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The Biology of Wetas, King Crickets and Their Allies
The Biology of Wetas,
King Crickets and Their Allies
Edited by
L.H. Field
Department of Zoology
University of Canterbury
New Zealand
CABI !"#$%&'%()
CABI Publishing is a division of CAB International
© CAB International 2001. All rights reserved. No part of this publication may be
reproduced in any form or by any means, electronically, mechanically, by
photocopying, recording or otherwise, without the prior permission of the copyright
owners.
A catalogue record for this book is available from the British Library,
London, UK.
A catalogue record for this book is available from the Library of Congress,
Washington DC, USA
Contributors ix
Foreword xi
David C.F. Rentz
Preface xiii
Introduction xv
Laurence H. Field
v
vi Contents
10 Sexual Selection and Secondary Sexual Characters of Wetas and King Crickets 179
Laurence H. Field and Neil A. Deans
III Ecology
13 The Gallery-related Ecology of New Zealand Tree Wetas, Hemideina femorata and
Hemideina crassidens (Orthoptera, Anostostomatidae) 243
Laurence H. Field and Graham R. Sandlant
IV Behaviour
20 The Reproductive Biology and the Eggs of New Zealand Anostostomatidae 379
Ian A.N. Stringer
VI Physiology
Index 529
Contributors
ix
x Contributors
Barry O’Brien, Department of Biological Sciences, University of Waikato, Private Bag 3105, Hamilton,
New Zealand. email: b.obrien@waikato.ac.nz
Robert Poulin, Department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand.
email: zoodw@zoo.latrobe.edu.au
David C.F. Rentz, Division of Entomology, CSIRO, Canberra, ACT 2601, Australia. email:
dave.rentz@ento.csiro.au
Graham R. Sandlant, Department of Zoology, University of Canterbury, Private Bag 4800,
Christchurch, New Zealand. email: zool146@it.canterbury.ac.nz
Greg Sherley, Science and Research Division, Science Technology and Information Service,
Department of Conservation, PO Box 10420, Wellington, New Zealand. email:
gsherley@doc.govt.nz
Ian A.N. Stringer, Ecology Group, Institute of Natural Resources, Massey University, PO Box 11222,
Palmerston North, New Zealand. email: i.stringer@massey.ac.nz
Rob B. Toms, Department of General Entomology, Transvaal Museum, Northern Flagship Institution,
PO Box 413, Pretoria 0001, South Africa. email: toms@nfi.co.za
Steve Trewick, Department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand.
email: mary.morgan-richards@stonebow.otago.ac.uk
Claudine L. Tyrrell, Department of Zoology, University of Otago, PO Box 56, Dunedin, New
Zealand. email: zoodw@zoo.latrobe.edu.au
Thomas J. Walker, Department of Entomology and Nematology, University of Florida, Gainesville,
FL 32611, USA. email: tjw@gnv.ifas.ufl.edu
David B. Weissman, Department of Entomology, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118, USA. email: gryllus1@juno.com
David A. Wharton, Department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand.
email: zoodw@zoo.latrobe.edu.au
Anna York, School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New
Zealand. email: b.lewis@auckland.ac.nz
Foreword
The wetas, king crickets and their allies are some A peculiarity of king crickets is the bizarre
of the most fascinating members of the insect production of secondary sexual characters, such as
world. Some are the heaviest of insects, dwarfing enlarged heads, mandibles and hind legs. These
even some small mammals in weight. Their large can be asymmetrical as well. Extensive behavioural
size and often ferocious nature always engenders studies are reported here. Odd weaponry, such as
public interest. This book has been long awaited. tusks and mandibular extensions and protrusions,
Orthopterists have always had ‘trouble’ dealing suggest bizarre courtship rituals unlike those
with these insects because they appear to bear no found in other orthopteroids.
