Demography of a population with a long history of poaching and the utility of the individual identification technique as a tool for monitoring intermittently studied elephant populations

In many elephant populations in Africa, adverse effects of poaching including altered age and sex structure and long calving intervals continue to negatively impact the rate of recovery. The study reported here characterized the demographic status of the elephant population in Meru National Park, Kenya, so as to determine the size, age and sex structure, and calving interval of the population and to compare its demographic parameters against those from the relatively stable elephant population at Amboseli. Additionally, a demographic study conducted 7 years earlier enabled us to determine the demographic performance of the population and to explicitly test the utility of the individual identification technique in monitoring intermittently studied populations. We found 392 elephants in the park. The proportion of older individuals was lower, and the calving interval was longer than estimates from Amboseli. We recognized 16% of elephants indexed in the previous study. Our results suggest that the individual identification technique may be useful in monitoring demographic changes including poaching in intermittently studied elephant populations. Overall, our results suggest that although the Meru elephant population showed signs of recovery, poaching continues to imperil its recovery.     

AGE CRITERIA FOR THE AFRICAN ELEPHANT

1. The paper is based on a collection of 385 lower jaws of the African elephant (Loxodonta a. africana Blumenbach) from western Uganda. 2. Thirty age groups are described and illustrated, which are related to the progress of eruption and wear of the six teeth in each side of the lower jaw. 3. Correct identification of individual teeth in the series is essential and several checks are described. Thus, when length is plotted against width for the six teeth the points fall into six well - defined groups. Frequency distributions of laminary indices and lamella numbers support the belief that teeth have been correctly identified, but overlapping, distributions preclude identification on these characters alone. 4. The presence of abnormal seventh molars in four jaws is described. 5. Chronological ages have been assigned to the relative age groups. Information on age and growth of captive animals has been considered and the general form of the growth curve established. Arbitrary estimates of the intervals between successive age groups were made and subsequently checked against growth and seasonal ridges on the roots. These indicate an upper age limit of about 60 years, which is compatible with fuller information on Indian elephants. 6. Growth curves support the validity of the ages assigned. Asymptotic heights at shoulder (∞) are respectively 317 cm and 298 cm for males from Murchison Falls and Queen Elizabeth National Parks. For females no distinction is made (because the sample from the latter area is small) and ∞ is 272 cm. 7. Growth in weight has also been established but shows no significant differences between the two populations. The mean maximum weight of females is 2766 kg (6100 lb), and of males 5450 kg (12000 lb). Growth apparently continues throughout life. 8. Tusk growth is analysed. There is a linear increase in weight of female tusks from eruption at 1–3 years up to the oldest group, with an indicated mean combined weight of 17.7 kg (39 lb) at 60 years. Male tusks show an increasing rate of growth throughout life to a mean combined weight of about 109 kg (240 lb) at 60 years. Big tusks are generally the result of prolonged growth; extremely big tusks probably result from prolonged and above average rate of growth. 9. Seasonal and annual incremental layers on the root 3 of the teeth are briefly described; these give an objective estimate of the chronological intervals between the relative age groups. 10. Field age criteria are presented which are derived from these growth curves. 11. The age at puberty in the female elephant is discussed and compared with earlier conclusions. There is evidence of a retardation in recent years in the Murchison Falls National Park (South bank) population and of a lengthening of the mean calving interval. This considerable depression of the reproductive rate, which is almost certainly density dependent, agrees with the observed lower recruitment in this population. 12. Survivorship curves constructed from material representing 325 natural deaths are presented for the two populations. If estimates of the age at puberty and the reproductive rate are taken into account, the expected differences in recruitment are found. 13. The calculated mean expectation of life is less than 15 years. 14. Mean individual weight is estimated at about 3800 lb.     

