Positional behavior of Pan troglodytes in the Mahale Mountains and Gombe Stream National Parks, Tanzania.

The positional behavior of habituated adult chimpanzees and baboons was observed for 784 hr in a year-long study. Comparisons between species were made to establish the distinctiveness of chimpanzee positional behavior and habitat use. Brachiation (sensu stricto, i.e., hand-over-hand suspensory locomotion) was observed in low frequencies among chimpanzees, and its significance for chimpanzee anatomy is judged slight. Although no significant differences were found between sympatric baboons and chimpanzees in the proportion of time spent in the terminal branches, or in the mean diameter of weight-bearing strata, chimpanzees exhibited evidence of a terminal branch adaptation in that they, unlike baboons, used postures among smaller supporting strata different from those used among larger supports. Among chimpanzees, unimanual arm-hanging was most common among the smallest strata and was associated with smaller mean and median support diameter than other postures. Unimanual arm-hanging was the only common behavior among chimpanzees that usually involved complete abduction of the humerus. A number of behaviors often subsumed under the label "quadrumanous climbing" were distinguished in this study. Compared to baboons and other cercopithecoids, chimpanzees did not show increased frequencies of large-stratum vertical climbing, and their vertical climbing did not involve significant humeral abduction. Arm-hanging (i.e., unimanual suspension) and vertical climbing distinguish chimpanzee positional behavior from that of monkeys.     

Effects of body size and social context on the arboreal activities of lowland gorillas in the Central African Republic

The objectives of this 27 month study were to document the positional behaviors used by lowland gorillas (Gorilla gorilla gorilla) in the Central African Republic and to compare the effects of body size dimorphism on the use of arboreal substrates. During this study, despite their great size, all gorillas used trees regularly. Predictions concerning the relationship of body size to arboreal behavior were generally upheld. Small branch and suspensory activities were rare for silverbacks. Females used smaller and multiple substrates and suspensory postures more frequently than males. Although females foraged in the periphery of trees, males stayed close to the cores and rarely used terminal branches. In addition to body size, this study found that party size, social rank, and tree structure all influence an animal's substrate choice and subsequent positional activities. Lone males typically remained in the cores of trees where substrates are large. Group males may have been forced to use all parts of trees because others were present. Lone males used small crown trees which provided easy access to terminal branch foods. Males and females foraging together used larger trees (containing more feeding sites) than single sex groups. Female positional behavior may have been affected by the presence of males. When apart from males, females used the cores of trees and larger substrates more than when foraging with males. As habitat and social context both influence substrate use, they should be considered essential components of body-sized based interpretations of the behavior of fossil or extant species. © 1995 Wiley-Liss, Inc.     

Body size effects on vertical climbing among chimpanzees

I analyzed observations from a yearlong study of the positional behavior of Pan troglodytesat the Mahale Mountains National Park to determine whether there are detectable differences in behavior between large and small individuals. Analysis was complicated by a weak correlation between body size and social rank. To factor out rank effects, I performed two types of analyses, depending on the type of data: (1) multiple regressions or (2) comparisons of similarly ranked animals of different body size. With social rank effects accounted for, larger males fed lower in the canopy, fed on the ground more often, fed preferentially among food tree species with smaller adult heights, and climbed significantly less often than smaller males did. Contrary to expectation, large males utilized smaller weight-bearing structures than small males did. These results suggest that large males minimized climbing versus optimizing support diameters, perhaps because vertical climbing is disproportionally expensive for larger animals. The large body weight of chimpanzees compared with other primates suggests that minimizing altitude changes, and therefore vertical climbing, is an important consideration in budgeting daily energy expenditures.     

Body size effects on vertical climbing among chimpanzees

I analyzed observations from a yearlong study of the positional behavior of Pan troglodytesat the Mahale Mountains National Park to determine whether there are detectable differences in behavior between large and small individuals. Analysis was complicated by a weak correlation between body size and social rank. To factor out rank effects, I performed two types of analyses, depending on the type of data: (1) multiple regressions or (2) comparisons of similarly ranked animals of different body size. With social rank effects accounted for, larger males fed lower in the canopy, fed on the ground more often, fed preferentially among food tree species with smaller adult heights, and climbed significantly less often than smaller males did. Contrary to expectation, large males utilized smaller weight-bearing structures than small males did. These results suggest that large males minimized climbing versus optimizing support diameters, perhaps because vertical climbing is disproportionally expensive for larger animals. The large body weight of chimpanzees compared with other primates suggests that minimizing altitude changes, and therefore vertical climbing, is an important consideration in budgeting daily energy expenditures.     

