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Air War Northern Ireland
 Air War Northern Ireland
Britain’s Air Arms and the ‘Bandit
Country’ of South Armagh
Operation BANNER 1969-2007

Steven Taylor
 First published in Great Britain in 2018 by
Pen & Sword Aviation
an imprint of
Pen & Sword Books Ltd
47 Church Street
Barnsley
South Yorkshire
S70 2AS

Copyright © Steven Taylor 2018

ISBN 978 1 52672 154 9

eISBN 978 1 52672 155 6
Mobi ISBN 978 1 52672 156 3

The right of Steven Taylor to be identified as Author of this work has been
asserted by him in accordance with the Copyright, Designs and Patents Act 1988.

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

All rights reserved. No part of this book may be reproduced or transmitted in any
form or by any means, electronic or mechanical including photocopying, recording
or by any information storage and retrieval system, without permission from the
Publisher in writing.

Pen & Sword Books Ltd includes the Imprints of Pen & Sword Aviation, Pen &
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For a complete list of Pen & Sword titles please contact

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E-mail: enquiries@pen-and-sword.co.uk
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 Contents

Acknowledgements
Image Credits
Map
Glossary
Prologue
Introduction
Chapter One Troubled Times
Chapter Two Bandit Country aka ‘The Independent Republic of
South Armagh’
Chapter Three ‘Everything was done by helicopter’
Chapter Four Pot-shots
Chapter Five Priority Target
Chapter Six ‘If it flies, it dies’. The SAM Threat
Chapter Seven Gifts from the Colonel
Chapter Eight Striking Back
Chapter Nine Home-Made Solutions
Chapter Ten Towards Peace
Appendix A Aircraft crashes/forced landings in South Armagh
during Operation BANNER
Appendix B Aircraft types operated by AAC, RN and RAF
during Operation BANNER
End Notes
Bibliography
 Acknowledgements

F
or their assistance with official records and historical
information, I wish to thank the staff at the National Archives,
Kew; the RAF Air Historical Branch; 38 (Irish) Brigade; the
MoD’s Army Secretariat; and the Police Service of Northern Ireland.
For permission to quote extracts from published sources, I would
like to thank The O’Brien Press Ltd (Insider and Enemy of the
Empire); Routledge (British Generals in Blair’s Wars © Jonathan
Bailey, Richard Iron and Hew Strachan, 2013); Flight International
magazine (extract reproduced with permission of Reed Business
Information Limited via PLSclear); HarperCollins Publishers Ltd
(Hellfire © Ed Macy, 2009); Guy Warner (Sycamores Over Ulster);
Headline (Heroes of the Skies © 2012 MAA Publishing Ltd. Extracts
reproduced by permission of Headline Publishing Group); Imperial
War Museum, London (various sound archives); Pan Macmillan
(Northern Ireland: Soldiers Talking © Max Arthur, 1987. Extracts
reproduced with permission of Pan Macmillan via PLSclear); Duncan
Rogers of Helion & Company (A Long Long War and Wasted Years,
Wasted Lives - The British Army in Northern Ireland Vol. 2); Stuart
Leasor (Rats: The Story of a Dog Soldier); and Bloomsbury
Publishing Plc (Provos: The IRA and Sinn Fein © Peter Taylor, 1997).
My thanks are also due to the following for their kind permission
to use the illustrations featured in this book: The International
Auster Club; Eamon Melaugh; Tony Crowley and the Claremont
Colleges Digital Archive – Murals of Northern Ireland; David
Townsend; Donald MacLeod; Museum of Army Flying; the Airborne
Assault Archives, Duxford; www.flyingmarines.com; Victor Patterson;
and BAe Systems.
 Finally, I wish to thank Richard Doherty and the team at Pen &
Sword.
 Image Credits

1. Map of Northern Ireland (© Queen Mary University of London)

2. Bristol F2B (public domain)
3. Auster WE551 (© The International Auster Club)
4. Westland Sioux AH1 over Londonderry, 1972 (© Eamon
Melaugh)
5. GPMG-armed Westland Sioux AH1 (Courtesy of
flyingmarines.com)
6. Bessbrook Mill (© Museum of Army Flying)
7. DHC-2 Beaver AL1 (© BAE Systems)
8. Aerospatiale Gazelle AH1 (Courtesy of flyingmarines.com)
9. IRA volunteers manning Browning M2 (© Victor Patterson)
10. Westland Wessex HC2 XR506 (© Aviation Photo Company)
11. Test-firing of FIM-92 Stinger (© US Department of Defense)
12. Russian DShK 12.7mm heavy machine gun (iStock)
13. Mural of Lynx being shot down (© Tony Crowley)
14. Westland Lynx hovering over South Armagh field (© Museum of
Army Flying)
15. View of ‘Romeo’ watchtower from door gunner’s position (©
Airborne Assault Archives, Duxford)
16. Britten-Norman Islander AL1 (© David Townsend)
17. Boeing Chinook HC2 airlifting section of watchtower (© Donald
MacLeod)
Map
 Glossary

