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GOS

FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

Special Issue: The Orchids in the Ottawa District

Volume 111, Number 1 January—March 1997

The Ottawa Field-Naturalists’ Club

FOUNDED IN 1879

Patron
His Excellency The Right Honourable Roméo LeBlanc, P.C., C.C., C.M.M., C.D.,
Governor General of Canada
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse infor-

mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain-
ing or restoring environments of high quality for living things.

Honorary Members

Edward L. Bousfield Anthony J. Erskine Don E. McAllister Hugh M. Raup
Irwin M. Brodo Clarence Frankton Stewart D. MacDonald Loris S. Russell
William J. Cody W. Earl Godfrey Verna Ross McGiffin Douglas B.O. Savile
Ellaine Dickson C. Stuart Houston Hue N. MacKenzie Pauline Snure
R. Yorke Edwards George F. Ledingham Eugene G. Munroe Mary E. Stuart
Thomas H. Manning Robert W. Nero Sheila Thomson
1997 Council
President: David W. Moore Stephen Bridgett Philip Martin
: : ; Ronald E. Bedford Cendrine Huemer
eS ee ee ee Fenja Brodo Patricia Narraway
, ; Barbara Gaertner Isobel Nicol
Recording Secretary: David Smythe William J. Cody Frank Pope
Corresponding Secretary: Lee Cairnie Francis R. Cook Stan Rosenbaum
Treasuier Ellaine Dickson Chris Traynor
; Alan German Ken Young
Jeff Harrison Eleanor Zurbrig

Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box
P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. For information on Club activities telephone (613) 722-3050.

The Canadian Field-Naturalist

The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal do not necessarily-reflect those of The Ottawa Field-Naturalists’ Club or any other agency.

Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1R0; (613) 269-3211

Copy Editor: Wanda J. Cook

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374

Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1J0

Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, 9302 Edinboro Road, Edmonton, Alberta

T6G 2A1

Associate Editors:
Robert R. Anderson Robert R. Campbell Anthony J. Erskine
Warren B. Ballard Brian W. Coad W. Earl Godfrey
Charles D. Bird Paul M. Catling William O. Pruitt, Jr.

Chairman, Publications Committee: Ronald E. Bedford

All manuscripts intended for publication should be addressed to the Editor with the exception of book reviews
which should go directly to Book Review Editor.

Subscriptions and Membership

Subscription rates for individuals are $23 per calendar year. Libraries and other institutions may subscribe at the rate of
$38 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $23 includes a subscription to The
Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $5.00 to cover postage.
Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to:
The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2.

Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Date of this issue: January-March 1997
(April 1997).

Back Numbers and Index

Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886,
and The Ottawa Naturalist, 1887-1919, and Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa
Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager.

Cover: Ram’s-head Lady’s-slipper (Cypripedium arietinum) is an orchid species with notable colonies in the Ottawa
District. Detail of drawing by Susan Laurie-Bourque. See species account pages 57-60.

THE CANADIAN

FIELD-NATURALIST

Volume 111

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THE OTTAWA FIELD-NATURALISTS’ CLUB

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The Canadian Field-Naturalist

Volume 111, Number 1 January—March 1997

The Orchids in the Ottawa District: Floristics, Phytogeography,
Population Studies and Historical Review

Joyce M. REDDOCH AND ALLAN H. REDDOCH
548 Rivershore Crescent, Gloucester, Ontario K1J 7Y7

Reddoch, Joyce M., and Allan H. Reddoch. 1997. The orchids in the Ottawa District: Floristics, phytogeography, popula-
tion studies and historical review. Canadian Field-Naturalist 111(1): 1-185.

The Ottawa District is the area within 50 km of the Peace Tower of the Parliament Buildings in Ottawa, Ontario, Canada.
Within these parts of eastern Ontario and western Quebec, 44 species of orchids have been recorded since 1856. Of these,
Six species are provincially rare in Ontario and 13 in Quebec. This report is presented as a baseline study from which to
design further research and prepare effective planning measures to protect wild orchid populations. It includes information
on identification, past abundance, population changes, development cycles and relative stability of colonies. The history of
collecting and recording (including the work of the Native Orchid Location Survey), principal orchid habitats, local distri-
bution patterns, rare species, colour forms and other topics are discussed in an introductory section. Detailed information
on these topics is presented for each species, when relevant, together with a brief description of the plant, the overwintering
shoot, capsules and seeds, blooming dates and colony sizes, all based on herbarium specimens, literature and 30 years of
field observations. An illustration and a spot distribution map accompany each species account. Long-lived colonies of
many species are described, and detailed population studies are included for Corallorhiza striata, Goodyera pubescens, G.
tesselata, Platanthera hookeri, P.orbiculata and Spiranthes cernua. The distributions of some species are shown to corre-
late with the Canadian Shield or the St. Lawrence Lowlands, or with calcareous rock, sandstone or sand deposits.

Key Words: orchids, distributions, floristics, historical review, phytogeography, population studies, spirality, Ottawa

District, Ontario, Canada.

