Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

The Effects of Primates and Squirrels on Seed Survival of a Canopy Tree,Afzelia quanzensis,in Arabuko‐Sokoke Forest,Kenya1

I examined the fate of seeds from ten focal trees of Afzelia quanzensis (Leguminosae), a canopy tree in the Arabuko‐Sokoke, Kenya. The study was conducted for one fruiting season, between August 1990 and July 1991. Yellow baboons (Papio cynocephalus), Syke's monkeys (Cercopithecus albogularis), sun squirrels (Heliosciurus rufobrachium), and bush squirrels (Paraxerus palliatus) were all observed to interact with A. quanzensis seeds at various stages of pod development. Baboons and squirrels consumed high percentages of seeds when they were still immature, but the seeds were still unavailable to Syke's monkeys at this stage. Baboons bit open the hard green pods and squirrels gnawed through the pods to extract the immature seeds (hereafter referred to as seed predation), but monkeys were unable to open the pods. Upon maturity, the pods opened slightly, revealing red arils that were sought by baboons, monkeys, and squirrels. Monkeys removed the highest percentage of mature seeds from these pods. These mammal dispersers ate the arils from the mature seeds and discarded the viable part that germinates (hereafter referred to as seed dispersal). My data indicate that baboons and squirrels are seed predators while monkeys are seed dispersers of A. quanzensis.     

Why do larvae of Utetheisa ornatrix penetrate and feed in pods of Crotalaria species? Larval performance vs. chemical and physical constraints

Larvae of Utetheisa ornatrix (L.)(Lepidoptera: Arctiidae) are found mainly inside unripe pods of several alkaloid‐bearing Crotalaria (Fabaceae) species. Although eggs are laid on the leaves, the larvae are usually found feeding on unripe seeds in the pods. In this work, we investigated the selective pressures that could explain why U. ornatrix larvae feed primarily on unripe pods with seeds and not on leaves. Our results showed that larval survivorship in the laboratory was unaffected by feeding on leaves or unripe seeds, and that larval development up to the pupal stage was better in larvae that fed on unripe seeds, although perforating unripe pods to reach seeds was costly in terms of survivorship. Females were also heavier when fed on unripe seeds, but there was no significant difference in the fecundity of females fed either of the two diets. Feeding on unripe seeds in pods had other benefits for U. ornatrix, including a lower predation rate for larvae that fed inside compared to larvae that fed outside the pods. Similarly, adults derived from larvae that fed on unripe seeds were preyed upon less frequently by the orb‐weaving spider Nephila clavipes than were adults that fed on leaves. The latter benefit may be closely related to the high concentration of pyrrolizidine alkaloids in unripe seeds, which is about five times more than in leaves. These alkaloids are sequestered by the larvae and transferred to adults, which then become chemically protected. However, this chemical defence does not protect the larvae against ants such as Ectatomma quadridens and Camponotus crassus. Pods with unripe seeds that confer physical protection to larvae and pyrrolizidine alkaloids that confer chemical protection to adults limit the use of leaves by U. ornatrix larvae.     

Intensity of pre‐dispersal seed predation in the invasive legume Leucaena leucocephala is limited by the duration of pod retention

Abstract The intensity of seed predation the invasive tropical legume Leucaena leucocephala by the bruchid Acanthoscelides macropthalmus was investigated in south‐eastern Queensland, Australia. The number of seeds damaged by A. macropthalmus as a proportion of total seeds available was found to increase the longer the pods remained on the tree. Seed predation ranged from a mean of 10.75% of seeds on pods that remained on the plant for 1 month and increased to 53.54% for pods that remained of the plant for 4 months. The low bruchid populations at high pod densities results in ‘predator satiation’. However, pods dehisce over time and the proportion of pods available over time to the bruchid correspondingly declines. By the time bruchid densities build up, most pods have dehisced and the seeds consequently escape predation. As a result the number of seeds lost to bruchid damage increases only marginally over time. Despite the levels of seed predation observed over the course of the study, the number of seeds in the soil seedbank almost doubled over time increasing from 8.5 seeds m^?3 to 15.5 seeds m^?3 over a 4‐month period. Levels of seed predation and addition of seeds to the soil seedbank were not correlated. The taxonomic (subspecies) status and apparency of host plants as measured by plant and patch traits (average plant height, density of podding plants and patch size) did not influence levels of seed predation. Pre‐dispersal seed predation studies need to take into account the pod/seed retention behaviour of the plant. The ability of the bruchid to regulate the invasiveness of Leucaena through influencing its demography is likely to be diminished if the insect populations cannot increase rapidly enough to use the seeds before pod dehiscence.     

