Male morphology and dishonest signalling in a fig wasp

Despite theoretical predictions, dishonest signalling has rarely been observed in aggressive interactions. We present evidence of such signalling in the nonpollinating. g wasp Philotrypesis sp. A ex Ficus rubiginosa. First, morphometric data indicated that an alternative 'atypical' male morph (17.8% of individuals) exists that tends to be larger in body size and has longer mandibles for a given body size than other 'typical' males. Second, behavioural observations suggested that males use mandible gape width (which depends on mandible length) as a cue to assess opponents before fights and retreat without escalating if they are unlikely to win, and, probably because their greater mandible gape width causes more opponents to retreat without escalating, that atypical males engaged in fewer fights than typical males for a given body size but had higher mating success. Third, atypical males were less likely to win fights than typical males of similar mandible length relative to opponents. In addition, we found that atypical males incur more injuries (greater receiver-dependent signal costs) than typical males of similar body size relative to rivals. We discuss the implications of our findings for future work on dishonest signalling. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Interference competition and high temperature reduce the ‘virulence’ of fig wasps and contribute to stability of a fig-wasp mutualism

Fig trees are pollinated by fig wasps, which also oviposit in female flowers. The wasp larvae gall and eat developing seeds. Although fig trees benefit from allowing wasps to oviposit, because the wasp offspring disperse pollen, figs must prevent wasps from ovipositing in all flowers, or seed production would cease, and the mutualism would go extinct. In Ficus racemosa, we find that syconia (‘figs’) that have few foundresses (ovipositing wasps) are underexploited in the summer (few seeds, few galls, many empty ovules) and are overexploited in the winter (few seeds, many galls, few empty ovules). Conversely, syconia with many foundresses produce intermediate numbers of galls and seeds, regardless of season. We use experiments to explain these patterns, and thus, to explain how this mutualism is maintained. In the hot summer, wasps suffer short lifespans and therefore fail to oviposit in many flowers. In contrast, cooler temperatures in the winter permit longer wasp lifespans, which in turn allows most flowers to be exploited by the wasps. However, even in winter, only in syconia that happen to have few foundresses are most flowers turned into galls. In syconia with higher numbers of foundresses, interference competition reduces foundress lifespans, which reduces the proportion of flowers that are galled. We further show that syconia encourage the entry of multiple foundresses by delaying ostiole closure. Taken together, these factors allow fig trees to reduce galling in the wasp-benign winter and boost galling (and pollination) in the wasp-stressing summer. Interference competition has been shown to reduce virulence in pathogenic bacteria. Our results show that interference also maintains cooperation in a classic, cooperative symbiosis, thus linking theories of virulence and mutualism. More generally, our results reveal how frequency-dependent population regulation can occur in the fig-wasp mutualism, and how a host species can ‘set the rules of the game’ to ensure mutualistic behavior in its symbionts.

Longevity, early emergence and body size in a pollinating fig wasp - implications for stability in a fig-pollinator mutualism

1. Fig trees (Ficus) are pollinated only by agaonid wasps, whose larvae also gall fig ovules. Each ovule develops into either a seed (when pollinated) or a wasp (when an egg is also laid inside) but not both. 2. Ovipositing wasps (foundresses) favour ovules near the centre of the enclosed inflorescence (syconium or 'fig'), leaving ovules near the outer wall to develop into seeds. This spatial stratification of wasps and seeds ensures reproduction in both partners, and thereby enables mutualism persistence. However, the mechanism(s) responsible remain(s) unknown. 3. Theory shows that foundresses will search for increasingly rare inner ovules and ignore outer ovules, as long as ovipositing in outer ovules is sufficiently slow and/or if inner ovules confer greater fitness to wasps. The fig-pollinator mutualism can therefore be stabilized by strong time constraints on foundresses and by offspring fitness gradients over variation in ovule position. 4. Female fig wasps cannot leave their galls without male assistance. We found that females in outer ovules were unlikely to be released. Inner ovules thus have added value to foundresses, because their female offspring are more likely to mate and disperse. 5. For those offspring that did emerge, gall position (inner/outer) and body size did not influence the order in which female pollinators exited syconia, nor did early emerging wasps enjoy increased life spans. 6. We also found that the life spans of female wasps nearly doubled when given access to moisture. We suggest that conflict resolution in the fig-pollinator mutualism may thus be influenced by tropical seasonality, because wasps may be less able to over-exploit ovules in dry periods due to time constraints.

