand and New Caledonia, and Donovan & Macfarlane (1984) on bees and pollination. These papers reported that on the basis of their flower visiting preferences, most Colletinae could be divided into three groups: those that visited Compositae (= Asteraceae), Leguminosae (= Fabaceae), or Myrtaceae. During this period Quinn (1984) surveyed the native bees in the Mackenzie Basin, MK, in the inland South Island. Some species were abundant, with an estimated 840,000 nests of Leioproctus fulvescens along 6.5 km of a little-used unsealed road.
Botanical view of native bees as pollinators
From the plant pollination point of view, until recently native bees were not mentioned (e.g., Cheesman 1873, 1875, 1877, 1878, 1880, Thomson 1879, 1881a), or were only occasionally mentioned (e.g., Thomson 1881b, 1927, Heine 1937, Garnock-Jones 1976, Godley 1979, Primack 1979, Bocher & Philipp 1985, Webb 1985). However, Primack (1978, 1983) reported tremendous numbers of Lasioglossum sordidum foraging on Discaria flowers, great numbers of Leioproctus and Hylaeus on flowering manuka, and abundant native bees on a wide range of mountain flora, and Webb (1994) found that Lasioglossum sordidum appeared to be an effective pollinator of Corokia contoneaster. Dugdale (1975) reported an observation by Dr E. J. Godley of small bees prising open the appressed corolla lobes of Loranthaceae, and of his own observations of similar bees doing the same thing to unexpanded flowers on racemes of Hebe gracillima. Godley (1979) identified the Loranthaceae as Elytranthe flavida (now Alepis flavida) and the bees as Prosopis agilis. Kelly et al. (1996) found the same bee (as Hylaeus agilis) opening flowers of Peraxilla tetrapetala. They further found that fruit set was increased above that in unopened buds. The female Leioproctus photographed by Robertson et al. (2005) on a P. tetrapetala flower which it has just partially opened is probably L. pango.