REPRODUCTION IN FEMALE WILD GUINEA-PIGS

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REPRODUCTION IN FEMALE WILD GUINEA-PIGS
                       J.   P. ROOD      and   BARBARA J. WEIR

    Department ofZoology, MichiganState University, East Lansing, Michigan, U.S.A.
    and Wellcome Institute of Comparative Physiology, Zoological Society of London,
                           Regent's Park, London, jV. W. 1
                                (Received 10th December 1969)
   Summary. The breeding characteristics of three species of wild guinea-
   pig (F. Caviidae) are reported. Cavia aperea, Galea musteloides and Micro-
   cavia australis were studied in Argentina in the field and in outdoor pens,
   and laboratory colonies of the two former species were also established
   in England. Pens of domestic guinea-pigs (Cavia porcellus) and of C. aperea
   \m=x\C. porcellus hybrids were maintained in Argentina for comparisons with
   C. aperea.
       C. aperea and G. musteloides gave birth in every month but there was a
    breeding peak in spring (September to December). Microcavia had a
    more restricted breeding season ; in the field study area, births occurred
   only between August and April. Gestation length in C. aperea was variable
    but the mode was at 61 days, while the modes of Galea and Microcavia
   were much shorter at 53 and 54 days, respectively. All three species
   exhibited a post-partum oestrus and Galea may experience a lactation
   anoestrus. Oestrous cycle lengths in C. aperea and Galea varied consider-
   ably but the mean length in Cavia was 20\m=.\6\m=+-\0\m=.\8days and in Galea it
   was 22\m=.\3\m=+-\1 \m=.\4days; in the latter species, the presence of a male in the same
   cage was necessary for the induction of oestrus. Average litter size was
   2\m=.\2for C. aperea, 2\m=.\6for Galea and 2\m=.\8for Microcavia. In the Argentine
   colonies, the age at first conception in C. aperea and Galea varied with the
   time of year of birth, but this variation was not maintained under the
   more equable laboratory conditions.

                                     INTRODUCTION
Although the domestic guinea-pig, Cavia porcellus, has been kept in captivity for
over 400 years, little information has been published on wild guinea-pigs.
Those studies which have been reported have mainly been concerned with the
taxonomy and distribution of cavies (Cabrera, 1954) and with the genetics of
the genus Cavia (Detlefson, 1914; Castle & Wright, 1916; Ubisch & Mello,
1940). These authors give some data on reproduction in the species which they
studied and Pearson (1951) gives a little information on Galea musteloides in
Peru. Nothing else appears to have been published on the reproductive biology
of any wild members of this family (Caviidae) of hystricomorph rodents. The
             *
                 Enquiries concerning reprints should be addressed to Dr     . J. Weir.
                                                393
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394                        J. P. Rood and Barbara J. Weir
literature on reproduction of the domestic guinea-pig is, of course, extensive
 (see Asdell, 1964; Marshall, 1952-66).
    Various other hystricomorphs have been studied, mainly in captivity, and
several unusual features of their reproductive physiology have been reported
 (Weir, 1967a). All except the coypu (Myocastor coypus) have a vaginal closure
membrane which is perforate at oestrus and parturition. Apart from the
sub-family Caviinae, all other hystricomorphs investigated have very long
pregnancies of over 100 days and this is not associated with a delay of implanta¬
tion such as is found in various other species (e.g. roe deer, stoat, bear, badger,
 armadillo). The oestrous cycle is also longer in other hystricomorphs than the
 16^-day cycle of the domestic guinea-pig. For example, both the chinchilla
 (Chinchilla laniger) and the plains viscacha (Lagostomus maximus) have a mean
 cycle length of 41 days (Weir, 1966; Weir, in preparation) and the cycle length
in the dasyproctids (Dasyprocta and Myoprocta) is about 35 days (Weir, 1967b;
 Kleiman, 1969).
    During the course of investigations into the comparative behaviour of wild
 guinea-pigs in Argentina (Rood, in preparation), information was obtained on
 the breeding characteristics of three species of different genera. Colonies of two
 of these species were also established in London and the breeding data compiled
 at both places are compared below. The species identifications for Galea
 musteloides (PI. 1, Fig. 2) and Microcavia australis (PI. 1, Fig. 1) are based on
 Cabrera (1954); but for Cavia aperea (PL 1, Fig. 3) the recommendation of
 Hiickinghaus (1962) has been followed.
                        MATERIALS AND METHODS
Observations in Argentina (December 1965 to February 1968)
   The three wild species (Cavia aperea, Galea musteloides and Microcavia australis)
were observed in the areas shown in Text-fig. 1 and limited trapping and
observations were also made in several other localities. Colonies of each of
these three species, and of the domestic guinea-pig (Cavia porcellus) and of
C. apereaxC. porcellus hybrids were maintained in large pens at Sierra de la
Ventana, Buenos Aires Province.
   Cavies were trapped live in the study areas by mesh, treadle-type traps
 (National Live Traps) placed in their runways. Each animal was individually
marked by toe clipping and with Nyanzol dye. Total length and weight were
recorded and compared with data on animals of the field and pen populations of
known age in order to assess approximate ages. Pregnancy was determined by
palpation.
   The pens were initially stocked with Galea and C. aperea trapped within a
40-km radius of Sierra de la Ventana; the Microcavia colony was established
with animals brought from the study area in southern Buenos Aires Province.
The original C. porcellus stock was donated and the hybrids were obtained by
crossing female C. porcellus with a C. aperea male caught in the wild. The colonies
were allowed to grow naturally until about twenty-five animals were present
and were then controlled by removing newly weaned animals. Such removals
were only necessary for the Galea population as, although all the species did