AAC Army Air Corps

A/C Aircraft
AFC Air Force Cross
AFM Air Force Medal
ARF Airborne Reaction Force
ASU Active Service Unit (IRA)
ATO Ammunition Technical Officer
AVRE Armoured Vehicle Royal Engineers
Bn Battalion
Brick Four-man British Army patrol
CASEVAC Casualty evacuation
3 CBAS 3 Commando Brigade Air Squadron
CO Commanding Officer
COP Close Observation Platoon
Dáil Dáil Éireann, the lower house of the Irish legislature
DFC Distinguished Flying Cross
DFM Distinguished Flying Medal
Dicker IRA look-out
DShK Russian-made 12.7mm heavy machine gun
DSO Distinguished Service Order
ECM Electronic Counter Measures
EOD Explosive Ordnance Disposal
Garda Police force of the Republic of Ireland
GOC General Officer Commanding
GPMG General Purpose Machine Gun
HLS Helicopter Landing Site
HMG Heavy Machine Gun
HQNI Headquarters Northern Ireland
IED Improvised Explosive Device
IVCP Illegal Vehicle Check Point
MANPADS Man Portable Air Defence System
MiD Mention in Despatches
NAS Naval Air Squadron
NIO Northern Ireland Office
OC Officer Commanding
OP Observation Post
ORB Operations Record Book
PIRA Provisional Irish Republican Army
PR Photo Reconnaissance
PVCP Permanent Vehicle Check Point
PSNI Police Service of Northern Ireland
RIC Reconnaissance Intelligence Centre
RPG Rocket-Propelled Grenade
RUC Royal Ulster Constabulary
SAM Surface-to-Air Missile
SF Security Forces
SH Support Helicopters
SHDNI Support Helicopter Detachment Northern Ireland
SHFNI Support Helicopter Force Northern Ireland
SLR Self-Loading Rifle
Sunray Radio call sign for British Army unit commander
Taoiseach Irish Prime Minister
TAOR Tactical Area of Responsibility
UDR Ulster Defence Regiment
USL Underslung Load
VCP Vehicle Check Point
Volunteer Member of IRA
WO2 Warrant Officer Class 2
Yellow CardInstructions on rules of engagement issued to all British
soldiers serving in Northern Ireland
 Prologue

Jonesborough, South Armagh 17 February 1978

T
he Aérospatiale Gazelle skimmed over the fields of South
Armagh, flying at a height of just fifty feet. The rolling
countryside flashing below was some of the most scenic in the
British Isles, but the officer sitting in the observer’s seat of the sleek
helicopter, Lieutenant Colonel Ian Corden-Lloyd, wasn’t interested in
taking in the sights. Instead, his mind was firmly focused on
reaching the village of Jonesborough, just a few hundred metres
from the border with the Irish Republic.
The 39-year-old commanding officer of the 2nd Bn Royal Green
Jackets was regarded as a rising star in the British Army. Born in
Durban, South Africa, he was commissioned into the 10th Princess
Mary’s Own Gurkha Rifles before transferring to the Royal Green
Jackets. During his first tour in Northern Ireland in 1971 he earned
the Military Cross and five years later was promoted to lieutenant
colonel and given command of the 2nd Battalion which, in December
1977, was deployed to South Armagh for a two-year resident tour.
South Armagh. Bandit Country. Throughout the long years of ‘the
Troubles’ this was the most dangerous part of Northern Ireland for
the British Army. For the Green Jackets, the tour was already proving
to be an eventful one. Within weeks of their arrival, PIRA had
launched a mortar attack on the joint RUC/Army base in the village
of Forkhill where they were stationed, injuring several soldiers. The
flatbed lorry from which the mortars were fired was soon found by
the soldiers and checked out by an ATO, who declared it free of
booby traps. But, as two RUC officers started up the lorry, intending
to drive it to another police station for forensic examination, the cab
exploded. Missed by the ATO during his examination was a small
explosive device, hidden inside the windscreen washer bottle.
Corden-Lloyd helped pull the injured policemen clear of the
burning wreckage and both went on to make full recoveries.
Fortunately, nobody had been killed in the double attack, but the
incident served as a stark reminder of the cunning and ruthless
nature of the enemy the men of the Royal Green Jackets were up
against in South Armagh.
The archetypal soldier’s soldier, Corden-Lloyd was highly
respected by his men. Andy McNab, the SAS soldier turned
bestselling author, who served under Corden-Lloyd in South Armagh,
called him ‘the best officer I’d ever met’, and he was known to his
men as a commander who led from the front.
So when on the afternoon of 17 February 1978 a radio call came
into the Operations Room at Bessbrook Mill, the Army’s main base in
South Armagh, reporting a major contact between one of his units
and PIRA gunmen near the border village of Jonesborough, Corden-
Lloyd didn’t hesitate. He immediately jumped into a Gazelle with his
battalion adjutant, Captain Schofield, and ordered the pilot, Sergeant
Ives, to fly him to the scene of the firefight.
The contact report was radioed in by a member of the Green
Jackets’ Close Observation Platoon, or COP. These were units made
up of the best men in each infantry battalion, whose job, as the
name suggests, was to gather intelligence by conducting covert
surveillance of terrorist suspects at close quarters, often remaining
in position for days at a time.
The Green Jackets’ COP was dug into concealed positions at the
southern end of Jonesborough, overlooking the Edenappa Road
leading into the staunchly republican village, where intelligence
indicated PIRA were planning to set up an IVCP.
After hours of waiting, it was beginning to look as if the terrorists
were not going to appear. Then, at around 1620 hours, the calm of
the late afternoon was suddenly shattered by the rattle of automatic
gunfire, as one of the COP team’s positions came under attack from
gunmen hidden behind a stone wall a short distance away. Once
again, PIRA’s South Armagh brigade had demonstrated its skill at
uncovering a covert observation post in its midst.
The firefight between the Green Jackets and PIRA was still raging
when Corden-Lloyd’s Gazelle arrived at the scene less than ten
minutes later, accompanied by a Scout helicopter carrying the
Airborne Reaction Force, consisting of a medic and three men of the
2nd Bn Light Infantry.
Unable to pinpoint the firing positions of the terrorists on the first
low pass over the area, Corden-Lloyd ordered Sergeant Ives to make
another pass. As he did so, a stream of 7.62mm tracer rounds
flashed by the Gazelle as a PIRA volunteer armed with a belt-fed
M60 machine gun turned his attention to the helicopter. Ives reacted
instinctively, pulling up and banking hard to the left in a desperate
attempt to evade the incoming fire.
The COP soldiers and those onboard the Scout watched in horror
as the helicopter then appeared to lose power and plummeted
downwards, hitting the ground and briefly rising up into the air again
before crashing back down for a second and final time, its spinning
rotor blades thrashing into the earth as it cartwheeled across a field.
Finally, the helicopter came to rest on its side 100 metres away from
the initial point of impact, a crumpled wreck.
Time seemed to stand still for a few seconds. Then, the four men
of the ARF hover-jumped from the Scout and raced over to the crash
site, under fire from the terrorist gunmen. Moments later two words
crackled over the COP unit’s radio, bringing a stunned silence to the
men: ‘Sunray down!’ Dave Pomfrett, one of the COP soldiers,
remembered the moment well. ‘We looked around for someone to
make sense of it,’ he recalled. ‘However, we all knew what it meant.’
Lieutenant Colonel Ian Corden-Lloyd had just become the most
senior British officer to be killed on active duty in Northern Ireland.
 Introduction

