ON THE QUALITY OF SVALBARD REINDEER PASTURE IN THE SUMMER AND AUTUMN

Late summer and autumn reindeer pasture plants from Adventdalen, Svalbard were analyzed for contents of fatty acids, energy content, protein, fibre, ether extract as well as content of macro minerals. F o o d intake of grazing reindeer in Adventdalen was estimated from fecal production. Large intake of high quality food seems to account for the growth and fattening of Svalbard reindeer during summer. K e y w o r d s : Re indeer , Svalbard , pasture qua l i ty , fatty acids, M i n e r a l s , f o o d intake. Rangifer4 (1): 16-23 S t a a l a n d , H . 1984. O m kval i te ten på reinbeite a v re inbei te på S v a l b a r d s o m m e r o g høs t . Sammendrag: Innholdet av fettsyrer, energi, protein, fiber, eterekstrakt og makromineraler ble analysert i reinbeiteplanter fra Adventdalen på Svalbard. Plantene ble samlet på ettersommeren. Forinn taket hos beitende rein i Adventdalen ble estimert ut fra fecesproduksjonen. Et stort inntak av for med h ø y kvalitet synes å kunne forklare vekst og fettlagring hos Svalbard-reinen om sommeren. Rangifer 4 (1): 16-23 S T A A L A N D , H . 1984. E la in t ie tee l l inen laitos, N o r j a n m a a t a l o u s k o r k e a k o u l u , 1432 A s N L H , N o r j a . H u i p p u v u o r t e n p o r o n l a i d u n t e n l a a d u s t a k e s a l l a j a s y k s y l l a . Yhteenveto: Rasvahappojen, energian, fiiberin, eetteriuutteen ja makromineraalien sisaltoa analysoitiin poronlaidunkasveissa Huippuvuorten Adventtilaaksosta. Kasvit kerattiin loppukesalla. Adventtilaaksossa laiduntavien porojen rehun kulunki arvioitiin lannan maarasta. Korkealaatuisen rehun suuri kulutus nayttaa vovan selittaa Huippuvuorten porojen kasvun ja rasvakerrostuman kesalla. v v v Rangifer 4 (1): 16 23 INTRODUCTION T h e s u m m e r grazing season for the Svalbard reindeer is short . T h e s n o w cover the g r o u n d to the end of M a y or early June , and permanent s n o w cover can be expected f r o m the end of September. I n the p e r i o d 1957-65 mean m o n t h l y temperatures i n J u n e , J u l y , A u g u s t and September were o n l y above z e r o ; 2.6, 6.0, 5.3 and 1.0°C (Steffensen 1969). T h u s both the p e r i o d of plant g r o w t h and the p e r i o d reindeer can graze fresh vegetation is short . Nevertheless the Svalbard reindeer t h r o u g h this short per iod manage to restore their depleted b o d y reserves of nutrients and to accumulate fat to an extent u n k n o w n i n most other ruminants (Reimers et a l . 1982). T h e qua l i ty of the range is apparently g o o d , g iv ing a h i g h p r o d u c t i o n rate o f volat i le fatty acids (Whi te and Staaland 1983), and the vegetation has h i g h levels of p r o t e i n and macro as w e l l as m i c r o minerals (Staaland et al . 1983). Studies o n grazing behaviour and plant selection have s h o w n that Svalbard reindeer extensively feed o n grasses w i t h h i g h n u t r i t i o n a l value l ike Dupontia sp. Alopecurus alpinus and Poa sp. t h r o u g h late summer and a u t u m n . A h i g h l y selected f o o d seems also to be the seeds of the viv iparous grasses of Poa, Descbampsia and Festuca (Punsvik et al . 1980, Persen et al . 1983). T h i s s tudy aims at examining the nutr i t iona l q u a l i t y of some selected forage plants grazed b y reindeer i n A d v e n t d a l e n , Svalbard, and further m o r e to give some estimates of f o o d intake i n the g r a z i n g p e r i o d J u l y to September. 