Consequences of pre-dispersal damage by insects for the dispersal and recruitment of mangroves

Herbivores may enhance plant recruitment, but such positive interactions may be overlooked in favour of obvious negative effects of herbivory on propagules. My objective was to determine whether larval insects that feed and develop within fruit of the mangrove Avicennia marina act as mutualist herbivores by increasing the dispersal of propagules without affecting their viability and emerging successfully as adults following dispersal of the propagule by water. Surveys revealed that frugivory is common throughout the mangrove forest, and fruit had up to six exit holes where larvae had emerged as adults. Larval insects did not affect the flotation of propagules with pericarps, a thin structure that provides buoyancy for dispersal by water. In contrast, after simulating germination by removing the pericarp, the majority of propagules with three exit holes floated on average for 20 h longer than those without exit holes, which sank immediately. Based on this evidence that frugivory could increase the dispersal potential of propagules, I predicted that propagules consumed by larval insects should disperse farther than undamaged propagules, and this was tested by quantifying the potential viability of propagules stranded on beaches at increasing distances (up to 20 km) from mangrove forests. Flies and moths emerged as adults after being transported tens of kilometres within mangrove propagules, revealing a novel mode of dispersal. Proportionally fewer potentially viable propagules were supplied to beaches at increasing distances from mangrove forests, however, indicating that larval insects negatively affect recruitment and are thus not acting as mutualist herbivores. Nevertheless, when transported back to the mangrove forest, seedlings established from propagules damaged by larval insects and stranded on beaches. Therefore, although frugivory does not preclude mangrove recruitment, its negative effects in the pre-dispersal environment may be intensified with increasing dispersal distance, thus limiting the long-distance supply of propagules and recruitment of mangroves.     

Seed predation by insects in tropical mangrove forests: extent and effects on seed viability and the growth of seedlings

Summary Although insects are known to be important seed predators in most terrestrial forests, their role in marine tidal (mangrove) forests has not been examined. Surveys at 12 sites in tropical Australia showed that between 3.1 and 92.7 percent of the seeds or propagules of 12 mangrove tree species had been attacked by insects. Seeds/propagules of six species (Avicennia marina, Bruguiera gymnorrhiza, B. parviflora, Heritiera littoralis, Xylocarpus australasicus and X. granatum) showed consistently high (>40%) levels of insect damage. Greater than 99% of H. littoralis seeds were attacked by insect predators. The survival and subsequent growth in height and biomass of insect-damaged and non-damaged control seeds/propagules of eight mangrove species were compared in shadehouse experiments. Mangrove species fell into 4 groups with regard to the effect of insect predators on their seeds and seedlings. Xylocarpus australasicus and X. granatum had significantly decreased survival (X 48 and 70%) and growth in height (X 61 and 96%) and biomass (X 66 and 85%). Bruguiera parviflora showed decreased survival (X 59%), but there was no effect of insects on the growth of surviving propagules. In contrast, there was no effect of insect damage on the survival of seedlings of Avicennia marina and Bruguiera exaristata, but decreased growth in height (X 22 and 25%) and biomass (X 22 and 26%). Survival and growth of seedlings of Rhizophora stylosa and Bruguiera gymnorrhiza were not affected. The influence of insect seed predators on the survival and growth of seeds of mangrove species in forests will depend on the relative abundance of seed-eating crabs and intertidal position in mangrove forests.This is Contribution No 499 from the Australian Institute of Marine Science     

Predispersal mortality and fecundity in the grey mangrove (Avicennia marina) in southeastern Australia

Abstract The predispersal mortality of floral buds and fruits in the grey mangrove (Avicennia marina) was examined over a range of populations for 3 years. The average survival of floral buds to anthesis was 34%. Moth larvae (subtribe Phycitina) attack and consume the contents of variable numbers of flower buds, but their activities did not influence bud survival, as experimental exclusion of larvae did not increase the number of buds surviving to anthesis. Fruit mortality is initially high with an average of 21% of floral buds surviving to immature fruit. Subsequent mortality is lower until on average 2.9% of floral buds survive to become viable propagules. Several cohorts of the phycitine moth larvae feed throughout the development of the fruit, and up to 62% of mature fruit show signs of larval attack. Experimental exclusion of moth larvae doubled the survival of fruits. However more than 75% of fruit mortality may be attributed to maternal regulation. Hail storms can also cause fruit mortality. The potential fecundity (number of ovules) of A. marina was measured over a range of age-size classes of trees and the realized fecundity (viable propagules) was estimated from average predispersal mortality. These estimates were subsequently verified in the field. A fecundity schedule was constructed with assumptions on reproductive frequency and reproductive life. The average annual supply of propagules over a lifetime is estimated to be about 247 viable propagules per tree.     