A
mid little fanfare, at midnight on 31 July 2007 Operation
BANNER, the British Army’s codename for its involvement in
Northern Ireland during the period commonly referred to as
the ‘Troubles’, officially came to an end. What began as a limited
deployment, intended to last only a few weeks to separate
nationalist and loyalist mobs after an outbreak of fierce sectarian
rioting in the summer of 1969, became the longest continuous
military campaign in the history of the British Army.
Initially deployed in a strictly peacekeeping capacity, within
months the Army’s role changed to one of counter-terrorism as
soldiers increasingly came under attack from terrorist groups, chief
among them being the Provisional IRA. For the next three decades
the military, along with the RUC, was engaged in countering a
savage urban and rural insurgency, in which thousands were killed
and injured, until the signing of the Belfast Agreement in April 1998
finally brought the prospect of a lasting peace to the war-weary
country.
The conflict in Northern Ireland has often been called a ‘corporals’
war’, for it was the junior NCOs leading their ‘bricks’ on patrol day in,
day out on the dangerous streets of Belfast and Londonderry, and in
the deceptively tranquil countryside of the border areas, who were
at the forefront of the conflict.
But, like Vietnam, Northern Ireland was also a helicopter war.
Throughout the long years of Operation BANNER, helicopter crews of
the Army Air Corps, Royal Navy and Royal Air Force flew in support
of the thousands of security forces personnel serving in the province.
They sustained isolated bases in the border areas, evacuated
casualties, carried out aerial surveillance, patrolled the waters in and
around Northern Ireland and even, towards the end of the conflict,
served in a limited offensive capacity.
And nowhere was the air war in Northern Ireland more intense
than in South Armagh, an area that became known as Bandit
Country, due to its long history of lawlessness and to the strength of
the IRA there. If the helicopter was a valuable tool in other parts of
Northern Ireland, in South Armagh it was absolutely essential.
From the mid-1970s the danger posed by roadside IEDs to the
security forces’ mobile patrols in South Armagh forced soldiers off
the roads and into the air. By the 1980s Bessbrook Mill, the Army’s
main base in County Armagh, had become the busiest heliport in
Europe, with an average of 600 flights in and out of the base per
week.
In the observation role helicopters also became a vital element of
the Army’s counter-terrorism strategy. Along with fixed-wing types
like the de Havilland-Canada Beaver and Britten-Norman Islander,
helicopters fitted with increasingly sophisticated surveillance
equipment played a key role in many of the security forces’
operations, leading to the capture of suspects and forcing the
terrorist groups to abandon many of their attacks. Small wonder that
the IRA soon came to fear and loathe the helicopter, one volunteer
in their East Tyrone brigade stating in his memoirs that he dreamed
of watching them ‘fall out of the sky in flames’.
Turning that dream into reality became an overriding
preoccupation of PIRA’s feared South Armagh brigade. Regarded by
the security forces as the most skilled and disciplined of PIRA’s units,
they were not slow in recognizing just how dangerously dependent
the British Army had become on helicopters in South Armagh, and
set about trying to exploit that vulnerability, with rotary-wing aircraft
becoming one of their prime military targets from the mid-1970s
onwards.
Only in the skies over South Armagh, PIRA believed, could an
outright strategic victory over their enemy be achieved. If they could
make the skies as dangerous for the security forces as they had the
roads, the British Army would be effectively paralyzed. ‘We felt that
if we could nullify the helicopter, we would be well on the way to
winning the war,’ said one volunteer.1
The British Army itself concurred with that assessment. The
Army’s official report analyzing military operations in Northern
Ireland during Operation BANNER, published in 2006, stated: ‘Any
loss of control of the air would have seriously impeded the conduct
of security force operations on the ground’.2
Besides the purely military value of downing helicopters, there
was also a secondary – and perhaps equally important – aim for
PIRA in carrying out attacks on helicopters. As one of the world’s
most media-savvy militant nationalist movements, the Provisional
IRA was well aware of the immense propaganda value to be gained
from shooting down military aircraft. The Provisionals were keen to
portray themselves as a legitimate guerrilla army, engaged in an
armed struggle against what they characterized as a force of colonial
occupation. Attacking high-profile military targets like helicopters not
only guaranteed PIRA widespread media coverage but also fitted in
perfectly with the image they wished to convey of themselves as
heroic freedom fighters to their supporters, both at home and
abroad.
This explains why the organization went to such lengths during
the three decades of the ‘Troubles’ to bring down helicopters. From
taking pot-shots with Tommy guns and Garand rifles in the early
years of the conflict to meticulously planned ambushes by large