Termination of reproduction in nonhuman and human female primates

We examined demographic records from 13 captive primate species and a human population to determine age-related changes in female reproduction. In most species age-specific fertility declined and interbirth intervals increased with age. Using an operational definition of termination of reproduction based on individual variance in interbirth intervals, a proportion of females in most nonhuman species had terminated reproduction before death. Compared to other primates, a greater proportion of chimpanzees and human females ceased reproduction, and humans, in particular, were reproductively inactive for relatively longer than would be expected from their body weight. These empirical data quantify the extent of reproductive termination and thereby extend hitherto anecdotal accounts of this phenomenon in primates.     

Termination of reproduction in nonhuman and human female primates

We examined demographic records from 13 captive primate species and a human population to determine age-related changes in female reproduction. In most species age-specific fertility declined and interbirth intervals increased with age. Using an operational definition of termination of reproduction based on individual variance in interbirth intervals, a proportion of females in most nonhuman species had terminated reproduction before death. Compared to other primates, a greater proportion of chimpanzees and human females ceased reproduction, and humans, in particular, were reproductively inactive for relatively longer than would be expected from their body weight. These empirical data quantify the extent of reproductive termination and thereby extend hitherto anecdotal accounts of this phenomenon in primates.     

Temporal changes in fecundity and age at sexual maturity of southern elephant seals at Marion Island

Our objective was to examine the effect of variation in reproductive parameters on the demography of southern elephant seals at Marion Island. We used age-specific capture probabilities of breeding females in a Cormack-Jolly-Seber context to derive reproductive rates. We found that age at maturity declined and fecundity rates increased as the population declined, indicating a compensatory response. Fecundity rates ranged from 0.03 to 0.29 among 3-year-olds (mean=0.16), 0.18 to 0.50 in 4-year-olds (mean=0.40), and 0.28 to 0.50 in 5-year-olds (mean=0.45). We think that a relative increase in food availability, concomitant with the population decline, promoted earlier sexual maturity correlated with more rapid growth of juveniles when population abundance was lower. It is suggested that the relative importance of fecundity in population regulation in elephant seals has been underestimated. Moreover, it appears that the onset of sexual maturity may be the first demographic variable to change in response to a change in population density.     

Age distribution of adult female screwworms (Diptera: Calliphoridae) captured on sentinel animals in the coastal lowlands of Guatemala

The age structure of screwworm, Cochliomyia hominivorax (Coquerel), populations was studied in the Guatemalan lowlands with the pteridine technique of age determination. Flies (95% were 4-15 d old) were captured on sentinel animals. Mean age was 7.5 d. The oldest fly captured was 21 d old. Most flies visiting wounds were reproductively mature females. Of these, about one-half oviposited once, one-third oviposited twice, and 1 in 10 completed three ovarian cycles (assuming a 3-d interval between clutches). Periodicity in adult capture rates may be related to the reproductive cycle. Results suggest that local populations are composed of a high proportion of synchronized cohorts.     

Demography of fluctuating prairie vole populations: comparison of demographic variables among phases of fluctuations

We tested for differences in the proportion of reproductively active males and females, proportion of the population composed of young and immigrants, and monthly survival (total, adult, young) among phases (trough, increase, and decline) and among habitats (alfalfa, bluegrass, and tallgrass) of 30 population fluctuations ofMicrotus ochrogaster Wagner, 1842 over 25 years in east-central Illinois USA. Total population survival and survival of adults and young were greatest during the increase phase, among fluctuations, irrespective of habitat. The proportion of reproductively active adult males and females was lowest during the decline phase, an effect of lower reproduction during the winter. These results suggest that phase-specific changes in survival were the primary demographic factor driving population fluctuations ofM. ochrogaster in our study sites. We conclude that small-scale spatially different population fluctuations may be explained by the same mechanisms that explain fluctuations within a population.     

