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SUFFOLK HYMENOPTERISTS AND THEIR ASSOCIATES ADRIAN KNOWLES Abstract This paper summarises the key Suffolk naturalists who have contributed to our understanding of the Hymenoptera of the county, with additional references to their associates, many of whom have been commemorated in the scientific names of their discoveries. The county has, over the years, been visited by many other distinguished hymenopterists who have contributed to our knowledge of the county’s fauna, but these people have not been included here. The biographies below are listed in order of birth year.

Nicholas Gwyn (1710-1797) Originally from Fakenham in Norfolk, Nicholas Gwyn studied under the notable philosopher Herman Boerhaave at the University of Leiden in The Netherlands (Anon.

1941). Later in life he settled in Tacket Street, Ipswich. Such was his bond with the young William Kirby (see below) with whom he had long studied botany, that it is noted he would, as a 75-year-old, walk the five miles to the Barham rectory to visit his friend. Around this time, he introduced Kirby to the study of entomology. This may be his main legacy, a not insubstantial one in hindsight, for little else appears to be known of his studies or work.

In naming ‘Melitta gwynana’ (now Andrena bicolor) after him, Kirby states “Memoriae botanici periti, tum et naturae, scrutatoris indefessi, mihi et omnibus Historiae Naturalis cultoribus semper amicissimum se praebentis, Nicolai Gwyn, M.D. Gippovicensis, hoc insectum dicatum volo”. This dedication honours the botanical knowledge and friendship bestowed upon Kirby by Gwyn, noting he would freely and widely share his natural history knowledge.

Rev. James Coyte (1749-1812) James Coyte was, for 27 years, minister at St Nicholas’ Church, Ipswich, though he also spent time in other Suffolk villages, including Polstead and Farnham. His brother was the botanist Dr William Beeston Coyte, whose renowned botanical garden only survives in the name of a street in Ipswich (Coytes Gardens, close to the Buttermarket). William Kirby named Andrena coitana (as Melitta Coitana) in his honour.

Kirby states, “This has been taken only by my ingenious friend the Rev. James Coyte, of Ipswich, by whose name I have called it...and whose cabinet contains many other very rare insects”. This claim of uniqueness was, in fact, an error of judgement by Kirby, as explained in his entry, below.

One such rare insect collected by Coyte may well have been the parasitic wasp Evania appendigaster (Evaniidae). The ‘Victoria County History of Suffolk’ accords to James Coyte the honour of having been the first person to collect it in Britain, noting, “it is very rare, and even now regarded as being hardly permanently established in our Islands”. Indeed, it is thought to be an Asiatic species, no doubt spread by trade alongside its preferred host: cockroaches. This discovery warranted observation in Donovan’s “The Natural History of British Insects” (Donovan, 1801), “It is not more Trans. Suffolk Nat. Soc. 56 (2020)

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remarkable for its singularity than rarity, for we have ventured to introduce it as an English insect on one authority only. A specimen of it was lately taken by the Rev. James Coyte of Ipswich, in Suffolk”.

Rev. William Kirby (1759-1850) Much has been written about William Kirby, including a summary of his works, influences and legacy provided by Armstrong (2015). Born in Witnesham on 19 September 1759, he died some 91 years later just two and a half miles away in Barham, on 4 July 1850. After studying at Cambridge University, he was ordained into the church in 1781. Subsequently spending some 68 years in the church, first as curate and then rector, he had plenty of time to get to know his parish and its wildlife, when not attending to the needs of his parishioners.

Here, we will dwell mainly upon his immediate work with Hymenoptera. He was in one sense a very local man, but he networked with the very highest scientific minds of his time. He has rightly been dubbed the ‘Father of Entomology’ although he did not turn his attention to the study of insects until he was 26, under the guidance of Nicholas Gwyn (see above). By the age of 43, in 1802, he produced one of the most important early works on British Hymenoptera: his catchily entitled “Monographia Apum Angliæ; or, An attempt to divide into their natural genera and families, such species of the Linnean genus Apis as have been discovered in England; with descriptions and observations. To which are prefixed some introductory remarks upon the class Hymenoptera, and a synoptical table of the nomenclature of the external parts of these insects.”