characters distinctive for species separation or sec- Larry Field has assembled a most interesting
ondary sexual characters have been misinter- array of topics dealing with the morphology,
preted. The insects included in this book are for reproduction, physiology, behaviour, biogeogra-
the most part members of the orthopterous super- phy, systematics and ecology of these insects. The
families Stenopelmatoidea and Gryllacridoidea, chapter on conservation should prove valuable for
collectively known as king crickets. However, the the continuity of measures leading to the contin-
common name ‘bugaboo’ has helped to confuse ued existence of threatened species. The review of
the situation with ‘king cricket’, being the com- the Gryllacrididae, noted above, should prove use-
mon name for quite different insects, depending ful in relating their biology, behaviour, etc. to those
on geography. This is the reason for including the of the true king crickets. This book will go far in
review of the Gryllacrididae here. All these insects helping to answer questions that have long puzzled
are of Gondwanan origin and are important ‘flag- orthopterists. If it stimulates further investigations
ship species’ for biogeographical studies. But, as a of these insects, it will have more than fulfilled its
result of human-initiated activities, including purpose.
introduced animals, such as rats and cane-toads,
and development, a number of species are threat- D.C.F. Rentz
ened with extinction. Canberra, January 2000
xi
Preface
This book was conceived at the Hilo conference of Philadelphia Academy of Science, and to John
the Orthopterists Society, where, owing to the Edwards for use of unpublished figures of tree
enthusiastic interest shown, I became strongly weta cercal sensory anatomy and histology. I thank
aware that no previous synthesis of knowledge the following for reviewing chapters and for
existed for this primitive group of orthopterans. proofreading: David Rentz, Ian Stringer, Richard
Encouraged by Win Bailey and Dan Otte, I set Holdaway, George Gibbs, John Edwards, Piotr
about gathering together all the published Naskrecki, Paul Edwards, Hiroshi Nishino, Neil
research, as well as work buried in government Deans, Frank Howorth and Barry O’Brien. My
coffers, and unpublished research on the wetas of thanks also to Tim Hardwick and Emma Critchley
New Zealand. It soon became obvious that the of CAB International for their enduring patience
scope should be extended worldwide, since addi- while the book was being written. The figures
tional unpublished and ongoing research existed were digitized by Catherine Collins and Craig
in other southern hemisphere countries and the Russell, two young and talented digital artists,
USA. The resulting book therefore represents a who never thought that they would be confronted
timely foundation for the ever increasing research with the exacting labour of making every last sci-
on these stenopelmatoid insects. entific detail correct. Finally, I owe especial grati-
I owe much to the contributing authors and to tude to my wife Mitsuyo for her patience and
past students and colleagues for their efforts in endurance of the stresses involved in this project.
making this book possible. I am particularly grate- This book was funded by a grant from the
ful to Dan Otte for offering to illustrate the male University of Canterbury and supported by the
secondary sexual characters of many of the facilities of its Department of Zoology.
strange anostostomatids in the collections of the
xiii
Introduction
Laurence H. Field
Department of Zoology, University of Canterbury, Private Bag 4800,
Christchurch, New Zealand
Experiences and Background insects a fascinating pursuit for both biologist and
lay person alike. Their attraction lies not in any
In every biologist there may be kindled a sense of cuddly or virtuous appearance but, instead, in
wonderment and discovery, something akin to that their eccentric and often bizarre features and
which fired the excitement of the 18th- and 19th- behaviour. You immediately sense that here is
century naturalists, when first encountering a large something unusual indeed, photogenic because the
king cricket, weta or Jerusalem cricket. Add to that insect’s enormous head and threatening mandibles
a night-time setting in a far-flung South Pacific are all out of proportion with its body, or because
island reserve, offshore from New Zealand, where it has stout, curved tusks projecting from its head,
your torchlight is roving across the low-hanging like some mythical beast in miniature, or because it
fronds of Nikau palms and suddenly picks up the emerges from a shovelful of desert sand into the
enormous tell-tale antennae of a giant weta the daylight like a hidden monster living a subter-
size of a small rat, perched on the green blades. If ranean life, unknown and unseen by the world at
you’re careful and don’t move suddenly, you can large.