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PLATE 1

                  Fig. 1. Microcavia australis
                                             ( 0-5).
                  Fig. 2. Galea musteloides ( 0-5).
                  Fig. 3. Cavia aperea ( 0-3).

(Facing p. 394)

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Reproduction infernale wild guinea-pigs               395
breed in   the pens, the reproductive success varied considerably and was highest
in Galea and lowest in Microcavia.
   The wire mesh pens were each 5 5 2 m high ; the mesh was extended 75 cm
underground to prevent escape but only the Microcavia were found to burrow.
The principal shelter in each pen consisted of four waterproof nest boxes which
were  kept supplied with fresh sawdust. Additional shelter was provided by an
inverted metal tub in the Microcavia pen and open wooden boxes in the other
pens. All the pens were supplied with branches for climbing. Commercial rabbit

       Text-fig. 1. Map of part of Argentina to show the areas of study and the sources of cavies
       investigated in captivity: (1) Cavia aperea study area; 'Estancia El Destino', 20 km S.E.
       Magdalena. (2) Microcavia australis study area; 'Estancia La Cristina', 14 km S.E.
       Cardenal Cagherò. (3) Galea musteloides study area ; 'Estancia La Sorpresa', 5 km E. Torn-
       quist, and source of G. Musteloides for London colony. A, Source of C. aperea for London
       colony. ·,   All three   species in   the   penned colonies, Sierra   de la Ventana.

pellets (Purina),   whole oats and water were always available; a selection of
grasses, carrots, lettuce, apples and potatoes was given once a day. Lawn grass
was grown under wire mesh frames to supplement the supply of green food.

Observations in London          (August      1967     to   August 1969)
  The C. aperea and G. musteloides colonies were derived from three C. aperea
(one male, two females) caught in the wild and from ten Galea (four males, six
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396                       J.   P. Rood and Barbara J. Weir
females) which were either from the pen colonies (see above) or had been
trapped immediately before their transfer to London. Both species were initially
kept in small groups in standard guinea-pig cages, but C. aperea is now housed in
floor pens and Galea in mesh cages with solid floors. Details of the housing and
husbandry are given by Weir (1970). The animals were checked daily for
reproductive condition and all events, such as vaginal opening, copulatory
plugs and litters, were recorded. Oestrous cycle length was defined as the
interval from the 1st day of vaginal opening of one cycle up to, but not including,
the 1st day of opening at the next cycle. The day of finding a copulatory plug
was recorded as Day 1 of pregnancy; most gestation lengths were, however,
determined from the interval between births of litters as there is an immediate
post-partum oestrus in these species.
  Standard statistical tests were applied and mean values are followed by the
standard error of the mean.