In this two-volume tome, he describes all the species that had come to his attention, assigning them names as he saw fit. It is perhaps a measure of the importance of this work that some 61 (22%) out of the now 278 species of bee known in Britain still bear his name as the authority who first described the species to the scientific community. As was the style at the time, these scientific descriptions provided by Kirby are given in Latin, which makes them rather hard work for us mere mortals to appreciate. For those wishing to consult this magnum opus, the two volumes can be downloaded as pdf files from internet library resources.

These 61 species first described by Kirby are listed in Appendix 1. For those unfamiliar with the niceties of taxonomic nomenclature, the reason his name now usually appears in parentheses is because he described the species under a different name: placing everything in the genus Melitta or Apis. The one exception is, therefore, Melitta tricincta, which has borne his name, unaltered, for over 200 years.

As is apparent from other biographies in this paper, and by reference to Appendix 1, Kirby was fond of acknowledging the assistance or scientific contribution of his peers or others close to him by naming species after them. In this respect, the bee now known as Andrena trimmerana has a rather convoluted story behind it. Kirby does not acknowledge the source of this name in his Monographia. However, he had a cousin (Sarah Kirby) who married a James Trimmer of Ealing, Middlesex. Kirby acknowledges the assistance of James in the Preface to his Monographia: “To the industry, and accuracy of observation of his ingenious relation Mr. James Trimmer, he

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owes the discovery of several nondescripts…” making it a distinct possibility that it was after James that the bee, one of these ‘nondescripts’, was named. However, one of Sarah and James’ grandchildren was named Kirby Trimmer, later to become a Reverend and author of the first Flora of Norfolk. He came to serve the parish at St Georges, Tombland, Norwich and, coincidentally or not, this was the same parish previously served by William Kirby’s brother-in-law, the Rev. Charles Sutton. It is also possible that the bee was named in honour of Kirby Trimmer, but certainly it is a ‘nod’ to this branch of Kirby’s extended family. One of Kirby Trimmer’s uncles, William Kirby Trimmer was also something of a natural historian, unearthing and studying fossils from the family brickworks clay pit in London.

Some of Kirby’s tributes appear to be merely honouring his friends, with no obvious common thread of interest in the Hymenoptera. Hence the Rev. Revett Sheppard is remembered in Kirby’s Apis (now Nomada) sheppardana.

Kirby did not just describe the commonplace. It must be viewed as a considerable act of chance that the only known British specimens of what we now call Lasioglossum laeve fell under his gaze. In the Monographia he notes “Barhamiae semel capta, iterum in Nacton, Suffolciae” which translates as ‘taken once at Barham and Nacton, Suffolk’. These two specimens, both females, are the only known British specimens of this species and reside in Kirby’s collection in the Natural History Museum, London (Else & Edwards, 2018). Further fortune lay in his discovery of Andrena nana at Barham, one of only five specimens ever recorded in Great Britain.

Having now not been met with for over 80 years, it is presumed extinct in the UK. Perusal of the species list in Appendix 1 gives an interesting insight into the fauna of East Anglia and how it has changed. A look at the current UK distribution map for the bee Andrena coitana shows it to have a distinctly western and northern bias, so one might not have expected someone in deepest Suffolk to be the first to describe it.

Kirby was not, however, the collector of the specimen he described under this name.

That honour went to the Rev. James Coyte of Ipswich (see above). However, Kirby was flawed in suggesting that Coyte alone had taken a specimen of this species. Kirby also described “Melitta Shawella” from a specimen (a female) taken at Barham, not realising that this, too, was Andrena coitana, the males of which differ in having distinct ivory-white facial markings, leading him to describe it as a separate species (Coitana).

Finally, Kirby would no doubt have taken quiet pride in being proved correct concerning his description of “Apis subcornuta”, although he would have been truly amazed by the methods used to prove him right. For many decades after his description, this insect languished as a sub-species of Nomada fulvicornis, but recent DNA analysis work has determined that it is sufficiently distinct to be considered a true species: now Nomada subcornuta.

Rev. Peter Lathbury (1760-1820) Peter Lathbury was educated at the free grammar school, Ipswich, and then Clare Hall, Cambridge. After being ordained, he officiated at Campsey Ash, Butley, Woodbridge, Nacton, Levington and finally Great Livermere, where he is buried. He

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also practised medicine and was highly regarded as a physician. His obituary (Anon, 1820) notes that, “He was very fond of Natural History, and had collections both of insects, shells, and in particular fossils.” He eventually became a fellow of the Linnean Society.

