A QUARTERLY JOURNAL OF NATURAL HISTORY FOR THE NORTH OF ENGLAND

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From Coney to Rabbit: the Story of a Managed Coloniser —

A. Henderson

The Horse-trough Ostracod Heterocypris incongriiens —

Geoffrey Fryer

Diet of Barn Owls Tyto alba in a Low land Area of North
Yorkshire — J. Roberts, G. W. Scott and S. Hull

Published by the Yorkshire Naturalists’ Union

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FROM CONEY TO RABBIT:

THE STORY OF A MANAGED COLONISER

A. HENDERSON

101

i

i

Presidential address presented to the Yorkshire Naturalists
at Beverley, 7 December, 1996

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HISTORY MUSEUM

“Thys hand kan ferette in konnyngherys"

John Lydgate
The Pilgrinuiiie of Man (1426)

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l Introduction

To find a rabbit native to the part of the world we nowadays call the British Isles, we would
r need Dr Who’s Tardis to carry us over lO.OOO years back bel'ore the close of the
I Pleistocene Age when, the lossil record shows, the rabbit became extinct here and
t throughout most of Europe, surviving the Ice Ages only in the milder region of the Iberian
f peninsula. This was the part of their known world, their mappamundi, that the ancient
L Greeks and Romans regarded as its home: in the words of Catullus, in the last century BC,
‘Celtiheria cnnicidosa' , the rabbit-teeming Spanish peninsula.

f From the Mediterranean to Britain

The animal's nuisance value was already recogni.sed. Catullus’ contemporary, the Greek
t historian Strabo, describes how the colonists of the Balearic Isles petitioned the Emperor
Augustus to rid them of this pest or find them.selves other territory. Later, in the 1st century
AD, Pliny the Elder recounts how the walls of Tarragon succumbed to undermining from
r rabbit-burrowing: a reverse story to that of the geese which saved Rome. However, the
„ quality of the animal’s meat and fur were reckoned to be redeeming features. Catullus,
; admittedly in liercely .satiric mood, describes the debauchee, Thallus, as 'moUior ainiadi
cupillo', softer than the rabbit’s fur. The Roman epicure chef, Apicius, includes rissoles of
' rabbit (prized less than those of peacock or pheasant, but above those of chicken or tender
■ sucking-pig) amongst his recipes. Awareness of these favourable characteristics and of the
animal's sporting potential led the Romans to place the rabbit alongside the hares which, as
' Varro de.scribes, they were already keeping in leporaria or hare-warrens. Eventually, when
more exotic animals were added to their populations, some of these warrens became, in
. effect, vivaria or virtually game-reserves; perhaps we should really think of them as
prototypes of Whipsnade with the rabbit and hare in lieu of wallabies. Although Roman
law stipulated that game was res nnllius (finders and catchers keepers), the existence of
• such private leporaria shows that this was not entirely without qualification. From the
' Mediterranean and southern France rabbit-uarrens spread steadily through much of
western Europe, reaching Germany (e.g.. the island warren at Schwerin-See in 1407)
(Zeuner, l%3) and Hungary (Margaret Island in Budapest, originally called Rabbits'
Island) (Bdkonyi, 1974). At some time in the I Ith or 12th century the Normans introduced
the animal into Britain, and by the mid- 1 3th century there were rabbit-warrens as far north
as Crail and Cramond in Scotland (Gilbert, 1979) and as far west as Ballysax and Rosslare
in Ireland (Hayes. 1965) (Fig. 1).

Early iconography

Data on the rabbit’s early days in Britain are scattered, often brief mentions in the diverse
historical record. Perhaps some of the most delightfully informative notions we can arrive
at come from its entry into ecclesiastic and other iconography. It is surely serendipitous
that this talk is being given in St Mary’s parish hall. Beverley. St Mary's being the home of

Nuiunilisi 122(1 ‘)97 )

102

From Coney to Rabbit: the Story of a Managed Coloniser

FIGURE 1

Distribution of the rabbit in Europe and North Africa.

Erom Biadi, E. ( 1993) Le lapin de garenne. Office National de la Chasse.

one ot the county’s most well-known figures of a rabbit (Fig. 2), .sculptured about 1325, but
probably better known now than then, as it is rumoured to be the source of inspiration for
Tenniel’s White Rabbit in Carroll’s Alice story, a distinction well-deserved by this proud
specimen of Oryctolagiis ciinicidus, the tunnel digging hare, as his Greek and Latin roots
define him. On late medieval tiles of the 13th century or just after (Williams, 1985),
unearthed during the restoration of Bangor cathedral in the 1890s, he is seen emerging
from his burrow in repeat pattern. At Kilpeck church, Hereford, one of seventy l2th
century carvings on the corbel table (Fig. 3) is possibly the earliest sculptural likeness of a
rabbit in Britain, certainly one of the most engaging and Disneyesque. Particularly
informative, however, is one of the hagiographic illustrations (Fig. 4) to Lydgate’s life of
the holy English king, St Edmund, which shows his miraculous recapitation. When the
king was attacked and overpowered, his head was severed from his body, and this narrative
■scene pictures both a guardian wolf presenting the lost head to a team of ministering
monks, and the moment of reattachment. The incident is reputed to have occurred in 835
AD, but the 1 5th century miniaturi.st has pleasingly and anachronistically set the event in a
1 5th century coneygarth or rabbit-warren, giving us one of the best intimations we have of
such early managed warrens. Other pictures of the time show woven wattle fencing,
brushwood or other thick hedging bounding enclosures where conies thrive. In Queen
Mary’s FNalter (Warner, I9I2) women are represented catching conies with ferrets and
nets, and engaging in what one can only call a concy-bashing exercise! Even amongst
ground game conies were usually accounted of low status by mediaeval nobility and
gentry, and it may be that machoism as well as male chauvinism considered it woman’s
work to catch conies for the table, etc., in the nearby coneygarth.

From Coney to Kcihhit: the Stoiy of a Managed Coloniser

103

FIGURE 2

Pilgrim Rabbit, St Mary's Church, Beverley; carved c. 1325, and reputed to be the
inspiration lor Lewis Carroll's White Rabbit. Photo: Chris Hairsine.

FIGURE 3

Rabbit and dog sculpture from the corbel table of Kilpeck church.
From; The Sculptures of Kilpeck: ©Reg Boulton, 1987.

104

From Coney lo Rabbit: the Story of a Managed Coloniser

FIGURE 4

The recapitation of St Edmund. A 1 5th century miniature, illustrating Lydgate’s
Life of St Edmund. By permission of the British Library: Harley MS 2278 f 67v.

Etymology

In Middle English the rabbit was known as ‘coney’ or ‘cunning’, derived from the Old
French nouns ‘connil’ and ‘connin’. A rabbit-warren or earth was, therefore, a ‘cunning-
erthe’, leading to the later commonly occurring form, ‘coneygartlT. This and the Latin
‘coniger’ (= ‘coneygarth’) lead to such common corruptions as ‘conigree’, ‘coninger’,
‘cunnery’, etc. At first, the term ‘rabette’ referred only to the young rabbit, ‘coney’
referring to the adult animal. Eventually the terms ‘coney’ and ‘rabbit’ became
interchangeable, until today, of course, ‘rabbit’ is our usual name for the animal at all ages.

Indication of warrhn .site.s

Cartographic names can be effective indicators of the sometime presence of a coneygarth
or rabbit-warren. Generally, such names as ‘coneygarth’. ‘conigree’, etc., ‘warren’, ‘warren
house, farm or hill’, etc., are signs of one-time managed rabbit activity, although such
double entendres as ‘coney’, very occasionally deri\ing from the Teutonic root ‘koenig’
(= ‘king’), and ‘warren’ after the Earls Warenne and their kin. are grounds for caution.
Such indicators can then be checked in likely documentary sources, such as Charters.
Patent Rolls, liu|uisitions post Mortem, tithe and enclosure maps with the accompanying
awards, estate maps, terriers, manor extents, leases, etc. References may also be found in
local and personal records, diaries, notebooks and correspondence, game registers, kitchen

103

From Coney to Rahhit: the Story of a Managed Coloniser

accounts, etc. The reliability ol initial cartographic indications is evidenced by the
frct|Licncy with which lurther documentary or held searches yield supporting proof of the
existence of a managed warren in a given area at a given time. Such sites are, indeed,
commoner than might at lirst be supposed. That part of Yorkshire lying west of York has
already yielded evidence ol 130 such one-time warren sites. Equally important methods of
detection and exploration are provided by lieldwork and the consultation of aerial
photographs. These can reveal the presence in a landscape of some of the warren features
and artefacts now to be described.

Warren structurh.s and artefacts

The situation ol warrens and the structures and artefacts within them, large and small, are
governed by topographic, geo-edaphic, economic and practical agricultural dictates, which
has led to considerable individualisation in their establishment and realisation. The large
county ol Yorkshire is fortunate in having representatives of a wide range of warren forms
and artefacts.

Boundaries

Hurrell ( 1979) has drawn attention to the earliest known record of rabbits in Britain, in 1 133
AD on Drake’s Island in Plymouth Sound, when ownership of the island was transferred to
Plympton Priory ‘cam cuniciili' |sic| (= with the rabbits). Several of our earliest warrens
were on islands, where an open or largely unenclosed warren was provided with natural
boundaries against rabbit escape and predator intrusion. Thus, the Countess of Arundell’s
‘conny warrin’ on the south-western point of Hayling Island (East. 1891) required the
erection of only a short length of artificial boundary (a mere seventh of the total perimeter).
The earliest record of a mainland warren cited by Hurrell is a deed of transfer for a
Dartmoor warren, in the area of the later Trowlesworthy Warren, vaguely dated as 'drawn
up' sometime between 1 133 and 1272’. There are certainly other records within this interim
(for instance, a royal order of 1240 (Cal. Cl. Rolls. Henry III 1237-1242) to allow W'. de
Ferrariis to take 20 live rabbits from the Wirral; another order of the same date for 100
rabbits to be taken by Hugo le Franceys from Dorking and Reigate; and orders of 1233 for
10 live rabbits and of 1242 for 30 or 40 rabbits from the cunnery in the park at Guildford).
Domesday Book does not mention any rabbit-warren, but interestingly lists one hare-warren
at Gelston, Lincolnshire, a precursor of numerous later hare-warrens, such as one on the Isle
of Purbeck during the reign of King John, when Purbeck was a forest with a ranger and two
warreners preventing poaching of 'coneys, hares, foxes and pheasants' (Hyland. 1978). one
at Hampton Court in the 16th century and .several in post-mediaeval Wiltshire.

Like insular warrens those on the mainland coast made use of the sealine as a natural
boundary. Such warrens, however, often entailed considerable boundarx' construction,
utilising whatever materials w'ere at hand locally. Warrens in general were mainly situated
on marginal economy land of low fertility. (Hence, later, the development of the vast
commercial warren complexes that occupied thousands of acres on Dartmoor, in the East
Anglian Breckland. on the East Riding Wolds and the North Riding Tabular Hills, where
they reached from Hutton-le-hole to Hackness (Harris & Spratt. 1991). They were
frequently on rougher upland sites or. if lowland, on sandy, gravelly or heathland areas.
Earthen or sod embankments were constnicted to .serve as convenient and necessar>'
boundaries in many such cases, topped by growing gorse or thorn below' a cap of turf (Figs.
3 & 6). Crackles (1993) has pointed to the importance of gorse as winter fodder for
managed rabbits, and notes the use of gorse faggots as bearding on the top of warren sod
walls, overhanging the inner face to prevent rabbit escape. In regions like Yorkshire and
Dartmoor, where stone was available, stone walls or revetted embankments were employed
as well. Like earthen embankments they might akso be used to construct smaller internal
breeding and feeding enclosures within the larger warren territory, such as the four
newtakes known as the Four Aces, still in good preservation and visible over long distance
at Headland Warren. Dartmoor.

106

From Coney to Rabbit: the Story of a Managed Coloniser

FIGURE 5

Peripheral warren wall, constructed of tree-stumps, stones, twigs and cloth, covered with
light non-clayey earth, and roofed with stout branches and a variety of plants.

Biadi, F. ( 1993) Le lapin de garenne. Office National de la Chasse.

FIGURE 6

Reconstructed warren, Knettishall Heath.

By permission of Suffolk County Council, Graphics Section (G. Wade, artist).

Lodges and other warren ttousES

The most upstanding feature of the warren landscape was the lodge or warren house. Since
the demise of warrens, such lodges as remain often stand in a landscape that has progressed
one way or another to woodland, and they have lost some ol the stark dominance that led
Shakespeare to describe Claudio as ‘melancholy as a lodge in a warren'. Forest lodces,
such as the Royal Hunting Ia)dgcs of King John and Queen Eli/.abeth were primarily

107

From Coney to Rcihhit: the Story of a Managed Coloniser

directed towards the pursuit of higher game and enjoyment of the chase. Queen Elizabeth’s
Lodge, indeed, is reputed to have had an open-sided first storey for viewing the full
panoply of the passing hunt in comfort. The role of belvedere or gazebo was doubtless one
played at times by many lodges, but usually, like the warren house proper, their major role
was a custodial one, providing a working base for the warrener and his assistants, and for
storage, gutting, packing, etc.

Thetford Warren Lodge, on brown calcareous Breckland sands, is still, despite being
gutted by fire early this century, a bleak and sombre edifice, a former centre of defensive
activity in a fortified warren liable to threat by armed poachers, its stout walls in the local
flint. The most distinctive lodge of all architecturally is the Rushton Triangular Lodge in
Northamptonshire (Fig. 7), a laudable curiosity (English Heritage, 1997), built between
1593 and 1597 by Sir Thomas Tresham in celebratirm and honour of the Holy Trinity, with
three walls on a triangular plan, each 33 feet long and with three windows with three gables
to each, three-storied and with a three-sided chimney on top. Clearly, a building of profound
devotional intent in its design; but one which the Rushton accounts refer to as the
Warreyner’s Lodge (Finch, 1956). Manuscripts in the Public Record Office show that the
warren was 300 acres and that there was also a neighbouring connegerie at nearby Pipewell.
Ferrets were kept for bolting the rabbits, and boxes of rabbits and skins were despatched
three times weekly to London, amounting to a lucrative business in contracts with poulterers
there. The official in charge of such a warren would enjoy real status, unlike those employed
merely for the ruder tasks of warrening. In Yorkshire, Norton Tower (Fig. 8) on a high

FIGURE 7

Rushton Triangular Lodge (The Warreyner's Lodge).
© English Heritage.

108

From Coney to Rabbit: the Story of a Matutf’ed Coloniser

escarpment edge south ol Rylestone, the Rylestone of Wordsworth's White Doe, is now bul
a ruin ot its former self. It stands on old warren land with a commanding outlook over the
surrounding terrain. Moorhouse (1991) has identified the site as a late mediaeval rabbit-
warren and hunting enclosure, probably owing its origin to a dispute over hunting rights
between the Norton family of Rylestone and the Cliffords of Skipton who held much land to
the east.

In other warrening areas, of course, much less grandio.se buildings can be found. One
example is Ewden Cote on the Broomhead Hall estate in the Ewden valley, north-west of
Sheffield. This building was used as a gutting house. Upper and lower chambers to the
south have well-flued fireplaces for drying out hanging skins, and the building doubled as
an effective shelter for shooting parties in inclement weather, as well as providing simple
living quarters for a working warrener and family or his assistants. It has a covered loading
bay for storage and despatch by cart of rabbits and skins.

Pillow mound.s

Among the best known features of old coneygarths are pillow mounds; low mounds,
usually rectangular in plan and surrounded by a ditch where earth has been thrown onto the
mound, and fashioned to encourage rabbit habitation, the main aim of creating such
mounds being the localisation of rabbit activity with obvious advantages for a busy
warrener. Some pillow mounds consist entirely of soft or gravelly earth; others include
varying amounts of stone as a constituent, larger flatstones sometimes being used to cap
man-made burrows to prompt further spontaneous burrowing by the rabbits. Rounded and
cairn-shaped mounds are also encountered. Moorhouse notes them near Norton Tower, and
there is a small rounded mound, a low cone to the view, alongside .some of the rectangular
mounds at Elasby. At Merrivale Warren, Dartmoor, among many rectangular mounds one
was observed that is horse-shoe shaped. Probably the finest array of pillow mounds in ^
Yorkshire is to be seen on top of Douthwaite Dale, north-west of Hutton-le-hole (Fig. 9).

Recently. Williamson and Loveday ( 1988) have studied a range of pillow mounds, inter
alia in the light of possible confusion with similar Neolithic earthworks, etc. Examining
pillow mounds at 190 sites, they found the commonest width (c 60% of samples) was
between 4 m and 6 m, with a minimum width between I m and 2 m. and a maximum width >
ol 16 m. Lengths varied from 8.4 m to 234 m. In comparison, pillow mounds seen in
Yorkshire fit comfortably with their analysis.

FIGURE 8

Norton Tower, Rylestone. Photo: .1. Taylor.

109

From Coney lo Rabbit: the Story of a Managed Coloniser

FIGURE 9

Array of pillow mounds on top of the escarpment. Douthwaite Dale, north of Hutton-le-hole.
By permi.ssion of the Royal Commi.ssion on Historic Monuments: RCHME.

© Crown Copyright; NMR 12098/76.

