Land biotas of the last interglacial/glacial cycle on Jameson Land, East Greenland OLE BENNlKE AND JENS BOCHER

Bennike. 0. & Bocher. J . 1994 (December): Land biotas of the last interglacial/glacial cycle on Jameson Land. East Greenland. Boreas, V O I . 23, pp. 479 487. OSIO. ISSN 0300-9483. Macrofossil plant and insect remains from nearshore marine sediments in Jameson Land, central East Greenland show that the land biotas of the last interglacial stage, the Langelandselv stage, were more diverse than those of the Holocene. Rich dwarf shrub heaths with a diverse assemblage of ericaceous plants occupied low land areas with copses of Betula pubescens on sheltered sites. Many southern extra-limital species were present, and the mean summer temperature was c. 5°C higher than today. The subarctic bioclimatic zone was displaced from southernmost Greenland/lceland to central East Greenland. The diverse beetle fauna was of palaearctic affinity and strikingly different from the Plio-Pleistocene and the Holocene Greenlandic beetle faunas. A few fossil assemblages from the Hugin S0 Interstade. which is correlated with oxygen isotope stage 5c (early last glacial stage), point to poor, perhaps entirely herbaceous vegetation with a mean summer temperature that was perhaps 3 4°C lower than today. Ole Bennike, Geological Surwy of Denmark. Thoraoej 8, DK-2400 Copenhagen N V , Denmark; Jens BBcher. Zriological Museum, University of Copenhagen. Universitetsparken 15, D K - 2 / 0 0 Copenhagen 0, Denniark; 23rd March. 1994 (reiiyed 29th June. 1994).

BOREAS

Until recently, no remains from terrestrial biotas in Greenland could confidently be referred to thc last interglacial/glacial cycle. However, from the Thule area, Northwest Greenland, a pre-Holocene occur- rence points to warmer summer tempcratures during deposition than at present (Bocher 1989a; Funder 1990; Bennike & Bocher 1992). This deposit was orig- inally correlated with oxygen isotope stage 5a, but is now correlated with stage 5e (Kelly, pers. comm., 1993). Another pre-Holocene occurrence in central North Greenland provisionally dated to the last inter-

glacial period is indicative of climatic conditions much like those of today (Meldgaard & Bennike 1989).

During the PONAM expedition to Jameson Land in 1990 several samples of sediment with organic detritus were collected in the Hesteelv and the Lange- landselv areas (Fig. 1). Seven samples were handed over to us for study (Bocher & Bennike 1991). During the PONAM expedition to Jameson Land in 1992 large amounts of additional material were sampled from southern Jameson Land, and the aim of this paper is to report on this material.

Scoresby Sund Hesteelv

L 20 km 1

Fig. 1. Map of southern Jameson Land showing PONAM-numbered locations of the samples listed in Tables 1 and 2.

4 x 0 OIc Hennike und Jens Riicher ROREAS 23 (1994)

The lowland vegetation of Jameson Land today is dominated by various types of dwarf-shrub heaths. The most common species is Cussiope tetragonu, but heaths rich in Betulcr ncinu occur in sheltered places. Poorly drained sites are covered by mires and mead- ows, and exposed sites by fall-tields. Snow-patch com- munities are of limited, scattered occurrence (Ray & Holt 1986).

At present, a total of only 33 species of beetles are found in entire Greenland and merely three species have been recorded from Jameson Land, although four species can be expected there from thcir overall modern distribution in East Greenland ( Bijcher 1988).

Material and methods The samples were wet sieved through 0.42 and 0.31 nim sieves, and the residue left on the sieves sorted tinder a dissecting microscope. Part of the material was washed through 0.6 and 0.35 mm sieves and flotated with kerosene ( i n accordence with Coopc 1986a). Sample size varies greatly. but a total of about one ton of sediment has been analysed, corresponding

to 74 samples. All come from the south and west coasts of Jameson Land, between Falsterselv and Hes- tech. For details of sample sites. stratigraphy, correla- tions and datings, see Hansen et al. (1994), Ingblfsson et u1. (1994), Landvik et ul. (1994), Lysi & Landvik (1994), Tveranger et al. ( 1994), Vosgerau rt (11. ( 1994), and summary by Funder et al. ( 1994).