Ecophysiological aspects of bruchid reproduction. I. The influence of pod maturity and seeds of Phaseolus vulgaris and the influence of insemination on the reproductive activity ofZabrotes subfasciatus

ABSTRACT.

• 1 Economically important bruchid beetles like Zabrotes sub-fasciatus (Boh.) are able to reproduce both in the field before the legume crop harvest – whilst the seeds are still enclosed within pods – and on seeds in storage sites.
• 2 To take into account this reproductive feature, the influence of pods at different stages of maturity and seeds of Phaseolus vulgaris (Lin.) together with the influence of insemination on the reproductive activity of Zabrotes subfasciatus were examined.
• 3 Female fecundity, insemination and ovarian production were studied under laboratory conditions approximating as closely as possible to the thermal and photoperiodic regimes of the Costa Rican region from which the experimental strain originated.
• 4 The three stages of pod maturity used in experiments (fresh green, fresh yellow and dry yellow pods, all non-dehiscent) induced copulations and significantly stimulated oogenesis. Fecundities of females in the presence of pods were low and did not differ from controls.
• 5 The influence of bean seeds on female reproductive activity was more pronounced than that of pods. Contact with seeds led to a marked increase in the number of sexual meetings, ovarian production and egg-laying activity. Under these conditions, eggs are laid directly on the larvae's food.
• 6 Insemination alone stimulated oogenesis but did not induce egg-laying.
• 7 Possible additive or synergistic interactions between host plant information perceived by females, the effects of egg-laying and insemination are suggested to explain the observed results.
• 8 The adaptive value of this form of reproductive regulation by host plant seeds and pods is discussed in terms of the natural history and oviposition behaviour of Zabrotes subfasciatus.

     

Chemical composition of pods and seeds of Dichrostachys cinerea as a basis of seed dispersal

The occurrence of very few seeds of Dichrostachys cinerea in the soil seed bank prompted the present study as to why animal browsers and seed predators should select the infructescence of this plant. The preferential and extensive browsing of this indehiscent legume is apparently not due to any exceptional nutritive value of the pods and seeds. However, it is probably the strong rich aroma of the infructescence which initially attracts animals to this food source. The structure of the infructescence, ensures that upon location a large number of pods and seeds are consumed. Ingestion of the pods will provide fibre, lipid, fatty acids, carbohydrates, amino acids, nitrogen and protein.     

Seed abortion in Pongamia pinnata (Fabaceae)

In Pongamia pinnata only one of the two ovules develops into a seed in most of the pods. Since pollen was not found to be limiting and reduced fertilization could not completely explain the observed frequency of seed abortion, it implied an effect of postfertilization factors. Aqueous extracts of developing seeds and maternal tissue (placenta) did not influence abortion in vitro, suggesting that abortion may not be mediated by a chemical. Experimental uptake of ¹⁴C sucrose in vitro indicated that both the stigmatic and the peduncular seed have similar inherent capacities of drawing resources, but the peduncular seed is deprived of resources in the presence of the stigmatic seed. This deprivation of the peduncular seed could be offset by supplying an excess of hormones leading to the subsequent formation of two seeds in a pod. The prevalence of single-seeded pods in P. pinnata seems therefore to be a result of competition between the two seeds for maternal resources. The evolutionary significance of single-seeded pods in P. pinnata is discussed with respect to possible dispersal advantage enjoyed by such pods.     