Use of older landscape eeatl'res

Older earthworks have not uncommonly been used later in the establishment of warrens. At
Dolebury Warren. Somerset (Oxford. 1986) linear pillow mounds cross the ancient
hillfort's interior. A shallow ditch surrounds each, and they reach a width of 10 m. and
range in length from L'i m to 100 m. with a present height up to 1 m. At the Iron Age
hillfort of Danebury, Hampshire, the area was. according to an account of 1678 (Cunliffe.
1983), ‘anciently and till since the memory of man a warren and let to several tenants at
rack rents and was entire to itself and (not a sheep common) had a lodge in it. Only since
the rabbits were destroyed they have fed the sheep there, and when it was a warren the
sheep many times went over it and not questioned because the warreners' rabbits
trespassed as much on their sheep-down". Here in Yorkshire, at Huddlestone Old Wood,
near Mickleheld, adjacent to Coney Garth Hill (shown on the 1850 O.S. map), lies a now
thickly wooded, irregularly hexagonal embankment system, about 0.25 km- in area, with
evidence of warrening u.se where a 0.5 km-long transverse internal embankment has been
trenched out, the trench then capped over w ith flatstoncs and heaped over with earth.

no

From Coney to Rabbit: the Story' of a Managed Coloniser

One of the most intriguing of Yorkshire warren sites is at Cockmoor Dykes, north of
Snainton on the Scarborough-Pickering road. These dykes (Spratt, 1989) form an eastern
part of the extensive system of ancient dykes on the upland limestone areas, which
delineate strip territories. Cockmoor Dykes is a most impressive array of twenty closely
parallel ditches and hanks on the Troutsdale escarpment. Spratt concluded that the six
larger dykes formed part of the original boundary system, but the fourteen smaller dykes
originated in the 18th century, a period of major warrening activity in this area. Today the
dykes are riddled with very old and new burrows, and there is a rabbit tip-trap (see below)
nearby.

Warrens have been established at a great variety of sites. At Pompocali, near Bardsey
(Henderson, 1979) an old stone quarry on the Carlton Grits is named as Coney Garth
Plantation on the map accompanying the Enclosure Award, and may be an instance where
spontaneous rabbit colonisation invited the installation of a managed warren by the local
landowner or tenant farmer. Earlier this century unemployed miners at Wilson Wood
Colliery, near Clapham, created a rabbit-warren (Phil Hudson, pers. comm.) by digging out
slag-heaps, inserting pipes and bringing in rabbits. The artihcial tunnels can still be seen,
though in some disrepair, and rabbits are still active on the site. One of the most
unexpectedly located of mediaeval warrens is mentioned in an accounts list of 1453-1454
(Morton, 1954) for Peterborough monastery: 'Item quatuor cunicitli de cimeterio Saiicti
Johannis Baptistae' (Item: four rabbits from St John’s graveyard), indicating the existence
of a warren in the church graveyard, now St John’s Close.

Rabbit and vermin trap.s

Specially designed traps for rabhit and vermin were made in some regions. Tip-traps
(known as ‘types’), first introduced in the late 18th century (Eus.sell, 1955), have been fully
de.scribed from the North Yorkshire Moors (Harris & Spratt, 1989), where the same term,
‘type’, is also used to designate the small stone-walled enclosures in which .some pit-traps
are located. Several types have been located on Bolton Abbey estate (detection of the first,
according to reliable anecdotal account, involved the successful use of the divining rod!).
In Strid Wood the types, sited immediately by .stone boundary walls, are associated with
small muce-holes or passages through the wall. Muce-holes (al.so known as ‘smouse-’,
‘smoose-’ or ‘smoot-holes’) are small, square, rectangular or round passageways through
the base of a wall, allowing rabbit transit, at Strid Wood guiding the rabbit into a narrow
tunnel with a trap-door which, when .set, plunged the rabbit into the pit below. Such pits
varied in size and construction, the smallest seen to date being at Cunnery Wood, Shibden
Hall, Halifax, where the width of the circular aperture is about 0.6 m. The sides of this pit
slope conically outward towards the base, as in types on the Yorkshire Moors. Types seen
at Bolton Abbey have rectangular pits with perpendicularly upright sides and the pits
approach 1 m in depth and width. Larger still are known, and rounded pits are not
uncommon. Muce-holes unassociated with types were used to encourage rabbit movements
during netting and other trapping.

Vermin traps developed specialised forms in some regions. Those on Dartmoor have
been de.scribed by Haynes (1970). There, in general, vermin were led into hutch traps by a
system of funnelling walls. At Headland Warren Dartmoor, traps were found without any
such guiding walls, relying on chance, baiting or strategic positioning by muce-holes in
enclosure walls.

Warreners, gamekeepers and poachers

Warreners, gamekeepers and the motley individuals who stole from them used similar
methods to take their quarry. A notice of appointment of gamekeepers for his manor of
Aldburgh from John Wentworth of Woolley, near Barnsley, summarises the methods and
implements used (Lawson-Tancred, 1937). It authorises the appointee ‘to take and levy for
my use all such guns, hows, grayhounds, setting dogs, lurchers or other dogs to kill hares
or coneys, feretts, tramelles, low bells, hayes or other nets, snares or other engines for the

h orn Coney to Rabbit: the Story of a Managed Coloniser 1 1 I

taking and killing ot coneys, hares, pheasants, partridges and other game as shall be used
within the precincts ot the said manor by any person or persons not having lands or
tenements or some lorm ot inheritance, in his own or his wife’s right, or having lease or
leases of 99 years or for any longer term of the clear yearly value of £150, other than the
son and heir of an Esquire or other person of higher degree.’

This list ot equipment is not radically different from that described in the 14th-century
journal of the French nobleman. Sire de Gouberville (Delort, 1978): "The coney hunt is
engaged in by the iKrbility to the same degree or more than the lower estate. The nobility
make use tor such a chase of spaniels, likewise small greyhounds. One group of the animals
is taken "in course” by last hounds. The others shelter in their burrow's. The hunters then put
purse nets over the mouths of the burrows, except for one through which they put in a
muzzled lerret or one with its teeth cut down (so that it doesn't become drunk with blood) or
then they light a fire of sulphur, orpiment and myrrh on the inside of a cloth or parchment
and they throw it into a vent-hole. The rabbits emerge en masse with the draught and are
caught in the nets. In 1479, six people (including the king’s jester) hunting for 2'h days and
3 nights were able to catch 72 rabbits, using 3 I'errets. For the poor, a different smoke
mixture took the place of dogs and ferrets, i.e. old woollen rags, parchment with sulphur,
orpiment and myrrh, powder in a pot provided with an iron pipe, branches of broom, linseed
and charcoal.’ Rabbits would be taken abundantly by such restrained methods and by
netting, ensuring that the fur was completely undamaged.

Poaching

Already by John Lydgate’s time, the 15th century, rabhit poaching was commonplace
enough for him to use it in his long poem. The Pilgrimage of Man. as descriptive of
Avarice’s second hand:

‘Thys hand, thogh men hadde sworn.

Kan robbe and bern away the corn
Out off bernys and garnerys;

Thys hand kan ferette in konnyngherys
Be nyhte tyme, whan men slepe;

Thys hand by hofys kan in crepe
And bern a-way what he may fynde.

And lyst to leue nothyng behynde.’

The history continues consistently as far as poaching is concerned. When Charles
Fothergill pas.sed through North Dalton in 1805. his diary records his satisfaction at finding
‘the whole country, tho so wild and open, entirely enclosed from Pocklington to Driffield
(Fothergill, 1984). Even the rabbit warrens near Driffield, which last time 1 was here were
tilled with rabbits and heath, now exhibited fields of waving corn’. Not owing to increase
of prolit by growing corn, ‘but a gang of poachers, amounting to 26 in number, having for
several years past committed so many and continual depredations insomuch as to consume
two thirds of the rabbits, it was thought proper to destroy the remainder and turn the
warrens into arable land, notwithstanding the ring-leaders of poachers were taken and tried
at the lent assizes when they were transported for 7 years'.

Dlspute.s

Encouragement by landowners and warreners and a resulting backlash from neighbouring
tenants and other affected locals have been a persistent feature of the rabbit's existence
since it became established here in significant numbers, and the animal has been a frequent
cau.se of .social and legal disputation. For instance, towards the close of the 15th century
during the reign of Flenry VII. amongst several depositions laid against Sir Roger Flastings
in the Pickering area (Turton. 1894). was the charge ‘that he of newe hath enclosed within
the fields of Thornton and Farmonby ii acres and more of arrabill grounde and annexed it
to a Cunnygerthe. to the gret hurt and distruccion of the King's tenantes and inhabitauntes
of Thornton and Farmonby’,

1 12 From Coney to Rabbit: the Story of a Managed Coloniser

Equally annoyed were the townspeople of Liscard. Cheshire, in 1754, when they arrived /
at what they termed a Rabbit Agreement against the local landowner, John Gough (Woods ;
& Brown, 1960). ‘Be it known to all men whom it may concern, they whose names are ;
here under subscribed do covenant, agree with one another to destroy all the Rapits on the i
Warren or Coney Borrow on the Common. Whereas John Hough do claim a property ot the I
Rapits in that Warren, which we hereafter mentioned do not allow the said John Hough to i
have . . . etc.’

Feelings could become even more heated. So, in the mid- 18th century, John Minshull .
(17--) of Sunderlandwick posted up a notice to his neighbours (now in the East Riding
Records Office) airing his grievances: This is to give Notice that upon Complaint of
Damages by my Tenants and Neighbours which the Rabbits or otherwise Conies has made u
upon the Ground or Premisses of the aforesaid Tenants and Neighbours . . . Now I do hereby j
certify that 1 shall not pay one Farthing for the Damages . . . And for the Insult of my i
Tenants in Complaining to Me for Satisfaction 1 am fully Determin’d to take the Ground out i
of their Possession and let it to the best Bidder who will promise not to make any Complaint i
upon the like Occassion [sic]: And further, if any of the Tenants should take Kill or Destroy 1
any Conie or Conies upon their Ground or Premisses either with Guns dogs Nets Snares or -
any other Invented Engine or Engines whatsoever shall suffer according to law as in that <
case. Item I do here [s/c] there is a Grumbling and Complaining with the Neighbours around (
me about their Dogs being Killed such as Mr Dowbegin for the loss of Captain; Robert jj
Tateson and Richard Gunlass for theirs: and William Chambers two Dogs . . . these two last II
dogs I suppose escaped only with peppered Arses . . . These complaints are insufferable and
a man of any Spirit cannot help resenting such ill Treatment from Neighbours: I therefore i
but not in any Passion tho Just Reason from the aforesd; Complainants or Complainers do >
affirm that I shall or one by my order: Take Kill or Destroy: any kind of dog or dogs . . . that I
shall go ten Yards out of the Road to wag his tail or Piss upon Sunderlandwick Moor. Item ..
alledging yet more if I cannot . . . destroy the aforesd; dog or dogs Neither with Guns Traps
or Person, I shall for these dogs Trotting over or Pissing on my Grass Prosecute the J
Proprietor ... as far as law will Admitt of — John Minshull.

Four north-eastern English warrens

/. Holy Island

Today, rabbits are found on 700 of the world’s islands (Flux, 1994) between latitudes 60°N
and 55°S, over 300 of them being around the British Isles. Monks, sailors, island farmers, 5
lighthouse keepers and others have contributed to this distribution. Monastic orders may (
have found the introduction particularly desirable and satisfying since the Church allowed ■'
the consumption of rabbit meat at times when only hsh was permissible, since young
rabbits, not long from the womb, were regarded as aquatic! (Biadi & LeGall. 1993).

The island nearest to the Yorkshire coast with an early warren is Holy Island; on the map
(Fig. 10) it resembles a Liebig’s condenser with the long neck generously Jammed at the
mouth. This long-necked peninsula, the Snook, and the northern coastal stretches to the
east of it are the sandiest parts of the island, useless for most crops, but an ideal location for
a rabbit-warren. The earliest warren reference traced so far for Holy Island is in 1377
(Raine, 1852) when it had a valuable rabbit-warren belonging to the see of Durham. The
Bishop of Durham shortly afterwards caused tithe of the rabbits there to be paid to the Prior |
of Holy Island, and in 1438 Thomas Monckton was appointed Keeperc of the Warren for
life, receiving an annual fee, paid by the farmers of the warren, and ‘a robe of the suit of
our esquires’, with the concomitant duty of supplying the Bishop’s table with rabbits as
required. In 1528 a 12-year lease of the rabbit-warren and borough of Holy Island was
charged at £4 yearly the warren and £3 yearly the borough. In 1739 there was a lease to
Jane Selby of Holy Island of the ‘rocks, ware, kett l.v/c | or seaweed in Holy Island, the
coney warren, the tithes of fish’, etc., and the harbour dues, rents or customs paid over nine
years, for £105 per annum. By 1759 the lea.se had increased to £130.

Frotti Coney to Rabbit: the Story of a Managed Coloniser

113

FIGURE 10

Map of Holy Island. Based on a map ©Frank. Graham.

The areas of the Snook and Links were described by Winch in 1822: ‘The uncultivated
part of the island is sandhills or links slightly held together by the creeping roots of the sea
lyme-grass, sea mat-grass, rushy wheat grass and sea carex, and is occupied as a rabbit-
warren.’ Another description reads: ‘Without woods, moorlands or rivulets. Lindisfarne
possesses a .scurvy flora. Cryptogamic plants are peculiarly scarce, with the exception of
the marine algae’. On the links are listed: Flat-headed sedge. Grass of Parnassus.
Brookweed, Seaside Centaury, Bloody Cranesbill, Common Storksbill. Scarlet Pimpernel
and Blue Fleabane. and the lone lichen listed from this part of the island, 'baeomyces
alcicornis' (= Cladonia foliacea).

Some ecological pros and cons of rabbit-grazing have been summed up with regard to
the smaller Fame Islands to the south, where rabbits seem to have been brought in by the
lighthouse-keepers as a handy food supply (Hirons. 1994). ‘Rabbits can have a beneficial
effect on the vegetation. They help maintain floral diversity by removing tall grasses, they
produce lawn-like areas of turf. Their wastes help fertilize the soil and disused rabbit-
bumows can be used by puffins for nesting. Their harmful effects are most conspicuous
when numbers become high: damage to vegetation by overgrazing and trampling; the
young shoots are damaged ... By eating seedlings rabbits prevent many species becoming
important components of the vegetation cover. By scraping and burrowing, the soil is
exposed and desiccates’.

2. Holderness

In coastal mainland regions, too. many areas of dune, sandy links and lowland were
favoured warren sites, for instance, the chains of warrens along the Lancashire coast, and
along the East Anglian sandlings. Some of Yorkshire’s earliest warrens were on the
Holderness lowlands in the region of Spurn’s predecessor. ‘Aide Ravenser’. A royal
mandate of 1303 (Cal. P. Rolls 1301-1307) empowers ‘Richard Oysel and Ralph de Lellay
to enquire touching malefactors in the King’s free warren at Brustwyk. Preston.
Kayngham, Little Humbre. Burton Pydese, Skipse. Esington. Skettling, Kilnese and

I 14 From Coney to Rabbit: the Stoiy of a Managed Coloniser

Wythornese; persons who carried away hares and rabbits from the said warren’. A royal
writ of 1306 to the same Richard Oysel, seneschal, states: "The King has two coneygarths, >
that is the west coneygarth, and the east coneygarth in Esington and Kilnese by Aide
Ravenser on Humber, with warren in the same. The King gives to the warrener keeping the
great west coneygarth, 26s 8d a year, that is 6d a week, and 6s 8d yearly for his robe; and r
to a lad (garcioni) keeping the small eastern coneygarth, 13s a year, that is 3d a week.’ i
Quite how much earlier rabbits were stocked in Holderness remains doubtlul, but among |
those assisting the steward and sheriff, as early as 1 150, were warreners and falconers with 4
sport in their care. In 1214 (English, 1979) there is mention of a warrener, Augustine, and a
little later of a staff including four warreners; while in the sheriff’s accounts of 1261-1264
the keeper of the warren receives the same 6s 8d for his robe as mentioned above. A late
13th to early 14th valuation of Kilnsea (at £30 to £40 net) says that £2 of this was for the
herbage of the warrens. Beside this herbage rent, money was received from rabbit sales !
(292 in 1296-1297; but only 38 in 1348-1349 when manorial income was reduced by the
sea encroaching). Crackles (1993) points out that it is not clear whether the warren herbage ,
consisted of Corse or Sea Buckthorn or both: and notes a bylaw at Keyingham in 1637 '
stipulating that the inhabitants should ‘dike and beard their land with thorites, furres or
such like’. The warren at Kilnsea was not closed down until the 18th century.

J. Woodluill Warren

Woodhall Warren, near Carperby, also variously named as Nappa Warren and Lady Hill 1
Warren, is situated on the north bank of the Ure, just west of Aysgarth (Eig. II). It I
occupies a glacial area containing a 38 m high drumlin. Lady Hill, near its centre i
(Yorkshire Dales National Park, 1996). To the north runs the Askrigg-Caiperby road. In
1251-2 (Whaley, 1890) Henry II made a grant of free warren at Woodhall (such a grant
conveyed the right to hunt all small game, such as foxes, hares, pheasants, partridges, i
squirrels, rabbits, etc., but is, of course, not per se proof of rabbit presence in the locality at
the time of the grant).

In 1 890 Whaley described the warren of siver-grey rabbits then operating at Woodhall as
very ancient, but a plan of 1756 which matches the area of the warren, gives no indication i
of warrening activity. Jeffery’s 1771 map marks the area as warren, but without any I
marking of boundaries. However, Alexander Fothergill in his diary writes on 21 February
1757: ‘. . . to Woodhall Park where Robert Dent and Robert Durham met mee to take
Hutchinson House to rebuild for the warrener’ (Fothergill, 1985). This is the house now
known as Warren House, situated in Hutchinson Close, and it appears that warrening may
have begun at Woodhall some time earlier. All the references traced would fit in with
Hartley and Ingilby’s statement that it was probably started by Thomas Metcalf of Nappa
Hall, who lived from 1687 to 1756. On Metcalf’s death there was considerable change on
the Woodhall estate.