Results and discussion Lists of fossil animals and plants are presented in Tables 1 and 2 and some fossils are shown in Figs. 2 and 3. Some of the samples contained remains of marine organisms, such as foraminifers, hydroids, bivalves and ostracodes, and most of the sampled scd- iments were deposited in shallow water marine envi- ronments. All remains of terrestrial and limnic plants and animals are thus allochthonous.

Most samples consist of very fine. often mica-rich sand that is dominated by hydrodynamically light re- mains such as moss fragments, Sci1i.v herhuceri leaves and sclerotia of Cetzococciini genpliiluni. whereas some samples consist of fine or medium sand character-

Fig. 2. SEM photographs of selected plant fossils from Jameson Land. 7 A. Sclugincllu sduginuirles (Selaginellaceae) sporophyl. ref. no. MCUH 22748. I : R. A1nir.c. cf. crI.spu ( Betulaccae) female catkin scale, M G U H 22249. 1 C. Violu pulusrris (Violaceae) seed. MGUH 22250.

D. .-Irc/os/tri,//J'/(J.S tdpino (Ericaceae) endocarp. MGIJII 22 171. _I E. Cussiupc, te/rugontr (Ericaceae) leaf. MGIJH 2225 I . r F. /.oi.w/ciwiu procuriihc~ru ( Ericaceae) leaf, MGUH 70732. 17 G. Dnprtrurrr nigrurir ( Empetraceae) leal; M G U H 20734. L H . Dryus uc/optv/ala ( Kosaccae) leaf. MGUH 22257. P 1. Turasacunr (Asteraccac) sp. achene, MGUH 22253. 1 J. Poramogcfon pu.dlus (Potamogetonaceac) endocarp, MGUH 72254. The scale bars are I mm long. The specimens are housed in the type collection of the Geological Muscurn, Copenhagen (MGUH).

ROREAS 23 ( I Y Y 4 ) Land hiotas. Jameson Land 481

Tuhlr 1. Macrofossil plant remains from Jameson Land.

Age’

Sample no.

+ + r r t + r - c c r r r r c c r c f r + + + + t + + + t t + + + + + + + + + t

r s r r - - c s r r r - - r r - - . . . . . . . . . . . . . . . . . . . .

- - s 1 - - - 3 - - ] _ _ - -

3 r c r r l c - s - - a r - r f r r - I - 1 r - I - . - - - - 3 - - - 2 - 1 - -

_ _ _ + . - + - - + + + + + - + - - - -

s c I s r - - .

I - ? ? s - - s s r r r - - r r r r s s r s r - I - - r r .. . s -

s = Scarce: r = rare: c = common: f = frequent: a = abundant; + = present: - = absent: I 9 = Absolute numbers ’ LI = Langelandselv interglacial stage; HI : Hugin S0 interstade.

Nomenclature follows B6cher e / d . ( 1978). One calyx in X2415. Ioc. 150.

ized by hydrodynamically somewhat heavier remains, mostly twig fragments and diaspores. Thus some of the variation in macrofossil content reflects tapho- nomic processes. The largest stick found was 20cm long and measured 16 mm in diameter.

The animal and plant remains show varying signs of physical erosion, and they are often rather poorly

preserved as a result of the formation of hydrated iron oxides.

Bryophyte remains are common in many samples. Remains of Bryopsida are by far the most common. but Sphagnopsida and Hepaticopsida are also rep- resented. The bryophyte remains were studied by L. Hedenas, Stockholm ( Hedenas 1994).

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1758

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s (F

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( LeC

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63)

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1827

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0. F

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, 17

80)

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179

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pre

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I LI

= L

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land

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cial

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ge; 1

11 =

llug

in S

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tade

: C

= C

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lar d

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Pal

aear

ctic

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trib

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n: G

= E

xtan

t in

Gre

enla

nd. I

n th

is p

aper

Gre

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BOREAS 23 (1994) Land biotas, Jameson Land 483

Fig. 3. SEM photographs of the beetle O!iorlijnchus nodomc. head ( A ) and pronotum (B).

