The Orb-Weaving Spiders of Canada and Alaska: Uloboridae, Tetragnathidae, Araneidae, Theridiosomatidae) the Insects and Arachnids of Canada

378 Pages·2001·10.15 MB·English

by  C. D. Dondale

D084_i-vi.qxd 19/03/2003 1:59 PM Page i
THE INSECTS
AND ARACHNIDS
OF CANADA
PART 23
The Orb-Weaving Spiders
of Canada and Alaska
Araneae: Uloboridae, Tetragnathidae,
Araneidae, Theridiosomatidae
Charles D. Dondale and James H. Redner
Eastern Cereal and Oilseed Research Centre
Ottawa, Ontario
Pierre Paquin
Département de sciences biologiques, Université de Montréal
Montréal, Québec
Herbert W. Levi
Museum of Comparative Zoology, Harvard University
Cambridge, Massachusetts
NRC Research Press
Ottawa 2003
D084_i-vi.qxd 19/03/2003 1:59 PM Page ii
© 2003 National Research Council of Canada
All rights reserved. No part of this publication may be reproduced in a retrieval system, or transmit-
ted by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior
written permission of the National Research Council of Canada, Ottawa, Ontario K1A0R6, Canada.
Printed in Canada on acid-free paper.
ISBN 0-660-18898-8 Electronic ISBN 0-660-19260-8
ISSN 0706-7313
NRC No. 44466
National Library of Canada cataloguing in publication data
Dondale, Charles D.
The Orb-Weaving Spiders of Canada and Alaska: Araneae: Uloboridae, Tetragnathidae, Araneidae,
Theridiosomatidae
(The Insects and Arachnids of Canada, 0706-7313; pt. 23)
Co-published by Agriculture and Agri-Food Canada, Research Branch.
Issued by the National Research Council of Canada.
Includes bibliographical references and an index.
ISBN 0-660-18898-8
1. Orb weavers — Canada 2. Orb weavers — Alaska
3. Spiders — Canada 4. Spiders — Alaska
5. Araneidae 6. Tetragnathidae
7. Uloboridae 8. Theridiosomatidae
I. Redner, James H.
II. Canada. Agriculture and Agri-Food Canada. Research Branch.
III. National Research Council Canada.
IV. Title.
V. Series: Insects and Arachnids of Canada.
QL458.42A73D65 2003 595.4’4 C2003-980111-X
NRC Monograph Publishing Program
Editor: P.B. Cavers (University of Western Ontario)
Editorial Board: H. Alper, OC, FRSC(University of Ottawa); G.L. Baskerville, FRSC(University of
British Columbia); W.G.E. Caldwell, OC, FRSC(University of Western Ontario); C.A. Campbell, CM,
SOM(Eastern Cereal and Oilseed Research Centre); S.Gubins (AnnualReviews); B. Ladanyi, FRSC
(École Polytechnique de Montréal); W.H.Lewis (Washington University); A.W. May, OC(Memorial
University of Newfoundland); G.G.E. Scudder, OC, FRSC(University of British Columbia); B.P. Dancik,
Editor-in-Chief, NRC Research Press (University of Alberta)
Inquiries: Monograph Publishing Program, NRC Research Press, National Research Council of
Canada, Ottawa, Ontario K1A0R6, Canada. Web site: www.monographs.nrc.ca
Correct citation for this publication: Dondale, C.D., J.H. Redner, P. Paquin, and H.W. Levi. 2003. The
Insects and Arachnids of Canada. Part 23. The Orb-Weaving Spiders of Canadaand Alaska(Araneae:
Uloboridae, Tetragnathidae, Araneidae, Theridiosomatidae). NRC Research Press, Ottawa, Ontario,
Canada. 371 p.
D084_i-vi.qxd 19/03/2003 1:59 PM Page iii
The Insects and Arachnids of Canada
Part 1. Collecting, Preparing and Preserving Insects, Mites and Spiders, com-
piled by J.E.H. Martin, Biosystematics Research Institute, Ottawa, 1977.
Partie 1. Récolte, préparation et conservation des Insectes, des Acariens et des
Araignées, comilé par J.E. H. Martin, Institut de recherches biosysté-
matiques, Ottawa, 1983.
Part 2. The Bark Beetles of Canada and Alaska (Coleoptera: Scolytidae), by
D.E.Bright, Jr., Biosystematics Research Institute, Ottawa, 1976.
Part 3. The Aradidae of Canada (Hemiptera: Aradidae), by R. Matsuda,
Biosystematics Research Institute, Ottawa, 1977.
