Group Release of Sanctuary Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) in the Haut Niger National Park,Guinea, West Africa: Ranging Patterns and Lessons So Far

The release of wild or captive-bred mammals within their historical ranges typically aims to reestablish populations in areas where they have become extinct or extirpated, to reinforce natural populations, or to resolve human–wildlife conflicts. Such programs, which also typically in parallel help foster the protection of the release site, concern a wide range of endangered mammalian species, including our closest living relatives: chimpanzees. In June 2008, the Chimpanzee Conservation Center (CCC), which is located in the High Niger National Park (HNNP) in Guinea, released a group of 12 chimpanzees (Pan troglodytes verus) comprised of 6 females and 6 males (8–20 yr old). The selected release site lies 32 km from the sanctuary in the Mafou, a core area of HNNP where wild chimpanzees are also known to occur. The purpose of this release was therefore to reinforce the natural chimpanzee population within the Mafou core area and to promote the protection of the HNNP. Nearly 2 yr postrelease, 9 chimpanzees still remain free-living. Two thirds of the release chimpanzees were equipped with VHF-GPS store-on-board tracking collars. We used data from retrieved collars to explore the release chimpanzees’ habitat use, individual day range, and core area use (50% and 80%) during the first year of the release. Males traveled significantly further than females. Although minimum day range did not differ between the sexes or vary seasonally, some release males were active for longer during the day than the females. Males also ranged over larger areas and used a wider network of core areas than the females. Habitat use was similar to that recorded in wild chimpanzees in the HNNP. As of September 2010, 2 males and 3 females form a group at the release site. Two of these females gave birth to healthy offspring respectively 16 and 20 mo postrelease. Another female successfully immigrated into a wild chimpanzee community. We suggest that the success of this chimpanzee release can be attributed to the CCC’s lengthy rehabilitation process and the savanna-mosaic habitat of the HNNP. This release demonstrates that under special socioecological circumstances, the release of wild-born adult chimpanzees of both sexes is a viable strategy, which can also function as an effective conservation tool.     

High Population Density of <Emphasis Type="Italic">Pan troglodytes verus</Emphasis> in the Haut Niger National Park,Republic of Guinea: Implications for Local and Regional Conservation

The Republic of Guinea harbours the largest population of the endangered western chimpanzees Pan troglodytes verus, and the conservation of this population is regarded as a regional priority. Chimpanzees occur in 3 of the 5 protected areas in Guinea but their conservation status is unknown. We conducted a chimpanzee census in 2001–2002 in the Haut Niger National Park (HNNP), the largest protected area in the country. We counted nests along a total of 103.83 km of transects in the Mafou forest, 1 of the 2 core areas of the park. We recorded a total of 823 nests, of which 38.1% were located in gallery forests, although this habitat type covers only 4.2% of the study site. Using a site-specific mean nest decay rate of 194 days, we estimated the mean density of chimpanzees in the study area at 0.87 weaned individuals/km2, the highest density recorded in any West African protected area. A survey conducted in 2008–2009 by other authors in the Mafou forest suggests that the chimpanzee population remained stable over the last 6 yr. Given the significance of the chimpanzee population of the HNNP at the national and international levels, we provide recommendations for its conservation over the long term. Our results highlight the importance of gallery forests as key habitat for chimpanzees in the savannah biome. We therefore recommend that more attention be paid to the conservation of this habitat in land management and conservation planning schemes. In particular, we recommend setting aside gallery forests as ecological corridors between nearby protected areas to maintain exchange between distant chimpanzee populations.     

Influence of chimpanzee predation on the red colobus population at Ngogo, Kibale National Park, Uganda

Frequent hunting of red colobus monkeys (Procolobus rufomitratus) takes place at all long-term chimpanzee (Pan troglodytes) study sites where both species are present. Red colobus are the most commonly selected prey of chimpanzees even when other monkey species are more abundant. In particular, the chimpanzee community at Ngogo, Kibale National Park, Uganda, preys heavily on red colobus monkeys: the chimpanzee hunting success rate is extremely high, and chimpanzees kill many individuals per successful hunt. Census data had suggested that the red colobus population is declining and that predation by chimpanzees may be contributing to this decline. In this paper, I address the impact of hunting on the red colobus population at Ngogo. To test the hypothesis that chimpanzee hunting is sustainable, I am using demographic data collected on red colobus monkeys over a period of 3 years, as well as fecundity and mortality data from previous studies of this species. I apply matrix models and vortex analyses using a sensitivity analysis approach to project future population development. Results show that current rates of hunting are not sustainable, but that chimpanzees are neither more “noble”, nor more “savage” than humans are, but that they also hunt to ensure maximum benefit without regard for the consequences for the prey population.     

