Is the Dark Bee Really Native to Britain and Ireland?

Is the Dark Bee really native to Britain and Ireland?

Dorian Pritchard

Introduction one another, but quite unique to Britain. This confirms my belief According to Irish mythology honeybees, wheat and rye were that the ancestors of our present native bees almost certainly first taken to Ireland all in the same boat by three Christian monks arrived in Britain prior to 8 ,500BP, seven millennia ahead of the some time after AD500. Brother Finan Cam took "the full of date suggested by Randall and Sheppard (2007). his shoe" of wheat, Brother Declan "the full of his shoe" of rye and Brother Modomnoc, under the orders of St David of Wales, Post-glacial colonization "the full of his bell" of bees (Mac Oengobann, c.805). This It is helpful to consider present-day British ecosystems from picturesque idea is extended into modern mythology by Randall the perspective of post-glacial colonization. The Last Glacial and Sheppard (2007) who claim that honeybees were brought Maximum peaked 24 - 21,000 years ago. At that time the land to mainland Britain by Christian monks in "AD500+". If their north and west of the English Midlands was covered with ice, opinion were true, honeybees would class as alien to Britain. in places two kilometres or more deep, while much of southern The concept is important, as non-native species should ideally Britain were covered in loess, while the southern shoreline was be excluded from nature reserves, whereas natives should be 60-120 metres below that at present. By —16,000BP the loess welcomed. So what is the truth? Are honeybees native or alien? and other soils had transformed through tundra to grassland And how long does any species have to reside in an area before and the climate was warming sufficiently for re-colonization by it classes as "native"? man. A species that arrived before recorded history unaided by Temperatures reached a relative high point around 14,500 man, has since survived continuously, would do so indefinitely years ago, coinciding with the Upper Late Palaeolithic human and without human intervention, and has become completely expansion into northern Europe. At that time Ireland, Orkney integrated into its environment would seem to qualify as and the British mainland were linked to the rest of Europe by "native". Perhaps lesser criteria would satisfy, but the idea that a broad plain of tundra and sparse grassland, but subsequently honeybees are incomers persists. An observation persuasive to the grassland was replaced by steppe woodland of birch, willow some biologists is that, in contrast to many other bee species, and pine. honeybees show little anatomical or behavioural adaptation to By 13,000 BP the ice sheets over northern Britain had largely specific British flowers, being adapted instead to a variety of melted, but smaller ice caps and other glaciers briefly returned pollen and nectar sources, a degree of versatility that would a thousand years later in the Younger Dryas, characterized by seem to damn them as alien everywhere. However, one adaptive abundant pollen of the Arctic-montane plant Dryas octopetala feature that is sometimes overlooked is the localised timing (mountain avens). of broodnest development in coordination with flowering of From about 11,500 BP British temperatures soared again and the major local forage. In this honeybees are similar to other the sea level continued to rise with the melting of the ice sheets. pollinating species. By perhaps 10-9.000 BP Ireland was cut off, although plants and Dates reported from prehistory are difficult to integrate as animals continued to cross for another 2,500 years (Mitchell, some were deduced by traditional geological or archaeological 1990). By 10,000 BP the British mainland was clothed in a techniques, some by radiometric dating and some by application mixed forest of hazel, oak and elm and by 9000 BP, the climate of dendrochronology and varve deposition patterns for having become warm and dry, hazel was so dense as to contribute recalibration of radiocarbon dates (see Andersen and Borns, 80% or more of all tree pollens. 1992; Hulme and Sherriffs, 1985). In this report I consider the Over the next few thousand years, temperatures remained Late-Glacial period from 16,000 BP to 11,500 BP (calibrated reasonably constant, but the sea level continued to rise, creating dates) to correspond to the Upper Palaeolithic period of human the English Channel at 8500-7,500 BP and preventing much culture in Britain. The subsequent archaeological phases are further natural ingress of animals and plants. The mixed oak Mesolithic, from 11,500 to 7,500 BP, Neolithic to — 4000 and hazel forest expanded to become the dominant vegetation BP, and Bronze Age to — 2,500 BP, followed by the Iron Age type by 6000 BP, when the sea reached more or less its present extending into the period of Roman occupation. level. That concept is however masked by isostatic readjustment There is archaeological evidence of honey bees being in Britain of the land following melting of the ice, as the land tilted and the in the Iron Age and even the Bronze Age, and this is reinforced south-east sank relative to mean sea-level. by palaeobotanical evidence that honeybees almost certainly thrived there several thousand years before the Christian era (see The arrival of honeybees below). As a consequence of the work of Friedrich Ruttner (1988) I conclude that the "British Dark Bee", the Northern honeybee, it is generally accepted that the Northern Honeybee (A.m. Apis mellifera mellifera (L), almost certainly extended its range mellifera) survived the later glaciations in refugia, notably in into Britain during the Mesolithic, unaided by Man and well southern France and Spain. There is evidence in the form of before the land bridge to Europe was lost to the rising North Sea, rock art, of honey- gathering by Mesolithic man near Bicorp at about 8500 BP (Cunliffe, 2001). Analysis of British honeybee in Valencia, eastern Spain at 9 - 8000 BP (Crane, 1975). The mitochondrial DNA reveals ancient queen lines that are similar to important question though is whether conditions were suitable 11 Is the Dark Bee really native to Britain and Ireland?