On 13 May 1756 Fothergill viewed the premises at Woodhall Ings to divide them
between George Metcalfe and Solomon Beezon. Almost twenty years later he records
obtaining a rent of £19. 7s from John Dinsdale for the Woodhall Park (so called), noting
that they want a good wall at the bottom of the Intake that they may propagate a breed of
rabbits there to keep down the whins, etc. On 28 Octttber 1774, about eight months later,
he records the same rent collected from John Dinsdale for Woodhall Warren. The 1856
O.S. map shows the warren extending nearer the bank of the Ure than on the 1913 map,
owing to construction of the railway along the river’s north bank. In 1894 James Storey
moved into Warren House with his family, taking charge of the warren in addition to his
role as keeper for the Vyners of Newby Hall, to whom ownership of the warren had passed
by marriage. The warren was famous for its valuable breed of silver-greys. Tuke (1794)
suggested they were brought from a Lincolnshire warren, having come there from Ireland.
Up to 1936 they were being sold to the felt hat industry. Anecdotal account now is that
many were .sold alive as breeding stock. Whaley (1890) tells of them being .sold for
coursing. Further back, in 1774 Fothergill records some concern over the possibility of live

Askrtgg to Cirperby road

I

From Coney to Ruhhit: the Story of a Managed Coloniser

115

FIGURE I 1

Map of Wooclhall Warren, North Yorkshire. Based on a map produced for the Yorkshire Dales National Park, and on O.S. of the area.
©Crown copyright; ED2743.1X. Key: 1-6 = rabbit types or tip-traps; 7 and 8 = buildings.

I 16 From Coney to Rabbit: the Story of a Managed Coloniser

rabbits being sold, and mentions an agreement that this should not be done by the Nappa
tenant of that time.

The warren was entirely enclosed by a 6' to 8' high stone boundary wall, its only gate
being one down by the warren house. Rabbits were taken either by type or netting. Six
types have been located within the warren, immediately by or somewhat subsumed into
inner field walls. Both methods of capture would avoid damage to the animals’ fur. A
distinctive feature of the northern boundary wall is a number of inset alcove-like vermin
traps, situated high up along the wall's inner face at intervals. On the fiat bases of these i
alcoves circular traps were placed so that when a marauding fox or stoat attempted to gain
exit from the warren, they were attracted by the step-like aperture below the coping. None
of the traps used in this specialised situation has been traced as yet, but they are known to i
have been broadly disc-like and to have been fastened by a metal chain to the mid-reach of i
the wall below the alcove, so that the victim was often left cruelly dangling. A small I
enclosure by the eastern boundary wall was used for growing turnips as rabbit-fodder, and |
these were stored in a small shed in the corner of the held. «

Warrening continued after the decline of the very prohtable siver-grey industry. Methods
of capture changed, however, when intact fur was no longer an overriding priority.
Ferreting became acceptable, and snaring replaced the use of the types, which had always
been labour-intensive. The Vyners disposed of the estate in 1933, and gave hrst chance of
buying to Robert James Raw, who obtained it for £775. The declining years of warrening
activity are fortunately documented in the privately owned diary of Francis Thornborrow
Dinsdale (19--). Between 15 September 1953 and 12 January 1954 he sold 1341 and a half
couples of rabbits, when they were selling at 5s 6d each. In April 1955 he notes that the
man who bought most of the rabbits paid 4s a couple in February, 3s. 6d a couple in March,
and only 2s 6d a couple in April. His first record of the advent of myxomatosis into the

warren is at the start of that year, 29 January 1955 : ‘Found one rabbit in warren

home field near henhouse.’ Among a few such subsequent entries: ‘13 March 1955. Saw

another rabbit under Stoneman (not in the warren) with m ('A ml due n. of warren)’.

The last reference of all to the warren is on 2 April 1955: ‘Ferreting at west side of big hill
at west end of north side. Would not bolt, but got 14 rabbits’.

4. Wharnciiffe Chase

Wharncliffe Cha.se was in mediaeval times an unenclosed chase (Hey, 1975), with free
warren granted in 1242. It was enclosed in the 1500s and a lodge was built on the crags in
1510, ultimately with kennels adjoining. From the 16th to mid- 1 9th century deer were the
main stock. A charge from the hnal years of the 16th century (in the rough Latin of the
day) against Reginald Thompson, labourer, states that ‘he illegally broke into and entered a
cha.se of Richard Wortley and unjustly and illegally without any authority or licence killed
and did to death cennon in English a deer tormento in English with a Gunne’. Erom 1820
sheep were kept and subsequently became the main concern. The earliest record traced so
far of a rabbit-warren on the chase is an area to the east named as ‘warrens’ on a 1774 map
in the Wharncliffe Muniments and under a tenant farmer, Robert Charlesworth. A
charming 18th-century map of the chase is uninformative about warrening activity.

Rabbits on the parts of the chase where there is most remaining evidence of warrening
were, it appears, primarily bred to provide interesting days hunting and shooting. They and
the sheep farmed with them assumed greater importance as deer-chasing declined. Today
four areas of the chase hold signs of warrening activity (Eig. 12). The oldest is the earthen
boundary embankment of the tenant farmer Robert Charlesworth’s ‘warrens', already |
mentioned, to the eastern side of the chase, now known as Warren Plantation and
overgrown. The second is a large area to the north of the chase, where there are several
pillow mounds which are of distinctive and substantial structure (Fig. 13). Square or
rectangular in plan they had sides 10 m or more in length, c. 1.5 m in height, revetted in
well-dressed stone, and with two or three small nuice-holes in the foot or side of each stone
face, allowing entry to what appears to have been a two-storey system of stone-and-earth

From Coney to Rabbit: the Story of a Managed Coloniser

1 17

FIGURE 12

Map of Wharncliffe Chase. Based on a map produced by the Ecological Advisory Unit,
Sheffield County Council; and on O.S. of the area. © Crown copyright; ED27433X.
Key: 1-6 = revetted earth-&-stone pillow' mounds (see Fig. 13); 7 = earthen feeding
mound; m indicates lengths of wall with muce-gates and muce-holes (see Fig. 14).

118

From Coney lo Rabbit: the Story' of a Managed Coloniser

n

f';

FIGURE 13

Remains of revetted pillow mound, Wharncliffe Chase. Photo; A. Henderson.

fashioned burrows within, intended to encourage rabbit habitation and further spontaneous
burrowing on the animals’ part. The overall capping of each mound was completed by
0.35 m-deep turve-layering. A nearby stretch of the western boundary wall has a well
preserved course of shallow flat throughstones projecting on the inner face Just below
coping level, originally aimed at preventing deer jumping to escape. The third area of
rabbiting interest is in the central area of the cha.se, where there is a large, burrow-pocked,
earthen mound c. 12 m x 15 m in rectangular plan, which was a feeding mound for rabbits,
where turnips, etc. could be laid out as fodder. This mound has lost the surrounding iron
rails, c. 2 m high, which once protected the rabbits’ feed from intrusion by the sheep and
any remaining deer on the chase. Lastly, over by the Lodge and towards Wharncliffe Farm
is an area where activity was directed mainly at the provision of pleasant days in the field
with the gun. Here rabbits were run to and fro for the sportsmen, additional stock being
transported from elsewhere on the estate in steel box-traps on shooting days. Remaining
evidence of this sport is in the two strategic lengths of field-wall in this neighbourhood
which contain, interspaced, four muce-holes (cylindrical pipe-lengths through the base of
the wall) and most unusually seven muce-gates (Fig. 14). These are low gates through the
wall, typically c. 0.9 m wide by 0.45 m high, the wall above them supported by two stout
oak beams almost I m long and 15 cm .square in section. These finely crafted gates show
the degree of importance attached to producing a good show of rabbits for the guns.

Conclusion

Since its establishment here as the Normans’ coney, Oryctolagns ciiniailiis has moved
gradually towards the nearly totally feral existence it enjoys nowadays. Its history here is
that of a peculiarly restless ecological entity; more so than in most mammals because of its
extreme opportunism, its ecological pla.sticity and its fecundity. It was brought to these
islands as a valuable item, the Normans recognising its desirability not only for the table
and clothing, but also as source of profit in home and ovcr.scas commerce. Many of the
vicissitudes in its history have been closely connected with man. For instance, the
establishment of feral colonies by escapees from warrens at ihe lime of the Black Death
was favoured by the diminution of custody and husbandry in warren and field. The
Agricultural Revolution of the 18th century, with new methods of land fertilization and

I- ram Coney to Rcihhit: the Story of a Mcnuiged Coloniser

119

FIGURE 14

Miice-gatc and miice-holc, Wharncliffe Chase. Photo: A. Henderson.

with winter fodders available, meant more land was rendered arable and stoek eoiild be
overwintered; by the 19th century there was therefore less demand for the rabbit as food.
Eneoiiraged by changes in the game laws, however, feral rabbits quiekly began to assume
pest proportions. By mid-Victorian times the great commercial warren complexes were in
decline; the coney of the Normans had become the British rabbit. Waiting in the wings
were the horrors of myxomatosis and haemorrhagial virus.

Rhi-hrences

Apicius. De Re Coquinaria 2, 2. 6.

Biadi, F. and Le Gall, A. ( 1993). Le Lupin de Garenne. Hatier.

Bdkdnyi, S. (1974). History of Domesticated Mammals in Central and Eastern Europe.

Akademiai Kiado, Budapest.

British Library. Harley. MS 2278 67. v.

Calendar of Close Rolls. Henry III. (1237-1242).

Calendar of Patent Rolls. Edward /( 1 30 1 - 1 307).

Catullus. Carmina 25, and 37.

Crackles, F. E. (1993). The story of gorse in Holderness. Holderness Countryside 39:12-
13.

Cunliffe, B. ( 1983). Anatomy of an Iron Age Hillfort. Batsford. London.

Delort. R. ( 1978). Le Commerce des Eourrures en Occident d la Fin du Moyen Age (vers
I SIX) _ yers 1450). Bihliotheque des Ecoles Francaises d'Athenes et de Rome 236:1-
1383.

Dinsdale. F. T. (19--). Diaries (privately owned).

East, R. (1891). E.xtracts from the Portsmouth Records. Portsmouth Corporation.
Portsmouth.

English. B. (1979). The Lords of Holderness 1086-1260 (University of Hull Publication).
O.U.P.. Oxford.

English Heritage. National Monument 17159. Schedule Entry Jan. 23, 1997.

120

From Coney to Rabbit: the Story of a Managed Coloniser

Finch, M. E. (1956). The wealth of five Northamptonshire Families 1540-1640. 1

Northamptonshire Record Society 19:1 -246. I

Flux, J. E. C. (1994). World distribution. In The European Rabbit (H. V. Thompson and I
C. M. King, eds): 8-21. O.U.P., Oxford. '

Fothergill, A. (1985). Alexander Fothergill and the Richmond to Lancaster Turnpike Road i|
(M. Hartley, J. Ingilby, D. S. Hall & F. P. Wenham, eds) North Yorkshire County Record I)
Office Publications 37: 1 -27 1 .

Fothergill, C. (1984). A Diary of Charles Fothergill 1805. Yorkshire Archaeological i
Record Series 142: 1-28 1 .

Fussell, G. E. (1955.) The coney warren a century ago. Yorkshire Life Illustrated (Jan, i
1955): 31.

Gilbert, J. M. (1979). Hunting and Hunting Reserves in Medieval Scotland. John Donald, l|
Edinburgh. j

Harris, A. (1990). The rabbit warrens of East Yorkshire in the 18th and 19th centuries. •
Yorkshire Archaeological Journal 62:429-443. J

Harris, A. and Spratt, D. A. (1989). Rabbit warrens on the Tabular Hills: archive of field ^
and documentary work 1987-1988. W.Y.A.S. MS1523.

Harris, A. and Spratt, D. A. (1991). The rabbit warrens of the Tabular Hills, North
Yorkshire. Yorkshire Archaeological Journal 63: 177-206. '

Hayes, R. J. (1965). Manuscript Sources for the History of the Irish Civili.sation. G. K.
Hall, Boston.

Haynes, R. G. (1970). Vermin traps and rabbit warrens on Dartmoor. Post Medieval
Archaeology 4: 147-164.

Henderson, A. (1979). Pompocali, a topographic curiosity. Naturalist 104: 75-87.

Hey, D. (1975). The Making of Modern Yorkshire. South Yorkshire County Council,
Sheffield.

Hirons, M. J. D. (1994). The flora of the Fame Islands. Transactions of the Natural History
Society of Northumbria 57: 69-1 14.

Hurrell, H. G. (1979). The little-known rabbit. Countryside 23: 501- 504.

Hyland, P. (1978). Purbeck, the Ingrained Island. Gollancz, Fondon.

Lawson-Tancred, T. ( 1937). Records of a Yorkshire Manor. Edward Arnold, London.

Minshull, J. (17--). Sunderlandwick: Notice of John Minshull. Held in the East Riding of
Yorkshire Archives Office, Beverley. DDGR 20/1.

Moorhouse, S. (1991). Norton Tower, Rylestone, North Yorkshire. C.B.A. Forum 1991.
Annual Newsletter of the CBA Group 4.

Morton, W. (1954). The Book of William Morton, Almoner of Peterborough Monastery
1448-1467. Publications of the Northamptonshire Record Society 16.

Oxford, M. J. (1986). Planning for Conseiwation of Wildlife and Archaeological Features
on a Nature Reserve in Britain. M.Sc Thesis, University of Calgary, Alberta.

Pliny the Elder. Natural History 8: 217; 226.

Sheffield City Archives ( 1774). A Map of Wharncliffe Moors. Wharnclijfe Muniments TAN
3L.

Raine, J. (1852). The History and Antiquities of North Durham. J. Bowyer Nichols,
London.

Spratt, D. A. (1989). Linear Earthworks of the Tabular Hills, North-east Yorkshire. J. R.
Collis Publications, Department of Archaeology and Prehistory, University of Sheffield,
Sheffield.

Strabo. Geography 3, 2, 6; 3, 5, 2.

Stuart, A. J. (1982). Pleistocene vertebrates in the British Isles. Longman, London.

Tuke, J. ( 1794). General View of the Agriculture of the North Riding of Yorkshire. Board
of Agriculture, London.

Turton, R. B. (1894). The Honor and Forest of Pickering. North Riding Records. New
Series 1:1-280.

Varro. Rerum Rusticarum 3, 12, 6-7.

121

The Horse-trough Ostracod Heterocypris incongruens

Warner, G. F. (1912). Queen Mary’s Psalter. Brotherton Library, Leeds. Speeial
Collections Art E-8q. Royal MS 2B VII in B.M.

Whaley, C. ( 1890). Parish of Askrigg. Skeffington, London.

Williams, S. W. (1895). Tiles found at the restoration of Bangor cathedral. Archaeologia
Cambrensis [5th Series j 1 1 : 107- 1 1 I .

Williamson, T. and Loveday, R. (1988). Rabbits or ritual? Artificial warrens and the
Neolithic long mound tradition. Archaeological Journal 145: 290-313.

Winch, N. J. (1822). Remarks on the geology of Lindisfarne or Holy Island. Annals of
Philosophy {December 1822): 169.

Woods, E. C. and Brown, F. C. (1960). The Rise and Progress of Wallasey 2nd edition.
Wallasey Corporation, Wallasey.

Yorkshire Dales National Park (1996) Woodhall Rabbit Warren, Carperby, North
Yorkshire. Archaeological Survey (E. Denison).

Zeiiner, F. E. ( 1963). A History of Domesticated Animals. Heinemann, London.

Acknowledgements

I am especially grateful to the staffs of the British Library, the Brotherton Library', the East
Riding Record Office, English Heritage, the National Commission on Historic Monuments,
the Office National de la Chasse, Sheffield County Council and Archives, and the Yorkshire
Dales National Park, for the provision of information and the waiving of several copyrights,
as well as other archive and record offices, local historians and archaeologists too numerous
for individual mention. I am indebted to many landowners for access to sites, and would like
also to thank all colleagues and friends for information and for help in the field.

THE HORSE-TROUGH OSTRACOD
HETEROCYPRIS INCONGRUENS

GEOFFREY FRYER

Institute of Environmental & Natural Sciences. Lancaster University, Lancaster LAI 4YQ
Introduction

The freshwater ostracod Heterocypris incongruens (Ramdohr) (= Cyprinotus incongruens),
a minute bivalved crustacean, has an apparently almost cosmopolitan distribution. It is
common in Britain and elsewhere in Europe. Ecologically tolerant, it favours small water
bodies of various kinds and has such a liking for horse-troughs and similar receptacles - a
liking seldom shared by other ostracods - that it merits the common name of the horse-
trough ostracod. Such a designation may serve to draw attention to this interesting
crustacean, to some of the phenomena that it displays, and to the intriguing problems that it
poses.

The animal and its lifestyle

Only females (Figs. 1 and 2), are known in Britain, where reproduction is entirely
parthenogenetic. They may attain a length of about 1.8 mm but are frequently smaller.
Yellowish in colour, they often become dark reddish brown dorsally as they get older. The
ovaries lie between the inner and outer lamellae of the carapace on each side and contain
orange eggs that can be dimly seen through the chitinous outer lamella. The surface of the
carapace (‘shell’) is smooth and polished. H. incongruens is an active animal that can swim
briskly by means of its antennules and antennae which can be extended outside the
carapace and are provided with long natatory setae (‘swimming bristles'). It can also crawl
by use of the antennae and first pair of thoracic limbs, both of which terminate in stout
claws. As a means of defence it can bring together the valves of its carapace and shut itself

Naturalist 122 (1997)

122

The Horse-trough Ostracod Heterocypris incongruens

FIGURE 3

Part of a cluster of eggs.

within like a tiny mussel. An omnivore, it can subsist on algae and organic detritus and can
also skeletonize leaves by rasping their .soft cells - lettuce leaves are rapidly degraded if a
large population is at work. It exploits windfalls of dead animal matter, and when it can
seize them, which it is not particularly well equipped to do, it kills and devours such small
animals as Daphnia.