Mechanical implications of chimpanzee positional behavior.

Mechanical hypotheses concerning the function of chimpanzee anatomical specializations are examined in light of recent positional behavior data. Arm-hanging was the only common chimpanzee positional behavior that required full abduction of the humerus, and vertical climbing was the only distinctive chimpanzee positional behavior that required forceful retraction of the humerus and flexion of the elbow. Some elements of the chimpanzee anatomy, including an abductible humerus, a broad thorax, a cone-shaped torso, and a long, narrow scapula, are hypothesized to be a coadapted functional complex that reduces muscle action and structural fatigue during arm-hanging. Large muscles that retract the humerus (latissimus dorsi and probably sternocostal pectoralis major and posterior deltoid) and flex the elbow (biceps brachii, probably brachialis and brachioradialis) are argued to be adaptations to vertical climbing alone. A large ulnar excursion of the manus and long, curved metacarpals and phalanges are interpreted as adaptations to gripping vertical weight-bearing structures during vertical climbing and arm-hanging. A short torso, an iliac origin of the latissimus dorsi, and large muscles for arm-raising (caudal serratus, teres minor, cranial trapezius, and probably anterior deltoid and clavicular pectoralis major) are interpreted as adaptations to both climbing and unimanual suspension.     

Sex differences in adult chimpanzee positional behavior: The influence of body size on locomotion and posture

Focal animal instantaneous sampling of adult male and female chimpanzee positional behavior was conducted during a 7-month study in the Tai Forest, Ivory Coast, in order to determine whether there are sex differences in the locomotion, posture, substrate use, and height preference of sexually dimorphic adult chimpanzees, and if so, whether these differences support predictions based on body size differences. Results indicate that as predicted, adult male and female chimpanzees differ in their arboreal locomotor behavior, with the larger males using less quadrupedalism and more climbing, scrambling, and aided bipedalism than females during feeding locomotion. There is a sex difference in height preference as well, with female chimpanzees consistently using more arboreal behavior than males, primarily during resting. Although it has been previously demonstrated that separate primate species of differing body size differ in locomotor and postural activities (Fleagle and Mittermeier, 1980; Crompton, 1984), this study clearly demonstrates that body size differences within a species can also be correlated with differences in locomotor behavior. These findings may influence how we interpret sex differences in body size of extinct species. © 1993 Wiley-Liss, Inc.     

Positional behavior in the Hominoidea

Quantitative studies on the positional behavior of members of the Hominoidea are compared in order (1) to identify consistencies across the superfamily, (2) to contrast ape positional behavior with that of Old World monkeys (forest-livingPapio anubis were chosen for study to reduce body size effects), and (3) to identify distinctive behaviors in each of the ape taxa. Differences in the way behaviors were sampled in the various studies necessitated considering posture and locomotion separately. Unimanual arm-hanging and vertical climbing were the most distinctive shared postural and locomotor modes among the apes (the gorilla excepted), constituting ≥5.0% and ≥4.9% of all behavior in each species. Arm-hanging and brachiation (sensu stricto) frequencies were the highest by far in hylobatids. Hand-foot hanging, bipedal posture, and clambering, an orthograde suspensory locomotion assisted by the hindlimbs, were more common in orangutans than in any other hominoid. Sitting and walking were observed in the highest frequencies in the African apes but were no more common than in the baboon. Relatively high frequencies of brachiation (sensu stricto) were reported for all apes except chimpanzees and gorillas. Brachiation and arm-hanging were kinematically different in apes and baboons, involving complete humeral abduction only in the former, whereas vertical climbing appeared to be kinematically similar in apes and baboons. It is concluded that the morphological specializations of the apes may be adaptations to (1) the unique physical demands of arm-hanging and (2) less kinematically distinct, but still quantitatively significant, frequencies of vertical climbing.     