Multiple paternity in common bean (Phaseolus vulgaris L., Fabaceae)

We report on two field experiments that were conducted in 1991 and 1992 at the South Coast Extension and Research Center, Irvine, CA, to study the incidence of multiple paternity in the common bean (Phaseolus vulgaris L.). Hypocotyl color and shikimate dehydrogenase (Skdh) isozymes were used as genetic markers. The white-seeded cultivar ‘Ferry Morse 53’ (FM 53) was used as the female parent. This cultivar is homozygous recessive (pp) for hypocotyl color. The pollen source parents were three homozygous dominant (PP) purple-hypocotyled, black-seeded cultivars. Three cultivars, ‘ICA Pijao,’ G4459, and the maternal parent FM 53, are of Mesoamerican origin and homozygous for the fast (F) allele at the Skdh locus. The other cultivar, Black Valentine, is of Andean origin and is homozygous for the slow (S) allele at the Skdh locus. Overall, 6 125 pods were obtained from 57 and 111 plants harvested individually in 1991 and 1992, respectively. All progeny, 28938 seeds, were scored for hypocotyl color at the seedling stage. The purple-hypocotyled seedlings were genotyped for the Skdh locus to identify their pollen parents. Multiple paternity was identified in all the pods with hybrid seeds (i.e., those of intercultivar crosses) at 5.8% and 8.1% in 1991 and 1992, respectively. All multiply sired pods produced both nonhybrid and hybrid seeds. As many as three successful fathers per pod were identified, but the number of markers limited measuring higher levels of multiple paternity. Most multiply sired pods (≈70%) were filled by nonhybrid seeds plus a single hybrid seed. Ovule position effect within multiply sired pods was inferred from the nonrandom distribution of hybrid seeds within a pod. On average, hybrid seeds occurred more frequently in ovules in position 7 (most basal) and in position 1 (most stylar) than in ovules in the middle positions of the pod.     

Stigmatic inhibition of pollen grain germination-its implication for frequency distribution of seed number in pods of Leucaena leucocephala (Lam) de Wit

Summary Seed number per pod is negatively skewed in Leucaena leucocephala. Post-fertilisation abortion of pods does not completely explain the skewed distribution. It is shown that a pre-fertilisation regulation effectively restricts the formation of pods with less than a certain number of seeds. The regulation is rendered through the inhibition of pollen grain germination by the stigmatic fluid when the pollen grain is less than a certain critical load in the stigma. Such a control is shown to have implications in the evolution of pollen package units, generation of male gametophytic competition and the economy of packing seeds in pods.     

Injury to oilseed rape caused by mirid bugs (Lygus) (Heteroptera: Miridae) and its effect on seed production

Injury by Lygus spp. to oilseed rape, Brassica napus L. and Brassica campestris L., was assessed based on laboratory and field studies in Alberta, Canada. The visible injuries consisted of lesions on the surfaces of stems, buds, flowers and pods similar to those described for other crops. They caused buds and flowers to abscise and seeds to collapse, and reduced the weight of healthy seeds produced per pod. The plants compensated for bud loss so that no net reduction in the number of pods occurred. However, in some situations the damage to buds resulted in a reduced seed yield that increased with the amount of injury. The plants also compensated for flower loss so that no net reduction in the number of pods occurred, but seed yield declined as injury increased. Plants did not compensate for seeds that collapsed as a result of lygus feeding. The feeding activity of lygus bugs reduced seed yield in oilseed rape.     

DEMOGRAPHIC ASPECTS OF FLOWER AND FRUIT PRODUCTION IN BEAN PLANTS,PHASEOLUS VULGARIS L.

In this paper we assess the relative contributions to total pod yield of cohorts of flowers of known age. Bean plant (Phaseolus vulgaris L.) growth was monitored and 'births' of individual flowers were recorded and their fates followed. For every cohort of flowers an array of yield measures was calculated, including the mean and total cohort dry weight of fruits, and the numbers of viable and aborted seeds and completely undeveloped ovules. When mature, pods were removed from half of the plants. Plants whose mature pods were removed produced significantly greater numbers of flowers, ovules and viable seeds and a greater total weight of pods than did untreated plants. However, the proportion of ovules giving rise to viable seeds and to aborted seeds and undeveloped ovules was the same in plants of both groups. Approximately 50% of ovules of marketable pods, in all plants, yielded viable seeds. The 12 cohorts of flowers contributed markedly different amounts to plant yield. Early and later cohorts contributed much less than did middle cohorts in terms of marketable pod dry weight and numbers of fully developed seeds, aborted seeds, and undeveloped ovules. Peak productivity was therefore obtained from the middle phase of flowering in these annual bean plants. The lower pod yield of later cohorts is interpreted as a result of competition for limited resources between maturing pods and new flowers. We consider possible causes for the various fates of flowers and ovules and discuss flowers as plant modules suitable for demographic examination.     

