Nm- Phytol. (1983) 95.

153-157 ^ 53
THE IDENTITY AND ORIGINS OF PICEA ABIES
(L.) KARSTEN FROMTHE CHELFORD
INTERSTADIAL (LATE PLEISTOCENE)
OF ENGLAND
By D. T. HOLYOAK
Quaternary Research Unit, Department of Botany, University of Reading,
Reading RG6 2AS, UK
{Accepted 27 April 1983)
SUMMARY
Pollen and mscrofossils of Picea from the Ghetford Ititerstadia! {Early Devensian) of England
are described. All leaves and cones were referred to P. abtes (L.) Karsten, and the unimodal
size distribution of pollen also suggests on!y this species was present. The fossil leaves are mostly
similar to those of P. abies subspecies obmata (Ledeb ) HuJten, but the cones are intermediate
between subspecies iihoiata and abies. Occurrence of forms approaching the north-east
European subspecies obm^ata does not necessarih' imply immigration oi Ptcea into England from
the east or north-east during the Early Devengian because similar intermediate forms also occur
in the Alps.
INTRODLCTION
Tlie Chelford Interstadial of the Early Devensian is distinctive in the Late
Pleistocene of Britain mainly because of the plentiful representation of Picea. In
contrast the preceding Ipswichian Interglacial had only ver>- low frequencies of
Picea pollen (West, 1980).
The Chelford Interstadial deposits have been correlated with those of the Brorup
Interstadia! of Denmark (Simpson and West, 1958). Occurrence of two Picea pollen
t^'pes has been reported from north-west European Brorup age deposits, and these
have been referred to P. abies (L.) Karsten and P. omorikoides Weber (Andersen,
1961; Zagwijn, 1961). PoUen from the Chelford Interstadial type site in Cheshire
was found however, to resemble that of P. abies (Birks, 1978). Leaves from deposits
correlated with the Chetford Interstadial at Beetley, Norfolk were identified as
P. abies cf. subspecies obovata (Ledeb.) Hulten by Phillips (1976). Whitehead
(1977) likewise identified five incomplete cones from Chelford as P. obotata Ledeb.
rather than P. abies. Birks (1978) showed that Picea pollen from Chelford was
similar in morphology' to his 'group B' , the predominantly Scandinavian and
Russian type of P. abies pollen (although this type was also found in one sample
from Yugoslavia). Subspecies obovata is the form of P. abies native in north-eastern
Europe (Tutin et al., 1964). so on the basis of the occurrence of the Scandinavian
type of pollen and the macrofossils attributed to subspecies obovata, Birks {op. cit.)
suggested that in the Early Devensian this subspecies had a larger range extending
westwards to Britain.
This paper reports results of study of additional Picea pollen and macrofossils
of Chelford Interstadial age, from the t>pe site (Farm Wood Quarry, Chelford),
Acre Nook Quarry close to the type site, and from Brimpton, Berkshire (Bryant,
)28.546X/83/o9oiS3+05 S03.00/0 Q 1983 Thf Ne* Phrtologist
154
D. T. HOLVOAK
Holyoak and Moseley, in press), leading to a reappraisal of the direction from which
these Picea may have reached Britain.
LEAVES
Phillips (1976) identified leaves from Beetley as P. abies cf. subspecies obotata
because they were more robust than is usual in subspecies abies, and mostly had
more rows of stomata. Leaves of subspecies obovata typicalJy have four stomatal
rows on each of the two \ entral faces, and three on each dorsal face, compared to
two or three (ventral) and two (dorsal) in subspecies abies. A majority of the fossil
leaves from Beetley had the larger numbers of stomatal rows characteristic of
subspecies obovata, although some had fewer.
Table 1. Measurements of leaves of Picea abies
Picta abies ssp. abies
Uppsala, Sweden (n = 20)'
Suffolk (cultiv.) (n = 20)*
Bedgeburj. V.K. (cultn.) (n = 50)
Ptcea abies ssp. obotata
Russia (n = 20)*
.Mtai Mts (n = 20)
Finnish Lapland (n = 20)*
N'E Finland (i = 50)
Fossil leaves
Bt!ey (n = 500)*
Brimpton (n = 50)
Acre Nook Quarry. Chelford (n = 50)
Farm Wood Quarrv, Chelford. (n = 50)
Stomatal rows
(one ventral face:
one dorsal face)
3: 2( 3: 2- 2: 1)
2: 2( 3: 2- 2: ) )
3: 2( 3: 2 2:1)
4: 3( 7: 5 2:1)
4: 3( 7: 5 2:1)
4: 3( 5: 4- 3: 3)
4: 3( 6: 4 2: t )
4: 3( 8: 4 2:1)
4: 3( 7: 6- 2: 1)
4: 3( 6: 4- 2: 1)
4: 3( 7: 4- 2: 1)
Leaf width
(mm)
0-7 (0 5 0-81
(^9(()-8 1 1)
0-8(0-7-1 1)
1-2(0-8 1-4)
1-0(0-8 1-6)
1 2(0-1* 1 6)
1-1 (0-8 1-5)
!-0(0-6-l-9)
1 3(0-8 1-5)
1-1 (07-1-6)
1-1 (0-8 16)
Leaf length
(mm)
10(8-13)
11 (9-13)
11 (813)
13(11 161
12(7 !5)
11 (8 14)
12(8 14)
10(7-14)
10(7 13)
11 (7-14)
11 (6 14)
Data from Phi]lip.s, 1976
Examination of 50 leaves from Brimpton (Sample 59J), 50 from Acre Nool
Quarry, Chelford and 50 from Farm Wood Quarry, Chelford showed that thes
also most often had four rows of stomata on each ventral face and three on ead:
dorsal face (Table 1). Numerous individual leaves in each of these three sampfe
had fewer rows and some had more. Leaf width and length were very variable ir
each sampJe; statistical treatment of measurements was avoided because a prevalenci
of short stout leaves in at least the Brimpton sample might have been due to mon
frequent breakage of long thin leaves. However, comparisons of the overlappins
ranges of stomatal row numbers show that the Chelford and Brimpton lea
samples resemble those from Beetley in suggesting more similarity to subspecie
obovata than to subspecies abies.
