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PubMed1988

Molecular cloning and characterization of the DNA of two papillomaviruses from monkeys.

Benign and malignant lesions from monkeys were analyzed for the presence of papillomavirus (PV) DNA. By hybridization with different PV DNA probes under conditions of lowered stringency, two tumors were found to contain PV-specific DNA sequences: (1) a cutaneous papilloma from a Colobus monkey; and, (2) a lymph node metastasis of a squamous cell carcinoma of the penis from a Rhesus monkey. Analysis of the DNA of the papilloma from the Colobus monkey indicated the presence of extrachromosomal DNA whereas analysis of DNA from the Rhesus tumor suggested the presence of integrated viral DNA. The physical size (7.8 and 8.1 kb), colinear alignment to HPV-5, and cross-hybridization with other PV types under low stringency indicate that the two genomic DNA clones represent new PV types that have been tentatively designated as Rhesus papillomavirus type 1 (RhPV 1) and Colobus guereza papillomavirus type 2 (CgPV 2). A putative viral-host DNA junction fragment was also isolated from the Rhesus genomic library. Nucleotide sequences very closely related to RhPV 1 were observed by in situ hybridization in a laryngeal carcinoma from the Colobus guereza monkey. This report communicates the finding of novel papillomaviruses associated with a benign cutaneous tumor and genital and laryngeal malignancies in non-human primates which may have significance as a putative system for the study of papillomavirus-induced genital and laryngeal malignancies in humans.

Animals↗
PubMed2006

Spatial distribution of primates in a mosaic of colonizing and old growth forest at Ngogo, Kibale National Park, Uganda.

Primate censuses were conducted in a mosaic of colonizing (two locations) and old-growth forests using line transect methods at the Ngogo study site, Kibale National Park, Uganda. Black and white colobus monkeys (Colobus guereza) were encountered more frequently in the colonizing forests than in the old growth forest, while chimpanzees (Pan troglodytes) were encountered more frequently in the old growth forest than in colonizing forests. Although not significant, results suggest that blue monkeys (Cercopithecus mitis) frequented colonizing forests more often than old growth forest. The encounter rates of mangabey (Lophocebus albigena), and redtail (Cercopithecus ascanius) groups were ambiguous with their density being higher in some colonizing forests but not others as compared to old-growth forest. No significant differences were detected for baboons (Papio anubis), L'hoest's (Cercopithecus lhoesti), and red colobus monkeys (Piliocolobus tephroscales). The conversion of forests to farmland is one of the major problems encountered in primate conservation. This study shows that secondary forests replacing anthropogenic grasslands have the potential of supporting some primate species such as black and white colobus, redtail monkeys, and possibly blue monkeys. Therefore, such areas should not be given up but should be conserved for the benefit of primates that can survive in secondary forests; as the forests mature further, primate species that are adapted to old growth forest will colonize the area provided there is a nearby source.

Animals↗
PubMed1999

Genetically engineered Mengo virus vaccination of multiple captive wildlife species.

Encephalomyocarditis virus (EMCV), has caused the deaths of many species of animals in zoological parks and research institutions. The Audubon Park Zoo, (New Orleans, Louisiana, USA) attempted vaccination of several species with a killed EMCV vaccine with mixed results. This paper reports an attempt at vaccination against EMCV using a genetically engineered, live attenuated Mengo virus (vMC0) at the Audubon Park Zoo and Miami Metro Zoo, (Miami, Florida, USA) from December 1996 to June 1997. Several species of animals were vaccinated with vMC0, which is serologically indistinguishable from the field strain of EMCV. Serum samples were taken at the time of vaccination and again 21 days later, then submitted for serum neutralization titers against EMCV. The vaccinate species included red capped mangebey (Cercocebus torquatus), colobus (Colobus guereza), angolan colobus (Colobus angolensis), ruffed lemur (Lemur variegatus ruber and Lemur variegatus variegatus), back lemur (Lemur macaco), ring-tailed lemur (Lemur catta), siamang (Hylobates syndactylus), diana guenon (Cercopithicus diana), spider monkey (Ateles geoffroyi), common marmoset (Callithrix jacchus), talapoin monkey (Cercopithecus talapoin), Brazilian tapir (Tapirus terrestris), Baird's tapir (Tapirus bairdii), Malayan tapir (Tapirus indicus), dromedary camel (Camelus dromedarius), bactrian camel (Camelus bactrianus), gerenuk (Litocranius walleri), guanaco (Lama glama guanicoe), black duiker (Cephalophus niger), Vietnamese potbellied pig (Sus scrofa), babirusa (Babyrousa babyrussa), collard peccary (Tayass tajacu), and African crested porcupine (Hystrix africaeaustralis). The vaccine response was variable, with high virus neutralizing antibody titer responses in some primate species and mixed to poor responses for other species. No ill effects were seen with vaccination.

