African Wild Dog (Lycaon pictus) ... Invastive Research, Reduced Longevity and Mortality ... Roger Burrows

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APPENDICES

INVASIVE RESEARCH, REDUCED LONGEVITY AND MORTALITY IN AN ENDANGERED SPECIES,
THE AFRICAN WILD DOG (Lycaon pictus),

with special reference to the extinction of the
Serengeti-Mara study population 1985-91.
Roger Burrows (December 2003)

Appendix 1. Cause of death in some handled free living and captive packs

			type of handling
Location	date	type of pack	r.c.	vaccinated	events following & n. packs or individs.	cause

Zimbabwe +	1986	captive	?	no data	pack died	rabies
Serengeti	1985-90	free living	yes	no	5 packs died	*rabies
Mara	1987-90	free living	yes	yes	3 packs died	rabies
Serengeti	1991	free living	yes	yes	5 packs died	**rabies
Namibia (Etosha )	?	captive	?	yes	6 of 7 pups died ('Vanguard' live vaccine)	CD
	1989	introduced	?	?	all 5 dogs died	lions
			fitted with contraceptive implants
	1990	captive	?	yes	all 11 dogs died	lions(6), rabies(4), 1?
Botswana
i. Moremi	1996	free living	***yes	?no	5 packs died in 3 weeks	?CD/ rabies
ii.Chobe	1994	free living	no data provided		10 of 12 dogs died in 2 weeks	CD
South Africa
i. Madikwe	1997	introduced	yes	yes	7 of 10 dogs died or 'put down'	rabies
	2000	introduced	adult yes	ad. yes pups no	8 of 11 pups died	rabies
ii. Kruger	1995- 2000	free living	yes	no	pop. decline of 60% & 30%of packs lost	unknown
			but adult pair in each pack immob./ bloodsampled n=72
Tanzania (Mkomazi)	2000-1	captive	no	yes CD$ rabies etc	49 of 52 dogs died in 4 packs in 2 months	CD
Zimbabwe (Hwange )	2003-	free living	yes	no	1 female died	rabies

+Kat et al (1995) claims that this pack was unhandled.

* Confirmed in a tissue sample from the only pack from which an adequate sample was taken

**. No tissue samples taken but 2 Mara packs died in December 1990 one with rabies confirmed

*** All radios expired within 19 months, dogs re-immobilized to replace. No tissue samples from 5 dead packs.

CD= Canine Distemper:- despite serological evidence of high levels of exposure of free living packs to the virus in Botswana, Selous, South Africa and Namibia, the only reported case of CD in free living wild dogs was in Botswana in 1994 when 10 individuals in a pack died; whether the pack was handled is unclear.

$ Inactivated CD vaccine used (van de Bildt et al 2002) cf. Etosha.

c. =radio-collared

Appendix 2. Non invasive Research

1. Photographic surveys and information from rangers provides reliable indications of size and changes in populations of wild dogs without the use of telemetry.

i. In Serengeti :- In the period 1965 to 1970, 163 different dogs (of all ages) were photographed and catalogued on the Serengeti plains (van Lawick- Goodall 1970) at a time when the population was claimed to be small but stable. This is comparable with the 162 different individuals in 12 different packs photographed in another 6 year period (1985-91 see Table 1) at a time when the wild dog population of the plains was claimed to be very low compared with that in the late 1960s and 1970.

Of the 12 packs photographed between 1986-91 a maximum of 5 packs were seen in any one year; that was in early 1991 (Burrows 1993). Based on published data (Appendix 5) the same number of packs were present on the Serengeti plains in 1970 as in early 1991.

From photographic evidence it can concluded there is no evidence for a significant change in the number of wild dog packs resident on the Serengeti plains from 1964-91 (Burrows et al 1994).

ii. In Kruger. As a result of a wild dog photographic competition tourists provided information which enabled a 60% decline in the population between 1995-2000 to be detected when telemetry had apparently failed to do so (M.G.L. Mills. http://www.parks-sa.co.za/conservation/scientific).

iii. In Selous estimates of the size of the population provided by rangers and game scouts before handling began in this woodland habitat in 1991 proved to be reliable and were later confirmed by radio- telemetry (Creel & Creel 1994).

