Reproductive seasonality in tropical satyrine butterflies: strategies for the dry season

Abstract.

• 1 In tropical savanna environments rainfall is often very seasonal, so that much of the year is characterized by a long and unpredictable dry season. Because the timing and availability of rain exerts a major influence on plant growth and production, many species during the dry period exhibit dramatic reduction in leaf quality. Accordingly, and kind of behaviour shown by phytophagous insects that synchronizes larval feeding with food availability will be adaptive.
• 2 The reproductive status of three Mycalesis butterflies was monitored over a 2-year period (1989–90) in north-eastern Queensland, Australia, at a lowland site (Cardwell, 18°16's, 146°02′E) which experiences a pronounced dry season. Females of these species and of five other satyrines (Ypthima, Hypocysta spp.) were also examined less intensively during the dry season in areas throughout northern and central Queensland, north of the tropic of Capricorn.
• 3 These relatively sedentary butterflies exhibit three different strategies for dealing with the unpredictable dry period and associated deterioration of larval food plants (grass). First, five species appear to breed continuously, though for most reproductive activity (mature egg number) declines markedly in the late dry season. Two of these (Hypocysta irius, H.metirius) are restricted to less seasonal and more favourable (wetter) areas but the three others (Ypthima arctous, H.adiante, H.pseudirius) occur widely in the relatively dry savanna, where they may specialize on grass in moister microenvironments. Second, two species (M.terminus, M.sirius) live in predictably moist habitats which are buffered from climatic extremes; they breed for much of the season but reproductive activity declines as the dry season progresses and may cease late in the season. Third, one species (M.perseus) is more opportunistic, breeding for only a limited interval during the favourable (wet) periods; during the long dry season adults contract to moist refugia and remain in reproductive diapause.
• 4 Spending the late dry season as an adult, either in diapause or with mature eggs, may improve the capacity to utilize new growth of grasses at the start of the favourable season, thereby enhancing population growth during good times. It may also provide additional flexibility to counter the temporal uncertainty of the dry season.
• 5 The strategy of residing in more equitable habitats or specializing on predictable foods may be the most restrictive in terms of distribution.

     

Seasonal allocation of dry mass and nitrogen in a fynbos endemic Restionaceae species Thamnochortus punctatus Pill.

Summary The annual growth pattern of male plants of the endemic, dioecious fynbos species, Thamnochortus punctatus, revealed sequential growth of the various below-and above-ground organs. Root and rhizome development were favoured in the wet winter months while culm elongation occurred in the warmer but drier spring to summer period. The reproductive phase of development followed in the hot dry summer and autumn. The asynchronous pattern of growth is suggested to enable the plant to maximize utilization of scarce resources within the confines of the environmental controls determined by the mediterranean-climate. Maximum root and rhizome activity in the rainy season enabled the plant to absorb and store nitrogen in mature culms for later above ground organ development. As the absorption of nitrogen from senscing material is essential for this pattern of organ development to assist in alleviating nutrient scarcity the high degree of nitrogen reabsorption, 35 to 70% for branches and culms respectively, shows the potential importance of asynchronous organ growth in overcoming nitrogen limitation of plant growth. Comparisons of dry mass and nitrogen allocation patterns indicate that developmental strategies employed by evergreen, perennial plants in climatically and edaphically constrained environments cannot be deduced from biomass allocation patterns alone as they do not appear to reflect strategies evolved to overcome nutrient limitation.     

Vegetative, reproductive, and physiological Adaptations to aridity of pitayo (Stenocereus queretaroensis, Cactaceae)

Pitayos fStenocereus spp.) are columnar cacti that produce attractive edible fruits under both wild and cultivated conditions. Until recently the study of pitayos has been relatively neglected even though they have been a crucial staple for the inhabitants of subtropical semiarid lands of Mexico since ancient times and this century have become an important fruit crop there. The main objective of this review is to relate seasonal aspects of vegetative and reproductive growth, CO₂ uptake, and carbohydrate resources to abiotic components of the environment. Flower and fruit production occur during the dry season in the late winter and spring, roots grow during the summer wet season, and stem elongation commences in the autumn at the beginning of the dry season. Vegetative growth thus does not coincide with reproductive growth, as it does for other fruit crops in temperate and tropical regions. The time offset of vegetative and reproductive activities reduces competitive sink effects, regulating assimilate partitioning to different organs, which may allow Stenocereus queretaroensis to adapt to water-limited environments. Reserve carbohydrates (starch, mucilage) accumulate in the early summer just after reproductive growth and in the early winter just after stem extension. Reducing sugars increase during the middle of the summer before stem elongation begins. The low rates of growth and of photosynthesis for S. queretaroensis are associated with low tissue levels of nitrogen, chlorophyll, and some micronutrients (Fe, Mn). Low levels of gibberellic acid may also contribute to its low growth rate. It does not respond morphologically to irrigation during the dry period, indicating a low plasticity for growth. Although S. queretaroensis is cultivated, its physiological and phenological activities closely resemble those of wild CAM species and other wild perennial plants, reflecting its rather recent domestication and the fact that selections were based mainly on fruit quality and adaptation to aridity rather than biomass productivity.     