active service units of a dozen or more volunteers, armed with
rocket-propelled grenade launchers and 12.7mm heavy machine
guns, their determination to shoot down British military aircraft
became an obsession, one that would lead PIRA to embark on an
international arms hunt spanning more than a decade to procure the
one weapon they were convinced could tip the balance of the war in
South Armagh in their favour: the heat-seeking surface-to-air missile
launcher.
 Despite all that effort, their ‘kills’ were few. The Provisional IRA
never did succeed in turning South Armagh into another Vietnam, as
they hoped; the loss of Lieutenant Colonel Corden-Lloyd’s Gazelle in
February 1978 was one of just six helicopters brought down, either
directly or indirectly, by hostile fire during the ‘Troubles’.
But the figures alone do not tell the whole story, for as this book
reveals there were many other attacks which came terrifyingly close
to success. That PIRA ultimately failed in its stated goal to halt
movement by air in South Armagh, as well as the other border
regions where guerrilla activity was also intense, was due in large
part to the skill and professionalism of the Army, Royal Navy and
RAF crews.
What follows in these pages is the untold story of that air war
between the years 1969 and 2007 when the Provisional IRA and the
British security forces fought their long, gruelling battle for control of
the skies over Bandit Country.
Another random document with
no related content on Scribd:
received a dorsally-placed accessory gland, open to the exterior by a
median aperture placed ventrally a little way behind the mouth.
Life-history.—The egg undergoes a large portion of its
development within the body of the mother. In Linguatula
taenioides, which lives in the nasal cavities of the dog, the eggs pass
away with the nasal excretions. If these, scattered about in the grass,
etc., be eaten by a rabbit, the egg-shell is dissolved in the stomach of
the second host and a small larva is set free. In Porocephalus
proboscideus and others, which inhabit the lungs of snakes, the eggs
pass along the alimentary canal and leave the body with the faeces.
They also must be eaten by a second host if development is to
proceed.
The larva which emerges when
the egg-shell is dissolved has a
rounded body provided with two
pairs of hooked appendages, and
a tail which is more or less
prominent in different species
(Figs. 259, 260). Each appendage
bears a claw, and is strengthened
by a supporting rod or skeleton.
Anteriorly the head bears a
boring apparatus of several
chitinous stylets. The various
internal organs are in this stage
already formed, though in a Fig. 259.—A late larval stage of
somewhat rudimentary state, and Porocephalus proboscideus, seen from
the side. Highly magnified. (From
it is doubtful if the anus has yet Stiles.) 1, primordium of first pair of
appeared. chitinous processes; 2, primordium of
By means of its boring second pair of chitinous processes; 3,
apparatus, and aided by its mouth; 4, ventral ganglion; 5,
hooked limbs, the larva now receptaculum seminis; 6, oviduct; 7,
ovary; 8, anus; 9, vagina.
works its way through the
stomach-walls of its second host,
and comes to rest in the liver or in some other viscus. Its presence in
the tissues of its second host causes the formation of a cyst, and
within this the larva rests and develops. In man, at least, the cysts
often undergo a calcareous degeneration, and Virchow states “dass
 beim Menschen das
Pentastomum am häufigsten von
allen Entozoen zu
Verwechselungen mit echten
Tuberkeln Veranlassungen giebt.”
The larva moults several times,
and loses its limbs, which seem to
have no connexion with the
paired hooks in the adult (Fig.
256). The internal organs slowly
assume the form they possess in
the adult. The larva is at first
quite smooth, but as it grows the
annulations make their
appearance, arising in the middle
and spreading forward and
Fig. 260.—Larva of Porocephalus backward (Fig. 259). In this
proboscideus, seen from below. Highly encysted condition the larva
magnified. (From Stiles.) 1, Boring, remains coiled up for some
anterior end; 2, first pair of chitinous months, according to Leuckart;
processes seen between the forks of the
second pair; 3, ventral nerve-ganglion;
six in the case of L. taenioides,
4, alimentary canal; 5, mouth; 6 and 7, and a somewhat shorter
[385]
period,
gland-cells. according to Stiles, in the case
of P. proboscideus.
The frequency of what used to
be called Pentastoma denticulatum (= the larval form of L.
taenioides) in the body of man depends on the familiarity of man
with dogs. Klebs and Zaeslin found one larva in 900 and two in 1914
autopsies. Laenger[386] found the larva fifteen times in about 400
dissections, once in the mesentery, seven times in the liver, and
seven times in the wall of the intestine. After remaining encysted for
some time it may escape, and begins wandering through the tissues,
aided by its hooks and annulations, a proceeding not unaccompanied
by danger to its host. Should the latter be eaten by some carnivorous
animal, the larva makes its way into the nasal cavities or sinuses, or
into the lungs of the flesh-eating creature, and there after another
ecdysis it becomes adult. If, however, the second host escapes this
