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Species and sex identification of western lowland gorillas (Gorilla gorilla gorilla), eastern lowland gorillas (Gorilla beringei graueri) and humans.

Methods for the identification of the sex and species of individuals from samples non-invasively taken from humans and gorillas were established. Amplification of a segment of amelogenin (AMG), which is an X-Y homologous gene, using two pairs of primers from human AMG, revealed both X- and Y-specific bands. The possibility of sex identification was examined by typing the AMG gene using hair and fecal samples from captive western lowland gorillas (Gorilla gorilla gorilla) in Japan and hair samples from wild eastern lowland gorillas (Gorilla beringei graueri) in the Kahuzi-Biega National Park, Democratic Republic of Congo, which were sexed by direct observation. Species-specific bands of AMG in gorillas and humans were identified by restriction fragment length polymorphisms analysis. These tests could be used for sexing unidentified individuals of wild western and eastern lowland gorillas, and screening contamination of human DNA from non-invasively acquired samples.

Amelogenin↗

Social structure and life-history patterns in western gorillas (Gorilla gorilla gorilla).

Life-history traits and ecological conditions have an important influence on primate social systems. Most of what we know about the life-history patterns and social structure of gorillas comes from studies of eastern gorillas (Gorilla beringei sp.), which live under dramatically different ecological conditions compared to western gorillas (Gorilla gorilla sp.). In this paper we present new data on western gorilla social structure and life histories from four study sites, and make comparisons with eastern gorilla populations. Data were obtained from two study sites with gorilla groups undergoing the habituation process (Lossi, Democratic Republic of Congo and Bai Hokou, Central African Republic) and two "bai" studies (Maya Nord and Mbeli Bai, Republic of Congo). The size and structure of these groups were similar to those seen in eastern gorillas. However, differences in the occurrence of various group transitions (group formations, changes between one-male and multimale composition, and group disintegrations) exist, and western gorillas notably exhibit much higher rates of male emigration and correspondingly fewer multimale groups compared to mountain gorillas. Certain phenomena have been observed only rarely, including predation by leopards. The preliminary data show no significant differences in birth rates between western gorillas and mountain gorillas. The ecological variability across gorilla habitats likely explains the flexibility in the social system of gorillas, but we need more information on the social relationships and ecology of western gorillas to elucidate the causes for the similarities and differences between western and eastern gorillas on the levels of individuals, social groups, and population dynamics.

Africa, Central↗

Relative growth, ontogeny, and sexual dimorphism in gorilla (Gorilla gorilla gorilla and G. g. beringei): evolutionary and ecological considerations.

Gorillas are the largest and among the most sexually dimorphic of all extant primates. While gorillas have been incorporated in broad-level comparisons among large-bodied hominoids or in studies of the African apes, comparisons between gorilla subspecies have been rare. During the past decade, however, behavioral, morphological, and molecular data from a number of studies have indicated that the western lowland (Gorilla gorilla gorilla) and eastern mountain (Gorilla gorilla beringei) subspecies differ to a greater extent than has been previously believed. In this study I compare patterns of relative growth of the postcranial skeleton to evaluate whether differences between subspecies result from the differential extension of common patterns of relative growth. In addition, patterns of ontogeny and sexual dimorphism are also examined. Linear skeletal dimensions and skeletal weight were obtained for ontogenetic series of male and female G.g. gorilla (n = 315) and G.g. beringei (n = 38). Bivariate and multivariate methods of analysis were used to test for differences in patterns of relative growth, ontogeny, and sexual dimorphism between sexes of each subspecies and in same-sex comparisons between subspecies. Results indicate males and females of both subspecies are ontogenetically scaled for postcranial proportions and that females undergo an earlier skeletal growth spurt compared to males. However, results also indicate that the onset of the female growth spurt occurs at different dental stages in lowland and mountain gorillas and that mountain gorillas may be characterized by higher rates of growth. Finally, data demonstrate lowland and mountain gorilla females do not differ significantly in adult body size, but mountain gorilla males are significantly larger than lowland gorilla males, suggesting mountain gorillas are characterized by a higher degree of sexual dimorphism in body size. Thus, although lowland and mountain gorillas do not appear to have evolved novel adaptations of the postcranium which correlate with differences in locomotor behavior, the present investigation establishes subspecies differences in ontogeny and sexual dimorphism which may be linked with ecological variation. Specifically, these findings are evaluated in the context of risk aversion models which predict higher growth rates and increased levels of sexual dimorphism in extreme folivores.

