Captive Husbandry and Conservation of the Madagascar Ploughshare Tortoise, Geochelone yniphora
by James O. Juvik, Duane E. Meier and Sean McKeown
A captive group of Madagascar ploughshare tortoises or angonoka (Geochelone yniphora) has been maintained at
the Honolulu Zoo since April 1971 (Juvik & Blan, 1974). There was little reproductive activity in this group until the
late 1970's when an additional adult female G. yniphora was secured on breeding loan from the San Antonio Zoo in
Texas. Between September 1879 and September 1982 this tortoise laid a total of 25 eggs in 6 clutches. However, none of
these eggs proved to be fertile. Also, over this period, the zoo herpetology staff developed techniques to encourage
successful natural mating by periodically isolating the males and females and then reintroducing them to one another for
short periods. More natural mating attempts followed the adoption of this management approach. McKeown (1982) and
McKeown et al. (1982) have described general captive husbandry, courtship behavior and egg laying for this species.
Figure 1. Hatchling Geochelone
yniphora; age one month (October 1983); carapace length 50 mm; weight 32 g.
During the summer of 1982, several apparently successful courtship and mating sequences were observed involving the
male and egg laying female G. yniphora. Between September 1982 and February 1983, the female laid a total of five
egg clutches at approximately one month intervals. The egg laying dates and clutch sizes were as follows: 22 September
1982, four eggs; 25 October 1982, seven eggs; 1 December 1982, five eggs; 6 January 1983, six eggs; and 19 February
1983, three eggs. All eggs were incubated in covered plastic containers with moist vermiculite at 28.9° C (+/- 1.0° C).
On 12 March 1983, rotting eggs of the 25 October clutch were opened and one revealed a dead embryo (approximately 19 mm
in length). This was the first evidence of egg fertility among the many egg clutches laid by the female since 1979.
Other eggs of the 1982-83 clutches were examined and candled periodically during the summer of 1983 (two additional
dead embryos were discovered). One egg of the 2 February 1983 clutch, that appeared to be developing during periodic
candling, hatched successfully on 16 September 1983 after an incubation period of 226 days. At birth, the baby G.
yniphora was 42 mm long and weighed 24.6 g.
Figure 2. Electroejaculation of G. yniphora at the Honolulu Zoo (13 October
1982). Left: tortoise secured upright in working position. Right: penis relaxed and protruded, 2.5 cm
diameter rectal probe (with electrodes) inserted dorsal to exposed penis into the cloaca a depth of 10-12 cm.
Because of egg infertility in G. yniphora clutches laid in Honolulu between 1979 and 1982, Zoo staff became
interested in the possible application of semen collection/evaluation and artificial insemination (AI) procedures then
being developed for turtles and tortoises (Platz et al., 1980). With technical expertise and equipment provided by Dr.
C. Platz, Jr. (Texas A&M University) and Dr. Greg Mengden (Australian Museum-Sydney), an AI program was initiated at the
Honolulu Zoo. In October 1982, Mengden visited Honolulu, and began initial experimental work with the Zoo's G.
radiata (radiated tortoise) group. The electroejaculator consisted of a variable voltage power source and a plastic
rectal probe 2.6 cm in diameter with three 3.0 mm electrodes arrayed longitudinally. After first fully testing semen
collection procedures on a male G. radiata, a small quantity of motile sperm was successfully collected from the
male G. yniphora on 13 October, 1982 and immediately injected into the egg-laying female. This was the only AI
procedure with the female prior to September 1983. The female G. yniphora had laid a clutch of 4 eggs on 23
September 1982 (all proved infertile). Twelve days after the first AI procedure, on 25 October 1982, the female laid
another clutch of 7 eggs. It was from this clutch that the first evidence of egg fertility was recorded when a single
dead embryo was discovered in March 1983. We conclude that the interval between AI and egg laying (12 days) for this 25
October 1982 clutch provided insufficient time to credit the AI procedure for the recorded egg fertility; fertilization
was probably produced by natural inseminations from active mating episodes that occurred during the summer and fall of
Evident fertility associated with additional dead embryos recorded in egg clutches laid on 1 December 1982 and 19 February 1983, and the live hatchling (also from the 19 February 1983 clutch), is thus equivocal with respect to the
possible role of the October 1982 AI procedure or concurrent natural fertility. In July 1983, one of us (McKeown) left
the Honolulu Zoo, and the zoo's herpetologist position remained unfilled for almost a year. During this period AI work
with G. yniphora was under the direction of the zoo veterinarian. After a further series of electroejaculation
procedures were undertaken in September 1983, the male G. yniphora stopped eating and died 30 days later
(necropsy was inconclusive but suggested kidney failure).
