Comparison of habitat quality and diet of Colobus vellerosus in forest fragments in Ghana

The forest fragments surrounding the Boabeng-Fiema Monkey Sanctuary (BFMS) in central Ghana shelter small populations of Colobus vellerosus. Little is known about these populations or the ability of the fragments to support them, despite the fact that these fragments represent potentially important habitat for the colobus in this region. We compared the diet of three groups of C. vellerosus in the fragments to two groups in BFMS. We also examined the differences in plant species composition and food abundance among fragments. The study took place from June to November 2003. Dietary data were collected using scan sampling. Plant species composition and food abundance were evaluated using tree plots and large tree surveys. As in BFMS groups, leaves constituted the highest proportion of the diet of fragment groups, yet the colobus in fragments fed on more lianas than did those in BFMS. Over 50% of all species observed eaten by colobus in the fragments were not consumed in BFMS groups during the same season. Food abundance was similar between fragments and BFMS, although species composition differed. There was no relationship between the density of colobus and the density of food trees or percentage of food species, suggesting that other factors may be influencing the number of colobus present. This study highlights the broad dietary range of C. vellerosus, which may be a factor allowing its survival in these fragments.     

Status of diurnal primate populations at the former settlement of a displaced village in Cameroon

To understand whether or not primate populations recover in areas of abandoned human settlements, data are required that allow for site‐specific comparison over time. Here, we present baseline information on the presence/absence and relative abundance of primate species at the abandoned settlement of the Ekundukundu village, relocated out of the Korup National Park in 2000. Between July 2007 and March 2008, 62 km of transects was surveyed for sightings and calls of primates. All eight species of diurnal primates reported in the KNP were confirmed: Cercopithecus nictitans ludio, C. mona, C. erythrotis camerunensis, C. pogonias, Procolobus pennantii preussi, Cercocebus torquatus, Mandrillus leucophaeus leucophaeus, and Pan troglodytes vellerosus. At old Ekundukundu, C. nictitans accounted for 65% of all primate group sightings. Overall, sighting frequency of primates (0.55 groups/km) was not significantly different from other park sectors surveyed by a previous observer (J. Linder) in 2004–2005. The data reported here will be useful in the long‐term monitoring of primate populations in regenerating forest habitats of earlier human settlements. Am. J. Primatol. 72:645–652, 2010. © 2010 Wiley‐Liss, Inc.     

High levels of infant handling by adult males in Rwenzori Angolan colobus (Colobus angolensis ruwenzorii) compared to two closely related species,C. guereza and C. vellerosus

Infant handling (holding or carrying) by adult males is rare in mammals; however, high levels have been reported in some primates. Though infant handling is a costly behaviour, there are many benefits that male handlers can accrue. Infant handling by males is most conspicuous in platyrrhines and tends to be uncommon in catarrhines. In the latter species, research on male–infant interactions has focused on low-cost behaviours, such as proximity and grooming. However, to better understand the evolution of infant handling by males, more data on its occurrence across the Primate order are essential, even in species where it is relatively uncommon. We compare the occurrence of infant handling by males in three closely related species of catarrhine: Colobus vellerosus, C. guereza, and C. angolensis ruwenzorii. We collected focal animal samples on infants to quantify infant handling rates and durations, and found that adult male C. a. ruwenzorii handled infants much more frequently and for much longer than males in the other two species. We discuss how C. a. ruwenzorii’s unique social organization may explain high levels of infant handling by adult males in this species. More long-term and detailed comparisons of infant handling across species and populations will shed light on how sociality has shaped the evolution of this behaviour in the Primate order.