Forty-four species of orchids have been found with-
in 50 km of the Parliament Buildings in Ottawa. This
total for such a small area is the consequence of an
interesting geological history and the resulting rich
diversity of habitats. It is also the product of the
coincidental overlap of the ranges of orchids of the
boreal forest, the deciduous forest, the prairie and the
coastal plain on an already rich Great Lakes - St.
Lawrence flora. In addition, human disruption of the
natural landscape over the past two centuries, while
destroying many habitats, has created new ones and
augmented others. Moreover, Ottawa, being the
national capital, has attracted an unusual concentra-
tion of scientifically-trained individuals, who for a
century and a half have studied the local flora.

The Ottawa District

The Ottawa District is half way between the equa-
tor and the north pole. It straddles the Ottawa River,
the north half in Quebec and the south half in
Ontario, and is centred on the cities of Ottawa in

It is our goal that this paper contribute to a better
understanding of floristic biodiversity in this region
and provide information that will permit effective
monitoring and management of wild orchid popula-
tions. We review the knowledge gained since 1856
and, in addition, report on our own observations and
population studies since 1966. The treatment of each
species emphasizes quantitative treatment of heights,
flower numbers, colony sizes and blooming dates, as
well as correlations of distributions with relevant
geological features. We report flower colours using
standardized colour nomenclature and describe habi-
tats, long-lived colonies and early collections. For
some species we report studies of annual population
variation and seasonal development.

Ontario and Hull in Quebec (Maps | and 2). The
Ottawa District is defined as the area within the cir-
cle of 50 km radius centred on the Peace Tower of
the Parliament Buildings in Ottawa (45°25°31°N,

Text and distribution maps: © 1997 Joyce M. Reddoch and Allan H. Reddoch; Line drawings: © 1997 Joyce M. Reddoch, Allan H. Reddoch
and Susan Laurie-Bourque; Other maps: © 1997 Joyce M. Reddoch, Allan H. Reddoch and Marcel Jomphe.

NW

THE CANADIAN FIELD-NATURALIST

Vol. 111

Map 1. Eastern Ontario and Western Quebec showing the Canadian Shield (shaded) and the St. Lawrence
Lowlands (white). The Ottawa District is the area within the circle of 50 km radius centred on the
Peace Tower in Ottawa. This map is adapted from an earlier version prepared by Joyce Reddoch and
Marc Guertin (Reddoch 1983a).

75°42’03”W). It was established by The Ottawa
Field-Naturalists’ Club in 1895 (as a 30-mile circle;
Anonymous 1895) as the common study area for all
fields of natural history. During the century since
then, many significant studies on the flora, fauna and
geology of the District have been published by
Ottawa Field-Naturalists’ Club members, writing in
their capacities as government scientists or as serious
amateurs (Reddoch 1981a, 1981b, 1995).

In the mid 1960s, the Native Orchid Location
Survey (see below) chose an Ottawa Study Area that

History of Orchid Collecting and Recording

The history of orchid studies in the Ottawa
District is essentially that of the general flora, for
only a few people have specialized in the study of
the local orchids. In turn, the history of local botany
is part of the history of Canadian botany, which has
been described recently by Pringle (1995). (The
plant lists mentioned below are summarized in
Appendix 1.)

included the Ottawa District and all of the land within
the rectangle between 45° and 46° N and 75° and
76°30’ W (Map 2; Reddoch 1977b). The Ottawa
Study Area (at 13 104 km?) is about 60% larger than
the Ottawa District (at 7854 km7). Although the pre-
sent work is concentrated on the Ottawa District, it
includes, as well, information about the orchids in the
surrounding Study Area. Although this surrounding
area contributes useful information on habitats and
colonies, no additional taxa except the acyanic form
of Cypripedium arietinum have been found within it.

The earliest travellers, such as Samuel de
Champlain in 1613, who was a good naturalist
(Macnamara 1926), and the collector Philip Whiteside
Maclagan in 1843 (Maclagan 1847; Dore 1983),
mentioned only a few plant species in the District
and did not report any orchids.

The first record of a local orchid is a painting by
Elizabeth Keen White in 1856 of Calypso bulbosa

1997

Stony Swamp
Conservation Area

Marlborough Forest

N
Carleton Rlace

\

76° 30°

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 3

Mer Bleue Bog
Conservation Area 6

-

Casselmam
\ i

\ Prescott (
and Russell 4
A
a c
a
S

As)
eh
s/f) \4
ce y ) Sa

Stormont
Dundas and
Glengarry

Grenville

Kempwille

15°

Map 2. The Ottawa District and surrounding Study Area. The Quebec section of the District is dominated by the
two Outaouais regional municipalities, la Communauté urbaine de |’Outaouais and les Collines-de-
l’Outaouais, while the Ontario part is primarily in the Regional Municipality of Ottawa-Carleton. About a
million people live in the District, three-quarters of them in Ontario.

collected at Ottawa by her husband William White,
later the first president of The Ottawa Field-
Naturalists’ Club (Dore 1965"). Subsequently, she
painted Galearis spectabilis in 1869, Cypripedium
parviflorum in 1870 and Goodyera repens in 1877,
as well as plants in other families. These paintings
were based on local collections that apparently were
not preserved.