Effect ofAspergillus parasiticus soil inoculum on invasion of peanut seeds

Environmental control plots adjusted to late season drought and elevated soil temperatures where inoculated at peanut planting with low and high levels of conidia, sclerotia, and mycelium from a brown conidial mutant ofAspergillus parasiticus. Percentage infection of peanut seeds from undamaged pods was greatest for the subplot containing the high sclerotial inoculum (15/cm² soil surface). Sclerotia did not germinate sporogenically and may have invaded seeds through mycelium. In contrast, the mycelial inoculum (colonized peanut seed particles) released large numbers of conidia into soil. Soil conidial populations of brownA. parasiticus from treatments with conidia and mycelium were positively correlated with the incidence of seed infection in undamaged pods. The ratio ofA. flavus to wild-typeA. parasiticus in soil shifted from 7:3 to 1:1 in the uninoculated subplot after instigation of drought, whereas in all subplots treated with brownA. parasiticus, the ratio of the two species became approximately 8:2. Despite high levels of brownA. parasiticus populations in soil, nativeA. flavus often dominated peanut seeds, suggesting that it is a more aggressive species. Sclerotia of wild-typeA. parasiticus formed infrequently on preharvest peanut seeds from insect-damaged pods.     

Effects of artificial defoliation on reproductive allocation in the common vetch,Vicia sativa (Fabaceae: Papilionoideae)

Previous field experiments have shown that, for the annual herb Vicia sativa, leaf area lost to herbivory results in reduced number of fruit (pods) produced per plant as well as fewer seeds per pod. We conducted a controlled garden experiment to determine the precise relationship between level of defoliation and various measures of maternal fitness through fruit and seed. We employed manual clipping of individual leaflets of newly produced leaves to 25% increments of damage (from 0% to 100%) over the entire period of development and flowering of these annual herbs, harvesting pods when filled but not dehisced. We found significant reduction in number of pods, number of seeds, total seed mass, and individual seed mass with leaf area lost. Even with the highest levels of defoliation over the life of the individual, plants still produced a substantial number of pods and seeds. Seeds produced by plants in all treatment groups showed similar percentages of germination and time to germination.     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

Seed production and seed predation in a patchy and time-varying environment. Dynamics of a milkweed — tephritid fly system

Summary Vincetoxicum hirundinaria is a longlived perennial herb. Its pod production was measured during seven years in about 40 small isolated patches in two study areas in southeastern Sweden. Total pod production in the two areas varied synchronously and up to about 200 times between years. This variation was mainly due to drought conditions as determined by the interplay of weather (sunshine h) and site characteristics.Populations of a univoltine tephritid fly, euphranta connexa, whose larva feeds on V. hirundinaria seeds in the maturing pods, were monitored for five and seven years in the two study areas respectively. Larval fly populations in the two areas varied 30 and 50-fold between years, with highs in years when pods were abundant and lows when pods were scarce. The percentage of pods attacked by E. connexa, however, varied in the opposite direction, with very high attack rates (about 100%) when pods were scarce and low rates (down to 10–20%) when pods were common. Thus the temporal tracking of food resources by E. connexa was poor, resulting in yearly variations in the amount of unattacked pods (and seeds) being about 2,00-fold.Many host plant patches were small and totally without pods in some years, causing frequent local extinctions of E. connexa. Patches were, however, readily colonized in later years when pods appeared again. Low overall attack rates in certain years were thus only to a minor extent explainable by host plant patches being uncolonized by the fly. The efficient spatial tracking of resources by the fly population allows this population system to be analyzed largely in terms of its temporal dynamics.Although E. connexa populations often exploit only a minor part of their potential larval resources, the rate of population change was closely correlated with these resources expressed on a per capita basis. Natural enemies only have negligible effects on E. connexa population trends. In this population system the herbivore (seed predator) population is thus controlled by its food resources, but weather-imposed temporal variations in resource set are too large for the herbivore population to closely track its resource base. This temporal tracking inertia has important implications for the long-term production of healthy seeds.     

