Alternative adaptations, sympatric speciation and the evolution of parasitic, inquiline ants

Inquiline ant species are workerless social parasites whose queens reproduce in colonies of other species alongside the host queens. Inquilines arise either when one non-parasitic species evolves into an inquiline parasite of another non-parasitic species (the interspecific hypothesis), or by the speciation of intraspecific inquilines from their host stock (the intraspecific hypothesis): it is unlikely that inquilines evolve from other forms of social parasite. This paper reviews the evidence for and against the inter-and intraspecific hypotheses. All inquilines are close phylogenetic relatives of their host species (loose ‘Emery's rule’), and some are their host's closest relative (strict ‘Emery's rule’). A problem for the interspecific hypothesis is how to explain the strict Emery's rule, because phylogenetic constraints on host choice are probably quite weak. By contrast, the intraspecific hypothesis has difficulty accounting for the parasites' sympatric reproductive isolation. Facultative polygyny, in which queens may found colonies alone or by adoption into an existing multi-queen colony, should promote the evolution of small intraspecific inquilines. This is because small colony-founding queens should preferentially seek adoption, which provides the opportunity to produce a sexual-only brood. We suggest that microgynes, i.e. miniature queens found in some polygynous ants, represent such parasites. We review the evidence that inquiline species have evolved intraspecifically from microgynes in Myrmica ants. The coexistence within a species of a monogynous (singly-queened) and a polygynous form is probably a phenomenon usually unconnected with inquiline evolution. The reproductive isolation of intraspecific inquilines plausibly arises from divergent breeding behaviour associated with the parasites' small size. Such divergence could involve either a temporal separation in mating episodes, with small parasites maturing early, or a spatial separation, with small males being sexually-selected to mate near the nest with small queens seeking adoption, instead of in mating aggregations. We conclude that inquiline species strictly following Emery's rule could have evolved by the intraspecific route. If so, such species provide evidence for West-Eberhard's “alternative adaptation” hypothesis that between-species diversity frequently stems from diversity within species. They also represent likely cases of sympatric speciation. We suggest work on the parasites' phytogeny, genetics, behaviour and mating biology to test these conclusions further.     

Influence of Complexation on the Uptake by Plants of Iron, Manganese, Copper and Zinc: I. EFFECT OF EDTA IN A MULTI-METAL AND COMPUTER SIMULATION STUDY

After growing barley (Hordeum vulgare L.) in nutrient solutionscontaining EDTA, uptake of the nutrient metals was determinedat three harvests and concentrations of the various chemicalspecies of each metal in the growth solutions was modelled bycomputer simulation. Complexation with EDTA had different effectson the uptake of the ions Fe³⁺, Mn²⁺, Cu²⁺, and Zn²⁺. At thehighest EDTA level (EDTA/Fe=2/l) the plants were chlorotic andgrowth was inhibited. This is attributed to a deficiency inZn rather than in Fe. The critical level of free Zn²⁺ requiredin nutrient solutions for healthy growth was found to be approximately10^–1010^–10 mol dm^–3, which is consistent withthat found by earlier workers for other plant species. Barleytolerated much lower levels of the free ions of copper and ironwithout exhibiting any obvious adverse effects. Key words: EDTA, micronutrients, trace metals, computer simulation, deficiencies, absorption, iron, manganese, copper, zinc     

Rice responses to drought under carbon dioxide enrichment. 2. Photosynthesis and evapotranspiration