Like many freshwater ostracods, but unlike most crustaceans, it attaches its bright orange
eggs to firm substrata, such as a piece of rotting wood, a dead leaf, or stone surface. This
applies to all eggs save the first batch produced, which are (always?) attached to the
carapace and cast off at the final moult, and to a few that may be shed free. The
subspherical eggs (Fig. 3), which have a tough, protective outer pellicle and two inner
membranes, are mostly deposited in clusters, often in parallel rows. A lifetime's production
may be up to at least 100 eggs, laid in small batches, but not all individuals are so
productive. Several animals often lay eggs in clo.se proximity and veritable crusts of eggs
can then be seen with the naked eye. These are usually one layer deep, but overlaying
sometimes occurs. Veins are attractive when dead leaves are used, and when only these
veins remain they often bear ‘ropes’ of eggs.

An egg hatches as a minute nauplius, the basic crustacean larva, hidden by a bivalved
shield. This has led to erroneous statements that ostracods are born as 'juveniles' not

123

The Horse-troitf’h Ostrucod Heterocypris incongruens

larvae. Alter passing through 8 larval and pre-adult stages, adulthood is reached, which,
depending on temperature, takes between about 5 weeks and 5 months. Egg laying begins
with the eighth moult. Adults may survive for several months.

An inordinate fondne.ss for horse-troughs

A preference for horse-troughs and similar receptacles is a noteworthy and well-known
feature of the ecology of H. incongruens. For example, although such troughs were not
specilically sought during a survey of the freshwater Crustacea of Yorkshire, it was found in
six, scattered over a wide area and, atypically, in a spring-fed metal trough supporting a
growth of charophytes (Fryer, 1993). Ostracods found in a wooden tub when 1 was a small
boy were certainly this species, which I have encountered in similar troughs elsewhere in
England, e.g. in Cumbria, Cheshire and Gloucestershire. Troughs are often, but not always,
turbid, green with algae, contain dead leaves or other decaying organic matter, are
sometimes foul, and must often suffer from oxygen deliciency. Indeed. Fox and Taylor
( 19,3.3) demonstrated that II. incongruens can not only Hourish at low oxygen concentrations
but that it prefers such conditions to well aerated water. Green ( 1939) showed it to be well
adapted to situations low in oxygen and even to be able to endure anaerobic conditions for at
least two weeks, and to lay eggs while so doing; he also found that it produces a yellow
pteridine pigment which he suggested may play a part in intermediate metabolism under
anaerobic conditions. Nevertheless, individuals often congregate near the surface, where
oxygen is most plentiful, and lay many of their eggs there.

Conditions differ much in different troughs. In Cumbria I have found H. incongruens at
Witherslack, on the Carboniferous limestone, in a trough made from limestone blocks,
somewhat less than 3 m- in area and up to almost half a metre deep when full. This is
clearly alkaline, is usually green with suspended algae, often foul, and newts breed in it.
Different conditions prevail in a smaller, glazed ceramic horse-trough near Lazonby that
intermittently receives clear water from .sedimentary Permian rocks. Different again is a
garden trough at Grasmere, less than 0.3 m- in area, of local volcanic rock, containing
clear, perhaps slightly acidic, water. However, like most ostracods, H. incongruens shuns
very acidic water, perhaps becau.se of the difficulty of Hnding enough calcium with which
to impregnate its carapace rather than avoidance of acidity as such. It was not seen in the
Southern Pennines during an exploration of stone troughs that receive cold springs. These
are modihed rheocrenes. sometimes have a carpet of bryophytes. and harbour a variety of
cold-loving micro-crustaceans. Here it may be deterred by cold, acidity, low calcium
levels, well oxygenated water, or a combination of these factors. It has, however, been
recorded in springs in Europe (see Klie (1938) for summary) though these are perhaps
largely limnocrenes (seepages), where the water spreads out and may quickly warm up
after leaving the ground. Indeed, this is precisely how Bronshtein (1947) describes its
occurrence in limnocrenes, where he specifically states that it lives 'far from the outlet of
the spring, and also well heated'. The spring-fed Yorkshire trough is an unexplained
e.xception.

A liking for house-troughs and similar habitats is shown by finds in the Channel Islands,
themselves isolated and not easy for freshwater animals to find. In Guernsey. 1 found it in
small stone troughs at St. Peter Port and Saumarez. Even more striking, on the much
smaller island of Sark 1 found it in three such troughs, one very' small and containing only a
few centimetres of water and. as troughs unfortunately now often do, much rubbish. Most
remarkable of all, H. incongruens has been found on the small, remote Easter Island in the
Pacilic Ocean (Dumont & Martens. 1996). being one of only 3 micro-crustaceans found
there, of which 2. possibly 3. of the others are suspected to be introductions. Of the several
sites in which it was found, one was ‘a stone basin' near the shore, another 'a eutrophic
stone basin used for watering horses'! How H. incongruens locates such sites is a mystery
to be considered. It has also been recorded in water-filled tree holes (Klie, 1938).

H. incongruens also occurs in what might be regarded as natural equivalents of horse-
troimhs In Northern Manitoba. Canada, it is common in rock pools that occupy small

124

The Horse-trough Ostrucod Heterocypris incongruens

depressions on quartzite bluffs. Of 157 sampled pools of mean area about 1 m^, it was
found in 67 (Havel et ai, 1990). Likewise Canning ( 1971 ) described a rock pool in Sweden
which supported a persistent population of H. incongruens as ‘a very typical biotope’ for
this species. This had a maximum volume of about 300 litres and often dried up
completely. Other sites are extremely diverse but seemingly always small, except perhaps
shallow rice-fields. Ponds and pools, including foul and turbid farm ponds, ditches and
muddy seepages are the sort of places favoured.

Eggs for all occasions |

In Britain all eggs are produced parthenogenetically. No male has ever been recorded and |
this is so for much of Europe, Northern Asia and N. America, although males do occur J
sporadically in Europe and have been found in N. and W. Africa and Palestine. It is
sometimes suggested that males become more prevalent in warm, southern latitudes, but
more evidence is needed. No males have been seen in Italian ricefields, where much work
has been done, but they are known from further north in Germany, Hungary and Poland.
Because they are produced by ameiotic parthenogenesis, (i.e. without any reduction of the
chromosome number from the diploid condition), and no genes are received from a male,
every egg has the same genetic constitution as its mother and all eggs produced by a given
individual are genetically identical. Nevertheless, even under the same conditions, eggs of
a single batch do not all behave alike. (Even genetically identical eggs may respond in
different ways to different conditions). Depending on temperature, many hatch within as
little as 3 to about 14 days. Others do so after a longer time interval. As this period is very
variable, it is not easy to define the types of eggs. By hatching within, to set arbitrary
limits, say 14 days, eggs reveal themselves as immediate developers. When they take
longer, they may be said to have entered a period of rest or diapause. However, among
these some may hatch within say 15 to 150 days, while others may become dry, in which
condition they can withstand desiccation and, like plant seeds, remain dormant for long
periods. These long-lived eggs are here referred to as dry resting eggs. In this condition
they are exceedingly resistant, as indeed are wet eggs, the latter, and presumably the
former, being able to withstand even freeze drying.

Sohn and Kornicker (1979) froze wet eggs for several days at c. -25°C, then freeze dried
them at -25 to -30°C in a 0.01 millibar near-vacuum for several more days. In one case
eggs were kept for a day in a near-vacuum of 0.001 mb. Such temperatures are similar to
those that previl at altitudes of about 6 km, the pressures like those at 80 to 100 km. As wet
eggs were used, dehydration must have been rapid and very rigorous. On immersion in
water, many eggs hatched (37% of those subjected to the greatest reduction in pressure did
so) and grew into adults, which produced viable eggs. Freeze drying dehydrates tissues
very effectively. Ability to survive this process is interesting as it is apparently by driving
out the last trace of water that the very heat-resistant eggs of the notostracan branchiopod
Triops granarius can be destroyed (Carlisle, 1968).

Resting eggs of freshwater calanoid copepods that become buried in lake sediments can
apparently remain viable for more than 300 years (Hairston et ai, 1995), and those of some
species of Daphnia may be able to do so for 125 years (Caceres, 1997). However, even if
viable, such eggs are so deeply buried that their chance of hatching is exceedingly remote.
Whether dried eggs of H. incongruens can survive so long remains to be explored, but
some ostracod eggs have survived for up to 30 years. Wet diapausing eggs of H.
incongruens appear to be less long-lived than dry resting eggs and their role seems
generally to be to tide the population over one period of adverse conditions, e.g. wintertime
in Britain, but more information is needed.

That H. incongruens produces both quick-hatching and resting eggs by parthenogenesis
is in itself interesting because in many small freshwater crustaceans e.g. branchiopods such
as Daphnia and its allies, and fairy shrimps, resting eggs are usually (but not invariably)
produced from sexual (fertilized) eggs. Furthermore, the production of quick-hatching
parthenogenetic, and resting fertilized eggs usually involves different underlying genetic

125

The Horse-troiif>h Ostnicod Heterocypris incongruens

and cytological mechanisms. In Daphnia tor example, parthenogenetic eggs, which have
the same number of chromosomes as the mother, are carried in the latter’s brood pouch and
there hatch quickly, being liberated as free-swimming juveniles. Production of sexual eggs
on the other hand involves a halving of the number of chromosomes during the making of
the egg, with fertilization by a male sperm (whose chromosome complement is half that of
the male) restoring that number to the usual (diploid) condition. The fertilized resting eggs.
Just two in this instance, are placed in a protective receptacle formed from the carapace
which is then cast off by the female and goes by the name of an ephippium. In H.
incongruens there is no such clear distinction between the production of the two kinds of
eggs, and a female can lay a mixture of quick-hatching and resting eggs. This remarkable
ability has not attracted the attention it deserves, and the mechanism whereby it achieves
this leat remains mysterious. The possibility that eggs react in the.se different ways
according to prevailing conditions can be ruled out by the fact that some eggs of a batch
may hatch quickly while others, from the same batch, may enter a period of dormancy.
Experiments to .sort out these problems face the difficulty that all eggs look alike. They can
only be classified after the event on the basis of their performance. In some freshwater
crustaceans, eggs that develop quickly have coarse-grained yolk and vacuoles are present,
while drought-resistant resting eggs have fine-grained yolk and no vacuoles. Whether this
applies to H. incongruens remains to be investigated, but it seems unlikely as females
would have to produce eggs of two kinds, sometimes as components of a single laying.
Like tho.se of other crustacean resting eggs, those de.stined to enter diapause (i.e. to ‘rest’)
develop to at least the gastrula stage or a little beyond. Eggs exposed to the air immediately
after laying will presumably perish if.they have not reached this .stage.

It is difficult to draw a clear distinction between immediately hatching and resting eggs
in H. incongriiens. Thus, Angel! and Hancock (1989) isolated 48 females, one per
container, and put each egg laid into a .separate container. Of 2.662 eggs so isolated, 842
(32%) hatched within 125 days. Of these. 309 hatched within 5 days and 88 more in the
next 5 days. In the third 5 day period only 15 eggs hatched and a rather similar hatch rate
continued up to about 60 days, then increased until, in the period between 76 and 80 days.
48 eggs hatched. The hatch rate then declined steadily and hatching had virtually ceased
when observations ended after 125 days. The remaining 68% of the eggs laid were
potentially resting eggs or had died.

Because the parentage of each egg was known it was possible to see how many eggs
from a particular female hatched within the first 10 days and how many did so within 125
days. There was enormous variation. For example, of the 99 eggs produced by the most
fecund female, 43 hatched within 10 days while none of the 83 eggs of another individual
did so. The position in the sequence of laying bears no relationship to the type of egg.
There is no apparent pattern: a female may produce diapausing eggs anywhere within the
sequence - or lay only such eggs. Eggs may be deposited at a steady rate, or in bursts of
activity. Whether any of the variability in egg behaviour can be explained by the fact that
the animals used belonged to different clones (see below) is not known. Certainly,
however, such remarkable behaviour as both rapid and slow hatching of eggs in a single
batch cannot be so explained.

These observations incidentally revealed great variation in fecundity - from 99 to 3 eggs
in this batch of 48 females. The animal that laid 99 eggs survived for 34 days after laying
began; the next most productive (96 eggs) for 61 days. The 3-egg female lived for 40 days
after laying its last egg but its life span is not recorded.

Parthenogenesis and clonal diversity

As all eggs, at least in Britain (and in most populations elsewhere), are produced by
ameiotic parthenogenesis, the offspring to which they give rise are genetically identical to
their mother. Long-held theoretical opinion has it that little genetic diversity is therefore to
be expected. Furthermore, deleterious mutations gradually accumulate in asexual lineages
because individuals with the fewest such mutations are. in the long term, liable to be lost

126

The Horse-trough Ostracod Heterocypris incongruens

by chance, with an inevitable decline in litness, or so the theory goes. Indeed, Lynch ( 1984)
demonstrated a rapid decline in genetic variation in Daphniii during periods ot parthen-
ogenetic reproduction, and Spitze (1991), working with a population of D. pulex in which
there is an alternation between periods of parthenogenesis and sexual reproduction, showed
that between 93 and 99% of the initial genetic variation can be lost in the course ot a
season of parthenogenesis spanning 8 to 10 generations. These losses are made up during
the annual bout of sexual reproduction. This cannot happen in permanently parthenogenetic
populations of H. incongruens, yet such populations persist and are widely distributed. To
this we shall return.

Because of the drawbacks associated with parthenogenesis it has been argued that
asexual species are less well endowed for long term survival than sexually reproducing
species. The latter generate enormous variability through repeated mating and the
inheritence of genetic material from two parents, no two of which, save monozygotic
twins, are alike. This means that genotypes of many kinds are available to cope with
environmental diversity, and selection for the fittest genotypes can occur. Historical
evidence in support of this view has been adduced from just one group of animals -
freshwater ostracods. Their calcite-impregnated carapaces preserve well in alkaline
situations, and the patterns of such remains deposited at several Holocene sites in Britain
and Europe between cl(),()()0 and 5,0()()B.P. - a period of considerable climatic change -
suggested that sexually reproducing forms had an advantage over those reproducing
asexually, and that the advantages of asexuality are generally short-term (Griffiths &
Butlin, 1995). Nevertheie.ss, the idea that obligate parthenogenesis leads, literally, to a dead
end can be challenged. Bdelloid rotifers, of which there are many species, all apparently
entirely parthenogenetic, have evidently existed for long periods. The same is possibly true
of a group of freshwater ostracods, the Darwinuloidea, that brood their young and whose
swollen brood pouch can be seen in fossils, of which there is a record of females only that
extends over at lea.st 70 million years. However, an alleged male of the extant Darwinula
stevensoni has been de.scribed which, if correct, is an embarrassment to an otherwise
fascinating story, though such rare males are regarded by Rossetti and Martens (1996) as
examples of non-functional atavisms (‘throwbacks’ to a former state) of little or no
significance in reproductive strategies.

More certain is that the theoretical belief that parthenogenetic animals show little
variability is in some cases erroneous. The technique of enzyme electrophoresis has shown
that there are many different clones of H. incongruens. The first to demonstrate such clones
in this, and indeed in any ostracod, were Sywula and Lorenc (1982). Electrophoresis
reveals some of the genetic variation of an organism in the form of bands on a gel, whose
pattern is the same in all individuals of a single clone, but different in different clones. The
bands reveal the genotype of an individual at several, but by no means all. its enzyme loci.
Clones revealed in this way are sometimes referred to as electrophoretic clones. When
patterns differ at several loci their possessors are termed multilocus genotypes. They are
morphologically identical, or at least this seems to be the case though there is no theoretical
reason why a difference, which would probably be small, should not be morphological.

Electrophoresis can only reveal differences in enzyme loci for which tests are possible:
other such differences probably exist. The technique may therefore not detect all the clones
that are present in a given species or population. The only example known to me of clones
of ostracods that can be distinguished by simple inspection concerns what in Britain is the
exclusively parthenogenetic Psychrodronnis olivaceus which is usually green in colour but
of which I found populations in two separate springs in Yorkshire that included both green
and white individuals (Fryer, 1993). Presumably the differences between clones include
perhaps a single gene for colour whose effects arc seen in this striking way rather than as
hidden differences revealed by electrophoresis. It is of course possible that both green and
white clones each embrace several electrophoretic clones.

In spite of theory, H. incongruens has at its disposal a degree of genetic variation.
Moreover, several clones sometimes co-exist. This increases the chances of survival or of

127

The Horse-trough Ostracod Heterocypris incongruens

building up large populations because conditions deleterious to one clone may be less so,
or even advantageous, to a dilTerent clone. That this is no mere hypothetical argument has
been convincingly demonstrated in Italian ricelields. Here obligate parthenogenetic
populations ol H. incongruens flourish. No males have ever been found there. Italian
ricelields are flooded from May to September. During the rest of the year small puddles a
few centimetres deep remain. Early investigations revealed 5 clones. Although the situation
later proved to be more complicated than at first thought, it is convenient initially to ignore
later lindings and refer to the original work, which remains valid. Of the 5 clones, each of
the 3 commonest favours, or flourishes best at, a particular temperature and have been
designated as winter, summer and intermediate clones. To simplify the story attention is
concentrated largely on the winter and summer clones.

In the early part of the year, the winter clone predominates or exists alone and the
summer clone is completely unrepresented. A few individuals of this clone appear in May
and by the end of July it has entirely replaced the winter clone (Rossi & Menozzi, 1990).
Experiments conlirm the adaptive differences between clones. Individuals of the various
clones reared at different temperatures differ strikingly in performance at extreme
temperatures. At 4°C only individuals of the winter clone survive: those of the summer
clone die within a few days. Those of the intermediate clone survive a little longer. At high
temperatures the advantage changes in favour of the summer clone, and at 32°C individuals
of the winter and intermediate clones die in a few days w'hile those of the summer clone
may survi\e for up to 60 days. (Life spans are in any case shorter at high temperatures).
Tested at 16°, 24°, 28° and 32°C, members of the winter clone laid most eggs at the lowest
temperature and very few at the highest, while the summer clone did the reverse, laying
scarcely any eggs at 16°, and most at 32°C. The intermediate clone laid most eggs at 24°C.
Likewi.se eggs of the winter clone hatched much better at I6°C than did those of the
summer clone: at 32°C the position was reversed. Of eggs laid by the w'inter clone, the
percentage of diapausing eggs rose rapidly at high temperatures (such eggs are an
insurance against adverse conditions) but fell in the summer clone. It is therefore clear that
the summer and winter clones of H. incongruens are adapted to life at the warm and cooler
temperatures at which, respectively, they occur in nature.