The first orchid collections to survive were not
made until two generations after the first settlkements

* An asterisk (*) after a date indicates unpublished item, listed sepa-

rately in Documents Cited section between Acknowledgments
and Literature Cited.

around 1800. Several orchids — Calopogon tubero-
sus, Liparis loeselii, Platanthera dilatata and
Pogonia ophioglossoides — were collected by
Braddish Billings Jr. from Dow’s Swamp near the
Billings home in 1860 (specimens at Queen’s
University (QK)). In 1861, John Kerr McMorine
(Ross 1984) began what was to become a large plant
collection that by 1867 included 13 orchids from
Ramsay, near Almonte, Ontario (QK and DAO; see
Appendix 1). In that year, Braddish Billings Jr.
(1867) published, in the Transactions of the Ottawa
Natural History Society, a list of some 400 plants
that he had collected in 1866 and 1867. This impres-
sive list, almost one third of the Ottawa flora, con-
tained eight orchid species, six of them new, bring-

4 THE CANADIAN FIELD-NATURALIST

ing his total to ten species. Unfortunately, this col-
lection, which James Fletcher (1888) had examined
in the Museum of the Ottawa Literary and Scientific
Society in 1888, has since disappeared.

In 1879, “fully forty gentlemen” founded The
Ottawa Field-Naturalists’ Club with “the express
purpose of working up the natural history of the dis-
trict” (Whyte 1880). Members of the Club, in partic-
ular James Fletcher and Robert B. Whyte, initiated a
period of enthusiastic and productive field work that
was to continue for some thirty years. Among the
botanical collectors were Henry M. Ami, W. Hague
Harrington, William Scott and H. Beaumont Small.
In 1880, Fletcher published his Flora Ottawaensis, a
list of 810 species that he had collected in 1879 with-
in 12 miles of the city, including 25 currently recog-
nized species of orchids. By 1888 he was able to
begin publication of a much enlarged and annotated
flora from within a circle of 30 miles radius. The
number of orchids grew to 31 species (Fletcher
1893). (See Map 3 for early collecting sites.)

Four more species were discovered between 1893
and 1905. Between 1901 and 1911, John Macoun,
who arrived in Ottawa in 1882, completed a
manuscript on the Ottawa flora before moving to the
west coast in 1912; however, it was never published
(Macoun 1901, circa 1911°, 1979).

Starting about 1907, Charles Macnamara of
Arnprior became the first person in the Ottawa
Valley to devote special attention to orchids
(Reddoch 1981c). He discovered and photographed,
but apparently did not collect, 22 species (see
Appendix 1). Much of his orchid work is summa-
rized in an unpublished manuscript (Macnamara
circa 1940°), although he did publish an early note
(Macnamara 1911). (We found nine of his pho-
tographs filed with the orchid collections at the
Smithsonian Institution (US).)

The period from 1912 to 1940 seems to have been
fairly quiet as far as orchid collections are con-
cerned; however, additional contributions were made
by such people as Faith Fyles, Herbert Groh, M. O.
Malte, A. E. Porsild, Frére Rolland-Germain and
Frere Marie-Victorin. Only one new orchid species
was discovered, the recently arrived Epipactis helle-
borine in 1930.

In the 1940s and 1950s, botanists at the
Experimental Farm (Agriculture Canada) began an
intensive program to strengthen the herbarium col-
lections. Orchid sheets of this time bear the names of
A. J. Breitung, J. A. Calder, W. J. Cody, W. G. Dore,
J. M. Gillett, W. H. Minshall, D. B. O. Savile, H. A.
Senn, J. H. Soper and M. N. Zinck. Some of these
botanists were active from as early as the 1930s and
as late as the 1970s and 1980s. Two new orchid
species, Platanthera lacera and the then-undescribed
Spiranthes casei, were discovered in the 1940s.

In 1950, some members of The Ottawa Field-
Naturalists’ Club, feeling the need to revive botani-

Vol. 111

cal field work within the Club, formed the Fern
Group. In 1955, the group wound up its study of
ferns and expanded its activities under the name of
Bog Group. Soon, this name seemed too restrictive
and a new name, Traill Group (after Catharine Parr
Traill, the Canadian pioneer and naturalist), was
adopted along with a stated interest in orchids,
among other subjects (Dill 1982). Orchid collectors
in the 1950s and early 1960s included A. W.
Anderson, I. Bayly, C. and E. Frankton, L. Jenkins
and E. G. Ross.

In 1958, J. M. Gillett prepared the first new check-
list of the Ottawa flora in over half a century. The
total number of orchid species in the District was
then 39 (including Listera australis, which had been
inadvertently omitted from Gillett’s list).

In the early 1960s, the arrival of E. W. Greenwood
brought about the founding of the Club’s Native
Orchid Location Survey, which is discussed below.
In 1977, J. M. Reddoch (1977b) published a list of
the orchids of the District, which by then numbered
42 after the discoveries of Listera auriculata and
Platanthera leucophaea, and with the description of
Spiranthes casei as a new species. The following
year, J. M. Gillett and D. J. White (1978) brought
out a revised checklist of the Ottawa flora containing
the same orchid species. In 1984, M. Runtz pub-
lished an annotated list of the 30 species of orchids
known in the Arnprior area, an increase of eight over
Macnamara’s treatment.