                                     RESULTS
                                    A. CAVIA SPP.
1. Annual reproductive cycle
   Under laboratory conditions, domestic guinea-pigs breed throughout the
year, although slight seasonal variations in reproductive performance have been
reported (Rowlands, 1949; Wright, 1960). Castle & Wright (1916) reported
that their colony of Cavia cutleri, obtained from Peru and maintained in the
United States, bred in all months, but thirty-eight litters were born from January
to June as against fifteen born from June to January.
   It is clear from Text-fig. 2 that, in Argentina (B, C and D), Cavia species
breed throughout the year, although there is an indication that reproductive
activity is reduced in C. aperea (B) during the autumn and winter months (April
to August). In the combined data for the 2-year period studied, 42% to 87%
of C. aperea females over 3 months old were pregnant in each month from
August to February whereas only 9% to 25% were pregnant from March to
July. In the field study area, five of thirteen females in March 1967, none of
twelve in August 1967 and four of four in January 1968 were visibly pregnant
or lactating. The incidence of winter breeding differed in the 2 years studied.
In the pen, several C. aperea females produced litters in the winter of 1966 but
none did so in the unusually cold winter of 1967. In the field, one of six females

trapped in August 1967 at Balcarce, Buenos Aires Province, was visibly
pregnant whilst all females trapped in the study area in August 1967 were non-
breeding.
  No seasonal pattern of breeding      was   apparent in the London colony of C.
aperea (Text-fig. 2A).
2. Oestrus
   Observations of the vaginal closure membrane in twenty-one laboratory
C. aperea (Text-fig. 3) gave a mean cycle length for sixty-seven cycles of 20-6+ 0-8
days. Females of all species conceived most frequently at the post-partum oestrus
and one laboratory C. aperea had six consecutive pregnancies resulting from

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Reproduction infernale wild guinea-pigs                 397
post-partum matings. In Argentina, the maximum number of litters produced by
a female in 1 year was five in the Fj hybrids, five in C. aperea and four in C.

porcellus. When conception did not occur at the post-partum oestrus, there was no
evidence of a lactation anoestrus in five of the laboratory animals.

        Text-fig. 2. Monthly distribution of births in species of Cavia: A, C. aperea in London
        (August 1967 to August 1969) ; B, C. aperea in Argentina (February 1966 to February 1968) ;
        C, C. porcellus in Argentina (February 1966 to February 1968); D, C. aperea C. porcellus
        Fj hybrids in Argentina (January 1967 to January 1968).

                 Text-fig. 3. Distribution of oestrous   cycle length in C. aperea in London.
3. Gestation
     The variation of       gestation length in Cavia species is shown in Text-fig. 4.
It   can   be    seen that the range is very much greater in laboratory C. aperea (A)
although        the mode at 60 days is near the mean of 60-9+0-3 days for C. aperea

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398                               J.   P. Rood and Barbara J. Weir
in  Argentina (B). In this small sample, the difference between the mean
gestation length afterpost-partum matings (62-7+ IT days) and after conceptions
at other oestrous periods (65-5+1-4 days) was not significant (P>0-05).
   Although there is no overlap for gestation lengths in the Argentine colonies
of C. aperea (B) and C. porcellus (C), recorded gestation lengths in large colonies

        Text-fig. 4. Distribution of gestation length in species oí Cavia: A, C. aperea in London
        (mean = 63-9±0-9 days); B, C. aperea in Argentina (mean 60-9 + 0-3 days); C,
                                                                             =

        C. porcellus in Argentina (mean     =67-0 + 0-8 days); C, C. aperea
Reproduction infernale wild guinea-pigs                                              399
4. Litter size and sex ratio
   In the present study, litter size was similar in all species (Text-fig. 5) and the
overall mean was 2-3.
   Of 117 C. aperea born in the pens in Argentina and in the laboratory, forty-
nine were males and sixty-eight were females.