As detailed investigations by Carvalho (1987, 1988) had revealed seasonal

electrophoretic clones in an almost entirely parthenogenetic population of the anomopod
’cladoceran' Daphnia magnet, the situation in H. incongruens is probably typical of
parthenogenetic crustaceans, and perhaps of parthenogenetic invertebrates in general. The
work on Daphnia not only revealed a seasonal succession of clones but showed that their
.survival and ability to flourish were clearly related to the temperatures at which they occur
in nature, and that egg production of different clones was also geared to appropriate
temperatures. This last point, w'hich also applies to H. incongruens. is important, as egg
production had long been thought to be related in a simple manner to such en\ ironmental
factors as food levels and temperature (which are of course important) and not to genotype
(i.e. not to the genetic attributes of their producer). In fact, for some clones a rise in
temperature results in a decline in egg production (though it rises in other clones) and not
an increase, as was generally assumed to be the ca.se. This is so also in H. incongruens
(Rossi et ai. 1991 ).

Although no morphological differences have been found to be associated with
electrophoretic patterns. Rossi and Menozzi (1990) found that while individuals of the
winter clone had a mean length of 1.55 mm. in the summer clone it was only 1.15 mm.
This is not surprising, as small crustaceans that develop at high temperatures tend to be
markedly smaller than tho.se that do .so in cooler conditions (in Ghana females assigned to
H. incongruens by Benzie (1984) had a length of between 0.8 and 0.93 mm), and this
appears to be a purely phenotypic elfect; tor example. Coker (1933) tound that individuals
of the copepod Acanthocyclops vernalis reared at 7-10°C are ‘invariably so much larger
than tho.se reared at I8-I9°C. and 28-30°C that the difference is obvious to the naked eye.
It is therefore surprising that Rossi and Menozzi (1990) say that significant size differences

128

The Horse-trough Ostracod Heterocypris incongruens

between the clones persisted through at least three generations reared at c. 20-24°C, which
suggests that the differences have a genetic basis. However, reared individuals of the
winter clone had a mean length of 1.26 mm, which is considerably less than in the field,
and consistent with development at higher than winter field temperatures. Had size
determination been entirely genetic these animals would have been the same size as their
parent. The summer clone had a mean length of 1.04 mm, again less than the field
population, but in this case, perhaps unexpected, as summer temperatures are sometimes
higher than those at which these individuals were reared. Again, however, an
environmental factor is clearly involved. Nevertheless, individuals of the winter clone were
still larger than those of the summer clone, which supports the likelihood of a genetic
effect, though the disparity was less than in the field. The general diminution in size of the
reared animals may in part be a result of keeping them in very small volumes (20 ml) of
water. It seems best to keep an open mind on this matter. While the rearing experiments
suggest that there may be a genetic element in the size difference between summer and
winter clones, they do not take into account what seems to be a general phenotypic effect
of temperature on size.

Temperature is not the only thing that changes seasonally in temperate climates. Day-
length does likewise and is used as a cue for changes in reproductive behaviour by several
small crustaceans. In some species of Daphnia short day-length plays a key role in
inducing a change from parthenogenetic reproduction to the production of ephippia (resting
eggs) and the males that are usually required to fertilize them. Shortening day-length also
serves as a cue leading to the production of resting eggs in certain copepods. However,-//.
incongruens seems to be the only ostracod for which information has been obtained on the
effect of day-length on reproductive behaviour. Using summer and winter clones from
Italian ricefields, Rossi and Menozzi (1993) showed that, when kept at 24°C in a regime of
16h light and 8h dark, individuals of the summer clone survived longer and produced more
eggs than those of the winter clone, which accords with their occurrence at a time of the
year with long days. When kept at 24°C in 12h each of light and dark there was no
significant difference in the survival of the two clones. However, of the eggs produced by
the winter clone, the percentage of diapausing eggs was greater, as was the actual
production of such eggs, than it was in the summer clone. Resting eggs are often produced
by small crustaceans as a response to unfavourable conditions (though these are not always
easy to define). Long days are favourable to the summer clone, but unfavourable to the
winter clone, so the reaction is ‘sensible’ even if the mechanism is not understood. We may
here, however, have a possible clue to the factors that regulate the production of diapausing
eggs in this ostracod, though it does not explain everything concerned with what is still a
mysterious matter.

Work on Daphnia provides supporting clues. Carvalho and Hughes (1983) showed that
two populations of D. magna, from different ponds, and evidently of different clones, kept
at the same density of individuals, at the same temperature, and given identical amounts of
food, nevertheless differed in the incidence of ephippium production over a range of what
were known to be inductive photoperiods. Other experiments also suggested that
ephippium production may be triggered by subtle differences in day-length in different
clones in a single pond.

By extending the investigation of clones to another ricefield many kilometres further up
the Po Valley, many more clones of H. incongruens were found. Indeed, although sampling
was less intense than at the first site, 26 multi-locus genotypes were found (Rossi et ai,
1993) and further work increased the number to 35 (Rossi et al, 1996). This difference in
clonal richness of the two populations is considered later. These clones also exhibit
.seasonal patterns. However, instead of one clone dominating a particular season, a cluster
of elones do so, to be replaced by other clusters at other times of the year. Nevertheless,
clones with the same seasonal cycle, e .g. summer clones, may employ different strategies.
Thus one summer clone produces almost entirely eggs that enter a period of diapause while
another produces only a small percentage of such eggs. This makes one wonder if the

129

The Horse-lrouf’h Ostracod Heterocypris incongruens

animals used by Angell and Hancock ( 1989), that behaved in different ways in this respect,
may have belonged to different clones. Such differences endow a species with the ability to
cope with a variety of environmental conditions, which is advantageous in the often
ephemeral habitats frequented by H. incon^nens. They may go some way towards
explaining the statement of Angell and Hancock that ‘no environmental stimulus is needed
to induce the production of dormant eggs’ and that the.se form a normal part of the output
of every female even though conditions at the time of laying are suitable for immediate
hatching. Problems, however, remain. If no cues that stimulate the production of
diapausing eggs are involved, it is possible that the production of both kinds of eggs by all
clones allows clonal succession to take place simply by crude climatic elimination ot
inappropriate seasonal clones. As we have seen, however, there is some evidence that cues
arc used, c.g. a long day-length appears to induce the production of diapausing eggs in
winter clones.

Likewise, while it seems logical to deduce from the incidence of seasonal clones that
their diapausing eggs hatch at the appropriate time of the year in response to particular
stimuli, Angell and Hancock maintain that ‘no special environmental stimulus is needed' to
induce hatching save that conditions are ‘favourable for development’. Again one can
envisage inevitable hatching and climatic elimination of inappropriate clones, though the
enormous variability of hatching makes it diflicult to see how any mechanism, such as a
day-degree effect, might operate. The observations ol Angell and Hancock were not
designed to answer such questions nor carried out under conditions suitable lor this. In fact,
Rossi et al. (1991) showed that both wet diapausing and dry resting eggs of summer and
winter clones of H. incongruens differed in their response to conditions at the time ol
hatching. Eggs from both clones were obtained from individuals kept at 28° and 32°C
(which happened to provide abundant material), subjected to the same treatments of light:
dark regimes and temperatures, and then tested for hatching abilities at 32° and 4°C. In all
cases eggs from the winter clone hatched more quickly and in greater numbers than did
those of the summer clone (which in some cases failed to hatch within the time span of the
experiments). Thus different responses to hatching stimuli in long-lived eggs of summer
and winter clones are indicated, but how these are to be interpreted remains problematic.
Indeed, it is mysterious why eggs of the winter clone should hatch at high temperatures.
That tho.se used were laid by females kept at high temperatures, at which they surxive
badly, may complicate an already complicated situation. It is also possible that in northern
areas, where H. incongruens over-winters either as wet diapausing or dry resting eggs, cues
for hatching may be more simple than in populations where a series of clones replace each
other throughout the year.

Enzyme "electrophoresis has also revealed clones of H. incongruens in Canada. Seven
such were found in populations living in tiny rock pools in Manitoba. Of 42 investigated
pools. 31 revealed only one clone. 10 pools each held 2 clones and one had 3 (Havel et al..
1990). These pools often lie in close proximity, which may facilitate dispersal. In Italy, in
each of five small temporary ponds only one clone was detected (Rossi et al.. 1993). No
assessment of the number of clones present in a single horse-trough has been made, but
most such populations probably consist of a single clone. The chance of a trough being
colonised more than once is even lower than is that for a single such event. Such sites
could however, receive more than one clone it a leal or other object derived from a larger
habitat were to arrive carrying a clump of eggs deposited by more than one female that
belonged to different clones.

How CLONAL DIVERSITY HAS BEEN ACHIEVED

Parthenogenesis is clearly derived from sexual reproduction and has arisen many times
among ostracods. The existence of numerous parthenogenetic freshwater species, and the
obvious success of .some of them, suggests that the alleged drawbacks of parthenogenesis,
namely an inability quickly to generate genotypic diversity, has been successfully
overcome by exploiting numerous clones. How has this clonal diversity arisen? Some of it

130

The Horse-trough Ostrcicod Heterocypris incongruens

has undoubtedly done so by mutation within asexual lineages, though genetic work |
suggests that mutation rates are not abnormally high. Some of it also seems to stem from n
the repeated suppression of sexuality in a single species. Because the genotypes of each n
bisexual ancestor would inevitably be different, so too is each derived asexual lineage.

Another route depends on the ability of females that normally reproduce by |
parthenogenesis to retain the ability to mate with males. Because eggs produced by I
parthenogenetic females are diploid (as are the fertilized eggs of bisexual animals), should |
a potentially parthenogenetic female mate with a male, its diploid (2n) eggs and the haploid d
(n) sperm of the male, will give rise to a triploid (3n) individual. Triploid parthenogenetic |
clones of H. incongruens have indeed been found in Canada, and are genetically similar to \
what would be expected of such a mating (Turgeon & Hebert, 1994). The finding of a 1
polyploid clone was itself interesting as it had long been thought that all parthenogenetic iji
ostracods were diploid, though triploid forms of Eucypris virens had in fact been found in |
France. Since the discovery of polyploid clones of H. incongruens, study of other species ij;
has revealed further cases. Tetraploid (4n) clones of H. incongruens have also been found d
and may be the result of matings between triploid females and males with haploid sperm. ■
However, as they could perhaps arise via an abnormal event in the development of eggs in I
a diploid lineage, their origin remains sub judice. Although the ricefield population in Italy I
that has at least 35 multi-locus genotypes is in an area in which males have never been I
found, it is possible that this clonal richness reflects recombination events in the past.
Whether any of these clones are triploid is unknown.

Most remarkable of all is the origin of variability discovered by Turgeon and Hebert
(1994). Near Churchill, Manitoba, parthenogenetic populations of H. incongruens co-exist
with bisexual populations of H. gluucus. Among the clones of H. incongruens were two
that were not only triploid but at four loci combined alleles otherwise unique to H. glaucus
with others typical of H. incongruens. This could be the result only of matings of
‘parthenogenetic’ females of H. incongruens with males of H. glaucus: indeed, such
copulations were observed on several occasions. It is, however, doubtful whether such
hybridization is of much importance in this instance. Such hybrid polyploid clones must
rarely be successful. If they were common, the genetic variability of H. glaucus found in
that area would generate a much greater variety of clones than in fact exists. Nevertheless
the phenomenon is of great interest in revealing a remarkable way in which the variability
of essentially parthenogenetic clones of H. incongruens (and perhaps other ostracods) can
be increased, and may be important in some cases.

Dispersal and related problems

How small freshwater crustaceans are dispersed has long attracted attention, but has
produced little concrete information. Dispersal of H. incongruens is mainly by dry resting
eggs. Other devices are almost certainly unimportant as active stages quickly die if
removed from water and dispersal of wet diapausing eggs is probably a rare event, except
perhaps in peculiar circumstances such as the Hushing of a rice field. Other ostracods have
been seen attached to the setae of aquatic bugs - I have myself recorded Cyclocypris laevis
so attached (Fryer, 1953) but there appear to be no reports of H. incongruens doing so, and,
at best, this could lead only to dispersal over short distances. Cyclocypris ovum has been
found clinging to the skin of toads and newts (Seidel. 1989), but again an inability to
survive for long out of water suggests that H. incongruens is seldom, if ever, so dispersed.
Longer survival may be possible if the curious case of living Cypridopsis vidua found by
Lowndes ( 1930) in the rectum of a frog is not an unusual event involving an ostract)d, as it
seems to be.

Although H. incongruens produces resting eggs, their dissemination appears to face
formidable problems. Most are lirmly attached to substrata, which must frustrate rather
than facilitate dispersal. A small proportion of eggs, however, lies among bottom detritus,
and others are attached to dead leaves or other iibjects that lend themselves to dispersal by
wind - but only if they become dry and possibly disintegrate. Not all habitats dry out

131

The Hor.se-lroitf>h Ostnicod Heterocypris incongruens

regularly, and winter ice in northern latitudes must also frustrate dispersal. From sites such
as horse-troughs, dispersal may be almost as dilhcult as colonisation. Nevertheless, wind is
almost certainly the main agent ol dispersal. Dried eggs are small and very light. Some
may be carried to high altitudes, where similar sized particles have been found, and where
conditions can be tolerated (p. 124). Any that do so may be dispersed over large distances
by the strong winds that blow there. Most eggs, however, are probably less widely
.scattered. Even if only a small proportion of the eggs deposited each year is so dispersed,
vast numbers must be scattered. For example, if in an area of I km- there is just one small
pond that in the course of a season produces 10^ adults of H. incongruens. and if only one
dry resting egg per individual is dispersed, this is equivalent to one egg per m- for that area.
11 only 10%, or 1%, ol this number of eggs is .scattered, this provides one egg per 10 m-, or
100 m-, respectively. Such calculations are, however, unrealistic. By ignoring the
properties ot resting eggs, which cannot be quantilied, they greatly underestimate their
potential as agents of dispersal. Dry resting eggs can survive for several years. During this
time they may ‘alight" many times and '.sample' many prospective sites.

On the other hand, the hazards are enormous. Many eggs must be lost by lodging in
terrestrial situations and some may hatch, and perish, if wetted by heavy rain, or if they
alight in small temporary puddles. On this matter, a possible major destroyer of resting
eggs, we have no information. Any eggs reaching the sea are also doomed. More may
perish here than on land. Even inoculation of large bodies of freshwater, which is more
likely than is that of small sites, and must often happen, may not be very helpful. //.
incongruens does not flourish there. In the survey of Yorkshire, for example, it was never
found in water bodies even as large as about 3,()(){) m-, and many sites were very small. It is
a reasonable deduction that individuals that hatch in large water bodies, and any offspring
to which they give rise, are quickly eliminated. As a species so tolerant of a wide range of
conditions can presumably cope with the physical environments of lakes and large ponds
and lind suitable food there, competition seems to be indicated. H. incongruens indeed
bears all the hallmarks of a fugitive species - which has good colonising abilities but is
easily outcompeted. Its colonising abilities are not in doubt - 1 have, for example found it
in a small pond dug about 15 months earlier (Fryer, 1985) - and its ability to reach such
remote places as Easter Island reveals its potential in this respect. Interspecific competition
in horse troughs and similar situations must be minimal! Keeping it in company with other
animals may throw light on its competitive abilities or deficiencies.

Comparisons with animals such as parasitic copepods of fishes are illuminating. How
the.se find mobile hosts, which may be rare species widely separated in three dimensional
space, pose what may seem to be even greater problems. However, amazing as is the
location of hosts by such parasites, they have advantages over Heterocypris. Their
dispersal stages can swim, and have sensory equipment that may guide them to a host,
either directly, or in respon.se to signals from already settled parasites, just as settling larvae
of such sedentary adult crustaceans as barnacles are chemically attracted to previously
settled sites. Even rare hosts may have habits that assi.st their parasites. They may assemble
to feed, or mate, or spend periods in sleep, and some parasites conceivably disperse only
for short distances. H. incongruens enjoys none of these advantages. It relies on chance
dispersal of only a small proportion of its propagules. which lack any means of guidance to
prospective sites. Compari.sons with plant .seeds are perhaps apposite though some seeds
are .scattered only in the immediate vicinity of the parent plant, and those that disperse
more widely, usually by wind, have devices to enhance wind-borne transport, or to
facilitate attachment to animal carriers. Target areas are also usually vastly larger than are
those of H. incongruens whose resting eggs have no specialisations for dispersal save small
size and light weight which, in the case of single eggs, are in fact probably ver}' efficient
aids. Wind-blown pollen finds much smaller targets, though there are usually many more of
them, often in close proximity, and pollen grains vastly outnumber eggs of Heterocypris.
Likewise, the number of eggs available for dispersal is minuscule in comparison with such
passively dispersed propagules as fungal spores.

132

The Horse-trough Ostracod Heterocypris incongruens

Dispersal in H. incongruens can in fact serve as a model of a widespread phenomenon.
How small crustaceans locate small, isolated, but favourable habitats is still mysterious,
though perhaps less so than supposed. The nearest relative of H. incongruens in Britain, H.
salina, frequents brackish waters, where dispersal is probably easy, but also colonises
inland saline sites, sometimes of small size. In Yorkshire, three small, widely separated
sites with water of high ionic content, but different chemical composition, have been
colonised (Fryer, 1993), and similar instances are known elsewhere. Formidable as are the
obstacles, such animals do find rare habitats, such as inland waters of high ionic content
located by H. salina, or small sites, such as horse-troughs, favoured by H. incongruens.
Long-lived resting eggs, each with the potential to sample several sites, provide much of
the explanation.