Animals↗
PubMed1997

Mapping homology between human and black and white colobine monkey chromosomes by fluorescent in situ hybridization.

We used in situ hybridization of chromosome specific DNA probes ("chromosome painting") of all human chromosomes to establish homologies between the human and the white and black colobus (Colobus guereza 2n = 44). The 24 human paints gave 31 signals on the autosomes (haploid male chromosomes homologus to human 14 and 15, 21 and 22, form colobine chromosomes 6 and 16, respectively. Reciprocal translocations were found between human chromosomes 1 and 10, 1 and 17, as well as 3 and 19. The alternating hybridization signals between human 3 and 19 on Colobus chromosome 12 show that in this case a reciprocal translocation was followed by a pericentric inversion. The hybridization data show that in spite of the same diploid number and similar Fundamental Numbers, the black and white colobine monkey differs from Presbytis cristata, an Asian colobine, by 6 reciprocal translocations. Comparisons with the hybridization patterns in other primates show that some Asian colobines have a more derived karyotype with respect to African colobines, macaques, great apes, and humans. Chromosome painting also clearly shows that similarities in diploid number and chromosome morphology both between colobines and gibbons are due to convergence.

Animals↗
PubMed1997

Genital papillomaviruses (PVs) and epidermodysplasia verruciformis PVs occur in the same monkey species: implications for PV evolution.

Portions of the genome from two different papillomaviruses (PVs) of the Abyssinian Colobus monkey were sequenced and phylogenetically analyzed. This revealed that the major evolutionary separation between genital PVs and epidermodysplasia verruciformis-associated PVs (EV-PVs) hitherto found only in human papillomaviruses (HPVs) also exists in animal PVs. The sequence of the long control region (LCR) of Colobus monkey PV type 2 (CgPV-2) reveals a small size and an arrangement of potential cis-responsive elements typical of the EV-HPVs; namely four binding sites for the viral E2 protein, with one of them being located within the L1 gene, a cluster of nuclear factor I (NFI)- and AP-1-binding sites and a 50-bp sequence upstream of the E6 gene consisting only of the nucleotides A and T. This level of conservation of functional elements within the highly variable LCR suggests that CgPV-2 could be adopted as a model for studying human skin cancer associated with EV-HPVs. Although isolated from the same monkey species, the other Colobus monkey PV, CgPV-1, is a typical genital PV as shown by E1 and L1 sequence comparisons. The presence of these two major phylogenetic divisions of PVs in both human and monkey hosts strongly suggests that this diversification predated the evolutionary split between monkeys and apes. In other words, at least two different groups of PVs have been evolving separately in their respective primate hosts for more than 22 million years with only moderate sequence changes since their genesis.

Animals↗
PubMed1983

Rise and fall of the delta globin gene.