The growth of Kandelia candel seedlings in mangrove habitats of the Zhangjiang estuary in Fujian, China

Propagules of Kandelia candel collected from the Zhangjiang estuary were planted in mangrove habitats along the intertidal gradient. The rooting rates of K. candel propagules varied spatially. The lowest rate occurred in Avicennia marina forest (69.7%). The rates were higher in K. candel forest (90.0%), at the fringe of the mangrove forest (89.3%) and on the bare tidal flat outside the mangrove forest (82.7%). After one year, the survival rates of seedlings planted under A. marina forest, K. candel forest, at the fringe of the mangrove forest, and on the bare tidal flat were 13.7%, 54.7%, 76.0%, and 34.7%, respectively. Among the surviving K. candel seedlings, those at the fringe of the mangrove forest and on the bare tidal flat had greater height, stem diameter, leaf number, leaf area, and biomass than those under A. marina and K. candel forests. These results demonstrated that establishment and growth of K. candel seedlings occurred successfully at the fringe of the mangrove forest, but were the worst under A. marina forest. The performance of K. candel seedlings was independent of physico-chemical characters of sediment. However, interspecies competition, propagule predation by insects and crabs, and the incident light had significant effects on seedling survival and growth.     

Propagule size and predispersal damage by insects affect establishment and early growth of mangrove seedlings

Variation in rates of seedling recruitment, growth, and survival can strongly influence the rate and course of forest regeneration following disturbance. Using a combination of field sampling and shadehouse experiments, we investigated the influence of propagule size and predispersal insect damage on the establishment and early growth of the three common mangrove species on the Caribbean coast of Panama: Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle. In our field samples, all three species exhibited considerable intraspecific variation in mature propagule size, and suffered moderate to high levels of predispersal attack by larval insects. Rates of insect attack were largely independent of propagule size both within and among trees. Our experimental studies using undamaged mature propagules showed that, for all three species, seedlings established at high rates regardless of propagule size. However, propagule size did have a marked effect on early seedling growth: seedlings that developed from larger propagules grew more rapidly. Predispersal insect infestations that had destroyed or removed a substantial amount of tissue, particularly if that tissue was meristematic or conductive, reduced the establishment of propagules of all three species. The effect of sublethal tissue damage or loss on the subsequent growth of established seedlings varied among the three mangrove species. For Avicennia, the growth response was graded: for a propagule of a given size, the more tissue lost, the slower the growth of the seedling. For Laguncularia, the response to insect attack appeared to be all-or-none. If the boring insect penetrated the outer spongy seed coat and reached the developing embryo, it usually caused sufficient damage to prevent a seedling from developing. On the other hand, if the insect damaged but did not penetrate the seed coat, a completely healthy seedling developed and its growth rate was indistinguishable from a seedling developing from an undamaged propagule of the same size. Similar to Avicennia, if an infestation did not completely girdle a Rhizophora seedling, it survived, but grew at a reduced rate. In summary, our experiments demonstrated that natural levels of variation in propagule size and predispersal damage by insects translate into significant differences in seedling performance in terms of establishment and/or early growth. Such differences are sufficiently large that they could influence the intensity and outcome of competitive interactions during forest regeneration.     

Leaf abscission phenology of a scrub oak: consequences for growth and survivorship of a leaf mining beetle

Brachys tessellatus is a leaf-mining beetle that attacks Quercus laevis (turkey oak), a deciduous scrub oak in the fall line Sandhills of the southeastern United States. This oak species varies substantially in leaf abscission phenology. In the fall of 1994 we examined leaf abscission patterns at three sites in central South Carolina and found that leaves containing active miners abscised earlier than leaves without active miners. To quantify the effect of leaf abscission phenology on the survivorship and body size of leaf miners within the abscised leaves, we marked naturally and manually abscised leaves, left leaves on the ground to over-winter, and then collected leaves before beetles emerged the following spring. We found that beetles inside leaves that were abscised early in the season were more likely to be killed by predators, and that they produced smaller pupae than beetles within leaves abscised later in the season. We also evaluated the relationship between adult beetle body size and fecundity; body size of B. tessellatus is positively correlated with lifetime fecundity and daily oviposition rate indicating that effects of leaf-abscission phenology on beetle body size likely translate into effects on beetle fitness. We discuss how B. tessellatus has adapted to variation of leaf abscission phenology of oaks by evolving developmental plasticity that permits successful completion of larval development over a range of season lengths.     