An important feature of dispersal in H. incongruens is that every individual of
parthenogenetic origin that arrives in a suitable habitat is a potential founder of a new
population. Were it necessary for a male and a female to colonise such sites as horse-
troughs, the chances of establishing populations there would be extremely low. As only one
coloniser is required, the chance of success is greatly increased. Herein lies a great
advantage of parthenogenesis, and one that appears to have received inadequate
recognition in theoretical discussions.

H. INCONGRUENS AS A POSSIBLE ANCIENT ASEXUAL

A general inference about asexual animals is that they generally persist for only short
periods of time. Genetic studies confirm that many of them are of recent origin and that
persistent lineages rely on the infusion of genetic variation from sexual relatives.
Nevertheless a few asexual lineages are believed to be ancient. It is difficult to decide
whether the genetic diversity of these is the result of mutation since they became asexual or
of intermittent input from sexual relatives. Using both mitochondrial DNA (which is
inherited entirely maternally) and allozyme data (technical details of which are unnecessary
here) attempts have been made to estimate the age of unisexual lineages of H. incongruens
(Chaplin & Hebert, 1997). An incidental, and completely unexpected, result was that at the
one site at which the two co-existed, H. incongruens displayed considerably more diversity
than the sexual H. glaucusl

Divergence between clones was found to be substantial in both the nuclear and
mitochondrial genomes. A strong correlation in the extent of nuclear and mitochondrial
divergence between pairs of clones suggests that H. incongruens is an ancient asexual
which has undergone considerable mutational divergence. Indeed by calculating the rate of
sequence divergence in mtDNA (and adopting a conservative value for a somewhat
contraversial procedure) it was estimated that it has persisted as an asexual for more than
5 million years. Acceptance of such a figure is provisional but there is a distinct possibility
that asexuality is indeed an ancient attribute of some lineages of H. incongruens. If so, this
remarkable ostracod has another claim to distinction.

An historical irony

H. incongruens has another modest, and unexpected, claim to fame. In view of the great
rarity of males it is ironical that it was in this species that sperms of ostracods were first
recorded. Ramdohr (1808), the describer of H. incongruens, saw them when making a
dissection but, although they came from the sperm duct, didn’t recognise them as sperms.
This is not surprising for they are of enormous length. Wingstrand (1988), who had access
to males from Palestine, gives their length as 1.3 mm, that is about the same length as the
animal itself. Ramdohr saw them straighten out into hair-like structures after their release
and wondered if they were ‘Blutgeftisse’ (blood vessels)! Not until almost 30 years later
did Wagner recognise them for what they were. Giant sperms of ostracods (some are
considerably longer than those of H. incongruens) have received much study but it is a
curious and little-known fact that they were first seen in a species in which most
populations consist entirely of females! It is surprising that Sars ( 1928), who notes that he

The Horse-trough Ostracod Heterocypris incongruens 133

never saw a male of H. incongruens, and who cites Ramdohr’s paper, does not mention the
latter’s observation.

SOMH UNSOLVED CONUNDRUMS

What seems on first acquaintance to be the simple life cycle of H. incongruens turns out in
fact to be very complex. Although much has been learned, unsolved problems remain. It is,
for example, curious that what have been taken to be genetically identical individuals, kept
under the same conditions, may differ considerably in such things as life span and
fecundity. A possible explanation is that in fact they belong to different, unrecognised,
clones, or less likely, the differences are due to infections of which the observer is
unaware, or even to mutations. More problematic is why genetically identical eggs
prorluced by a single female, laid at the same time or within a few days of each other, and
kept under the same conditions, may either hatch within a few days or remain dormant for
long periods. How these different programmes for development operate and, even more
remarkable, how a single parthenogenetic female can lay a mixture of such eggs, remain
unknown.

The exi.stence of genetically different clones has been firmly established, and clear
ecological differences (particulary seasonal requirements) have been demonstrated between
clones, but there are still problems as to how, if at all, clones are cued to produce the kinds
of eggs appropriate for their particular life cycles, or whether they rely on the production of
their mysterious mixtures of eggs who.se products are simply sorted by climatic conditions.
The same is true of stimuli that induce hatching of diapausing eggs whose nature remains
ob.scLire and difficult to investigate. While there is evidence that eggs of different sea.sonal
clones differ in this respect, the differences are not really understood, nor is their
significance in nature. There is also the question of what determines whether dry resting
eggs hatch when they first become wet, or require more than one such wetting. The value
of variability in this respect is self-evident, but how it is achieved remains obscure.

The genetic nature of populations in horse-troughs remains to be investigated. While one
is tempted to predict that there will usually be only one clone per trough, this remarkable
ostracod has produced so many surprises that it would be rash to make this assumption. It
would also be interesting to know whether adjacent troughs share the same clone(s). A
place like Sark with at least three trough populations in close proximity, yet well isolated
from adjacent populations, may provide clues about patterns of dispersal.

Another problem is how H. incongruens maintains genetic variability. Rapid loss of
genetic variation in parthenogenetically reproducing Daphnia has been demonstrated see
p. 126). Many populations of Daphnia can restore this variability by periodic bouts of
sexual reproduction. This is not an option in unisexual populations of H. incongruens. e.g.
in Britain, yet these populations persist and are clearly successful. Some populations of D.
pulex consist of obligate parthenogens. As a group these harbour as much variation in
quantitative traits as do those that reproduce sexually. This seems to be due to the high rate
of origin of obligate parthenogenesis. This cannot be the case in H. incongruens in areas
where there are no bisexual populations from which parthenogenetic forms could be
repeatedly derived, and certainly not if some lineages are ancient asexuals. Nevertheless, in
some areas in which males are unknown, as in certain Italian ricefields. a rich diversity ot
electrophoretic clones may be present. How do these manage to survive so much better
than, apparently, do obligately parthenogenetic Daphnia"!

The roles of environment and heredity in the detemiination of the size of individuals in
different seasonal clones remain to be clarified. Early observations on such clones
overlooked environmental influences but nevertheless .suggest that genetic differences
between clones may be involved.

More information is needed on the structure of the eggs, on how they are attached to
objects, whether there are physical differences between slow- and fast-developing eggs
(which at present can only be recogni.sed on the basis of their performance), on the nature
of the yolk granules, on embryology, on what happens to eggs when they become dry and

134

The Horse-trough Ostracod Heterocypris incongruens

when they are rehydrated. What gives dry eggs their amazing resistance is a problem 4
common to a wide range of resting eggs, and is not really understood.

Finally, one wonders what it is about horse-troughs and similar receptacles that makes |
them so attractive to this common ostracod.

Concluding remarks i

The life cycle and ecology of H. incongruens present many problems. When to these are it
added others posed by the variation in the eggs, the amazing resistance of resting eggs, and i
their dispersal, it becomes apparent that we really know very little about some important a
aspects of the life of this interesting animal. The horse-trough ostracod is truly worthy of 1
attention. !

I thank Prof. G. A. Carvalho who kindly commented on a preliminary draft of this paper, ij

References |

Angell, R. W. and Hancock, J. W. (1989). Response of eggs of Heterocypris incongruens j
(Ostracoda) to experimental stress. J. Crust. Biol. 9: 381-386.

Benzie, J. A. H. (1984). Small scale diurnal migrations by Heterocypris incongruens
(Ramdohr, 1808) Ostracoda: Cyprididae) in a temporary pool, Ghana, West Africa. J.
Crust. Biol. 4: 63-65.

Bronshtein, Z. S. (1947). Fre.shwater Ostracoda. Fauna of the USSR Crustaceans Vol. 2.
No. 1 (In Russian. Eng. Transl. 1988).

Caceres, C. E. (1997). Interspecific variation in the abundance, production, and emergence
o\' Daphnia diapausing eggs. (In prep.)

Carlisle, D. B. (1968). Triops (Entomostraca) eggs killed only by boiling. Science. 161: 1

279. '

Carvalho, G. R. (1987). The clonal ecology of Daphnia inagna (Crustacea: Cladocera) II (
Thermal differentiation among thermal clones. J. Anim. Ecol. 56: 469-478.

Carvalho, G. R. (1988). Differences in frequency and fecundity of PGI-marked genotypes |
in a natural population of Daphnia magnet Straus (Crustacea: Cladocera) Funct. Ecol. 2: *
453- 462.

Carvalho, G. R. and Hughes, R. N. ( 1983.) The effect of food availability, female culture-
density and photoperiod on ephippia production in Daphnia nuigna Straus (Crustacea:
Cladocera) Ereshwat. Biol. 13: 37-46.

Chaplin, J. A. and Hebert, P. D. N. ( 1997). Cyprinotus incongruens (Ostracoda): an ancient
asexual? A/o/ec. Ecol. 6:155-168.

Coker, R. E. (1933). Influence of temperature on size of freshwater copepods (Cyclops).
Int. Rev. Hydrohiol. Hydrogr. 29: 406-436.

Dumont, H. J. and Martens, K. (1996). The freshwater microcru.stacea of Easter Island.
Hydrobiologia ilS: 83-99.

Eox, H. M. and Taylor, A. E. R. (1955). The tolerance of oxygen by aquatic invertebrates.
Proc. R. Soc. LondB. 143: 214-225.

Fryer, G. (1953). Notes on certain British freshwater crustaceans. Nauralist 78 : 101-109.
Fryer, G. (1985). Crustacean diversity in relation to the size of water bodies: some facts
and problems. Ereshwat. Biol. 15: 347-361.

Fryer, G. ( 1993). The freshwater Crustacea of Yorkshire: a faunistic and ecologiccti sun’ey.
Yorks. Nat. Union & Leeds Phil. Lit. Soc.

Ganning, B. (1971). On the ecology of Heterocypris salinus. H. incongruens and
Cvpridopsis etculeata (Crustacea: Ostracoda) from Baltic brackish-water rockpools. Mar.
Biol. 8: 271-279.

Green, J. (1959). Pigmentation of an ostracod. Heterocypris incongruens. J. E.\p. Biol. 36'
575-582.

Grilfiths, H. I. and Butlin, R. K. (1995). A timescale for sex versus parthenogenesis:
evidence from subfossil ostracods. Proc. R. Soc. B. 260: 65-71.

Hairston, N. G. Jr., van Brunt. R. A., Kearns, C. M. and Engstrom, D. R. ( 1995). Age and

135

The Horse-irau^h Ostracod Heterocypris incongruens

survivorship of diapaiising eggs in a sediment egg bank. Ecology 76: 1 706- 1711.

Havel, J. E.. Hebert, P. D. M. and Delorme, D. (1990). Genotypic diversity of a.sexual
Ostracoda from a low arctic site. J. Evol. Biol. 3: 391-410.

Klie, W. (1938). Krebstiere oder Crustacea. Ill Ostracoda. Muschelkrebse. TTie Tienvelt
Deiitschlands. 34: 230 pp.

Lowndes, A. G. (1930). Living ostracods in the rectum of a frog. Nature Lond 76: 958.

Lynch, M. ( 1984). The limits to life-history evolution in Duphniu. Evolution 38: 465-482.

Ramdohr, F. A. ( 1 808). Uber die Gattung Cypri.s Miill. und drei zu derselben gehorige neue
Arten. Maguzin Ges. miturf Freunde Berlin 2: 83-93.

Rossetti, G. and Martens, K. (1996). Redescription and morphological variability of
Darwinula stevensoni (Brady and Robertson, 1870) (Crustacea. Ostracoda). Bull. Inst. R
Sci. Nat. Belg. Biol. 66: 73-92.

Rossi, V. and Menoz/.i, P. (1990). The clonal ecology of Heterocypris incongruens
(Ostracoda). Oikos 51: 388-398.

Rossi, V. and Menozzi, P. (1993). The clonal ecology of Heterocypris incongruens
(Ostracoda): life history traits and photoperiod. Eunct. Ecol. 7: 177-182.

Rossi, V.. Gandolfi, A. and Menozzi. P. (1996). Egg diapause and clonal structure in
parthenogenetic populations of Heterocvpris incongruens (Ostracoda). Hvdrohiologia
320: 45-54.

Rossi, V., Giordano, P. and Menozzi, P. (1993). Genetic variability in parthenogenetic
populations of Heterocypris incongruens (Crustacea, Ostracoda). In Ostracoda in the
earth and life .sciences (ed. McKenzie, K. G. and Jones, P. J.), I llh Int. Synip. Ostracoda
369-383.

Rossi, V., Rossi, M. C. and Menozzi. P. (1991). Life strategy differences among
electrophoretic clones of Heterocypris incongruens (Crustacea, Ostracoda) Verb.
Intermit. Verein. IJninol. 24: 2816-2819.

Sars, G. O. (1928). An account of the Crustacea of No nvay. Vol. 9 Ostracoda. Bergen
Museum.

Seidel. B. (1989). Phoresis of Cyclocypris ovum (Jurine) (Ostracoda, Podocopida,
Cyprididae) on Boinhina variegata variegata (L) (Aniira. Amphibia) and Triturus
vulgaris (L) (Urodela. Amphibia). Crustaceana 57: 171-176.

Sohn, 1. G. and Kornicker, L .S. (1979). Viability of freeze-dried eggs of the freshwater
Heterocvpris incongruens. In N. Kristie, ed. VII Int. Symp. Ostracoda 1-3.

Spitze, K. (1991). Chaoborus predation and life history evolution in Daphnia pulex:
temporal pattern of population diversity, fitness and mean life-history'. Evolution 45: 82-
92.

Sywula. T. E. and Lorenc. R. ( 1982). Esterase polymorphism as a marker of interpopulation
differentiation in Cvprinotus incongruens (Ramd). (Ostracoda. Crustacea) Bull. Acad.
Poln. Sci. 29: 317-322.

Turgeon, J. and Hebert, P. D. N. (1994) Evolutionary interactions between sexual and all-
female taxa of Cvprinotus (Ostracoda:Cyprididae). Evolution 48: 1855-1865.

Wingstrand, K. G. (1988). Comparative spermatology of the Crustacea Entomostraca 2.
Subclass Ostracoda. Biol. Skrift. Kongel. Danske. Vidensk. Selskab. 32: 1-118.

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Based on their observations and e.xperience. as well as a detailed sur\ey of the literature,
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are and discuss ways of improving their con.servation potential. Particular attention is paid

1 36 Book Reviews j

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137

DIET OF BARN OWLS TYTO ALBA
IN A LOWLAND ARE:A OF NORTH YORKSHIRE

J. ROBERTS, G. W. SCOTT' AND S. L. HULL
School of Science and Management, University College Scarborough, Filey Road,

Scarborough, YOl I 3AZ

Abstract

Samples of Bam Owl Tyto alba pellets were collected from a lowland area of North
Yorkshire during the spring and summer of 1995. Microscopic analysis of their contents
provides information about the diet of the study population and highlights the nutritional
importance of small mammals which account for 95% of all prey taken. The field vole
Microtus agrestis was recorded as being the most important single species in the diet,
accounting for half of all prey consumed.

Introduction

Analysis of the contents of regurgitated owl pellets offers a convenient insight into both the
diet of the species of owl concerned and also into the constitution of the community upon
which they prey. The Barn Owl is, in Britain, at the northern limit of its range and has in
recent times become a subject of conservation concern. The species is now restricted to just
a few strongholds in the British Isles (Gibbons et ai, 1993), with many populations
dependent to some degree upon the provision of artificial nest sites. In this study we
investigate the diet of such a population inhabiting the mixed agricultural land of the Vale
of Pickering, North Yorkshire.

' Methods

Study site and pellet collection

.345 pellets were collected from barn owl nest sites during regular ringing and monitoring
work (carried out under license), and from sites within their hunting ranges. The areas
concerned are all within the Vale of Pickering and at approximately 20 m above sea level.
The study area is a mosaic of mixed agricultural land with few large human settlements and
little woodland. It is crossed by a number of water courses ranging in size/volume from
rivers such as the Derwent, to streams and drainage ditches. The majority of the area's field
boundaries support little more than a 2 m belt of unmanaged grassland.

Collection of pellets was carried out twice. A spring collection was made in April 1995
and it is assumed that these pellets contained prey consumed during late winter and early
spring. The second sample was collected (where possible from the same sites) during the
summer months (June and July 1995) and is assumed to have contained prey consumed
during late spring and early summer.

Pellet analysis

The length of all pellets was recorded. Small fragments of pellet from a single site were
‘reconstructed" to produce a pellet of average length. Pellet width was not found to vary
greatly and was not .systematically recorded (though we estimate that all pellets were 25-28
mm in diameter). After measuring, the dry pellets were gently teased apart and their
contents retrieved for identification. Obvious material was removed (skulls, long bones
etc.) and the matrix of the pellet was then examined under a stereo-microscope in order that
smaller remains (in.sect parts etc.) might be collected.

Identification of prey items

Skull and jaw remains w'ere used to identify mammalian and avian prey and their numbers

'Author for correspondence (email: grahams@uesearb. ae.uk

NcinmiUst 122(1997)

138

Diet of Barn Owls Tyto alba in a Lowland area of North Yorkshire

were established by a skull count (Corbett & Harris, 1993; Glue, 1974; Julian &
Altringham, 1994). As Barn Owls are known to decapitate avian prey regularly prior to
ingestion (Glue, 1967), leather material, a sacrum, a sternum or long bones were used as
indicators of avian presence. It was not possible to separate the remains of Wood Mice
Apodemus sylvaticus and Yellow-necked Mice Apodemiis flavicolliis and for the purposes
of this study the two are grouped as wood mouse.

The lower jaws and pelves of shrews Sorex spp. were retained during sorting and used to
age and sex individuals so that an assessment of shrew prey population structure might be
made. Shrews were aged according to tooth wear and sexed according to pelvis shape after
Yalden and Morris (1990). When e.stablishing the frequency of individuals in each age/sex
class, jaws/pelves were paired, with each additional jaw/pelvis counting as one individual.