     

Development of the integument of Dasypus hybridus and Chaetophractus vellerosus,and asynchronous events with respect to the postcranium

The integument of extant armadillos (Xenarthra, Cingulata) is a unique organ in which complex glandular systems are associated with pilose follicles, dermal ossifications, and cornified scales. Up to date, papers have focused on neither comparative morphology of the skin (dorsal and ventral) nor chronology of the development of interspecific homolog structures. In order to clarify the way in which events occur during development of the integument structures, maturity of other tissues (e.g. skeletal tissues) should be considered. Therefore, we will be able to identify events that have been pre- or post-displaced during ontogenetic development. The aim of this paper is to describe in a developmental and comparative framework the integumentary system of neonates of Dasypus hybridus and Chaetophractus vellerosus. In order to understand the morphology of the different integumentary structures serial histological sections were prepared. Staining techniques included H–E, Masson Trichrome, PAS, orcein and reticulin. To study ossification of postcranial elements, the specimens were cleared and double-stained with alcian blue and alizarin red. Determinations of ossification centers and their progress were recorded through the early uptake of alizarin. The dorsal dermis of neonates from D. hybridus is clearly differentiated into a superficial and deep layer, as in fetuses of Dasypus novemcinctus. In C. vellerosus, however, these layers could not be identified. This suggests a less connective tissue differentiation in the latter species at this stage. Osteoderms in D. hybridus are well differentiated unlike C. vellerosus where no condensations of osteoprogenitory cells are observed. Conversely, pilose follicles and glandular tissues are less developed in D. hybridus. Regarding postcranial elements, ossification centers are less advanced in C. vellerosus than D. hybridus, this is particularly notorious for the vertebral column, sternal, and pelvic girdle elements. Asynchronies between neonates of both species observed on integumentary and postcranial skeletal tissues could match with specific adaptive strategies related to distribution in different environments, and/or different postnatal care.     

Male takeover in Colobus vellerosus at Boabeng-Fiema Monkey Sanctuary, central Ghana

We describe a case of male takeover in the ursine black-and-white colobus (Colobus vellerosus). In April 2001, an all-male group attacked and eventually invaded our uni-male study group. Aggression increased following the takeover and the former resident male, severely wounded, became peripheral. The youngest immature received severe aggression from the new males but survived. The immatures mother intervened in most instances of this aggression. Eventually, the former resident male re-established relationships with some of the females and concurrently intervened to protect the immature. Defeated males that stay in their group can contribute to the protection of infants born during their tenure from infanticidal males. The females mated with the new males. Takeovers may be a means by which males acquire groups of females in C. vellerosus.     

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.     

Tropical rain forest conversion and perspectives in the conservation of wild primates (Alouatta and Ateles) in Mexico

The original distribution of the tropical rain forest and of the populations of Alouatta palliata, Al. pigra, and the two subspecies of Ateles geoffroyi in southern Mexico have been reduced by at least 90% in the last 40 years as a result of conversion of natural habitat to pasture and agricultural fields. This dramatic conversion has been caused mainly by the rapid growth of the human population in the southern states of the country. In the region of Los Tuxtlas in southern Veracruz, where the only longitudinal popululational and ecological studies of A. palliata and At. g. vellerosus have taken place, only 15% of the original extension of the tropical rain forest remains today. The intensive destruction of suitable primate habitat in this region has resulted in an accelerated process of extinction of the primate species. It is estimated that only about 200 At. g. vellerosus and about 1200 Al. palliata exist in the remaining small portion of their original habitat. Today, the distribution of the three primate species in Mexico is intensively and extensively fragmented, and only five potential foci for conservation exist in the country. Urgent action is required to protect the primate populations in the region of Los Tuxtlas and at the other four foci, as some of these populations may disappear by 1995.     