Growth data for the 1983 G. yniphora hatchling are presented in table 1.
Table 1. Growth of captive born Geochelone yniphora in the Honolulu Zoo, 1983-1990.
CARAPACE STRAIGHT LENGTH (mm)
16 SEPT 1983
27 SEPT 1983
21 MAR 1984
22 SEPT 1984
30 MAR 1984
16 SEPT 1985
12 MAR 1986
10 AUG 1986
5 NOV 1987
19 SEPT 1988
15 SEPT 1989
23 JULY 1990
Figure 3 illustrates a normal growth curve (made linear by log transformation of the weight values), with weight and
age highly correlated (R=0.997; p<0.001) at age 2 years and above.
Figure 3. Relationship between age and weight (log scale) of the captive born G. yniphora in the Honolulu Zoo. Regression line fitted to data points beginning at two years of age.
Figure 4. Growth rates for the Honolulu captive bred G. yniphora in comparison with wild
tortoises (age estimated from annual growth rings). Wild tortoise data from Juvik et al., 1981 and Curl et al.,
If this growth curve was maintained over the next few years, the regression equation (log Y = 2.045 + 0.213X)
predicts that typical adult weight (10-15 kg) would be achieved at between 9 and 10 years of age. Figure 4 illustrates
the relationship between age and carapace length for the Honolulu captive-raised G. yniphora in comparison with
growth of wild tortoises in the Baly Bay region (age estimated from annual growth rings). The linear regression
equations yield annual growth increments of 25.77 mm/yr for the captive tortoise compared with 14.96 mm/yr in the wild.
At the captive tortoise's current age (6 years, 10 months) its shell length (225 mm) is 57% larger than that predicted
(143 mm, from regression) for a wild tortoise of comparable age.
Figure 5. Honolulu born G. yniphora at age 2.5 years (carapace length 106 mm). Abnormal "doming" of shell evident.
Gaymer (1968) in an analysis of comparative growth in Geochelone gigantea (Aldabra tortoise) found an even
larger difference in growth rates between captive-raised and wild populations. At age 7 years, the average carapace
length of captive-raised G. gigantea was 94% larger than in wild populations. This extreme difference may be
explained by generally suppressed growth in high density wild populations experiencing intense intraspecific competition
for food and shade (see Merton et al., 1976). Such competition-related growth suppression should not be a significant
factor in wild G. yniphora which now exist at very low densities.
Zovickian (1973) reported that sexual maturity in the closely related G. radiata is achieved at carapace
straight length of 330 mm (male) to 340 mm (female). If these values are roughly transferable to G. yniphora
(with a slight upward adjustment to compensate for the overall larger size of this species, i.e., 350 mm), then, based
on the above regression equations, the Honolulu captive should reach maturity at year 12, compared to year 20 for wild
In retrospect, the growth rate achieved in the Honolulu Zoo, captive-raised G. yniphora was too fast, and led
to distortions in allometric growth. At one year of age the captive G. yniphora exhibited abnormal shell
"doming". This species naturally possesses a high domed shell, particularly in adult females, where shell height
averages 62% of carapace straight length (Juvik et al., 1981). By age 3 years, the captive G. yniphora shell
height reached 76% of carapace length, although by age 6 this value had retreated to 73%. This shell deformity is not
only of aesthetic concern, particularly if carried into adulthood. Excessively domed shell geometry in G. yniphora
could be expected to lead to reduced reproductive success.
Our captive growth results in G. yniphora indicate that allometric relationships must be closely monitored
during the first few years. Diet and environmental factors should be manipulated to ensure that captive growth rates do
not get too far ahead of "natural" (wild) rates so as to produce shell distortions.
Figure 6. Ratio of shell height to carapace length in the Honolulu captive born G. yniphora
and wild tortoises (data from Juvik et al., 18981 and Curl et al., 1984)
Shortly after the departure of the supervising herpetologist (McKeown) to the Fresno Zoo in 1983, the diet for
juvenile tortoises which had consisted of chopped butter lettuce, romaine, kale, grated carrots, tomato and papaya was
modified by the Honolulu Zoo veterinarian. Additional fruit in the form of banana and apple were added along with
moistened (dry) dog food and later, ZU/PREEM Marmoset diet. We believe that the abnormal growth of the captive raised
G. yniphora may be related to both the additional fruit, as well as the dog chow and marmoset diet provided during
the first two years of its life.
