Leaf miners: The hidden herbivores

Leaf mining is a form of endophagous herbivory in which insect larvae live and feed within leaf tissue. In this review we discuss aspects of leaf miner ecology, and the current evidence for three hypotheses relating to the evolution of this feeding guild. We also present a summary of the literature coverage relating to these herbivores, which have been relatively poorly studied compared with insects that feed externally such as sap suckers and leaf chewers. The majority of published studies concern leaf miners from the northern hemisphere, with a general focus on those species considered to be agricultural, forestry or horticultural pests. In a more detailed case study, we examine aspects of leaf miner ecology of Australian species. At least 114 species have been recorded as leaf miners in Australia in four orders: Coleoptera, Lepidoptera, Diptera and Hymenoptera. Lepidoptera and Diptera are the most speciose orders of Australian leaf miners; Hymenoptera are represented by a single endemic genus and half of all coleopteran miners are species introduced for biological control. Both the known number of leaf‐mining species in Australia and the known number of hosts have increased in recent years following new targeted surveys. Leaf miners in Australia occur in many habitats and feed on a wide variety of host plants in at least 60 families although most individual species are monophagous. Although much of the research on leaf miners in Australia has focused on species that are commercially important pests or biological control agents, studies on fundamental aspects of leaf miner ecology are increasing. We identify a number of research questions aimed at better understanding the ecology of leaf miners in Australia and elsewhere.     

Python phylogenetics: inference from morphology and mitochondrial DNA

We used nucleotide sequences from four mitochondrial genes and structural features of the mitochondrial control region, combined with a revised, previously published, morphological data set to infer phylogenetic relationships among the pythons. We aimed to determine which of two competing hypotheses of relationships of the genera Aspidites and Python best explains the evolutionary and bioegeographical history of the family. All analyses of the combined data recover a set of relationships in which (1) the genus Python is paraphyletic with the two east Asian species, P. reticulatus and P. timoriensis , as the sister lineage to the seven Australo-Papuan python genera. We support recognition of a distinct genus for the P. reticulatus  +  P. timoriensis clade; (2) the remaining species of the genus Python form a clade which is the sister lineage to the remainder of the family; (3) the genus Aspidites is embedded among the Australo-Papuan genera. The seemingly primitive characteristics of Aspidites may be better interpreted as reversals or specializations that have accompanied a switch to burrowing in this genus. Resolution of the relationships among the Australo-Papuan lineages is weak, possibly because of rapid diversification early in the history of the radiation. We assessed the tempo of the Indo-Australian python radiation using a maximum likelihood framework based on the birth–death process. We find strong support for elevated speciation rates during the period when Australia collided with the proto-Indonesian archipelago. The data support an origin for pythons outside Australia, followed by a radiation into Australia during the mid-Tertiary.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 603–619.     

Seedling survival in a northern temperate forest understory is increased by elevated atmospheric carbon dioxide and atmospheric nitrogen deposition

We tested the main and interactive effects of elevated carbon dioxide concentration ([CO₂]), nitrogen (N), and light availability on leaf photosynthesis, and plant growth and survival in understory seedlings grown in an N‐limited northern hardwood forest. For two growing seasons, we exposed six species of tree seedlings (Betula papyrifera, Populus tremuloides, Acer saccharum, Fagus grandifolia, Pinus strobus, and Prunus serotina) to a factorial combination of atmospheric CO₂ (ambient, and elevated CO₂ at 658 μmol CO₂ mol⁻¹) and N deposition (ambient and ambient +30 kg N ha⁻¹ yr⁻¹) in open‐top chambers placed in an understory light gradient. Elevated CO₂ exposure significantly increased apparent quantum efficiency of electron transport by 41% (P<0.0001), light‐limited photosynthesis by 47% (P<0.0001), and light‐saturated photosynthesis by 60% (P<0.003) compared with seedlings grown in ambient [CO₂]. Experimental N deposition significantly increased light‐limited photosynthesis as light availability increased (P<0.037). Species differed in the magnitude of light‐saturated photosynthetic response to elevated N and light treatments (P<0.016). Elevated CO₂ exposure and high N availability did not affect seedling growth; however, growth increased slightly with light availability (R²=0.26, P<0.0001). Experimental N deposition significantly increased average survival of all species by 48% (P<0.012). However, seedling survival was greatest (85%) under conditions of both high [CO₂] and N deposition (P<0.009). Path analysis determined that the greatest predictor for seedling survival in the understory was total biomass (R²=0.39, P<0.001), and that carboxylation capacity (V_cmax) was a better predictor for seedling growth and survival than maximum photosynthetic rate (A_max). Our results suggest that increasing [CO₂] and N deposition from fossil fuel combustion could alter understory tree species recruitment dynamics through changes in seedling survival, and this has the potential to alter future forest species composition.     

