Andrew Lack

Relationships between flowering phenology, plant size and reproductive success in Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.

Genetics of colonizing and established populations of Primula veris

Primula veris, a long-lived iteroparous herb which has declined in abundance in southern England since the 1940s, has recently been observed colonizing disturbed sites. The genetic structure of P. veris populations in the Oxford region was investigated to determine whether colonizing populations differ from those longer established. Nine enzyme systems, revealing 19 presumptive loci, were screened in 11 established and seven colonizing populations. Levels of variation were low in all populations, and all populations except one approximated panmixia. The proportion of the total variation among populations was small (FST=0.039) as were genetic distances between populations. No significant differences in genetic structure were recorded between established and colonizing populations. The lack of differentiation within and between populations is surprising for a long-lived, insect pollinated species with gravity dispersed seeds, for which gene flow would be expected to be spatially restricted.

Phosphoglucose isomerase (EC 5.3.1.9) isozymes in diploid and tetraploid Polygala species: evidence for gene duplication and diversification

Phosphoglucose isomerase (PGI, EC 5.3.1.9) isozymes were examined by electrophoresis in the three common British species of Polygala, the diploids P. serpyllifolia and P. calcarea and the tetraploid P. vulgaris. All three species share the most anodal band which is probably the form of the enzyme occurring in the plastids (PGI-I). Individuals of the two diploid species each have one further band, probably the cytosolic form of the enzyme (PGI-2), with two phenotypes in each species, one of which is common to both. Individuals of P. vulgaris have one, three or six PGI-2 bands and 10 different homozygous phenotypes were observed. The commonest phenotypes in most populations show three bands with a central heterodimeric band staining more intensely than the outer two bands. Although P. vulgaris is a predominantly self-fertilising species with no vegetative spread, many populations or sub-populations consist almost entirely of three-banded phenotypes. This indicates that the three-banded phenotypes are “fixed heterozygotes” resulting from the duplication of the Pgi-2 locus in the tetraploid. Heterozygotes between different three-banded forms show six bands in which the staining intensities are consistent with random pairing of the subunits. Plants showing silencing of one of the duplicate alleles were found in two small isolated areas in West Glamorgan. The biochemical diversity produced by the multiplicity of enzyme morphs in the tetraploid P. vulgaris may have contributed to its success. It is the most common, widespread and ecologically wide-ranging species in the genus in Europe.

Peter G. Ayres: Shaping ecology: the life of Arthur Tansley

To be interested in conservation is to be interested in ecology, and to be interested in ecology one inevitably comes across the towering figure of Sir Arthur Tansley. Peter Ayres is surely wrong when he states that ‘‘few beyond a relatively small group of professional ecologists are aware of the priceless practical legacy that he left to everyone who cares about wildlife and its conservation’’. My surprise is that this is the first full-length biography of him. In fact it is not particularly long and, in nearly every way, all the better for it. Ayres takes a chronological look at Tansley’s life and legacy. As such he gives us a balanced portrait of the great man. Arthur Tansley was born in 1871 in London to liberalminded parents with strong views and intense interest in workers’ education. He fell in love with the English countryside and its plants when sent to boarding school in Sussex, aged 11. These two features shaped his professional life, both within the universities in which he worked and taught, and in the other activities in which he was involved. He was astonishingly clear-sighted and, coupled with intense personal ambition as well as consummate skills as an organiser, what he started nearly always worked. He saw ecology as the means to study and to conserve the British countryside and, in so doing, created the ecology we know today. He saw his place as a pioneer, made sure he met all the influential people and, naturally, made some enemies in doing so. Tansley is now remembered for three main things: founding the New Phytologist in 1902, founding the British Ecological Society (BES), the world’s first, in 1913 (here Ayres adds little to Sheail’s (1987) account), and publishing the enormous The British Islands and their Vegetation in 1939. Of course there is much else, from his study of psychology, including writing a successful book, during a professional and personal crisis in the 1920s to his leading role, during retirement, with the nature conservation movement. He was involved with almost all the main ecological movements. He saw ecology as, fundamentally, a botanical activity. Despite this bias, he quickly recognised the importance of animal ecology as formulated by, especially, Charles Elton. The two men evidently held each other in high regard. Ayres singles out the attempted organising of a Students’ British Flora to be Tansley’s one main failure of clear-sightedness. Tansley admitted that he was no taxonomist, but in fact even this led to his young colleague, Roy Clapham, putting together the Flora we have known for so long as CTW (1952). One thing that rather surprised me from this book is that Tansley does not come across as a keen naturalist. The great naturalist-ecologists come from the next generation, born in the early twentieth century. Tansley’s vision was more all-encompassing: a concern for the countryside certainly, but not for the minutiae that a naturalist examines. Tansley was chairman of the Nature Reserves Committee of the BES, which led directly to the founding of the Nature Conservancy in 1949. He saw nature conservation as essential to the progress of ecology and saw National Nature Reserves as places that should be reserved for scientific ecological work, and shielded from public amenity. How times change! He could be generous and financed a number of projects of colleagues that he admired. He also could be a staunch friend, for example though strongly disagreeing professionally, he never let this come between his friendship for A. Lack (&) Department of Biological and Medical Sciences, Oxford Brookes University, Headington, Oxford OX3 0BP, UK e-mail: ajlack@brookes.ac.uk

Lynn V. Dicks, David A. Showler and William J. Sutherland: Bee conservation—evidence for the effects of interventions