Captive Breeding of Geochelone yniphora in Madagascar
Throughout the 1970's and early 1980's, the Malagasy Government (Direction des Eaux et Forets: DEF) maintained a captive breeding facility for confiscated native land tortoises at the Ivoloina Forestry station in Toamasina (Tamatave)
on the island's humid (windward) east coast.
On a visit to the Ivoloina facility in January 1984, the senior author found the tortoise collection to include 43
adult G. radiata (21 male, 22 female) and 7 adult G. yniphora (5 male, 2 female), all confined together in
a small thatched shed (area 22.5 m, or 2.2 tortoises/sq. m). In spite of space limitations the tortoises appeared well
cared for and in good health. The G. radiata had been breeding successfully for several years. About 50-60 eggs
were laid annually and incubated in closed wooden boxes of sand under ambient temperature conditions. Each year, 20-30
hatchlings emerged after an incubation period of 8-12 months (A. Befeno, personal communication, 1984). In January 1984,
there were 144 captive bred young G. radiata at the Ivoloina facility, ranging in age from recent hatchlings to 5
year olds. The largest captive bred animals (1979 hatch) had carapace straight lengths of 82-125 mm (X=105 mm; N=6), and
noticeable pyramiding of the carapace scutes.
In contrast to the prolific reproduction of G. radiata, no egg laying was reported for the G. yniphora
as Ivoloina. Although confined in cramped quarters with adult G. radiata, it was observed that G. yniphora
males made clear interspecific discrimination as both episodes of male combat and courtship in G. yniphora were
noted during the brief visit.
Because the Ivoloina site did not possess climatic conditions comparable (i.e., distinct wet-dry season contrasts) to
that found in the natural range of G. yniphora, it was earlier recommended in the Species Recovery Plan (Juvik et
al., 1982) that the Ivoloina animals be relocated to a west coast forestry facility in the Mahajanga area (some 100 km
east of Baly Bay) where general environmental conditions were more appropriate.
In 1986, with support from the World Wildlife Fund, the Jersey Wildlife Preservation Trust (JWPT) in cooperation with
the Malagasy DEF, established a G. yniphora captive breeding facility at the Ampijoroa Forestry station near
Mahajanga. In September 1986, the Ivoloina G. yniphora (5 male, 2 female) were transferred to Ampijoroa. More
recently, additional captive held G. yniphora from villages in the Soalala area have been donated to the
Ampijoroa breeding program, bringing the adult tortoise population at the facility to 20 animals (8 males, 12 females)
in October, 1991. The first captive breeding success at Ampijoroa was achieved in 1987 with the emergence of a single
hatchling G. yniphora. By the end of 1989 a total of 21 captive hatchlings had been produced, including 12 in
1989 alone, from 7 clutches totaling 25 eggs (Reid et al., 1989). In 1990 there were a further 10 hatchlings, and for
1991, there are 55 eggs under incubation (Reid, pers. comm. November, 1991). Eggs have been left to incubate in situ in
outdoor tortoise pens, with wire screen covers placed over nest sites to provide protection from predators. Natural
incubation periods have ranged from 168-266 days and are apparently correlated with wet-dry season onset and duration
Wild Populations of G. yniphora: Status and Prospects
Figure 7. Location map, northwestern Madagascar
Based on field surveys during the early and mid 1970's, Juvik et al. (1981) documented the presence of wild G.
yniphora populations at four different dry forest locations in the Baly Bay area:
1. North Ankoro (tortoise seen in wild, see Andrianarivo, 1977)
2. Cape Sada (several tortoises seen in wild)
3. East of Andanivato (fresh tortoise dung)
4. Forest patches between Anky and Antanandava (fresh tortoise dung)
More recent surveys in 1983 and 1986 (Curl et al., 1984, 1985 and Curl, 1986a) collected considerable anecdotal
information (and documented a substantial number of captive tortoises in villages) but failed to uncover or confirm any
new wild populations beyond those listed above.