Program of the fifty-sixth annual meeting of the American Association of Physical Anthropologists to be held in New York City Vista International Hotel April 2–5, 1987

Male orangutans (Pongo pygmaeus) weigh about twice as much as females. Observations in northern Sumatra of adults of both sexes (one male, three females) feeding in the same trees reveal the effects of body size dimorphism on feeding behavior: The male tended to use larger branches than the females, and to employ above-branch postures (sitting and standing) with greater frequency. The females employed suspensory under-branch postures more often. When feeding techniques were variable, the male tended to pull in branches to detach food with the mouth, whereas the female plucked more fruit by hand. By controlling for postcranial morphology and habitat structure, these results provide the first rigorous quantitative test of predictions about the effects of body size on primate positional behavior, and raise further questions about sexual diethism in feeding postural behavior of primates of varying absolute size.     

Positional behavior,selective use of habitat substrate and associated non-positional behavior in free-rangingColobus guereza (Rüppel, 1835)

Positional behavior (posture and locomotion), associated social/maintenance behaviors and selective utilization of available habitat structure were recorded for free-rangingColobus guereza occupying a remant montane forest near Limuru, Kenya. This research design focuses on relationships between anatomy, behavior, and the environment and provides a framework for interpreting adaptations inColobus. Quantitative analyses show daily activity patterns. These monkeys spend most of their time in above branch postural activities, primarily in sitting postures. Locomotion accounts for less than one-fifth of positional behavior. In both postural and locomotor sequences, the forelimbs may be placed above the shoulder complex, often in abducted positions. Suspensory positional behavior is rare. Most activity occurs within Zone II, flexible branches of medium diameter and less than 45° in grade. Selective use of the substrate and preferred positional modes are related to variables such as social/maintenance activities, weight distribution of the link system and substrate configuration. These data suggest that, (a) black and white colobus do not exhibit behavior associated with so-called “semibrachiation” and (b) bio-behavioral adaptations involving foraging and feeding strategies may account, in part, for observed morphological differences among Old World monkeys.     

How feeding competition determines female chimpanzee gregariousness and ranging in the Taï National Park, Côte d'Ivoire

Socioecological theory suggests that feeding competition shapes female social relationships. Chimpanzees (Pan troglodytes) live in fission–fusion societies that allow them to react flexibly to increased feeding competition by forming smaller foraging parties when food is scarce. In chimpanzees at Gombe and Kibale, female dominance rank can crucially influence feeding competition and reproductive success as high‐ranking females monopolize core areas of relatively high quality, are more gregarious, and have higher body mass and reproductive success than low‐ranking females. Chimpanzee females in Taï National Park do not monopolize core areas; they use the entire territory as do the males of their community and are highly gregarious. Although female chimpanzees in Taï generally exhibit a linear dominance hierarchy benefits of high rank are currently not well understood. We used a multivariate analysis of long‐term data from two Taï chimpanzee communities to test whether high‐ranking females (1) increase gregariousness and (2) minimize their travel costs. We found that high‐ranking females were more gregarious than low‐rankers only when food was scarce. During periods of food scarcity, high rank allowed females to enjoy benefits of gregariousness, while low‐ranking females strongly decreased their gregariousness. High‐ranking females traveled more than low‐ranking females, suggesting that low‐rankers might follow a strategy to minimize energy expenditure. Our results suggest that, in contrast to other chimpanzee populations and depending on the prevailing ecological conditions, female chimpanzees at Taï respond differently to varying levels of feeding competition. Care needs to be taken before generalizing results found in any one chimpanzee population to the species level. Am. J. Primatol. 73:305–313, 2011. © 2010 Wiley‐Liss, Inc.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. II. Influence of Male Rank and Possible Competition for Partners

Allogrooming contributes to the development and maintenance of social relationships, including those that involve alliances, in many primate species. Variation in relatedness, dominance rank, and other factors can produce variation in the value of others as grooming partners. Several models have been developed to account for variation in the distribution of grooming in relation to dominance ranks. These start from the premise that individuals are attracted to high-ranking partners, but time limits, direct competition, and prior grooming engagement between high-ranking individuals can constrain access to them. Sambrook et al. (1995) formalized some of these models and showed the importance of taking group size variation into account when assessing them. Chimpanzees form multimale communities in which males are the philopatric sex. Males commonly associate and groom with each other; they also form dominance hierarchies and form alliances that influence dominance ranks and mating success. Both male rank and the rank distance between partners are significantly correlated with the distribution of grooming between males in an extremely large chimpanzee community at Ngogo, Kibale National Park, Uganda, that has more males than any other known community. High-ranking males had more grooming partners than mid- or low-ranking males. Grooming predominantly went up the dominance hierarchy, but was also concentrated among males that were close in rank. Rank and rank distance apparently both affected grooming independently of reciprocity in grooming and independently of the frequency with which males associated in temporary parties. However, the data do not clearly indicate how constraints on access to partners might have operated. Published data from a smaller chimpanzee community at Mahale show no rank or rank distance effect on male grooming. These results and earlier, conflicting findings on the association between dominance rank and grooming in male chimpanzees indicate that variation in group size, i.e., the number of males per community, probably influences the strength of any such effects, as happens for grooming between females in several cercopithecine species. Data on coalitions at Ngogo support the argument that high-ranking males are valuable social partners, and similarity in strategies of alliance formation may influence the distribution of grooming.     