Stratigraphically, the fossil assemblages belong to the Langelandselv interglacial stage and to the Hugin S0 Interstade. correlated with the last interglacial - E stage and the earliest Weichselian, respectively (Fun- der e f NI. 1994). D

2.0

Tlir t,utigi~luti~i.srlt: intrrgluciul

This stage is correlated with the last interglacial stage, the Eemian or substage 5e in the oxygen isotope record ( Funder e f al. 1994). The assemblages (Tables 1 and 2) are characterized by high frequencies of southern ex- tralimital species. Remains of dwarf shrubs dominate the fossil assemblages, and their diversity shows that rich dwarf-shrub heaths with a diverse assemblage of ericaceous plants grew in southern Jameson Land. One of the most unexpected finds is that of Betulu puhrscms. a northern tree birch, and a closer study of its remains was undertaken. The birch nutlets rarely

* a . . . . . . . . . .

. a . a a - . . . . . . ...... . .

1 .o

I I I ,

1.5 2.0 2.5 3.0 Length (mm)

Fig. 4. Scatter plot of the length breadth ratios of the central body of fossil birch nutlets. The large dots mean that two nutlets were of the same size. The mean length is 2.30 m m and the mean breadth 1.45 mm.

retain their lateral wings, which makes positive identifi- cation difficult. The central body of the nutlets has been analysed biometrically. Measurements of the length- breadth ratio reveal that only one size population is present in the material, although the variation is fairly large (Fig. 4). Comparison of the mean length-breadth ratio with modern samples of dwarf birches (Betula nana and B. glandulusu) and the tree birch B. puhescens shows that the fossil nutlets are referable to the latter taxon (Fig. 5 ) . The shape of the female catkin scales (Fig. 6) also agrees with B. puhescens, and the same applies to those nutlets that retain the lateral wings. No remains of Brtula sect. Nanue were found. In the absence of tree trunks it is impossible to say whether the Brfula puhescens remains represent trees or shrubs,

since this species, although a member of the tree birches (Betual sect. Afhae), today grows north of the arctic tree line, where it forms copses. We suggest that Befulu puhescens copses occupied well-drained sheltered local- ities. Alnus cf. crispa is another copse-forming species, and Selaginella selaginoides and Viola Palustris may also have grown in such sites. Most of the herbs represented have a fairly wide ecological amplitude and are found in dwarf-shrub heaths as well as in fell-fields. The Dryas leaves are narrow and referred to the wind adapted type of plants growing in exposed situations.

The common presence of Salix herhacea leaves indi- cates that snow-patch vegetation was widespread and that snowfall was heavier than today.

484 Ole Bennike und Jrns Biicher

. I .4

A 3 2

B. pubescens

rn B.nana

A B. glandulosa

1.0 2.0 Length (rnm)

Fi,y. 5. Mean length-breadth ratio of the central body of the fossil birch nutlets from Jameson Land ( 5 and 7 ) compared with n~odcrn samples of three different birch species. I from Smiland, Sweden; 2 from C'harcot Land. East Greenland: 3 from Akia. South Grcen- land; 4 from Ssndre Strornljord, West Greenland; 6 from Fnjos- kndalur, Iceland; 8 from Tunugdliarfik. South Greenland: 9 from Hi iy l jc ldct . Troms. northern Norway. Each of the modern samples represents 50 nutlets from one or a few catkins from one plant. All intcrnicdiatc forms can he found between the birch species. but such intcrmcdiatc forms were not used f o r measurements.

Fi,.y. 6. Thrcc fossil lciilale birch catkin scales from Janicson Land. MGtIII 10736 38.

Most of the plant species grow in well-drained dry or moist localities. and beetle species such as Amurtr trlpinrr. C\wiitidis r:aporurioriini, Byrrlzus jasciatus, Nephirs h is ip i tus . Otiorliynchus urc/ic~us and the ly- gacid bug Nj~siu.7 groerilandiciis are eurytopic in fairly dry plant communities, whereas Otiorhyni*hus noiiosus prcfcr moderately humid hcaths. The majority of the C'uri,.~ species probably stem from mires, meadows or othcr poorly drained sites, and the beetle species EUC- riworiim teniic, Olophrunz consin~ile and Stenus car- hoiiurius are hygrophilous, associated with similar damp biotopcs, while Eluphru.7 rirherciilatits and Sirii- plourria nietdlic'ir are usually found on freshwatcr shores.