The effect of altered sink-source relations on photosynthetic traits and matter transport during the phase of reproductive growth in the annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis>

Annual plants transport a large portion of carbohydrates and nitrogenous compounds from leaves to seeds during the phase of reproductive growth. This study aimed to clarify how reproductive growth affects photosynthetic traits in leaves and matter transport within the plant in the annual herb Chenopodium album L. Plants were grown in pots and either reproductive tissues or axillary leaves were removed at anthesis. Matter transport was evaluated as temporal changes in dry mass (as a substitute of carbohydrates) and nitrogen content of aboveground organs: leaves, axillary leaves, stems and reproductive tissues. Photosynthetic capacity (light-saturated photosynthetic rate under ambient CO₂ concentration), nitrogen, chlorophyll and soluble protein content were followed in the 20^th leaf that was mature at the start of the experiment. Removal of reproductive tissues resulted in accumulation of dry mass in leaves and axillary leaves, and accumulation of nitrogen in stem as nitrogen resorption from leaves and axillary leaves proceeded with time. Removal of axillary leaves proportionally reduced dry mass and nitrogen allocation to reproductive tissues, thus affecting the quantity but not quality of seeds. Removal treatments did not alter the time course of photosynthetic capacity, nitrogen, chlorophyll or soluble protein content during senescence in the 20^th leaf, but changed the photosynthetic capacity per unit of leaf nitrogen according to demand from reproductive tissues. Together, the results indicate that reproductive tissues affected carbon and nitrogen economy separately. The amount of carbon was adjusted in leaves through photosynthetic capacity and carbohydrate export from them, and the amount of nitrogen was adjusted by transport from stem to reproductive tissues. The plant’s ability to independently regulate carbon and nitrogen economy should be important in natural habitats where the plant carbon-nitrogen balance can easily be disturbed by external factors.     

Plant compositional change over time increases with rainfall in Serengeti grasslands

Previous work from the Plains of Serengeti National Park (SNP) suggested that plant species composition (i.e. the composite measure of species' identities and abundances) is unchanged on the temporal scale of decades ( Belsky 1985 ), raising the possibility that African grazing ecosystems may be resistant to climate change. In this study, grassland plant composition was measured three times between 2000 and 2007 at eight permanent sites spanning the SNP environmental gradient. Confirming previous work, plots in the Serengeti Plains were relatively stable with respect to plant composition through time. In contrast, sites in all other regions of the ecosystem showed greater change in plant species composition during the study. Three factors, annual rainfall, dry season rainfall, and fire frequency, were linearly correlated (all r>0.73) with the amount of composition change at the sites. A path model, which accounted for the strong correlation between annual and dry season rainfall and for the known effect of annual rainfall on fire frequency, was used to elucidate which factors were associated with composition change and which were spuriously correlated. The path model demonstrated that change in plant composition was positively related to annual rainfall, but not dry season rainfall or fire frequency. However, dry season rainfall was positively associated with changes in plant species identity, one component of compositional change, across sites over time. Because climate models predict both increased wet season and decreased dry season rainfall in East Africa, these findings raise uncertainty about how these grasslands will respond to future changes. This study and others ( Belsky 1985 ) suggests that plant assemblages adapted to low rainfall in Serengeti may exhibit compositionally stability, while the same may not be true in regions of higher average rainfall and larger species pools.     