Evaluation of various parts of the paw paw tree, Asimina triloba (Annonaceae), as commercial sources of the pesticidal annonaceous acetogenins.

Various plant parts of the paw paw tree (Asimina triloba Dunal, Annonaceae) were extracted and partitioned to concentrate the mixture of acetogenins into a standardized pesticidal extract (F005). A bioassay with brine shrimp larvae (Artemia salina Leach) was used to determine the relative potencies of the various extracts. The small twigs (0-0.5 cm diameter) yielded the most potent extract (LC50 = 0.04 ppm); the stem wood (LC50 = 4.9 ppm) and leaves (LC50 = 53.7 ppm) yielded the poorest activities. The unripe fruits, seeds, root wood, root bark, and stem bark were notably potent and, generally, yielded > 2% of their dry weight as F005. The smaller diameter stems were more potent than the larger stems. We conclude that, by pollarding the trees, the entire twigs and small branches of paw paw could be processed to produce a potent acetogenin mixture; this biomass could be made available in quantities needed for commercialization of the pesticidal product and could be renewable through regrowth from the parent trunk and larger branches.     

The allometry of the weight of fruit on trees and shrubs in Barbados

Summary Branch sampling of branch diameter and fruit crop on 22 species of Barbadian trees and shrubs provided sufficient data to build regressions between plant size and fruit crop weight. Orchard plants bear much more fruit than wild, feral or garden plants of similar size, but this difference disappears in multiple regression of fruit crop weight (F in g, fresh mass) on branch or stem diameter (D in cm) and individual fruit weight (W in g): F=22D^1.2 W^0.57. This explains 89% of the variation in F and successfully predicts crop weight for wild tropical and temperate trees and shrubs, but underestimated the crops on commercial, temperate, fruit trees by an order of magnitude. Comparisons of crop weight for feral, wild, and garden plants (F_f) using a simple regression F_f=47D^1.9 show that crop weight is a minor load relative to branch weight for larger branches. Although fruit crops represent a declining proportion of total plant weight as plants become larger, the crops become larger relative to leaf and twig weight and in this sense, reproductive investment increases in larger plants. Finally, our equations, combined with the self-thinning rule, suggest that stands of large species of fruit plants produce more fruit per unit of land area than stands of small ones.     

Life history and locomotion in Cebus capucinus and Alouatta palliata

As an individual matures from birth to adulthood, many factors may influence the positional repertoire. The biological and behavioral changes that accompany a growing individual are expected to influence foraging strategy, social status and interaction, diet, predator avoidance strategies, and ultimately positional behavior as a behavioral link between anatomy and the environment. In this work, positional behavior is considered as an important feature of life history in juvenile and adult white‐faced capuchins (Cebus capucinus) and mantled howling monkeys (Alouatta palliata) inhabiting the same tropical forest in Costa Rica. During growth and development ontogenetic changes in body size, limb proportions, and motor skills are likely to influence locomotion and posture through the arboreal canopy. I collected data on positional behavior, activity, branch size, branch angle, and crown location during a 12‐month period at Estación Biológica La Suerte in northeastern Costa Rica. Life history timing and differences in rates of growth did not predictably influence the development of adultlike positional behaviors in Cebus and Alouatta. Young Cebus resembled the adult pattern of positional behavior by 6 months of age while howlers exhibited significant differences in several positional behavior categories through 24 months of age. The positional repertoire of both species revealed similarities in the types of modes used during feed/forage and travel in juveniles and adults. Data presented here suggest that the environment exerts different pressures on growing Cebus and Alouatta that may relate to diet, energy expenditure, foraging skill, and/or social learning. Am J Phys Anthropol, 2009 © 2009 Wiley‐Liss, Inc.     