2. Data collection using faecal analysis

DNA studies of free living wild dogs and other species using faecal samples from known individuals of known social status can produce valuable information on relationships of individuals (K. Leigh per comm.) It is also possible to measure changes in hormone levels (Creel et al 1997, East et al 1997, Hofer & East 1998, Goymann et al 2001).

Appendix 3 a-d. Number of packs and individuals claimed on the Serengeti plains in the late 1960s and the claimed series of population crashes pre 1985 is based on a misinterpretation of data:-

Appendix 3a Estimate of the population of wild dogs for the 'ecological unit' based on observation on the plains pre 1970.

George Schaller studied lions on the plains of Serengeti from 1966-69 in the wet seasons between December to May each year a period co-incident with the denning period of the wild dogs when the packs are most easily observed; but in the dry season he worked primarily in woodland (Schaller 1972). Serengeti wild dog packs travel widely over their extensive home ranges during the dry season and are difficult to locate and follow as they often travel long distances at night (Burrows 1995). Schaller thought that the wild dogs left the plains and retreated with their migrant prey into the woodlands during the dry season. This was not the case plains packs avoid woodland whenever possible and no packs resident (i.e. denning) in woodlands in the Serengeti around the plains were located either pre 1970 or subsequently (Frame & Frame 1981). Both observations are surprising, if as is claimed, woodland is the preferred habitat of Lycaon (McNutt & Boggs 1996).

The invaluable data which Schaller produced pre 1970 including his estimated population densities therefore relate to wild dog packs and groups on the open plains in the wet season when the packs are confined to the area around the den.

Schaller guessed that during his study period (June 1966-September 1969), some 200-250 dogs, (including older pups not at dens which made up 1/3 of the population) existed in the 'ecological unit' which at that time was 25,500 km 2. This guess was an extrapolation based on a population density derived from his knowledge of the number of packs and individuals resident on the plains in the wet season.

Kruuk & Turner (1967), also basing their knowledge of the plains wild dogs, estimated that possibly 150 wild dogs (age unspecified ) occurred in the National Park, which at that time was about 12,500 km2. (a PD of 1: 84 km2).

In his Table 65 Schaller (1972) reported that a total of 33 'packs' of differing composition were seen during the period 1966-69 on the Serengeti plains but:-" As packs changed their size and composition it was difficult to recognise some after a lapse of a few months" and as a result:- "Some individuals may have been counted several times…. (Schaller 1972 p 324). Schaller included single sex groups in his tally of 'packs'.

As Schaller reported, a healthy wild dog pack changes its composition frequently. This is due to successful reproduction, frequent emigration (extent of this unknown to Schaller) plus some immigration and mortality. There were not 33 different packs present on the plains between 1966-69. In his Table 56 Schaller provides 15 examples of different pack compositions, including that of 13 'groups' with a basic 'pack' structure (i.e a potential breeding unit of at least one adult pair) and 2 single sex groups. In the sample he selected there were 153 individuals : 100 adults and 53 older pups (6-12 mo.) as this was a period of high reproductive success.

Schaller did not identify each dog in each pack located and it later became apparent that data from at least 1 pack (the Genghis ) was included for both 1966 and 1967 (Schaller's Table 65). How many packs and thus number of individuals in them, other than in the 'Genghis Pack', were multiple counted in this way is unknown. Unless each individual is know and checked with photographs showing both sides of each individual multiple counting of wild dogs in a pack is a common field problem and individuals and groups also emigrate between packs.

However, in both 1964 and 1967 there were 5 known 'packs' (one in 1967 consisted of young females only - i.e. not a 'true' pack) on the Serengeti plains. Wild dogs were described as scarce and rare judging by the infrequency with which they were seen but the population up to September 1969 was described as stable with no oscillations and had changed little since the 1950s (Kuhme 1965, Schaller 1972). Between 1965-69 a maximum of 5 packs were seen in any one year on the Serengeti Plains and based on the estimated number of adults and older pups (comprising 1/3 of the population) the population density (PD) was 10-12 per 1000 km2 of which 7-8 were adults (data based on Schaller 1972 ). This compares well with Kruuk & Turner's estimated PD of approximately 12 per 1000 km2 age of individuals unspecified.