Survey of the reproductive cyclicity status of Asian and African elephants in North America

The Asian and African elephant populations in North America are not self‐sustaining, and reproductive rates remain low. One problem identified from routine progestagen analyses is that some elephant females do not exhibit normal ovarian cycles. To better understand the extent of this problem, the Elephant TAG/SSP conducted a survey to determine the reproductive status of the captive population based on hormone and ultrasound evaluations. The survey response rates for facilities with Asian and African elephants were 81% and 71%, respectively, for the studbook populations, and nearly 100% for the SSP facilities. Of the elephants surveyed, 49% of Asian and 62% of African elephant females were being monitored for ovarian cyclicity via serum or urinary progestagen analyses on a weekly basis. Of these, 14% of Asian and 29% of African elephants either were not cycling at all or exhibited irregular cycles. For both species, ovarian inactivity was more prevalent in the older age categories (>30 years); however, acyclicity was found in all age groups of African elephants. Fewer elephant females (～30%) had been examined by transrectal ultrasound to assess reproductive‐tract integrity, and corresponding hormonal data were available for about three‐quarters of these females. Within this subset, most (～75%) cycling females had normal reproductive‐tract morphologies, whereas at least 70% of noncycling females exhibited some type of ovarian or uterine pathology. In summary, the survey results suggest that ovarian inactivity is a significant reproductive problem for elephants held in zoos, especially African elephants. To increase the fecundity of captive elephants, females should be bred at a young age, before reproductive pathologies occur. However, a significant number of older Asian elephants are still not being reproductively monitored. More significantly, many prime reproductive‐age (10–30 years) African females are not being monitored. This lack of information makes it difficult to determine what factors affect the reproductive health of elephants, and to develop mitigating treatments to reinitiate reproductive cyclicity. Zoo Biol 23:309–321, 2004. © 2004 Wiley‐Liss, Inc.     

Growth and age determination of African savanna elephants

Understanding the population dynamics of savanna elephants depends on estimating population parameters such as the age at first reproduction, calving interval and age-specific survival rates. The generation of these parameters, however, relies on the ability to accurately determine the age of individuals, but a reliable age estimation technique for free-ranging elephants is presently not available. Shoulder heights of elephants were measured in 10 populations in five countries across southern and eastern Africa. Data included shoulder height measurements from two populations where the age of each individual was known (i.e. Addo Elephant National Park, South Africa and Amboseli National Park, Kenya). From the known-age data, Von Bertalanffy growth functions were constructed for both male and female elephants. Savanna elephants were found to attain similar asymptotic shoulder heights in the 10 populations, while individuals in the two known-age populations grew at the same rate. The Von Bertalanffy growth curves allowed for the accurate age estimation of females up to 15 years of age and males up to 36 years of age. The results indicate that shoulder height can serve as an indicator of chronological age for elephants below 15 years of age for females and 36 years of age for males. Ages derived from these growth curves can then be used to generate age-specific population variables, which will help assess the demographic status of savanna elephant populations across Africa.     

Possible involvement of ecdysteroids in photoperiodically induced suppresion of ovarian development in a Japanese strain of the migratory locust,Locusta migratoria

In a Japanese population of Locusta migratoria, adult females become reproductively inactive under crowding and long days (LD) and reproductively active under crowding and short days (SD). The identity and titre of ecdysteroids in the haemolymph and ovaries from adult females reared under SD and LD were investigated by RIA/HPLC. The effects of exogenous juvenile hormone (JH) III treatments on the termination of such reproductive arrest and ecdysteroid contents in LD females were also examined. In general, ecdysteroid titres in both haemolymph and ovaries were significantly higher in reproductively active SD females than in reproductively inactive LD females. A clear difference was also observed in oocyte growth between SD and LD individuals. JH III applications (three consecutive topical applications, 150 μg per insect per day from day 3) stimulated ovarian development in LD females and significantly increased the haemolymph and ovarian ecdysteroids to a level comparable to that of reproductively active SD adult females.     

A possible role for the thyroid in reproductive inhibition in laboratory populations of the prairie deermouse (Peromyscus maniculatus)

Serum thyroxine concentrations were determined and found to be reduced in reproductively inhibited deermice selected from controlled growth laboratory populations. Daily injections of 1 microgram thyroxine in reproductively inhibited male deermice were shown to significantly increase testis weight and the number of spermatids with acrosome formation. Daily feeding of thyroxine to a controlled growth population promoted an increase in the mean weights of the reproductive organs of both sexes that was intermediate between reproductively inhibited and capable deermice. Also, two females in the population produced the first surviving young after 250 days of inhibition.     