fate, the larvae re-encyst themselves, and then if swallowed they are
said to bore through the intestine
of the flesh-eater, and so make
their way to their adult abode.
Systematic.[387]—The
Pentastomida are a group much
modified by parasitism, which
has so deeply moulded their
structure as to obscure to a great
extent their origin and affinities.
The larva, with its clawed limbs,
recalls the Tardigrades and
certain Mites, e.g. Phytoptus,
where only two pairs of limbs
persist, and where the abdomen Fig. 261.—Encysted form of
is elongated and forms a large Porocephalus protelis, × 1, lying in the
proportion of the body. The mesentery of its host. (From Hoyle.)
resemblances to a single and
somewhat aberrant genus must
not, however, be pressed too far. The striated muscles, the ring-like
nature of the reproductive organs and their ducts, perhaps even the
disproportion both in size and number of the females to the males,
are also characters common to many Arachnids.
The Pentastomida include three genera, Linguatula, Fröhlich,
Porocephalus, Humboldt, and Reighardia, Ward.[388] The first two
were regarded by Leuckart as but sub-genera, but Railliet[389] and
Hoyle[390] have raised them to the rank of genera. They are
characterised as follows:—
Linguatula, body flattened, but dorsal surface arched; the edges of
the fluke-like body crenelated; the body-cavity extends as diverticula
into the edges of the body.
Porocephalus, body cylindrical, with no diverticula of the body-
cavity.
Reighardia, devoid of annulations, transparent, with poorly
developed hooks and a mouth-armature.
The following is a list of the species with their primary and
secondary or larval hosts:—
i. Linguatula pusilla, Diesing, found in the intestine of the fresh-
water fish Acara, a South American genus of the Cichlidae. This
is possibly the immature form of L. subtriquetra.
ii. L. recurvata, Diesing, found in the frontal sinuses and the
trachea of Felis onca.
iii. L. subtriquetra, Diesing, found in the throat of Caiman
latirostris and C. sclerops, perhaps the mature form of L. pusilla.
iv. L. taenioides, Lamarck, found in the frontal sinuses and nasal
chambers of the dog and ounce, and in the nasal cavities of the
wolf, fox, goat, horse, mule, sheep, and man, and in the trachea
of the ounce. The immature form has been found in or on the
liver of the cat, guinea-pig, and horse; in the lungs of the ox, cat,
guinea-pig, porcupine, hare, and rabbit; in the liver and
connective tissue of the small intestine of man; and in the
mesenteric glands of the ox, camel, goat, sheep, antelope, fallow-
deer, and mouse.
v. Porocephalus annulatus, Baird, found in the lungs of the
Egyptian cobra, Naja haje; the immature form is thought to live
encapsuled in a species of Porphyrio[391] and in the Numidian
Crane.
vi. P. aonycis, Macalister, from the lungs of an Indian otter taken
in the Indus.
vii. P. armillatus, Wyman, found in the adult state in the lungs of
certain African pythons, and in the lion; in the larval form it
occurs encysted in the abdomen of the Aard-wolf, the mandril,
and man—usually in negroes. Its migrations in the body of its
second host sometimes cause fatal results.
viii. P. bifurcatus, Diesing, found in the body-cavity of certain
snakes, and in the lungs of boa-constrictors and the legless
lizard, Amphisbaena alba. Possibly an immature form.
ix. P. clavatus, Lohrmann, found in the lungs of the Monitor lizard.
x. P. crocidura, Parona, found in the peritoneum of the “musk-rat”
Crocidura in Burmah. Probably a larval form.
xi. P. crotali, Humboldt, found in the lungs, body-cavity, kidneys,
spleen, and mesentery of many snakes and lizards, and of the
lion and leopard. The immature forms occur in the liver and
abdominal cavity of species of opossum, armadillo, mouse,
raccoon, bat, and marmoset.
xii. P. geckonis, Dujardin, found in the lungs of a Siamese gecko.
xiii. P. gracilis, Diesing, found free in the body-cavity or
encapsuled on the viscera and mesenteries of South American
fishes, snakes, and lizards,
xiv. P. heterodontis; Leuckart, found encapsuled in the abdominal
muscles and mesentery of a species of Heterodon.
xv. P. indicus,[392] v. Linst., found in the trachea and lungs of
Gavialis gangeticus.
xvi. P. lari, Mégnin, found in the air-sacs of the Burgomaster or
Glaucous gull, Larus glaucus of the Polar seas.
xvii. P. megacephalus, Baird, found embedded in the flesh of the
head of an Indian crocodile, C. palustris, the “Mugger.” Probably
a larval form.
xviii. P. megastomus, Diesing, found in the lungs of a fresh-water
tortoise, Hydraspis geoffroyana.
xix. P. moniliformis, Diesing, found in the lungs of pythons.
xx. P. najae sputatricis, Leuckart, found encapsuled in the
abdominal muscles and peritoneum of the cobra, Naja
tripudians. Probably a larval form.
xxi. P. oxycephalus, Diesing, found in the lungs of crocodiles and
alligators.
xxii. P. platycephalus, Lohrmann, habitat unknown.
xxiii. P. subuliferus, Leuckart, in the lungs of the cobra Naja haje.
xxiv. P. teretiusculus, Baird, found in the lungs and mouth of
certain Australian snakes.
xxv. P. tortus, Shipley, found in the body-cavity of a snake,
Dipsadomorphus irregularis, taken in New Britain.
xxvi. Reighardia, sp., Ward, found in the air-sacs of Bonaparte’s
gull and the common North American tern.
 PYCNOGONIDA