A role for parasites in stabilising the fig-pollinator mutualism

Mutualisms are interspecific interactions in which both players benefit. Explaining their maintenance is problematic, because cheaters should outcompete cooperative conspecifics, leading to mutualism instability. Monoecious figs (Ficus) are pollinated by host-specific wasps (Agaonidae), whose larvae gall ovules in their "fruits'' (syconia). Female pollinating wasps oviposit directly into Ficus ovules from inside the receptive syconium. Across Ficus species, there is a widely documented segregation of pollinator galls in inner ovules and seeds in outer ovules. This pattern suggests that wasps avoid, or are prevented from ovipositing into, outer ovules, and this results in mutualism stability. However, the mechanisms preventing wasps from exploiting outer ovules remain unknown. We report that in Ficus rubiginosa, offspring in outer ovules are vulnerable to attack by parasitic wasps that oviposit from outside the syconium. Parasitism risk decreases towards the centre of the syconium, where inner ovules provide enemy-free space for pollinator offspring. We suggest that the resulting gradient in offspring viability is likely to contribute to selection on pollinators to avoid outer ovules, and by forcing wasps to focus on a subset of ovules, reduces their galling rates. This previously unidentified mechanism may therefore contribute to mutualism persistence independent of additional factors that invoke plant defences against pollinator oviposition, or physiological constraints on pollinators that prevent oviposition in all available ovules.

The phenology and potential for self-pollination of two Australian monoecious fig species

In this preliminary study, the reproductive phenology of two monoecious fig species, Ficus racemosa and F. rubiginosa, was examined in tropical Australia. Syconia (inflorescences) occurred on both species all year round, but pre-floral and interfloral syconia were much commoner than the wasp-receptive and wasp-emitting phases in both species. The temporal overlap of the wasp-receptive and wasp-emitting phases on a single tree indicated that self-pollination was possible in both species and that pollinators may sometimes persist through multiple generations on one tree. This sexual phase overlap was commoner in F. rubiginosa than in F racemosa. The two species also differed in their general within-tree asynchrony, with a higher diversity of phases on F. rubiginosa than on F. racemosa. The time from syconium initiation to ripening was very similar in F. rubiginosa (mean = 48.51 days) and F. racemosa (mean = 43.53 days). However, there was much more variation within and between trees for F. rubiginosa. In addition, the wasp-receptive phase was found to last up to 5 days (rnean = 4.38) in F. rubiginosa. Such longevity should contribute substantially to local pollinator population persistence. Future work should use genetic studies to determine whether self-pollination is common in these fig species.

Fighting strategies in two species of fig wasp

Although theory exists concerning the types of strategies that should be used in contests over resources, empirical work explicitly testing its predictions is relatively rare. We investigated male fighting strategies in two nonpollinating. g wasp species associated with Ficus rubiginosa figs. In Sycoscapter sp. A, males did not assess each other before or during fights over mating opportunities. Instead,fights continued until the loser reached an energetic cost threshold that was positively correlated with its body size (fighting ability) and retreated. In Philotrypesis sp. B, pre fight assessment was indicated, with males attacking competitively inferior rivals to remove them from the competitor pool ( they then continued to do so until they reached a cost threshold that was again positively correlated with body size). Using data on species ecology, we discuss our findings with respect to theory on when different fighting strategies should evolve. We argue that the type of strategy used by a. g wasp species is determined by its relative benefits in terms of inclusive fitness. (c) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Diverse mariner-like elements in fig wasps

Mariner transposable elements are widespread and diverse in insects. We screened 10 species of fig wasps (Hymenoptera: Agaonidae) for mariner elements. All 10 species harbour a large diversity of mariner elements, most of which have interrupted reading frames in the transposase gene region, suggesting that they are inactive and ancient. We sequenced two full-length mariner elements and found evidence to suggest that they are inserted in the genome at a conserved region shared by other hymenopteran taxa. The association between mariner elements and fig wasps is old and dominated by vertical transmission, suggesting that these 'selfish genetic elements' have evolved to impart only very low costs to their hosts.

Evidence for inclusion of South African endemic Nylandtia in Muraltia (Polygalaceae)

Nylandtia (Polygalaceae) is a small South African genus of two or more species distributed mainly in the Cape region. Previous studies based on anatomical, morphological and molecular data have already revealed a close relationship between Nylandtia and Muraltia, a genus of 117 species that is nearly endemic to South Africa. New evidence from molecular studies of family Polygalaceae and genus Muraltia shows that Nylandtia is derived from genus Muraltia, and is nested in Muraltia subgenus Psiloclada. These results have prompted a morphological re-evaluation of the genera Nylandtia and Muraltia, supporting the conclusion that the two species of Nylandtia currently recognized, N. spinosa and N. scoparia, should be included in subgenus Psiloclada within the genus Muraltia. Only one nomenclatural change is necessary: Muraltia spinosa (L.) E Forest & J. C. Manning, comb. nov.

A new species of Myosotis L. (Boraginaceae) from Cameroon, with a key to the Tropical African species of the genus

Myosotic cameroonensis Cheek & R Becker (Boraginacene) is described from Cameroon. Its conservation status and taxonomic affinities are assessed and an updated key to the Tropical African species of the genus is presented.

