Wildflower Meadows

contents

To all the volunteers who have come from different walks of life to learn botanical identification skills and have shared their expertise during many happy hours spent surveying meadows.

First published in Great Britain in 2012 by Papadakis Publisher

An imprint of New Architecture Group Limited Kimber Studio, Winterbourne, Berkshire, RG20 8AN, UK

Introduction

6

Chapter 1

Archaic grassland indicator plants

26

Chapter 2

The management of hay meadows

48

Chapter 3

Hay meadow constants

60

Chapter 4

Meadow vetchling meadows

86

Chapter 5

Lady’s Bedstraw meadows

112

Chapter 6

Heath grass meadows

132

Chapter 7

Orchids & other rarer meadow flowers

154

Chapter 8

Meadow flowers of wet places

174

Chapter 9

The conservation of wildflower meadows

202

Index

222

A CIP catalogue of this book is available from the British Library.

Bibliography

224

Printed and bound in China

Acknowledgments

224

Tel. +44 (0) 1635 248833 info@papadakis.net www.papadakis.net Publishing Director: Alexandra Papadakis Design Director: Aldo Sampieri Editor: Sheila de Vallée Editorial Assistant: Juliana Kassianos

Front cover: A Dorset meadow in May Back cover: Pyramidal orchid flowering in a lady’s bedstraw meadow Frontis: Ragged robin flowering with meadow buttercups in a wet meadow Title page: Green-winged orchids

All images are © John Pilkington with the exception of the images listed below and those in the public domain.

Page 217 © copyright The Board of Trustees of the Royal Botanic Gardens, Kew; page 219 courtesy John Prodger

We gratefully acknowledge the permission granted to use these images. Every possible attempt has been made to identify and contact copyright holders. Any errors or omissions are inadvertent and will be corrected in subsequent editions.

ISBN 978 1 906506 26 1 Copyright © 2012 Margaret Pilkington and Papadakis Publisher All rights reserved.

Margaret Pilkington hereby asserts her moral right to be identified as the author of this work.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording or any other information storage and retrieval system, without prior permission in writing from the Publisher.

Wildflower_00_001-005_Prelims_AS_2.inx 4-5

9/2/12 16:26:26

contents

To all the volunteers who have come from different walks of life to learn botanical identification skills and have shared their expertise during many happy hours spent surveying meadows.

First published in Great Britain in 2012 by Papadakis Publisher

An imprint of New Architecture Group Limited Kimber Studio, Winterbourne, Berkshire, RG20 8AN, UK

Introduction

6

Chapter 1

Archaic grassland indicator plants

26

Chapter 2

The management of hay meadows

48

Chapter 3

Hay meadow constants

60

Chapter 4

Meadow vetchling meadows

86

Chapter 5

Lady’s Bedstraw meadows

112

Chapter 6

Heath grass meadows

132

Chapter 7

Orchids & other rarer meadow flowers

154

Chapter 8

Meadow flowers of wet places

174

Chapter 9

The conservation of wildflower meadows

202

Index

222

A CIP catalogue of this book is available from the British Library.

Bibliography

224

Printed and bound in China

Acknowledgments

224

Tel. +44 (0) 1635 248833 info@papadakis.net www.papadakis.net Publishing Director: Alexandra Papadakis Design Director: Aldo Sampieri Editor: Sheila de Vallée Editorial Assistant: Juliana Kassianos

Front cover: A Dorset meadow in May Back cover: Pyramidal orchid flowering in a lady’s bedstraw meadow Frontis: Ragged robin flowering with meadow buttercups in a wet meadow Title page: Green-winged orchids

All images are © John Pilkington with the exception of the images listed below and those in the public domain.

Page 217 © copyright The Board of Trustees of the Royal Botanic Gardens, Kew; page 219 courtesy John Prodger

We gratefully acknowledge the permission granted to use these images. Every possible attempt has been made to identify and contact copyright holders. Any errors or omissions are inadvertent and will be corrected in subsequent editions.

ISBN 978 1 906506 26 1 Copyright © 2012 Margaret Pilkington and Papadakis Publisher All rights reserved.

Margaret Pilkington hereby asserts her moral right to be identified as the author of this work.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording or any other information storage and retrieval system, without prior permission in writing from the Publisher.

Wildflower_00_001-005_Prelims_AS_2.inx 4-5

9/2/12 16:26:26

Whether they are hay meadows or grazed meadows, the meadows in this book are what we call ‘permanent’ grassland to distinguish them from short-term sown grassland (known as a ‘ley’) which is ploughed up and re-sown every few years. Many of them will not have been ploughed within living memory and some, like ancient woodland, may never have had their soil disturbed. There is a suite of plants associated with old or archaic grassland and these are featured in the next chapter. kaleidoscopic meadows

Meadows look very different at different times of year as the various grasses and wildflowers come into bud, flower and fruit. Over winter, wildflower meadows look rather like any other grassy area giving an overall impression that grassland

12

Wildflower Meadows

Wildflower_00_006-025_Introduction.inx 12-13

habitats occur in abundance. This is one reason why even those who care passionately about the conservation of the countryside are often unaware of biodiversity loss in grassland habitats. Many meadow plants disappear below ground in winter and the indigenous grass species are difficult to identify without their flower spikes.

As spring progresses, the flowers of lady’s smock (Cardamine pratensis) and field woodrush (Luzula campestris) appear, and meadows begin to reveal their true colours. This is the best time of year to look for the curious adderstongue fern (Ophioglossum vulgatum) before the tiny green leaves and the spore-producing ‘tongue’ are hidden in the lengthening grass sward.

The bright yellow flowers of the much maligned dandelion provide a welcome early nectar source

above left: Catkin of male pussy willow flowers with bright yellow anthers on the end of long white stalks. At the base of each flower there is a dark-looking nectary – an essential food source for insects emerging from hibernation at the end of winter. The anthers contain pollen which is collected by colony-making queen bumblebees.