A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

Pollination activity by blowflies and honeybees on onions in breeders' cages

Honeybees (Apis mellifera) and three species of blowflies (Calliphora vomitoria, Lucilia caesar and L. sericata) were observed on mass-pollinated populations of onions (Allium cepa) in 4·4 × 3·6 m × 2-2·5 m cages in June-August 1977. Pollination activity at temperatures from 14° to 28°C was compared on the basis of mean time per flower touched on each umbel visit. Honeybees did not forage below 16°C. Above 16°C their mean time per flower was short (1·4 s) and varied little with temperature. For blowflies, it decreased markedly from 12·1 s at 14–15·5°C to 2·7 s at 26°C and above, largely because at low temperatures flies spent long periods quiescent or grooming rather than actively feeding. When flowering was not completely synchronous between cultivars, honeybees were more selective than blowflies, but where flowering was synchronous, both types of pollinator visited the two cultivars at random. For the size of cage used, neither type of insect had a distinct advantage as a pollinator of onions, despite their different behaviour patterns.     

Onion pollination by blowflies and honeybees in large cages

Three experiments to investigate the effect of pollination by blowflies (Calliphora and Lucilia spp.) and honeybees (Apis mellifera) on seed yield and crossing level were made using a mixture of plants from two open-pollinated cultivars of onions (Allium cepa) in breeders' cages. No consistent differences were found between pollinators. In two experiments, fly pollination gave significantly higher yields than bee pollination but in the third, bee pollination gave slightly more seed. Crossing between cultivars, assessed by progeny testing on one cultivar, varied between 22-7 and 55-9%. Differences in crossing level attributable to the pollinators were not consistent and were not or only just statistically significant, but there were significant and substantial differences between the experiments themselves. Where more umbels were produced by the red-bulbed cultivar than by the yellow, higher crossing levels were found in the yellow cultivar. The highest crossing levels were found in the only experiment where insects were not introduced until peak flowering time, but where seed yield was comparatively low. Crossing levels were lowest where the umbel numbers were approximately equal. Individual umbels studied showed that while seed set was usually heaviest at and just before the mid-flowering date, this period could correspond with the lowest levels of crossing during the flowering period, particularly if flowering of the two cultivars was not completely synchronous. In field plots, approximately 75% of crossed seeds sown survived to give harvestable bulbs, but fewer than 50% of selfed seeds did so. In an experiment in which crossing levels from different pollinators were similar but survival levels differed significantly, it was deduced that bees performed more crossing within a cultivar than flies.     

Nectar Production and Floral Characteristics of Tropaeolum majus L. Grown in Ambient and Elevated Carbon Dioxide

Tropaeolum majus (nasturtiums) were grown from seed in growthcabinets, under 380 and 750 ppmv CO₂. Elevated CO₂significantlyincreased nectar secretion rate, both in flowers milked of nectardaily and in once sampled, 3-d-old flowers. Elevated CO₂didnot affect time to flowering, total number of flowers produced,pollen to ovule ratio, or the total or individual concentrationsof nectar amino acids. The dry weight and longevity of individualflowers was also unchanged. Nectar sugar content was unchangedby elevated CO₂in a subset of flowers used to assess the 3-d-oldnectar volume. This subset did not show the same increase innectar volume under elevated CO₂as the full set, resulting inthe concentration of sugars remaining unchanged. Overall, thequantity rather than the quality of the nectar changed underelevated CO₂while flower characteristics remained constant,implying that the identity of pollinators may remain the samewhile foraging behaviour (e.g. number of visits per plant, distancetravelled) may change in the future. Copyright 1999 Annals ofBotany Company Tropaeolum majus, nasturtiums, elevated CO₂, nectar, phenology, floral characteristics, amino acids, pollinator-plant interactions.     