Beginning in the 1970s and continuing to the pre-
sent, general floristic studies have declined in the
District because of decreasing institutional support.
In the 1970s and 1980s, members of the
Conservation Committee of The Ottawa Field-
Naturalists’ Club were prompted by planning exer-
cises at various government levels to identify and
learn more about significant areas. Based on their
research, they wrote briefs and published articles in
The Ottawa Field-Naturalists’ Club’s local journal,
Trail & Landscape. In the 1980s and 1990s, the
National Capital Commission, the Ontario Ministry
of Natural Resources and the Regional Municipality
of Ottawa-Carleton contracted for more intensive
surveying to evaluate candidate natural environment
areas in their respective jurisdictions. These surveys
generated a large number of valuable unpublished
reports; collections made in support of these surveys
were deposited in CAN and DAO. Among the col-
lectors of orchids in the past 25 years are M. L.
Anderson, S. G. Aitkin, D. F. Brunton, S. J.
Darbyshire, A. W. Dugal, D. Gagnon, J. M. and
A. H. Reddoch, M. J. Shchepanek, D. J. White and
A. Vogg.

Now, in 1996, the number of orchids known in the
Ottawa District has risen to 44 with the addition of
two species that have been recognized anew,
Platanthera macrophylla and P. huronensis, in a
flora that totals about 1400 species.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 5

Wakefield ; N ic aicove (North Wakeriigy, 9 Favourite Excursion FV to high Fas
(La Péche) Gracefield and Paugan Falls an d Cc ol lecting Sj tes %
%

D

showing roads and railway lines

Cascades %, used at the turn of the century
Meach's ~~

3 Buckingham

ZG

0 [5 [10 15]
KILOMETRES

41 :
. A East Templeton Station Cumberland
gg. ye fee oe. Kettle Island
ors Kings \ King's Mere %__ lronsides’ Duck Is. 18
me Sim ee 13. @—~ McKay's Lake (Hemlock Lake)
Fairy” — 7% 1214 Green's —_ Blackburn Station
a = 3% Beechwood
lake 4 a4 Creek Nove Leonard

Carlsbad Springs

6 7 wi : — ¢ ?
Britannia Billings (1. Chelsea Grove (Gilmour's Grove) }## silat ace
Bridge . Chelsea Ravine, Old Chelsea Domini pring
. Lake Flora

1
iD

4 t

4. Beaver Meadow (e)

5. Chaudiére Falls Casselman
6

7

8

. Blueberry Point
Black . Central Experimental Farm Pi
. Dow's Swamp
Rapias 9. Billings Bush, Rideau Park ;
10. Patterson's Creek Woods "ig,
11. Stewart's Bush ey

: 12. New Edinburgh, Rideau Hall
: 13. Rockliffe
14. MacKay's Grove an
15. Victoria Sulphur Springs ,

16. Borthwick's Springs

Long 17. Poplar Ridge
_ East Templeton Wharf Russell i

iy
JON PROD py

Map 3. Favourite Excursion and Collecting Sites a century ago. This map is adapted from a 1979 version prepared by Joyce
Reddoch and Marc Guertin (Reddoch 1979b). Dashed lines show roads and lines with hatch marks show railways.

Native Orchid Location Survey Although the benefits of voucher specimens were
The arrival of Ed Greenwood in the Club brought recognized, they decided, in the interests of orchid
a new level of enthusiasm, organization and knowl- conservation and minimization of the effort, not to
edge to the study of Ottawa orchids, which led to the make collections to document the many locations
founding of the Club’s Native Orchid Location recorded, except for new species and special situations.
Survey in 1966. Following the preliminary work of | Taxonomic revisions appeared that split new or rein-
the Traill Group, the Survey set out to locate and _ stated species from the concepts being used by the
map accurately the orchid colonies in the Ottawa Survey. These species, Platanthera grandiflora, P.
District as well as in the rest of Canada (Greenwood huronensis, P. macrophylla and Spiranthes casei,
1967). Their goal was to make an inventory of the were added to the Survey’s list and all records for
species present and their abundances for purposes of them were carefully verified. All of them have
conservation, land-use planning and orchidological proved to be appreciably less abundant than the
studies. After Ed left Ottawa in 1969, the position of | respective species from which they were split: P.
Coordinator of the Survey passed to Hue psycodes, P. hyperborea, P. orbiculata and S.
MacKenzie, then to Bill Petrie in 1975, and finally to cernua. As far as possible, we reexamined sites that
us in 1977. had already been recorded under the latter names to
The members of the Survey were mostly serious determine the correct identities. In some cases, reex-
amateurs who were familiar with the current taxono- amination was no longer possible but it is evident on
my and nomenclature. They reported the location of a statistical basis that, given the relative scarcity of
each orchid colony to within 100 m using the _ the species added to the list, these cases would con-
Universal Transverse Mercator Coordinates from tribute only a very few errors to the distribution
National Topographic Maps. Varieties and forms maps of the more abundant taxa.
were not recorded formally but were discussed The accuracy of the location data also allowed us
among members. to monitor the progress of colonies and to correlate

6 THE CANADIAN FIELD-NATURALIST

plant locations with mapped soil and rock types.
Thus the Survey data can be regarded as comple-
mentary to the herbarium collections in providing
many more colony locations for each species. To
safeguard the colonies, the detailed location data has
been treated as confidential. Copies of the dataset
have been deposited at the National Herbarium of
Canada (CAN) and the Herbarium of Agriculture
Canada (DAO).