The type locality of <Emphasis Type="Italic">Pan troglodytes vellerosus</Emphasis> (Gray, 1862), and implications for the nomenclature of West African chimpanzees

We show that, in 1862, Richard Burton collected the type specimen of Pan troglodytes vellerosus not on Mount Cameroon, as has been generally assumed, but in Gabon. Therefore, P. t. vellerosus is not the correct name for the chimpanzee population of western Cameroon and southern Nigeria, if that population is taxonomically distinct. As First Reviser, we choose the name Pan troglodytes ellioti for this population of chimpanzees, based on Anthropopithecus ellioti named by Matschie [Matschie P (1914) Neue Affen aus Mittelafrika. Sitzungsber Ges Naturforsch Freunde Berlin 1914:323–342] from a specimen in the Humboldt Museum, Berlin, collected in Bascho (=Basho), Cameroon, and given to the museum in 1905.     

Historical demography of a wild lemur population (Propithecus verreauxi) in southwest Madagascar

The human colonization of Madagascar is associated with the extinction of numerous lemur species. However, the degree to which humans have negatively influenced the historical population dynamics of extant lemur species is not well understood. This study employs genetic and demographic analyses to estimate demographic parameters relating to the historical population dynamics of a wild lemur population, Verreaux’s sifaka (Propithecus verreauxi). The genetic analyses are used to determine whether this population experienced a historically recent (i.e., within the last 2000 years) population bottleneck, as well as to estimate the historical population growth rate and the timing of any changes in population size in the past. In addition, a retrospective demographic analysis is used to determine sources of variation and covariation in the sifaka life cycle and how variation in life-cycle transitions contributes to variation in population growth rate. The genetic analyses indicate that the sifaka population did not experience a recent population bottleneck; however, the historical population growth rate was negative, indicating that the ancestral population size was much larger than the current size. The timing of the ancestral population decline has a point estimate of 2300 years ago, but with large credible intervals: 3611–1736 years ago. This point estimate corresponds with the first evidence for human arrival to Madagascar. Climatic variation has also likely influenced past (and current) population dynamics due to stochastic annual rainfall patterns and climatic desiccation, the latter of which began in southwestern Madagascar around 4000 years ago. Variation in the survival of 2-year-old animals as well as large adult females makes the largest contribution to variation in population growth rate. In the absence of more explicit models pertaining to historical population dynamics, it is difficult to attribute the negative population growth rate of this species solely to a single factor (e.g., hunting, habitat destruction).     

Hunting and Occasional Consumption of Prey Items by Chimpanzees at the Primate Foundation of Arizona

Researchers have documented hunting by wild chimpanzees (Pan troglodytes) for >30 yr. However, no one had reported the behavior in captive chimpanzees. The Primate Foundation of Arizona (PFA) is a biomedical research facility, and though every effort is made to ensure that rodents and other indigenous animals do not enter cages, an occasional small mammal or bird enters the outside cage area. We document 18 known successful hunts over 2.5 yr by ≥8 captive chimpanzees housed at PFA. The most common prey were rock squirrels (Cittallus variegatus; n = 8) and birds (n = 7). The most common killing method was a bite to the head (n = 9), indicating that the deaths were deliberate and not accidental. Two adult males, 1 wild- and 1 captive-born, together accounted for 11 of the 18 hunting episodes. The chimpanzees generally captured and killed the prey in their outdoor enclosures. The results have implications for the surveillance of outdoor areas in other facilities and the potential for disease or introduction of parasitic vectors, and are also an interesting comparison to wild chimpanzee behavior.     