Lipid Metabolism in the Brown Marine Algae Fucus vesiculosus and Ascophyllum nodosum

Lipid metabolism and environmental effects on this process havebeen studied in the marine brown algae Fucus vesiculosus andAscophyllum nodosum. These algae showed very similar patternsof lipid metabolism during 24 h incubations. Labelling from[1-¹⁴C]acetate showed the major labelled lipids to be the ß-alanineether lipid and the neutral lipid fraction in both algae. Ofthe glycolipids, only sulphoquinovosyldiacylglycerol was welllabelled and the phosphoglycerides were all poorly labelled.The major labelled fatty acids were palmitate and oleate, againin both algae, although Fucus vesiculosus also showed significantlabelling of stearate and behenate. Although the amount of fattyacid labelling increased with time, the proportion of labelin palmitate and oleate remained approximately constant. Verylong chain fatty acids (arachidic, behenic) were increasinglylabelled with time. Lowered incubation temperatures decreased labelling of the saturatedfatty acids. Cu²⁺ increased the proportion of oleate labelledin both algae, and of linoleate in Fucus vesiculosus. This cationdecreased the percentage labelling of stearate and myristatein Ascophyllum nodosum. Lipid metabolism in Ascophyllum nodosumwas more sensitive to raised Cu²⁺ levels than in Fucus vesiculosus Key words: Acyl lipid metabolism, Fucus vesiculosus, temperature effects, Ascophyllum nodosum, copper pollution     

Differential effects of elevated nutrient and sediment inputs on survival,growth and biomass of a common larval fish species (Dorosoma cepedianum)

1. Elevated allochthonous inputs of nutrients and sediments to aquatic ecosystems are associated with eutrophication and sedimentation. Reservoirs receive substantial subsidies of nutrients and sediments from catchments due to their large catchment : lake area ratios. We examined the effect of elevated subsidies of sediments and/or dissolved nutrients on the success (survival, growth, biomass and condition factor) of larval gizzard shad (Dorosoma cepedianum), a widespread and dominant omnivorous fish in reservoir ecosystems. 2. We simulated allochthonous agricultural subsides by manipulating dissolved nutrients and sediment inputs in a 2 × 2 factorial design in experimental mesocosms. We predicted that larval fish success would be greater under elevated nutrients. However, we propose two alternative hypotheses with respect to the overall effect of allochthonous sediment inputs. If sediment inputs negatively affect larval gizzard feeding success, larval success would be highest when only nutrients are added and lowest when only sediments are added (+N > +N+S ≥ C > +S). If high turbidity enhances larval foraging activity (due to greater contrast between prey and background), we predict that larval success would be highest when both subsidy types (nutrients and sediment) are elevated, intermediate when either nutrients or sediments are added and the lowest when no subsidies are added (+N+S > +N ≥ +S > C). 3. Our results indicate that elevated nutrient and sediment conditions enhanced larval gizzard shad biomass, but the overall nutrient addition effect was greater than the sediment addition effect (+N ～ +N+S > +S > C). We observed differential effects of nutrient and sediment inputs on larval survival, growth and condition factors. 4. The enhancement of fish biomass in elevated nutrients (+N, +N+S) relative to control conditions was associated with improved gizzard shad survival and not greater growth. The enhancement of fish biomass in the elevated sediment treatment (+S) relative to the control conditions was caused by an increase in survival that more than compensated for a negative effect of sediment addition on growth. 5. Our findings support the recommendation that reservoir management practices must consider the links between land use practices and food web dynamics. Our results suggest that reduction of subsidies of nutrients and sediments to productive reservoirs would decrease survival of larval gizzard shad due to lower food availability.