Intermale affiliative behavior in ringtailed lemurs (Lemur catta) at the Beza-Mahafaly reserve, Madagascar

Intermale affiliative behavior was studied in three groups of naturally occurring ringtailed lemurs over a one-year period. The adult males in the sample exhibited affiliative behavior with all other males in their social groups; but affiliative behavior between some male-male dyads occurred markedly more often than between others. These dyads are called “preferred partnerships.” The formation of preferred partnerships did not appear to be influenced by male dominance rank or age-class. Furthermore, these partnerships were of a short-term nature, and none persisted throughout the entire study period. The following factors may explain the absence of rank or age effects, and the brevity of preferred partnerships: (1) frequent fluctuation in the male dominance hierarchy in each study group; (2) the fact that higher-ranking males may not offer specific benefits to lower-ranking males; and (3) the fact that male dispersal affected the male membership of each group over the 12-month study period. The focal males engaged in significantly more affiliative behavior during the lactation period compared with the other reproductive seasons. Migration partners may offer each other predator protection during the transfer process, and although males that transfer together do not engage in alliance or coalition behavior towards resident males, males transferring in pairs or threesomes are in a better position to spot and defend themselves against attack by resident males. Migration partners also provide social contact for each other while in the process of immigration.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. I. Partner Number and Diversity and Grooming Reciprocity

Allogrooming serves many social functions in primates. Grooming can help individuals to service social relationships generally, sometimes reciprocally, and may be particularly important in the development and maintenance of alliances. However, time constraints limit the number of partners with whom one individual can groom enough to maintain cooperative relationships. As a result, the size of its grooming network may reach an asymptote as the size of its group increases, and it may distribute its grooming less equally among potential partners. Chimpanzees live in multimale, fission-fusion communities; males are philopatric, and commonly associate and groom with each other. Males form within-community alliances that influence dominance rank and access to mates, and allies groom with each other regularly; males also cooperate in aggression between communities. The chimpanzee community at Ngogo, in Kibale National Park, Uganda, is unusually large and has more males than any other known community. Field data show that adult Ngogo males groomed far more with other adult males than with females or with adolescent males, in contrast to a previous report (Ghiglieri, 1984). Adolescent males groomed adults much more than the reverse; males groomed and were groomed by females about equally. Individual males groomed mostly with a small number of other males. On average, males at Ngogo had only slightly more male grooming partners overall and had the same number of important partners as those of males in a much smaller community in the Mahale National Park, Tanzania, and they distributed their grooming less equitably. These results fit those expected if limits on available grooming time cause males to have a loyalty problem as the number of potential grooming and alliance partners increases. Despite differences in the extent and equitability of their grooming networks, males at both Ngogo and Mahale showed reciprocity in grooming. Grooming reciprocity has been demonstrated for captive chimpanzee males, but the Ngogo findings are the first demonstrations of reciprocity in wild communities.     

Cognitive research in zoo-housed chimpanzees: influence of personality and impact on welfare

We monitored chimpanzee welfare during the introduction of on-exhibit cognitive research training and testing, as measured by behavior and interest in such training, and related individual variation to personality assessments. We observed 11 chimpanzees (six males; five females) over a 16-month period and compared their behavior across three conditions: (1) Baseline (nontraining/research situations) and (2) an on-going, off-exhibit program of Husbandry Training and (3) Research Pod Activities, on-exhibit, group training for cognitive testing. There was considerable individual variation in their interest levels during research sessions; females and those scoring higher for Openness were present more frequently (including those who actively participated and those who observed others participating), but interest did not vary in relation to rates of self-directed behaviors (SDBs), rank, or the level of social disruptions within the group (i.e. large-scale displays or fights). The frequency of SDBs was predicted by the Neuroticism personality factor, but did not differ across baseline and training contexts, indicating that these activities do not negatively impact welfare. We also explored vigilance as an indicator of social uncertainty, but social monitoring did not differ in relation to either social context or rank. Finally, we explored how the specific characteristics of the research context impacted on SDBs; namely, social context, reward contingency, and visual access to keepers. SDBs increased only when visual access to keepers was restricted, suggesting that visual contact reduced uncertainty in novel training contexts. Overall, the introduction of a cognitive research program did not compromise welfare, and the chimpanzees' repeated interest and willingness to participate suggests that the research was enriching.     