16 RANGIFER, 4 (1), 1984 MATERIAL AND METHODS Some of the i m p o r t a n t forage plants u t i l i z e d b y the Sva lbard reindeer d u r i n g late summer and a u t u m n were col lected i n A u g u s t and September 1981 i n A d v e n t d a l e n o n western Spitsbergen. F u r t h e r samples for comparat ive analyses of fatty acid c o m p o s i t i o n were col lected i n J u l y 1982 at Slettrust near T y i n i n alpine areas of South N o r w a y . D e p o s i t e fat ( r u m p fat) f r o m Svalbard reindeer was col lected f r o m animals shot near Isf jord also o n western Spitsbergen i n the fal l of 1982. A l l samples were f r o z e n as soon as possible and kept f r o z e n at -20° t i l l prossessed. Samples of dehydrated fat and g r o u n d , dehydrated plant material were b o i l e d at 8 0 ° C i n a s o l u t i o n of 2 % concentrated sul fur ic acid i n m e t h a n o l . H e x a n e was added to the s o l u t i o n before in ject ion in to the gass c h r o m a t o g r a p h for determinat ion of f ract ional concentrat ions of l o n g chained fatty acids. These analyses were carried out at the A n a l y t i c a l l a b o r a t o r y of the D e p a r t m e n t of A n i m a l n u t r i t i o n , A g r i c u l t u r a l U n i v e r s i t y of N o r w a y . E n e r g y content (Cal/g D . M . ) was determined b y b o m b c a l o r i m e t r y , and d r y matter i n v i t ro digest ib i l i ty was determined according to the methods descr ibed b y T i l l e y and T e r r y (1963). I n o c u l u m f r o m r u m e n fistulated sheep fed hay and pelleted rat ions was used for the i n v i t ro studies. A l l other analyses were carried out by standard methods used at the C h e m i c a l research laboratory at the A g r i c u l t u r a l U n i v e r s i t y of N o r w a y (see Staaland et a l . 1983). Feca l p r o d u c t i o n was measured b y f o l l o w i n g the reindeer at close range, c o u n t i n g their fecal d r o p p i n g s and col lect ing and w e i g h i n g t h e m . A l s o f requency of u r i n a t i o n as w e l l as resting and g r a z i n g t ime was recorded. Size of animals i.e. l i k e l y range of b o d y weight , was estimated f r o m experience. D r y matter content of feces was determined after d r y i n g at 105°C for 12 hrs . R E S U L T S Fatty acids T h e d o m i n a t i n g fatty acids in plant material f r o m Svalbard is palmit ic ac id ; C : 1 6 . 0 (15-21%) and l ino len ic ac id ; C : 1 8 . 3 (13-36%). In Luzula confusa l ino le ic ac id ; C : 1 8 . 2 is f o u n d in highest concentrat i o n . In mosses the C : 1 8 acids are less d o m i n a t i n g and there is a large rest of n o n ident i f ied acids. There seems to be no major difference between Svalbard plant and m o u n t a i n plants f r o m N o r w a y i n content of fatty acids (F ig . 1). A s w o u l d be expected a large p r o p o r t i o n of the unsaturated acids are apparently saturated w h e n passing t h r o u g h the r u m e n ( C h u r c h 1976). T h u s l i n o l e i c and l i n o l e n i c acids are o n l y smal l fractions of the back fat acids. C o n t r a r y stearic, C : 1 8 . 0 , and ole ic acid C : 1 8 . 1 are f o u n d i n h i g h concentrat ions . A l s o the u n i d e n t i f i e d rest f rac t ion is smal l c o m p a r e d to i n plant material . T h e dominance of p a l m i t i c , stearic and oleic acids i n fat deposits is t y p i c a l for ruminants (see C h u r c h 1976) and has also p r e v i o u s l y been demonstrated i n Svalbard reindeer (R ingberg et al . 1980). Proximate food analyses H i g h p r o t e i n content was f o u n d i n Polygonum viviparum and i n Alopecurus alpinus. A l s o the v iv iparous grass seeds had h i g h levels of p r o t e i n (Table 1). Luzula confusa had very l o w p r o t e i n and sugar levels, and h i g h crude f ibre content . Seeds and the t w o grasses Dupontia pelligera and Alopecurus alpinus had the highest energy content . T h e v i v i p a r o u s grass seeds had also v e r y h i g h d r y matter d igest ib i l i ty , above 8 0 % . Luzula confusa h a d the lowest d igest ib i l i ty , about 4 0 % . Mineral content T h e minera l content of the plants agrees w i t h prev ious studies (Staaland et al . 