Sustainability of culture-driven population dynamics

We consider models of the interactions between human population dynamics and cultural evolution, asking whether they predict sustainable or unsustainable patterns of growth. Phenomenological models predict either unsustainable population growth or stabilization in the near future. The latter prediction, however, is based on extrapolation of current demographic trends and does not take into account causal processes of demographic and cultural dynamics. Most existing causal models assume (or derive from simplified models of the economy) a positive feedback between cultural evolution and demographic growth, and predict unlimited growth in both culture and population. We augment these models taking into account that: (1) cultural transmission is not perfect, i.e., culture can be lost; (2) culture does not always promote population growth. We show that taking these factors into account can cause radically different model behavior, such as population extinction rather than stability, and extinction rather than growth. We conclude that all models agree that a population capable of maintaining a large amount of culture, including a powerful technology, runs a high risk of being unsustainable. We suggest that future work must address more explicitly both the dynamics of resource consumption and the cultural evolution of beliefs implicated in reproductive behavior (e.g., ideas about the preferred family size) and in resource use (e.g., environmentalist stances).     

Changes in Demographic Parameters of <Emphasis Type="Italic">Macaca fuscata</Emphasis> at Takasakiyama in Relation to Decrease of Provisioned Foods

Though researchers have studied population dynamics extensively in many free-ranging primate populations, information on the relationship between food abundance and demographic parameters based on long-term data is sparse. We examined changes in demographic parameters in relation to decreased provisioning of foods based on data gathered for >50 yr in a provisioned, free-ranging Japanese macaque (Macaca fuscata) population at Takasakiyama, Oita Prefecture, in the southern region of Japan. At Takasakiyama, population size increased linearly because of heavy provisioning in the 1950s and 1960s. The provisioning of food to macaques decreased from 1965. We examined changes in the physique index of full adult females, primiparous age, birth percentage, infant mortality rate, population density, and annual population growth rate relative to the decrease of provisioned foods between the 1970s and 1990s. As a result of the drastic decrease in the amount of foods provisioned, the physique index of full adult females decreased, primiparous age increased, birth percentage decreased, and infant mortality rate increased, such that annual population growth rate diminished substantially. Ours is the first study to demonstrate quantitatively the relationship between provisioned food abundance and demographic parameters based on long-term data.     

Skeletal evidence of probable treponemal infection in free-ranging African apes

The prevalence and patterning of inflammatory lesions of the skeleton were investigated in samples of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) curated at the Powell-Cotton Museum, Birchington, UK. One hundred and two chimpanzees (42 adults and 60 subadults) and 126 gorillas (50 adults and 76 subadults) comprise the samples. Twenty per cent of chimpanzees and 14% of gorillas were affected with a disseminated inflammatory skeletal condition caused by infection. The lesions appear to have originated as localized patches of new bone deposition on the surface of long bones and to have progressed to infection of the bone cortex and marrow. Although female prevalence of involvement exceeds that of males in both species, the differences are not statistically significant. The age distribution of affected animals indicates that the disease began in some animals as early as 2 yr of age. Given the skeletal and demographic prevalence and patterning of the lesions as well as the ecology and behavior of these animals, the most likely diagnosis of the condition is a yaws-like treponemal infection.     

Female Coalitions Against Male Aggression in Wild Chimpanzees of the Budongo Forest

In the wild, female chimpanzees (Pan troglodytes) are subject to male aggression that at times can be prolonged or particularly violent. There are no reports of cooperative retaliation to such aggression, a strategy observed in the congeneric Pan paniscus, from the wild despite >4 decades of detailed behavioral study across a number of populations and its occurrence among captive female chimpanzees. If the reports from captivity represent an inherent capacity, then the absence of similar reports from wild populations suggests that females may be able to form coalitions only under appropriate ecological and demographic conditions. During a study of male and female aggressive interactions among chimpanzees of the Budongo Forest, Uganda, wild adult female chimpanzees sometimes formed coalitions with one another to retaliate against male aggression. This may be possible because these females tend to be more gregarious than in other populations of East African chimpanzees, as other studies of the same population have suggested; the extent and variation of female chimpanzee social strategies may, therefore, need reconsideration. Further, my observations strengthen the argument that at least some of the differences between chimpanzees and bonobos may be more of degree than of kind.     