Animals↗

Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla).

The geographical distribution of genetic variation within western lowland gorillas (Gorilla gorilla gorilla) was examined to clarify the population genetic structure and recent evolutionary history of this group. DNA was amplified from shed hair collected from sites across the range of the three traditionally recognized gorilla subspecies: western lowland (G. g. gorilla), eastern lowland (G. g. graueri) and mountain (G. g. beringei) gorillas. Nucleotide sequence variation was examined in the first hypervariable domain of the mitochondrial control region and was much higher in western lowland gorillas than in either of the other two subspecies. In addition to recapitulating the major evolutionary split between eastern and western lowland gorillas, phylogenetic analysis indicates a phylogeographical division within western lowland gorillas, one haplogroup comprising gorilla populations from eastern Nigeria through to southeast Cameroon and a second comprising all other western lowland gorillas. Within this second haplogroup, haplotypes appear to be partitioned geographically into three subgroups: (i) Equatorial Guinea, (ii) Central African Republic, and (iii) Gabon and adjacent Congo. There is also evidence of limited haplotype admixture in northeastern Gabon and southeast Cameroon. The phylogeographical patterns are broadly consistent with those predicted by current Pleistocene refuge hypotheses for the region and suggest that historical events have played an important role in shaping the population structure of this subspecies.

Africa South of the Sahara↗

Flow cytometric sorting of fresh and frozen-thawed spermatozoa in the western lowland gorilla (Gorilla gorilla gorilla).

We adapted flow cytometry technology for high-purity sorting of X chromosome-bearing spermatozoa in the western lowland gorilla (Gorilla gorilla gorilla). Our objectives were to develop methodologies for liquid storage of semen prior to sorting, sorting of liquid-stored and frozen-thawed spermatozoa, and assessment of sorting accuracy. In study 1, the in vitro sperm characteristics of gorilla ejaculates from one male were unchanged (P > 0.05) after 8 hr of liquid storage at 15 degrees C in a non-egg yolk diluent (HEPES-buffered modified Tyrode's medium). In study 2, we examined the efficacy of sorting fresh and frozen-thawed spermatozoa using human spermatozoa as a model for gorilla spermatozoa. Ejaculates from one male were split into fresh and frozen aliquots. X-enriched samples derived from both fresh and frozen-thawed human semen were of high purity, as determined by fluorescence in situ hybridization (FISH; 90.7%+/-2.3%, overall), and contained a high proportion of morphologically normal spermatozoa (86.0%+/-1.0%, overall). In study 3, we processed liquid-stored semen from two gorillas for sorting using a modification of methods for human spermatozoa. The sort rate for enrichment of X-bearing spermatozoa was 7.3+/-2.5 spermatozoa per second. The X-enriched samples were of high purity (single-sperm PCR: 83.7%) and normal morphology (79.0%+/-3.9%). In study 4 we examined frozen-thawed gorilla semen, and the sort rate (8.3+/-2.9 X-bearing sperm/sec), purity (89.7%), and normal morphology (81.4%+/-3.4%) were comparable to those of liquid-stored semen. Depending on the male and the type of sample used (fresh or frozen-thawed), 0.8-2.2% of gorilla spermatozoa in the processed ejaculate were present in the X-enriched sample. These results demonstrate that fresh or frozen-thawed gorilla spermatozoa can be flow cytometrically sorted into samples enriched for X-bearing spermatozoa.

Analysis of Variance↗

Management of reactive arthritis in a juvenile gorilla (Gorilla gorilla gorilla) with long-term sulfasalazine therapy.