Graminivory and Fallback Foods: Annual Diet Profile of Geladas (<Emphasis Type="Italic">Theropithecus gelada</Emphasis>) Living in the Simien Mountains National Park,Ethiopia

All primates show some dietary flexibility, particularly during food shortages. Foods consumed during times of scarcity (i.e., fallback foods) strongly influence the ecology and evolution of a species. Geladas (Theropithecus gelada) eat primarily graminoid leaves (i.e., grasses and sedges), but also consume other diet items (e.g., underground storage organs), especially in the dry season. We investigated the feeding ecology of wild geladas in the Sankaber region of the Simien Mountains National Park, Ethiopia across 12 mo. We asked how the gelada diet in this region, which is disturbed by livestock and agriculture, correlated with food availability and whether underground foods are fallback foods. We quantified the monthly diets of adults from eight reproductive units using instantaneous scan sampling, and seasonal aboveground and underground food availability using point-intercept transects and soil core sampling. Geladas primarily consumed graminoid leaves year-round (76.3% of the annual diet, 36.2–93.2% of the monthly diet) but spent considerable time consuming underground foods in the dry season (14.0% of the annual diet, 11.1–49.7% of the diet across dry season months). Graminoid consumption increased with graminoid availability, and underground food consumption decreased with graminoid availability. Underground food availability did not vary significantly between the dry and wet season sampling months, supporting the hypothesis that underground foods are fallback foods for geladas. We then compiled data from gelada feeding studies and found that underground foods are an important dry season diet item across study sites, but geladas rely more heavily on underground foods in habitats more heavily influenced by humans. Understanding the range and effects of primate dietary flexibility in human-modified habitats will contribute to a better understanding of how changing environments shape primate ecology and evolution.     

Reproductive phenology, life-forms, and habitats of the Venezuelan Central Plain

Reproductive phenology of 171 plant species belonging to 57 families of angiosperms was studied according to life-forms in four habitat types in a savanna-forest mosaic on the Venezuelan Central Plain. Flowering, unripe fruit, and mature fruit patterns were affected significantly according to life-forms and habitats respectively. Production of flowers, unripe fruits, and mature fruits showed marked seasonality for all habitats except for the forest. Flowering peaked during the rainy season, and fruiting peaked toward the end of the rainy season. The savanna and the disturbed area had similar proportions of species that flowered over the year. The percentage of species with unripe fruits produced throughout the year was more seasonal for the disturbed area than for the other habitats. Mature fruit patterns showed an increase during the late rainy season for the ecotone and savanna. A large number of herbaceous (annual and perennial) and liana species flowered during the wet season, and a smaller fraction flowered during the dry season; and trees, shrubs, and epiphytes increased flowering activity during the dry season. Unripe fruit patterns were similar to those of flowering for all life-forms, however, tree species were less seasonal. Mature fruit production by shrubs peaked in the period of maximum rainfall, while the peak for perennial herbs was in the late rainy season and the peak for annual herbs was during the transition between the rainy season and the dry season. The largest proportion of tree and liana species with ripe fruits occurred during the dry season. Differences among phenological patterns in habitats were caused mainly by life-forms and promote a wider distribution of reproductive events in habitats and overall community in the Venezuelan Central Plain.     

N, P, and C stoichiometry of Eranthis hyemalis (Ranunculaceae) and the allometry of plant growth

We report the nitrogen (N), phosphorus (P), and carbon (C) stoichiometry for each of the five organ-types (leaves, aerial stems, reproductive organs, roots, and tubers) of 17 actively growing Eranthis hyemalis plants differing in size (as measured in g C). We also report the N, P, and C stoichiometry of 20 winterized tubers, which are the only perennial organs of this species. Comparisons between whole-plant and winterized N/C and P/C levels indicate that N was resorbed from aerial organs and stored in tubers by the end of the growing season. Leaves were substantial reservoirs for N and P. With few exceptions, N scaled isometrically with respect to C for each organ-type, whereas P scaled as the 3/4 power of C. Thus, N is proportional to P(3/4), which is proportional to C regardless of organ-type. Additionally, annual growth rate G of shoots (leaves and aerial stems) scaled as the -3 power of leaf N/P quotients such that G was proportional to the 3/4 power of leaf P. We suggest that these scaling relationships (together with previously reported allometric trends across herbaceous species) show that growth is constrained by organ-specific N and P allocation patterns (presumably to proteins and ribosomes, respectively).     