Invertebrates were identified from chaetae (earthworms), or heads and elytra (beetles)
and their numbers were approximated.

Estimates of the mass of mammalian prey items consumed (relative mass being an
indicator of the relative nutritional value of prey types) were obtained from the available
literature (Julian & Altringham, 1994; Yalden & Morris, 1990). The mass of invertebrate
prey items was estimated by the authors. If avian remains could not be identified to a
species, an average mass of 15g was recorded after Julian and Altringham ( 1994).

Rh.sults

A total of 345 pellets was collected, from which 1,616 prey items were retrieved. Pellets
ranged from 20 mm to 90 mm in length (mean ± standard deviation = 49.5 ± I 1 .2).

Sample I (hereafter spring) produced a total of 589 prey items from 122 pellets. The
highest number of items recovered from a single spring pellet was 10 - a combination of
only Common Shrew Sorex araneus and Pygmy Shrew Sorex miiintiis. The mean weight of
spring prey items was estimated as I3g. Sample 2 (hereafter summer) consisted of 223
pellets containing 1,027 prey items. The highest numher of items recovered from a single
summer pellet was 13, again a mixture of Common Shrew and Pygmy Shrew with a single
Field Vole; this was also the largest pellet analysed (90 mm x 28 mm). The mean weight of
summer prey items was estimated as 15g. Two of the summer sample pellets comprised
solely vegetable material (grass and straw). It was initially assumed that such material might
indicate the presence of invertebrate remains but none was found. Similar pellets have been
reported by Shawyer (1994) but an explanation of their existence remains elusive.

Table 1 summarises the data collected (spring and summer combined). A total of 10
species of mammals was recorded in the diet of the Barn Owl population. Of these. Field
Vole and Common Shrew were the most commonly recorded prey items and together
accounted for more than 70% of the total prey intake (in term of numbers of animals
caught). In terms of their relative nutritional importance (proportion of the total prey
weight) Field Voles appear to be disproportionately important, accounting for 51.8% of
prey weight (in comparison with their numerical importance) with Common Shrew, Wood
Mouse and Bank Vole Clethrionotnys glareolus making up the bulk of the remainder of the
diet. All other species appear relatively unimportant (although of course they clearly are
important if considered as a single unit).

Figures la and Ib illustrate the seasonal importance of the various prey types. Figure la
reveals that Common Shrew accounted for 24.7%, Wood Mouse for 9% and Field Vole for
51% of the total prey items recorded in spring. Whilst the Field Vole still accounted for
51% of the diet in summer (Figure Ib), Common Shrew decreased in importance (30% to
16.9%) and the Wood Mouse increa.sed in importance from 9% in spring to 18.4% in
summer. The summer diet is more varied than the spring diet (x~ test to compare the
relative frequencies of prey types in the two samples; x* = 53.73, d.f = 12, p<().()()l ) (Sokal
and Rholf, 1991). In both samples the single most important species is the Field Vole (c.
52% by weight). Of the other important species. Bank Vole appears relatively constant in
value (8-10% by weight).

139

l'>iel of Barn Owls Tyto alba in a Lowland area of North Yorkshire

TABLE 1

Summary ol barn owl diet Irom pellet analysis (pooling spring and summer samples).
Prey weight (%) is the prey by weight, expressed as a % of the weight of all prey.

All values for prey weights are from Yalden and Morris (1990).

Beetle weight was estimated by the authors. Bird weights were ealculated assuming
a mean weight of 15g (Altringham et al., 1994).

N.

%of

mean

total

prey

prey

total

item

prey

weight

prey

weight

weight

as a 9c

(g)

(g)

of diet

• MICTROTINES
( Field Vole

M icrotiis a f> res Its

571

35.3

21

1 1991

51.8

i Bank Vole

CleihriononiYs glareoliis

123

7.6

16

1968

8.5

' Water Vole

A rvicola terrestris

1

0.2

100

100

0.4

■ MURID.S
^ Wood Mouse

Apodennis sxlvaticiis

193

1 1.9

14

3474

15

1 House Mouse

Mas niiisciiliis

13

0.8

12

156

0.7

1 Harvest Mouse

Micromxs niinutiis

5

0.3

5

25

0.2

! Brown Rat

Rcitliis norvef’iciis

3

0.2

60

180

0.8

• SHREWS
t Common Shrew

Sore.x arciiietis

563

34.8

8

4504

19.4

' Pygmy Shrew

Sore.x niiniitiis

1 12

6.9

4

448

1.9

Water Shrew

Neoniys fodiens

17

LI

12

204

0.9

1 BIRDS

Total

5

0.3

80

0.3

L Chaflinch x 2

Friiigilla coelehs

44

: Pa Ills species x 2

P. ccieriileiis or ater

21

1 Unidentilied x 1

15

) BEETLES
'Scarabaeidae
' Summer Chafer

Aniphiinallon solstitialis

4

0.2

1

4

0.01

' Silphidae

Necrodes littoralis
Necrophorus hiimator x 2

ANNELIDA
' Earthworms

Liinihriciis species

6

0.4

5

30

0.1

1 TOTAL

1616

100

23168

100

The population structure of the shrews recovered from the spring (20b individuals. 309?-
of the total spring prey) and summer (210 individuals, 30*^ of the summer total prey) pellet
samples indicates that spring pellets contain a higher proportion of older animals, whifst
younger animals account for a higher proportion of the prey in the summer pellets (x’ =
30.67. d.f = 3. p<().001). Both samples contained a predominance of male animals (x- =
1 1 .95, d.f = I . p<0.001 ) although in summer the sex bias was not quite so pronounced, with
males accounting for 659c of the summer sample and 809^ of the spring sample.

Dlscussion

A puncture wound at the back, of most of the recovered mammal and bird skulls was
assumed to have been the killing method employed most frequently by the owls of this

140

Diet of Barn Owls Tyto alba in a Lowland area of North Yorkshire

(b)

Wcxxl Mouse
18.4%

ComuKii Shrew
16.9%

Bank Vole
7.8%

Field Vole
51.9%

IVgrav' Shrew
1 ,4%

(Xhers

3,2%

Others include:
Water Vole 0.6%
Water Shrew 0.8%
Har%’cs4 Mouse 0.1%
House Mouse 0.7%
Rat 1.2%

Bird 0.4%

Beetles 0.1%
I-arthwimns 0, 1 %

FIGURE 1

Prey items identified in (a) spring pellet sample analysis and (b) summer pellet
sample analysis. Prey items represent a % by weight of total prey weight.

study population, although Raezyinski and Ruprecht ( 1974) have recorded that damage to
skulls is usually less common in the prey of Barn Owls when compared to species such as
the Tawny Owl Strix alitco or Little Owl Athene noctiia. Such damage, which facilitates
skull digestion, made it necessary to identify some remains from jaws and long bones. It
was also noted that a number of pellets contained under-digested material, a situation
which probably reflects high prey abundance where the stimulation of the next meal
triggers premature regurgitation (Lowe, 1980).

As with the majority of previous studies of Barn Owl diet, the Field Vole was identified
as being the main prey species of this population in both seasons. Although more shrews
than voles were caught in spring, the Field Vole consistently provided the highest
nutritional value (as a percentage of the total prey weight). The relative proportions of
voles and shrews in Barn Owl diets have been discussed by Webster (1973) (who
suggested that a fall in dietary vole numbers results in a disproportionately large numerical

141

Diet of Barn Owls Tyto alba in a Lowland area of North Yorkshire

increase in shrew prey, ensuring that the owls achieve their nutritional requirement), and by
Bunn et al. (1982), who suggest that a vole/shrew ratio relationship is typical of British
pellet analysis. We observed a relatively constant (by weight) vole component to the diet of
our study population but did note a shift in the relative value of shrews and Wood Mice,
presumably a response by the birds to the need to balance primary and secondary prey
consumption in order to maintain nutritional intake.

In addition to the shift in the relative importance of shrews in the diet of the owls,
examination of the population structure of the shrews captured shows that in spring, when
shrew numbers were higher, more males than females were recorded and the majority of
the animals were older individuals. In summer more males were recorded but the bias was
lower and the percentage of younger animals was higher. Similarly Brown (1981) has
reported that 61% of the shrews recovered from Barn Owl pellets in May are male. Corbett
and Harris (1993) have suggested that males are more likely to be caught than females
because they spend more time above ground vigorously defending their territories and are
thus exposed to a higher predation risk; this is particularly the case in spring. The younger
animals in the summer sample are presumably put at predation risk during post-weaning
dispersal involving both sexes (May onwards; Butterlield et al., 1980), hence the relative
increase in the number of females recorded.

In some parts of the Barn Owls’ range (and especially on arable land) the Brown Rat
Rattits norvegietts is a major component of the diet (Glue, 1974). As rats can weigh more
than 5()()g tCorbett & Harris, 1993), it is likely that attacks would concentrate upon
younger animals. In this study the rat did not represent a major prey component, and the
animals which were recorded were relatively small. Similarly. House Mouse Mas musculus
and Harvest Mouse Microtnys inimitus, whilst important in some studies of the diets of
owls were of little importance in this study, probably as a result of the relatively small
amounts of suitable habitat for these species in the study area.

In common with previous work (Glue. 1974), birds formed a relatively unimportant
proportion of the diet of the owls. The species caught were common woodland/hedgerow
birds. Coal Tit Pants ater. Blue Tit P. caeriileiis and Chaffinch Fringilla coelebs. and the
slight increase in their importance in summer may be due to the increased availability of
fledglings. Similarly, invertebrates did not form a major component of the diet.
Earthworms were recorded from three pellets and beetles were recorded only from the
-summer sample. Shawyer (1994) has suggested that invertebrates, particularly beetles
(Mikkola. 1983), may be consumed by adult birds during the raising of young when most
mammalian prey must be given over to their chicks.

Acknowledgments

We would like to express our gratitude to Pawd Willet and David Jardine who provided the
pellets used in this study.

References

Brown, D. J. (1981). Seasonal variations in the prey of some Barn Owls in Gwnynedd.
Bird Studs- 2S: 139-146.

Bunn. D. S.. Warburton, A. B. and Wilson. R. D. S. (1982). The Bant Owl T. & A. D.
Poy.ser. Carlton.

Butterfield, J.. Coulson, J. C. and Wanless. S. (1980). Studies of the distribution of Pygmy
and Common Shrews in Northern England. J. ZooL. London. 195; 169-180.

Corbett. G. B. and Harris, S. eds. (1993). The Handbook of British Mammals. 3rd edition.
Blackwell, Oxford.

Gibbons, W. G.. Reid. B. R. and Chapman. R. A. (1993). The New Atlas of Breeding Birds
in Britain and Ireland: 1 988- 199 1. T. & .A. D. Poyser, Carlton.

Glue. D. E. ( 1967). Prey taken by the Barn Owl in England and Wales. Bird Study 14; 169-
183.

Glue. D. E. (1974). Eood of the Barn Owl in Britain and Ireland. Bird Study 21; 200- 210.

142 Book Reviews

Julian, S. and Altringham, J. D. (1994). Bat predation by a Tawny Owl. Naturalist 119: 49-
56.

Lowe, V. P. W. ( 1980). Variation in digestion of prey by tbe Tawny Owl (Strix aluco). J.
ZooL, London 192; 283-293.

Mikkola, H. (1983). Owls of Europe. T. & A. D. Poyser, Carlton.

Raczyinski, J. and Ruprecht, A. L. (1974). The effect of digestion on osteological
composition of owl pellets. Acta Ornithol. 14: 25-38.

Shawyer, C. R. (1994). The Barn Owl. Hamlyn, London.

Sokal, R. R. and Rholf, F. J. ( 1991 ). Biometry. 2nd edition. Freeman. New York.

Webster, J. A. (1973). Seasonal variation in mammal contents of Barn Owl castings. Bird
Study 20: 185-196.

Yalden, D. W. and Morris, P. A. (1990). The analysis of owl pellets Occasional
Publications of the Mammal Society No. 13. The Mammal Society, London.

BOOK REVIEWS

The Making of the Cretan Landscape by Oliver Rackham and Jennifer Moody.
Pp. xvii + 237 with numerous line drawings & maps, -I- 4 pp. of colour plates. Manchester
University Press. 1997. £50.00 hardback, £19.99 paperback.

A remarkable natural landscape with a rich archaeological heritage. Through held
observations, archaeology, ancient and medieval documents, place-names, dendro-
chronology and standing buildings, the authors demonstrate how the island’s wonderful
and dramatic features have been moulded and modified by centuries of human activity.

Charles Waterton: a Biography by Brian W. Edginton. Pp. xi -t- 254, with 40 b/w plates.
Lutterworth Press, Cambridge. 1996. £25.00, hardback.

The eccentric but nevertheless gifted Yorkshire naturalist Charles Waterton (1782-1865)
was a, if not the, pioneer of nature conservation, converting the 250 acre estate around his
home, Walton Hall near Wakefield, into probably the world’s first nature reserve.
However, he is best remembered for his Wanderings in South America, a best-selling work
which has never been out of print since its first publication in 1825.

Five biographies of Waterton and innumerable papers about him have been published
prior to this one. The latest undoubtedly contains new information and fresh interpretations,
but unfortunately it lacks co-ordination. Information spills out in an illogical manner: the
author is easily side-tracked, thus often spoiling the chronological sequence through the use
of quotations apparently included in order to .score points rather than for any real relevance
to the author’s argument. The author is also prone to journalese and slang, as exemplified by
such expressions as ‘little Lord Waterton’, ‘young massa’, ‘boffins’, ‘bubble bubble’ and
‘load of old bull’. The author’s lack of a scientific background is all too evident: he writes of
‘animals’ rather than ‘mammals’, uses incorrect Latin nomenclature and poor terminology
(e.g. rain acid) and his interpretation is inadequately researched. Spelling mistakes and
inconsistencies within the text and index have also been noted.

It has long been recognised that Waterton is a very unreliable source and his
recollections can be untrustworthy and contradictory. This latest biographical treatment of
him has .some merit in trying to set the record straight, but all-in-all the presentation lacks
authority due to the haphazard manner in which the information is provided. Nevertheless,
those interested in the history of natural history and naturalists in general, particularly those
in Yorkshire, will find much to interest them in this biography.

MRDS

143

Obituary - George Rowland Edwards - 1910-1996

OBITUARY

GEORGE ROWLAND EDWARDS
1910-1996

George Edwards was born in Halifax on 16th February, 1910 and died in Eastleigh,
Hampshire on 19th July, 1996. He was the youngest of a family of live, with a sister and
three brothers. His sister remembers that he was always fascinated with wildlile and in his
earliest diaries in 1925 and 1926 he mentioned animals and flowers he had seen as well as
his daily bird list. These diaries he wrote up to the day he died; they provide an excellent
record of his travels at home and abroad. He was one ol the founder members of the Halifax
Zoological Group in 1932. The moorland reservoirs, especially the Gorple group and White
Tlolme, and the Calder valley, especially Elland sewage farm, were visited regularly by
members of the Group in Spring and Autumn. These visits showed there were annual
movements of waders through the area and accounts ol these were published in The

• Naturalist. In 1947, a ringing station was established at High Royd sewage farm, the first
inland site in Britain. Then, in 1949, just after the Group merged with the Halifax Scientific

■ Society, there was excitement when a Desert Wheatear was lound on the local moors near
the appropriately named farm, ‘Egypt’.

By the early 1930s he was travelling more widely in the county and in Britain. In 1934, on
a trip to Scotland, he recorded 13th June as Temminck’s Stint day when the first nest lor
t Britain was found in Speyside. In the same year, he visited Spurn tor the first time, followed
by a further seven visits up to 1939, including one with some of the Halifax Zoological
Group in 1936, an account being published in The Naturalist.

He soon developed an interest in photography, probably encouraged by his elder brother,
and, in 1933, turned to cinephotography for birds. With Vernon Crapnell, his friend and
.regular companion with a similar interest, he must have been one of the first wildlife
cinephotographers in Yorkshire. A year later, he was capturing Temminck s Stint on film.
‘-Nearer home, from a mill roof in Stainland. he filmed a rookery and, at High Royd sewage

• fann. the display of Redshanks. Before the 1939-45 war he worked for the family tirm of

■ Edwards and Sons, a furnishing business housed in a two-storey building with a tower. He
used this tower as a studio for titling and editing films and a place for making photographic

' hides.

During the war he served in the Auxiliary Fire Service, based in Halifax and seconded to
f Bradford and Hull during their air raids. After the war George decided to take up a career
■filming wildlife. In the late 1940s and early 1950s he worked as a field assistant to Eric
; Hosking, making films in the summer and giving shows of his own films and those he had
made for the R.S.P.B. in the winter. With Eric Hosking and Stuart Smith he also studied the
aggressive display of Ringed Plovers and Oystercatchers with the use of stuffed specimens
and a mirror and the reactions of some passerine species to a stuffed Cuckoo. Papers
describing this work were published in British Birds in 1947-1950. In January' 1953 he
joined the staff of the R.S.P.B. to set up a film unit, making films and giving shows until he
left in January. 1957. He was the ideal man for the job at that stage. In 1957 he was assistant
warden on Skokholm for seven months. In 1958 it was back to filming with a five-month
trip on a tanker from Holland to the Persian Gulf and back. In December 1958. he made two
round-the-world trips in the southern oceans as Purser's writer, and in December 1959 and
November 1960. he was filming for the Falkland Islands Dependencies’ Survey (now the
British Antarctic Survey). In 1963. he made three trips on British India cruise .ships, two into
the Mediterranean and one to Madeira and North Africa, when he showed slides and films to
school parties and also demonstrated there could be things to see at sea between the ports of

call! In 1964 there was a similar trip to Reykjavik and Norway. , •

From late 1964 until he retired in 1975. he made films for the Anglia TV Survival

■ Series. These took him to di.stant and remote places: to the Falklands and Antarctica again,
to Tristan da Cunha. Aldabra. Natal. Trinidad and Tobago. Panama and Venezuela and

Naturalist 122(1997)

144

Obituary - George Rowland Edwards - 19 JO-1 996 4

Israel, as well as various assignments in Britain including St. Kilda, Orkney and Hilbr
Island. He also made a film, 'The Spider’s Web’ for the Yorkshire Naturalists’ Trust (nov
the Yorkshire Wildlife Trust).