Only queen bees survive through winter and each queen has to collect sufficient pollen to rear the first cohort of worker bees in her newly established colony. above right: Catkin of flowers on a female pussy willow tree. Each flower is topped with a yellow stigma and has a prominent dark-looking nectary at its base. This provides a much-needed source of nectar for female bumblebees emerging from hibernation at the end of winter. Bumblebees are active at temperatures as low as five degrees centigrade, so are flying in March when not much is flowering in the countryside. left: Lady’s smock, one of the first wildflowers to appear in meadows. opposite: An attractive landscape in the High Weald of Sussex in April, but the bright green field surrounded by ploughed land is a sown grass ley like the field in the foreground which lacks wildflowers except for dandelions and white clover.

Introduction 13

10/2/12 09:27:03

Whether they are hay meadows or grazed meadows, the meadows in this book are what we call ‘permanent’ grassland to distinguish them from short-term sown grassland (known as a ‘ley’) which is ploughed up and re-sown every few years. Many of them will not have been ploughed within living memory and some, like ancient woodland, may never have had their soil disturbed. There is a suite of plants associated with old or archaic grassland and these are featured in the next chapter. kaleidoscopic meadows

Meadows look very different at different times of year as the various grasses and wildflowers come into bud, flower and fruit. Over winter, wildflower meadows look rather like any other grassy area giving an overall impression that grassland

12

Wildflower Meadows

Wildflower_00_006-025_Introduction.inx 12-13

habitats occur in abundance. This is one reason why even those who care passionately about the conservation of the countryside are often unaware of biodiversity loss in grassland habitats. Many meadow plants disappear below ground in winter and the indigenous grass species are difficult to identify without their flower spikes.

As spring progresses, the flowers of lady’s smock (Cardamine pratensis) and field woodrush (Luzula campestris) appear, and meadows begin to reveal their true colours. This is the best time of year to look for the curious adderstongue fern (Ophioglossum vulgatum) before the tiny green leaves and the spore-producing ‘tongue’ are hidden in the lengthening grass sward.

The bright yellow flowers of the much maligned dandelion provide a welcome early nectar source

above left: Catkin of male pussy willow flowers with bright yellow anthers on the end of long white stalks. At the base of each flower there is a dark-looking nectary – an essential food source for insects emerging from hibernation at the end of winter. The anthers contain pollen which is collected by colony-making queen bumblebees.

Only queen bees survive through winter and each queen has to collect sufficient pollen to rear the first cohort of worker bees in her newly established colony. above right: Catkin of flowers on a female pussy willow tree. Each flower is topped with a yellow stigma and has a prominent dark-looking nectary at its base. This provides a much-needed source of nectar for female bumblebees emerging from hibernation at the end of winter. Bumblebees are active at temperatures as low as five degrees centigrade, so are flying in March when not much is flowering in the countryside. left: Lady’s smock, one of the first wildflowers to appear in meadows. opposite: An attractive landscape in the High Weald of Sussex in April, but the bright green field surrounded by ploughed land is a sown grass ley like the field in the foreground which lacks wildflowers except for dandelions and white clover.

Introduction 13

10/2/12 09:27:03

bumblebees. In the final chapter, we look at what is being done to conserve wildflower meadows.

receptable (ovary within)

Many wildflower plants are important nectar sources for adult butterflies; others provide nectar and pollen for bees and other beneficial insects; some show intimate connections between the flower structure and their insect pollinators; still others are eaten by the caterpillar stage of meadow butterflies or moths. The photographs in the following pages, illustrating the different wildflowers found in each type of meadow, have been chosen to reveal the connections between the wildflowers and a range of meadow insects.

sepal, one of five anther filament petal, one of five stigma

A diagram of the flower of agrimony to illustrate the flower parts of a typical wildflower. There are two whorls of petal-like structures: the outermost, which are green in agrimony, are sepals and inside the sepals are the showy petals which are yellow in agrimony. Then there are one or more whorls of stamens, the male parts of the flower. Each stamen consists of a pollen-bearing anther on the end of a stalk called a filament. There are 10 to 20 stamens in agrimony flowers; not all are shown in this diagram.

And finally in the centre of the flower are the female flower parts which consist of a stigma which receives the pollen, a stalk of appropriate length to place the stigmatic surface where it will encounter pollen and an ovary which contains the ovules destined to become seeds after fertilisation by pollen. In the agrimony flower there are two stigmas. The female parts of the flower – stigma, style and ovary – are known collectively as carpels. opposite above: Birdsfoot trefoil providing nectar for a bumblebee in a mid June meadow.

style opposite below: Dandelions in a sown grass ley providing an early source of nectar for queen bumblebees emerging from hibernation.

understanding how flowers are pollinated

In order to understand the intimate connection between meadow flowers and pollinators such as bees and hoverflies, we need to be able to talk about the parts of a flower. For many of the wildflowers in this book, it is an incredible story and an appreciation of how pollination is effected can add immeasurably to our enjoyment of the flowers. So let us start with the parts of a fairly simple flower in the rose family and then we can elaborate on this as we deal with the many variations on this theme found in flowers in subsequent chapters.

In insect pollinated flowers, the pollen is transferred from the anthers of one specimen of a particular species of wildflower to the stigma of another specimen of the same species by insects which are attracted to the flowers by a nectar or pollen reward. The pollen grains are rough and sticky so that they stick to the insect’s body and then are transferred onto the receptive surface of the stigma as the insect brushes against it. In order

20

Wildflower Meadows

Wildflower_00_006-025_Introduction.inx 20-21

to ensure that the pollen which becomes attached to the stigma is from a different specimen (so called ‘cross pollination’) the stigmatic surface is either not receptive to pollen from the same flower (selfincompatible) or matures at a different time to the anthers. Some flowers hedge their bets by also having a period in which self pollination can take place if cross pollination has failed to happen. surviving and reproducing in the meadow. how meadow plants maintain their presence in the meadow.