5. Age at onset of reproduction
  Although a few exceptional female guinea-pigs kept under laboratory
conditions have been reported to come into first oestrus at 20 days (Reid, 1958;

     Text-fig. 5. Distribution of litter size in species of Cavia: A, C. aperea in London (mean =
     2-5 young/litter); B, C. aperea in Argentina (mean          =  2-1 young); C, C. porcellus in
     Argentina (mean 2-1 young); D, C. aperea JxC.porcellus $ in Argentina (mean 2-3
                        =                                                                             =

     young) ; E, Hybrid C. aperea C. porcellus FjS in Argentina (mean = 2-5 young).
                                              Table 2
            age at first conception in relation to season of birth in
                                  Cavia species

                                             Estimated age at   conception (days)
                                                                  C. aperea        C.   porcellus
               Month born        C. aperea     C.   porcellus          Fts               F2s
           March                                    143
           June                                     102, 109
           July                                     45
           August                                                    73                 73
           September               62,30                                                36, 33
           October                 89               31               29
           November                                                  48,29              43

Ibsen, 1950), the majority probably do not do so until they are over 30 days
old. Young, Dempsey, Hagquist & Boling (1939) recorded a mean age at first
heat of 67-8 days (range 33 to 134 days). In wild forms, Castle & Wright (1916)
reported that female C. cutleri could breed at 2 months and Ubisch & Mello
(1940) recorded two female C. rufescens which littered at 3 months and thus
conceived at about 1 month of age.
   The estimated ages at first conception Cavia females born in the pens during
different months are shown in Table 2. Of the ten females born from September
to November, six (representing all forms studied) conceived at their first
oestrus when 29 to 36 days old. The three animals, all C. porcellus, born between

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400                            J.   P. Rood and Barbara J. Weir
March and June did not conceive until they were more than 3 months old suggest¬
ing that onset of reproduction may be delayed in autumn- and winter-born
females.
   In the laboratory colony of C. aperea, mean age at first vaginal opening in
twenty-seven animals was 58-6+3-0 days but only three females conceived at
this first oestrus, the youngest being 27 days of age. The mean age at conception
in ten animals was 84-4+9-9 days and first conception occurred up to the fifth
oestrus after puberty. There was no evidence in the laboratory colony that the
age at puberty or conception varied according to the time of year of birth.

                                     B. GALEA     MUSTELOIDES
 1. Annual reproductive cycle
    The field observations suggested that wild populations of Galea may produce
litters throughout the year and that most litters are born in spring and summer.
All five adult females caught in December 1965, two often in June 1966 and
three of seven in April 1967 were pregnant or lactating. In the pens in Argentina
(Text-fig. 6A) and in the laboratory in England (Text-fig. 6B), births occurred in

      Text-fig. 6. Monthly distribution of births in Galea musteloides: A, In Argentina in the pens
      (February 1966 to February 1968); B, In London (August 1967 to August 1969).

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Reproduction infernale wild guinea-pigs             401
every month. In the Argentine colony, more than 50% of females over 3 months
of age   were   pregnant in any          one  month but at least 90 % were pregnant from
September February,
               to                   indicating increased reproductive activity in spring and
early summer.
2. Oestrus
    Galea also has a vaginal closure membrane which is open for 1 to 6 days at
oestrus and parturition, although closure is very rapid after mating has taken
place. The majority of animals mate at the post-partum oestrus and once maturity
is reached, most females are perpetually pregnant. A female caught in the wild
in Argentina produced twelve litters in 21 months and in London, a female had
fifteen consecutive pregnancies.

                                                                                        —r-r
                    12   14   16   18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48
                                               Length of cycle (days)
     Text-fig. 7. Distribution of oestrous cycle length in laboratory G. musteloides; black
     columns denote cycles in the presence of a vasectomized male, open columns denote other
     cycles.