He knew well Sarah Trimmer (nee Kirby, William Kirby’s cousin) having put much effort into the education of poor children, which was also her vocation. It is perhaps inevitable, then, that he struck up a friendship with William Kirby; the Rev. Lathbury was noted as one of the key Suffolk entomologists who aided Kirby in his studies. In the Preface to Kirby’s Monographia, he notes, “To the Rev. Peter Lathbury, he is indebted for much assistance every way.” In honour of this, Kirby named “Apis lathburiana” after him. Indeed, it appears that the Rev. Lathbury may have caught the specimen near Woodbridge. This is another interesting record and observation from Suffolk. Now known as Nomada lathburiana, this bee is a ‘cuckoo’ or cleptoparasite in the nests of Andrena cineraria. In recent times this Andrena has been a rare insect in eastern England, though more common and widespread in the west, so a species less likely to be encountered by East Anglian entomologists at that time, if this distribution pattern held true back then. In his Monographia, Kirby interestingly observes that Andrena cineraria was “not rare near London” but that Lathbury had once taken a male in Suffolk.

Rev. John Stevens Henslow (1796-1861) John Stevens Henslow was born in Rochester, Kent but spent the early part of his adult life at Cambridge, where he studied botany and mineralogy. In 1833 he was appointed vicar of a parish in Berkshire but remained at Cambridge other than during the university vacations. However, in 1839 he moved to Hitcham in Suffolk to where he had been appointed rector two years earlier.

Whilst known primarily as a botanist, he studied many aspects of natural history.

Morley (1899) writes “Henslow was a great Naturalist and a good Entomologist. He paid considerable attention to the Social Wasps and Bees of his parish; very few of his investigations into this subject have, however, we regret to say, descended to us”.

Charles John Paget (1811-1844) Sir James Paget (1814-1899) James Paget was born in Great Yarmouth, Norfolk. After a career in the navy was thwarted by lack of finances, he became, at the age of 16, an apprentice to a general practitioner. This started what was to become a very distinguished career in medicine, but he became an avid naturalist in his leisure time.

In 1834 he published, with his brother Charles, a summary of the flora and fauna to be found in the environs of Great Yarmouth (Paget & Paget, 1834). This includes a limited list of Hymenoptera, including from sites in the old vice county of East Suffolk.

However, with only one species each of Andrena and Halictus (which at the time embraced all species now included within Lasioglossum) this is hardly an exhaustive survey of the area, compiled by two persons only in their early 20s. Their brief list is not, however, without interest. They describe the Shrill Carder-bee (Bombus sylvarum), now extinct in Suffolk, as “common”.

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Rev. Edwin Newson Bloomfield (1827-1914) Edwin Newson Bloomfield was born in Great Glemham, Suffolk and went to Dedham Grammar School. He then studied at Cambridge for 18 years, before becoming rector at Guestling, Sussex in 1862. Given his location, it is no mean feat that he authored the “Lepidoptera of Suffolk” in 1890 and its 1900 Supplement. He was primarily a lepidopterist but was apparently no less skilled in Diptera, Hymenoptera, Neuroptera and Orthoptera. One has to wonder how much time he actually spent in Sussex, because he was also well advanced in his preparation of the manuscript for the first review of Suffolk aculeate Hymenoptera, before handing it over to Claude Morley to finish (as Morley, 1899). In his preface to this book, Morley notes that they had been intending to publish the work in the Transactions of the Norfolk and Norwich Naturalists’ Society – these were the days before the Suffolk Naturalists’ Society had been formed! It is widely held that Bloomfield was responsible for nurturing Morley’s interest in Hymenoptera.

Rev John Hocking Hocking (c.1834-1903) John Hocking was born near Penzance, Cornwall, in humble circumstances (his father being an agricultural labourer). Having entered the church in Cornwall, he then had a chaplaincy in India between 1862 and 1880. He eventually settled in Suffolk, serving the church at Copdock and then Debenham before returning to Copdock for the rest of his life. His obituary (Anon, 1904) states that “he was an enthusiastic collector of Lepidoptera, but reserved in his habits, so that his brother entomologists seldom knew much of his doings”.