Dorian Pritchard

for honeybees (Ruttner, Milner and Dews, 1990; Crane, 1999). Wild honeybees also need rock cavities, buildings or substantial hollow trees for nest sites secure from predators. In lowland regions nest sites can be provided by hollow broadleaved trees, such as oak, elm, lime (Tilia spp.) and beech, these species, along with hazel and cherry, being recognised as defining suitable territory for honeybees (Ruttner, Milner and Dews, 1990). However, honey bees have also been known to utilise old woodpecker nests in birch trees of lesser dimensions (Carreck, N. personal communication). Mixed oak forest with hazel and abundant lime in the south was the main cover of Britain throughout the Mesolithic (Andersen and Borns, 1992). Lime and hazel were both present in Norfolk at 8250 BP (Godwin, 1975 — see Crane, 1999) and hazel, but not lime, in Perthshire (Hulme and Sherriffs, 1985 — see Crane, 1999). Rackham (1986; Fig. 52) illustrates oak-hazel forest at 6,500BP extending as far north as Skye in the west and Buchan in the east. On such evidence Crane (1999) concluded honeybees could have been present in Britain before 7500 BP, in line with Limbrey's previous conclusion that honeybees arrived in Britain along with hazel and lime (Limbrey, 1982). Mixed oak and hazel forest are indicated in the fossil record by pollen, acorns and hazelnuts, as well as traces of their associated fauna, red deer (then a forest species), roe deer and wild pig. A Mesolithic site on the island of Risga in Loch Sunart, Argyll yielded bones of red deer and wild pig and there are numerous 1 early Mesolithic sites along the valley of the Bann in Northern Ireland, where many young pigs were butchered and quantities of hazelnuts, wild pear and crab apple collected (Cunliffe, 2001). Considerable quantities of hazelnuts shells were found among pollen of alder, small-leafed lime (Tilia cordata), oak and elm at a Mesolithic site at Oakhanger in the Weald and the pollen record shows similarly favourable honeybee habitat extending Fig. 1. Western Europe at 9000-8600BP, during the Mesolithic throughout much of Europe, including France. Such evidence period, showing the extent of continuous broad-leafed forest (An- substantiates the concept, widely held among archaeologists, dersen and Borns, 1992) and the probable path of migration of that oak-hazel forest with its normally associated fauna extended Apis me/lifera mellifera (L.) from its glacial refuge in southern virtually unbroken from southwest France as far as southwest France and/or Spain (Ruffner, 1988) and across the land bridge Scotland at least a thousand years before the land bridge closed to Britain. The dark blue areas represent land-locked water. (see Fig. 1; Andersen and Borns, 1994; Cunliffe, 2001). As the climate ameliorated it would be expected that forest Legend: (left to right; top to bottom) i) Birch forest. ii) Broad- animals sequestered in southern glacial refuges would have leafed or hazel forest. iii) Mixed birch-pine-spruce forest. iv) moved north with the advancing forest. However, it is worth Mixed pine or spruce and broad- leafed or hazel forest. v) Mixed considering that if honeybees had instead remained in their pine or spruce and hazel forest vi) Alpine vegetation. glacial refuge until the land bridge itself was fully forested, how far would they have had to travel to reach England and how long would they have taken? for a sufficient length of time for honeybees to move north and across the land bridge to Britain, before that route was lost to the From Toulouse to Dover is about 800 km. On the conservative rising sea. estimate that a wild colony might swarm once every second year and that swarms might travel an average of 1.5 km to their In present-day Britain A.m.mellifera survives in the wild to just new homes, starting from Toulouse and travelling "in a beeline" north of Inverness (Scottish Beekeepers'Association literature), they could have reached Britain in 1100 years. The available while hazel (Corylus avellana) is present as far north as Orkney time window between the commencement of amelioration of and Shetland. This supports the accepted concept that the the British climate (-11,500 BP) and the flooding of the English presence of hazel broadly indicates suitable climatic conditions Channel (-8,500 BP) was several times that long. 12 Fig. 2. Western Europe at 7500-2500BP, during the Neolithic and Bronze Age, showing the vegetation cover (Andersen and Borns, 1992), the major human migration routes into Britain at that time (Oppenheimer, 2006) and sites where evidence of honeybees and/or beekeeping have been found.