THE ANNUAL CYCLE IN A TROPICAL WET FOREST HUMMINGBIRD COMMUNITY

In the Caribbean lowlands of Costa Rica, rainfall is moderately seasonal, although even in the driest month over 100 mm of rain usually fall. Flowering of hummingbird food plants shows a peak in the dry season (February-April) and another in the early wet season (July-September), with a severe flower shortage at the end of the rains (November-December). The dry season peak involves largely canopy epiphytes, the wet season peak large herbs of light gaps and edges and forest understory plants. This study examines the responses of the associated community of 22 species of hummingbirds (of which 13 breed, and 12 are common for at least parts of most years) to these spatial and temporal patterns of resource availability. Nearly all common breeding species show a peak of reproductive activity in the dry season, coinciding with the first flowering peak, followed by a discrete moulting season that coincides with the wet season peak of flowering. Of the three species with extended breeding seasons, the two species of hermit, Phaethornis, show moult-breeding overlap to varying degrees on an individual basis. In a number of species moult and breeding appear antagonistic. The annual peak of body weight and fat deposits in all species occur during the second flowering peak, approximately corresponding to the moult. The annual minima of body weight and fat occur in the lean season and the breeding season respectively. The lack of concordance of these two possibly reflects the use of muscle protein as a nutrient source during the lean season. Several species show pronounced habitat shifts through the year, with the sexes sometimes occupying different microhabitats, especially during the dry season. At least five species show pronounced seasonal migrations, largely or entirely leaving La Selva for part of the year. Overall hummingbird numbers are greatest early in the rainy season, lowest in the lean season, with the non-hermits (Trochilinae) showing a more pronounced annual cycle of numbers than the hermits (Phaethorninae). Comparisons with other tropical lowland hummingbird-flower communities are made with respect to the roles of flowers as proximate and ultimate factors regulating the annual cycles and affecting the population biology of the birds.     

Reproductive Phenology of Woody Species in a North Australian Tropical Savanna1

Interspecific and interannual variation in reproductive phenology was quantified for 50 common species of trees and shrubs from a mesic savanna near Darwin, northern Australia. The presence of buds, flowers, and fruit was noted over a 30-month period, from September 1992 to February 1995. Surveys were undertaken at monthly intervals for the less common species, and at bimonthly intervals for ten of the common trees and tall shrubs. The majority of species flowered each year at about the same time. There was no evidence of sub-annual or continuous regimes of reproductive phenology. There was no supra-annual carryover of seed-bearing fruit in the canopy of any species. The peak flowering periods were the mid to late dry season (July–August) and the transition between the dry season and the wet season (October–November). The two dominant trees–Eucalyptus miniata and E, tetrodonta– flowered during the dry season, thereby providing resources for some elements of the vertebrate fauna. Flowering and fruiting were uncommon at the end of the wet season (February/March), although two species that flower and fruit at this time (E. porrecta and Terminalia ferdinandianas may provide resources to consumers at a time when floral or fruit resources are otherwise scarce. Because the peak of reproductive activity takes place during the late dry season, fruit maturity and seed dispersal have occurred prior to the onset of the rainy season for most species, and germination and seedling establishment potentially may take effect in response to the first rains. Late dry season fires, which tend to be extensive and intense, are a potential threat to the floral and fruit reserves within these savannas.     

Herbaceous production in South India—limiting factors and implications for large herbivores

This study’s goal was to better understand the growth pattern and limitations of the herbaceous production that supports South India’s rich large herbivore grazer assemblage. We conducted a fully factorial nitrogen and water (three levels each) treatment field experiment in the herbivore rich South Indian Western Ghats region to determine the seasonal pattern and the extent to which nitrogen and water availability limit herbaceous production. Graminoid production was found to be nitrogen limited. Despite low rainfall, additional water did not significantly increase overall biomass production nor extend growth in the dry season. Accumulated standing biomass was highest in the late wet season (November) and lowest in the dry season (May). Leaf nitrogen was highest in the early wet season (June) and lowest in the late dry season (March). Grazing had a positive effect on grass production by extending the growing season. Biomass production and graminoid leaf nitrogen concentration levels in the study area were similar to other tropical areas in the world. Also similar to other tropical large herbivore areas, the dry season poses an annual challenge for large herbivores in the study area —particularly the smaller bodied species—to satisfy their nutrient requirements.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

The evolution of reproductive strategies in dasyurid marsupials: implications of molecular phylogeny

