Anne C. Gaskett

Orchid pollination by sexual deception : pollinator perspectives

The extraordinary taxonomic and morphological diversity of orchids is accompanied by a remarkable range of pollinators and pollination systems. Sexually deceptive orchids are adapted to attract specific male insects that are fooled into attempting to mate with orchid flowers and inadvertently acting as pollinators. This review summarises current knowledge, explores new hypotheses in the literature, and introduces some new approaches to understanding sexual deception from the perspective of the duped pollinator. Four main topics are addressed: (1) global patterns in sexual deception, (2) pollinator identities, mating systems and behaviours, (3) pollinator perception of orchid deceptive signals, and (4) the evolutionary implications of pollinator responses to orchid deception, including potential costs imposed on pollinators by orchids. A global list of known and putative sexually deceptive orchids and their pollinators is provided and methods for incorporating pollinator perspectives into sexual deception research are provided and reviewed.

Changes in male mate choice in a sexually cannibalistic orb-web spider (Araneae: Araneidae)

In theory, male mate choice should occur when the costs of copulation, in terms of future mating opportunities, are high. The criteria males use to choose mates may change depending upon male mating history and the potential for future matings. We examine male mate choice in the St. Andrews Cross Spider (Argiope keyserlingi Araneae: Araneidae). Laboratory experiments revealed that death and injury caused by female sexual cannibalism limits males to a maximum of two copulations. We assessed the mate choices of virgin and mated males for females of different reproductive status. We used field and laboratory choice bioassays involving airborne and web-based pheromones. In field experiments, wild males were strongly attracted to webs built by laboratory-raised virgin females. Webs from mated females did not attract males. Male mate choice was affected by male reproductive status: while virgin males strongly preferred penultimate and virgin females to mated females, mated males were apparently indifferent to females of different mating status. Such post-copulatory changes in male mate choice have not been previously documented, and may reflect a decreased potential for future mating.

Quantifying Variation in the Ability of Yeasts to Attract Drosophila melanogaster

Yeasts that invade and colonise fruit significantly enhance the volatile chemical diversity of this ecosystem. These modified bouquets are thought to be more attractive to Drosophila flies than the fruit alone, but the variance of attraction in natural yeast populations is uncharacterised. Here we investigate how a range of yeast isolates affect the attraction of female D. melanogaster to fruit in a simple two choice assay comparing yeast to sterile fruit. Of the 43 yeast isolates examined, 33 were attractive and seven repellent to the flies. The results of isolate-versus-isolate comparisons provided the same relative rankings. Attractiveness varied significantly by yeast, with the strongly fermenting Saccharomyces species generally being more attractive than the mostly respiring non- Saccharomyces species (P = 0.0035). Overall the habitat (fruit or other) from which the isolates were directly sampled did not explain attraction (P = 0.2352). However, yeasts isolated from fruit associated niches were more attractive than those from non-fruit associated niches (P = 0.0188) regardless of taxonomic positioning. These data suggest that while attractiveness is primarily correlated with phylogenetic status, the ability to attract Drosophila is a labile trait among yeasts that is potentially associated with those inhabiting fruit ecosystems. Preliminary analysis of the volatiles emitted by four yeast isolates in grape juice show the presence/absence of ethanol and acetic acid were not likely explanations for the observed variation in attraction. These data demonstrate variation among yeasts for their ability to attract Drosophila in a pattern that is consistent with the hypothesis that certain yeasts are manipulating fruit odours to mediate interactions with their Drosophila dispersal agent.

Male mate choice and patterns of paternity in the polyandrous, sexually cannibalistic orb-web spider Nephila plumipes

Studies that investigate patterns of paternity in polyandrous species typically employ double-mating trials, in which the paternity share of each male is established by either the sterile male technique or using genetic markers. However, polyandrous females may mate with more than two males and, in some species, triple-mating trials produce different patterns of paternity from double-mating trials. We investigated patterns of paternity share in triple-mating trials of the sexually cannibalistic orb-web spider Nephila plumipes. These experiments reveal little quantitative changes to paternity share when more than two males mate with the female; the third male apparently diluted the fertilisation success of the second male but not of the first male. Sexual cannibalism had little impact on the fertilisation success of the first male, but greatly increased the fertilisation success of the third male. When offered a choice, males did not prefer to mate with virgin over mated females, but males that chose virgin females were significantly heavier than those that chose mated females. ZOce and er w Mlg et

The volatile organic compounds of introduced and native dung and carrion and their role in dung beetle foraging behaviour

1. The decomposition of biological material produces a plethora of volatile organic compounds (VOCs), which are implicated in the foraging behaviour of coprophagous and necrophagous insects. Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) have an acute olfactory system used to locate food resources. Accordingly, identification of food resource VOCs potentially used in food location is integral to understanding dung beetle foraging ecology.

Convergent evolution of sexual deception via chromatic and achromatic contrast rather than colour mimicry

The Orchidaceae is characterised by the repeated evolution of sexual deception, one of the most specialised pollination strategies. In orchids, sexual deception involves long-range pollinator attraction via mimicry of female insect sex pheromones. At close range, visual signals involving colour mimicry, contrast to the background, and exploitation of pollinator sensory biases could attract pollinators, but remain largely untested. Here we focus on a remarkable system in which species from two only distantly related sexually deceptive orchid genera with strikingly different flowers (Drakaea livida and three species of Caladenia) share the same pollinator, males of the thynnine wasp Zaspilothynnus nigripes. We used spectral reflectance measurements and modelling to investigate pollinator perception of colour, including the first examination of overall colour patterns in flowers via colour pattern geometry analyses. Rather than closely matching the colours of female Z. nigripes, these orchids had strong chromatic and achromatic contrast against their backgrounds. For Caladenia, the sepals and petals show high contrast, while in D. livida, which has diminutive petals and sepals, it is the labellum that contrasts strongly against the background. Despite varying in colour, the Caladenia species all had strong within-flower contrast between a UV-bright central target (column and labellum) and a corolla of radiating stripes (petals and sepals). The colour pattern geometry analyses also indicated that the orchids’ overall colour patterns are highly conspicuous against their backgrounds. Contrast, UV, and target patterns could all enhance detection, and exploit pollinators’ innate preferences. Since colour contrast may function with a range of colours and floral forms, attracting pollinators via contrast rather than visual mimicry may be a critical but previously overlooked process facilitating the evolution of sexual deception.