A juvenile western lowland gorilla (Gorilla gorilla gorilla) experienced recurrent fever, lethargy, diarrhea, and/or arthritis starting at age 6 mo. During an episode at age 15 mo, Shigella sp. was isolated from diarrheic feces. At age 41 mo, reactive arthritis was diagnosed. In addition, the gorilla's growth was retarded. All arthritic attacks were managed symptomatically prior to age 4 yr, at which time a severe episode precipitated the implementation of therapy with sulfasalazine, an arthritis suppressive medication. Examination 27 mo later revealed cessation of progressive joint pathology although the animal exhibited decreased range of motion in most joints. The gorilla has been on sulfasalazine therapy for 4 yr without lameness. Growth has resumed, and there has been no radiographic evidence of progressive joint degeneration. Immunogenetic analysis of whole blood obtained at age 68 mo identified the gorilla major histocompatibility class I allele, Gogo-B*0101, which has limited nucleotide sequence similarity to HLA-B27, an allele associated with postinfection reactive arthritis in humans. Sulfasalazine therapy effectively managed reactive arthritis in this gorilla and should be considered for similarly frequently affected animals. Juvenile gorillas, in populations with a history of clinical shigellosis and/or postdiarrhea arthritis, may benefit from prophylactic sulfasalazine therapy after episodes of bacterial enterocolitis. Sulfasalazine therapy should be considered in all gorillas, juvenile and adult, experiencing confirmed Shigella sp.-associated enterocolitis.

Animals↗

Single-trial learning of "what" and "who" information in a gorilla (Gorilla gorilla gorilla): implications for episodic memory.

Single-trial learning and long-term memory of "what" and "who" information were examined in an adult gorilla (Gorilla gorilla gorilla). We presented the gorilla with a to-be-remembered food item at the time of study. In Experiment 1, following a retention interval of either approximately 7 min or 24 h, the gorilla responded with one of five cards, each corresponding to a particular food. The gorilla was accurate on 70% of the short retention-interval trials and on 82% of the long retention-interval trials. In Experiment 2, the food stimulus was provided by one of two experimenters, each of whom was represented by a card. The gorilla identified the food (55% of the time) and the experimenter (82% of the time) on the short retention-interval trials. On the long retention-interval trials, the gorilla was accurate for the food (73%) and for the person (87%). The results are interpreted in light of theories of episodic memory.

Animals↗

Event memory and misinformation effects in a gorilla (Gorilla gorilla gorilla).

Event memory and misinformation effects were examined in an adult male gorilla ( Gorilla gorilla gorilla). The gorilla witnessed a series of unique events, involving a familiar person engaging in a novel behavior (experiment 1), a novel person engaging in a novel behavior (experiment 2), or the presentation of a novel object (experiment 3). Following a 5- to 10-min retention interval, a tester gave the gorilla three photographs mounted on wooden cards: a photograph depicting the correct person or object and two distractor photographs drawn from the same class. The gorilla responded by returning a photograph. If correct, he was reinforced with food. Across three experiments, the gorilla performed significantly above chance at recognizing the target photograph. In experiment 4, the gorilla showed at-chance performance when the event was followed by misinformation (a class-consistent, but incorrect photograph), but significantly above-chance performance when no misinformation occurred (either correct photograph or no photograph). Although the familiarity can account for these data, they are also consistent with an episodic-memory interpretation.

Animals↗

Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task.

Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.

Animals↗

Hand preference for food processing in wild western lowland gorillas (Gorilla gorilla gorilla).

Western lowland gorillas (Gorilla gorilla gorilla) have not previously been represented in studies of laterality in wild great apes. The discovery of swampy clearings frequented by gorillas in northern Congo has provided the first opportunity to redress this imbalance. Hand preference data are presented from 33 gorillas in seated and standing postures, covering the procurement and processing of 2 to 4 plant species. Levels of hand preference exhibited were low. When data from all postures and plant species were pooled, 33% of gorillas showed hand preferences in excess of chance. In the standing posture, more gorillas exhibited significant left-hand preferences than right, but an overall population-level bias was not evident.

Animals↗

Survey of gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes troglodytes) in Southwestern Cameroon.