He would not warrant an entry in this article, were it not for a frenetic bout of Hymenoptera collecting during 1900 and 1901. The Ipswich Museum collection includes numerous species assigned to the “Hocking/Harwood” collection, with Hocking’s material all coming from ‘Copdock’ in those two years. It is possible that he had been encouraged by Claude Morley to undertake collecting in order to update Morley’s (1899) original review of Suffolk Hymenoptera for the soon to be published volume 1 of the “Victoria County History” for Suffolk. This was obviously curtailed by Hocking’s death in 1903 and perhaps a period of ill-health before that, which prevented any further collecting beyond 1901.

It has not been possible to verify all the records attributed to Hocking, but Morley (1936) cites some very significant records. The mining bee Andrena tridentata is now considered likely to be extinct in Britain, but Hocking is accredited with a record from Copdock (one of three for Suffolk). This is a surprising record for this part of Suffolk since this species is a denizen of dry heathland and sandy grassland. Further significant records include: Andrena hattorfiana (a mining bee) – although widely recorded in Breckland, Hocking’s record from Copdock is the only one cited by Morley (1936), with only one other record currently known from vice county East Suffolk.

Coelioxys quadridentata (a cuckoo-bee) – this nationally rare bee is known from Suffolk on account of five old records in Morley (1936). Hocking’s specimen from Copdock is in the Ipswich Museum collection.

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Crossocerus capitosus (digger wasp) – one of only four Suffolk records, not seen in the county since 1996 (when recorded twice, at Elveden and Dunwich Heath).

Lasioglossum zonulum (a mining bee) – one of only three Suffolk records, Hocking recorded it at Hollesley in 1899, a rare record from outside his main location and period of activity. It went un-recorded again in Suffolk until 2003.

Nomada sexfasciata (cuckoo-bee) – the only record known to Morley (1936) was Hocking’s observation at Copdock, along with its host Eucera longicornis. I am not aware of any other Suffolk records and both this species and its host are thought to be extinct in the county.

Osmia pilicornis (a mining bee) – an undated specimen taken from near Copdock

by Hocking resides in the Ipswich Museum collection and has been verified by the author. This bee is only known from about six records from Suffolk but now thought to be extinct in the county and, indeed, is very rare nationally.

William Henry Tuck (1840-1922) William Henry Tuck was a son of the Rev William Gilbert Tuck, Rector of Tostock, near Bury St Edmunds. He had a younger brother, Julian George, who developed a keen interest in birds. Julian followed his father into the church and also served the parish of Tostock as his father had done before him. William was born in Moulton, Norfolk, but the family eventually settled in Tostock. By 1891, William was living with his widowed mother at Tostock House. In 1895 he published the first known attempt at a list of aculeates for any part of Suffolk, being the environs of Tostock (Tuck, 1895). In this paper’s introduction he notes that his labours began c.1891. During this time, he appears to have had a good working relationship with Edward Saunders and J.B. Bridgman of Norfolk, but little else appears to be known of how he came to learn the Hymenoptera. In the concluding remarks to this 1895 paper he also announces his retirement from the study of Hymenoptera, thus ending his brief reign as a pioneering Suffolk Hymenopterist!

During this short but intense period of study, he made some notable observations.

I know of no modern records for the ant Lasius umbratus away from its stronghold in the Brecks and, to a lesser extent, in the Sandlings, but he notes, “For over forty years we have had a colony in the house which emerge from under a doorstep facing west, early in September, on hot days, when all the sexes appear.” He notes the presence of the small digger wasp Passaloecus insignis, for which there are only a handful of modern records in the county. He lists, frustratingly without further comment, the rare digger wasp Crossocerus vagabundus. Despite Morley’s (1935) doubtful assertion that it was “quite common”, Tuck’s record is, to the best of my knowledge, the only precise record for this species in Suffolk. As with other sweeping generalisations of his, it is suspected that Morley’s identification was a little lazy.

Other rarities claimed by Tuck from around Tostock include Ancistrocerus antilope, Andrena coitana, Melitta leporina, Coelioxys quadridentata, Megachile circumcincta, Stelis phaeoptera and Eucera longicornis.

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Lt-Col Charles George Nurse (1862-1933) Charles’ father was the Rev. G.T. Nurse of Barnham and Euston, in west Suffolk, and it was in Barnham that Charles was born. He was educated at King Edward VI School, Bury St Edmunds, and the Royal Military College (Anon, 1934). He made his name in the army in India (1895-1909), where he also collected widely, including within Lepidoptera and Hymenoptera. Colonels Cameron and Bingham later described over 200 species of Hymenoptera from his collections, further those that Nurse had described himself. He was elected a fellow of the Entomological Society of London in 1895 and a special life fellow in 1932.