This book is the first in a projected series by the Conservation Evidence Project ‘building into a comprehensive summary of evidence on the effects of conservation interventions for all biodiversity throughout the world’. Quite a claim, and clearly one that will always be ongoing and need updating. The project was set up because the authors felt that conservation intervention has so often been on an ad hoc basis. A proper assessment of evidence was needed to provide a rigorous baseline for all conservation interventions. In the interests of spreading the information as widely as possible, the project provides a website with a peer-reviewed open access journal and including this entire book: http://www.conservationevidence.com/Attachments/ 2931ddc6-c4c1-486b-85ac-df7d48298565.pdf. They invite contributions that detail any evidence, positive or negative, for an intervention in the name of conservation. As such it seems a most laudable project and this is its first booklength fruit. The book runs to only 133 text pages but is divided into twelve chapters and a total of 68 sections. These detail the threats from developments, agricultural change (the one long chapter), pollution, transport, biological resources, fire, non-native and native species and positive measures of providing artificial nest sites, rearing wild bees and education. The authors have found a comprehensive list of detailed studies that genuinely demonstrate evidence of impact. Each study is examined in turn, and the book frequently states: ‘‘...replicated control trials... show...’’. Most usefully, they raise the negative evidence too. Potential threats, problems or solutions that people think are important, or could make an impact, are dealt with briefly with the oft-repeated phrase ‘‘We have captured no evidence for...’’. This surely will stimulate researchers to see whether there is any evidence for any of these issues, for example, connecting areas of natural or semi-natural habitat, re-planting forest, or reducing fertilizer run-off from fields. Almost inevitably most studies are from Europe, especially Britain, and North America, but other parts of the world are represented. There is a strong emphasis on bumblebees, mainly because of the availability of evidence, but there are some interesting studies on solitary bees and a few on honeybees where conservation is the thrust. Some of the evidence is unsurprising, but it is valuable to have it presented in this way; some is more unexpected, such as heathland being no better than restored sites, and restored flower-rich grassland actually being better than old hay meadows. We have heard a great deal about the potential effect of alien species, but evidence for effects on bees seems to be remarkably absent. The most surprising lack of evidence is for any impact on the native bee faunas of the Americas by the introduced Africanized honeybee. Surely this is simply lack of controlled research? The book is short and strictly about the evidence. And, even within this, they state ‘Key Messages’ at the head of each chapter, each a summary of the conclusions, and then, unnecessarily, either repeating these at the start of each section or writing a slightly different, usually longer, summary. References are given in each section of each chapter. This inevitably leads to some overlap though the authors have tried to keep it to a minimum. So is this evidence-based approach the way forward? Emphatically ‘yes’, but it is inevitably limited in scope. The founders of the project draw the parallel with medicine and how that has been revolutionised within the last few A. Lack (&) School of Life Sciences, Oxford Brookes University, Gipsy Lane Campus, Headington, Oxford OX3 0BP, UK e-mail: ajlack@brookes.ac.uk

J. E. Peña, J. L. Sharp and M. Wysoki (eds), Tropical Fruit Pests and Pollinators: Biology, Economic Importance, Natural Enemies and Control

This book consists of 13 unequal chapters by a total of 28 authors and is, fundamentally, a compilation of information about the invertebrate pests of certain tropical fruits and how to control them. After an Introduction the fruits treated are, in order of length, avocado (71 pp.), tropical citrus (45 pp.), banana (44 pp.), pineapple (39 pp.), passion fruit (29 pp.), litchi and longan (29 pp.), mango (27 pp.), papaya (26 pp.), Annona spp. (25 pp.), guava (19 pp.) and minor tropical fruits, comprising durian, mangosteen, rambutan, carambola and Barbados cherry (16 pp.). A final chapter discusses quarantine. There are statistics on where these fruits are grown, quantities and importance for the economy in each main country. The pests considered are largely insects, include mites and nematodes but no vertebrates. Their distributions, life cycles and effects are described along with a comprehensive treatment of control measures, chemical, biological and integrated pest management from many countries. Several chapters have extensive lists of pests in tables, while others treat them in the text, but all the chapters are well laid out with many headings and subheadings. There are numerous references for each chapter. There are some obvious omissions (e.g., breadfruit, cucurbits, cocoa) and no real reason is given for this. The title indicates that it is about pollinators too but the treatment of pollinators is, to say the least, uneven. There is a fairly extensive treatment for avocado, a few pages for passion fruit and Annona, just one page for mango, one for litchi/longan and nothing for the others. This seems poorly thought out and, to be frank, pollination and pollinator details is another book as there is a great deal of information on some of these. The interesting material on pollination that is here gets lost because it is so patchy. Finally the index is seriously inadequate, though this may not matter too much since the book is well laid out itself and easy to navigate. The book is about pests and how to limit or stop their infestations, and in this respect has nothing to say about insect conservation. Pollinators are vital for some of these crops, but the short sections here are mainly about honeybees, with brief reference to other species. As such the message becomes a reiteration of the rather depressing fact that honeybees dominate the pollinator fauna over most of the world including many parts in which they are not native. No indication is given here about their effects on the native pollinator fauna. The book will prove most useful to anyone with an interest in growing the fruits discussed here, as it is a single source of almost everything they would need to know about invertebrate pests and their control. The title should really read ‘‘Tropical Fruit Pests and Their Control: A Compendium’’ or some such, more accurately reflecting the contents and the use of the book.