Blood concentrations of gonadotrophins, prolactin and gonadal steroids in males and in non-pregnant and pregnant female African elephants (Loxodonta africana)

No seasonal variation in any of the hormones measured was apparent in males or females. Testosterone levels in males increased around puberty (10-11 years) and remained significantly higher in adult than prepubertal males. This was not accompanied by any significant change in levels of LH, FSH or prolactin. In non-pregnant females there was no apparent difference in levels of LH, FSH or prolactin with age. There was a significant increase in progesterone around puberty (12 years) but there was considerable overlap in values between prepubertal and adult females. During pregnancy, progesterone levels were significantly higher than in non-pregnant females with maximum levels occurring at mid-pregnancy (9-12 months). However, there was considerable overlap in values between non-pregnancy and pregnancy. Concentrations of LH and FSH decreased significantly during mid-pregnancy while prolactin levels increased dramatically during pregnancy; after 7 months of gestation until term levels were always at least 8 ng/ml greater than in any non-pregnant female. It is suggested that this consistent increase in plasma/serum levels of prolactin can be used to diagnose pregnancy in the elephant.     

Population characteristics of pallid sturgeon (Scaphirhynchus albus (Forbes & Richardson, )) in the Lower Missouri River

Population characteristics of pallid sturgeon Scaphirhynchus albus in the lower Missouri River are relatively unknown. Therefore, data collected from the Nebraska Game and Parks Commission Pallid Sturgeon Population Assessment Program was synthesized to (i) document the population structure of pallid sturgeon by origin (hatchery‐reared or wild), gender, and reproductive readiness, (ii) document the minimum size and age‐at‐maturity by gender, and (iii) document the fecundity rates of the fish that were successfully spawned in the hatchery. During this 4‐year study (2008–2011), relative abundance for wild and hatchery‐reared pallid sturgeon collected with gill nets did not significantly change whereas relative abundance for wild fish using trot lines declined significantly. The proportion of hatchery‐reared pallid sturgeon increased annually, with the population being composed primarily of hatchery‐reared fish. The proportion of reproductively ready females to non‐reproductively ready females was 1 : 2.0, compared to male ratios at 1 : 0.9. Minimum fork length‐at‐maturity was estimated for females at 788 mm and for males at 798 mm. Minimum age‐at‐maturity for hatchery‐reared released fish was age‐9 for females and age‐7 for males. Highest relative fecundity, based on the ovosomatic index, was 10% with an overall mean of 7%. The number of eggs per ml (egg size) was not correlated with fork length (P = 0.0615) or weight (P = 0.0957). Relative condition factor (Kn) for females was significantly different by reproductive condition (P = 0.0014) and Kn for males did not differ between reproductive conditions (P = 0.2634). Detecting shifts in population characteristics are essential not only to understand population dynamics since hatchery inputs and natural perturbations continue to change the population structure but also to assess species recovery efforts to ensure long‐term species sustainability.     

How immunocontraception can contribute to elephant management in small, enclosed reserves: Munyawana population as a case study

Immunocontraception has been widely used as a management tool to reduce population growth in captive as well as wild populations of various fauna. We model the use of an individual-based rotational immunocontraception plan on a wild elephant, Loxodonta africana, population and quantify the social and reproductive advantages of this method of implementation using adaptive management. The use of immunocontraception on an individual, rotational basis stretches the inter-calving interval for each individual female elephant to a management-determined interval, preventing exposing females to unlimited long-term immunocontraception use (which may have as yet undocumented negative effects). Such rotational immunocontraception can effectively lower population growth rates, age the population, and alter the age structure. Furthermore, such structured intervention can simulate natural process such as predation or episodic catastrophic events (e.g., drought), which regulates calf recruitment within an abnormally structured population. A rotational immunocontraception plan is a feasible and useful elephant population management tool, especially in a small, enclosed conservation area. Such approaches should be considered for other long-lived, social species in enclosed areas where the long-term consequences of consistent contraception may be unknown.     