Flowers, phenology and pollination of the endemic New Zealand greenhood orchid Pterostylis brumalis

ABSTRACT Greenhood orchids are most diverse in New Zealand and Australia, with at least some species pollinated by sexual deception of male fungus gnats. Preliminary investigation of endemic New Zealand Pterostylis brumalis via field phenology, hand-pollination and insect trapping revealed P. brumalis is self-compatible but relies on insects for cross pollination. Natural fruit-set was low, averaging 2.6% across two populations. High fruit-set in hand-pollinated flowers (66.7%) indicates severe pollinator limitation. In comparison, only one collected insect, a female Mycetophila vulgaris, bore orchid pollinia. Potential pollinator attractants (colour and scent) were analysed. UV reflectance may enhance pollinator detection of the flower against the background, but visual modelling generated conflicting results: a categorical fly vision model indicated excellent camouflage against herbivory, but a generic tetrachromat vision model suggested flowers are easily distinguishable against the background. Both models have flaws that create issues with drawing conclusions, and identification of the pollinator is required to understand their visual system further. Scanning electron microscopy (SEM) revealed putative scent-releasing osmophores (lateral sepals) and uniform papillate trichomes (labellum) that may mechanically direct pollinators into the floral trap. The trap is a pressure sensitive labellum that temporarily traps the visitor inside the flower.

Pollination and insect visitors to the putatively brood-site deceptive endemic spurred helmet orchid, Corybas cheesemanii

Abstract Deceptive orchids typically employ food or sexual deception of male insects. Brood-site deception, when flowers are pollinated by female flies fooled into attempting oviposition, is less well characterized for orchids but is probably common among New Zealand terrestrial orchid species. Helmet orchids (Corybas, Nematoceras and related genera) are considered specialist brood-site deceivers that mimic mushrooms and are pollinated by ovipositing female fungus gnats (Mycetophilidae). We monitored pollination in the endemic spurred helmet orchid, Corybas cheesemanii, and trapped insects visiting orchids and sympatric mushrooms. Flowering occurred from mid-May to early July, each flower lasted 23.14 d, 78.5% of the population flowered (147/194 plants), and c.25% of these set seed naturally (50% of bagged orchids set seed via autonomous self-pollination). Traps caught mostly flies (Mycetophilidae, Anisopodidae and Lauxaniidae), but orchid, mushroom and control traps did not attract significantly different fauna. Seven mycetophilids (of both sexes) visited both orchids and mushrooms, but we found no compelling evidence that C. cheesemanii was pollinated by these. Only some mycetophilid individuals caught above orchids were small enough to enter orchids, but none carried orchid pollinia. If C. cheesemanii is fungus gnat-pollinated, the range of species and sexes attracted suggests a more generalist pollination strategy than assumed.

Dung beetles in an avian-dominated island ecosystem: feeding and trophic ecology

Globally, dung beetles (Scarabaeidae: Scarabaeinae) are linked to many critical ecosystem processes involving the consumption and breakdown of mammal dung. Endemic New Zealand dung beetles (Canthonini) are an anomaly, occurring at high abundance and low diversity on an island archipelago historically lacking terrestrial mammals, except bats, and instead dominated by birds. Have New Zealand’s dung beetles evolved to specialise on bird dung or carrion, or have they become broad generalist feeders? We test dietary preferences by analysing nitrogen isotope ratios of wild dung beetles and by performing feeding behaviour observations of captive specimens. We also use nitrogen and carbon stable isotopes to determine if the dung beetle Saphobius edwardsi will consume marine-derived carrion. Nitrogen isotope ratios indicated trophic generalism in Saphobius dung beetles and this was supported by behavioural observations where a broad range of food resources were utilised. Alternative food resource use was further illustrated experimentally by nitrogen and carbon stable isotope signatures of S. edwardsi, where individuals provided with decomposed squid had δ15N and δ13C values that had shifted toward values associated with marine diet. Our findings suggest that, in the absence of native mammal dung resources, New Zealand dung beetles have evolved a generalist diet of dung and carrion. This may include marine-derived resources, as provided by the seabird colonies present in New Zealand forests before the arrival of humans. This has probably enabled New Zealand dung beetles to persist in indigenous ecosystems despite the decline of native birds and the introduction of many mammal species.

Exotic species enhance response diversity to land-use change but modify functional composition

Two main mechanisms may buffer ecosystem functions despite biodiversity loss. First, multiple species could share similar ecological roles, thus providing functional redundancy. Second, species may respond differently to environmental change (response diversity). However, ecosystem function would be best protected when functionally redundant species also show response diversity. This linkage has not been studied directly, so we investigated whether native and exotic pollinator species with similar traits (functional redundancy) differed in abundance (response diversity) across an agricultural intensification gradient. Exotic pollinator species contributed most positive responses, which partially stabilized overall abundance of the pollinator community. However, although some functionally redundant species exhibited response diversity, this was not consistent across functional groups and aggregate abundances within each functional group were rarely stabilized. This shows functional redundancy and response diversity do not always operate in concert. Hence, despite exotic species becoming increasingly dominant in human-modified systems, they cannot replace the functional composition of native species.