Legend: (left to right; top to bottom) i) Birch forest. ii) Broad- leafed or hazel forest. iii) Mixed birch and pine or spruce forest . iv) Mixed pine or spruce and broad- leafed or hazel forest. v) Pine and /or spruce forest. vi) Alpine vegetation.

1. Dun Aonghusa, Aran Islands. Evidence of lost-wax casting c. 3200-2600BP (Cunliffe, 2001). 2. Area where Ballintober bronze swords were probably made by lost-wax casting during the Bronze Age c.4000-2500BP (Op- penheimer, 2007). 3. Jahlshof, Shetland. Evidence of lost-wax casting c. 3200- 2600BP. 4. Ashgrove, Fife. Beaker of honey or mead c. 3000BP (Dick- son, 1978; Dickson and Dickson, 2000). 5. Egtved, Denmark. Traces of drink derived from honey and grain in birch-bark bucket, c. 3030BP (Dickson, 1978). 6. Lichterfelde, near Berlin. Log hive, c. 3030BP. Ii i 7. Hayes, Middlesex. Remains of two worker bees c. 4000- 3750BP and 16 honey bees from the Roman era (Robinson, 2007). 8. Dover, Kent. Bronze Age boat caulked with beeswax, c. 3550BP (Clark, 20 04). 9. Runneymede, Berkshire. Remains of worker bee (Robin- son, 2000) and traces of bees wax and honeydew on pottery, c. 4650BP (Crane, 1990; Needham and Evans, 1987). 10. Eastern Spain. Rock painting of honey-gatherer, c. 9000BP (Crane, 1975).