CONES
Whitehead (1977) identified five partly incomplete Picea cones from Chelford as
P. oboz-ata Ledeh. on the basis of the rounded cone scales. However, m.'
observations show that the majority of cones from Chelford have damaged conf
scales. In samples of 99 cones from Farm Wood Quarry and c. 30 from Acre Nw'
Interstadial Picea abies
155
" " ^' " "''P '*^ '^'"'''' "*"' subspccifs ahies/ohmata. Chelford Interstadial (Farm
suh ^'^'^- Chelford, Cheshire; Coll Dr. P Worsley). t o views of same cone, (b) Ptcea ab,es
K I M ' , ' "' ' "*" ' "<' Chelford Interstadial (.Acre .Nook Quarry. Chelford, Cheshire- Coil Dr
(Loiu., ^ ** "*'" ' " * ' "f '^^ ""'"' "'" f"^'' <''> ^"''' " ' " " subspecies ahmata. modern
Q()if m 7 * ^ " ' ' " ' ' ^" " " ^' ''^'^'* ^ ' ' ^"' ""' ' ' ' ' ^' ' "^"""' ' P'"dS. (e) Aff a6, aubspccics
Kewi m P ' ' ^"' ' "^"' ' ' Bedgehury National Pinetum, England: per Royal Botanic Gardens,
'0 (i) P " " s"hspecies abies. modern (Haute-Savoie, France: Coll. D T. Holyoak). (g)
Kea omortka. modern (cultivated at Bedgebury National Pinetum, England: per Royal
Botanic Gardens. Kew).
156
D. T. HOLYOAK
Quarr\' the small fraction of cone scales that had survived completely intact were
found to be less rounded than in typical subspecies obovata [Fig. l(a), (b) cf. (c),
(d)], but instead to approach some subspecies abtes in form of the cone scales [cj.
Fig.' 1(0]- Furthermore, the intact cones from Chelford were larger than is usual
in many populations (not all) of subspecies abies, which itself usually has larger
cones than the 6 to 8 cm typical of subspecies obovata.
POLLEN
Picea pollen from Early Weichsehan interstadial deposits in north-western Europe
has been found to include both P. abies type and smaller grains that Andersen
(1961) and Zagwijn (1961) attributed to P. omorikoides Weber. The latter species
is based on Pleistocene fossils from Germany; the very detailed original description
(Weber, 1898) shows that it is similar to P. omorika (Pancic) Purkyne, which is
known as a native tree only in central Yugoslavia (Jalas and Suominen, 1973), and
It may be most simply regarded as conspecific with that species. Aleaf tentatively
attributed to P. omorika has been reported from the Pastonian of Norfolk (Wilson,
1973), but there appears to be no Late Pleistocene record of macrofossils from
north-western Europe.
Measurements of Picea pollen from Brimpton (biozone Br-C) and the Chelford
Interstadial type site show unimodal size distributions and give no indication that
P. omorika might bave occurred (Fig. 2). Cones of P. omorika [Fig. l(g) to (j)] and
leaves (Weber, 1898, Taf. XI) are distinctive and nothing resembling them was
found in either the Chelford or Brimpton samples.
20-
0 -
20 r
lOf
n
ID)
n
16 20
24 28 32 36 40
Mi cromet er l i nes
Fig. 2. Measurements of pollen (height of air-sacs) of Picea sp. (a) Brimpton. n = 60; (h) Cheiford
n = 148; (cj modem P. at)ies (Norway), n = 176; <d) modem P. omorika (Yugoslavia), n = 80.
DI SCUSSI ON AND CONCLUSI ONS
The P. abies from the Chelford Interstadial thus appears to have had leaves mosti>
similar to those of subspecies obovata, cones intermediate between subspeci
obovata and abies, and pollen (fide Birks, 1978) of a predominantly Scandinavia^
type. However, although subspecies obovata is the north-east European foi^
occurring mainly in western Siberia (USSR), it would be premature to
Interstadial Picea abies 157
this form colonized Britain from the east or north-east during the Early Devensjan.
Forms more or less intermediate between subspecies abies and subspecies obovata
are well known in the Alps [Tutin et al., 1964; Weihe, 1972; Fig. 1 (f)] and Poland
^Srodon, 1967), as well as in Fennoscandia. In their combination of leaf, cone and
pollen characters the Cheiford Interstadial Picea do not correspond closely to any
of these living intermediate popuiations. However, some of the morphological
characters of the leaves, cones or pollen might be affected primarily by local
environmental factors (e.g. daylength, humidity, temperature) rather than being
under simple genetic control. Consequently, immigration oi Picea in the Chelford
Interstadial might have been from the north-east, east or south-east, and the
characteristics of fossil pollen, leaves and cones may not allow discrimination
between these different origins.
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