Most meadow plants are perennial in their life habit. This means that an individual plant lives for more than one year in contrast to an annual plant which grows from a seed, produces flowers and seeds and then dies over the space of one year. A grassy sward makes it difficult for an annual plant to find an open space on the surface of the soil to germinate and grow into a tiny seedling, but never-the-less there are a few annuals which are characteristic of meadows. Mole hills and worm casts provide opportunities, and aftermath grazing

Introduction 21

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bumblebees. In the final chapter, we look at what is being done to conserve wildflower meadows.

receptable (ovary within)

Many wildflower plants are important nectar sources for adult butterflies; others provide nectar and pollen for bees and other beneficial insects; some show intimate connections between the flower structure and their insect pollinators; still others are eaten by the caterpillar stage of meadow butterflies or moths. The photographs in the following pages, illustrating the different wildflowers found in each type of meadow, have been chosen to reveal the connections between the wildflowers and a range of meadow insects.

sepal, one of five anther filament petal, one of five stigma

A diagram of the flower of agrimony to illustrate the flower parts of a typical wildflower. There are two whorls of petal-like structures: the outermost, which are green in agrimony, are sepals and inside the sepals are the showy petals which are yellow in agrimony. Then there are one or more whorls of stamens, the male parts of the flower. Each stamen consists of a pollen-bearing anther on the end of a stalk called a filament. There are 10 to 20 stamens in agrimony flowers; not all are shown in this diagram.

And finally in the centre of the flower are the female flower parts which consist of a stigma which receives the pollen, a stalk of appropriate length to place the stigmatic surface where it will encounter pollen and an ovary which contains the ovules destined to become seeds after fertilisation by pollen. In the agrimony flower there are two stigmas. The female parts of the flower – stigma, style and ovary – are known collectively as carpels. opposite above: Birdsfoot trefoil providing nectar for a bumblebee in a mid June meadow.

style opposite below: Dandelions in a sown grass ley providing an early source of nectar for queen bumblebees emerging from hibernation.

understanding how flowers are pollinated

In order to understand the intimate connection between meadow flowers and pollinators such as bees and hoverflies, we need to be able to talk about the parts of a flower. For many of the wildflowers in this book, it is an incredible story and an appreciation of how pollination is effected can add immeasurably to our enjoyment of the flowers. So let us start with the parts of a fairly simple flower in the rose family and then we can elaborate on this as we deal with the many variations on this theme found in flowers in subsequent chapters.

In insect pollinated flowers, the pollen is transferred from the anthers of one specimen of a particular species of wildflower to the stigma of another specimen of the same species by insects which are attracted to the flowers by a nectar or pollen reward. The pollen grains are rough and sticky so that they stick to the insect’s body and then are transferred onto the receptive surface of the stigma as the insect brushes against it. In order

20

Wildflower Meadows

Wildflower_00_006-025_Introduction.inx 20-21

to ensure that the pollen which becomes attached to the stigma is from a different specimen (so called ‘cross pollination’) the stigmatic surface is either not receptive to pollen from the same flower (selfincompatible) or matures at a different time to the anthers. Some flowers hedge their bets by also having a period in which self pollination can take place if cross pollination has failed to happen. surviving and reproducing in the meadow. how meadow plants maintain their presence in the meadow.

Most meadow plants are perennial in their life habit. This means that an individual plant lives for more than one year in contrast to an annual plant which grows from a seed, produces flowers and seeds and then dies over the space of one year. A grassy sward makes it difficult for an annual plant to find an open space on the surface of the soil to germinate and grow into a tiny seedling, but never-the-less there are a few annuals which are characteristic of meadows. Mole hills and worm casts provide opportunities, and aftermath grazing

Introduction 21

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archaic grassland indicator plants

W

ildflower meadows in lowland Britain have their origins in hay production. It is difficult for us to appreciate today, with our oil driven economy, how important hay was in the days of horse-drawn transport, and when ploughing was done by oxen. A large number of animals had to be maintained over winter when there was no fresh grass in the fields. It was crucial to have a good crop of hay and at the end of the 19th century an average of 4.5 million tons of hay were produced each year. Up until the middle of the last century the best land was given over to hay production, and hay meadows with their profusion of wildflowers were a familiar sight. Now they are vanishing at an unprecedented rate, their demise locked away in the memories of a passing generation. Here is a farmer in Sussex recalling a riverside hay meadow called Iron Gates in the late 1940s: “The Iron Gates, in my day we always made hay… Because the grass there was a very fine grass and made beautiful soft hay, very suitable for feeding to calves.”

The meadow had been full of cowslips until he applied chemical fertiliser:

A typical landscape of old meadows along the River Hooke in Dorset. Up until the middle of the last century, hay meadows such as this were common alongside rivers and streams providing a connected landscape for meadow plants and pollinating insects.

26

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Wildflower_01_026-047_Archaic grassland indicator plants.inx 26-27

“When I came here, the Iron Gates was full of cowslips and as soon as I applied some fertiliser, the cowslips disappeared completely and have never come back.”

Archaic Grassland Indicator Plants 27

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archaic grassland indicator plants

W

ildflower meadows in lowland Britain have their origins in hay production. It is difficult for us to appreciate today, with our oil driven economy, how important hay was in the days of horse-drawn transport, and when ploughing was done by oxen. A large number of animals had to be maintained over winter when there was no fresh grass in the fields. It was crucial to have a good crop of hay and at the end of the 19th century an average of 4.5 million tons of hay were produced each year. Up until the middle of the last century the best land was given over to hay production, and hay meadows with their profusion of wildflowers were a familiar sight. Now they are vanishing at an unprecedented rate, their demise locked away in the memories of a passing generation. Here is a farmer in Sussex recalling a riverside hay meadow called Iron Gates in the late 1940s: “The Iron Gates, in my day we always made hay… Because the grass there was a very fine grass and made beautiful soft hay, very suitable for feeding to calves.”

The meadow had been full of cowslips until he applied chemical fertiliser:

A typical landscape of old meadows along the River Hooke in Dorset. Up until the middle of the last century, hay meadows such as this were common alongside rivers and streams providing a connected landscape for meadow plants and pollinating insects.

26

Wildflower Meadows

Wildflower_01_026-047_Archaic grassland indicator plants.inx 26-27

“When I came here, the Iron Gates was full of cowslips and as soon as I applied some fertiliser, the cowslips disappeared completely and have never come back.”