The effects of chilling and forcing temperatures on spring synchrony between larch casebearer and tamarack

1. Spring phenological synchrony can be important for tree-insect interactions. Depending on the magnitude and direction of phenological shifts, overwintering insects could be affected in many ways, for example, facing starvation or having to contend with increased chemical or physical defences of host trees. If temperature has different influences on the phenology of trees and insects, climate change can alter spring phenological synchrony.
2. In this experiment, we exposed tamarack seedlings and larch case bearer larvae from Minnesota, USA, to a variety of chilling and forcing temperatures and measured spring phenology (twig bud break and larval activation). We additionally measured case bearer performance on seedlings that were exposed to different forcing × chilling levels, tracking larval survivorship to adulthood.
3. Warmer forcing enhanced larval activation and bud break, but larval development slowed down past 21°C. Higher chilling temperatures accelerated bud break, but the effect was inconclusive for larvae. There was no chilling × forcing interaction for either species. Spring activity accelerated more quickly with increases in temperature for larvae than for seedlings, resulting in increased phenological synchrony at warmer temperatures. Activation rates for overwintering larvae were highest at 27°C, while survivorship to adulthood following spring activation was highest at 21°C. At temperatures at or beyond 27°C, no larvae reached adulthood.
4. Warmer winters and springs will likely initially increase spring synchrony between tamarack and larch case bearer, exposing larvae to younger, potentially more nutritious foliage, but extremely warm spring temperatures may decrease survivorship of larvae to adulthood.

     

The excavation and cultivation in containers of mature grey mangroves,Avicennia marina

The grey mangrove, Avicennia marina, grows in intertidal habitat that is under pressure from expanding human settlement and industry along coastlines in the tropics and subtropics. Inappropriate clearing, and also the dieback of large stands of A. marina associated with pollution, have created an interest in methods for revegetation and also the need for a better understanding of the species physiological tolerances including to herbicide residue. Eco-toxicological studies have so far been exclusively on seedlings because of perceived difficulties cultivating mature plants in controlled environments. Reforestation projects are often associated with low rates of establishment, because of the susceptibility of propagules, seedlings and saplings to wind and wave erosion, as well as flooding and desiccation associated with the natural inter-tidal environment. We report on the successful excavation, and cultivation in containers with an automated irrigation system of eight mature A. marina plants. It is possible to transplant mature mangroves and achieve significant vegetative growth and also flowering and fruiting within the first year. It is difficult to compare the likely costs of transplanting a more limited number of mature plants to many propagules. However, we suggest that in situations where reforestation with young propagules, seedlings or saplings has failed because of excessive wash, drowning or smothering, the transplanting of mature mangroves could be a better option.     

Seed growth suppression constrains the growth of seed parasites: premature acorn abscission reduces Curculio elephas larval size

Abstract.  1. The abscission of seeds infested by insects is common in many plants and has been proposed as a defensive mechanism, although its negative consequences for insects have rarely been assessed.
2. We assessed the consequences of seed abscission studying the interactions between the holm oak Quercus ilex and the chestnut weevil Curculio elephas , its main pre-dispersal seed predator. Female weevils oviposit into the acorns and the larvae must complete their development inside a single acorn feeding on the cotyledons. The growth of the infested acorns is suppressed because they are prematurely abscised.
3. Female weevils oviposit along the acorn growing period; hence, the size of the infested acorns increased with date. The growth of the larvae inside the smaller, early infested, acorns was constrained because food (i.e. cotyledons) was frequently depleted. Larval size increased with the date along with the size of the infested acorns, but it declined slightly in the latest dates as a result of the higher conspecific competition provoked by the larger number of larvae per acorn.
4. The present results demonstrate that premature seed abscission by Q. ilex had negative consequences for C. elephas , as a smaller acorn size reduced food availability and constrained larval size, a key insect life history trait. At the same time, it is suggested that the growth suppression of infested acorns may condition the oviposition phenology of these insects according to their body size. These results need to be considered in further research in the context of the evolutionary significance of premature seed abscission as a defensive mechanism.     