BY

D’ARCY W. THOMPSON, C.B., M.A. Trinity College

Professor of Natural History in University College, Dundee
 CHAPTER XXI
[393]
PYCNOGONIDA

Remote, so far as we at present see, from all other Arthropods, while yet
manifesting the most patent features of the Arthropod type, the Pycnogons
constitute a little group, easily recognised and characterised, abundant and
omnipresent in the sea. The student of the foreshore finds few species and seldom
many individuals, but the dredger in deep waters meets at times with prodigious
numbers, lending a character to the fauna over great areas.
The commonest of our native species,
or that at least which we find the oftenest,
is Pycnogonum littorale (Phalangium
littorale, Ström, 1762). We find it under
stones near low water, or often clinging
louse-like to a large Anemone. The squat
segmented trunk carries, on four pairs of
strong lateral processes, as many legs,
long, robust, eight-jointed, furnished each
with a sharp terminal claw. In front the
trunk bears a long, stout, tubular
proboscis, at the apex of which is the
mouth, suctorial, devoid of jaws; the body
terminates in a narrow, limbless,
unsegmented process, the so-called
“abdomen,” at the end of which is the
anal orifice. The body-ring to which is
attached the first pair of legs, bears a
tubercle carrying four eye-spots; and Fig. 262.—Pycnogonum littorale, Ström, × 2.
below, it carries, in the male sex, a pair of
small limbs, whose function is to grasp
and hold the eggs, of which the male animal assumes the burden, carrying them
beneath his body in a flattened coherent mass. In either sex a pair of sexual
apertures open on the second joints of the last pair of legs. The integument of body
and limbs is very strongly chitinised, brown in colour, and raised into strong bosses
or tubercles along the middle line of the back, over the lateral processes, and from
joint to joint of the limbs. The whole animal has a singular likeness to the Whale-
louse, Cyamus mysticeti (well described by Fr. Martins in 1675), that clings to the
skin of the Greenland Whale as does Pycnogonum to the Anemone, a resemblance
close enough to mislead some of the older naturalists, and so close that Linnaeus,
though in no way misled thereby, named it Phalangium balaenarum. The substance
of the above account, and the perplexity attending the classification of the animal,
are all included in Linnaeus’s short description:[394] “Simillimus Onisco Ceti, sed
pedes omnes pluribus articulis, omnes perfecti, nec plures quam octo. Dorsum
rubrum, pluribus segmentis; singulis tribus mucronibus. Cauda cylindrica,
brevissima, truncata. Rostrum membranaceum, subsubulatum, longitudine pedum.
Genus dubium, facie Onisci ceti; rostro a reliquis diversum. Cum solo rostro absque
maxillis sit forte aptius Acaris aut proprio generi subjiciendum.... Habitat in mari
norvegico sub lapidibus.”[395]
The common Pycnogonum is, by
reason of the suppression of certain
limbs, rather an outlying member than a
typical representative of the Order, whose
common characters are more strikingly
and more perfectly shown in species, for
instance, of Nymphon. Of this multiform
genus we have many British species, some
of the smaller being common below tide-
marks, creeping among weeds or clinging
like Caprellae with skeleton limbs to the
branches of Zoophytes, where their
slender forms are not easily seen. In
contrast to the stouter body and limbs of
Pycnogonum, the whole fabric of
Nymphon tends to elongation; the body is
drawn out so that the successive lateral
processes stand far apart, and a slender
neck intervenes between the oculiferous
tubercle and the proboscis; the legs are
produced to an amazing length and an
extreme degree of attenuation: “mirum
tam parvum corpus regere tam magnos
pedes,” says Linnaeus. Above the base of
the proboscis are a pair of three-jointed
appendages, the two terminal joints of
which compose a forcipate claw; below
Fig. 263.—Dorsal view of Nymphon brevirostre, and behind these come a pair of delicate,
Hodge, × 6. Britain. palp-like limbs of five joints; and lastly,
on the ventral side, some little way
behind these, we find the ovigerous legs
that we have already seen in the male Pycnogonum, but which are present in both
sexes in the case of Nymphon. At the base of the claw which terminates each of the
eight long ambulatory legs stands a pair of smaller accessory or “auxiliary” claws.
The generative orifices are on the second joint of the legs as in Pycnogonum, but as
a rule they are present on all the eight legs in the female sex, and on the two
hindmost pairs in the male. One of the Antarctic Nymphonidae (Pentanymphon)
and one other Antarctic genus less closely related (Decolopoda) have an extra pair of
legs. No other Pycnogon, save these, exhibits a greater number of appendages than
Nymphon nor a less number than Pycnogonum, nor are any other conspicuous
organs to be discovered in other genera that are not represented in these two: within
so narrow limits lie the varying characters of the group.
In framing a terminology for the parts and members of the body, we encounter an
initial difficulty due to the ease with which terms seem applicable, that are used of
more or less analogous parts in the Insect
or the Crustacean, without warrant of
homology. Thus the first two pairs of
appendages in Nymphon have been
commonly called, since Latreille’s time,
the mandibles and the palps (Linnaeus
had called them the palps and the
antennae), though the comparison that
Latreille intended to denote is long
abandoned; or, by those who leaned, with
Kröyer and Milne-Edwards, to the
Crustacean analogy, mandibles and
maxillae. Dohrn eludes the difficulty by
denominating the appendages by simple
numbers, I., II., III., ... VII., and this
method has its own advantages; but it is
better to frame, as Sars has done, a new
nomenclature. With him we shall speak of
the Pycnogon’s body as constituted of a
trunk, whose first (composite) segment is
the cephalic segment or head, better
perhaps the cephalothorax, and which Fig. 264.—Nymphon brevirostre, Hodge. Head,
terminates in a caudal segment or from below, showing chelophores, palps, and
abdomen; the “head” bears the proboscis, ovigerous leg.
the first appendages or “chelophores,” the
second or “palps,” the third, the false or
“ovigerous” legs, and the first of the four pairs of “ambulatory” legs. The chelophores
bear their chela, or “hand,” on a stalk or scape; the ambulatory legs are constituted
of three coxal joints, a femur, two tibial joints, a tarsus, and a propodus, with its
claws, and with or without auxiliary claws.
The Body.—The trunk with its lateral processes may be still more compact than
in Pycnogonum, still more attenuated than in Nymphon.
In a few forms (e.g. Pallene, Ammothea, Tanystylum, Colossendeis) the last two,
or even more, segments of the trunk are more or less coalescent. In Rhynchothorax
the cephalic segment is produced into a sharp-pointed rostrum that juts forward
over the base of the proboscis. The whole body and limbs may be smooth,
tuberculated, furnished with scattered hairs, or sometimes densely hispid.
 Fig. 265.—A, Colossendeis proboscidea, Sabine, Britain; B, Ammothea echinata,
Hodge, Britain; C, Phoxichilus spinosus, Mont., Arctic Ocean. (The legs omitted.)