After the first decade, the amount of new data
being reported was declining as contributors moved
on to new interests. We have continued to accept
new reports to the present; however, we could not
find the volunteers to carry on the project on a
national scale. From its inception to the present, the
Survey has collected some 3500 sight records for 43
species in the Ottawa Study Area with a total of
12 000 records for all of Canada.

Vol. 111

Those contributing to the Survey in the District
(and Study Area) during the first decade were R.
Anstee, C. Appleton, D. F. Brunton, A. W. Dugal, A.
J. Erskine, C. & E. Frankton, D. Gagnon, E. W.
Greenwood, S. E. Hamill, A. Hanes, J. F. Keddie, J.
D. Lafontaine, R. M. MacDonald, H. N. & E.
MacKenzie, W. Petrie, A. H. & J. M. Reddoch, A. E.
Richards, M. E. Stuart, E. C. D. Todd, D. J. White
and R. E. Whiting. In addition to some of the above,
the following provided records during the past two
decades: B. T. Aniskowicz, E. Beaubien, R. A.
Bracken, P. M. Catling, D. G. Cuddy, E. O. Dodson,
M. Gawn, S. E. Godkin, F. E. Goodspeed, R. Killeen
& S. Kettley, R. A. Layberry, M. H. S. Light & M.
MacConaill, A. Mason, H. McLeod, T. Mosquin, P.
J. Narraway, P. E. Rothschild, M. W. P. Runtz and
A. Vogg.

The Orchids of the Ottawa District (following Dressler (1993))

CYPRIPEDIOIDEAE Lindley

Cypripedium acaule Aiton

Cypripedium arietinum R. Brown
Cypripedium parviflorum Salisbury
var. pubescens (Willdenow) Knight
var. makasin (Farwell) Sheviak
Cypripedium reginae Walter

SPIRANTHOIDEAE Dressler
CRANICHIDEAE Endlicher
GOODYERINAE Klotzsch

Goodyera pubescens (Willdenow) R. Brown in Aiton
Goodyera repens (Linnaeus) R. Brown in Aiton

var. ophioides Fernald
Goodyera tesselata Loddiges

SPIRANTHINAE Lindley

Spiranthes casei Catling & Cruise

var. Casel

Spiranthes cernua (Linnaeus) L. C. M. Richard
Spiranthes lacera (Rafinesque) Rafinesque

var. lacera

Spiranthes lucida (H. H. Eaton) Ames
Spiranthes romanzoffiana Chamisso

ORCHIDOIDEAE
ORCHIDEAE
ORCHIDINAE

Amerorchis rotundifolia (Banks) Hultén
Coeloglossum viride (Linnaeus) Hartman
var. virescens (Muhlenberg) Luer
Galearis spectabilis (Linnaeus) Rafinesque
Platanthera blephariglottis (Willdenow) Lindley
var. blephariglottis
Platanthera clavellata (Michaux) Luer
var. clavellata
Platanthera dilatata (Pursh) Lindley ex Beck

var. dilatata

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

Platanthera flava (Linnaeus) Lindley
var. herbiola (R. Brown) Luer
Platanthera grandiflora (Bigelow) Lindley
Platanthera hookeri (Torrey) Lindley
Platanthera huronensis (Nuttall) Lindley
Platanthera hyperborea (Linnaeus) Lindley
var. hyperborea
Platanthera lacera (Michaux) G. Don in Sweet
var. lacera
Platanthera leucophaea (Nuttall) Lindley
Platanthera macrophylla (Goldie) P. M. Brown
Platanthera obtusata (Banks ex Pursh) Lindley

Platanthera orbiculata (Pursh) Lindley
Platanthera psycodes (Linnaeus) Lindley

EPIDENDROIDEAE Lindley
NEOTTIEAE Lindley
LIMODORINAE Bentham

Epipactis helleborine (Linnaeus) Crantz

LISTERINAE Lindley

Listera auriculata Wiegand

Listera australis Lindley

Listera cordata (Linnaeus) R. Brown

var. cordata

VANILLEAE Blume or other tribe
POGONIINAE Pfitzer

Pogonia ophioglossoides (Linnaeus) Jussieu

MALAXIDEAE Lindley

Liparis loeselii (Linnaeus) L. C. M. Richard
Malaxis monophylla (Linnaeus) Swartz
var. brachypoda (Gray) Morris & Eames

Malaxis unifolia Michaux

CALYPSOEAE Dressler

Aplectrum hyemale (Muhlenberg ex Willdenow) Nuttall
Calypso bulbosa (Linnaeus) Oakes

var. americana (R. Brown) Luer
Corallorhiza maculata (Rafinesque) Rafinesque

var. maculata

var. occidentalis (Lindley) Ames
Corallorhiza striata Lindley
Corallorhiza trifida Chatelain

ARETHUSEAE Lindley
ARETHUSINAE Lindley

Arethusa bulbosa Linnaeus

BLETIINAE Bentham

Calopogon tuberosus (Linnaeus) BSP.