Remains of freshwater plants are relatively rare, but represcntcd by the aquatic plants Poturnogeton and 1lippitri.s uztlgrrris and the semiaquatic plant Menyan- rhcs rrlfdiata. The tadpole shrimp Lepihrus was found in wine samples, and watcr insects are repre- scntcd by the beetle genera 1Iyriroporu.s. Agubiis and C ' c i l ~ ~ m i h i ~ r v s and by midgc (Chironomidae) larvae. The

BOREAS 23 (1994)

plants and animals are indicative of ponds or lakes in the region.

Betula puhescens can endure a mean summer tem- perature slightly below 10°C. Alnus cf. crispa, Se- luginelb selaginoides and Violu palustris have northern range limits in the low arctic zone. The present iso- lated occurrence of S. selaginoides in East Greenland south of Scoresby Sund is connected with a hot spring (Halliday el al. 1974). Potantogeton pusillus is com- mon in West Grecnland, where it has a northern limit of distribution at 72'45" (Bocher ef ul. 1978). The lack of this water plant in East Greenland is pre- sumably a consequence of delayed immigration. Two of the identified plant species. Menyunfhes trifoliutu and Loiseleuria procurnhens, have a northern range limit in Jameson Land today. Thus the most warmth- demanding plant species of the fossil assemblages in B. puhesrens, which indicates an increase in mean summer temperatures of about 5-C during the last interglacial maximum as compared with the present, and 3-4'C as compared with the mid-Holocene warm period when dense dwarf-shrub heaths - but without Betitla puhescens and Alnus cf. crispa - were wide- spread in Jameson Land (Funder 1978). This means that during the Langelandselv interglacial the subarc- tic bioclimatic zone was displaced from southernmost Greenland/Iceland to central East Greenland.

The modern ranges of the fossil beetle species are indicated in Fig. 7. It appears that in spite of very different distributions all the species are found in the subarctic zonc (the Befula 7onc) of the boreal region.

There are some indications that Brrulu puhe.wn.v and othcr southern extralimital plants became rare or absent towards the end of the Langelandsclv intcr- glacial, but the beetle fauna does not substantiate this trend. It is possible that the dccrcasc in southern extralimital plants is due to increasingly colder cli-

HA LA SA B NT T ST

Elaphrus tuberculafus Amara alpina Cymindis vaporarium Hydroporus morio Hydroporus planus Colymbetes dolabratus Agabus bipustulafus Eucnecosum tenue Olophrum consimile Stenus carbonarius Nephus bisignatus Coccinella hieroglyphica Ofiorhynchus arcticus Otiorhynchus nodosus Lepyrus arcticus Lepyrus nordenskioeldi Nysius groenlandicus

Fix. 7. Present climatic ranges of the intcrglacial insect species from Jameson Land. HA =high arctic; LA =low arctic: SA 2 subarctic ( =northern boreal); R - boreal; NT = northern temperate; T = temperate; ST = subtropical.

BORF.AS 23 (1994) Land biotas, Jumeson Land 485

i Betula pubescens Spermatophyta, Betulaceae r S>

Viola palusfns Spermatophyta, Violaceae

Men) Sperrnatophyta, Menyanthaceae ,

Alnus cnspa Spermatophyta, Betulaceae

Fi,g. 8. Circumpolar maps showing the modern geographical ranges of some o f the fossil plants recovered, based on Hulten & Fries (1986), and 01‘ one beetle. based on Holdhaus & Lindroth ( 1939) and Riicher (1988). Arrows indicate Jameson Land.

486 Ole Bennike ond Jens Biicher ROREAS 23 (1994)

mates towards the end of the last interglacial stage, but it is also possible that the decrease reflects tapho- nomic processes.

The Ilugin SH interstade This interstade is correlated with oxygen isotope stage 5c (Funder r t ul. 1994). The assemblages here are characterized by low diversity and the common occur- rence of Suxijiragci oppositifoliu with rare, usually worn, remains of more warmth-demanding plants that are presumably dcrivcd from older deposits. S(i.vi- fruga oppositifoliu is found throughout Greenland. but becomes increasingly important towards the north, where it is perhaps the most ubiquitous and abundant of all vascular plants. No remains have been found in assemblages from the Langelandselv interglacial stage. Another hardy plant that occurs is Pupuorr sect. Scupijloru. The vegetation during the Hugin Sa inter- stade may have been entirely herbaceous, and we suggest that the mean summer temperature was L'.