Whilst Nurse will primarily be remembered for his work on the Indian fauna, he also made an impact during his retirement back in England. By the early 1900s he was residing at Timworth Hall, near Bury St Edmunds. Whilst here, he discovered Rhopalum gracile (described as Crabro kiesenwetteri) as new to Britain, from Ampton and West Stow (Nurse, 1913). Morley (1935, 1936) quotes liberally from Nurse’s records in his review of the county’s Hymenoptera. Towards the end of his life he moved to Tunbridge Wells in Kent, where he died on November 5 1933.

Claude Morley (1874-1951) There’s no two ways about it: Claude Morley was “a character”! Much has been written about his idiosyncratic lifestyle and mannerisms (e.g. Aston 1999, 2000; Simpson 1989, 1990), but here we will restrict ourselves to his work within the Hymenoptera.

Morley was born in Blackheath, London, but appears to have moved around widely as a child, being schooled at Beccles, Peterborough and then Epsom College.

He was undoubtedly a precocious talent, so that by 19 he was being published in “The Entomologist” and elected to the Entomological Society of London at the age of 21.

At 19 he was at Ipswich Museum, largely studying geology under Dr John Ellis Taylor.

Later, he was on the staff of British Museum (Natural History), working on groups within the parasitic Hymenoptera.

His largest work was the five volume “Ichneumons of Great Britain”, published between 1903 and 1914. It has to be said that time has not been kind to Morley’s work in this respect. Two of the earliest members of the Suffolk Naturalists’ Society were George Kloet and Walter Hincks, later famous in the entomological world for their ‘Check List of British Insects” series published by the Royal Entomological Society. In their original introduction to Part 4 covering Hymenoptera they state, “it is well known that the systems adopted by Morley and Schmiedeknecht are quite out of date and, in part at least, erroneous” (from Aston, 2000). However, it must be remembered the enormity of the task Morley set himself and the debates concerning the taxonomy of this complex and large group that remain to this day. Few men would have had the calibre and knowledge to even attempt such a task.

As noted above, Edwin Bloomfield is likely to have nurtured Morley’s interest in Hymenoptera and provided much of the information that was to go into his first major publication on Suffolk aculeate Hymenoptera (Morley, 1899). This county summary underwent major revisions for the Victoria County History of 1911 and

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updates in 1935 and 1936, providing two references that I use regularly, to this day.

It almost seems like an act of heresy, then, to criticise the great man. On two accounts I would not be writing this paper were it not for him. Firstly, there might not be a Suffolk Naturalists’ Society at all to publish it and secondly, as its current County Recorder for Hymenoptera, I would not be in a position to review his work and that of his forebears. However, one does get the impression that his acclaimed talent may have got the better of him towards the end. Aston (2000) notes that, “he could, it was said by knowledgeable contemporaries, identify any British insect on sight”.

However, one wonders if anyone would have had the temerity to question one of his pronouncements in his presence! This is an excessively bold claim that would surely be beyond the scope of any one naturalist.

Yet close examination of his two major works (Morley, 1935, 1936) and scrutiny of his collection in the Ipswich Museum reveals chinks in his armour that ought not to be there. For example, in the review of bees in the 1936 Transactions he states that the mining bee Andrena florea is “abundant in spring”, bracketing this species under this comment with Andrena rosae. It is beyond reasonable doubt that he was referring to Andrena trimmerana not florea. Andrena trimmerana is very closely related to A. rosae and has variable patches of red on the anterior abdominal segments, as A. florea does. However, the national distribution map for Andrena florea shows that, until 2015, it was unknown anywhere in England much north of the river Thames, being found in the extreme south of Essex. For it to be “abundant in spring” in Suffolk in the early part of the 20th Century, but completely unrecorded elsewhere in East Anglia and the east Midlands, seems highly unlikely. There were 12 specimens under the name “florea” in the Ipswich Museum collection, comprising four different species, none of which was actually florea.

This, and other inconsistencies, inevitably make one doubt the veracity of any published record and, to that end, work is currently in progress to catalogue Morley’s collection in the museum and to capture validated records from his very numerous specimens.

References Anon. (1820). Obituary: The Christian Remembrancer, Volume 2. pp. 573-575. F.C. & J. Rivington, London.