Comparing survey and multiple recruitment–mortality models to assess growth rates and population projections

Estimation of population trends and demographic parameters is important to our understanding of fundamental ecology and species management, yet these data are often difficult to obtain without the use of data from population surveys or marking animals. The northeastern Minnesota moose (Alces alces Linnaeus, 1758) population declined 58% during 2006–2017, yet aerial surveys indicated stability during 2012–2017. In response to the decline, the Minnesota Department of Natural Resources (MNDNR) initiated studies of adult and calf survival to better understand cause‐specific mortality, calf recruitment, and factors influencing the population trajectory. We estimated population growth rate (λ) using adult survival and calf recruitment data from demographic studies and the recruitment–mortality (R‐M) Equation and compared these estimates to those calculated using data from aerial surveys. We then projected population dynamics 50 years using each resulting λ and used a stochastic model to project population dynamics 30 years using data from the MNDNR's studies. Calculations of λ derived from 2012 to 2017 survey data, and the R‐M Equation indicated growth (1.02 ± 0.16 [SE] and 1.01 ± 0.04, respectively). However, the stochastic model indicated a decline in the population over 30 years (λ = 0.91 ± 0.004; 2014–2044). The R‐M Equation has utility for estimating λ, and the supporting information from demographic collaring studies also helps to better address management questions. Furthermore, estimates of λ calculated using collaring data were more certain and reflective of current conditions. Long‐term monitoring using collars would better inform population performance predictions and demographic responses to environmental variability.     

Recovery dynamics and viability of the white-tailed eagle (<Emphasis Type="Italic">Haliaeetus albicilla</Emphasis>) in Germany

Understanding the effect of protection measures on recovery of endangered populations is crucial for assessing the efficiency of management plans. Following the ban of DDT, PCB and other detrimental chemicals in the 1970s, the German white-tailed eagle population recovered rapidly. Using nest monitoring data, dead recovery data and population dynamics models, we examined both short and long-term viabilities of the population. Between 1991 and 2005 the proportion of breeding pairs with an active nest among territorial pairs was α = 0.88 ± 0.007. Annual breeding success was γ = 0.62 ± 0.014. Mean fledged brood size was b = 1.50 ± 0.010 fledged/successful breeding pair. The analysis of dead recovery data of 1,273 individuals ringed as nestling between 1991 and 2006 allowed survival rates to be estimated. We needed to distinguish between “young” (5 years old) and “old” (older than 5 years) adults. Furthermore the annual survival rate was higher for juvenile and immature birds (0.921 ± 0.024) than for adults (s _“young” = 0.760 ± 0.097 and s _“old” = 0.875 ± 0.079), a pattern presumably explained by territorial fights. Finally, an age structured model was used to examine the joint effect of population regulation, environmental and demographic stochasticities. All results indicated that only pessimistic scenarios could lead to substantial extinction probabilities within 100 years. We conclude that management measures were successful in removing the main extinction factors of the population and that future conservation efforts need to focus on the remaining ultimate extinction causes.     

Responses of wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) toward seismic aftershocks in the Mahale Mountains National Park,Tanzania

This is the first report documenting the responses of wild chimpanzees (Pan troglodytes schweinfurthii) to seismic activities. During our long-term fieldwork in Mahale Mountains National Park, Tanzania, a high-intensity earthquake with a Richter magnitude of 6.8 occurred at 15:19 hours local time on 5 December 2005. During the main tremor, the chimpanzees displayed the “wraa” call, “scream,” and “pant bark” or “bark” vocalizations. Many mild aftershocks followed the main tremor, and the wild chimpanzees displayed a variety of responses to these. In several cases, they climbed trees or stopped activities such as grooming, moving, and feeding. These responses are similar to those previously reported in nonhuman primates. During the observations, a unique behavior, one never reported before was exhibited by a female chimpanzee. She placed her right palm on the ground giving the impression she was inspecting the trembling of the ground.     

Reproductive Parameters of Female<Emphasis Type="Italic">Pan paniscus</Emphasis> and <Emphasis Type="Italic">P. troglodytes</Emphasis>: Quality versus Quantity