Appendix 3 b. Number of wild dogs and packs on the Serengeti plains in 1970

For 1970 in a similar Serengeti plains area to that observed by Schaller it was reported that there were 12 packs and 95 adults in a 3000 km2 study area, (Malcolm 1979, Frame et al 1979), a PD 1:35km2 or 28-29 adults per 1000 km2 and so an apparent population explosion between 1969-70 with a near tripling of the adult population, and more than double the number of packs present on the plains in the mid -late 1960's. Clearly these highly anomalous data needs to be explained.

In 1970 a then young and inexperienced field worker, based near Ndutu on Lake Lagarga, just south of and outside Serengeti National Park,followed one particular pack (the Genghis) which was the subject of a film. Working within a very limited area around Ndutu it was claimed that in 1970 there were over 100 adults and 12 packs on the Serengeti Plains (Malcolm & van Lawick 1975, Malcolm 1979).

Malcolm (1979) agreed with Schaller's estimate for the total number of dogs in the 'ecological unit' and claimed a similar number existed in 1970. However Schaller included older pups in his guess, Malcolm included only adults.

Both Schaller & Malcolm based their population estimates on the PD on the Serengeti plains. If Malcolm's PD for the plains in 1970 really was 1:35 km2 then the number of adults in Schaller's 'ecological unit' must have dramatically increased from 200-250 (134 - 167 adults) up to 1969 to over 700 adults in 1970. Clearly this is highly improbable.

The first mention of 12 packs on the Serengeti plains in any one year in the literature was in 1975:-

'There are no less than 12 packs of wild dogs which may occasionally include the short grass plains around Lake Lagarja, in the Serengeti as part of their nomadic circuit" (van Lawick 1973 p 10). van Lawick does not claim these were all seen in any one year. [These 'packs' included single sex dispersing groups].

The number of packs and adults in 1970 in the 3000km2 study area was later 'revised' by Malcolm to 77 adults in 10 packs plus one all male group. However even these revised data include at least 30 unknown individuals who were only seem once or twice, at the periphery of the study area and were not catalogued and not resident year round in the area, and included unpublished data for 2 other packs which contain another 10 adults (J Malcolm per. comm. & see this Appendix 6).

Of the 10 packs reported to have been seen in 1970, just 4 were known to be resident ( i.e. denning) in the 3000 kms2 study area. Of the 3 packs which bred in 1970 one was a lone pair which split away temporarily from another pack (the Genghis). No pups born in 1970 are known to have survived.

For 1970 published data (Frame et al 1979) show a maximum of 5 packs on the plains included a lone pair from a larger pack a resident all male group and a newly located pack whose denning area was unknown i.e. a maximum of 5 potential (if immigrant females had joined the all male group) denning packs in 1970 (see Appendix 6).

Unfortunately Schaller's data on the number of individuals on the plains and van Lawick's anecdotal statement relating to 12 packs has been misinterpreted and this is most probably the basis for the claim that 12 packs and 95 adults were living year round on the Serengeti plains in 1970 in a 3000 km2 study area. (Malcolm 1979, Frame et al 1979).

Neither Schaller nor van Lawick reported the presence of 12 packs in any one year. Between 1967-1972, twelve packs were identified (Malcolm 1979) and named with sometimes different names being given to the same pack by different observers (see Appendix 6) In only some of these packs were all the individuals catalogued. There were a maximum of 12 packs reported over a 5 year period not all in 1970 (Malcolm 1979).

Later analysis of the published data and other research suggests that the most likely number present in the Serengeti plains 3000km2 study area in 1970 was around 30 resident adults in 4 -5 packs and so comparable with Schaller's estimate for the number of adults and breeding packs pre 1970 (Schaller 1972, Burrows 1995 and see Appendix 6).

The Serengeti plains are considered to be 'not good habitat' for Lycaon (Frame & Frame 1981) and it is suggested that even in more favourable habitats (claimed to be woodland) a reserve of 2,300 km2 could only support just 6 packs and a maximum of 30 adults. The validity of the published 1970 data is clearly highly questionable.

There is no evidence that 12 packs and 95 adults were resident on the Serengeti Plains in 1970 or a population density of adults of 1: 35km2

Appendix 3 c. A claimed population crash in 1971-73

It is claimed that in 1971-73 the wild dog population was not handled but fell from 96-49 with 5 of 12 packs disappearing, with disease implicated (Creel 1992, Creel et al 1997 ).