This paper examines the molecular basis for the origin and subsequent silencing of the delta globin gene during the evolution of higher primates. We cloned and sequenced the delta gene from representatives of the two most distantly related types of Old World monkeys in which the gene is silent. The silent delta genes of these two monkeys (rhesus and colobus) are highly homologous to each other (97.5%) and to the functional human delta gene (94%). The presumed mutation that silenced the delta gene is therefore unlikely to be obscured by later mutations at the same site. This silencing mutation could have affected transcription since we find no delta transcript in rhesus bone marrow. Furthermore, the delta globin genes from colobus and rhesus monkeys are many times less efficient than the human delta gene in a cell-free transcription assay. The two Old World monkeys share only three nucleotide substitutions in the first 100 base-pairs of the DNA sequence 5' to the site for initiation of transcription when compared to human delta. It is likely that the in vitro silencing of the delta gene was brought about by one (or a combination) of these three substitutions. Three analysis shows how the delta sequences are related to those for other beta-like genes, including the colobus beta gene whose sequence we report. One part of the delta gene appears to be derived from a (pseudo?) beta gene that diverged from beta before the divergence of mice and men. The other part is closely related to the beta gene of higher primates. The mosaic nature of the delta gene is apparently due to a partial conversion which took place between the pseudo beta and the functional beta gene before the divergence of the lineages leading to Old World monkeys and hominoids.

Animals↗
PubMed1989

Evolution and transcription of old world monkey globin genes.

Although apparently inactive in the whole animal, the delta globin genes from three species of Old World monkey (rhesus, baboon and green monkey) are all functional in an in vitro transcription assay. Their activities in vitro are similar to that of the functional human delta gene. A fourth monkey gene, from the colobus monkey, is transcribed approximately fivefold less efficiently than the others. This reduced in vitro activity results from a 20 base-pair deletion, which removes the normal site of mRNA initiation. When the deletion is repaired by site-directed mutagenesis, transcriptional activity increases to the level observed for the other delta genes. We also report the complete nucleotide sequences of the colobus beta, colobus delta, and rhesus delta genes. Sequence comparisons show that the delta and beta genes in the same species have not exchanged genetic information since the divergence of the human and monkey lineages. Phylogenetic analysis of these sequences affirms that the Old World monkey delta genes are evolving more rapidly than their functional counterparts. Moreover, the rate of replacement substitutions has risen to equal that of non-coding DNA, as expected for genes no longer under selective constraint.

Amino Acid Sequence↗
PubMed1998

Some gastro-intestinal parasites of zoonotic (public health) importance commonly observed in old world non-human primates in Kenya.

A study was undertaken to categorise some gastro-intestinal (GIT) parasites commonly observed in Kenyan non-human primates (NHPs) on the basis of their health implications for humans. Six species of locally available non-human primates, namely olive baboons (Papio cyanocephalus anubis), Vervet monkey (Cercopithecus aethiops), Sykes monkey (Cercopithecus mitis), Black and white colobus (Colobus abyssinicus), Debrazzas monkey (Cercopithecus neglectus) and Grey and Black mangabeys (Cercocebus torquatus and Cercocebus albigena) which were imported from Zaire (Democratic Republic of Congo) were sampled. Simple laboratory methods involving microscopic examination of stained faecal smears were used. Wet faecal smears stained with iodine and unstained controls were used for conventional parasites while acid fast staining was employed to detect Cryptosporidium oocysts. Both helminths and protozoan parasites were detected in varying rates in all primate species. Trichuris sp. was the most frequent helminth followed by Strongyloides fulleborni, Strongyles sp. and Schistosoma mansoni in that order. Entamoeba coli was the most common protozoan followed, respectively, by Balantidiun coli and Entamoeba histolytica. All primate species examined were infected with all the parasites listed except the black and white colobus. Cryptosporidium was found in both clinically normal and diarrhoeic baboons and vervets. Most taxa of parasites observed could prejudice human welfare directly through infection and causation of illness and indirectly through increased cost of livestock production and decreased availability of animal proteins. The potential of some of the agents to cause opportunistic infections in immuno-compromised persons was suggested as a likely threat to man's well-being. This would warrant such person's exemption from high risk operations at primate and other animal facilities in developing countries. Further, specific studies are needed to provide data on the epidemiology, socio-economic impact and pathogenicity of the primate parasites to other species of animals and man.

Animals↗
PubMed1997

The primate community of the Lopé Reserve, Gabon: diets, responses to fruit scarcity, and effects on biomass.