Seasonal changes in ovarian steroid hormone concentrations in the large hairy armadillo (Chaetophractus villosus) and the crying armadillo (Chaetophractus vellerosus)

Knowledge of armadillo reproductive physiology is essential for developing ex situ and in situ assisted reproductive techniques for propagating and/or controlling populations of these animals. The present study included assessment of fecal sex steroids by radioimmunoassay, determining reproductive status via monitoring ovarian activity (in the wild) and therefore reproductive status, in wild females of the large hairy armadillo (Chaetophractus villosus) and the crying armadillo (Chaetophractus vellerosus) in the southern hemisphere. Plasma and fresh fecal progesterone concentrations were not significantly correlated in either species. However, in both species, there was a significant positive correlation between plasma progesterone and dry fecal progesterone concentrations (r = 0.82, P < 0.05 and r = 0.60, P < 0.05, respectively). Dry fecal progesterone and estradiol concentrations were measured in one captive C. villosus (average baseline progesterone and estradiol concentrations 28.72 ± 11.75 ng/g dry feces and 3.04 ± 0.80 ng/g dry feces, respectively) and one captive C. vellerosus (average baseline progesterone and estradiol concentrations 14.05 ± 3.03 ng/g dry feces and 3.46 ± 1.20 ng/g dry feces, respectively) to detect hormonal peaks over 1 y; these occurred from late fall to early summer. Feces from wild C. villosus and C. vellerosus were also collected over 1 y to determine progesterone peaks, which occurred in winter and spring in both species (with no peaks during the summer or fall). Accordingly, C. villosus and C. vellerosus had a seasonal reproductive pattern. The significant correlations between dry fecal and plasma progesterone concentrations validated this method for monitoring reproductive status in these species.     

Scramble Competition Among <Emphasis Type="Italic">Colobus vellerosus</Emphasis> at Boabeng-Fiema,Ghana

We investigated the occurrence of scramble competition among Colobus vellerosus at Boabeng-Fiema, Ghana. If scramble competition had an impact on feeding efficiency among females, we expected a positive relationship between group size and the proportion of time spent feeding, day journey length, or home range size assuming resource availability is similar among the groups compared. We collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000–August 2001) on 2 study groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also collected ranging data on group movements at half-hour intervals. The large group (WW1) had a significantly longer day journey length than the small group (B1), and females in the large group spent a significantly greater proportion of time feeding in the wet season, a period of low food availability, which suggests it may be a bottleneck period when food resources are scarce and Colobus vellerosus is close to being energy limited. The proximity data suggested females may be able to reduce or adjust for competition by having fewer neighbors when they feed and by spreading out when in a larger group. However, we found no relationship between home range size and group size or that females spent a greater proportion of time feeding than adult males did. Our results highlight the need to factor in differences in food availability when investigating scramble competition. Though equivocal, our results suggest scramble competition occurs among Colobus vellerosus, leading us to suggest there was a match with the potential competitive regime, i.e., food distribution.     

Leaf-swallowing in Nigerian chimpanzees: evidence for assumed self-medication

A field study in Gashaka, Nigeria, adds the fourth subspecies of chimpanzee, Pan troglodytes vellerosus, to the list of African ape populations in which leaf-swallowing occurs. Unchewed herbaceous leaves of Desmodium gangeticum (Leguminosae-Papilionoideae) were recovered in 4% of 299 faecal samples of wild chimpanzees and clumps of sharp-edged grass leaves in 2%. The ingestion is believed to serve self-medicatory purposes because the leaves had a rough surface or were sharp-edged (which could be related to parasite control through a self-induced increase of gut motility), were not chewed, were excreted whole (indicating that they were not ingested for nutritional purposes), leaf-swallowing was restricted to the rainy season (during which time parasite re-infections are more common), and parasitic worms (Oesophagostomum stephanostomum) were found together with the leaves. Submitted to: Primates, May 2005; revision: 26 Oct 2005     

Ontogenetic criteria to distinguish vertebral types on the debated xenarthran synsacrum