Archaic Grassland Indicator Plants 27

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catsear

Hypochoeris radicata

left: The splashes of yellow amongst the oxeye daisies in this June meadow are flowering heads of catsear. The ‘flower’ consists of a head of long thin strap-like florets all of which contain nectar and pollen: a bountiful supply of food for the many kinds of insects which visit the flower-head.

One or more branched flower stalks each with several flower-heads grow out from the centre of conspicuous rosettes of hairy pimply leaves. above: Like knapweed the ‘flower’ of catsear is really a collection of flowers or florets each with male and female flower parts, but this time the florets are asymmetrical and strap-like.

The anthers are united into a long thin tube surrounding the style and pollen is shed into this tube before it opens. Unopened anther tubes are visible in the centre of the flower. Pollen is pushed out of the top of the anther tube in response to the stimulation of a visiting insect.

When all the pollen has gone, the style emerges and the two arms of the stigma separate to expose the upper surface for the collection of pollen which may be from another plant.

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Wildflower_01_026-047_Archaic grassland indicator plants.inx 42-43

Archaic Grassland Indicator Plants 43

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catsear

Hypochoeris radicata

left: The splashes of yellow amongst the oxeye daisies in this June meadow are flowering heads of catsear. The ‘flower’ consists of a head of long thin strap-like florets all of which contain nectar and pollen: a bountiful supply of food for the many kinds of insects which visit the flower-head.

One or more branched flower stalks each with several flower-heads grow out from the centre of conspicuous rosettes of hairy pimply leaves. above: Like knapweed the ‘flower’ of catsear is really a collection of flowers or florets each with male and female flower parts, but this time the florets are asymmetrical and strap-like.

The anthers are united into a long thin tube surrounding the style and pollen is shed into this tube before it opens. Unopened anther tubes are visible in the centre of the flower. Pollen is pushed out of the top of the anther tube in response to the stimulation of a visiting insect.

When all the pollen has gone, the style emerges and the two arms of the stigma separate to expose the upper surface for the collection of pollen which may be from another plant.

42

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Wildflower_01_026-047_Archaic grassland indicator plants.inx 42-43

Archaic Grassland Indicator Plants 43

9/2/12 16:52:21

After cutting the hay has to be turned. Until after the second world war this was largely done by hand or with a horse-drawn roller and chain. Albert remembered helping the nuns from the local priory turn the hay during the war: above: Along with a revival in traditional countryside crafts, demonstrations of the agricultural use of heavy horses are popular.

These heavy horses are a popular attraction as they harrow a meadow at Wakehurst Place in Sussex. A similar set-up would have been used in the past to turn hay with a roller and chain. opposite above: Old horse-drawn mower, in production until the mid-1930s and subsequently converted for use behind a tractor. opposite right: Cutting hay by hand-scythe in a small meadow in Devon, August 2010. The swinging action of the scythe is laying the hay in regular lines or windrows with a cut strip of meadow in between.

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Wildflower Meadows

Wildflower_02_048-059_The management of hay meadows.inx 52-53

“There’s a field up here on the left hand side. Well the nuns would be out there in the summer turning the hay. I used to go from the farm here, with a horse and a roller and a chain and do it for them.”

Horse-drawn mowers superseded the hand-scythe and by the late 1940s, tractors had largely replaced the horses. These tractors, though, were quite unlike today’s models.

“The difference in the power of tractors … most of the time I was here, all the work was done with those little grey Ferguson tractors, but today they’re just a museum piece. They cost four hundred pounds new.

Today a new tractor would cost what, thirty five thousand? “ (97 year-old farmer interviewed in 2008)

Extra labour was brought in to make hay while the fine weather lasted. Here is Derek, interviewed in 2007 recalling his childhood in the 1950s.

“Our hay making was all done by hand … there was me grandfather, me father and me grandfather’s brother-in-law that used to work on the farm, but then when it came to haying in the summertime, we had three other people that came in for a bit of casual work and they worked down the local timber yard. So they would come in sort of about half past five and work through to about ten o’clock. Last orders at the Sheffield Arms was always about half past ten so it was always the last half an hour that they’d go down and have a drink. And grandfather would supply them with sort of supper.”

The Management of Hay Meadows 53

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After cutting the hay has to be turned. Until after the second world war this was largely done by hand or with a horse-drawn roller and chain. Albert remembered helping the nuns from the local priory turn the hay during the war: above: Along with a revival in traditional countryside crafts, demonstrations of the agricultural use of heavy horses are popular.

These heavy horses are a popular attraction as they harrow a meadow at Wakehurst Place in Sussex. A similar set-up would have been used in the past to turn hay with a roller and chain. opposite above: Old horse-drawn mower, in production until the mid-1930s and subsequently converted for use behind a tractor. opposite right: Cutting hay by hand-scythe in a small meadow in Devon, August 2010. The swinging action of the scythe is laying the hay in regular lines or windrows with a cut strip of meadow in between.

52

Wildflower Meadows

Wildflower_02_048-059_The management of hay meadows.inx 52-53

“There’s a field up here on the left hand side. Well the nuns would be out there in the summer turning the hay. I used to go from the farm here, with a horse and a roller and a chain and do it for them.”

Horse-drawn mowers superseded the hand-scythe and by the late 1940s, tractors had largely replaced the horses. These tractors, though, were quite unlike today’s models.

“The difference in the power of tractors … most of the time I was here, all the work was done with those little grey Ferguson tractors, but today they’re just a museum piece. They cost four hundred pounds new.

Today a new tractor would cost what, thirty five thousand? “ (97 year-old farmer interviewed in 2008)

Extra labour was brought in to make hay while the fine weather lasted. Here is Derek, interviewed in 2007 recalling his childhood in the 1950s.

“Our hay making was all done by hand … there was me grandfather, me father and me grandfather’s brother-in-law that used to work on the farm, but then when it came to haying in the summertime, we had three other people that came in for a bit of casual work and they worked down the local timber yard. So they would come in sort of about half past five and work through to about ten o’clock. Last orders at the Sheffield Arms was always about half past ten so it was always the last half an hour that they’d go down and have a drink. And grandfather would supply them with sort of supper.”