   Spontaneous oestrous cycles are, therefore, uncommon and only three cycles
of 17, 17 and 18 days were recorded in Argentina. In England, the mean cycle
length of fifty-six observations on twenty-six animals (Text-fig. 7) was 22-3+1-4
days and most of these cycles occurred before the first conception (see below).
When females were isolated from males, either singly or in groups, spontaneous
cycles did not occur even if the male was separated only by a mesh partition.
For example, a pregnant female was removed from the male and littered on
23rd January 1968; no periods of vaginal opening were observed until the male
was re-introduced on 14th May 1968 and a fertile oestrus occurred 2 days later.
To follow consecutive cycles in individual females, it was necessary to use a
vasectomized male. All four females run with this male then cycled at regular
intervals. The mean length of twenty-one cycles in the presence of the vasectom¬
ized male was 19-0+1-2 days and this was not significantly different from the
mean length for
                   sporadic cycles. It is clear that the presence of the male is very
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402                      J. P. Rood and Barbara J. Weir
important in this species for the initiation of oestrus. Adult males 'chin-rump
follow' a female with increasing frequency as she approaches oestrus, suggesting
that this pattern may be instrumental in inducing a pro-oestrous condition
(Rood, 1970).
   In five laboratory females which did not conceive at the post-partum oestrus,
the mean interval to the next oestrus was 41-8+4-0 days. This was significantly
different (P
Reproduction infernale wild guinea-pigs                 403
5 Onset of reproduction
   In Argentina, nine females born from June to December conceived at a mean
age of 58 days (range 28 to 93 days) while six females born from February to
April did not conceive until a mean age of 167 days (range 113 to 255 days).
This suggests that females born in the autumn do not breed until the following
spring, while females born in the spring breed that summer.
   Under laboratory conditions, there was no evidence of any seasonal effect of
the time of birth on age at first conception. In thirty-four animals, all of whom
were housed with a male from birth, the mean age at conception was 60-6 days
(range 17 to 152 days) ; twenty-one of these conceived at the first oestrus. The
mean age at puberty (as determined by first vaginal opening) in fifty-two females
was 48-5 days (range 15 to 170 days).

     Text-fig. 9. Combined distribution of litter size of G. musteloides in     Argentina      and
     London.

   Three females were housed at 21 days of age with an adult male in an indoor
cage in Argentina and were checked twice daily for vaginal opening. One came
into oestrus and mated at 32 days of age and the other two were then separated
from the male by a mesh partition. No vaginal opening occurred within the
next 20 days, but when the females were replaced with the male both came into
oestrus within 2 days (at the ages of 53 and 54 days). Although these values
fall within the normal range for the interval to puberty, the experiment indicated
that the initiation of puberty may also be dependent upon the presence of a
male (see above).

                                 C. MICROCAVIA AUSTRALIS

1. Annual   reproductive cycle
   In a field study area, litters were born from August to April with the majority
of births between September and December (Rood, 1970). In the pens, litters
were born from October to February and none of the females was pregnant

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404                            J.   P. Rood and Barbara J. Weir
from March to July. Therefore, both the field and pen data indicate that
Microcavia has a more restricted breeding season than Cavia or Galea.
   Three females resident in the pen in the winter of 1966 all produced their
first spring litters within 3 days of each other (from 7th to 10th October) and
the only survivor of the winter of 1967 also gave birth in this period (on 7th
October), suggesting that reproductive activity may be triggered by a specific
environmental stimulus.

2. Oestrus
  Microcavia also has a post-partum oestrus and the vaginal closure membrane is
perforate for about 5 days after parturition. One female came into oestrus and
mated 15 days after giving birth and three other animals, which were thought to
have aborted upon capture, conceived within 15 days of introduction into the
pen.

3. Gestation
   This was estimated from the interval between consecutive litters and the
mean length for six pregnancies was 54-2+0-5 days (range 53 to 55 days).
Gestations of 51 and 52 days were also recorded but the young appeared to be
premature and died within a few hours of birth.
        size and sex ratio
4. Litter
 The distribution of litter size is shown in Text-fig. 10. In the pens, twenty-one
males and nineteen females were born.

                    Text-fig. 10. Distribution of litter size in Microcavia australis.

5. Onset    of reproduction
  Information on the age at which breeding first commences is limited since
mating of young females at normal oestrus was not observed in the pen colony
and only one female born in the pen subsequently produced a litter. This

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Reproduction infernale wild guinea-pigs             405
parturition took place when the female was 138 days old and, therefore, con¬
ception occurred when she was about 85 days old. In the field study area, two
young females born in the spring were observed mating when they were 40 to
50 days old (Rood, 1970).