The reference given is Malcolm (1979). No wild dog research involving general monitoring of packs and mortality data collection was carried out in Serengeti from 1971-74. Opportunistic records were collected mainly from a small area around Ndutu with months of little activity. Malcolm was present for just 3 months in 1971 and 2 months in 1972 (Malcolm 1979).

For the periods 1969-71 and 1974-79 annual adult survival was 0.825 any, mortality was inferred from the disappearance of known single adult dogs but 2 adults died in the Kuhme pack from unknown causes (Frame & Frame 1981, Burrows 1995). In 1971 the lodge manager at Ndutu reported in 1971 that a pack was 'looking sick', they often do when they are not as after wallowing in a muddy pool (Frame & Frame 1981).

The assumption made that disease related high adult mortality played an important role in regulating the wild dog population in Serengeti in the late 1960s and 70s is ill founded. As the researcher involved in the 1970s later agreed "the data are thin" (J.Malcolm pers. comm.).

There is no evidence either for the loss of 5 packs between 1971-73, (see Appendix 6) or significant adult disease related mortality, but at least 3 packs which were not identified were shot in Serengeti, one in 1970 at the boundary of the National Park with a Game reserve, a second in 1972 when 25-30 were shot in the middle of the Park and in 1973 an entire pack was shot near Seronera (Malcolm 1979, Frame 1976, Burrows 1995).

Creel's claimed loss of 5 packs from disease and so a population crash in 1971-73 is based on the highly inflated estimate of the number of wild dog adults and packs claimed to be resident on the Serengeti plains in 1970 which has been uncritically accepted There is no evidence for a crash in 1971-73 or population 'bottlenecks' or that the Serengeti wild dog could be described as a 'roller coaster' population (contra Creel 1992).

Appendix 3 d. A claimed Serengeti population crash in 1975-76

Ginsberg et al's 1995 PVA model is based on what are claimed to be a series of significant crashes ( >40%) in the 'Serengeti' population, the first in 1975-76 before intensive invasive handling began. This putative population crash from 60-30 adults, is crucial to the model but a closer examination of the published data ( Frame et al 1979) of the yearly maximum number of adults in each pack seen in the study area does not support such a claim. Fortunately the researchers involved also provided data on the study population per quarter year (see Table 2). It is clear that there were not 60 individual study adults recorded at any one time in the Serengeti study area in 1975.

Again, as for the claimed crash in 1971-74, disease is claimed to be the cause of a putative crash in 1975-76. Again there is no data support as no ailing dogs were seen in the Serengeti between 1974-77 and just 13 of 79 known adults died giving an annual survival 0.825 (Malcolm 1979, Burrows 1995). Deaths were inferred from the disappearances of single adult dogs (mainly age related see data in Malcolm 1979) excluding probable emigrants which also disappeared (Frame et al 1979, J. Malcolm pers. comm., Burrows 1995).

The number of individuals seen in 1975 is the maximum number of adults in each pack during the year and included a rare sighting of one large peripheral pack which did not den in the study area. When seen in late 1974 this pack (the Spitfire) contained 4 adults and 11 pups present. The pups were not included in the population count but by the second quarter of 1975, as the pups were >12 months old and so adults, they were now included in the count by the researchers. Therefore the pack appeared to rapidly expand from 4 to 15 adults during the course of one quarter year (Appendix 6).

The apparent loss of individuals and 2 packs in 1976 was mainly due to the failure of the researchers to see two other peripheral packs which denned outside their study area. One was not seen again in the study area after the first quarter of 1975 when it had 3 members. The second a previously unknown pack containing 18 adults when first seen in August 1973. Its home range was not defined and its denning area unknown. When last seen this pack consisted of 12 adults (4 adults and 7 yearling males and a lone adult female). Five yearling females from this pack had emigrated to join an all male study group in January 1974. The pack was last seen in the study area in the first quarter of 1975. This gave the appearance from the raw yearly data that 2 packs and 15 individuals had died/disappeared. There is no evidence that they did. There were not 60 adults at any one time in the Serengeti study area in 1975: there was a maximum of 46 in 1975 (Table 2).

Table 2. Total verified population of adult wild dogs observed in the Serengeti plains study area 1974-76 (data from Frame 1977 Table 3 & Frame et al. 1979 Table 1). Data given at 3 month intervals; amg =all male group.