3-4 C lower than at present. The only bcetlc re- covered is Aniuru ulpinu, which is probably the most hardy ground beetle on Earth. However, i t was found in only one sample, and it may be redeposited from 1,angclandselv interglacial sediments, where i t is common.

Biugc~ogrtiphy

A total of 38 taxa of vascular plants have been identified, of which 13 represent woody plants, and 4 arc limnic. No extra-Greenlandic plants have been idcntified, but several are southern extralimital (Fig. 8). and a few are presently confined to West Green- land (Ledum pahistre and Potarmgeton pusillus).

In contrast, of the 20 beetle species identified, only six live i n Greenland today. Two of these belong to the present fauna of ccntral East Greenland (Col~mhr~i~.s ~ioluhru tus and Byrrhus ,fii.sciritir.s), but four additional species arc found i n Greenland today (Table 2). In two cases the fossil localities on Jameson Land are well north (Otiorhynchuy arcticus) or far north (0. norio.vus) of the modern Greenlandic distribution.

One intriguing problem concerning the modern Greenlandic beetle fauna is the prevalence of palaearc- tic species. Of the total of 33 (probably) indigenous species, one is cosmopolitan, 16 are circumpolar or holarctic, 12 are palaearctic, one is amphi-atlantic, and merely one (Tvlicus .suhcanu.s, Byrrhidae) is ne- arctic ( Bocher 1988). Considering the proximity of Greenland to North America this situation is very strange and has aroused intense discussion ( Lundbeck 1891; Henriksen & Lundbeck 1917; Lindroth 1957. 1970, 1979; Danks 1981; <'oope 1979; Downes 1988; Bijcher 1988). One solution to the problem has been presented by Coope (1986b), who suggested an inva- sion from Europe to South Greenland by rafting over

a brackish North Atlantic Ocean on frozen flood refuse with a deflected Gulf Stream at the Younger Dryas- Holocene transition (Ruddiman & McIntyre 1981; Jansen ct al. 1983). One problem with this suggestion is that most lowland areas in South Greenland are thought to have been heavily glaciated at this time.

The fossil beetle fauna shows that also during the last integlacial stage, species with a modern palaearc- tic range dominated the Greenlandic fauna (Table 2 ) . Nine or possibly ten of the 18 identified species are today palaearctic and the remainder are circumpolar, whereas none is nearctic. This result suggests that invasions from northern Europe to Greenland might have been a common event during interglacials.

The fossil beetle fauna deviated strongly from the modern. For instance, two species of Coccinellidae are found both as fossils and living, but the species are different. Four species of Carabidae are known from Greenland today, but none of them is the interglacial carabid species. This striking dissimilarity may reflect the extremely unlikely possibility of successful disper- sal from (mainly) Europc.

Only four of the species found as fossils in Jameson Land are known from the Plio-Pleistocene Kap Kab- cnhavn beetle fauna from northernmost Greenland (Bocher 1989b). Thus the last interglacial fauna can- not be regarded simply as the last survivors from the Plio-Pleistocene Greenlandic beetle fauna.

In the discussion of the origin of the terrestrial beetle fauna of Grccnland, it has frcquently been argued that part of the fauna could have survived the Pleistocene glacial stages in rcfugia (e.g. Lindroth 1957, 1970). The dissimilarity of the Holocene, Eemian and Plio-Pleistocene beetle faunas does not support such a view. However, it is possible that some of the less warmth-demanding vascular plants and insects could have survived the glacial stages in unglaciated areas.

.4cknow~letl~emen/.s. - We are indebted to Anders Nilsson. IJniver- sity of Urncb, Tor idcntilication of Dytiscidac spp., and to ticnri Goulet, Biosystematics Research Centre, Ottawa. for identification of F:laplrrus ruhercularus. We express our gratitude to the PONAM participants for samples. company in the held and discussions. 0. B. Berthelsen, B. W. Rasmussen and J. Fuglsang are thanked for technical assistance. S. Funder. Copenhagen. kindly read an early draft of the paper.

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