The proboscis varies much in shape and size. It may be much longer or much
shorter than the body, cylindrical or tumid, blunt or pointed, straight or (e.g.
Decolopoda) decurved; usually firmly affixed to the head and pointing straight
forwards; sometimes (Eurycide, Ascorhynchus) articulated on a mobile stalk and
borne deflexed beneath the body.
Chelophores.—The first pair of appendages or chelophores are wanting in the
adult Pycnogonum, Phoxichilus, Rhynchothorax, and Colossendeis.[396]
In Ammothea and its allies they are extremely rudimentary in the adult, being
reduced to tiny knobs in Tanystylum and Trygaeus, and present as small two-
jointed appendages in Ammothea; in this last, if not in the others also, they are
present in complete chelate form in the later larval stages.
 Fig. 266.—A, B, Chelophores of Ascorhynchus abyssi, G.O.S. A, Young; B, adult.
(After Sars.) C, Anterior portion of Ammothea hispida, Hodge, Jersey: late larval
stage (= Achelia longipes, Hodge), showing complete chelae. D, Chela of
Eurycide hispida, Kr.

In Eurycide, Ascorhynchus, and Barana they are usually less atrophied, but yet
comparatively small and with imperfect chelae, while in some Ascorhynchi (A.
minutus, Hoek) they are reduced to stumps.

Fig. 267.—Chelae of species of Nymphonidae: A, Nymphon brevirostre, Hodge;

B, Boreonymphon robustum, Bell; C, Chaetonymphon macronyx, G.O.S.; D,
Nymphon elegans, Hansen.

In Pallenopsis the scape of the chelophore consists of two joints, as also in

Decolopoda and some Ascorhynchus: in Nymphon, Phoxichilidium, Pallene, and
Cordylochele of one only; in all these the terminal portion or “hand” forms a
forcipate “chela,” of which the ultimate joint forms the “movable finger.” In some
species of Nymphon the chela is greatly produced and attenuated, and armed with
formidable serrate teeth on its opposing edges; in others it is shortened, with blunter
teeth; in Boreonymphon robustum the
claws are greatly curved, with a wide gap
between. In this last, and in
Phoxichilidium, the opposing edges are
smooth and toothless. In Cordylochele
the hand is almost globular, the movable
finger being shortened down, and half
enclosed by the other.
Palpi.—The second pair of
appendages, or palps, are absent, or all
but absent, in the adult Pycnogonum,
Phoxichilus, Phoxichilidium, Pallene, and
their allies. In certain of these cases, e.g.
Phoxichilidium, a knob remains to mark
their place; in others, e.g. Pallenopsis, a
single joint remains; in a few Pallenidae a
sexual difference is manifested, reduction
of the appendage being carried further in
the female than in the male. The Fig. 268.—Proboscis and chelophores of
composition of the palps varies in the Cordylochele longicollis, G.O.S. (After Sars.)
genera that possess them. In Nymphon
there are five joints, and their relative
lengths (especially of the terminal ones) are much used by Sars in defining the many
species of the genus. The recently described Paranymphon, Caullery, has palps of
six or seven joints. In the Ammotheidae the number of joints ranges from five or six
in Tanystylum to nine (as a rule) in Ammothea and Oorhynchus, or ten, according
to Dohrn, in certain species of Ammothea. Colossendeis and the Eurycididae have a
ten-jointed palp, which in this last family is very long and bent in zigzag fashion, as
it is, by the way, also in Ammothea. The terminal joints of the palp are in all cases
more or less setose, and their function is conjecturally tactile.
Ovigerous Legs.—Custom sanctions for these organs an inappropriate name,
inasmuch as it is only in the males that they perform the function which the name
connotes.[397] They probably also take some part, as Hodgson suggests, in the act of
feeding.
Fig. 269.—Eurycide hispida, Kr., showing
stalked proboscis and zigzag palps.

Fig. 270.—Ovigerous legs of A, Phoxichilus spinosus, Mont.; B, Phoxichilidium

femoratum, Rathke; C, Anoplodactylus petiolatus, Kr.; D, Colossendeis
proboscideus, Sab.
 In Pycnogonum, Phoxichilus,
Phoxichilidium, and their immediate
allies they are absent in the female; in all
the rest they are alike present in both
sexes, though often somewhat smaller in
the female than in the male. They are
always turned towards the lower side of
the body, and in many cases even their
Fig. 271.—Terminal joints of ovigerous leg of point of origin is wholly ventral. The
Rhynchothorax mediterraneus, Costa. number of joints varies: in

Phoxichilidium five, Anoplodactylus six,

Phoxichilus seven; in Paranymphon
eight; in Pycnogonum nine, with, in
addition, a terminal claw; in the
Ammotheidae from seven (Trygaeus) to
ten, without a claw; in Pallenidae ten,
with or without a claw; in
Rhynchothorax, Colossendeis, Eurycide,
Ascorhynchus, Nymphon, ten and a claw.
The appendage, especially when long, is
apt to be wound towards its extremity
into a spiral, and its last four joints
usually possess a peculiar armature. In
Rhynchothorax this takes the form of a
stout toothed tubercle on each joint; in
Colossendeis of several rows of small Fig. 272.—Nymphon brevirostre, Hodge.
Terminal joints of ovigerous leg, with magnified
imbricated denticles; in Nymphon and “tooth.”
Pallene of a single row of curious serrate
and pointed spines, each set in a little
membranous socket.
 Fig. 273.—Nymphon strömii, Kr. Male carrying egg-masses on his ovigerous
legs.

Fig. 274.—Terminal joints (tarsus and propodus) of legs. 1, Chaetonymphon

hirtum, Fabr.; 2, N. strömii, Kr.; 3, Nymphon brevirostre, Hodge; 4, Ammothea
echinata, Hodge; 5, Ascorhynchus abyssi, G.O.S. (All after Sars.)