1974

1975

1976

Frame 1977

[44

41]

[39

37]

[32

30]

n Packs

cf Frame et al. 1979

but give data for

4 + 1 amg

4 packs

5 packs

* Data provided for only 4 packs; 3 of which were peripheral packs rarely seen, and did not den in the study area.

As the researcher involved observed with reference to the Serengeti decline 1975-76.

" there are several reasons to suspect that the decline in the population is more apparent than real" (Frame 1977).

The pack is the basic reproductive unit in wild dogs and the size of the breeding population is directly related to the number of packs in the study population (Woodroffe & Ginsberg 1999).

The claimed 1975/76 wild dog population crash is an artefact of data presentation. Ginsberg et al's PVA model based on a series of crashes pre and post the use of routine handling (i.e. 1985) is fatally flawed.

Appendix 4. Based on misrepresentation of field data and anecdotal statements a number of erroneous or misleading statements commonly appear e.g. :-

1. In the late 1960s .."some 153 wild dogs ranged the Serengeti Plains". The reference given is Schaller 1972 i.e. Table 65 (Estes 1992 p 413). Estes implies that there were 153 individual adults present as residents on the plains in any one year. Schaller's Table lists 153 individuals seen between 1966-69 some of which were counted more than once (see Appendix 3a). Schaller's total also contained 53 pups, whereas the demographic data provided by researchers post 1969 related only to adults with older pups excluded ( Frame et al 1979, Burrows et al 1994).

However based on this error of data interpretation Estes suggests that when the mean number of adults in the study packs fell to 3.6 in the late 1970s the decline in the sub-population may have been irreversible.

2. It is claimed that the number of adults living on the plains year-round (i.e. resident) in 1970 was around 110 (Malcolm & van Lawick 1975 and pers. comm. to Hanby & Bygott 1979 in Sinclair &Norton-Griffiths 1979).

3. "Wild dog have declined on the plains from over one hundred in the 1960s to only thirty in 1977" (Sinclair & Norton-Griffiths p.13, 1979).

4. Relating to the increase in migrant and resident herbivore species on the plains in the 1970s:- "The latter was accompanied by an increase in lion and hyaena populations, which in turn, decreased the smaller population of wild dogs" ( Sinclair 1979).

5. "The increase in predator numbers has already been proposed as a likely cause [ of the population decline of wild dogs in the 1970s] in Serengeti (Sinclair 1979a),and has not yet been discounted " (Sinclair 1995).

6. The plains were perhaps no longer a suitable habitat for wild dog (Sinclair & Norton Griffiths 1979, Frame & Frame 1981).

7. The population density for Serengeti wild dogs between 1967-79 is claimed to be 0.015 adults/km2 (Creel & Creel 1996). However not only is Schaller's estimate taken to be based on numbers of adults pre 1970 but it also appear that the highest figure given by Schaller was taken as the basis for their calculation of PD up to 1970 i.e. 1 dog per 85 km2. Creel & Creel then take Malcolm's PD (in Frame et al 1979) for 1970 (1:35 adults/ km2) and Frame et al's PD data for 1977 (1:200 adults/km2 ) and average the results (see below).

Population density estimates 1967-177

Category of dog	years(s)	1/km2	n/km2

adults plus older pups	1966-69	1:85-102	0.012-0.009	Schaller (1972)
adult only	1970	1:35	0.028	Malcolm (1979)
adult only	1977	1:200	0.005	Frame et al (1979)

	Average 1967-78		0.015	Creel & Creel (1996)

Frame et al's data was based on a study area of 5,200 km 2 in which in 1977 they counted 26 adults.

This was NOT the entire population of Serengeti, the National Park or the ecosystem as is implied by some commentators (e.g Estes 1991). In 1976 it was claimed that there were at least 7 more packs in the areas around their study area (Frame &Frame 1976).

Errors made in the estimation of population density and causes of mortality in Serengeti have not only led to totally unfounded claims e.g. 1- 7 above, but are the basis of a plethora of scientific papers particularly post 1991 relating to the fate of Serengeti wild dogs, most based on false assumptions and uncritical acceptance and use of raw data.

Appendix 5. Survival less than 12 months following handling.