The present natural distribution of the different A.m. subspecies 3700BP) are from the Perry Oaks excavations at Hayes in in Europe accords closely with mean temperatures in July Middlesex, where two honeybee workers were found in separate (Pritchard, 2006). This suggests it is predominantly summer excavation pits (Robinson, 2007). The same site yielded 16 temperatures that limit their spread (see also Crane, 1999). The honeybees dating from Roman times. A single worker bee was evidence of flower pollen and beetle exoskeletons reveals mean found in a late Bronze Age deposit at Runnymede in Berkshire July temperatures in Britain to have been similar to current mean (Robinson, 2000), as well as traces of beeswax and honeydew temperatures (15 -20°C) since 9,500 BP (Andersen and Borns, from silver fir (Abies alba) on pottery dating from the Neolithic 1994) and perhaps a degree or so warmer during the Post-Glacial (c. 5000-4650BP; Crane, 1990; Needham and Evans, 1987). Climatic Optimum of the early Holocene. At 13°C it is however A beaker containing pollen of the small-leafed lime (Tilia sufficiently warm in present-day Inverness (Tansley, 1965) for cordata), meadowsweet (Filipendula ulmaria) and heather foraging and swarming of northern bees. It would be quite (Calluna vulgaris) was found in a Bronze Age burial (c.3000 unreasonable to suppose therefore that A.m. mellifera would not BP) at Ashgrove in Fife and thought to have contained honey have moved into Britain in the available time window. Indeed or mead (Dickson, 1978; Dickson and Dickson, 2000), although archaeological evidence suggests it definitely did. this was probably imported, as there is no evidence small-leafed lime grew north of the Scottish Border at that time or since Direct archaeological evidence (Rackham, 2003). A Bronze Age boat caulked with moss and Unlike beetle elytra, the remains of dead honeybees rarely beeswax and dated to circa 3550BP was excavated at Dover survive for long. Nevertheless several British Bronze Age sites (Clark, 2004), although either the boat or the wax could have have yielded honeybee remains. The earliest (dated to 4000- originated on the Continent.