Climate, Geology and Vegetation

Climate

The Ottawa District is a region of warm, moist
summers and cold, snowy winters (data from Crowe
1984). In the centre of the District, the mean daily
maximum and minimum July temperatures are
26.2°C and 14.8°C, while the corresponding January
temperatures are -6.2°C and -15.1°C. Mean annual
precipitation is 843 mm, while the mean precipita-
tion of July alone is 86 mm. The mean frost free

period is 152 days, extending from, on average, 7
May to 7 October.

Although the above data give an average picture of
the climate close to the Ottawa River, the weather is
highly variable from day to day, month to month and
year to year. The climate is influenced also by such
factors as vegetation, elevation, water bodies and
man-made features. On the whole, year-round precip-
itation is greater to the north and less to the south of

8 THE CANADIAN FIELD-NATURALIST

the Ottawa River, while temperatures are somewhat
cooler to the north and warmer to the south.

Geology

The Ottawa District lies in two physiographic
regions, the Canadian Shield and the St. Lawrence
Lowlands (Maps | and 2; account derived mainly
from Bélanger and Harrison (1980) and Chapman
and Putnam (1984)).

The Canadian Shield forms the uplands of the
north half of the District (and Study Area) and the
western part of the Study Area, as well as the Carp
Ridge. The bedrock belongs to the Grenville
Province of the Precambrian Period. It is a complex
of metamorphosed and igneous rocks of which the
most common types are marble, quartzite, gneisses,
granites and syenites. Much of the Shield is hilly
with extensive areas of thin soils and exposed rock.
Between the Ottawa and Gatineau Rivers, Gatineau
Park occupies some of the highest land in the
District, its position accentuated by the 130 to 250 m
high Eardley Escarpment facing south over the
Ottawa River valley. The Eardley Escarpment is the
product of some of the faults that created and shaped
the Lowlands of the Ottawa Valley in Paleozoic
times.

The Lowlands are covered with thick, flat-lying
layers of Paleozoic rock on top of Cambrian sand-
stone, which in turn lies on the basement Precambrian
rocks. The Paleozoic rocks are mainly Ordovician
limestones, dolomites, sandstones and shales.

Glacial deposits from the most recent
(Wisconsinan) ice sheet are scattered across the
District. Following the retreat of the ice sheet about
12 000 B. P., deep layers of clays and sands were
deposited by the Champlain Sea, the marine inunda-
tion that covered the Lowlands, the lower parts of
the Shield and far up the valleys of some northern
rivers, especially the Gatineau. Sand and gravel
beaches were left to mark successively lower water
levels as the sea receded. Subsequent freshwater
deposits of sands and clays were laid down by the
ancestral Ottawa River in a broad swath adjacent to
its present course, especially in the southeast.

Principal Orchid Habitats

The Ottawa District has a great diversity of land
cover types, those of importance to orchids being
forests, wetlands, shores and open areas. General
descriptions of these types are given below; additional
details are contained in the individual species
accounts.

Forests

The forest vegetation in the Ottawa District is
mainly deciduous or mixed with only a small propor-
tion of coniferous forest. The moisture regimes of

Vol. 111

Geological features that are particularly relevant
to the distributions of some orchids in the Ottawa
District are calcareous bedrock (marble and lime-
stone), sandstone bedrock and sand deposits.

Vegetation

The Ottawa District lies in the Great Lakes - St.
Lawrence Forest Region of Rowe (1972). This
region is often combined with Rowe’s Acadian
Forest Region under the designation of Mixed Forest
Region (The Times Atlas of the World (1975)). The
Great Lakes Forest Region is defined by the overlap-
ping ranges of Eastern White Pine (Pinus strobus),
Red Pine (Pinus resinosa), Eastern Hemlock (Tsuga
canadensis) and Yellow Birch (Betula alleghanien-
sis), while the Acadian Region is additionally char-
acterized by Red Spruce (Picea rubens). All but Red
Pine and Red Spruce are common in the District;
these two are known mainly from the Shield, the lat-
ter only rarely. Other species characteristic of the
Mixed Forest Region are also representative of the
(Eastern) Deciduous Forest Region to the south and
the Boreal Forest Region to the north; for example,
Sugar Maple (Acer saccharum), Red Maple (Acer
rubrum), Red Oak (Quercus borealis), Largetooth
Aspen (Populus grandidentata) and White Elm
(Ulmus americana) in the former, and White Spruce
(Picea glauca), Black Spruce (Picea mariana),
Balsam Fir (Abies balsamea), Trembling Aspen
(Populus tremuloides), Balsam Poplar (Populus bal-
samifera) and White Birch (Betula papyrifera) in the
latter.

There is a gradual tendency for the vegetation in
the District to have an increasingly boreal compo-
nent and decreasingly southern component towards
the northern and western reaches; however, the con-
tinuous boreal forest does not begin for another 150
km or so north of the District (Rowe 1972).