Inhibition of premature leaf abscission by a leafminer and its adaptive significance

We tested the possibility that a lepidopteran leafminer, Coptotriche japoniella Puplesis and Diskus, inhibits the host plant Eurya japonica Thunberg from abscising mined leaves prematurely to increase its survivorship in immature stage. We monitored abscission patterns of mined leaves with sacrificed larvae, mined leaves with living larvae, and unmined leaves from April to July 2004 and 2005 until leafminers emerged as adults. Unmined leaves rarely abscised before July. Mined leaves with sacrificed larvae fell at a constant rate after May, abscising significantly more than unmined leaves. In contrast, mined leaves with living larvae rarely fell before adult emergence; afterward they abscised rapidly. We also examined larval/pupal survivorship and mortality sources on the ground and trees after leafminers completed larval development. Leafminers on the ground suffered a higher mortality from predation than those on trees, and thus they emerged as adults on the ground less successfully. These findings suggest that the leafminer C. japoniella prevents the host plant from abscising mined leaves prematurely until adult emergence, thereby increasing their survivorship.     

Quantitative character variations of cambial derivatives in mangroves and their functional significance

Quantitative character variations of xylem cambial derivatives during secondary growth of the trunk are described for five representative mangrove species: Rhizophora stylosa (Rhizophoraceae), Bruguiera gymnorrhiza (Rhizophoraceae), Kandelia candel (Rhizophoraceae), Sonneratia alba (Sonneratiaceae) and Avicennia marina (Avicenniaceae). Two variation patterns in tracheary element length were revealed among these species. For R. stylosa, A. marina and S. alba, both vessel elements and fibers showed an increase in length during the early stages of secondary growth, then tended to be constant in later growth. In the other two species, little change occurred in the length of either vessel elements or fibers throughout the thickening growth period. Variation patterns in tracheary element length appeared to correspond with the different mangrove species' adaptations to their habitats. In addition, these five species exhibited diverse variation patterns in quantitative characters of the rays as well as in other quantitative characters of the vessels and fibers during secondary growth of their trunk.     

福建漳江口红树林区秋茄幼苗生长动态

通过福建漳江口红树林自然保护区内8个样方24个小样方人工种植600个秋茄胚轴,在3a时间内对秋茄胚轴建立、幼苗生长以及环境因子进行定期观测。研究结果表明：林缘空地的秋茄生长状况良好,白骨壤林内最不利于秋茄幼苗的生长。潮位、盐度、底质土壤理化因子不是造成该样地各样方间秋茄幼苗生长差异的主要原因。动物取食、光照状况以及种间竞争是限制秋茄生长的主要环境因子。秋茄胚轴在长根前易于随潮水漂走,底质土壤中自骨壤致密的根系抑制了秋茄胚轴的定植,导致白骨壤林内秋茄幼苗漂走的数量最多。昆虫和螃蟹等动物的取食是导致林内已经固着生长的秋茄幼苗大量死亡的最主要原因,而林外被取食的幼苗个体极少。此后秋茄幼苗能否继续成长,主要取决于幼苗所接受到的光照条件。3a后,在荫蔽的树冠下,秋茄幼苗无法存活;而在林外,秋茄幼苗已经长成幼树。在林外滩涂上迅速生长的互花米草,也将影响秋茄幼苗的更新和生长。     

Delayed inducible resistance against a leaf-chewing insect in four deciduous tree species

Summary Both mechanical damage to mountain birch foliage and rearing of moth larvae on the trees reduced the growth of Epirrita autumnata larvae reared on these trees in the following year. The effects of physical damage and some other cues from insects were additive. On bird cherry the performance of Epirrita larvae was equal on untreated trees and on trees artificially defoliated in the previous year, but larval growth was reduced on previously insect-damaged branches. With mountain ash just physical damage per se reduced the performance of Epirrita larvae. On Salix phylicifolia there were no significant differences in the growth or survival of Epirrita on untreated control bushes and on bushes with partial larval damage during the previous year. Among untreated control trees the growth and survivorship of Epirrita were higher on fast-growing willow and bird cherry than on the slow-growing mountain birch. Mountain birch and mountain ash, the two deciduous tree species adapted to nutrient-poor soils, showed delayed inducible resistance triggered by defoliation (artificial or insect-made). This supports the hypothesis that delayed inducible resistance may be a passive response due to nutrient-stress caused by defoliation. On the other hand, the additional increase in the resistance of mountain birch triggered by specific cues from insects suggests that this response may be an evolved defense against leaf-eating insects.     