coax the nearly extinct giant insect on to your The Stenopelmatoidea is a superfamily of
palm, which it fills entirely with its leg span. In ensiferan Orthoptera, commonly known as wetas,
another land, you press your memory into action king crickets, Jerusalem crickets and raspy crick-
as you walk carefully at night, trying to recall the ets. The group encompasses a huge variety of
vague trail through the wet and overhanging cricket-like insects, many of which are flightless
vegetation of the North Queensland montane and which seem rather closer to their ancient ori-
rainforest where you laid the rotting pineapple bait gins than do the more modern grasshoppers,
stations the previous afternoon. Suddenly your crickets and katydids (long-horned crickets). Most
light beam encounters a bait covered with a swarm are nocturnal and infrequently encountered,
of waving antennae on the muddy ground up unless attracted by bait or sought by excavation or
ahead. They belong to a group of long-legged night lighting. In New Zealand, Australia and
‘white-knees’, the flightless king cricket Penalva. North America, certain species are abundant
But just on the edge is lurking a huge-headed male enough to be studied in their habitats or main-
of the giant Australian king cricket Anostostoma, tained in laboratory colonies. Much of the present
looking prehistoric and bizarre, with great curved volume stems from 25 years of research by the edi-
mandibles, which make the whole head seem tor, and his students and colleagues, on many
nearly as long as the body. aspects of the biology of New Zealand wetas.
These are the real-life experiences that make An explanation of the term ‘weta’ is necessary
the study of this superfamily of orthopteran at the outset. This Maori word refers generically
xv
Exploring the Variety of Random
Documents with Different Content
Yes, Daddy.
Millman
[Examining an envelope.]
Here’s an envelope from your aunt Mary—and look what’s in it!
[He draws out a yellow-backed bill.]
David
You take care of it for me, Daddy.
Millman
[Putting the envelope in his pocket.]
Here’s an electric train from Santa. Switches, and stations, and coal-
cars—I declare, it’s a wonderful train. Isn’t it, Halligan?
Halligan
It is that, sir.
Millman
You ought to be a very happy boy, Davy.
David
Yes, Daddy.
Millman
[Glancing at his watch again.]
I must go now. You don’t mind, do you, Davy? Important cablegrams
—letters—
David
I understand, Daddy.
Millman
[Already in the doorway.]
By the way, there’s a present from me downstairs—a new limousine.
Later on you’ll go riding in it.
David
[Rushing to him with pathetic eagerness.]
Will you come too, Daddy?
Millman
Sorry, Davy—haven’t time.
[He pats the boy’s head and nods pleasantly to the
others.]
Good morning.
[The door closes behind him, and David, as if struck by a
blow, withdraws again into his shell.]
Vicky
A fine gentleman, Mr. Millman!
Halligan
That he is!
Vicky
Did you see the check he gave me?
[Shows it.]
Halligan
Just take a look at this one!
Vicky
[Fervently.]
The salt of the earth!
Halligan
A fine gentleman!
David
[Who has been motionless, now turns to Halligan, in
joyous assent.]
A fine gentleman, isn’t he, Halligan?
[Proudly.]
When my father walks along the street everybody stops to look at
him! I can see the people nudging each other, and saying, “There
goes Mr. Millman.” And the newspapers send men here to take his
picture—but father’s too busy to let them do that. And when I go out I
hear everybody say, “There goes Millman’s boy.” And all the
policemen touch their caps and talk to me.
Halligan
Yes, I’ve seen that many a time.
David
And it’s all because daddy’s such a fine man!
Halligan
[Finding the statement difficult to correct.]
Yes; I guess that’s the reason.
[He is in distress.]
Vicky
[Coming to the rescue.]
Now you may look at the tree, Master David.
David
[Suddenly unenthusiastic, barely glancing at the tree.]
It’s very nice.
Vicky
Is that all you can say about it?
David
It’s just as nice as the one we had last year—and that was the nicest
I ever saw.