1983). A s s h o w n before, Svalbard vegetation is part iculare ly h i g h i n N a and C I c o m p a r e d to some N o r w e g i a n plants . V i v i p a r o u s grass seeds are apparently relat ively l o w i n C a , whereas Equisetum arvense is a generally g o o d minera l source, part iculare ly h i g h i n S (Table 2). Fecal production and food consumption T h e frequency of defecation and u r i n a t i o n o n s u m m e r and fa l l pasture is h i g h (Table 3), g i v i n g a h i g h p r o d u c t i o n of feces pr . day . In large animals (100 kg) the rate of fecal p r o d u c t i o n is i n the order of 1.6 to 2.0 k g d r y matter p r . day. In smaller animals the fecal p r o d u c t i o n is l o w e r . If the d r y matter d iges t ib i l i ty is assumed to be about 6 0 % the d r y matter f o o d c o n s u m p t i o n i n adult large reindeer w o u l d be i n the order of 4-5 k g pr . day (Table 3) w h i c h again, w h e n tak ing i n t o cons iderat ion the water content of fresh vegetat i o n , w o u l d indicate an intake of 14 18 k g fresh vegetation pr . d a y . Average d r y matter content of plants analysed (Table 1) is 28 ± 8 % . RANGIFER, 4 (1), 1984 17 o 5 û . Ü J t/5 > 'g +| fi o> ^ .s? s


INTRODUCTION
The summer grazing season for the Svalbard reindeer is short.The snow cover the ground to the end of May or early June, and permanent snow cover can be expected from the end of September.In the period 1957-65 mean monthly temperatures in June, July, August and September were only above zero; 2.6, 6.0, 5.3 and 1.0°C (Steffensen 1969).Thus both the period of plant growth and the period reindeer can graze fresh vegetation is short.Nevertheless the Svalbard reindeer through this short period manage to restore their depleted body reserves of nutrients and to accumulate fat to an extent unknown in most other ruminants (Reimers et al. 1982).
The quality of the range is apparently good, giving a high production rate of volatile fatty acids (White and Staaland 1983), and the vegetation has high levels of protein and macro as well as micro minerals (Staaland et al. 1983).Studies on grazing behaviour and plant selection have shown that Svalbard reindeer extensively feed on grasses with high nutritional value like Dupontia sp.Alopecurus alpinus and Poa sp.through late summer and autumn.A highly selected food seems also to be the seeds of the viviparous grasses of Poa, Descbampsia and Festuca (Punsvik et al. 1980, Persen et al. 1983).
This study aims at examining the nutritional quality of some selected forage plants grazed by reindeer in Adventdalen, Svalbard, and further more to give some estimates of food intake in the grazing period July to September.

Fatty acids
The dominating fatty acids in plant material from Svalbard is palmitic acid; C:16.0 (15-21%) and linolenic acid; C:18.3 (13-36%).In Luzula confusa linoleic acid; C:18.2 is found in highest concentration.In mosses the C:18 acids are less dominating and there is a large rest of non identified acids.
There seems to be no major difference between Svalbard plant and mountain plants from Norway in content of fatty acids (Fig. 1).
As would be expected a large proportion of the unsaturated acids are apparently saturated when passing through the rumen (Church 1976).Thus linoleic and linolenic acids are only small fractions of the back fat acids.Contrary stearic, C:18.0, and oleic acid C:18.1 are found in high concentrations.
Also the unidentified rest fraction is small compared to in plant material.The dominance of palmitic, stearic and oleic acids in fat deposits is typical for ruminants (see Church 1976) and has also previously been demonstrated in Svalbard reindeer (Ringberg et al. 1980).

Proximate food analyses
High protein content was found in Polygonum viviparum and in Alopecurus alpinus.Also the viviparous grass seeds had high levels of protein (Table 1).Luzula confusa had very low protein and sugar levels, and high crude fibre content.Seeds and the two grasses Dupontia pelligera and Alopecurus alpinus had the highest energy content.
The viviparous grass seeds had also very high dry matter digestibility, above 80%.Luzula confusa had the lowest digestibility, about 40%.