The Management of Hay Meadows 53

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left: Great burnet flowering with oxeye daisies in Askrigg Bottom, a northern hay meadow (MG3) in Wensleydale, North Yorkshire. This upland type of hay meadow is almost completely restricted to a few valleys in northern England in Durham, North Yorkshire and the Lake District. above: Snakeshead fritillaries flowering in the famous North Meadow at Cricklade. This type of lowland flood-plain hay meadow (MG4) is only found in a few scattered sites in the Midlands, and in southern Britain where seasonally-flooded land has been managed for centuries in the same traditional way. North Meadow is still cut for hay and then grazed in the autumn as common land. overleaf: As the early flowers disappear, a mass of flowering common knapweed brightens up this Devon meadow in late July. A rich source of nectar, common knapweed occurs in great abundance in some meadows.

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Wildflower_03_060-085_Hay meadow constants.inx 64-65

Hay Meadow Constants 65

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left: Great burnet flowering with oxeye daisies in Askrigg Bottom, a northern hay meadow (MG3) in Wensleydale, North Yorkshire. This upland type of hay meadow is almost completely restricted to a few valleys in northern England in Durham, North Yorkshire and the Lake District. above: Snakeshead fritillaries flowering in the famous North Meadow at Cricklade. This type of lowland flood-plain hay meadow (MG4) is only found in a few scattered sites in the Midlands, and in southern Britain where seasonally-flooded land has been managed for centuries in the same traditional way. North Meadow is still cut for hay and then grazed in the autumn as common land. overleaf: As the early flowers disappear, a mass of flowering common knapweed brightens up this Devon meadow in late July. A rich source of nectar, common knapweed occurs in great abundance in some meadows.

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Hay Meadow Constants 65

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common daisy

Bellis Perennis oxeye daisy

Leucanthemum vulgare

top left: Flower-head of oxeye daisy with visiting bee getting amply dusted with the copious pollen while probing for nectar. The ‘flower’ of both common daisy and oxeye daisy is really a head of tiny flowers. In the centre are symmetrical tubular flowers each composed of five yellow petals with stamens and stigmas inside. The head of tubular flowers is made more conspicuous for visiting insects by the encircling ring of white ray florets. These are asymmetrical and strap-like and do not produce any pollen. The yellow florets of oxeye are similar to the tubular knapweed florets described in chapter 1. When the flowers open, the stigmas protruding from the central florets are already dusted with pollen and the stamens have withered. A huge range of insects – bees, beetles, flies, butterflies and moths – visit oxeye daisy flowers for nectar secreted at the base of the petal tube and become covered in pollen. After several days the style opens out to expose three receptive stigmatic surfaces to capture pollen from another flower, but the receptive surfaces will also accept pollen from the same plant so self pollination is frequent. A large quantity of seed is produced which can germinate in the autumn or the spring and can remain viable in the soil for several years. This enables the plant to act like an arable weed and exploit areas where the soil has been disturbed such as roadside verges and heaps of spoil, but makes it less successful in a meadow context. It may be that the disturbance of the sward by aftermath grazing is essential to maintain the presence of oxeye daisy in meadows. I have noticed a decline in meadows managed by hay cutting without grazing.

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top right: Common daisy flowering with meadow buttercup in a May meadow. The ordinary common daisy, a familiar plant of garden lawns and playing fields, and the much larger oxeye daisy are both diagnostic species for MG5a. Common daisy comes into flower really early in the year, sometimes at the beginning of March, and where the grass is cut or grazed at regular intervals it goes on flowering into autumn. Oxeye daisy on the other hand has a much more restricted flowering period and is particularly associated with June meadows. above: Oxeye daisy flowering with common spotted orchids, red clover and birdsfoot trefoil in a June meadow.

opposite: Oxeye daisy is often sown into roadside verges, roundabouts and garden meadows.

When sown it may be almost too successful producing a boring uniform sea of white flowers.

Here, it has formed small patches of white in a natural meadow, adding contrast to the tapestry of colours amongst the grass. The basal leaves of this perennial plant can be found close to the ground all through winter, but as the flowering stem elongates and the vegetation of the meadow becomes taller, photosynthesis is taken over by leaves on the stem. The short rhizomes lead to little vegetative spread, and although large quantities of seed are produced oxeye daisy seems to be decreasing in meadows.

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common daisy

Bellis Perennis oxeye daisy

Leucanthemum vulgare

top left: Flower-head of oxeye daisy with visiting bee getting amply dusted with the copious pollen while probing for nectar. The ‘flower’ of both common daisy and oxeye daisy is really a head of tiny flowers. In the centre are symmetrical tubular flowers each composed of five yellow petals with stamens and stigmas inside. The head of tubular flowers is made more conspicuous for visiting insects by the encircling ring of white ray florets. These are asymmetrical and strap-like and do not produce any pollen. The yellow florets of oxeye are similar to the tubular knapweed florets described in chapter 1. When the flowers open, the stigmas protruding from the central florets are already dusted with pollen and the stamens have withered. A huge range of insects – bees, beetles, flies, butterflies and moths – visit oxeye daisy flowers for nectar secreted at the base of the petal tube and become covered in pollen. After several days the style opens out to expose three receptive stigmatic surfaces to capture pollen from another flower, but the receptive surfaces will also accept pollen from the same plant so self pollination is frequent. A large quantity of seed is produced which can germinate in the autumn or the spring and can remain viable in the soil for several years. This enables the plant to act like an arable weed and exploit areas where the soil has been disturbed such as roadside verges and heaps of spoil, but makes it less successful in a meadow context. It may be that the disturbance of the sward by aftermath grazing is essential to maintain the presence of oxeye daisy in meadows. I have noticed a decline in meadows managed by hay cutting without grazing.