Floral abortion and pollination in four species of tropical mangroves from northern Australia

We assessed natural rates of floral abortion in four common mangrove species from northern Australia and subsequently manipulated pollination experimentally. Sonneratia alba J. Smith exhibited the highest rate of fruit set of the four species (23%), indicating this mangrove was best able to utilise the natural pollination opportunities provided. Fruit set in S. alba appeared, however, to be pollinator limited, as large increases in fruit set occurred after manual cross-pollination of flowers. Avicennia marina (Forsk.) Vierh. had the highest rate of natural pollination, but fruit set was lower (15%) and appeared to be impeded by resource limitations. Although a range of insects visited Ceriops australis (C.T. White) Ballment, T.J. Sm & Stoddart, the rate of fruit set was low (3%) and the capacity for flower fertilisation limited, despite evidence of autogamy in this species. There was an indication of both resource and pollinator limitation in C. australis. Rhizophora stylosa Griff. exhibited limited fruit set (0.5%), possibly due to limiting maternal resources and the lack of adaptation of flowers to either animal or wind pollination. Large increases in fruit set were recorded after manual cross-pollination of R. stylosa flowers. R. stylosa and C. australis, characterised by resource rich propagules with long periods of development, both aborted a large proportion of propagules during the fruit maturation process.     

The demography of the temperate mangrove [Avicennia marina (Forsk.) Vierh.] at its southern limit in New Zealand

An expanding monoculture of the mangrove Avicennia marina var. resinifera growing at its southern limit in New Zealand was modelled using a Leslie matrix technique which assumed exponential population growth. The predictions from the model were partially verified from sequential aerial photographs of the population. The results add to a growing body of practical experience in the application of matrix techniques to plant demography. Modelling indicates that survivorship of the young tree size class is the main determinant of the population growth rate; annual seed production is relatively unimportant. Avicennia trees produce relatively few large viviparous propagules, each of which has a high probability of establishment. Seedlings are retained in a ‘seedling bank’ for several years. The λ value of 1.2228 reflects the colonizing ability of the species, and is compared with published values for other species.     

Establishment and growth of mangrove seedling in mangrove forests of southern Thailand

The demography and growth of mangrove seedlings in southern Thailand were studied for an 18-month period. Plots for measurement of seedlings were established using gradients of light and surface soil texture. The number of established propagules peaked from April to August, the peak forRhizophora being earlier than that forBruguiera. Under the mangrove forest canopy, the survivorship of seedlings established during May to December 1982 was 33% in June 1983. Light conditions and soil texture did not strongly affect initial seedling establishment, and the relationship between species seed size and water depth at a particular site influenced that were able to become established there. The elongation growth was large during the rainy season and in open sites. The growth ofR. apiculata Blume,B. parviflora Wight & Arn. andB. cylindrica Blume in open sites was more than ten times that of seedlings established in the shade. It has noted that light conditions affected the growth of young trees (>1 year old) to a greater degree than that of current-year seedlings.     

The intertidal distribution of the grey mangrove (Avicennia marina) in southeastern Australia: The effects of physical conditions,interspecific competition,and predation on propagule establishment and survival

Abstract The upper and lower limits of the distribution of mature Avicennia marina lie between mean high water and mean sea level in open estuaries in southeastern Australia. Newly established seedlings are highly variable in abundance, but are rarely found in the saltmarsh or on mudflats. Their distribution is unlikely to be limited by dispersal because propagules disperse into the saltmarsh and to intertidal mudflats, but their establishment may be limited by physicochemical conditions, interspecific competition and predation. The model that physicochemical conditions control the intertidal limits of establishment of seedlings was accepted for propagules stranding in the saltmarsh but rejected for those stranding on mudflats. No seedlings established on saltmarsh sediments but similar numbers of seedlings established within light gaps in adult mangrove stands and on intertidal mudflats. The model that interspecific interaction with freeliving macroalgae (Hormosira banksii) reduces the establishment of seedlings on mudflats covered with macroalgae or in stands with a ground cover of macroalgae was accepted. Under controlled conditions five times as many propagules established on cleared ground compared with ground covered with macroalgae. Predators also reduce seedling establishment, but the model that they preferentially act on propagules stranding on the mudflat was rejected. The low number of seedlings found on mudflats without macroalgae appears to relate to wave and current effects on establishment and the effects of waterlogging or fouling on survival.     