Mineral content
The mineral content of the plants agrees with previous studies (Staaland et al. 1983).As shown before, Svalbard vegetation is particularely high in Na and CI compared to some Norwegian plants.
Viviparous grass seeds are apparently relatively low in Ca, whereas Equisetum arvense is a generally good mineral source, particularely high in S (Table 2).

Fecal production and food consumption
The frequency of defecation and urination on summer and fall pasture is high (Table 3), giving a high production of feces pr.day.In large animals (100 kg) the rate of fecal production is in the order of 1.6 to 2.0 kg dry matter pr.day.In smaller animals the fecal production is lower.If the dry matter digestibility is assumed to be about 60% the dry matter food consumption in adult large reindeer would be in the order of 4-5 kg pr.day (Table 3) which again, when taking into consideration the water content of fresh vegetation, would indicate an intake of 14 -18 kg fresh vegetation pr.day.Average dry matter content of plants analysed (Table 1) is 28 ± 8%.
RANGIFER, 4 (1), 1984  In between this grasses are also found Equisetum arvense which is also used by the reindeer (Punsvik et al. 1980, Persen et al. 1983).Further observations have also revealed that the seeds of the viviparous grasses, Deschampsia alpina, Poa spp.
and Festuca vivipara are selected by the reindeer.
In late autumn grasses with viviparous seeds can often only be found in localities not reached by the  3).The main error encumbered is that the body weights of the animals have to be judged by experience from the weight of shot animals.From the amounts of feces produced food intake can be estimated provided the digestibility is known.From studies on in vitro digestibility (Table 1; Staaland et al. 1983) and from feeding trials (Staaland and 0ritsland in prep.)60% dry matter digestibility might be a fairly good estimate on summer pasture.The dry matter intake of adult bucks would then be in the order of 4-4.9 kg or, at a medium body weight of 100 kg, 126-155 g/BW 0 : 75 per day.For the 50 -60 kg animals (medium body weight 55 kg) 148 g/BW 0 : 75 and for the yearlings (35 kg body weight) 104 g/BW 0 : 75 (Table 3).These values are lower than previous estimates made by measuring the ruminal VFA production in summer grazing Svalbard reindeer, 187 g/BW 0 : 75 (White and Staaland 1983).
However, a small increase in dry matter digestibility to e.g.65% would increase the 68 and 33 g/kg per day (White and Staaland 1983).
If these values are accepted it can also be calculated that the intake of energy and nutrients are high in summer grazing reindeer on Svalbard; due to high quality forage and large intake.The mean energy content of all plant samples from Svalbard (Table 1) is 4320 cal/g dry matter.(Klein 1970;Renning 1976Renning & 1979;;Punsvik et al. 1980;Staaland et al. 1983).
The Svalbard reindeer apparently lives a more sedentary life than continental reindeer/caribou.This is obviously an effect of several factors.The lack of predators and insect harassment, as well as a cool summer climate make movements to snow patches and high altitude less important.Since the Svalbard reindeer graze singularely or in small groups and are spaced out over the available grazing terrain, each area can support the same group og animals through prolonged periods of time or throughout the whole year.This may reduce intraspecific competition, reduce the need of migration and ultimately save energy and increase available grazing time.
Thus although the primary production may seem low (Brzoska 1976;Brattbakk and Ranning 1978) the high quality of the forage together with the sedentary behaviour of the Svalbard reindeer may explain these animals capability to grow new tissue and store fat during summer and to maintain high density populations (Reimers et al. 1980).
of the important forage plants utilized by the Svalbard reindeer during late summer and autumn were collected in August and September 1981 in Adventdalen on western Spitsbergen.Further samples for comparative analyses of fatty acid composition were collected in July 1982 at Slettrust near Tyin in alpine areas of South Norway.Deposite fat (rump fat) from Svalbard reindeer was collected from animals shot near Isfjord also on western Spitsbergen in the fall of 1982.All samples were frozen as soon as possible and kept frozen at -20° till prossessed.Samples of dehydrated fat and ground, dehydrated plant material were boiled at 80°C in a solution of 2% concentrated sulfuric acid in methanol.Hexane was added to the solution before injection into the gass chromatograph for determination of fractional concentrations of long chained fatty acids.These analyses were carried out at the Analytical laboratory of the Department of Animal nutrition, Agricultural University of Norway.Energy content (Cal/g D.M.) was determined by bomb calorimetry, and dry matter in vitro digestibility was determined according to the methods described by Tilley and Terry (1963).Inoculum from rumen fistulated sheep fed hay and pelleted rations was used for the in vitro studies.All other analyses were carried out by standard methods used at the Chemical research laboratory at the Agricultural University of Norway (see Staaland et al. 1983).Fecal production was measured by following the reindeer at close range, counting their fecal droppings and collecting and weighing them.Also frequency of urination as well as resting and grazing time was recorded.Size of animals i.e. likely range of body weight, was estimated from experience.Dry matter content of feces was determined after drying at 105°C for 12 hrs.