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top right: Common daisy flowering with meadow buttercup in a May meadow. The ordinary common daisy, a familiar plant of garden lawns and playing fields, and the much larger oxeye daisy are both diagnostic species for MG5a. Common daisy comes into flower really early in the year, sometimes at the beginning of March, and where the grass is cut or grazed at regular intervals it goes on flowering into autumn. Oxeye daisy on the other hand has a much more restricted flowering period and is particularly associated with June meadows. above: Oxeye daisy flowering with common spotted orchids, red clover and birdsfoot trefoil in a June meadow.

opposite: Oxeye daisy is often sown into roadside verges, roundabouts and garden meadows.

When sown it may be almost too successful producing a boring uniform sea of white flowers.

Here, it has formed small patches of white in a natural meadow, adding contrast to the tapestry of colours amongst the grass. The basal leaves of this perennial plant can be found close to the ground all through winter, but as the flowering stem elongates and the vegetation of the meadow becomes taller, photosynthesis is taken over by leaves on the stem. The short rhizomes lead to little vegetative spread, and although large quantities of seed are produced oxeye daisy seems to be decreasing in meadows.

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Lady’s Bedstraw meadows

I

n this chapter, we look at the plants which occur most frequently in the second sub-community of crested dogstail – common knapweed grassland; the lady’s bedstraw sub-community (MG5b).

Most of these preferential species also occur in MG5a meadows, but at a much lower frequency. Four of them also occur in MG5c meadows. Sheep’s fescue grass is the only one which is only present in MG5b meadows.

As in the previous chapter, we will be trying to build up a picture of these meadows and so will include the most frequently occurring associate species, as well as the diagnostic or preferential species which can be used to identify MG5b grassland. The most characteristic wildflower is lady’s bedstraw (Galium verum) which occurs with yarrow (Achillea millefolium), ragwort (Senecio jacobaea), agrimony (Agrimonia eupatoria), salad burnet (Sanguisorba minor) and hoary plantain (Plantago media). The most characteristic grass is yellow oatgrass (Trisetum flavescens) which occurs with quaking grass (Briza media), creeping bent (Agrostis stolonifera), crested hairgrass (Koeleria macrantha), and sheep’s fescue (Festuca ovina).

There is one sedge (glaucous sedge) which is preferential and one moss (rough-stalked feather moss) which is particularly associated with these meadows.

Lady’s bedstraw meadows become golden in July with masses of tiny yellow bedstraw flowers which are sweetly scented. The aroma fills the air.

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Many of these plants, such as lady’s bedstraw, salad burnet and quaking grass, also occur in chalk grassland and so are associated in our minds with a more alkaline soil. However, plants growing in chalk grassland may be as much dependent on the lack of nutrients which prevent fast growing competitor plants taking over as on the pH of the soil. This is true for wildflower meadows

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Lady’s Bedstraw meadows

I

n this chapter, we look at the plants which occur most frequently in the second sub-community of crested dogstail – common knapweed grassland; the lady’s bedstraw sub-community (MG5b).

Most of these preferential species also occur in MG5a meadows, but at a much lower frequency. Four of them also occur in MG5c meadows. Sheep’s fescue grass is the only one which is only present in MG5b meadows.

As in the previous chapter, we will be trying to build up a picture of these meadows and so will include the most frequently occurring associate species, as well as the diagnostic or preferential species which can be used to identify MG5b grassland. The most characteristic wildflower is lady’s bedstraw (Galium verum) which occurs with yarrow (Achillea millefolium), ragwort (Senecio jacobaea), agrimony (Agrimonia eupatoria), salad burnet (Sanguisorba minor) and hoary plantain (Plantago media). The most characteristic grass is yellow oatgrass (Trisetum flavescens) which occurs with quaking grass (Briza media), creeping bent (Agrostis stolonifera), crested hairgrass (Koeleria macrantha), and sheep’s fescue (Festuca ovina).

There is one sedge (glaucous sedge) which is preferential and one moss (rough-stalked feather moss) which is particularly associated with these meadows.

Lady’s bedstraw meadows become golden in July with masses of tiny yellow bedstraw flowers which are sweetly scented. The aroma fills the air.

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Many of these plants, such as lady’s bedstraw, salad burnet and quaking grass, also occur in chalk grassland and so are associated in our minds with a more alkaline soil. However, plants growing in chalk grassland may be as much dependent on the lack of nutrients which prevent fast growing competitor plants taking over as on the pH of the soil. This is true for wildflower meadows

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greater butterfly orchid

bitter vetch

Platanthera chlorantha

Lathyrus montanus

right: Greater butterfly orchid growing with common spotted orchids, common knapweed and rough hawkbit in a wildflower meadow at Pentwyn Farm in Gwent. Although greater butterfly orchid was probably common in hay meadows throughout the country in the past, its occurrence in hay meadows and pastures is limited now to the north and west of Britain. It still lingers in scrub and the edge of woodland on the chalk in the south-east, but seems unable to move from here out into meadows. below: The greenish-white flowers of greater butterfly orchid almost glow in the dusk and the heady scent from the flowers is stronger at night attracting large night-flying moths such as noctuids. There is a strap-like lip on which the moth alights and a long curved spur filled with nectar into which the tongue of the moth can fit. The other two petals form a hood above the opening to the spur. Two prominent viscidia can be seen lying on either side of, and just above,

the opening to the spur. The positioning of the viscidia is such that a moth pushing its tongue into the mouth of the spur to suck up nectar ends up with a viscidium stuck fast to one of its large compound eyes, As the sticky substance hardens, the orientation of the pollinium changes so that it points inwards and is in the right position to deposit pollen on the centrally placed stigma in the next flower visited. Although the pollen masses are loosely attached and come off readily on the stigma, the viscidium is likely to remain attached to the hapless insect for the rest of its life. The pollination mechanism is very efficient and up to 90% of capsules produce seed.