Part 4. The Anthocoridae of Canada and Alaska (Heteroptera: Anthocoridae),
by L.A. Kelton, Biosystematics Research Institute, Ottawa, 1978.
Part 5. The Crab Spiders of Canada and Alaska (Araneae: Philodromidae and
Thomisidae), by C.D. Dondale and J.H. Redner, Biosystematics
Research Institute, Ottawa, 1978.
Part 6. The Mosquitoes of Canada (Diptera: Culicidae), by D.M. Wood,
P.T.Dang and R.A. Ellis, Biosystematics Research Institute, Ottawa,
1979.
Part 7. Genera of the Trichoptera of Canada and Adjoining or Adjacent United
States, by F. Schmid, Biosystematics Research Institute, Ottawa, 1998.
Partie 7. Genera des Trichoptères du Canada et des États adjacents, par
F.Schmid, Institut de recherches biosystématiques, Ottawa, 1980.
Part 8. The Plant Bugs of the Prairie Provinces of Canada (Heteroptera: Miri-
dae), by L.A. Kelton, Biosystematics Research Institute, Ottawa, 1980.
Part 9. The Sac Spiders of Canada and Alaska (Araneae: Clubionidae and
Anyphaenidae), by C.D. Dondale and J.H. Redner, Biosystematics
Research Institute, Ottawa, 1982.
Part 10. The Spittlebugs of Canada (Homoptera: Cercopidae), by
K.G.A.Hamilton, Biosystematics Research Institute, Ottawa, 1982.
Part 11. The Genera of Larval Midges of Canada (Diptera: Chironomidae),by
D.R. Oliver and M.E. Roussel, Biosystematics Research Institute,
Ottawa, 1983.
Part 12. The Families and Subfamilies of Canadian Chalcidoid Wasps
(Hymenoptera: Chalcidoidea), by C.M. Yoshimoto, Biosystematics
Research Institute, Ottawa, 1984.
iii
D084_i-vi.qxd 19/03/2003 1:59 PM Page iv
Part 13. Carrion Beetles of Canada and Alaska (Coleoptera: Silphidae and
Agyrtidae),by R.S. Anderson and S.B. Peck, Biosystematics Research
Institute, Ottawa, 1985.
Part 14. The Grasshoppers, Crickets, and Related Insects of Canada and Adja-
cent Regions (Ulonata: Dermaptera, Cheleutoptera, Notoptera, Dic-
tuoptera, Grylloptera, and Orthoptera), by V.R. Vickery and
D.K.McE. Kevan, Biosystematics Research Institute, Ottawa, 1986.
Part 15. The Metallic Wood-Boring Beetles of Canada and Alaska (Coleoptera:
Buprestidae), by D.E. Bright, Biosystematics Research Institute,
Ottawa, 1987.
Part 16. The Horse Flies and Deer Flies of Canada and Alaska (Diptera:
Tabanidae),by H.J. Teskey, Biosystematics Research Institute, Ottawa,
1990.
Part 17. The Wolf Spiders, Nurseryweb Spiders, and Lynx Spiders of Canada
and Alaska (Araneae: Lycosidae, Pisauridae, and Oxyopidae), by
C.D.Dondale and J.H. Redner, Biosystematics Research Centre,
Ottawa, 1990.
Part 18. The Flower Flies of the Subfamily Syrphinae of Canada, Alaska, and
Greenland (Diptera: Syrphidae), by J.R. Vockeroth, Centre for Land
and Biological Resources, Research Branch, Ottawa, 1991.
Part 19. The Ground Spiders of Alaska and Canada (Araneae: Gnaphosidae),
by N.I. Platnick and C.D. Dondale, Centre for Land and Biological
Resources, Research Branch, Ottawa, 1992.
Part 20. The Genera and Subgenera of the Sawflies of Canada and Alaska
(Hymenoptera: Symphyta), by H. Goulet, Centre for Land and Biolog-
ical Resources, Research Branch, Ottawa 1992.
Part 21. The Weevils of Canada and Alaska. Volume 1 (Coleoptera: Cur-
culionoidea, excluding Scolytidae and Curculionidae),by D.E. Bright,
Centre for Land and Biological Resources, Research Branch, Agricul-
ture Canada, Ottawa, 1993.