reindeer.The importance of Graminae in the reindeer diet in Adventdalen have been confirmed by botanical analyses of rumen content(Staaland   et al. 1983).Both in summer and winter mosses are ingested in large quantities.In a recent paper,Prins (1982)  argue that mosses are eaten by animals in cold environments because of high content of the poly-unsaturated arachido-nic acid.Some of the positive effects in cold climate would be lowering the melting points of fat in the extremities and protection of cell, membranes against cold-However, Svalbard reindeer, one of Prins' examples, even though consuming, large amount of mosses (Staaland et al. 1983) may have little benefit in this respect from eating mosses, since unsaturated fatty acids usually are hydrogenated when passing through the rumen.These acids will therefore enter the small intestine in a saturated form.This seems to be confirmed in the present study since the rest fraction, several long chained unsaturated fatty acids as well as C:18.3 (linolenic) and C:18.2 (linoleic) are almost absent from back fat deposits.This is in contrast to non ruminants like e.g.Willow and Rock ptarmigans (Lagopus lagopus and L. mutus), where both linolenic, linolic and arachidonic acids constitutes large fractions of body fats (Tanhuanpaa and Pulliainen 1969).Linoleic acid is the only truly essential fatty acid in the diet (McDonald et al.1978).There seems to be no major difference in fractional content of this acid between Svalbard plants and mountain plants from Norway (Fig.1).Scarcity of this component in the diet of Svalbard reindeer seems unlikely.In particular the viviparous seeds from grasses are with very high digestibility and high energy and protein content.Some grasses, in particular Alopecurus alpinus seem to be high quality forage.Also plants like Oxyria digyna and Equisetum arvense can be important forage because of their high mineral and protein content(Staaland et al. 1983; tables 2 and 3).In contrary the abundant Northern Woodrush, Luzula confusa is very low in protein, sugar and digestibility, but high in fibre.E. g. low sugar could be a reason for not selecting this species(Arnold and Dudzinski 1978).The food intake of wild grazing reindeer is difficult to estimate.Because of the apparent tameness of the Svalbard reindeer, they can relatively easily be followed for hours at close distance and urination and defecation observed.During such observation periods they continue grazing, lie down ruminating and apparently behave normally.They may even sometimes come close (5-10 m) to have a good look at the biologist.After defecation the fecal droppings are usually easy to find, due to the open landscape and short vegetation.By this method it is therefore relatively easy to obtain a good estimate of fecal production in Svalbard reindeer during summer (Table calculated dry matter intake to 178 g/BW°: 75 per day.The obtained values are in good agreement with data obtained from pen-fed Svalbard reindeer, about 3.0 kg/dry matter, i.e. approximately 112 g/BW 0 : 75 per day (Nilssen and Ringberg 1980).The food intake for Svalbard reindeer thus appears considerably higher than estimates made for 20 RANGIFER, 4 (1), 1934 Norwegian reindeer at Lødingen in Northern Norway and for caribou at Prudhoe Bay in Alaska;

Mineralkonsentrasjoner (mMIkg tørrstoff) i beiteplanter samlet i ! Adventdalen i midten av september 1981.
The wet meadows of Svalbard, which are prime summer habitats are more dominated by Graminae than by sedges and rush than in similar wet areas (bogs) in Norwegian reindeer habitats.Also the apparent large intake of forbs and viviparous seeds in Svalbard reindeer appears important.Available plants are furthermore high in nutrient.The a food of high digestibility and high nutrient content.There are several factors both in the habitat and in the behaviour of the Svalbard reindeer which may enhance growth and fattening processes.