Forests cover much of the Shield in the District
but are confined to rocky or difficult locations in the
mainly agricultural and urban Lowlands. According
to Agriculture Canada’s Plant Hardiness Zones in
Canada map (1991), the Lowlands part of the
District is in zone 5a and the Shield is in zone 4b.

forests suitable for most orchids are in the mesic to
wet-mesic range. Much of the forest in the District is
semi-mature (45 - 70 years old). There are only small
pockets of forest over 75 years old.

Most forest-dwelling orchids in the District are
found in mesic deciduous forests of semi-mature,
shade-tolerant hardwoods with a composition
approaching that of the climax deciduous forest in
the region. Sugar Maple is the principal tree species;
it is usually accompanied by a selection of other

1997

deciduous trees including Beech (Fagus grandifo-
lia), White Ash (Fraxinus americana), Hop
Hornbeam (Ostrya virginiana), Basswood (Tilia
americana), Red Maple, Yellow Birch and Black
Cherry (Prunus serotina). These forests are often
described as “rich” because they have a large diver-
sity of plant species including many spring wild-
flowers. This diversity results from the abundance of
humus and nutrients in the soil and the moderate
amount of light still penetrating the tree canopy.
There is even more light in and around forest edges
and clearings that result from windfalls, cutting or
rocky outcrops.

Mixed forests differ from deciduous forests by the
addition of one or more of Eastern Hemlock, Eastern
White Pine, Eastern White Cedar (Thuja
occidentalis), White Spruce and Balsam Fir. Most of
the orchids that grow in the deciduous forest are also
found in the mixed forest. Some species more char-
acteristic of coniferous forests are confined to conif-
erous groves in mixed forests.

Orchids characteristic of deciduous and mixed
forests include Coeloglossum viride, Corallorhiza
maculata, C. striata, C. trifida, Cypripedium parvi-
florum var. pubescens, Epipactis helleborine,
Galearis spectabilis, Goodyera pubescens, G.
repens, G. tesselata, Platanthera hookeri, P. hyper-
borea, P. macrophylla and P. orbiculata. Aplectrum
hyemale is an extirpated species of this habitat.

Moist depressions and stream floodplains in
mixed forests are important habitats for Listera
auriculata, Platanthera grandiflora and P. psycodes.
The Red Maple — Trembling Aspen — Yellow Birch
forests with occasional conifers on the mesic to wet-
mesic sand plains of the southeast are another impor-
tant habitat for Platanthera grandiflora.

Coniferous forests are present in the District either
as separate stands or as groves in otherwise decidu-
ous forests. Forests of Eastern White Cedar, White
Spruce, Eastern White Pine and Balsam Fir are com-
mon on the limestone plains of the Lowlands and on
some parts of the Shield. Sandstone outcrops,
Precambrian ridges and areas of sand are often cov-
ered with forests in which Eastern White Pine is
dominant. The floors of coniferous forests are thick-
ly carpeted with needles and are almost devoid of
plants except for Wild Lily-of-the-valley (Maianthe-
mum canadense), a few ferns and club-mosses, and
certain orchids, including Corallorhiza maculata, C.
trifida, Cypripedium acaule, C. arietinum, Goodyera
tesselata and Malaxis unifolia.

Pine plantations on deep sand shelter a number of
orchids. The Larose Forest, established in 1928 with
extensive plantings of Red Pine, White Pine and
White Spruce (Reid 1979), supports some large
colonies of Cypripedium acaule, Malaxis unifolia
and Spiranthes lacera, as well as a few Corallorhiza
trifida and S. romanzoffiana. In addition, Goodyera

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 9

tesselata and Platanthera hookeri have appeared in
similar plantations elsewhere in the District.

Wetlands

The three wetland classes that are important habitats
for orchids in the Ottawa District are swamps, fens
and bogs. About 60% of the local species grow in
wetlands. The Canadian Wetland Classification
System (Tarnocai 1988) defines these wetlands as
follows:

“A swamp is a mineral wetland or a peatland with
standing water or water gently flowing through pools
or channels. The water table is usually at or near the
surface. There is pronounced internal water move-
ment from the margin or other mineral sources;
hence the waters are rich in nutrients. If peat is pre-
sent, it is mainly well-decomposed wood, underlain
at times by sedge peat. ... The vegetation is charac-
terized by a dense cover of deciduous or coniferous
trees or shrubs, herbs, and some mosses.”

‘A fen is a peatland with the water table usually at or
just above the surface. The waters are mainly nutri-
ent-rich and minerotrophic from mineral soils. The
dominant materials are moderately to well decom-
posed sedge and/or brown moss peat of variable
thickness. ... The vegetation consists predominantly
of sedges, grasses, reeds and brown mosses with
some shrubs and, at times, a sparse tree layer.”

“A bog is a peatland, generally with the water table
at or near the surface. The bog surface, which may
be raised or level with the surrounding terrain, is vir-
tually unaffected by the nutrient-rich groundwaters
from the surrounding mineral soils and is thus gener-
ally acid and low in nutrients. The dominant materi-
als are weakly to moderately decomposed Sphagnum
and woody peat, underlain at times by sedge peat. ...
Bogs may be treed or treeless, and they are usually
covered with Sphagnum spp. and ericaceous shrubs.”