Vicky
[Taking up the gifts.]
Boxing gloves!
David
[Tentatively, knowing what to expect.]
May I put them on?
Vicky
Some day, perhaps—not just yet. Sleds!
David
May I go coasting on them?
Vicky
Next year, maybe—not now. A trumpet?
David
May I blow on it?
Vicky
Mercy, no! Not until it has been boiled.
David
But that’ll spoil the paint.
Vicky
Better to spoil the paint than to ruin your health.
David
[Disappointed.]
Oh, all right.
Halligan
[Coming to the rescue in his turn.]
I have a list of the presents here.
[He produces a long sheet.]
Two railroad trains—complete.
David
[Not boasting; simply stating facts.]
I have four already.
Halligan
Two phonographs.
David
I have three—and I can’t play more than one at once.
Halligan
Your cousin Willy sent you a set of books.
David
And what did I send him?
Halligan
[Referring to the list.]
You sent him a Boy Scout outfit.
David
Why didn’t he keep the books and send me the outfit?
Halligan
[Avoiding the question.]
Your aunt Genevieve sent you a bicycle.
David
[Interested despite himself.]
Oh, that’s nice!
[He moves toward it.]
Vicky
[Interposing.]
You may ride on it when you’re older.
David
But not now?
Vicky
[With real solicitude.]
You might hurt yourself, Master David.
David
[Crestfallen; turning to Halligan.]
Don’t read me any more, Halligan.
[Christmas is a complete failure so far as he is concerned.
Vicky realizes it, and directs his attention to the gifts.]
Vicky
Look: a tennis racket.
David
[Despite himself.]
That’ll be fun this summer!
Halligan
It will that!
David
I may play with it?
Vicky
All you like.
David
Good! Watch my overhand!
[He swings the racket.]
Vicky
[Taking it from him.]
Not in the house, Master David; you’ll break something.
David
I knew there was a string tied to it.
Vicky
[Indicating.]
Golf clubs.
David
For the summer?
Vicky
Naturally for the summer.
David
[Nodding.]
I see; it’s winter now.
[He sees a pair of ice skates, and takes them up.]
Do these have to wait for the summer, too?
Halligan
[Impressively.]
You’re to go to the rink with them this afternoon.
David
That will be nice!
Vicky
They’re a gift from us, Master David—
Halligan
From me and her.
David
[Sincerely.]
Thank you, Vicky.
[He kisses her.]
Thanks, Halligan.
[He shakes hands.]
You couldn’t have given me anything I’d like better.
[He feels the edge.]
They’re sharp, aren’t they?
Halligan
I saw to that.
Vicky
[Alarmed.]
Be careful, Master David!
David
[Smiling.]
You aren’t going to get them away from me!
[He gives them to Halligan, and turns back to the tree.]
Look!
Halligan
[Following his glance.]
Candy!
Vicky
[Horrified.]
Candy? Who sent it?
Halligan
It’s not down on the list.
Vicky
[Kneeling to examine it.]
There’s no card.
David
I guess it came from Santa Claus.
Vicky
[Reluctantly.]
Of course you mayn’t eat it.
David
That’s all right, Vicky; I don’t mind.
Vicky
[Who has opened the box.]
There’s no card inside, but it looks lovely.
David
I mayn’t have any, Vicky; eat it yourself.
Vicky
Do you really want me to?
David
Of course, Vicky.
Vicky
[Eating and smacking her lips.]
They’re good! Have one, Mr. Halligan?
Halligan
I don’t mind if I do.
[He eats a candy.]
David
[Watching with interest.]
What do they taste like?
Vicky
Chocolate—
[Taking another.]
Halligan
With strawberry cream inside—
[Taking another.]
Vicky
This one has a cherry.
Halligan
This has a walnut.
David
Santa Claus makes good candy, doesn’t he? Some day, when I’m
older, he’ll make some that I can eat. I’d like that!
Vicky
Another, Mr. Halligan?