Data from Ginsberg et. al. (1995a) most mortality claimed to be due to 'natural causes'

	% Radio -collared	% radio implanted	% blood sampled only

Botswana	25		33
Hwange	31		ND
Kruger	40	53	21
Mara (Kenya)	33		70
Selous (Tz)	20		ND

Appendix 6. Resident (i.e. denning), Peripheral (non denning) and unknown packs/groups of wild dogs on the South East Serengeti Plains 1969-77 (no data for 1972&73) pd = based on published data from Frame et al (1979) & Frame (1977). Data for 1971 James Malcolm (JM) pers comm.

YEAR	1970a pd	1970b new data JM.	1971J.M..& .pd	1972-73ND	1974pd	1975pd	1976pd	1977pd

a. RESIDENT PACKS
Genghis	16(7)	16(7)	9(2)		7(0)	10(4)	5(0)	4(0)
Genghis a.	2(0)	2(0)
Genghis b.			2(0)
$Herod=Flavian	7 (5)	7(4)	2(amg)
Kuhme	ND	[5]	ND		5(0)	4(0)	5(0)	2 (0)
$Pimpernel=Nettle	4 (amg)	4(3)	6(0)
$Cassidy=Seronera	ND	[5]	4(0)		7(0)	8(5)	6(0)	3(0)
Plains					3(0)amg	7(0)	5(0)	5(0)
Falcon								4

Packs (including resident amgs)	4 3+amg	6	5 4+amg		4 3+amg	4	4	5
Resident adults %(ylgs)	29 54	39 36	23 9		22 0	29 31	21 0	18 0

b.PERIPHERAL PACKS

Plane a	8(10)	8(10)	9		ND
Plane b			5		ND
Semetu					9(6)	3(0)	ND	ND
Spitfire					4(0)	15(11)	8(4)	4(0)
Simba					12(0)	**12(0)	ND	ND
peripheral packs	1	1	0		3	3	1	1
peripheral ad	18	18			25 (6)	30(11)	8(4)	4(0)

c.UNKNOWN PACKS &amgs number of adults	0	a[4] b[9] c[7] [10] amg	0		0	0	0	d 3
unknown adults	ND	30	0		0	0	0	3
unknown packs	0	3	0		0	0	0	1
Total adults a+b+c % yearlings	47 47	77 42	37 6		47 13	59 40	29 8	25 0
Packs & amgs	5	7	7		7	7	5	6
. non resident amgs	0	1	0		0	0	0	0

TOTAL INDIVIDUALS a+b+c in year	47	77	37		47	59	29	25
TOTAL PACKSa+b+c in year	5 4+amg	10 +amg	7 6+amg		7 6+amg	7	5	7
Compare above with:-
individuals Frame et al	95	95	ND	ND	49	62	29	26
packs Frame et al.	12	12	ND	ND	7	8	5	7

Figures in brackets:- ( ) yearlings; [ ] new unpublished data; ND =No Data; amgs = all male groups resident in a home range after the female(s) have left/died; $ Some packs were given different names by different observers.

Peripheral= packs which did not den in the study areas , JM = James Malcolm;

* Plane 1969 with 10 pups. Adult sex ratio unknown, pack not catalogued; 10 pups approx. 6 mo in early 1970

** No data for Simba pack in 1975 in Frame et al but possibly seen in first quarter of year (L.Frame 1977).

*** Frame et al 1979 (Table 1) claim 8 packs in 1975 but this is an error due to counting an all female group twice.

Unknown = packs/groups reported once or twice in study area in 1970 by J.Malcolm (pers comm.) but for which there are no published data (cf revised data of 87 in 10 packs and 1 amg. J.Malcolm pers comm. in Burrows (1995).

Identity of 3 packs shot (1970-73) not known

Appendix 7. Decline of rinderpest a possible factor in the 1970s decline in Serengeti Lycaon mean pack size

A dramatic decline in the reproductive success of Serengeti wild dog (but not in the number of packs) in the late 1960s and the 1970s was coincident with reports of the death of wild dog pups with clinical symptoms suggesting Canine Distemper (CD) (Schaller 1972). In the 1960s Spotted hyaena were found to be seropositive for Rinderpest (RV) although no associated mortality was reported (Sachs & Staak 1966). The suspected CD in wild dog affecting mainly wild dog pups and young adults. There is no evidence of high adult mortality or that between 1971-73, 5 of 12 packs disappeared with disease implicated (contra Creel 1992). Three packs were shot (Burrows 1995). The successful vaccination of domestic stock against rinderpest in the 1950s led to the virtual elimination of the virulent strain of rinderpest virus from wild life by 1963. The viruses causing rinderpest and canine distemper are very closely related and exposure to rinderpest virus is known to provide cross resistance to CD in domestic dogs, although with no serum antibodies reported (DeLay et al 1965).