Dasyurid marsupials show a remarkable diversity of reproductive patterns ranging from aseasonal polyoestry to restricted annual breeding in which males synchronously die after a brief mating season. Previous studies have categorized dasyurid reproduction into six strategies, defined on the basis of five life-history characters. We provide an up-to-date summary of reproductive traits in dasyurid species and examine the evolution of these characters on a phylogeny for the family recently obtained from DNA sequence data. Our results suggest that reproductive evolution in modern dasyurids is characterized by a basal separation of subfamily lineages employing Strategy II (monoestrous females, restricted breeding season, 11 months to maturity; Dasyurinae) and Strategy V (polyoestrous females, extended breeding season, 8–11 months to maturity; Sminthopsinae). Strategies I (male die-off) and III (facultative polyoestry) appear to have arisen several times from Strategy II or V ancestors, and Strategy IV appears to have arisen within Sminthopsis from a Strategy V ancestor. Strategy VI (aseasonal breeding) has arisen independently in each of the four major dasyurid lineages (tribes), and is highly (but not perfectly) correlated with New Guinean endemism. This scenario is not strongly affected if reproductive characters are optimized on an alternative phylogeny more consistent with morphology-based opinions on species relationships. When evaluated in light of current habitat associations and geographic distributions, the reproductive data suggest that the Miocene diversification of modern dasyurids may have been correlated with the invasion of dry forest or woodland habitats.     

Stem tilting in the inter‐tropical cactus Echinocactus platyacanthus: an adaptive solution to the trade‐off between radiation acquisition and temperature control

While plants require radiation for photosynthesis, radiation in warm deserts can have detrimental effects from high temperatures. This dilemma may be solved through plant morphological attributes. In cold deserts, stem tilting keeps reproductive organs warm by increasing radiation interception at the cost of decreased annual light interception. Conversely, little is known about stem tilting in warm deserts. We hypothesised that stem tilting in Echinocactus platyacanthus prevents high temperatures near the apex, where reproduction occurs. The study was conducted in the warm, inter‐tropical portion of the Chihuahuan Desert, Mexico. We found that cacti preferentially tilted towards the south, which reduced temperatures of reproductive organs during the hot season, but increased total annual near‐apex PAR interception. Tilting also maximised reproduction, a likely consequence of temperature control but perhaps also of the difficulty in translocating photosynthates in cacti; therefore, annual energy acquisition near floral meristems may be largely allocated to reproduction. Unlike plants of higher latitudes, in inter‐tropical deserts sunlight at noon comes either from the north or the south, depending on the season, and thus stem tilting may more strongly affect total annual radiation received in different portions of the stem. Inter‐tropical cacti can synchronise reproduction with irradiance peaks if flowering occurs in a specific (north or south) portion of the stem; also, they effectively solve the conflict between maximising annual PAR interception and minimising temperature at the hottest time of day. Notably, the two inter‐tropical cacti in which stem tilting has been studied successfully solve this conflict.     

Soil extracellular enzyme activities correspond with abiotic factors more than fungal community composition

Soil extracellular enzymes are the proximal drivers of decomposition. However, the relative influence of climate, soil nutrients and edaphic factors compared to microbial community composition on extracellular enzyme activities (EEA) is poorly resolved. Determining the relative effects of these factors on soil EEA is critical since changes in climate and microbial species composition may have large impacts on decomposition. We measured EEA from five sites during the growing season in March and 17 sites during the dry season in July throughout southern California and simultaneously collected data on climate, soil nutrients, soil edaphic factors and fungal community composition. The concentration of carbon and nitrogen in the soil and soil pH were most related to hydrolytic EEA. Conversely, oxidative EEA was mostly related to mean annual precipitation. Fungal community composition was not correlated with EEA at the species, genus, family or order levels. The hyphal length of fungi was correlated with EEA during the growing season while relative abundance of taxa within fungal phyla, in particular Chytridiomycota, was correlated with the EEA of beta-glucosidase, cellobiohydrolase, acid phosphatase and beta-xylosidase in the dry season. Overall, in the dry season, 35.3 % of the variation in all enzyme activities was accounted for by abiotic variables, while fungal composition accounted for 27.4 %. Because global change is expected to alter precipitation regimes and increase nitrogen deposition in soils, EEA may be affected, with consequences for decomposition.     