We investigated intra- and interspecific differences in life history and reproductive parameters in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We compare the parameters of wild and captive females in order to shed light on the influence of habitat or specific differences or both on reproduction. We present new and additional information on reproductive parameters from captive bonobos and chimpanzees. Captive chimpanzees birth more live offspring and have a shorter interbirth interval, but experience higher infant mortality than captive bonobos. Although captive bonobo females tend to start reproduction at a younger age than chimpanzees, this is effectively only so for wild-born females of both species. Ultimately both species reach the same rate of production of offspring surviving to 5 yr. These results contrast with data from the wild. Wild bonobos tend to have higher reproductive success, a higher fertility rate and a shorter interbirth interval than wild chimpanzees. Reproduction is similar for wild and captive bonobos, which suggests that they are producing at their maximum under both conditions. Overall captive chimpanzees perform better than their wild conspecifics, probably because of lower feeding competition. Infant survival is the only specific difference not affected by captivity. Bonobo infants survive better, which suggests that chimpanzee infants are more at risk. We argue that the interspecific variation in reproductive parameters in captivity is related to the different influence of captivity on reproduction and different pressures of external sources of infant and juvenile mortality.     

Density and distribution of western chimpanzees around a bauxite deposit in the Boé Sector,Guinea‐Bissau

The Boé sector in southeast Guinea‐Bissau harbors a population of western chimpanzees (Pan troglodytes verus) that inhabits a mosaic of forest and savanna. The Boé sector contains a substantial bauxite deposit in a region called Ronde Hill, and there are plans for the construction of a mine, which may endanger the chimpanzee population. In 1‐week survey in May 2013, we used the standing crop nest counts method to obtain the number of chimpanzee nests and from that estimate the density and abundance of chimpanzees. We carried out five 1‐km line transects that covered the bauxite deposit and surrounding valleys. We used density surface modeling to analyze habitat preferences, then predicted chimpanzee nest density and distribution based on environmental variables. We found the projected location of the mine partially coincides with an area of high predicted abundances of chimpanzee nests and is surrounded by highly suitable areas for chimpanzees (northeast and southwest). We conclude the mine could have significant direct and indirect effects on this population of chimpanzees whose impacts must be carefully considered and properly mitigated if the mine is built.     

Do chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) use cleavers and anvils to fracture <Emphasis Type="Italic">Treculia africana</Emphasis> fruits? Preliminary data on a new form of percussive technology

Wild chimpanzees (Pan troglodytes) are renowned for their use of tools in activities ranging from foraging to social interactions. Different populations across Africa vary in their tool use repertoires, giving rise to cultural variation. We report a new type of percussive technology in food processing by chimpanzees in the Nimba Mountains, Guinea: Treculia fracturing. Chimpanzees appear to use stone and wooden “cleavers” as tools, as well as stone outcrop “anvils” as substrate to fracture the large and fibrous fruits of Treculia africana, a rare but prized food source. This newly described form of percussive technology is distinctive, as the apparent aim is not to extract an embedded food item, as is the case in nut cracking, baobab smashing, or pestle pounding, but rather to reduce a large food item to manageably sized pieces. Furthermore, these preliminary data provide the first evidence of chimpanzees using two types of percussive technology for the same purpose.     

Menstrual cycles continue into advanced old age in the common chimpanzee (Pan troglodytes)

A long postreproductive lifespan may distinguish women from all other female primates. A long-held consensus among reproductive scientists has been that our closest living relative, the chimpanzee (Pan troglodytes), experiences menstrual cycles until death. However, a recent study of biannual assessments of gonadotropins, but lacking observations of menstruation, concluded that menopause occurs in chimpanzees between 35 and 40 yr of age. A separate report, but on wild chimpanzees, documented fertility through the 40-44 age range in all populations studied. These contradictory reports pose questions about differences between wild and captive populations and about assessments of menopause. The present study revisits this controversy by analyzing longitudinal records of anogenital swelling and menstruation in 89 female chimpanzees aged 6 to 59 yr (n = 2386 records on cycle length), monitored for most of their adult lives at the Yerkes National Primate Research Center. Twenty of these chimpanzees were observed past 39 yr of age; all 20 displayed menstrual cycles beyond this age, as confirmed by at least two observations of menses about 35 days apart. Three of these were older than 50 yr and still displayed menstrual cycles. Only the oldest female appeared menopausal, with cycles of anogenital swelling ceasing 2 yr prior to her death at age 59. Random-effects statistical modeling reveals a slight decrease in cycle length until 20 yr of age and a slight lengthening thereafter. Mean cycle length across the lifespan is 35.4 days. Our findings, based upon actual observations of menstrual cycles, suggest that menopause in the chimpanzee is rare, occurring near the end of the lifespan.     