The minute seeds are easily dispersed on the wind, but will only germinate if the right fungus is present in the soil. Even then, it will be eight years from when the seed germinates until it produces the first flowering spike. Fortunately, butterfly orchids like spotted and marsh orchids produce new tubers from a bud at the base of the flowering stem and in good years more than one tuber may be produced from each flowering stem.

right: The bright crimson flowers of bitter vetch soon turn blue. They contain nectar and are pollinated by bees. Although very similar to vetches (Vicia), bitter vetch has a winged stem and fewer leaflets in the leaves, so is really a vetchling (Lathyrus) like meadow vetchling. The end of the leaf terminates in a small point rather than a tendril, so the plant does not have the same ability to scramble up through tall meadow vegetation as meadow vetchling. It tends to have a northern and western distribution, but turns up fairly frequently in Sussex in Wealden meadows on sandy soils where it flowers early before the meadow vegetation becomes tall. A perennial plant spreading with long rhizomes, it is not dependent on producing lots of seed. In Sussex and Lincolnshire it is an ancient woodland indicator species which reflects its dependence on undisturbed soil and an inability to colonise new sites.

grass vetchling

Lathyrus nissolia right and below: This amazing plant has one or two brilliant crimson pea-flowers which make it suddenly visible amongst the tall meadow vegetation of mid summer. Before it comes into flower it is the most frustrating plant to find. The leaves are long and thin, looking very similar to the surrounding grass and since the plant is an annual it occurs in different places in the meadow each year. The flowers are short-lived being replaced by long green pods containing 15 to 20 seeds and once again the plant is all but invisible.

It is not easy being an annual competing with the perennial life habit of the vast majority of meadow plants, so it is perhaps not surprising that this is an uncommon flower. Like orchids, it has a special fascination because it appears unpredictably and the flower is stunning. For the specialist there is the added fascination that the long thin leaves are technically not leaves at all, but phyllodes: leaf stalks which have become flattened and have taken over the function of photosynthesis.

above: Butterfly orchid has two large shiny green leaves at the base of the plant, one slightly above the other, from which the tall flowering stem arises.

There are a few narrow leaves on the stem and a delicate head of creamy white flowers with long strap-like lips and even longer thread-like spurs.

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greater butterfly orchid

bitter vetch

Platanthera chlorantha

Lathyrus montanus

right: Greater butterfly orchid growing with common spotted orchids, common knapweed and rough hawkbit in a wildflower meadow at Pentwyn Farm in Gwent. Although greater butterfly orchid was probably common in hay meadows throughout the country in the past, its occurrence in hay meadows and pastures is limited now to the north and west of Britain. It still lingers in scrub and the edge of woodland on the chalk in the south-east, but seems unable to move from here out into meadows. below: The greenish-white flowers of greater butterfly orchid almost glow in the dusk and the heady scent from the flowers is stronger at night attracting large night-flying moths such as noctuids. There is a strap-like lip on which the moth alights and a long curved spur filled with nectar into which the tongue of the moth can fit. The other two petals form a hood above the opening to the spur. Two prominent viscidia can be seen lying on either side of, and just above,

the opening to the spur. The positioning of the viscidia is such that a moth pushing its tongue into the mouth of the spur to suck up nectar ends up with a viscidium stuck fast to one of its large compound eyes, As the sticky substance hardens, the orientation of the pollinium changes so that it points inwards and is in the right position to deposit pollen on the centrally placed stigma in the next flower visited. Although the pollen masses are loosely attached and come off readily on the stigma, the viscidium is likely to remain attached to the hapless insect for the rest of its life. The pollination mechanism is very efficient and up to 90% of capsules produce seed.

The minute seeds are easily dispersed on the wind, but will only germinate if the right fungus is present in the soil. Even then, it will be eight years from when the seed germinates until it produces the first flowering spike. Fortunately, butterfly orchids like spotted and marsh orchids produce new tubers from a bud at the base of the flowering stem and in good years more than one tuber may be produced from each flowering stem.

right: The bright crimson flowers of bitter vetch soon turn blue. They contain nectar and are pollinated by bees. Although very similar to vetches (Vicia), bitter vetch has a winged stem and fewer leaflets in the leaves, so is really a vetchling (Lathyrus) like meadow vetchling. The end of the leaf terminates in a small point rather than a tendril, so the plant does not have the same ability to scramble up through tall meadow vegetation as meadow vetchling. It tends to have a northern and western distribution, but turns up fairly frequently in Sussex in Wealden meadows on sandy soils where it flowers early before the meadow vegetation becomes tall. A perennial plant spreading with long rhizomes, it is not dependent on producing lots of seed. In Sussex and Lincolnshire it is an ancient woodland indicator species which reflects its dependence on undisturbed soil and an inability to colonise new sites.

grass vetchling

Lathyrus nissolia right and below: This amazing plant has one or two brilliant crimson pea-flowers which make it suddenly visible amongst the tall meadow vegetation of mid summer. Before it comes into flower it is the most frustrating plant to find. The leaves are long and thin, looking very similar to the surrounding grass and since the plant is an annual it occurs in different places in the meadow each year. The flowers are short-lived being replaced by long green pods containing 15 to 20 seeds and once again the plant is all but invisible.

It is not easy being an annual competing with the perennial life habit of the vast majority of meadow plants, so it is perhaps not surprising that this is an uncommon flower. Like orchids, it has a special fascination because it appears unpredictably and the flower is stunning. For the specialist there is the added fascination that the long thin leaves are technically not leaves at all, but phyllodes: leaf stalks which have become flattened and have taken over the function of photosynthesis.

above: Butterfly orchid has two large shiny green leaves at the base of the plant, one slightly above the other, from which the tall flowering stem arises.

There are a few narrow leaves on the stem and a delicate head of creamy white flowers with long strap-like lips and even longer thread-like spurs.

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an ideal system for managing a hay meadow (hanging meadow)

What is the very best way of managing a flower-rich meadow? The Royal Botanic Gardens, Kew at Wakehurst Place in Sussex, the home of the Millennium Seed Bank, is showing the way with their management of Hanging Meadow. The meadow is part of the Loder Valley nature reserve which lies adjacent to Wakehurst Place Gardens and is managed by Kew. There was anecdotal evidence of a decline in the number of common spotted orchids in this beautiful flower-rich meadow following several years when the aftermath had not been grazed. So a flock of sheep was purchased in 2008 and aftermath grazing re-introduced. Special light-weight equipment was also purchased for cutting and turning the hay and for making mini hay bales suitable for feeding to the sheep over the winter months. The effect of the ‘improved’ management is being carefully monitored in marked squares to ensure that the wildflowers are responding in the expected way.