13 Is the Dark Bee really native to Britain and Ireland?

Dorian Pritchard

That beekeeping was practised in Northern Europe as early as Historical evidence the Bronze Age is demonstrated by a log hive, dated to 3030 BP, Strabo (7BC) reported that the Greek adventurer, Pytheas of excavated at Lichterfelde, a suburb of Berlin, similar to those Marsalias (Marseilles) visited and circumnavigated Britain still used in rural areas of Central Europe (Lehmann, 1996). in 344BC. He reported that the inhabitants of a northern land The analysis of human DNA reveals considerable human called Thule enjoyed a drink made from fermented honey and traffic at that time across the North Sea between what are now grain (Fraser, 1958). Thule was described as six days' sail Denmark and Eastern England (Fig. 2; Oppenheimer, 2007). north of Britain, where the sea froze, but volcanoes made it If honeybees were naturally absent from Britain in the Bronze boil. It was probably Iceland. This suggests that honey was Age, settlers from mainland Europe would surely have taken known in Northern Europe over 800 years before the Christian them there, establishing their presence in the British fauna, if monks. Indeed such a drink, contained in a birch bark bucket, artificially, some 3000 years ago. was placed among grave goods at Egtved in Denmark thousands Records from more recent times include the head of a worker of years earlier, in the Bronze Age (see Fig. 2; Dickson, 1978). Apis mellifera is the only western European species capable of honeybee in peat deposits at Hardwick-with-Yelford,Oxfordshire, supplying honey in large quantities. at an Iron Age site dating from around 220 BC (Robinson, 1984; Allen and Robinson, 1993). Kenward (2005) lists six British The first historical record of honeybees in Britain is a Roman sites from Roman times where samples of members of the document (c. AD100) found at Vindolanda, a fort beside genus Apoidea (probably A.m.m.) have been found, and one Hadrian's Wall in Northumberland, which mentions "lini in Carlisle where a specimen was identified as Apis mellifera. mellari" or "honey cloths" that were probably used for filtering Wings of honeybees taken from the excavations at Coppergate honey (Crane, 1999). in York and dated to 975-935AD were in a sufficiently good state of preservation for morphometric analysis, confirming their Evidence for survival of the native bee to the present subspecies identity as A.m. mellifera (Ruttner et al., 1990). Around the time of the First World War the honey bee population of Britain was devastated by a mysterious disease originating in Taken together, these findings confirm the undoubted presence the Isle of Wight when, according to official statistics, 90% of of Apis mellifera mellifera in Britain well before the modern era. Britain's honeybees met their demise [Butler, 1954]. One of the The earliest recorded importation of foreign (Italian) honeybees most eminent and influential of Britain's beekeepers, Brother is AD1859 (Fraser, 1958). Adam, later senior apiarist at Buckfast Abbey, interpreted this figure as 100% and throughout his very long life expressed the Cultural evidence from prehistory erroneous view that the native British honeybee was extinct (e.g. Crane (1999) noted the close similarity of Irish, Breton and Adam, 1987a). Welsh words for obscure aspects of honey bee behaviour. For example a second swarm, or first cast, containing a virgin queen Nevertheless Cooper (1986) surveyed and studied the and drones, is in all three given the unusual term "bull swarm", survivors and recorded their external appearance, behavioural "tarwheit" in Welsh, "tarbsaithe" in Irish and "tarvhed" in Breton. characteristics and distribution. He founded the Village Bee (In Welsh "taranu" means "to thunder") This suggests a detailed Association to promote their conservation, which evolved into knowledge of honey bee behaviour was acquired by Celtic the Bee Improvement and Bee Breeders' Association (BIBBA). speaking people prior to their arrival in Britain - and retained Members of BIBBA in collaboration with the German scientist, after their arrival. Their route was from Iberia, in the Bronze Friedrich Ruttner, have led subsequent efforts to study and Age or earlier, i.e. before 2500 BP (see Fig. 2; Oppenheimer, conserve the native bee and to formulate a highly definitive set 2007). They also could have taken hives of bees with them. of characters by which A.m.m. can be distinguished from the other subspecies. These include general body colour, proboscis A major feature of Bronze Age culture was metallurgy and length, tomentum width, body hair length, cubital index and the casting of intricately shaped objects in bronze, notably Irish discoidal shift, the latter two referring to measurements on the swords of the Ballintober pattern and socketed axe heads by the patterning of forewing veins (Ruttner, 1988; Ruttner, Milner and thousand. There is dispute over the extent to which the "lost- Dews, 1990; Dews and Milner, 1993). wax"or "cire perdue" process was used (Hodges, 1958), but this technology, which utilises the then unique properties of beeswax, Recent surveys report honey bees of native type in 14 counties was well established in Britain before the beginning of the Iron of England and Wales and all but two of the Scottish Regions Age (Wilcox, 2008). The relics of crucibles and clay moulds (Pritchard, 2006; Stoakley and Stoakley, unpublished). Samples used for lost-wax casting of axes, swords, spearheads, bracelets, from at least six of these have had their identity as A.m.m. ornaments and tools and dating from the late Bronze Age (3,200- confirmed by DNA analysis. 