Nitrogen allocation in Mojave Desert winter annuals

Summary Nitrogen contents and allocation were examined in winter annuals at two Mojave Desert sites near Boulder City, Nevada. Application of 10 g N m^-2 as NH₄NO₃ increased production 0- to 7-fold in species growing on a sandy soil (an Entisol) but fertilizer had no effect on plants on an alluvium (an Aridisol). Tissue nitrogen comprised 0.09–3.5% of dry weight with the lowest concentrations found in vegetative organs of nitrogen-responsive plants. During development, nitrogenpoor species showed only minor changes in nitrogen concentration and allocation compared with more nitrogen-rich species. Maximum reproductive nitrogen allocation varied among species from 43 to 67%, while reproductive biomass allocation was 31 to 51%. Fertilizer increased reproductive biomass allocation by 7 to 16%, reproductive nitrogen concentrations by 120 to 260%, and eaf and root nitrogen concentrations by 200 to 615% in nitrogen-deficient plants. Nitrogen-poor plants appear to allocate nitrogen to reproduction at the expense of vegetative organs throughout the life cycle.     

The productive and reproductive biology of flowering plants

The life cycles, programme of energy expenditure and allocation to reproduction, and the reproductive efforts of three wildAllium species, i.e.,A. Victorialis ssp.platyphyllum, A. monanthum, andA. Grayi, all native to Japan, were studied and compared. Furthermore, their adaptive strategies were discussed from the point of view of life history strategy. First, the reproductive systems, number of male and female gametes borne, and the number and size of propagules produced were critically investigated. In order to estimate the crude reproductive efficiency (sensu Harper and Ogden, 1970) of these species, sequential harvests were taken and the plants were divided into their component structures, dried and weighed. The quantity of dry weight allocated to sexual or vegetative reproduction was obtained by weighing the seeds, bulbils, or bulblets produced at the end of the season. A. Victorialis ssp.platyphyllum showed a rather low reproductive effort. However, the mean seed output per plant was 34.8±16.8 and the productivity appeared very constant every season. Thus, in the natural populations young plants are borne and recruited every season by means of sexual reproduction. A. monanthum was found to be characterized by annual type dry matter economy. The sexuality and reproductive systems of this species turned out to be extremely complex, and ten different reproductive types were distinguished. The exceedingly low efficiency of sexual reproduction in this species is apparently supplemented by vegetative propagation. The dry matter allocation to daughter bulbs at final harvest was very high; whereas the allocation to sexual reproduction was extremely low. InA. Grayi (a polyploid complex of 4X, 5X, and 6X), a surprisingly high amount of the total annual net assimilate is allocated to the bulbils and bulblets. On the other hand, sexual reproductive effort in this species is exceedingly low, even in obligate amphimictic plants. Thus, the recruitment of individuals in a population of this species appears to be largely dependent on vegetative reproduction. Considering the number of bulbils produced in the scape heads, their dispersibility, germinability, and rapid growth after sprouting, the bulbils evidently possess a function almost comparable to seeds. This species no doubt possesses an adapative strategy to unstable, open habitats exposed to frequent disturbances. It is concluded that the life history strategies of plants, as characterized here in this paper for three wildAllium species, have doubtlessly differentiated by adapting to the respective ecological backgrounds of their habitats.     

Reproduction in the agile wallaby Macropus agilis (Gould) in the tropical lowlands of the Northern Territory: opportunism in a seasonal environment