Halligan
I don’t mind if I do.
[He pauses, and looks toward the door.]
Now, it isn’t up to me to say what I’m thinkin’, and nobody knows that
better than myself, but it’s cruel not to let him have a taste.
Vicky
Mr. Halligan!
Halligan
Who ever heard of candy hurtin’ anybody?
Vicky
Orders are orders, and they’ll be followed!
[She relents, and, about to put the lid on the box, offers it
a last time to Halligan.]
More?
Halligan
[Swallowing hard and passing his hand over his forehead.]
No, thanks.
Vicky
[Rising suddenly and tottering.]
Mr. Halligan—I don’t feel well. Please get me some water.
Halligan
[Alarmed; hastening out of the room.]
Right away!
David
What’s the matter, Vicky? Vicky, dear?
Vicky
It’s nothing, Master David. It’ll pass away in a minute.
[She sways, and David steadies her.]
I feel dizzy—very dizzy—all of a sudden.
David
Sit down, Vicky.
Vicky
[Catching the back of a chair.]
I can’t imagine what’s wrong. Nothing like this has ever happened to
me before. Oh, dear! Oh, dear! Why doesn’t Halligan bring the
water? Why doesn’t he bring it? I’m so dizzy—so dizzy.
[From the hall at the right there is the sound of a heavy
fall, accompanied by the crash of breaking glass.]
David
[Alarmed.]
Halligan fell!
Vicky
[Reeling toward the door.]
Mr. Halligan! Mr. Halligan! I’m afraid I’m going to faint.
[On the threshold a sudden suspicion comes to her, and
she pulls herself together with a heroic effort.]
David! Davy, boy. Don’t touch the candy!
[She collapses on the threshold.]
David
[On his knees at her side.]
Vicky! Vicky, dear! Answer me, Vicky!
[During the last few seconds the window has been raised,
and Slim has come into the room.]
Slim
[To Bill, who follows.]
It woiked.
Bill
De kid didn’t eat de candy.
Slim
De udders did—dat suits me.
David
[Rising to confront the newcomers.]
What are you doing here?
Slim
We come after yuh—
Bill
[Pushing Slim to one side easily.]
Nuttin’ to get excited about, kid; yer lady friend’s all right—see?
[He leads David back into the room; Slim thrusts the door
shut, and locks it.]
She’s just daydreamin’—takin’ a little cat nap. It won’t hoit her a bit—
honest! She’ll feel fine when she wakes up.
David
What happened to Halligan?
Bill
He’s daydreamin’, too. De two of ’em are daydreamin’ togedder—
nice an’ sociable-like—see? Dey’re dreamin’ about de little boidies
singin’ in de tree tops. Ain’t dat pretty?
[Slim has come forward. Bill waves a hand.]
Meet my friend Slim.
David
[Extending a hand.]
How do you do?
Slim
[Shaking hands.]
Pleased to meet-cha.
Bill
Slim an’ me—we’re gonna look after yuh fer a w’ile.
David
Yes?
[He looks up at Bill with sudden recognition.]
You don’t have to tell me who you are!
Bill
[Worried.]
I don’t?
David
I’ve seen you before!
Bill
Yuh know my name?
David
Of course! Who doesn’t?
[He pauses while Bill plainly shows his anxiety.]
Why, you’re Santa Claus!
Slim
[Overcome and relieved.]
W’at? W’at did yuh say?
David
[Laughing.]
You’re Santa Claus, and you know you are!
Slim
Ha! Ha!
[He breaks into guffaws.]
Bill
[Poking Slim violently in the midriff with his elbow.]
Yuh guessed it right de very foist time, kid. John W. Santa—dat’s
me!
[He tidies his impossible beard and whiskers.]
David
I knew you right off!
Bill
Yuh sure did!
David
[Intensely interested, catching his hand.]
Did you have a cold trip coming here?
Bill
W’at’s dat?
David
Wasn’t it cold, coming all the way from the North Pole?