The presence of a synsacrum formed by the fusion of vertebrae that come into closed contact with the ilium and ischium is a feature that characterizes the clade Xenarthra. Nevertheless, the proper identity of each vertebral element that forms it is a matter of discussion. In this article, we provide ontogenetic information about skeletal ossification of the xenarthran synsacrum and define the position of the sacrocaudal limit within it. We analyzed the synsacrum of 25 specimens of nonadult and 101 adult armadillos and anteaters: Dasypus hybridus, D. novemcinctus, Chaetophractus vellerosus, C. villosus, Tamandua tetradactyla, and Myrmecophaga tridactyla. Two sets of vertebrae were identified: an anterior set, often attached to the iliac bones, in which transverse processes are originated mainly from an expansion of the base of the neural arches, and secondarily from a lateroventral ossification center. A posterior set is characterized by a series of vertebrae along which extra lateral ossifications (described here for the first time) are developed and form exclusively the transverse processes. Among armadillos, the sacrocaudal limit is set between the last vertebrae attached to the iliac bones and the first vertebrae that form the dorsal border of the sacroischial fenestra. In addition, anterior free caudals also showed extra lateral ossifications forming exclusively the transverse processes, supporting the notion that more posterior synsacrals are in fact caudal vertebrae that were incorporated to the synsacrum. In pilosans, the sacrocaudal limit is set between the first vertebrae that come into contact with the ischial bones and the immediately anterior one. However, the pattern of homologies is obscured by the low resolution in the ontogenetic sequence when compared to that of armadillos. J. Morphol. 276:494–502, 2015. © 2014 Wiley Periodicals, Inc.     

Primate surveys in Côte d'Ivoire's Sassandra–Bandama interfluvial region with notes on a remnant population of black‐and‐white colobus

We conducted surveys in Côte d'Ivoire's Bandama–Sassandra interfluvial region to assess the abundance of primates and investigate further a population of black‐and‐white colobus monkey whose taxonomic position has been debated. Primates were encountered in five of eight forests surveyed. The most common primates were Campbell's guenons and lesser spot nosed monkeys. Roloway monkeys were located in one reserve (Dassioko FR), sooty mangabeys in two (Dassioko and Port Gauthier FR) and olive colobus in three reserves (Dassioko, Port Gauthier, Bakadou) as well as one forest patch was considered a sacred grove (Grébouo I FG). Red colobus and chimpanzees were never observed. The only black‐and‐white colobus encountered were located in a forest adjacent to the Grébouo I village. The coat of this population is similar to that of Colobus vellerosus; however, vocal data are more complex. In one vocal attribute, the Grébouo I colobus is similar to Colobus polykomos while in another it is similar to C. vellerosus. We hope that by investigating the taxonomy of the Grébouo 1 colobus population, highlighting the conservation value of sacred forests and advocating for more active law enforcement in protected reserves, we can prevent the extirpation of all primates within the Bandama–Sassandra interfluvial zone region.     

Morphology of female genital tracts in Dasypodidae (Xenarthra,Mammalia): a comparative survey

Previous works about comparative spermatology in Dasypodidae determined that sperm morphology is a striking variable among genera. It was suggested that this sperm feature may be related to specific morphologies of the female reproductive tract. The present comparative study of the morphology of the female genital tract from seven species corresponding to six genera of Dasypodidae is aimed to determine the main similarities and differences between the species and to establish a possible correlation with the sperm shapes and sizes. Genital tracts were studied macroscopically and histologically. Dasypus hybridus has disk-shaped ovaries and the cortex occupies almost all the organ with a single oocyte in each follicle. Tolypeutes matacus, Chaetophractus villosus, Chaetophractus vellerosus, Zaedyus pichiy, Cabassous chacoensis and Clamyphorus truncatus possess ovoid and elongated ovaries, with both longitudinally polarized cortex and medulla, and the peculiar presence of several oocytes in the same follicle. D. hybridus and T. matacus have a simple pear-shaped uterus, but in the other species the uterus is pyramid shaped and bicornuate. The uterine cervix is very long in all studied species. Only T. matacus presents a true vagina as in most eutherian mammals; on the other hand, in the other species a urogenital sinus is observed. The structure of female reproductive tracts in Dasypodidae contains a mixture of assumedly primary and other derived features. According to the different morphologies of the regions analyzed, a classification of the female genital tracts in three groups can be performed (group 1: Dasypus; group 2: Tolypeutes; group 3: Chaetophractus, Zaedyus, Cabassous, Clamyphorus) and a correlation between each group and a specific sperm morphology can be established.     