Up to 1963 when rinderpest was common in Serengeti in young wildebeest and buffalo, wild dogs, like the Maasai's domestic dogs ate RV infected wildlife. It is conceivable that this naturally immunised both species against subsequent exposure to the CD virus which causes high domestic dog pup mortality. Such protection would have ceased within a few years of the rinderpest decline as wild dog females born post rinderpest failed to pass on maternal protection to their pups. Thus CD could have caused the very poor reproductive success of the Serengeti wild dog population and other wild dog populations over a wide area, including the Mara between 1970-78.

It was the lack of young adults combined with a lack of immigrants which led to a slow decline in adult numbers on the plains (Appendix 6) not the increase in the Spotted hyaena population commuting to the plains in the wet season the resident population remained stable (Burrows et al 1994).

The sporadic re-emergence of rinderpest in domestic animals and wildlife in the late 1970s and early 1980s and the existence of sub-lethal strains in domestic stock (Dobson 1995) may again have provided protection for wild dogs and domestic dogs against CD virus. This would explain the resurgence of the Serengeti-Mara and other wild dog populations which occurred post 1978, following improving reproductive success. All blood samples taken between 1985-91 from Serengeti and Mara wild dogs were seronegative for exposure to canine distemper virus; the only confirmed disease related mortality during this period was from rabies.

It can be concluded that between 1970-77 there was a non significant decline in the number of adults in the study packs due to very poor reproductive success post 1969 but no decline in the number of study packs resident in the study area (Frame et al 1979, Burrows et al 1994 & 1995 & see Appendix 6). Based on Schaller's data up to 1969 the resident Serengeti adult population in 1970 in a 3000 km2 study area was likely to be 21-24 this compares well with the 29 based on the published data for 1970 (see Appendix 6)

Given a wild dog packs ability to suddenly expand when conditions are favourable and the ability of a pack to exist for a number of years even without good reproductive success as in Serengeti in the 1970s, the pack, rather than the number of individuals, should be taken as the basic unit of a wild dog population (Woodroffe & Ginsberg 1999).

Appendix 8. Sequence in the Serengeti -Mara study packs extinction 1986-91

All these study packs contained radio collared dogs when they died/disappeared. Fate of a 14th pack (Border Rovers or Triangle Pack) an unvaccinated study pack with home range on Tanzania /Kenya Border is unknown but was last reported in January 1991 when the pack was healthy although both radio-collared dogs had disappeared (Burrows 1993).

uv = unvaccinated packs.
v = packs containing vaccinated dogs
* radio-collar removed
italics = Mara Packs

Packs and date of extinction

1.	Pedallars uv	June	1986	7. Intrepids v.	December	1990
2.	Naabi uv	August	1988	8. Ole Sere v.	December	1990
3.	Aitong v.	September	1989	9. Ndoha. v.	February	1991
4.	Ndutu uv	November	1989	10. T. Blazers nv.	May	1991
5.	Lemuta uv.	February	1990	11. Salei v.	May	1991
6.	Mountain uv.	August	1990	12. N. Barafu v.	June	1991
				13. M&S* v.	June	1991

Table 1. Numbers of wild dogs photographed 1985-91 on the Serengeti Plains
There were 161 different individuals in 12 different packs and 6 groups.
The maximum number of packs seen in any one year was 5 as in early 1991

P=Packs; bip =born in pack; emi = emigrating groups; m=male; f=female

Pack/Group	n seen	'new' dogs

Plains P	37	37
Naabi P	7	3
Naabi bip	33	33
Pedallers P	13	10
Salei P	4	2
Salei bip	18	18
Ndoha P emi. f.	5	0
emi. m. 4 & 3	7	7
Barafu P	2	0
Lemuta P	3	2
Aitong f.	5	5
Mountain P	7	7
bip	13	13
T.Blazers P	6	0
2m	2	2
2f	2	2
Moru Track P	7	7
Ndutu P	17	13
N.Barafu P	6	0
3f.	3	0
M&S P	2	0
Totals	199	161

Table 2. Total verified population of adult wild dogs observed in the Serengeti plains study area 1974-76 (data from Frame 1977 (Table 3), & Frame et.al. 1979 (Table 1). Data given at 3 month intervals.