International migrations toward the city of Parma (Northern Italy): Evolution during a decade (1989–2000)

The population of Parma can be considered a paradigm in order to describe the rapid demographic transformations characterizing Northern Italy communities, in which immigration flows play undoubtedly an important role. The present study uses data from the ‘Osservatorio Biodemografico’ (Biodemographic Observatory), a data bank set up in 1989 with the aim to analyze longitudinally the dynamics of that urban population. This data bank traces individual and family life-histories since 12 years, updating them year by year. The presence of migrants has significantly risen in this last decade: at the beginning of 1989, foreign people living in Parma were 1.341 (751 men and 590 women; 0.78% of the resident population); in the year 2000 foreign residents were 6.491 (3.583 men and 2.908 women; 3.9% of the resident population). Immigrants can be analyzed since their arrival in the city (or, better, since their inclusion in the Population Register) according to their characteristics and modalities of arrival, duration of permanence in Parma, co-residential patterns, dynamics of family rejoining. Numerous profiles, characterized by different and very complex individual and familial trajectories, have been described. Actually, despite of a common region of origin, migratory trajectories differ greatly one each other, in consequence of complex dynamics involving, sex age and kinship. We wish to thank the Fyssen Foundation, thanks to which the post doctoral studies of Dott.ssa Pellegrino are carried out.     

Substantial molecular variation and low genetic structure in Kenya’s black rhinoceros: implications for conservation

Kenya’s black rhinoceros population declined by more than 98% from 20,000 individuals in the 1970s to around 400 individuals in 1990 due to the effects of poaching, at which time the surviving individuals were isolated in a series of demographically inviable subpopulations. An initial management exercise translocated the survivors into four high security sanctuaries to control poaching and enhance breeding, and this measure successfully arrested the decline. Subsequently, new sanctuaries were established and the metapopulation size reached 650 animals by 2008. However, translocations and the current management strategy that partitions the metapopulation into ‘montane’ and ‘lowland’ rhinoceros may have substantial consequences at the population level and their impact on population genetic diversity has not been investigated. In this study, 12 of the 16 extant subpopulations were analysed using 408 bp of mitochondrial control region sequence (n = 170) and nine microsatellite loci (n = 145). Both markers detected moderate to high genetic diversity (h = 0.78 ± 0.027, n = 170; H_O = 0.70 ± 0.087, n = 145) consistent with previous studies on Diceros bicornis michaeli. However, mtDNA and nDNA diversity varied substantially between subpopulations. The results suggest that the Masai Mara is more differentiated, inbred and isolated than other subpopulations. It also suggests that there are neither distinct montane and lowland groups nor other detectable historical barriers to gene flow. Instead the large majority of genetic diversity was partitioned at the level of individuals; highlighting the need to conserve as many individuals as possible. Future translocations should consider the genetic profile of individuals and the demographic history of both the donor and recipient subpopulations.     

Using recruitment age, territorial fidelity and dispersal as decisive tools in the conservation and management of peregrine falcon ( Falco peregrinus ) populations: the case of a healthy population in Northern Spain

Methods of evaluating population trends have recently received particular attention because of perceived declines in several species during the twentieth century. We have studied demographic traits of the peregrine falcon (Falco peregrinus) population in Bizkaia (Northern Spain) for 11 years. This species suffered a severe decline in the 1950s and 1960s and started to recover in the 1980s and 1990s, although the recovery trends differed between areas. In our study area the peregrine falcon density is one of the highest found in Spain. The frequency of juvenile breeders was 2.0%, the mean age at first breeding being 3.7 calendar years for males and 4.0 cy for females. The territorial fidelity was at least 3.4 years for males and 3.7 years for females. Females dispersed on average 80.5 km and males 51.8 km during their pre-breeding movements, whilst distance between birthplace and breeding territories was on average 108.5 km for females and 64.5 km for males. We studied the relationships between adult turnover, recruitment age, territorial fidelity and dispersal in a healthy population in order to establish population dynamics. A combination of these parameters, not just age at first breeding, could be used as a potential early warning signal indicating future changes, prompting their consequences to be evaluated. This approach could lead to the reclassification of the large-scale health of a population. Its utilization would allow resources to be directed into helping “ailing” subpopulations, detecting causes of decline, and developing adequate recovery strategies.