Positional behavior of long-tailed macaques (Macaca fascicularis) in northern Sumatra

Although the majority of extant primates are described as "quadrupedal," there is little information available from natural habitats on the locomotor and postural behavior of arboreal primate quadrupeds that are not specialized for leaping. To clarify varieties of quadrupedal movement, a quantitative field study of the positional behavior of a highly arboreal cercopithecine, Macaca fascicularis, was conducted in northern Sumatra. At least 70% of locomotion in travel, foraging, and feeding was movement along continuous substrates by quadrupedalism and vertical climbing. Another 14-25% of locomotion was across substrates by pronograde clambering and vertical clambering. The highest frequency of clambering occurred in foraging for insects, and on the average smaller substrates were used in clambering than during quadrupedal movement. All postural behavior during foraging and feeding was above-substrate, largely sitting. Locomotion across substrates requires grasping branches of diverse orientations, sometimes displaced away from the animal's body. The relatively low frequency of across-substrate locomotion appears consistent with published analyses of cercopithecoid postcranial morphology, indicating specialization for stability of limb joints and use of limbs in parasagittal movements, but confirmation of this association awaits interspecific comparisons that make the distinction between along- and across-substrate forms of locomotion. It is suggested that pronograde clambering as defined in this study was likely a positional mode of considerable importance in the repertoire of Proconsul africanus and is a plausible early stage in the evolution of later hominoid morphology and locomotor behavior.     

Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela

I observed leaping behavior in the white-faced saki (Pithecia pithecia) and the black-bearded saki (Chiropotes satanas satanas) for 15 and 10 months, respectively, as part of a larger study of positional behavior in the tribe Pitheciini. I used focal animal instantaneous sampling to observe the two species on separate islands in their natural habitat at Guri Lake, Venezuela. Leaping behavior correlates with patterns of forest use and body size, and differences between the species relate more to habitat preferences than to habitat differences per se. Pithecia usually chose vertical or highly angled supports of lower tree portions for take-off and landing, and took off from a stationary posture. Chiropotes took off from the main crown or terminal branches, gaining momentum from locomotor movement before performing a leaping take-off. Pithecia's vertical body orientation and longer leap distance allowed it to assume a mid-flight tuck to prepare for a hindlimb-first landing onto a solid support, and to absorb landing forces with its relatively longer hindlimbs. Chiropotes remained more pronograde throughout its leaps, and minimized landing forces by landing on all four limbs onto numerous flexible supports in the terminal branches. The smaller-bodied P. pithecia is specialized for vertical clinging and leaping, and exhibits behavioral and morphological parallels with other vertical clingers and leapers. The larger C. satanas is a generalized leaper that lacks morphological specializations for leaping. Pithecia's use of solid supports in the lower tree portions allows it to move quietly through the forest-one of a suite of behaviors related to predator avoidance. This example of variation within one behavioral category has implications for devising locomotor classifications and interpreting fossil remains.     

不同密度下凤仙花重要形态性状与花朵数的关系

对30cm×30cm和50cm×50cm种植密度下的凤仙花(Impatiens balsamina L.)的地径、株高、冠幅、叶面积、花期、花径、一级分枝数、二级分枝数、一级分枝部位高和二级分枝部位高10个重要形态性状与花朵数的关系差异进行了相关性及通径分析,为凤仙花栽培与性状选择提供科学基础。结果表明:30cm×30cm种植的凤仙花的一级分枝数、二级分枝部位高都与花朵数呈显著正相关,地径、冠幅、二级分枝数与花朵数呈极显著正相关,对花朵数的直接效应大小顺序为:二级分枝数>一级分枝数>花径>花期;50cm×50cm种植的凤仙花,其地径、冠幅和二级分枝数都与花产量呈显著正相关,对花朵数的直接效应大小顺序为:二级分枝数>冠幅。二级分枝数对两种密度栽培的凤仙花花产量都起着重要作用。不同种植间距下凤仙花采用了不同的适应方式以获得最大花朵数目,进而获得最大数量的种子,获得较多的繁殖机会。在30cm×30cm下,是通过获得较多一级分枝数;在50cm×50cm下,是通过增加冠幅来产生较大花产量,而且与前者相比,植株茎秆粗壮,对霜霉病的抗性也强,因此,在栽培中,应以适合的低密度栽培比较好。