Future climate change is projected to include a strong likelihood of continued increases in atmospheric carbon dioxide concentration ([CO₂]) and possible shifts in precipitation patterns. Due mainly to uncertainties in the timing and amounts of monsoonal rainfall, drought is common in rainfed rice production systems. The objectives of this study were to quantify the effects and possible interactions of [CO₂] and drought stress on rice (Oryza sativa, L.) photosynthesis, evapotranspiration and water-use efficiency. Rice (cv. IR-72) was grown to maturity in eight naturally sunlit, plant growth chambers in atmospheric carbon dioxide concentrations [CO₂] of 350 and 700 μmol CO₂ mol^–1 air. In both [CO₂], water management treatments included continuously flooded controls, flood water removed and drought stress imposed at panicle initiation, anthesis, and both panicle initiation and anthesis. Potential acclimation of rice photosynthesis to long-term [CO₂] growth treatments of 350 and 700 μmol mol^–1 was tested by comparing canopy photosynthesis rates across short-term [CO₂] ranging from 160 to 1000 μmol mol^–1. These tests showed essentially no acclimation response with photosynthetic rate being a function of current short-term [CO₂] rather than long-term [CO₂] growth treatment. In both long-term [CO₂] treatments, photosynthetic rate saturated with respect to [CO₂] near 510 μmol mol^–1. Carbon dioxide enrichment significantly increased both canopy net photosynthetic rate (21–27%) and water-use efficiency while reducing evapotranspiration by about 10%. This water saving under [CO₂] enrichment allowed photosynthesis to continue for about one to two days longer during drought in the enriched compared with the ambient [CO₂] control treatments.     

Diving behaviour of the Shy Albatross Diomedea cauta in Tasmania: initial findings and dive recorder assessment

The diving behaviour of the Shy Albatross Diomedea cauta was investigated using archival time-depth recorders (TDRs) and maximum depth gauges (MDGs). Data from birds carrying multiple devices and from diving simulations indicated that the degree of correspondence between TDRs and MDGs varied with the dive depth, duration and frequency, as well as with body placement. The MDGs were the most reliable when the diving depth was greater than 0.5 m, when the diving frequency was low and when gauges were placed on the birds' backs. The TDRs were used during late incubation and early chick rearing in 1994. Fifty-two dives (0.4 m) were recorded during 20 foraging trips of 15 individuals. The majority of dives were within the upper 3 m of the water column and lasted for less than 6 s. However, dives to 7.4 m and others lasting 19 s were recorded. The albatrosses dived between 07.00 h and 22.00 h, with peaks in their diving activity near midday and twilight. Mean diving depth varied throughout the day. with the deepest dives occurring between 10.00 h and 12.00 h. Two dive types were identified on the basis of the relationship between dive depth and descent rate. Plunge dives were short (5 s), and the birds reached a maximum depth of 2.9 m. Swimming dives were both longer and deeper. The characteristics of Shy Albatross plunge dives were similar to those of gannets Morus spp., which are known to be proficient plunge divers. Swimming dives suggest that Shy Albatrosses actively pursue prey underwater.     

Insect diversity: facts,fiction and speculation*

Biologists are still trying to grasp the global dimensions of the phylum Arthropoda and its major class the Insecta, in spite of the fact that over a million species of arthropods have been described. The canopy of rain forest trees is believed by many to hold the key to the immense diversity of insects. In recent years the use of knock-down insecticides to sample insects from rain forest canopy has revealed information on the canopy's arthropod inhabitants and community structure. The sampling techniques involved are outlined and data reviewed on taxonomic and guild structure, species abundance, body size and biomass of insects, and the faunal similarity of trees. Calculations by Erwin (1982), based on knock-down insecticide studies of the beetle fauna of one species of Central American tree, suggest there may be 30 million species of tropical forest arthropods. Reanalysis of these calculations, using additional data, produces a range of possible estimates from about 10 to 80 million. The unknown range of plant host-specificities of tropical insects is the main weakness of this method of calculation. Assessment of the faunal importance of the canopy in relation to that of other rain forest biotopes requires comparative quantitative studies. The preliminary results of one such simple study suggest that over 42 million arthropods may be found in a hectare of Seram rain forest (at the time of study), and that 70% occur in the soil and leaf litter and 14% in the canopy. They also suggest that Collembola and Acarina are the dominant groups in this hectare, and that there are as many ants as all the other insects (excluding Collembola).     