                                  DISCUSSION
The breeding record for the penned Cavia species indicated that, under identical
conditions, the most successful breeders were the hybrids. The four hybrid Ft
females were continuously pregnant after they reached maturity whereas some
of the C. porcellus and C. aperea females were not. C. porcellus appeared better than
C. aperea but the reproductive performance of the latter was lowered by four
females which lived in the pens for 6 months without breeding. The C. aperea
also appeared to be more sensitive to climatic conditions because in the very
cold winter of 1967, breeding ceased in the pens as well as in the field study area.
The capture of a pregnant female in the middle of the milder winter of 1966
shows that winter breeding does occur when conditions are favourable. Wild
Cavia may have been affected by food shortage but this explanation was not
applicable to the penned animals which were fed regularly.
   The Argentine populations of Galea are similar to the Cavia species in the
characteristics of the annual reproductive cycle, although Pearson's (1951)
data suggest that G. musteloides in Peru may be a seasonal breeder as breeding
condition was found in males and females trapped in December, but not in
animals trapped in July and September. There was no peak of breeding activity
in the laboratory Galea colony, probably because, under the constant environ¬
mental conditions, females became sexually mature at approximately the same
age, regardless of the time of year of birth. In the Argentine colony, winter
breeding was restricted to females which conceived at the post-partum oestrus.
   The southern population of Microcavia australis studied has a restricted breeding
season and it would be interesting to know if the Cavia species which live on the

Altiplano also breed at a limited season of the year. The indications are that the
breeding season of cavies may be determined by climatic conditions, most
probably by temperature.
   The occurrence of a post-partum oestrus and ovulation is a common feature of
the Caviinae. It has been found in the three species described in this paper and
inC. cutleri (Castle & Wright, 1916) andC*. rufescens (Ubisch & Mello, 1940). In
favourable conditions, therefore, females of these species are constantly pregnant
after reaching maturity. The rapidity of re-impregnation must offset the rela¬
tively long gestation period and, together with the fact that the young are
precocious at birth, must contribute greatly to the success of these species in
competition with the more prolific myomorph rodents which have shorter
pregnancies. This post-partum characteristic is used extensively to ensure
maximum output of young in most large-scale breeding colonies of domestic
guinea-pigs.
   The possible association, in Galea, of a male sexual pattern (the chin-rump
follow) with the initiation of oestrus, and possibly also of puberty, causes
speculation as to whether a similar phenomenon will be discovered in Microcavia
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406                        J. P. Rood and Barbara J. Weir
australis since this species also exhibits this distinctive behaviour pattern (Rood,
 1970). The introduction of a male has also been shown to affect the time of onset of
oestrus in the mouse (Whitten, 1956) but in this species, unlike Galea, the females
undergo spontaneous oestrous cycles in the absence of a male and the excreta
is effective in modifying the cycle. It will be interesting to determine if a
post-partum ovulation occurs in the absence of a male in Galea. This species was
the only one studied by us which may have a lactation anoestrus, a phenomenon
also found in another hystricomorph rodent, the chinchilla (Weir, 1967a).
   The oestrous cycle lengths recorded for C. aperea and Galea are very variable
and it may be argued that these are abnormal because the animals were disturbed
under laboratory conditions. However, the similarity of all the data compared
for the English and Argentine colonies suggests that the laboratory studies are
valid. It has also been shown that most cycles in the chinchilla are 30 to 50
days (Weir, 1966) and in the plains viscacha, only 75% of cycles are between
25 and 55 days (Weir, in preparation). Therefore, this variation is probably
normal for other wild and newly domesticated species of hystricomorph rodent.
                                                     Table 3
            comparison of litter size in species of the sub-family caviinae

                                                         Litter size
                    Species
                                                     Mean         Range                         Authors

Microcavia australis                                  2-8         1 to   5      Present study
Galea musteloides                                     2-6         Ito    5      Present study
Cavia aperea (Argentina)                              2-1         1 to   5      Present study
Cavia aperea (London)                                 2-5         1 to   5      Present study
C. cutleri                                            1-89        1 to   4      Castle & Wright       (1916)
C. rufescens                                          1-35                     Detlefson (1914)
C. porcellus                                          2-1         1 to 5       Present study
C. porcellus                                          2-58        1 to 8       Haines (1931)
C. porcellus (MRC stock,      Frant)                  4-0         1 to 10      Rowlands (1949)
C. porcellus (MRC stock,      Pirbright)              4-3                      Hoyland (1965)
C. porcellus (MRC stock,      Frant)                  3-5         1 to 13*     Sutherland
                                                                               (personal communication, 1969)
C. aperea   C. porcellus   hybrid F1s                 2-5         1 to 6       Present study
                                *
                                    The largest viable litter contained nine young.