A study on the distribution and population density of the western lowland gorilla (Gorilla g. gorilla) and the central chimpanzee (Pan t. troglodytes) was undertaken between December 1997 and August 2000 in the Campo and Ma'an Forests in southwestern Cameroon. The aim of this survey was to estimate the densities of the apes in different parts of the area, to assess the importance of the region for the conservation of these endangered species and to determine the influence of human activities such as logging and hunting. The survey was based on night nest counts on a total of 665.5 km of line transects. The overall density in the Campo Forest was estimated at 0.2 gorillas/km(2) and at 0.63-0.78 chimpanzees/km(2). The overall density of chimpanzees in the Ma'an Forest was estimated at 0.8-1 individuals/km(2). Gorilla density in this area was too low to allow an estimation. The highest gorilla nest density was found in secondary forest. The gorilla density in unlogged forest was significantly lower. Chimpanzees showed a clear preference for less disturbed areas. In unlogged forest, old secondary forests (logging more than 23 years ago) and areas of recent logging with large remaining patches of primary forest, significantly higher densities were calculated than inside the more heavily exploited logging concession. In areas with both logging and high hunting pressure both species were rare or even absent. The Campo Ma'an area is considered a very important area for the conservation of gorillas and chimpanzees. Conservation measures are urgently required to reduce the impact of logging and hunting. The creation of the Campo Ma'an National Park in January 2000 is an important measure to preserve the unique biodiversity in this so far hardly protected area.

Animals↗

Food transfers in immature wild western lowland gorillas (Gorilla gorilla gorilla).

The transfer of food items between individuals has been described in primates as serving an informative purpose in addition to supplementing the diet of immature individuals. This behaviour has yet to be described in western lowland gorillas (Gorilla gorilla gorilla), and results are presented here of observations of food transfers in immature gorillas at Mbeli Bai, Republic of Congo. The frequency of food transfers decreased with increasing immature age, while the frequency of independent feeding and processing of food increased. Transfers between mothers and infants were the most frequent, with infants attempting to take items from the mother. These attempts were not always successful and the item was relinquished on less than 50% of attempts. Mothers also took items from their offspring. The results point to the functional significance of food transfers in western lowland gorillas being informational. In a bai environment, where one species forms the majority of a visiting gorilla's diet despite other species being available, the initiation of food transfers by immatures is proposed to serve the purpose of familiarising them with which species, and which parts of those species, may be eaten.

Age Factors↗

Gorillas' (Gorilla gorilla gorilla) spatial memory in a foraging task.

The spatial memory of 2 gorillas (Gorilla gorilla gorilla) was explored in a simulated foraging task. Trials consisted of 2 parts separated by a delay. In the 1st part, half of the total number of food sites were baited with a highly preferred food, and the subject was allowed to search, find, and consume these items (search phase). During the delay the same locations were again baited. After the delay the animal was reintroduced to the test enclosure and allowed to search through the sites again (re-search phase). In Experiment 1, an adult gorilla was very accurate in remembering locations that had previously contained food at delay intervals of 24 hr or more. In Experiment 2, a juvenile gorilla was also accurate in remembering locations that had previously contained food at delays up to 10 min. The adult gorilla appeared to use a counting strategy during the search phase to minimize the number of sites visited.

Animals↗

Chronic hypertension with subsequent congestive heart failure in a western lowland gorilla (Gorilla gorilla gorilla).

Chronic severe subclinical systemic hypertension was diagnosed in a 28-yr-old male western lowland gorilla (Gorilla gorilla gorilla). Thoracic radiography, electrocardiography, and echocardiography revealed an enlarged heart with a hypertrophied left ventricle, mitral regurgitation, and a persistent left bundle branch block. Enalapril, later combined with nifedipine, was of some value in reducing the hypertension, with partial reversal of cardiac enlargement and resolution of the bundle branch block. Two years after initiation of treatment, the gorilla developed lethargy and dyspnea. The diagnosis of heart failure was confirmed under anesthesia; the gorilla did not recover and was euthanized. Postmortem examination confirmed congestive heart failure with chronic, fibrosing cardiomyopathy similar to that in other gorillas.

Animals↗

Diagnosis of hypothyroidism in a western lowland gorilla (Gorilla gorilla gorilla) using human thyroid-stimulating hormone assay.

Primary hypothyroidism was diagnosed in a 26-yr-old female western lowland gorilla (Gorilla gorilla gorilla) on the basis of serum levels of thyroxine (T4), free T4, and thyroid-stimulating hormone (TSH) measured by human immunoassays. Compared with clinically normal gorillas, the TSH level (107 mlIU/L) was markedly elevated, and T4 (<14.0 nmol/L) and free T4 (5.0 pmol/L) levels were decreased. Thyroid hypofunction could explain the weight gain, unsettled appetite, anxious behavior, lethargy, and poor intraspecies interactions shown by this gorilla. The antibodies in the commercial immunoassay used in this study apparently cross-reacted with gorilla TSH. Supplementation with levothyroxine sodium was initiated and was followed by a marked decrease in circulating TSH and a noticeable improvement in the animal's physiologic status and activity level.