Part 22. The Genera of the Aphids of Canada (Homoptera: Aphidoidea and
Phylloxeroidea), by R.G. Foottit and W.R. Richards, Research Branch,
Agriculture Canada, Ottawa, 1993.
iv
D084_i-vi.qxd 19/03/2003 1:59 PM Page v
Table of Contents
Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vi
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Definition of orb-weaving spiders . . . . . . . . . . . . . . . . . . . . . . . . 12
Nature of orb webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Ecribellate webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Cribellate webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Production and uses of silk. . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Venoms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Habitats and diets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Family Uloboridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Key to genera of Uloboridae. . . . . . . . . . . . . . . . . . . . . . . . . 34
Genus HyptiotesWalckenaer. . . . . . . . . . . . . . . . . . . . . . . . . 34
Genus UloborusLatreille . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Tetragnathidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Key to genera of Tetragnathidae. . . . . . . . . . . . . . . . . . . . . . . 46
Genus GlenognathaSimon. . . . . . . . . . . . . . . . . . . . . . . . . . 47
Genus LeucaugeWhite. . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Genus TetragnathaLatreille . . . . . . . . . . . . . . . . . . . . . . . . . 53
Genus PachygnathaSundevall. . . . . . . . . . . . . . . . . . . . . . . . 88
Genus MetellinaChamberlin & Ivie. . . . . . . . . . . . . . . . . . . . 107
Genus MetaC.L. Koch . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Family Araneidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Key to genera of Araneidae. . . . . . . . . . . . . . . . . . . . . . . . . 121
Genus MastophoraHolmberg . . . . . . . . . . . . . . . . . . . . . . . 129
Genus MangoraO. Pickard-Cambridge . . . . . . . . . . . . . . . . . . 133
Genus MicrathenaSundevall. . . . . . . . . . . . . . . . . . . . . . . . 141
Genus GeaC.L. Koch. . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Genus ArgiopeAudouin . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Genus CyclosaMenge . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
Genus NeosconaSimon. . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Genus AculepeiraChamberlin & Ivie . . . . . . . . . . . . . . . . . . . 175
Genus Larinioidesdi Caporiacco. . . . . . . . . . . . . . . . . . . . . . 182
Genus AraniellaChamberlin & Ivie . . . . . . . . . . . . . . . . . . . . 191
Genus AraneusClerck . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Genus CercidiaThorell. . . . . . . . . . . . . . . . . . . . . . . . . . . 261
Genus EustalaSimon. . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
Genus SingaC.L. Koch. . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Genus HypsosingaAusserer . . . . . . . . . . . . . . . . . . . . . . . . 280
v
D084_i-vi.qxd 19/03/2003 1:59 PM Page vi
Genus ZygiellaF.O. Pickard-Cambridge. . . . . . . . . . . . . . . . . . 294
Genus AcanthepeiraMarx in Howard . . . . . . . . . . . . . . . . . . . 309
Genus LariniaSimon. . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
Genus MetepeiraF.O. Pickard-Cambridge . . . . . . . . . . . . . . . . 315
Family Theridiosomatidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
Genus TheridiosomaO. Pickard-Cambridge . . . . . . . . . . . . . . . 329
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
Acknowledgments
The authors are grateful to Donald J. Buckle, who contributed many speci-
mens and locality data to this work and also gave a critical evaluation of the manu-
script, and to Nadine Dupérré, who made the 52 original drawings. Raymond
Hutchinson kindly revised the keys. Brent Opell provided a valuable review.
vi
D084_1-32.qxd 14/03/2003 2:58 PM Page 1
Introduction
This manual treats the Canadian orb-weaving spiders and their webs. Previ-
ous parts in the series dealt with crab spiders (Dondale and Redner 1978), sac spi-
ders (Dondale and Redner 1982), wolf, nurseryweb, and lynx spiders (Dondale
and Redner 1989), and ground spiders (Platnick and Dondale 1992). This part is
based on a series of generic revisions by Gertsch (1964), Muma and Gertsch
(1964), Levi (1968-2002), Berman and Levi (1971), and Coddington (1986c). We
also incorporated some locality data of verified specimens from the Canadian
National Collection of Arachnids.