The diets of all diurnal primates (Gorilla g. gorilla, Pan t. troglodytes, Mandrillus sphinx, Colobus satanas, Cercocebus albigena, Cercopithecus nictitans, C. pogonias, C. cephus) in the Lópe Reserve, central Gabon, are described from qualitative and quantitative data collected over 10 years. A total of 397 foods were recorded, of which 91% were from plants. The diet of seven of the eight species were numerically dominated by fruit, the exception being Colobus satanas with a diet dominated by seeds. When proportion of feeding time was examined, fruit remained the dominant food class for six species, while Cercocebus albigena, like the black colobus, spent most time feeding on seeds. The number of foods recorded per species ranged from 46-220, but dietary breadth of the four species that have not been subjects of detailed study was certainly underestimated. Almost half of the foods (46%) were eaten by only one species, and only four (1%) were recorded for all eight species. At Lópe, fruit is scarce for 2-3 months during the annual dry season, and fruit crop failures can prolong this period of scarcity to as many as 8 months in some years. When fruit is scarce, the diets of all the species changed to include more nonfruit foods, but chimpanzees and the three species of guenons maintained a fruit dominated diet. The total biomass of the Lópe primate community is 374 kg/km2, which is low compared to other sites in tropical Africa. Within the Lópe community, biomass correlates negativity with degree of frugivory, suggesting that fruit availability may be critical, but this does not explain the low total biomass compared to other sites. Neither floristic diversity nor the composition of the primate community at Lópe can account for the biomass. While historical events such as climate-induced changes in forrest cover, disease, or impact of human activities cannot be ruled out, we conclude that the most likely explanation of low primate biomass is the occurrence of an ecological "catastrophe" in the fairly recent past from which populations have not yet recovered. We believe that the most likely scenario was climate-mediated disturbance of flowering of important fruit food species. Prolonged fruit scarcity would have caused mortalities and affected the most frugivorous species more severely.

Animals↗
PubMed1997

Fatal infections with Balamuthia mandrillaris (a free-living amoeba) in gorillas and other Old World primates.

Balamuthia mandrillaris is a newly described free-living amoeba capable of causing fatal meningoencephalitis in humans and animals. Because the number of human cases is rapidly increasing, this infection is now considered an important emerging disease by the medical community. A retrospective review of the pathology database for the Zoological Society of San Diego (the San Diego Zoo and San Diego Wild Animal Park) for the period July 1965 through December 1994 revealed five cases of amoebic meningoencephalitis, all in Old World primates. The infected animals were a 3-year, 10-month-old female mandrill (Papio sphinx), from which the original isolation of B. mandrillaris was made, a 5-year-old male white-cheeked gibbon (Hylobates concolor leucogenys), a 1-year-old female western lowland gorilla (Gorilla gorilla gorilla), a 13-year, 5-month-old male western lowland gorilla, and a 6-year-old female Kikuyu colobus monkey (Colobus guereza kikuyuensis). Two different disease patterns were identified: the gibbon, mandrill, and 1-year-old gorilla had an acute to subacute necrotizing amoebic meningoencephalitis with a short clinical course, and the adult gorilla and colobus monkey had a granulomatous amoebic meningoencephalitis with extraneural fibrogranulomatous inflammatory lesions and a long clinical course. Indirect immunofluorescent staining of amoebas in brain sections with a Balamuthia-specific polyclonal antibody was positive in all five animals. Indirect immunofluorescent staining for several species of Acanthamoeba, Naegleria fowleri, and Hartmanella vermiformis was negative. Direct examination of water and soil samples from the gorilla and former mandrill enclosures revealed unidentified amoebas in 11/27 samples, but intraperitoneal inoculations in mice failed to induce disease. Attempts to isolate amoebas from frozen tissues from the adult male gorilla were unsuccessful.

Amebiasis↗
PubMed1999

Demographic influences on the hunting behavior of chimpanzees.

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation.

Animals↗
PubMed2000

Diurnal primate densities and biomass in the Kakamega Forest: an evaluation of census methods and a comparison with other forests.