Legs.—The four pairs of ambulatory legs are composed, in all cases without
exception, of eight joints if we exclude, or nine if we include, the terminal claw. They
vary from a length about equal to that of the body (Pycnogonum, Rhynchothorax,
Ammothea) to six or seven times as much, perhaps more, in Nymphon and
Colossendeis, the fourth, fifth, and sixth joints being those that suffer the greatest
elongation. The seventh joint, or tarsus, is usually short, but in some Nymphonidae
is much elongated; the eighth, or propodus, is usually somewhat curved, and usually
possesses a special armature of simple or serrate spines. The auxiliary claws,
sometimes large, sometimes small, lie at the base of the terminal claw in
Ammotheidae, Phoxichilidae, in Phoxichilidium, in most Pallenidae, in nearly all
Nymphonidae. Their presence or absence is often used as a generic character,
helping to separate, e.g., Pallene from Pseudopallene and Pallenopsis, and
Phoxichilidium from Anoplodactylus; nevertheless they may often be detected in a
rudimentary state when apparently absent. The legs are smooth or hirsute as the
body may happen to be.
Fig. 275.—Legs of A, Pallene brevirostris, Johnston; B, Anoplodactylus
petiolatus, Kr.; C, Phoxichilus spinosus, Mont.; D, Colossendeis proboscidea,
Sabine; E, Ammothea echinata, Hodge, ♂.
 Fig. 276.—Boreonymphon robustum, Bell. Male with young, slightly enlarged.
Faeroe Channel.

Glands.—In some or all of the appendages of the Pycnogonida may be found

special glands with varying and sometimes obscure functions. The glands of the
chelophores (Fig. 280, p. 522) are present in the larval stages only. They consist of a
number of flask-shaped cells[398] lying within the basal joint of the appendage, and
generally opening at the extremity of a long, conspicuous, often mobile, spine (e.g.
Ammothea (Dohrn), Pallene, Tanystylum (Morgan), Nymphon brevicollum and N.
gracile (Hoek)). They secrete a sticky thread, by means of which the larvae attach
themselves to one another and to the ovigerous legs of the male parent. In Nymphon
hamatum, Hoek, the several filaments secreted by the separate sacculi of the gland
issue separately. In Pycnogonum the spine on which the gland opens is itself
prolonged into a long fine filament, and here, according to Hoek, the gland is in all
probability functionless and rudimentary. Hoek has failed to find the gland in
Ascorhynchus, and also in certain Nymphonidae (e.g. Boreonymphon robustum,
Bell), in which the young are more than usually advanced at the time of hatching.
The gland has also been described by Lendenfeld and others in Phoxichilidium,
whose larvae do not cling together but live a parasitic life; in this genus the long
spine or tubercle is absent on which the orifice is usually situated, and, according to
Lendenfeld, the secretion issues from many small orifices set along the opposing
edges of the chela. Of the two species described by Dohrn as Barana castelli and B.
arenicola, the former has the spine of inordinate length, more than twice as long as
the whole body, chelophore and all; while in the latter (which species rather
resembles Ascorhynchus) the spine is altogether absent.
In the palps and ovigerous legs of the adult are found glandular bodies of a hollow
vesicular form with a simple lining of cells, the vesicle being divided within by a
septum with a central orifice, the outer and smaller half opening to the exterior.
These glands are probably of general occurrence, but they have been but little
investigated. They lie usually in the fourth and fifth joints of the palp, and the third
and fourth joints of the ovigerous leg. Hoek describes them in Discoarachne
(Tanystylum) as lying within the elongated third joint of the palp, and opening by a
sieve-plate at the end of the second joint. In Ammothea (Dohrn) and Ascorhynchus
(Hoek) they open on a small tubercle situated on the fifth joint of the palp. In
Nymphon, Hoek describes them as opening by a small pore on the fourth joint of the
ovigerous leg. Dohrn failed to find them in Pycnogonum, but in Phoxichilus,
Phoxichilidium and Pallene he discovered the glands appertaining to the palps,
though the palps themselves have disappeared in those genera; he has found the
glands also in Ammothea, in larvae that have not yet attained their full complement
of legs.
The males in nearly all cases are known to possess glands in the fourth joints or
thighs of all the ambulatory legs, and these glands without doubt act as cement-
glands, emitting, like the chelophoral glands of the larvae, a sticky thread or threads
by which the eggs and young are anchored to the ovigerous legs. In some species of
Nymphon and of Colossendeis Hoek could not find these, and he conjectures them
to be conspicuous only in the breeding season. While in most cases these glands
open by a single orifice or by a few pores grouped closely together, in Barana,
according to Dohrn, and especially in B. arenicola, the pores are distributed over a
wide area of the femoral joint.[399] In Discoarachne (Loman) and Trygaeus they
open into a wide chitinised sac with tubular orifice. While the function of these last
glands and of the larval glands seems plain enough, that of those which occur in the
palps and ovigerous legs of both sexes remains doubtful.
In their morphological nature the two groups of glands are likewise in contrast,
the former being unicellular glands, such as occur in various parts of the integument
of the body and limbs of many Crustacea; while the latter are segmentally arranged
and doubtless mesoblastic in origin, like the many other segmental excretory organs
(or coelomoducts) of various Arthropods.
By adding colouring matters (acid-fuchsin, etc.) to the water in which the animals
were living, Kowalevsky demonstrated the presence of what he believed to be
excretory organs in Phoxichilus, Ammothea, and Pallene. These are small groups of
cells, lying symmetrically near the posterior borders of the first three body-
segments, and also near the bases of the first joints of the legs, dorsal to the
alimentary canal.[400]
Alimentary System.—The proboscis is a very complicated organ, and has been
elaborately described by Dohrn.[401] It is a prolongation of the oral cavity, containing
a highly developed stomodaeum, but showing no sign of being built up of limbs or
gnathites. The mouth, situated at its apex, is a three-sided orifice, formed by a
dorsal[402] and two lateral lobes; and hence the proboscis has been assumed by some,