The regeneration niche of the grey mangrove (Avicennia marina): effects of salinity,light and sediment factors on establishment,growth and survival in the field

Field observations of seedlings and saplings of Avicennia marina showed patterns that correlated with salinity, light and sediment. Models that account for these observations were subsequently tested in a series of field experiments. Establishment varied within an estuary under controlled conditions but was not related to salinity or sediment type. Seedling survival was uniform over 3 years regardless of position in estuary and sediment type. Seedling densities and survival under canopies or in canopy gaps were not significantly different. However, seedling growth and density of saplings were greater in canopy gaps. Experiments involving manipulations of canopies showed no differences in seedling survival under canopies or in light gaps, but addition of slow-release fertilizer enhanced growth and survival in canopy gaps and under canopies. Long-term comparison of areas denuded of a canopy and with sediment disturbance showed enhanced establishment and survival when compared with areas with canopy gaps but with undisturbed sediments. Overall there appears to be no restriction to establishment of propagules within mangrove stands other than the supply of propagules and tidal or wave action. In contrast, recruitment to the sapling stage appears to be restricted by light and sediment resources. We suggest that propagules need to establish in a regeneration niche for seedling recruitment to the sapling stage. This differs from the view that seedlings in the under-storey are analogous to a seed pool in the soil.     

Mangrove restoration without planting

Mangrove forests are disappearing at an alarming rate. Mangrove planting is the most common method of restoring these forests. However, this approach is not often successful, especially when the reasons for mangrove degradation were not removed prior to planting new seedlings or propagules. A successful mangrove restoration project may not necessarily include a planting phase. When the stressors are removed and suitable environmental conditions such as correct hydrology and calm area, particularly on exposed coasts, are provided, natural regeneration processes could recover mangroves from degradation. This paper describes an approach to mangrove restoration applied to an exposed shoreline on the west coast of Peninsular Malaysia. In this method a detached breakwater was erected seaward at the shoreline to shelter the restoration area from wave action, prevent the ongoing erosion and promote sediment deposition raising the elevation of the substrate to the target elevation. Survival of Avicennia marina seedlings transplanted in the sheltered area was significantly affected by sediment burial (P < 0.05). However, about eight months after construction of the breakwater, the sedimentation rate decreased and the substrate became stable. Subsequently, natural recruits appeared on the site. We postulate that waterborne seeds or propagules were available from adjacent stands in the study area but wave exposure reduced the possibility of natural recruitment. Thus the breakwater presence provided favourable conditions which could attract mangrove recruits, facilitating reestablishment and natural recovery of the mangrove ecosystem without planting.     

Occurrence of Amblycerus species in Cordia trichotoma seeds and their influence on germination

Forest species can have their seeds damaged by granivorous insects, especially by those in their larval stage. In this context, this study aims to report the occurrence of Amblycerus species in Cordia trichotoma seeds, to describe their main damage to seeds and effects on germination, as well as their associated hymenopteran parasitoids. Therefore, seven trees were selected in the municipality of Taquaruçu do Sul, RS, Brazil. Fruits were collected weekly from the medium third of the tree crown, from the beginning of their formation until total dehiscence. To examine the damage caused by granivorous insects within the fruits, 15 fruits from each tree were sectioned with a scalpel. Furthermore, 10 fruits from each tree were stored individually in clear plates to verify the occurrence and identification of granivorous insect species. Evidence of the damage caused to seeds was verified through the germination test by comparing preserved and damaged seeds, with four repetitions of 25 seeds each. The species Amblycerus longesuturalis and Amblycerus profaupar (Chrysomelidae: Bruchinae) were found associated with fruits of C. trichotoma. Female insects predominantly laid eggs on the superior part between the marcescent calyx and the fruit, and larvae perforated the fruit tegument to start consuming seed embryos and reserves. Bruquine larvae are parasitized by Hymenoptera of Bracon, Mirax, Omeganastatus and Triapsis genera. In conclusion, the germination of C. trichotoma seeds is significantly affected by emergence orifices caused by granivorous species.