Predicting the Competitive Regime of Female <Emphasis Type="Italic">Colobus vellerosus</Emphasis> from the Distribution of Food Resources

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.     

Aspects of Male Competition in <Emphasis Type="Italic">Colobus vellerosus</Emphasis>: Preliminary Data on Male and Female Loud Calling,and Infant Deaths After a Takeover

Male Colobus vellerosus are the main participants in intergroup encounters, and lead incursions in neighboring groups during which they attack infants. Extragroup copulations, all-male groups, and male takeover occur in the species. Here, we provide additional information on behaviors associated with male reproductive competition in Colobus vellerosus. We examined 1 resident male loud calling and participation in intergroup encounters in relation to a takeover. We also report a second case of takeover that led to the death of the former resident male and the death of 2 male infants, presumably as a result of aggression from the all-male group. The new resident male wounded the third infant of the group, which apparently died after its mother abandoned it. During the period characterized by the attacks on the infant and after its disappearance, females initiated and participated in loud call bouts with the new resident male. We examine the possible functions of female loud calling, and suggest that in this context, it might force the resident male to call along to indicate his presence.     

Social correlates of fecal testosterone in male ursine colobus monkeys (Colobus vellerosus): the effect of male reproductive competition in aseasonal breeders

Male testosterone (T) levels are thought to be linked with the mating system, degree of parental care, and male–male aggression in reproductive contexts (The ‘challenge hypothesis’; Wingfield et al., 1990). In many species though, T increases associated with mating behavior cannot be separated from those associated with male–male aggression. We tested the challenge hypothesis on aseasonally breeding ursine colobus (Colobus vellerosus), where male–male competition is intense outside of mating contexts. Fecal samples (N = 109) were collected from > 27 subadult and adult males in seven groups during 13-months of research in Ghana in 2004–2005. Fecal T (fT) levels were determined by enzyme immunosorbant assays. Behavioral data was collected using focal-animal and ad libitum sampling. The number of receptive females in each group did not positively correlate with male fT. There was a trend for adult males to have higher fT than subadult males; however there was no effect of rank on fT. The level of male–male aggression experienced was positively correlated with fT and individual males showed higher mean fT during ‘challenge’ than during ‘non-challenge’ periods. The number of male incursions experienced positively correlated with fT whereas the number of between-group encounters did not. Males attempt to gain reproductive opportunities during incursions, thus these results support the ‘challenge hypothesis’ in C. vellerosus. Outside of mating contexts, higher male fT levels are associated with increased aggression. Male parental investment in the form of infant defense was associated with increased fT, rather than the decline expected from other forms of paternal care.     

Divergence in gut microbial communities mirrors a social group fission event in a black‐and‐white colobus monkey (Colobus vellerosus)

Host behavior and social factors have increasingly been implicated in structuring the composition of gut microbial communities. In social animals, distinct microbial communities characterize different social groups across a variety of taxa, although little longitudinal research has been conducted that demonstrates how this divergence occurs. Our study addresses this question by characterizing the gut microbial composition of an African Old World monkey, the black‐and‐white colobus (Colobus vellerosus), before and after a social group fission event. Gut microbial taxonomic composition of these monkeys was profiled using the V‐4 hypervariable region of the bacterial 16S ribosomal RNA gene, and pairwise‐relatedness values were calculated for all individuals using 17 short tandem repeat loci and partial pedigree information. The two social groups in this study were found to harbor distinct microbial signatures after the fission event from which they emerged, while these communities were not divergent in the same individuals before this event. Three genera were found to differ in abundance between the two new social groups: Parabacteroides, Coprococcus, and Porphyromonadaceae. Additionally, although this fission happened partially along lines of relatedness, relatedness did not structure the differences that we found. Taken together, this study suggests that distinct gut microbial profiles can emerge in social groups in <1 year and recommends further work into more finely mapping the timescales, causes, and potentially adaptive effects of this recurring trend toward distinct group microbial signatures.     