Orb-weaving spiders build the two-dimensional wheel-shaped webs often
seen in hedges, meadows, and weedy waysides. These webs have long inspired
wonder as objects of beauty. They are most noticeable when covered by dew or
fog, and it was such a sight that moved Fabre (1905), the poet-naturalist of
Provence, France, to write: “Considérons, par une matinée brumeuse, le réseau
qui vient d’être construit pendant la nuit. À cause de leur hygrométrie, les gluaux
se sont chargés de gouttelettes et, fléchissant sous le poids, sont devenus autant
de chaînettes, autant de chaplelets de gemmes limpides, gracieux chapelets
rangés en ordre exquis et retombant en courbes d’escarpolette. Si le soleil perce
le brouillard, l’ensemble s’illumine de feux diaprés et devient splendide giran-
dole.” But the orb web, however attractive to the human eye, is really an aerial
trap capable of arresting the flight of an airborne insect and entangling the prey
on the sticky threads of the web. The owner of the web detects the struggles of
the insect, and attacks. Witt (1968) was aware of this scenario when he compared
the nocturnal construction of the web to the preparation of a gallows. Fabre
(1905), too, recognized the true nature of the web: “Il faut manger pour avoir de
la soie, il faut avoir de la soie pour manger, et surtout pour ourdir le dispendieux
cocon de la famille.” Hence the web is a kind of investment in which protein is
expended, and the “pay-off” is food which ideally will not merely replace the pro-
tein used in the web but will provide sufficient energy for the production of the
spider’s offspring. There is solid evidence in orb weavers for a direct relationship
between foraging success and reproductive success (Sherman 1994).
1
D084_1-32.qxd 14/03/2003 2:58 PM Page 2
Methods
The collector can collect individuals of some orb weavers from herbs and
shrubs (Argiopespp., Mangoraspp., Neosconaspp.) with a sweeping net. He can
dislodge those that inhabit tree foliage (Cyclosa spp., Araniella spp., Metellina
spp., Araneus nordmanni, Uloborus glomosus, Hyptiotes spp.) on a sheet held
beneath the branches as the latter are hit with a series of sharp blows from a stick.
He can collect individuals that build on cliffs, among boulders, or on fences and
buildings (Araneus diadematus, Larinioidesspp., Zygiellaspp., Aculepeiraspp.,
Theridiosoma gemmosum) by holding a sheet or net beneath the web and induc-
ing the spider to drop; the specimen can then be steered into a vial. Collecting dif-
ficulties with litter dwellers (Pachygnatha spp., Hypsosinga spp.) can be over-
come with the use of pitfall traps set in the ground (Martin 1977).
Rearing of orb weavers requires much time and patience. Witt and Reed
(1968) attempted to provide natural conditions of light, temperature, and humid-
ity in the laboratory using 16 hours of light daily and varying temperatures and
humidities. Houseflies were used to feed the stock, and the cages were of glass
and screen over aluminum frames. The three species reared by these authors were
Araneus diadematus, Zygiella x-notata, and Larinioides sclopetarius. Manage-
ment of web production, however, was not always successful, particularly in win-
ter.
Web photography is facilitated by the application of a fine mist to the threads
with an atomizer. Alternatively, the photographer can take advantage of a dewy
or foggy day. Or one can make the threads more visible by means of a dusting of
corn starch, in conjunction with a piece of dark material placed behind the web.
Eberhard (1976) gives further methods.
Identification of orb weavers is not easy, and it requires close attention to
anatomical detail. The largest representatives can sometimes be identified by
their size and color, but all identifications should be confirmed by microscopic
examination of the external genitalia (see below). Juveniles should not be named
to species because of the existence of closely related species, distinguishing of
which can only be made with confidence on adult material. Juveniles can, how-
ever, be sometimes reared to maturity (see above). Specimens destined for a col-
lection can be killed in 75% alcohol. Storage is in the same liquid after one or two
changes of alcohol. Leaving the specimens (and alcohol) in a refrigerator for a
few days initially seems to prevent separation of the body wall from the internal
organs (James Berry and Brent Opell, personal communication). The specimens
are stored in vials, with collection data and name of the species, and then placed
in preserving jars fitted with rubber rings. Specimens are examined under a
binocular microscope having magnifications ranging from 10X to 50X. If one
needs to see the spermathecae, he should first cut off the entire epigynum and
immerse it for a few minutes in oil of cloves to clear the structures inside it. After
2
D084_1-32.qxd 14/03/2003 2:58 PM Page 3
it is examined, the epigynum is then stored in a microvial and this in turn in the
larger vial used to hold the rest of the spider’s body. Examination of palpi is made
easy if the examination dish contains a layer of fine sand, fine glass beads, or
black silicon carbide as a cushion. The specimen must be completely immersed
in the alcohol in order to prevent light reflections.
In this manual, the sizes of specimens are indicated as follows: for samples
of 1–3 specimens, the individual measurements are given; for 4–7 specimens, the
mean and range are given; and for 8–10 specimens, the mean and standard devi-
ation are given. The maps include localities from revisions relevant to the United
States of America. Official common names of species are given where available.
3