Reproductive biology of bisexual and all-female populations of chaetodontid fishes from the southern Great Barrier Reef

Synopsis The reproductive biology of three species of chaetodonts from One Tree Reef (southern GBR) is described by analysis of gonads and population structure. Reproductive biology is related to age by a capture-markrecapture study of growth. For Chaetodon rainfordi there was a significant habitat effect. On the reef slope, females outnumbered males by 5.6: 1 and both sexes became reproductively active by the age of 2 years. In the lagoon, no males were found and females were reproductively inactive. For C. plebius, males were essentially missing from all samples and no reproductively active females were found. For Chelmon rostratus (1 :1 sex ratio) some males had mature testes but there was no evidence of female reproductive activity. This species is likely to be gonochoristic, but the reproductive mode of the other two species remains ambiguous.     

Seasonal changes in age class structure and reproductive status of house mice on Marion Island (sub-Antarctic)

Feral house mice on sub-Antarctic Marion Island become reproductively active (males scrotal, females with perforate vaginas or pregnant) at an age >60 days and breed until death, which may occur at more than 13 months. Breeding is strongly seasonal; pregnant or lactating females were found only from October to May. A substantial proportion of mice old enough to breed in one summer overwinters to form a significant component of the breeding population the following summer but it is unlikely that any survive a second winter. The onset of breeding is closely synchronized with increasing day length but occurs about 2 months before mean temperature at the ground surface starts to increase significantly. Cessation of breeding is more closely associated with declining temperatures in late summer. For both males and females, the best correlation between reproductive activity and any of the temperature parameters measured was with average maximum temperature 1 cm above the ground. Competition for macroinvertebrate prey increases sharply in early winter due to high mouse numbers. The breeding season in 1991/1992 and 1992/1993 was at least 2 months longer than in 1979/1980, because the mice started breeding earlier, and stopped breeding later, in 1991/1993. The later cessation of breeding in 1991/1993 was despite the fact that there was a greater competition for macroinvertebrate prey, and that mean air temperatures during the early winter months were lower, than in 1979/1980.     

Seasonal changes in the effects of arginine vasotocin and stretch on Anolis uterine contractions in vitro

We determined the effects of acute stretch on spontaneous and arginine vasotocin (AVT)-driven contractions of the Anolis carolinensis uterus in vitro. Whole uteri from reproductively inactive females (October) were placed in a bath of oxygenated 32 degrees C Anolis "Ringer's." Two initial tensions were utilized, 1.5 g or 15 g, the latter being an estimate of the tension on the wall of a uterine compartment. Uteri were then exposed to either saline or AVT (50 ng/ml), and spontaneous or AVT-driven contractions were recorded for 20 min with the use of a strain gauge and physiograph. A similar experiment was performed on uteri from reproductively active females in the summer (June). Our results indicate that the effects of acute stretch and AVT on uterine contractility were qualitatively similar in summer and fall. That is, AVT induced a tonic contraction; stretch decreased the duration of the tonic contraction; the saline-treated uteri exhibited spontaneous rhythmic contractions; AVT increased the amplitude of the rhythmic contractions, but only at the lower tension; there were no effects of AVT on the timing (contraction interval, duration, rest interval) of the rhythmic contractions; and stretch increased the frequency of the rhythmic contractions. Season greatly influenced the magnitude of these contractile phenomena. Uteri tested during the breeding season exhibited greater distensibility, an increase in the amplitude and duration of the AVT-driven tonic contraction, and an increase in the frequency of both spontaneous and AVT-driven rhythmic contractions because of a decrease in both contraction duration and rest interval.(ABSTRACT TRUNCATED AT 250 WORDS)     