    The most obvious reproductive character for which domestic guinea-pigs
have been selectively bred is litter size. In 1931, Haines reported that the mean
litter size in his large sample of guinea-pigs was 2-58 (range 1 to 8) while the
Dunkin-Hartley, or MRC, strain which originally had a mean litter size of 3-0
(Dunkin, Hartley, Lewis-Fanning & Russell, 1930) was developed at Frant to
produce 4-0 young per litter (Rowlands, 1949). Recently, the Pirbright strain
of the MRC stock has been reported as having a mean litter size of 4-3 (Hoy-
land, 1965). A comparison of the reported variations in litter size of different
species and strains of Caviinae is shown in Table 3.
    The observation that, in the domestic guinea-pig, smaller litters were pro¬
duced after longer pregnancies was recorded by Rowlands (1949) and by Goy
et al. (1957), and it is interesting to note a similar trend in C. aperea, although this
was a small sample. Since domestic guinea-pigs have been bred to produce

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Reproduction infernale wild guinea-pigs              407
larger litters, it is surprising that there has been no apparent shortening of
gestation length. The pregnancies of other Cavia species are shorter than that of
C. porcellus (see Table 4), and mating a female C. porcellus to a C. aperea male
appeared to shorten pregnancy by several days. This has been found for all
crosses of C. porcellus with wild cavies (Pictet & Ferrerò, 1951 ; Ubisch & Mello,

 1940). It is noteworthy that Galea and Microcavia have gestation lengths (52-7
and 54-2 days respectively) which are not only shorter than those Cavia species
but are the shortest known within the Hystricomorpha.
   The youngest animals to conceive in the present study were a 17-day-old
Galea and a 27-day-old C. aperea. These values represent an extreme (see Table2),
but Ubisch & Mello (1940) reported that C. rufescens can breed at 1 month of age
and conceptions in domestic guinea-pigs less than 21 days old occur in large
colonies (Sutherland, personal communication). The loss of a seasonal effect on
the onset of reproduction in laboratory C. aperea and G. musteloides probably
reflects the fact that the environmental conditions                           were more            constant      than

                                                Table 4
                        reported gestation lengths in species of cavia

                                            Gestation   length (days)
                       Species              Mean              Range                       Author
        C. aperea                            62-4           59 to 74         Present study
        C. cutleri                           63-3           56 to 69         Castle & Wright (1916)
        C. rufescens                                        62 to 65         Ubisch & Mello (1940)
        C. porcellus                         680            59 to 72         Rowlands (1949)
        C. rufescens ¿* x C. porcellus Çj                   63 to 67         Detlefson (1914)
        C. aperea
408                             J.   P. Rood and Barbara J. Weir
Ferrero's colony. However, C. aperea would seem to be a more probable ancestor
of C. porcellus than, for example, C. rufescens, in which the male offspring of
crosses with C. porcellus are sterile (Detlefson, 1914; Ubisch & Mello, 1940).
Crosses of C. cutleri males with C. porcellus females have been reported to be
fertile (Castle & Wright, 1916), and Hiickinghaus (1962) considers that
C. porcellus, C. aperea and C. cutleri are all conspecific.

                                       ACKNOWLEDGMENTS
The field work in Argentina was supported by United States Public Health
Service Fellowship Grant 5 F2 HD-28, 615-01 from the Institute of Child
Health and Human Development, and a Training Grant GM-1751 from the
Institute of General Medicine. Hazel Rood assisted in all phases of the investiga¬
tion and Dr John King, Michigan State University, also contributed greatly to
the study through his co-operation and interest. We are also indebted to the
owners of the estancias on which study areas were established, namely Sr G.
Walter, Sr C. Olson and Sr R. Pearson, and to the Institute Nacional de
Tecnologia Agropecuaria, Buenos Aires, for their donation of the domestic
guinea-pigs used in the study.
   The laboratory work in England was financed by the Ford Foundation, and
the expedition to Argentina during 1967 when the cavies were obtained was
supported by the Royal Society, the Wellcome Trust and the Medical Research
Council. It is impossible to thank individually the many kind people who assisted
the expedition, but B.J.W. would like to thank particularly the Rumboll
family for their help in Argentina and Dr I. W. Rowlands for his constant
encouragement of the project.

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