Ontogenetic variation in the stratum granulosum of the epidermis of Chaetophractus vellerosus (Xenarthra,Dasypodidae) in relation to the development of cornified scales

The epidermis of mammals is characterized by having a stratum granulosum that produces an orthokeratotic stratum corneum, different from the typical reptilian parakeratotic stratum. Nonetheless, some mammals show distinct degrees of parakeratosis in epidermal regions with few or no pilose follicles (e.g., areas subjacent to cornified scales). With respect to the epidermis and the development of cornified scales in the Dasypodidae, previous studies have supported the presence of a continuous stratum granulosum without any variations during ontogeny. This condition, in which the cornified scales develop without a loss of the stratum granulosum, was interpreted as primitive for eutherians. The present contribution expands the knowledge on the epidermis of Chaetophractus vellerosus in distinct ontogenetic stages in order to determine whether the cornified scales show the same developmental pattern as in other eutherians. The presence of a stratum granulosum in C. vellerosus neonates and its reduction in more advanced ontogenetic stages, in direct relationship with cornified scale development, supports the hypothesis that the partial parakeratosis in the xenarthran integument is secondary, as in other eutherians, and can be interpreted as a derived character state.     

Effect of Group Size on Activity Budgets of Colobus vellerosus in Ghana

Group size influences foraging efficiency in several primates. We examined the activity budgets of 3 groups of Geoffroy's pied colobus (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana to determine whether larger group size induces scramble competition. We studied 2 groups (B1; N = 7-8 and WW; N = 31-33) occupying slightly overlapping home ranges from August to November 2000. We observed the third group, B2 (N = 15-16), comprising B1 and 7 male invaders in the same home range as B1 from August to November 2001. By comparing groups belonging to the same population and occupying sligthly overlapping or similar home ranges, we were able to control, to a certain extent, for differences in food distribution. We recorded a total of 3353 scans, yielding 14,886 activity records, over 73 days. As with other black-and-white colobus, resting was their most common activity (59%). Intergroup comparisons suggest that time spent feeding, resting and moving did not vary in relation to group size. However, intragroup comparisons between the sexes show that females in the large group spent more time feeding than males did, whereas this was not the case in the small group, which suggests that scramble competition may be occurring among female Colobus vellerosus at BFMS. It is also possible that this may be due to greater nutritional requirements because of a higher proportion of infants in the large group. In fact, the proportion is quite similar between the two groups, lending support to the idea that females in the two groups had comparable nutritional demands due to lactation. This suggests that increased feeding in females in the large group was partly an effect of scramble competition. Group size and group composition also influenced the frequency of social behavior. There was more grooming in the large group, and it was performed mostly by females. The distribution of activities throughout the day was similar to the pattern reported for other black-and-white colobus.     

A contribution towards resolving the nomenclature of Citharexylum (Verbenaceae). II. Remarks on implicit typifications and lectotypification of names linked to Mesoamerican taxa

During ongoing taxonomic studies in Citharexylum (Verbenaceae), sixteen names were found in need of typification or typification remarks. As a result, six names (C. brachyanthum, C. danirae, C. flabellifolium, C. lucidum, C. rugendasii and C. tristachyum f. urbanii) are here lectotypified. Furthermore, remarks on implicit typifications by Harold Moldenke for another ten names (C. altamiranum, C. berlandieri, C. hexangulare, C. macradenium, C. ovatifolium, C. pauciflorum, C. pterocladum, C. recurvatum, C. schulzii and C. stenophyllum) are provided.