During all the time to 1971, Halifax was his home base but, whenever he returned U
Yorkshire between his travels, he spent most of his time, sometimes lengthy periods, a
Spurn where the Bird Observatory had been established in 1945 and which he helped ;
great deal in the early days. He also stayed at Eastleigh with his sister and moved his homt
base there in July, 1971. After his retirement he visited Spurn annually up to 1986 and
again, in 1990 and 1991, his final visits.

Rowland, George’s middle name, was chosen by his father as he was an admirer o
Rowland Hill, the 'stamp man’. There might have also been a painting connection a:
George’s father painted in his spare time. George kept up the interest in stamps and he wa;
certainly an accomplished artist. He painted many of the sea-birds he saw on his travels bu
his paintings became really well known at Spurn, not only for the ones he did there, somt
of which have been used on the covers of various publications, but also for the others h(
brought for sale. Many Spurn regulars must have examples of his work.

George had always had time to encourage an interest in birds and wildlife, especially ir
younger people. Frank Murgatroyd of Sowerby Bridge will always be grateful for hi;
encouragement during the early days of the Halifax Zoological Group, taking him oui
locally and further afield; Frank was on the 1936 trip to Spurn. Dennis Furnell who worker
at the R.S.P.B. in the 1950s remembers his time with George seemingly as if it was
yesterday. 'He was a small man with a big personality and a big voice. He was a great
influence and encouragement. He was a one-off, in the forefront of cinephotographers, the
first ol the really fine wildlife cameramen.’ Between 1952 and 1956, he organised a week
at Spurn each year for members of the Junior Bird Recorders’ Club (now the Young
Ornithologists’ Club).

The following appreciation by Roy Crossley, who grew up in Halifax, must express
some of the feelings of all who knew George:

'With his slightly lop-sided grin, his distinctive voice and somewhat
unconventional life-style George Edwards, or 'G.R.E.’ as we often called him, was
a charismatic figure who had a considerable influence on those of my generation in
the formative years of our youthful bird-watching.

Days spent birding with George were always enlivened with an endless fund of
anecdotes, usually amusing and often about the great and good of contemporary
British ornithology, for George seemed to know everyone who mattered in those
days. He inspired us Halifax youngsters with tales of birds beyond the hills and
valleys of our industrial West Riding, and those of us in the Halifax Junior Bird
Watchers’ Group in the 1940s who fell under the spell of his leadership have cause
to be grateful for his guidance and teaching, for he was a first rate ornithologist and
tutor.

Later on, when National Service took me to the Cotswolds, George provided
introductions to the local big-wigs who helped me to get the best out of my birding
there and later, when I went to Egypt, he was the person to whom I sent
descriptions of unidentified birds, being without adequate books in those pre-field-
guide days, and there was usually a prompt and always helpful reply, together with
news from the home birding front.

When I returned in 1953, George had gone to London to work for the R.S.P.B.
and we rarely met after that. But some years ago I heard that he was in brief
residence at Spurn and I went down the next free day that I had and managed to
find him at Warren Cottage. We had a chat about old times and, apart from looking
a bit older, he hadn’t changed a bit - the old sparkle I knew and loved as a teenager
was still there; he still had comic tales to tell!’

We mourn the passing of a line ornithologist and amiable companion who will long be
remembered and extend our sympathies to his sister and family.

John Cudworth

Book Reviews

145

BOOK RKVIEWS

tOcsert Sea: Fauna of the Saudi-Arabian Red Sea Coast by Hagen Schmid. Pp. 203,
ull colour photographs throughout. Hagen Schmid & Gazelle Book Services. Lancaster.
>nd edition, 1997. £28.99.

This is an unashamedly glossy book designed, primarily, to make one gasp at the sheer
leauty of the Saudi Arabian Red Sea. There is virtually no text, but the photographs are
ruly excellent. The book also has an interesting and attractive arrangement whereby the
photographs (one per page) do not hll the entire page, but are placed on a background ot
aim black and white images showing desert sands, corals or palm leaves.

It is a shame, perhaps, that the rationale in producing the book appears to have been to
! iccommodate the best available photographs and not to provide a more systematic
:overage of this fascinating region. There are, for example, nearly 130 images ot reel tish.
' lut only about 35 photographs of other reef organisms or scenes. This despite the tact that
he Red Sea has a fantastic (and photogenic) diversity of marine invertebrates (see, for
•';xample, Peter Vine’s excellent Keel Sea hiverlehrales). The final section ot photographs
- noves onto the land, with some fascinating photographs of wildlife, people and scenery
Vom the coastal zone.

Although good held guides do exist describing the marine and terrestrial launa ol the
'^ed Sea, it is still rather disappointing that the text is so minimal. Short paragraphs,
►scattered somewhat haphazardly throughout the book, do tell some of the interesting stories
1 about particular creatures or scenes, but the lack ot structure to the book leaves one feeling
hat much is missing. Overall however, this is a book about pictures, and as such is a great
■iiiccess: they are all works of art, and fantastically reproduced.

MDS

[ Loral Reef Fishes: Caribbean, Indian Ocean, and Pacific Ocean including the Red
'Sea by Ewald Lieske and Robert Myers. Pp. 400, including 175 colour plates. Princeton
University Press. 1996. Hardback $35.00.

This book, originally published in 1994 as the Collins Pocket Guide to Coral Reef Fish.

: oresents an excellent field-guide to coral reef fishes anywhere in the world.

A short introductory text provides a useful background summary, describing the reef
, environment and various aspects of reef fish biology, ecology, biogeography and behaviour.

' There are over 4.000 coral reef fish worldwide, but a large number of these are very small or
cryptic and are therefore unlikely to be seen, or at least noticed, by most observers. This
vvork deliberately ignores such species and provides a remarkably comprehensive coverage
af the remainder. Over 2.000 species are described and illustrated, with depictions of
Jifferent forms - male, female and immature, or geographical varieties - for some species.
Text descriptions are necessarily brief (10-15 species described per page), but useful and
iecurate. The illustrations themselves are very good indeed, with colours being very true to
life in almost all ca.ses. and with remarkable attention to detail, particularly for the salient
identifying features. (N.B. The text and the illustrations for plates 134 and 135 have been
switched. This error is present in all three editions so far published!)

Books on reef fish in particular countries or regions are increasingly available, many
oeing well illustrated with colour photographs. This book, however, fills a niche for
miateur and professional visitors to reefs, combining in one compact volume what has
previously required several, usually expensive, works. The full identification of many reef
fish from underwater sightings alone is often difficult, but not impossible. Using this book
as a first point of reference '"will be sufficient for many users, while others may wish to
-combine it with other works, particularly those showing reef fish in situ with colour
photographs, to provide a more secure base for reef fish identification.

MDS

146

Book Reviews

Basic Geological Mapping by John Barnes. Pp. x + 133, with 45 maps and diagrams anc
23 b/w photographs. 3rd edition. John Wiley. 1995. .£1 1.99.

This sturdy, conveniently sized paperback provides a wealth of good advice on how to
measure, record, and write reports of geological field observations. It is clearly targeted at
students with a grasp of basic geological concepts but little experience in field recording
and who need to collect their own field data.

It provides sound information on suitable equipment and commonly available types of
geological maps and briefly outlines the principles of cadestral surveying. There are
excellent chapters covering field measurements and techniques of rock description and on
maintaining an effective fieldbook. Advice is given on preparing reports and geological
sections and the whole is rounded off with guidance on field safety.

The recent trend in geological, geographical and environmental sciences towards
reduced field teaching suggests that there should be a substantial number of students who
would benefit from this book, as its simple, clear common sense approach is appropriate
not only to geologists, but to a wide range of people who would profit from advice on how
to collect field data.

DEC

The Birdwatcher’s Guide to Hawai’i by Rick Soehren. Pp. 215, with many b/w and 19
colour photographs and 7 maps. University of Hawai’i Press. 1996. $17.95.

Another book in the now very popular series of birdwatching guides. The Hawai’ian
archipelago is made up of 132 islands and this guide deals with six of the eight main ones
(two being private and out of bounds) which make up over 99% of the total land area. Each
island is dealt with in detail, including a check list of birds for each and listing the best
birdwatching locations (O’ahu 13, Kaua’i 6, Big Island (Hawai’i) 8, Maui 6, Malaka’i 4
and Lana’i 3). Each site chapter is very comprehensive, including acce.ss details and the
species most likely to be seen at any particular season. There are short chapters on the
habitats, conservation, accommodation, transport and organised hikes. Written by an
American who knows the area very well, the book is inevitably aimed at the American
market and it is a pity that few British birdwatchers will venture to this remote part of the
world, particularly so now that this guide is available, but. for those lucky enough to get
there, for whatever reason, this guide is es.sential.

JRM

The Birds of the Northallerton and Richmond Area by N. J. Morgan. Pp. 76, with line
drawings. Privately published, Ainderby Steeple, 1996. £3.00 paperback.

This booklet arose from the author’s keen interest in the area and his desire to produce a
base-line document for future researchers and he is to be commended for his effort.

The introduction is followed by brief details of the 13 major birdwatching sites in the
region (with map references) and a short review of the area’s birdwatchers, past and
present. The classified list takes up 67 of the 76 pages and deals with 260 species, 130 of
which have bred. Each species is thoroughly documented with status, favoured sites and
historical details, and local names are given for some.

A most worthwhile project and having regard for the amount of research required for
even a small booklet like this, well worth the bargain price of £3.00.

JRM

Contributors

147

Microfungi on Land Plants: an Identitication Handbook by Martin B. Ellis and J.
'Pamela Fdlis. Pp. x + 868, ind. 213 pages of line drawings. Richmond Publishing, Slough.
1997. £60 hardback, £4().()() paperback.

A much revised and enlarged edition of a standard work published in 1985 by Croom Helm
which, due to its success, went out of print in 1992. Its return will be welcomed by
mycologists and plant pathologists for it is unparalleled in terms of its identification value
in both the field and laboratory. Although concentrating on the British Isles, it will prove
useful in many north temperate countries, especially in Europe and North America.
Through keys, descriptions and exquisite line drawings, this work deals with c. 3500
■ipecies, including ru.sts, smuts, powdery mildews, discomycetes, pyrenomycetes,
loculoascomycetes, hypomycetes and coelomycetes in sufficient detail for accurate
determination. A glossary and indexes to fungi and host plants are also provided. This
indispensable landmark in mycological literature is both a practical guide and a reference
uturce and is therefore strongly recommended to both field workers and libraries.

MRDS

CONTRIBUTORS

Limbert. M. 88-92

Mather, J.R 29-30,146
Medd,T. F. 62-63,65
Messenger, J . E. 13-18

Norris, A. 27-28,31-32

■Abbott, P. P. 64 -65
Archer, M. E. 45-52, 59, 97-99

Bartley, D. D. 53-56
Beaumont, H. E. 30-31
Birks,J.D. S. 13-18
'Bunce, H. O. 33-43

Collingwood, C. 56

■ Cook. P. J. 43-44
Coppins, A. M. 11-12
Cotton. D.E. 99-100.146

■ Crackles, F. E. 44. 6 1 -62. 67-68, 79-83

■ Crossley, R. 60. 93-97
Cudworth, J. 143-144

Davison. A. 13-18
Denton. M. L. 69-79

Emsley, K. 24-26

Grant, D. R. 61-67

Hale. W. H.G. 31
Henderson, A. 101-121
Hull. S. 137-142

Oybak. E. 53-56

Roberts. J. 137-142

Scott. G.W. 137-142

Seaward. M. R. D. 18. 57-59. 100. 147

Skidmore, P. 28-29

Spalding, M. D. 145

Vaughan. N. 3-11
Vaughan. R. 3-11.32,33-43

Wardhaugh. A. A. 19-23
Wilmore, G. T. D. 65-67

Yalden. D. W. 83-88

148

Index

INDEX

\

Book Reviews

11-12, 18,24-26,28-32,56,59,99-100, 135-136, 142, 145-147. i

Botany

Maritime Fescues in S.E. Yorkshire, 43-44; Field note, 44; Botanical report tor 1996,
61-67; Wansford to Driffield site of special scientific interest, 67-68; The vegetation ot
Spurn (1946-1996), 79-83.

Bryology

Lateglacial and postglacial moss records from a valley bog in the Ingleborough region,!
53-56. I

Coleoptera

The beetles of Spurn Peninsula: an update, 69-79.

Crustacea

The Horse-trough Ostracod Heterocypris incon^ntens, 121-135.

Diptera !

Two recently described flies in Yorkshire, 60; Notes on the Marsh Flies ot Yorkshire, |
93-97.

I

Hymenoptera

The aculeate wasps and bees of two calcareous localities in Watsonsian Yorkshire, 45-52;
Recorder’s Sixth Report of the aculeate Hymenoptera in Watsonian Yorkshire, 97-99.

Lichenology

Progress in the study of the Yorkshire lichen flora. 57-59. t

Mammals

A 1994 Pine Marten record for Lancashire, including preliminary genetic analysis, 13-18;
The current status of the Water Vole in the Peak District. 83-88; From Coney to Rabbit: the
story of a managed coloniser, 101-121.

Molluscs

Terrestrial Molluscan fauna of Wilton Wood, 19-23.

Obituary I

John Armitage ( 1 900- 1 996), 27-28; George Rowland Edwards (1910-1 996), 1 43- 1 44. ^

I

Odonata

The White-faced Dragonfly on Thorne Moors, 88-92

Ornithology

Changes in populations of Kittiwakes and Common Guillemots breeding on the
Flamborough Headland, 1967-1994. 3-11; Snipe 32; The disappearance of the Stone-
Curlew as a breeding bird in Yorkshire, 33-43; Diet of Barn Owls in a lowland area of
North Yorkshire 137-142.

Palacobotany

l.ateglacial and postglacial moss records from a valley bog in the Ingleborough region,
53-56

Irish

Naturalists’ Journal

The Irish Naturalists’ Journal, successor to the Irish Naturalist,
commenced publication in 1925. The quarterly issues publish papers
on all aspects of Irish natural histor>’, including botany, ecology,
geography, geology and zoology. The Journal also publishes
distribution records, principally for cetaceans, fish, insects and
plants, together with short notes and book reviews.

Current subscription rates for four issues (including postage) are - £IR15.00
(£14.00stg); Students IR£4.00 (£3.50stg). Further details may be obtained from Ms
Catherine Tyrie, Ulster Museum, Botanic Gardens, Belfast BT9 SAB.

YORKSHIRE BIRD REPORTS 1961 to 1990

COPIES ARE STILL AVAILABLE
for 30p. each (postage and packing free)

Note:- 1972, 1983 and 1986 are sold out
and for 1 973, 1 987, 1 989 and 1 990 - 6 or fewer copies are available

Apply to:

ATHOL J. WALLIS, 51 RED SCAR LANE, SCARBOROUGH Y012 5RH

Telephone: 01723 361657

‘The Naturalist’ is available in microform

UNIVERSITY MICROFILMS INTERNATIONAL

For further information contact;

UMI, ATTN: Box 38, PO Box 1346,

300 North Zeeb Road.

Ann Arbor. MI 48 1 06- 1 346,

USA

Printed in Great Britain by Titus Wilson & Son, Kendal

ISSN 0028-0771

Latest publication of the Yorkshire Naturalists’ Union

THE FRESHWATER CRUSTACEA OF YORKSHIRE

a faunistic & ecological survey

GEOFFREY FRYER

The crustacean fauna of Yorkshire reflects the great physiographic diversity
of the region. Adopting an ecological approach, this book considers the
Yorkshire fauna in relation to climate, topography, geology, soils and water
chemistry, always keeping in mind that it is dealing with living organisms
whose habits, requirements and physiological limitations determine exactly
where they live.

Matters covered include the ecological background; faunal assemblages
and their regional attributes; an analysis of the factors that determine
distribution patterns, many of which are mapped; wide geographical aspects;
and conservation. Large areas, such as the Pennines, Howgill Fells, North
Eastern uplands and the lowland plains are surveyed. So too are localised
regions including Whernside, the Malham area, lowland heaths, and the largest
lakes, as well as habitats such as upland tarns, seepages, cold springs, small
lowland ponds, inland saline waters. Notes are given on every species
recorded, including parasitic forms.

Price £16.00 (plus £2.00 per copy p.&p.) Special offer to members of
the Yorkshire Naturalists’ Union £13.50 (plus £2.00 p.&p.)

Please make cheques payable to Yorkshire Naturalists’ Union.

Available from: Professor M. R. D. Seaward, Department of Environ-
mental Science, University of Bradford, Bradford BD7 IDP.

PUBLICATIONS FOR SALE

A Fungus Flora of Yorkshire. 1985. 296 pp. Hardback. £10.00 incl. p&p.

Butterflies and Moths of Yorkshire. 1 989. 380 pp. Paperback. £1 7.50 incl.
p&p. Unbound. £12.15 incl. p&p.

Mammals of Yorkshire. 1985. 256 pp. £7.50 incl. p&p.

Provisional Keys to British Plant Galls. 1986. 85pp. £5.50 incl. p&p.

First Yorkshire Lepidoptera Report \n 'ARGUS' Spring 1997. £2.50 incl.

p&p.

Moths and Butterflies of Spurn, 1 995. 1 24 pp. £6 incl. p&p.

Cheques should be made payable to Y.N.U.

From: Mrs ). Payne, 15 Broad Lane, Cawood, Selby, North Yorkshire, Y08 OSQ

Tele|)hone: 01 757 268242

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