Line-transect surveys were conducted at the Isecheno study site in the Kakamega Forest, western Kenya to estimate diurnal primate densities. The estimates from several different methods of analysis of census data were compared to "true" density values based on home range size and overlap for two species. The Whitesides method [Whitesides et al., 1988], which incorporates species-specific mean group spread into its formula for estimating transect width, provided the most accurate density estimates. The importance of including as many groups as possible when calculating density from home range size and overlap is demonstrated with long-term data from Colobus guereza and Cercopithecus mitis. Colobus guereza group density at Isecheno was much lower than that published from a recent brief study [von Hippel, 1996]. Cercopithecus mitis group density has fallen while overall population biomass appears to have remained stable over 20 years of study. Isecheno has the second highest diurnal primate biomass of the ten Guineo-Congolian rainforest sites for which biomass data are available, despite having the lowest primate species richness. Within the Guineo-Congolian rainforest system, primate biomass appears to vary to some extent between ecogeographic regions: two of three mid-elevation East African sites have high biomasses, two of two lowland West African sites have intermediate biomasses, and four of five lowland Central African sites have low biomasses. There is a strong positive correlation between total colobine biomass and total primate biomass at the ten Guineo-Congolian rainforest sites.

Animals↗
PubMed2007

Plant DNA sequences from feces: potential means for assessing diets of wild primates.

Analyses of plant DNA in feces provides a promising, yet largely unexplored, means of documenting the diets of elusive primates. Here we demonstrate the promise and pitfalls of this approach using DNA extracted from fecal samples of wild western gorillas (Gorilla gorilla) and black and white colobus monkeys (Colobus guereza). From these DNA extracts we amplified, cloned, and sequenced small segments of chloroplast DNA (part of the rbcL gene) and plant nuclear DNA (ITS-2). The obtained sequences were compared to sequences generated from known plant samples and to those in GenBank to identify plant taxa in the feces. With further optimization, this method could provide a basic evaluation of minimum primate dietary diversity even when knowledge of local flora is limited. This approach may find application in studies characterizing the diets of poorly-known, unhabituated primate species or assaying consumer-resource relationships in an ecosystem.

Animals↗
PubMed1995

Positional behavior in five sympatric Old World monkeys.

Observations of positional behavior and habitat use were recorded on focal individuals of five species of Old World monkeys at Kibale Forest, Uganda, through the dry season of 1990 and 1991. Cercopithecus ascanius, Cercopithecus mitis, Cercocebus albigena, Colobus badius, and Colobus guereza commonly utilize five similar types of positional behavior (i.e., quadrupedalism, leaping, climbing, sitting, and standing), but in varying frequencies and situations. As a group, colobines use oblique supports and leap more often, and cover greater linear distances during leaps than do cercopithecines. Colobines also prefer to sit (about 90% of all postures), while cercopithecines stand more frequently. Body size differences between the sexes of a species are not reflected in positional behavior. The two small-bodied species climb more and leap less often than the three larger species, which is the reverse of what we would expect. Leaping is the most common method of crossing open spaces within the canopy; but most spatial gaps and leaps are over short distances, usually one meter or less. All five species, regardless of body size or the availability of forest supports, prefer medium-sized supports. Incorporating our work from Uganda with previous investigations of positional behavior reveals few consistent trends with respect to body size or habitat use across primates.

Animals↗
PubMed2005

Oesophagostomum bifurcum in non-human primates is not a potential reservoir for human infection in Ghana.

In northern Togo and Ghana, human infection with the parasitic nematode Oesophagostomum bifurcum is of major health importance. Elsewhere, oesophagostomiasis is considered a zoonotic infection, non-human primates being the natural host. We examined 349 faecal samples of the olive baboon, mona monkey and black and white colobus monkey from two geographically distinct areas in Ghana, outside the region endemic for O. bifurcum in humans. Using both microscopy and species-specific PCR, we found a high prevalence of O. bifurcum (75-99%) in olive baboons and mona monkeys. The majority of the test-positive faecal samples contained large numbers of larvae after copro-culture (>100). No O. bifurcum was detected in the faeces of the black and white colobus monkeys. Observational studies on the behaviour of the non-human primates, focusing on defecation, food consumption and the sharing of habitat with the local human population, indicated favourable conditions for zoonotic transmission. Given that no human infection with O. bifurcum has been reported from either study area, the present findings support the hypothesis that O. bifurcum from humans in the north of Ghana, and O. bifurcum from olive baboons and/or mona monkeys are distinct.