Patterns of peristome reduction and ornamentation in African Entodontaceae

The Entodontaceae is characterised by erect capsules with inset and substantially reduced peristomes. Differences in peristome morphology and ornamentation delimit both genera and species within the family. The Entodontaceae, as presently recognised, consists of four genera, Entodon, Erythrodontium, Mesonodon and Pylaisiobryum, which show a variety of peristome ornamentation patterns, and exhibit differing levels of peristome reduction. This variation encompasses an unusual combination of structurally reduced peristomes, which retain massive exostome ornamentation, and residual peristome function. As part of a taxonomic revision of the family in Africa, generic patterns of peristome reduction and ornamentation are described. Inter- and intra-specific variation patterns of both the exostome and endostome have been investigated using detailed SEM studies of material from Africa and beyond. Structural reduction of the peristome is associated with high levels of intra-specific variation in peristome ornamentation patterns. Common trends in these patterns of variation are identified within the family. Putative hybrid sporophytes have been identified in Entodon, and the taxonomic implications of these observations are considered for a family where species delimitation has in the past often depended on minor variation in peristome structure or ornamentation patterns. (author abst.)

Factors controlling the distribution of ozone in the West African lower troposphere during the AMMA (African Monsoon Multidisciplinary Analysis) wet season campaign

Ozone and its precursors were measured on board the Facility for Airborne Atmospheric Measurements (FAAM) BAe 146 Atmospheric Research Aircraft during the monsoon season 2006 as part of the African Monsoon Multidisciplinary Analysis (AMMA) campaign. One of the main features observed in the west African boundary layer is the increase of the ozone mixing ratios from 25 ppbv over the forested area (south of 12 degrees N) up to 40 ppbv over the Sahelian area. We employ a two-dimensional ( latitudinal versus vertical) meteorological model coupled with an O-3-NOx-VOC chemistry scheme to simulate the distribution of trace gases over West Africa during the monsoon season and to analyse the processes involved in the establishment of such a gradient. Including an additional source of NO over the Sahelian region to account for NO emitted by soils we simulate a mean NOx concentration of 0.7 ppbv at 16 degrees N versus 0.3 ppbv over the vegetated region further south in reasonable agreement with the observations. As a consequence, ozone is photochemically produced with a rate of 0.25 ppbv h(-1) over the vegetated region whilst it reaches up to 0.75 ppbv h(-1) at 16 degrees N. We find that the modelled gradient is due to a combination of enhanced deposition to vegetation, which decreases the ozone levels by up to 11 pbbv, and the aforementioned enhanced photochemical production north of 12 degrees N. The peroxy radicals required for this enhanced production in the north come from the oxidation of background CO and CH4 as well as from VOCs. Sensitivity studies reveal that both the background CH4 and partially oxidised VOCs, produced from the oxidation of isoprene emitted from the vegetation in the south, contribute around 5-6 ppbv to the ozone gradient. These results suggest that the northward transport of trace gases by the monsoon flux, especially during nighttime, can have a significant, though secondary, role in determining the ozone gradient in the boundary layer. Convection, anthropogenic emissions and NO produced from lightning do not contribute to the establishment of the discussed ozone gradient.

In-service training needs in an African context: a study of headteacher and teacher perspectives in the Gucha District of Kenya

Improving the quality of teaching is an educational priority in Kenya, as in many developing countries. The present paper considers various aspects of in-service education, including views on the effectiveness of in-service, teacher and headteacher priorities in determining in-service needs and the constraints on providing in-service courses. These issues are examined though an empirical study of 30 secondary headteachers and 109 teachers in a district of Kenya. The results show a strong felt need for in-service provision together with a firm belief in the efficacy of in-service in raising pupil achievement. Headteachers had a stronger belief in the need for in-service for their teachers than did the teachers themselves. The priorities of both headteachers and teachers were dominated by the external pressures of the schools, in particular the pressures for curriculum innovation and examination success. The resource constraints on supporting attendance at in-service courses were the major problems facing headteachers. The results reflect the difficulties that responding to an externally driven in-service agenda creates in a context of scarce resources.

Post-partum depression and infant growth in a South African peri-urban settlement

Aim: To examine the association between maternal post-natal depression and infant growth. Background: Infant growth has recently been shown, in two studies from South Asia, to be adversely affected by maternal depression in the early post-partum period. It is uncertain whether a similar association obtains in developing countries outside Asia. Method: A sample of 147 mother–infant dyads was recruited from a peri-urban settlement outside Cape Town and seen at 2 and 18 months post partum. Results: No clear effect of post-partum depression on infant growth was found. Although maternal depression at 2 months was found to be associated with lower infant weight at 18 months, when birthweight was considered this effect disappeared. Conclusions: Possible explanations for the non-replication of the South Asian findings are considered.

The mother-infant relationship and infant attachment in a south African peri-urban settlement

A sample of 147 mother-infant dyads was recruited from a peri-urban settlement outside Cape Town and seen at 2- and 18-months postpartum. At 18 months, 61.9% of the infants were rated as securely attached (B); 4.1% as avoidant (A); 8.2% as resistant (C); and 25.8% disorganized (D). Postpartum depression at 2 months, and indices of poor parenting at both 2 and 18 months, were associated with insecure infant attachment. The critical 2-month predictor variables for insecure infant attachment were maternal intrusiveness and maternal remoteness, and early maternal depression. When concurrent maternal sensitivity was considered, the quality of the early mother-infant relationship remained important, but maternal depression was no longer predictive. Cross-cultural differences and consistencies in the development of attachment are discussed.