2,600) were found at both Dun Aonghusa in the Aran Islands off Following the I.O.W. epidemic honeybees of several races Galway Bay and Jahlshof in Shetland (Cunliffe, 2001). This (subspecies) were imported in quantity into Britain, including industry would have created an industrial and military demand A.m.m. from Holland and France. By most criteria the for beeswax, so it is possible that metal casting in the Bronze descendents of the latter are indistinguishable from native Age could have provided an early commercial incentive for British bees, but can however be discriminated by analysis of development of beekeeping in these islands. their DNA. DNA analysis has also detected introgression of 14 foreign DNA into the genomes of identifiable colonies among several samples from England and Scotland and two from populations of true British dark bees. Jensen et al (2005) carried Ireland. One Irish and two English East Coast samples show out such an analysis; they emphasized the distinctiveness of the close similarity to samples from mainland Europe, suggesting native British bee from those of other European populations and they could have originated there. However, several English, stressed the urgent need to preserve its genetic integrity. Scottish and one Irish sample appear to be absolutely unique to the British Isles, although most show relationship to one another The evidence of DNA and, less closely, to others in mainland Europe. One English Four categories of DNA can potentially yield information on sample is fundamentally unique. Samples collected in Tasmania the ancestry of Britain's honeybees. These are chromosomal from a stock derived from Yorkshire honeybees taken there coding DNA, chromosomal non-coding DNA, mitochondrial in 1834 AD have mtDNA identical to that of their relatives in coding DNA and mitochondrial non-coding DNA. Yorkshire. Since the sequencing of the honeybee genome (Honeybee Not all extant stocks have yet been examined, but unique British Genome Sequencing Consortium, 2006) it has become and Irish samples are most easily explained as representing stocks possible to utilise minor variations called single nucleotide that arrived there before the closure of the land bridges, evolved polymorphisms, or SNPs, in the nuclear coding sequences, or subsequently in isolation and are truly native to the British Isles genes. This approach is very new, but supports the concept that (see Pritchard, 2006). A.m.mellifera advanced into N and NW Europe from a refuge in DNA analysis therefore provides a completely independent the SW (Whitfield et al, 2007). approach that leads to the same conclusion: that the British The category utilized in forensic "DNA fingerprinting" is Isles are home to descendents of honey bees that evolved there a chromosomal non-coding fraction called "microsatellite in isolation from the rest of Europe and are therefore native to DNA", which is also found in honeybees. This consists of these islands stretches of repeats of short sequences of nucleotides at several sites near the ends of chromosomes. The number of repeats Conclusions varies between individuals and is usually inherited unchanged In answer to the question: "is the British Dark Bee (A.m. (see Pritchard and Korf, 2008). The different subspecies of mellifera) native to Britain?" we can only answer in the honeybees characteristically have different combinations of affirmative. Palaeobotanical study reveals that environmental the repeat series, which allows us to understand the ancestral temperatures and vegetation cover were appropriate for a origins, evolutionary past and relationships of honey bee stocks sufficient period of time during the post-glacial period for (Garnery et al, 1998b). honeybees to enter the British Isles (see Fig. 3). It is mitochondrial DNA of both categories that has so far been Archaeological studies provide inferential evidence of the most informative. This occurs as rings in each mitochondrion, honeybees in Britain during the Mesolithic, Bronze, Iron and independent of that in the chromosomes. It is transmitted to Roman periods, as well as actual specimens of Apis mellifera the next generation only by queens and consequently there is from the Bronze and Iron Ages and Roman times. Whether or no reassortment between DNA of maternal and paternal origins. not honeybees arrived in Britain under their own muscle power, The mitochondrial genome contains a non-coding sequence as seems most likely, or were brought in by Neolithic or Bronze of DNA of unknown function, which has evolved rapidly and Age human immigrants is of little import today, as the British diversified considerably among the different queen lines, which ecosystem has undoubtedly evolved in their presence for at least allows us to place honeybee populations in major categories 3000 years. designated I, II, III, IV, etc.(Garnery et al, 1998a). The situation with respect to Ireland is more problematic, as the MtDNA also contains a small number of genes, one of the most Irish Sea formed perhaps 2000 years before the English Channel important of which is that for cytochrome C. This enzyme is so and as yet there seem to be no archaeological records of Irish crucial to survival that only very minor mutational changes are honeybees. However, the DNA of (at Ieast) one Irish stock, the tolerated by natural selection. Nevertheless, over time single archaeological evidence of lost-wax casting at Dun Aonghusa, base changes occur and accrue and can be most informative the abundance of Irish bronze swords and the linguistic evidence (Itenov and Pedersen, 1991). argue that the northern honeybee, A.m.mellifera, has probably Detailed examination of the DNA of an individual can therefore been in Ireland also at least since the Bronze Age. inform us about the distant maternal origin of individual queens, Do native British and Irish Dark Bees still survive? Yes: DNA and potentially their relationships to all other queens descended analysis identifies stocks of the northern honeybee currently from a common ancestor. extant in Britain and Ireland that are genetically unique to these islands. The only sensible conclusion can be that at least some of By exploitation of these features research groups led by Lionel the British and Irish living specimens of A.m.m. are truly native Garnery in Versailles and Bo Vest Pederson in Copenhagen have to both Britain and Ireland, as defined in the opening paragraphs. succeeded in elucidating the relationships between many honey (See also Carreck, 2008 and Pritchard, 2008.) bee populations (Cornuet and Garnery, 1991; Pedersen, 1996; Jensen and Pedersen, 2005; Pritchard, 2006). These include 15 Is the Dark Bee really native to Britain and Ireland?

Dorian Pritchard

ACKNOWLEDGEMENTS 113.

I wish to thank Norman Carreck and Dr Angus Lunn for critical Fraser, H.M.1958. History of Beekeeping in Britain. International comments and a valuable exchange of ideas. Bee Research Association: London, UK, 106pp.

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