Bill
Well, it wasn’t so bad after we got to a Hunner an’ Twenty-fift’ Street
—
David
[Fascinated.]
No?
Bill
Den de goin’ was pretty good.
David
But before you got there?
Bill
It was a wee bit chilly.
Slim
It was sixty below.
David
Sixty below what?
Bill
Not below nuttin’. Just below—see?
[He gesticulates vividly, placing his hand parallel to the
floor at the level of his ankles.]
Dat was w’ere I caught cold. I gotta sneeze.
Slim
Now!... Now!... Now!
David
Oh, let him sneeze!
[Bill sneezes.]
God bless you!
Bill
Much obliged.
David
That’s all right. I always say “God bless you” when anybody
sneezes.
Slim
[Returning to the main topic.]
We’re gonna take care of yuh—me an’ Bill.
Bill
We’re gonna take yuh fer a long ride.
David
Are we going to the North Pole?
Bill
Foider den dat.
[Approaching him.]
But you gotta keep quiet!
David
[With a nod of comprehension.]
I know; you don’t want to frighten the reindeer.
Slim
[Alarmed.]
De w’ich?
David
Blixen and Vixen—
Bill
W’at?
David
—And Prancer and Dancer—
Slim
[Decidedly worried, to Bill.]
Did youse see any of dem guys w’en yuh come in?
Bill
Maybe dey was under cover.
[To David.]
Say, kid, w’ere do dey keep?
David
[Puzzled.]
Keep?
Bill
W’ere do dey hang out? W’ere do dey park? W’ere’s deir stampin’-
ground?
David
Oh, outside!
Bill
[To Slim.]
I told yuh de house was watched!
David
[Quoting some book.]
“Drawing Santa Claus from his home in the North, reindeer, swifter
than the wind, swift as light—”
Slim
[Beginning to understand.]
Hey! I get him now! He’s talkin’ about a noo kind of flivver!
David
“Swifter even than dreams, sturdy and strong, champing at their bits,
sparks coming from their nostrils—”
Bill
[Nodding.]
De kid’s got de right dope, Slim.
[To David.]
Dey’re waitin’ fer us outside: balloon tires, an’ four-w’eel brakes, an’
sparks just w’ere yuh said. Come on.
David
[Going toward the window.]
They’re in a big hurry, aren’t they? They know we’re coming, Santa
Claus. They can’t wait for us! I hear them shaking their sleigh bells!
[Sleigh bells are audible.]
Bill
[Much alarmed.]
Do yuh hear dat, Slim?
David
[With glee.]
Sleigh bells!
Slim
Shh!
[They hide in corners of the room. David cannot
understand their actions; he looks about, puzzled.
Then a ten-year-old girl, wearing a harness covered
with sleigh bells, appears at the window.]
Bessie
[Softly.]
Bill!
[More loudly.]
Bill! Bill!
[She spies David.]
Is he here?
[David nods silently, and indicates with his thumb where
Bill is hiding. Bessie climbs into the room, bells
jangling.]
Bill, we found yuh!
[Climbing through the window come seven more children,
in decreasing sizes. The smaller ones are helped by
the larger, and the smallest, which is but an infant, is
carried by one of the others.]
A Babble of Talk
Hey, give us a hand, Pete!
Look out!
Yuh’ll fall!
Mind de baby!
Gimme a good push!
Hey, you, Woodrow, quit yer crowdin’!
Up yuh go!
[Now that they are all in the room, we may pause to
inspect them. They are all badly dressed. Their
clothes are torn and shabby; their stockings are full of
holes; and they average about three quarters of a
glove to every hand. But they are all extraordinarily
happy, and not at all shy about showing it. And being
Bill’s younger brothers and sisters, they are as tough
as the proverbial nails.]
Bill
[Emerging from his concealment, looking decidedly
sheepish.]
How did youse get here?
Bessie
We seen yuh go, an’ we run after yuh.
Bill
All of youse?
Bessie
[Nodding.]
Maggie carried de baby.
Maggie