1974

1975

1976

Frame 1977

[44

41]

[39

37]

[32

30]

n Packs

cf Frame et al. 1979

but give data for

4 + 1 amg

4 packs

5 packs

* Data provided for only 4 packs; 3 of which were peripheral packs rarely seen, and did not den in the study area.

Frame & Frame's data was based on a study area of 5,200 km 2 in which in 1977 they counted 26 adults. This was NOT the entire population of Serengeti National Park, or the ecosystem as is implied by some commentators. In 1976 it was claimed that there were at least 7 more packs in the areas around their study area (Frame &Frame 1976).

As the researcher involved observed with reference to the Serengeti decline since 1975-76.

" there are several reasons to suspect that the decline in the population is more apparent than real" (Frame 1977).

Table 3. Sporadic deaths of unvaccinated study pack/group in Serengeti (n= 6) and Mara (n=1) post radio collaring 1985-90. Other study and non study packs in same area survived.

Pack survival = survival in months from handling to last sighting or to first deaths r.c. = radio-collared
older = more than 2 years old. bs =blood sampled , ylg = yearling (M) = Mara Group

pack	year r.c	age of dog r.c	pack survival/	cause	comments

Pedallars	1986	older male	<3	rabies supected	dead & dying at den the empty r.c found.
Naabi	1988	ylg female	<4	unknown	the empty r.c. found pack had been exposed to rabies pre the r.c. event
Ndutu	1989	older male	<3	unknown	a female collared as ylg in Naabi pack was bs at same time the 2 empty collars found
Henry group	1989	older male	<3	unknown	left Ndutu Pack early 89 seen with another dog before empty collar found near 2 Ndutu pack collars
Buffalo (M) Girls	1989 females	dispersing	<3	unknown	sister survived as Ole Sere alpha female
Lemuta	1990	older male	<1	unknown	not seen again post collaring the 2 females in pack were from Aitong Pack (Kenya) vaccination status unknown.
Mountain	1990	older f. May	<3	rabies	empty r.c. from f. found pups last to be seen alive as in Aitong pack 1989 rc m died underground
		old m. June	<2

Table 4. Multiple handling of packs both at vaccination and subsequent radio collaring of individuals post vaccination. Survival of pack in months following last anaethetization event in pack to first deaths.

All packs which became extinct contained vaccinated dogs.

ym = yearling male dart vacc = vaccinated against rabies by dart gun. bs = blood sampled rc = radio collared

Pack	year	age	survival	cause	n.vacc	comments

Aitong Mara	1989	3 pups	<2	rabies	>=4	pack part vaccinated since 1987 2 of the 3 r.c. pups were vacc. and 8 dogs anaetherized for bs a few months pre pack death a vacc dog died from rabies
Ole Sere Mara *	1990	?	<4	rabies	>=1	Rabid male was r.c as yearling in Serengeti (Ndoha Pack) in 1987. Collar battery ended early 1990. This male was dart vacc in early 1990 in the Mara.
Interepids Mara	1990	ym	<3	?	11/11	dead & dying dogs found no samples taken all pack members vaccinated in Mara
Ndoha Serengeti	1991	ym	<1	?	12/12	a female collared and vacc adults in Oct 90 was unsuccessfully darted. Alpha male not seen post vacc in Oct 1990. Empty collar found. pack not seen again post collaring
Salei Serengeti	1991	ym	<3	?	9/9	pup r.c Feb 91 dart vacc Oct 90 the empty collar found. alpha male seen alone in June.

Table 5. Packs in which most individuals were vaccinated but not handled again post vaccination.

ylg = yearling, r.c. = radio-collar , vacc. = vaccination

Pack	year	age m	survived months	cause	n. vacc	comments

Trail B Serengeti	1991	*ylg	<6	?	6/6	2 empty r.c. found
New Barafu Serengeti	1991	ylg	<6	?	5/8	neither collar found
M&S lone pair Serengeti	1991		<=8	?	2/2	no collars when lost collar removed from male Sept 90 when vacc.

* male in a group of 3 blood sampled in Nov 1990 before males joined 3 females to form pack

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