Prospects for genetically modified crops

Genetically modified (GM) crops have been in use commercially around the world for almost a decade. This review covers the successes and failures of GM crop varieties in that time, the current status of GM crop adoption and the traits that are being used. It also describes some of the GM crops that might come on to the market in the next decade. The barriers in the way of GM crop development in Europe, including consumer hostility, the difficulty in gaining official approval and discriminatory labelling laws are discussed.     

A new genus of frogmouth (Podargidae) from the Solomon Islands – results from a taxonomic review of Podargus ocellatus inexpectatus Hartert 1901

The frogmouth taxon occurring on the Solomon Islands has been known as Podargus ocellatus inexpectatus , a subspecies endemic to four islands in the Solomon Islands of a species that also inhabits New Guinea and Australia. Our morphological, osteological and molecular studies support recognition of inexpectatus at the species level, and further reveal that it merits placement in its own genus, which we describe here. Compared with the two other extant podargid genera, inexpectatus does not seem to be more closely related either to Batrachostomus (confined to the Indo-Malayan faunal region) or to Podargus of the Papuan–Australian faunal region. We also review the specimen history of the Solomon Islands Frogmouth, and discuss what little is known about its ecology, natural history and distribution.     

Endemic Cyprus Warbler Sylvia melanothorax and colonizing Sardinian Warbler Sylvia melanocephala show different habitat associations

Anthropogenic habitat change and assisted colonization are promoting range expansions of some widespread species with potential consequences for endemic fauna. The recent colonization of Cyprus by breeding Sardinian Warblers Sylvia melanocephala has raised concerns that it might be displacing the closely related and endemic Cyprus Warbler Sylvia melanothorax. Habitat associations of both species were examined using models of abundance within the 95% density kernel of the Sardinian Warbler’s range and also outside this range for Cyprus Warbler. Within the Sardinian Warbler’s range, the two species were associated with subtly different scrub habitats. Outside the Sardinian Warbler’s range, the Cyprus Warbler differed again in its habitat association, but this probably resulted from marked differences in habitat extent and availability in different parts of the island rather than from competitive displacement, as none of the habitat or land‐use elements differentially associated with Cyprus Warblers was positively associated with Sardinian Warbler occurrence. This suggests that the Sardinian Warbler has exploited a different niche, rather than displacing the endemic species, and has perhaps benefitted from changing land‐use patterns, particularly recent fallows and abandoned agriculture, in contrast to the stronger association of Cyprus Warblers with semi‐natural scrub.     

Ecological specialization in mycorrhizal symbiosis leads to rarity in an endangered orchid

Terrestrial orchid germination, growth and development are closely linked to the establishment and maintenance of a relationship with a mycorrhizal fungus. Mycorrhizal dependency and specificity varies considerably between orchid taxa but the degree to which this underpins rarity in orchids is unknown. In the context of examining orchid rarity, large scale in vitro and in situ germination trials complemented by DNA sequencing were used to investigate ecological specialization in the mycorrhizal interaction of the rare terrestrial orchid Caladenia huegelii. Common and widespread sympatric orchid congeners were used for comparative purposes. Germination trials revealed an absolute requirement for mycorrhisation with compatibility barriers to germination limiting C. huegelii to a highly specific and range limited, efficacious mycorrhizal fungus. DNA sequencing confirmed fidelity between orchid and fungus across the distribution range of C. huegelii and at key life history stages within its life cycle. It was also revealed that common congeners could swap or share fungal partners including the fungus associated with the rare orchid but not vice versa. Data from this study provides evidence for orchid rarity as a cause and consequence of high mycorrhizal specialization. This interaction must be taken into account in efforts to mitigate the significant extinction risk for this species from anthropogenically induced habitat change and illustrates the importance of understanding fungal specificity in orchid ecology and conservation.