Animals↗
PubMed2006

Prediction of parasite infection dynamics in primate metapopulations based on attributes of forest fragmentation.

Although the effects of forest fragmentation on species and ecological processes have been the focus of considerable research in conservation biology, our capacity to predict how processes will be altered and which taxonomic or functional groups will be most affected by fragmentation is still poor. This problem is exacerbated by inherent temporal and spatial variability in fragment attributes. To improve our understanding of this interplay, we examined how various fragment attributes affect one potentially important ecological process, parasite infection dynamics, and considered how changes in this process affect host metapopulations. From August 1999 to July 2003 we surveyed red colobus (Piliocolobus tephrosceles) metapopulations inhabiting nine fragments (1.2 to 8.7 ha) in western Uganda to determine the prevalence and richness of strongyle and rhabditoid nematodes, a group of potentially pathogenic gastrointestinal parasites. We used noninvasive fecal flotation and sedimentation (n = 536) to detect parasite eggs, cysts, and larvae in colobus fecal samples. To obtain an index of infection risk, we determined environmental contamination with Oesophagostomum sp., a representative strongyle nematode, in canopy (n = 30) and ground vegetation plots (n = 30). Concurrently, physical (i.e., size, location, and topography) and biological (i.e., tree diversity, tree density, stump density, and colobine density) attributes were quantified for each fragment. Interfragment comparisons of nine potential factors demonstrated that an index of degradation and human presence (tree stump density) strongly influenced the prevalence of parasitic nematodes. Infection risk was also higher in the fragment with the highest stump density than in the fragment with the lowest stump density. These results demonstrate that host-parasite dynamics can be altered in complex ways by forest fragmentation and that intensity of extraction (e.g., stump density) best explains these changes.

Animals↗
PubMed1977

The chromosomal location of rDNA in selected lower primates.

Hybridization in stiu was used to identify the chromosomes that carry rDNA in representative lower primates, including the baboons, Papio cynocephalus and Papio hamadryas; the colobus monkey, Colobus polykomos; the tree shrew, Tupaia glis; the lemur, Lemur fulvis; the saki, Pithecia pithecia; the marmoset, Saguinus nigricollis, and the spider monkey, Ateles geoffroyi. The marker chromosome, common to the Cercopithecines studied to date, carries the rDNA in the baboons. Another marker chromosome carries rDNA in a South American species, the spider monkey. A multichromosomal distribution of rDNA was demonstrated in the tree shrew, lemur, saki, and marmoset. None of the rDNA-containing chromosomes in the prosimians and New World monkeys show homology to the chromosomes that carry rDNA in the Hominids, Pongids, or Old World monkeys.

Animals↗
PubMed2000

Catarrhine photopigments are optimized for detecting targets against a foliage background.

The colour vision of many primates is trichromatic, whereas that of all other mammals is thought to be dichromatic or monochromatic. Moreover, the triplets of cone pigments in different catarrhines (Old World apes and monkeys) are strikingly similar in their spectral positions. We ask whether the selective advantage of trichromacy lies in an enhanced ability to find edible leaves or fruit. Further, we ask whether any factor in these two search tasks has constrained the particular set of cone spectral sensitivities observed in all catarrhines. We measured the spectral properties of the natural environments of six primate species in Uganda: Pan troglodytes, Cercopithecus mitis, Cercopithecus ascanius, Lophocebus albigena, Colobus guereza and Colobus badius. We concentrated on the fruit and leaves in their diets and the leaves of the trees that make up the background against which these diet items must be found. We plotted these measured stimuli in colour spaces appropriate for each primate species, and found that both frugivory and folivory are facilitated by the extra dimension of colour vision found in catarrhines but lacking in most other mammals. Furthermore, by treating the task of searching for food as a signal-detection task, we show that, of all possible combinations of cone sensitivities, the spectral positions of the actual primate pigments are optimal for finding fruit or young leaves against the background of mature leaves. This is because the variance of the chromaticities of the mature leaves is minimised in one channel of the primate's colour vision, so allowing anything that is not a mature leaf to stand out.

Algorithms↗