Diversification of seed arrangement induced by ovule rotation and septum formation in Leguminosae

The plants of Leguminosae, which comprises 731 genera, bear monocarpellary flowers. The fruit, known as a pod, generally contains a row of seeds, and the internal structure appears to be simple. In Leguminosae, the seeds are generally arranged longitudinally along the suture of the pod but, in a few genera, namely, Cassia, Indigofera, and Senna, they are arranged transversely. Transverse seed arrangement facilitates the accommodation of a considerable number of seeds and has been considered to be established by rotation of seeds during development. We studied the development both these patterns of seed arrangement, i.e., longitudinal and transverse, in Senna and Indigofera in order to elucidate the structural changes occurring in ovaries (or pods) and determine the forces responsible for rotation of ovules (or seeds) by dissecting ovaries and pods at different developmental stages. Our results showed that at an early developmental stage the ovules were obliquely oriented or transversely arranged; later, the direction of their growth was restricted by the ovary walls or neighboring ovules, and therefore the obliquely oriented ovules rotated inward or outward. Thus, it was clarified that the processes involved in the formation of the internal structure of pods are regulated in a complex manner. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

CHANGES IN WEIGHT,NITROGEN ACCUMULATION,RESPIRATION AND PHOTOSYNTHESIS DURING GROWTH AND DEVELOPMENT OF SEEDS AND PODS OF PHASEOLUS VULGARIS

The growth and development of pods and seeds of Phaseolus vulgaris are described by the following parameters: fresh and dry weight, length, ratio of fresh weight to length and nitrogen content. Pod and seed development are independent and sequential. Pod photosynthesis contributes only marginally to pod development, but may be significant in refixing CO₂ released in respiration. Pods and seeds do not seem to act simply as competing sinks. Nitrogen accumulation in the seeds begins only when it has ceased in the pods. Ways in which regulation may be achieved are discussed. Manipulation of nitrogen fertilization to enhance seed growth is suggested.     

Discrimination Against 13CO2 in Leaves,Pod Walls,and Seeds of Water-stressed Chickpea

The rate of photosynthesis (P _N) in leaves and pods as well as carbon isotope content in leaves, pod walls, and seeds was measured in well-watered (WW) and water-stressed (WS) chickpea plants. The P _N, on an area basis, was negligible in pods compared to leaves and was reduced by water stress (by 26%) only in leaves. WS pod walls and seeds discriminated less against ¹³CO₂ than did the controls. This response was not observed for leaves as is usually the case. Pod walls and seeds discriminated less against ¹³CO₂ than did leaves in both WW and WS plants. Measurement of carbon isotope composition in pods may be a more sensitive tool for assessing the impact of water stress on long-term assimilation than is the instantaneous measurement of gas exchange rates.     

Invasive on invasive – On the feeding ecology of the Ring-necked Parakeet,Psittacula krameri,in Oman (Aves: Psittacidae)

The Ring-necked Parakeet, Psittacula krameri, an invasive bird species in Oman was found to be feeding on the immature green pods of another invasive tree species, Prosopis juliflora, commonly known as Mesquite. This parakeet strips the pods and consumes the soft seeds within. The green pods collected from the base of trees were a mixture of stripped pods showing various levels of feeding damage and un-stripped pods without any damage. The percentage of stripped pods was significantly higher than that of un-stripped pods. The mean percentage number of pods in the five levels of feeding damage (10, 25, 50, 75 and 100%) significantly differed from each other. The un-stripped and undamaged pods indicate wasteful feeding activity. The comparison of the five levels of feeding damage (10, 25, 50, 75 and 100%) and the undamaged condition (0%) to the mean number of pods as percentages suggests that wasteful feeding is not a random event; the reasons for this relationship are discussed.     

DISPERSAL OF HETEROMORPHIC SEEDS IN AMPHICARPAEA BRACTEATA (FABACEAE)

The dispersal of the chasmogamous, aerial cleistogamous, and subterranean cleistogamous seeds of Amphicarpaea bracteata was examined. The chasmogamous and aerial cleistogamous seeds are ballistically dispersed. Chasmogamous seeds were dispersed farther than the aerial cleistogamous seeds due to the height advantage of the chasmogamous pods. There was no difference in the firing angle or the initial velocity of the seeds discharged from the two aerial pod types. The subterranean cleistogamous seeds are “dispersed” the shortest distance by the elongation of runners. Differential dispersal of the three types of seeds may be a factor involved in the evolution of cleistogamy in A. bracteata.