In January 1990, Don Reid, the Madagascar based conservation officer with JWPT, spent two weeks in the Baly Bay region surveying G. yniphora populations and potential protected area sites. Reid was unable to visit the forest
locality north of Ankoro (Anjaha) on the west side of Baly Bay but visits to this area in April, 1991 by local Malagasy
nature protection officials confirmed the continued presence of a significant G. yniphora population in the
Belambo forest to the north and east of Ankoro. This habitat area may well prove to be as important as Cape Sada for
G. yniphora conservation and is currently the focus of ongoing field work.
Reid (1990b) also interviewed villages at Antanandava concerning the status of G. yniphora populations inhabiting remaining forest patches to the north. These dry forest areas were reported to have been heavily degraded by
fire and cattle grazing over the past few years, with villagers convinced that no tortoises remained in the area. Reid
(1990b) did find one adult G. yniphora in the wild, in a small (30 hectare) dry forest "island" (surrounded by
palm/grass savanna) in the Betheta/Anamboho area. Reid (1990b) also visited Cape Sada and confirmed the earlier
recommendations of Juvik et al. (1982) and Curl et al. (1985) that Cape Sada is the most appropriate site for initial
protected area status based on the following factors:
a) 200-250 hectares of suitable tortoise habitat (little affected by burning) which could potentially support up to
several thousand tortoises;
b) the absence of a permanent human population or intensive human utilization of the area;
c) ease and economy of constructing a pig/cattle barrier fence across the Cape Sada peninsula (about 2 km); and
d) the convenience of access (by boat) from the administrative center of Soalala.
Juvik et al. (1982) gave specific design specifications on a mesh and barbed wire pig fence for Cape Sada, based on
successful fencing techniques used to control feral pigs in Hawaii Volcanoes National Park. According to Reid (1990b)
local Malagasy authorities expressed some concern that villagers near Cape Sada might well find alternative uses for
valuable fencing materials. This potential problem can possibly be overcome by employing local villages in fence
constructing and maintenance and providing each nearby village with some fencing material for pig traps and garden
In a recent uncritical review of the literature on G. yniphora conservation, Burke (1990) draws several
unsupported and errant conclusions which only detract from ongoing conservation efforts for the species. Perhaps
confused by contradictory assertions on the persistence of vegetation boundaries around Baly Bay (Curl et al., 1984,
1985 and Curl 1986b), Burke (1990) concludes there is little evidence for ongoing tortoise habitat degradation, and that
"the situation of the wild angonoka (G. yniphora) may be somewhat less precarious than previously thought." Such
contentions are merely "wishful thinking" in view of recent field work results (Reid, 1990b), and our own (Kiester and
Juvik, in preparation) assessment of regional habitat change based on comparative analysis of early aerial photography
(1949) and recent hand-held Space Shuttle photography and Landsat thematic mapping images (both from December, 1990).
Although preliminary remote sensing analysis indicates that some forest fragments have disappeared over the 41 year
interval, other forest patches have retained their boundaries virtually unchanged (e.g., the Antanandava Forest). Ground
surveys, however, indicate that even in these persistent forest fragments human induced habitat degradation is
diminishing the quality if not the quantity of the areas for native wildlife. These conclusions are consistent with the
recent work of Andrianarivo (1990) who studied dry forest cover changes from satellite data for a nearby area (Mahajamba
Bay) in western Madagascar. Although Andrianarivo found little net change in total forest cover between 1973-84, ground
surveys revealed qualitative forest ecosystem decline in the form of reduced tree basal area and lower densities for
Burke (1990) also asserts that Cape Sada is unsuitable for a nature reserve because its small size (200-250 hectares
of tortoise habitat) would probably not support a viable tortoise population. If pigs and cattle were effectively
excluded (by fencing) from Cape Sada, G. yniphora would remain the only significant terrestrial herbivore in an
ecosystem which could easily support several tortoises per hectare, or a total population exceeding 1,000 animals. It
should be noted that much larger tortoises (G. gigantea) exist at average densities of 17/hectare on the south
coast of Aldabra Atoll, some 600 km to the north of Baly Bay (Hnatiuk et al., 1976). We agree that Cape Sada comprises
less than ideal habitat size, however there is simply no realistic alternative at the present time given current land
use trends and cultural practices in the region.
The authors wish to thank Dr. Lee Durrell and Mr. John Hartley of the Jersey Wildlife Preservation Trust for access
to unpublished reports and photographs.
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From: Proceedings of the First International Symposium on Turtles & Tortoises: Conservation and Captive Husbandry, 127-137, 1991.