Analysis of female reproductive traits in Angus beef cattle divergently selected for blood serum insulin-like growth factor I concentration

Insulin-like growth factor-I (IGF-I) is an anabolic polypeptide involved in reproductive performance in several species. The objectives of this study were to determine relationships of pregnancy rate, and age of heifers at puberty and at first calving with serum IGF-I concentration in Angus beef cattle. Data were obtained from an ongoing divergent selection experiment for IGF-I concentration involving purebred Angus cows. The IGF-I concentrations measured at Days 28, 42, and 56 of the 140-day postweaning test are abbreviated as IGF28, IGF42, and IGF56, respectively. Pregnancy rate did not differ between high and low IGF-I line females (P=0.95; n=2618), but high line heifers tended to be 4.02+/-2.18 days younger (P=0.07; n=281) at first calving. Residual correlations of age of heifers at first calving (AFC) with IGF-I measurements were not significant. The linear and quadratic terms for regression of AFC on IGF-I concentrations were also non-significant. Contrast analysis showed no difference in age at puberty between the high and low IGF-I line heifers (5.3+/-6.4 days earlier in the high line; P=0.43; n=51). Residual correlations of age of heifers at puberty with IGF28, IGF42, IGF56, and mean IGF-I were -0.30 (P=0.03), -0. 22 (P=0.12), -0.35 (P=0.01), and -0.34 (P=0.01), respectively. The observed relationships between female reproductive traits and IGF-I concentration in Angus beef cattle suggest complex and multiple roles for IGF-I in reproduction.     

Spatio‐temporal variability in reproductive ecology of sand flathead,Platycephalus bassensis,in three Tasmanian inshore habitats: potential implications for management

Temporal and spatial variability in gonad development, duration of spawning period, and size/age at maturity were investigated in sand flathead, Platycephalus bassensis. A 3‐year study (2001–2003) revealed that variation in gonad weight with somatic weight was a function of an interaction between season and study location (Coles Bay, Georges Bay, and Tamar River estuary). Highest gonad weight was recorded in Coles Bay in early summer, which in comparison to Georges Bay, was approximately 50% higher and occurred 3 months later. Tamar River mature individuals were reproductively inactive during the spawning season. The proportion of mature individuals at different stages of maturity differed significantly among the three locations across all times. Coles Bay individuals were reproductively active from October to March, while in Georges Bay females with hydrated oocytes were seen in September but were absent from the population by November. Interannual variation in initiation and duration of spawning activity was evident in each location. Apart from male size at maturity, for both sexes the size and age at maturity (L₅₀) was higher in Georges Bay compared to Coles Bay. The results emphasize the necessity of temporal and spatial management based on population differences in reproductive ecology.     

Selective disturbance by elephants (<Emphasis Type="Italic">Loxodonta africana</Emphasis>) on eight tree species in a West African savannah

African elephants (Loxodonta africana) are ecosystem engineers in African savannahs, but their role in the modifications of the populations of trees by means of their browsing activities has been poorly studied so far in West Africa. We studied the disturbance of elephants to eight selected species of trees in the Nazinga Game Ranch (Burkina Faso), in 54 transects at the end of dry season 2008. We fitted simple models describing the number of dead individuals for each tree species in relation to their initial population density, and in relation to dung-pile density, in the assumption that there should be a positive relationship between elephant density and dung-pile density. Generalized Linear Model analyses and regression analyses showed that the more dense the woodland cover, the least the percentage of damaged plants by elephants. For each plant species, the initial density and density of elephant dung-piles explained a high proportion of the variance in the density of dead individuals. Stochastic models, generated by a purposely created simple computer program written in GW-BASIC programming language, predicted changes in tree and shrub abundance under different assumptions about elephant numbers. The models suggested that elephant browsing may cause considerable change in the selected plant populations, especially with regard to such species as Acacia gourmaensis, Vitellaria paradoxa and Maytenus senegalensis. These changes may possibly increase the meat harvest from controlled hunting activities, thus improving the income for surrounding communities.