Animals↗

A new southern limit for the distribution of African great apes: sympatric western lowland gorilla (Gorilla gorilla gorilla) and central chimpanzee (Pan troglodytes troglodytes) confirmed in Mayombe National Park, Angola.

The distribution of African great apes has remained unconfirmed regarding their southern limit, particularly on the western side of the continent. IUCN maps include the Mayombe forest of Angola as part of the estimated distribution of western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes). However, until now no published evidence-based records had confirmed the continued presence of both species. The Mayombe forest is a key biodiversity hotspot and a potentially important stronghold for the conservation of great ape populations in Africa. Here, we report the first systematic evidence of both species in the Mayombe National Park, Cabinda, Angola. In 2023, a grid of camera traps was systematically deployed, producing the first visual records of gorillas and chimpanzees. Building on these findings, in 2024, a pilot survey including ad libitum field observations was carried out along exploratory trails to maximise data collection. The combination of these records identified a hotspot of great ape activity where six transects were established, and systematic direct and indirect evidence was documented. Chimpanzees were recorded more times across a broader range of evidence categories, while gorillas appeared less and seemed more spatially restricted. Notably, both species were detected at overlapping sites but never simultaneously, indicating sympatric coexistence with spatio-temporal partitioning. These findings confirm the southernmost predicted distribution of both species for this part of Africa, filling critical gaps in the understanding of great ape evolution and biogeography, and providing a baseline for the first demographic and ecological census of great apes in Angola.

Animals↗

Population and group structure of western lowland gorillas (Gorilla gorilla gorilla) at Lokoué, Republic of Congo.

During a 17-month study at the Lokoué clearing in Odzala National Park, Republic of Congo, we identified 377 western lowland gorillas. This population included 31 solitary males, 37 breeding groups, and eight nonbreeding groups. Its age- and sex-class structure was similar to those observed at two other clearings in the same forest block. However, the size of breeding groups varied with site (either clearing or forest sites). At Lokoué, breeding groups (mean size: 8.2 gorillas; range: 3-15) included a single silverback male and, on average, 3.2 adult females. Nonbreeding groups (mean size: 5.5; range: 2-15) were devoid of adult females. Five of the nonbreeding groups were composed predominantly of blackbacks, subadult males, and juveniles, and thus fit the definition of all-male groups previously observed in mountain gorillas. Our study confirms that 1) one-male breeding groups are the norm in western gorillas, and 2) all-male groups occur in this species. Despite frequent changes in members due to migrations of the males, the persistence of these all-male groups indicates that they may play an important role in the life of migrating males. Variations in population structure, and group composition and type among gorilla populations are discussed. However, a further understanding of the evolution of group-living in gorillas requires detailed ecological studies conducted in parallel with studies of the population structure and dynamics of these groups.

Age Factors↗

Peripartum sex steroid profiles and endocrine correlates of postpartum maternal behavior in captive gorillas (Gorilla gorilla gorilla).

An association between pregnancy levels of estrogen and progesterone and maternal behavior has been demonstrated in several taxonomic orders of nonprimate and primate mammals, but has not so far been investigated in the gorilla. In this study we investigated whether prepartum titers of urinary estrone conjugates (E1C) or pregnanediol-3alpha-glucuronide (PdG) were related to postpartum maternal behavior in eight multiparous Western lowland gorilla females (Gorilla gorilla gorilla) housed in four zoological gardens. Urine samples were collected from each study animal for 14 days prepartum and 14 days postpartum, and measures of maternal responsiveness were scored during the first 15 days postpartum. Urine samples were assayed with radioimmunoassay for E1C and PdG. Results for the peripartum profiles of urinary E1C, as well as postpartum profiles of PdG, agree with previous findings for the gorilla, while results for late-pregnancy profiles of urinary PdG were inconclusive in confirming a prepartum increase or decrease. Neither prepartum levels of E1C or PdG, nor the E1C/PdG ratio were found to be related to measures of postpartum maternal behavior. This lack of association between late-pregnancy E1C titers per se and postpartum maternal behavior is contrary to findings in nonprimate and other primate species.

Animals↗