Hanging Meadow in 2009 shortly before new management was instigated with the use of lightweight hay-making equipment and aftermath grazing with sheep. Following several years without aftermath grazing, the common spotted orchids were struggling to maintain their presence.

top: Cutting the hay in Hanging Meadow in 2010 using lightweight, pedestrian equipment. centre: Using lightweight, pedestrian equipment to turn the hay in Hanging Meadow. bottom: Using a pedestrian mini-baler in Hanging Meadow.

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an ideal system for managing a hay meadow (hanging meadow)

What is the very best way of managing a flower-rich meadow? The Royal Botanic Gardens, Kew at Wakehurst Place in Sussex, the home of the Millennium Seed Bank, is showing the way with their management of Hanging Meadow. The meadow is part of the Loder Valley nature reserve which lies adjacent to Wakehurst Place Gardens and is managed by Kew. There was anecdotal evidence of a decline in the number of common spotted orchids in this beautiful flower-rich meadow following several years when the aftermath had not been grazed. So a flock of sheep was purchased in 2008 and aftermath grazing re-introduced. Special light-weight equipment was also purchased for cutting and turning the hay and for making mini hay bales suitable for feeding to the sheep over the winter months. The effect of the ‘improved’ management is being carefully monitored in marked squares to ensure that the wildflowers are responding in the expected way.

Hanging Meadow in 2009 shortly before new management was instigated with the use of lightweight hay-making equipment and aftermath grazing with sheep. Following several years without aftermath grazing, the common spotted orchids were struggling to maintain their presence.

top: Cutting the hay in Hanging Meadow in 2010 using lightweight, pedestrian equipment. centre: Using lightweight, pedestrian equipment to turn the hay in Hanging Meadow. bottom: Using a pedestrian mini-baler in Hanging Meadow.

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BIBLIOGRAPHY Atherton I, Bosanquet S ad Lawley M (editors), 2010.

Mosses and Liverworts of Britain and Ireland. British Bryological Society.

Peterken G (editor), 2005. Flower in the Field.

Parish Grasslands Project, Hewelsfield and St Briavels.

Banks C, 2011. Farm Diversification into Energy Production by Anaerobic Digestion. RELU Policy and Practice Note, number 26.

Poux X, 2010. Why Biodiversity should top the CAP Reform Agenda.

La Canada 24, pp.2-4.

Cope T and Gray A, 2009. Grasses of the British Isles.

Botanical Society of the British Isles, London.

Proctor M and Yeo P, 1973. The Pollination of Flowers.

New Naturalist. Collins, London.

Flower C, 2008. Where Have all the Flowers Gone? Papadakis, London.

Quinn T, 2005. Heaven Upon Earth. Swan Hill Press, Shrewsbury.

Gamble D and St. Pierre T (editors), 2010.

Hay Time in the Yorkshire Dales. Yorkshire Dales Millennium Trust.

Rodwell JS, 1992. British Plant Communities. Volume 3 Grassland and Montane Communities. Cambridge University Press, Cambridge.

Grime JP, Hodgson JG and Hunt R, 1990.

The Abridged Comparative Plant Ecology. Chapman and Hall, London.

Hennell T, 1936 (2nd edition). Change in the Farm.

Cambridge University Press, Cambridge.

Hill MO, Preston CD and Roy DB, 2004. PLANTATT – Attributes of British and Irish Plants. Centre for Ecology and Hydrology, Scotland.

Lloyd C, 2004. Meadows. Cassell Illustrated, London.

Mabey R, 1996. Flora Britannica. Sinclair-Stevenson, London.

Marren P, 1998. Fungal Flowers: the Waxcaps and their World.

British Wildlife 9 (3) pp.164-172.

O’Reilly J, 2010. The State of Upland Hay Meadows in the North Pennines.

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Rose F, 1989. Grasses, Sedges, Rushes and Ferns. The Penguin Group, London.

Rose F, 2006. The Wild Flower Key (revised expanded edition).

The Penguin Group, London.

Short B, Watkins C and Martin J (editors), 2007. The Front Line of Freedom: British Farming in the Second World War. British Agricultural History Society, Exeter.

Summerhayes VS, 1951. Wild Orchids of Britain. New Naturalist.

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Thomas J and Lewington R, 1991. The Butterflies of Britain and Ireland.

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Board of Agriculture, London.

Acknowledgments We would like to thank the meadow owners and nature reserve managers who made us so very welcome on many occasions especially James Hitchcock of the Worcestershire Wildlife Trust and Rupert Browning of Mid Sussex District Council.

Also colleagues on the River Ouse Project especially Will Pilfold and Nick Steer for their meticulous land-use research, and the oral historian Andrew Holmes for his skill in seeking out and recording wonderful gems of meadow history. And the many farmers who shared their expertise and knowledge with us in oral history interviews and informally in the field, especially Ben Courage who kindly allowed us to reproduce his family snap-shots of farming in the 1950s and 60s.

We are very grateful to the Board of Trustees of the Royal Botanic Gardens, Kew who converted Ben Courage’s photos into the splendid digital images used in this book and who

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also kindly allowed us to use three of their photographs of haymaking in Hanging Meadow, Wakehurst Place. And to Mike Samson, the archivist at Blundells School, who drew our attention to the map of Long Meadow.

We gratefully acknowledge The Leverhulme Trust who funded the project Integrating History and Ecology to Sustain a Living Landscape and enabled us to put on record recent farming history with a full-time oral historian. And the Royal Society Committee on the Public Understanding of Science (COPUS) who funded the setting up of the Meadow Management Field Trial.

And finally we would like to thank the team at Papadakis especially Alexandra Papadakis and Juliana Kassianos for being a joy to work with, and for creatively supporting our ideas for the book.

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