The agile wallaby, Macropus agilis, abounds on the tropical lowlands of the Northern Territory (12° 30′-13°S). The climate is markedly monsoonal and has an annual mean rainfall of 135 cm. During the rainy season, much of the wallaby's major habitat, the extensive coastal plains adjacent to the rivers, becomes flooded. The agile wallaby breeds continuously in captivity but not necessarily so in the wild, although fecundity was very high indeed. Reproduction was more successful on fertilized, nitrogenous pastures of farmland than on grasses of lower nutritive value. Breeding declined least (5%) in the dry season droughts as pastures dwindled and dried out, and most (> 45% in one year) during the rainy season in bushland where the dry season feeding grounds were inundated. The more quickly that the open riverine plains were flooded after the rains began, the higher the loss of breeding and the mortality of pouch-young. The mortality rates of young were similar on both improved and native pastures, however, with only about 35% of new-born surviving to 7 months in the pouch. Wallabies on the improved pastures were at higher densities than on the native pastures which may have influenced that mortality. That females became anoestrous during pregnancy, subsequently gave birth but did not have the usual post-partum oestrus makes the agile wallaby similar to the desert red kangaroo, M. rufus, in its method of entering anoestrus. Also as in that species, anoestrous lactating females can return to oestrus later. The agile wallaby is only the second macropodid marsupial for which these adaptations have been described. Such an adaptation in a desert species has a clear reproductive advantage which is not obvious in this tropical macropodid. Unlike the agile wallaby, the Asiatic waterbuffalo, (Babulis babulus) and domestic cattle, (Bostaurus and B. indicus) which have been introduced to its range breed and give birth on the rains. Such is the reproductive pattern in the tropical ruminants of East Africa also. Seasonal breeding is phylogenetically possible in macropodid marsupials because southern species. M fuliginosus, M. Eugenii, M. r. rufogriseus and Setonix brachyunis, do so in the mediterranean environment there. They time their breeding so that young leave the pouch when pastures are flush after the winter rains. The suggested reason for these differences in breeding of macropodids is seasonal unpredictability. Despite the enormous difference in rainfalls, weekly variability in pasture growth throughout the year is actually the same in monsoonal northern Australia as in the desert, and greater than in southern Australia. Environmental unpredictability may therefore be as important as the overall level of pasture growth in determining reproductive strategies of macropodid marsupials. For the tropical agile wallaby, the unpredictability can cause an absolute shortage or absolute abundance of food in the usual dry season depending on the length of the rainy season and on unseasonal cyclonic weather. It can also cause a relative shortage of food in the rainy season due to flooding of the best feeding grounds. It is concluded that reproduction in the wild mainly depends on the quantity and quality of the food supply.     

Geographical variation in reproductive investment across avian assemblages in Europe: effects of environmental drivers differ between altricial and precocial species

Reproductive traits provide information about the ways by which available resources are allocated during breeding. We tested for environmental drivers of large scale geographical patterns in assemblage mean clutch size, number of broods and overall reproductive investment per breeding season in European birds. We combined data about geographical distribution with published information about reproductive traits, and calculated mean trait values for avian assemblages occurring in 50 × 50 km grid cells. In total, we employed data from 499 species and 2059 assemblages. As the time available for breeding and the amount of food limit the reproductive effort, we related the geographical variation in reproductive traits to the length of breeding season, normalized difference vegetation index (NDVI) as a surrogate of resource availability, and its seasonality. Geographical patterns in traits may differ between reproductive modes, thus we performed the analyses separately for altricial Passerines (n = 203) and precocial non‐passerine species (n = 164) and controlled for the effect of taxonomy. Large clutches dominated in areas with high NDVI and, in precocial birds, also in areas with high annual seasonality and a long breeding season. High number of broods and high overall reproductive investment dominated in areas with a long breeding season, and high number of broods was found also in areas with low annual seasonality, but only in precocial species. High overall reproductive investment dominated in highly productive areas and also in areas with low annual seasonality in both groups. The increase in reproductive investment is caused mostly by an increase in the number of broods related to the length of season and partly by increase in clutch size related to NDVI. We found a negative correlation between clutch size and the number of broods in Passerines, which might suggest a trade‐off between these traits. Processes behind trait patterns differ between altricial and precocial species.     

Reproductive allocation of biomass and nitrogen in annual and perennial Lesquerella crops

* BACKGROUND AND AIMS: The use of perennial crops could contribute to increase agricultural sustainability. However, almost all of the major grain crops are herbaceous annuals and opportunities to replace them with more long-lived perennials have been poorly explored. This follows the presumption that the perennial life cycle is associated with a lower potential yield, due to a reduced allocation of biomass to grains. The hypothesis was tested that allocation to perpetuation organs in the perennial L. mendocina would not be directly related to a lower allocation to seeds. * METHODS: Two field experiments were carried on with the annual Lesquerella fendleri and the iteroparous perennial L. mendocina, two promising oil-seed crops for low-productivity environments, subjected to different water and nitrogen availability. * KEY RESULTS: Seed biomass allocation was similar for both species, and unresponsive to water and nitrogen availability. Greater root and vegetative shoot allocation in the perennial was counterbalanced by a lower allocation to other reproductive structures compared with the annual Lesquerella. Allometric relationships revealed that allocation differences between the annual and the perennial increased linearly with plant size. The general allocation patterns for nitrogen did not differ from those of biomass. However, nitrogen concentrations were higher in the vegetative shoot and root of L. mendocina than of L. fendleri but remained stable in seeds of both species. * CONCLUSIONS: It is concluded that vegetative organs are more hierarchically important sinks in L. mendocina than in the annual L. fendleri, but without disadvantages in seed hierarchy.