Anon. (1904). Obituary. Entomologists’ Monthly Magazine. Second Series 15: 19.

Anon. (1934). Obituary. C.G. Nurse. The Entomologist. 67: 23-24.

Anon. (1941). News for Naturalists. Trans. Suffolk Nat. Soc. 4: 234-240.

Armstrong, P. H. (2015). William Kirby (1758-1850): Eminent Suffolk Naturalist. Trans.

Suffolk Nat. Soc. 51: 87-94.

Aston, A. (1999). “Ever thine, Claude Morley”. Trans Suffolk Nat. Soc. 35: 122-123.

Aston, A. (2000). The Fields his Study. Trans. Suffolk Nat. Soc. 36: 47-52.

Donovan, E. (1801). The Natural History of British Insects; explaining them in their several states, with the periods of their transformations, their food, economy etc.

Vol. 10. Bye & Law, London.

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Else, G. R. & Edwards, M. (2018). Handbook of the Bees of the British Isles. Two vols.

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Kirby, W. (1802). Monographia apum Angliæ; or, an attempt to divide into their natural genera and families, such species of the Linnean genus Apis as have been discovered in England: with descriptions and observations. (2 Vols.). Ipswich.

Morley, C. (1898). A list of the Hymenoptera-Aculeata of the Ipswich District. The Entomologist, 31: 12-17 and 38-41.

Morley, C. (1899). The Hymenoptera of Suffolk. Part 1 Aculeata. James H. Keys, Plymouth.

Morley, C. 1935. The Hymenoptera of Suffolk, Portio Prima. Trans. Suffolk Nat. Soc., 3: 17-52.

Morley, C. (1936) The Hymenoptera of Suffolk, Portio Secunda. Trans. Suffolk Nat. Soc. 3: 132-162.

Morley, C. (1948). Names of Suffolk Naturalists 1400-1900, with an alphabetic index.

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Nurse, C. G. (1913). Two species of aculeate Hymenoptera new to Britain.

Entomologists’ Monthly Magazine. Second Series, 24: 83-84.

Paget, C. J. & Paget, J. (1834). Sketch of the Natural History of Yarmouth and its Neighbourhood. Longman, Rees & Co., London.

Simpson, F. (1989). Claude Morley and the Suffolk Naturalists’ Society. Trans. Suffolk Nat. Soc., 25: 1-6.

Simpson, F. (1990). Memories of the Suffolk Naturalists’ Society. Trans. Suffolk Nat. Soc., 26: 1-4.

Tuck, W. H. (1895). A list of the Aculeate Hymenoptera of a Suffolk village. Trans. Nor. Nat. Soc., 6: 36 et seq.

Adrian Knowles Jessups Cottage, London Road, Capel St Mary, Suffolk, IP9 2JR

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Appendix 1 Bees of the UK Described by Kirby (1802) as New to Science Andrena angustior Andrena barbilabris Andrena bimaculata Andrena chrysosceles Andrena clarkella Andrena coitana Andrena denticulata Andrena dorsata Andrena fuscipes Andrena labialis Andrena minutula Andrena nana Andrena nigriceps Andrena nigroaenea Andrena ovatula Andrena proxima Andrena tibialis Andrena tridentata Andrena trimmerana Andrena varians Andrena wilkella Bombus barbutellus Bombus cullumanus Bombus jonellus Ceratina cyanea Chelostoma campanularum Coelioxys inermis Hylaeus dilatatus (= annularis) Lasioglossum fulvicorne Lasioglossum laeve Lasioglossum laevigatum

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Lasioglossum leucopus Lasioglossum malachurum Lasioglossum minutissimum Lasioglossum nitidiusculum Lasioglossum quadrinotatum Lasioglossum sexnotatum Lasioglossum smeathmanellum Lasioglossum villosulum Lasioglossum xanthopus Megachile circumcincta Megachile ligniseca Megachile maritima Megachile willughbiella Melitta tricincta Nomada flavoguttata Nomada flavopicta Nomada goodeniana Nomada lathburiana Nomada leucophthalma Nomada marshamella Nomada sheppardana Nomada subcornuta Osmia leaiana Osmia spinulosa (=Hoplitis spinulosa) Osmia xanthomelana Panurgus banksianus Sphecodes geoffrellus Sphecodes monilicornis Sphecodes phaeoptera Sphecodes punctulatissima