Swamps are scattered across the District, but they
are especially abundant on the Lowlands. The most
productive swamps from the point of view of orchids
are calcareous and are characterized by various com-
binations of the following trees: Eastern White
Cedar, Black Ash (Fraxinus nigra), Red Maple,
Yellow Birch, White Spruce, Tamarack (Larix lari-
cina) and Balsam Fir. The mesic to wet swamp
floors are partially to heavily shaded; they are car-
peted with rich layers of mosses interspersed with
bare patches covered with fallen leaves. Cinnamon
Fern (Osmunda cinnamomea), Foamflower (Tiarella
cordifolia), Wild Sarsaparilla (Aralia nudicaulis),
One-flowered Wintergreen (Moneses uniflora),
Shinleaf (Pyrola elliptica) and Twinflower (Linnaea
borealis) are some typical swamp plants.
Characteristic orchids are Corallorhiza trifida,
Cypripedium parviflorum, C. reginae, Liparis loe-
selii, Malaxis monophylla, M. unifolia, Platanthera

10 THE CANADIAN FIELD-NATURALIST

clavellata, P. hyperborea, P. obtusata and P. psy-
codes. Corallorhiza striata, Cypripedium acaule, C.
arietinum, Goodyera pubescens, G. repens, G. tesse-
lata, Listera cordata, Platanthera huronensis and P.
orbiculata are occasionally found. Calypso bulbosa
is a swamp-dweller that is rare in the District.
Liparis loeselii, Malaxis monophylla and
Platanthera clavellata are infrequent inhabitants of
alder-willow swamps.

Fens are relatively uncommon in the Ottawa
District; they are usually components of wetland
complexes that include swamps and occasionally
marshes (Reddoch 1983b). Calcareous sedge fens
are the most significant fen type in the District. They
are dominated by sedges (Carex lasiocarpa and
sometimes C. livida) and typical fen mosses.
Tamarack, Eastern White Cedar and Black Spruce
occur in scattered clumps in some sedge fens and
provide the tree layer in treed fens. (Treed fens are
not well represented in the District; some sedge fens
are rimmed by treed fen habitat.) The sedge fens are
intermediate to rich fens with pHs in the range 6.0 to
7.0. (Poor fens (pH 4.6 - 5.5) are included under
bogs below.) Six sedge fens rich in orchids have
been studied in some detail (Reddoch 1979a, 1984,
1989). Four of the fens are underlain by limestone
on the Lowlands in Ontario and two are underlain by
marble on the Canadian Shield, one in Quebec and
the other at the western edge of the Study Area in
Ontario. See Appendix 2 for more details and a list
of the plants recorded in these fens.

Two other important peatlands that contain
swamp and fen habitats are the Leitrim Wetlands on
the Lowlands (Dugal 1990, 1992, 1993) and
Chilcott’s Swamp on the Shield. Chilcott’s Swamp,
on the edge of Johnston Lake in Quebec, harbours at
least 14 species of orchids. It was first explored by
naturalists in 1892 (Whyte, Craig, and Cowley 1893)
and has been visited from time to time since then
(Fyles 1912; Anderson 1959; André Sabourin, per-
sonal communication 1992).

Dow’s Swamp and Lake Flora are two Lowlands
swamps, no longer extant, that likely contained
pond-edge fens. Dow’s Swamp (Reddoch 1978b;
IlIlman 1980) was a peatland south of Dow’s Lake in
a depression that drained to the Rideau River, and
Lake Flora was a 4-ha pond on I’Ile de Hull (Nagy
1974) that is now Parc Fontaine. Early naturalists
recorded 13 orchids from Dow’s Swamp and six
from Lake Flora.

True bogs are uncommon in the District. The
largest is the 25 km* Mer Bleue Bog, which, like
Alfred Bog east of the District (Cuddy 1983), devel-
oped in undrained sections of channels abandoned
by an ancestral Ottawa River. The Mer Bleue is a
domed bog with peat up to 4 m deep. It is a mosaic
of treed bog, shrub bog, poor fen and marsh, the
result of natural development as well as of some
impact from fires, logging, drainage projects, prac-

Vol. 111

tice bombing and beaver dams. Black Spruce and
Tamarack are the typical trees, with Grey Birch
(Betula populifolia) in some open areas; Leatherleaf
(Chamaedaphne calyculata) and Labrador Tea
(Ledum groenlandicum) are the common shrubs.
The Mer Bleue is the first place in Canada where
Listera australis was found. This orchid is a species
of the poor fen habitat, as is Platanthera blephar-
iglottis, the other rare orchid in the Mer Bleue. In
1995 the peatland was accepted as the 33rd
Canadian wetland in the Ramsar Convention on
Wetlands of International Importance.

There are a number of small bogs and poor fens in
the District. The bogs have developed in small
depressions, while the poor fens usually take the
form of sedge mats at the edges of lakes and ponds.
Cypripedium acaule is the characteristic orchid of
the true bog habitat; Calopogon tuberosus and
Pogonia ophioglossoides are typical poor fen
species, while Arethusa bulbosa occurs infrequently
in this latter habitat.

Shores

Flooding, wave action and ice scraping along
lakes and rivers maintain habitats suitable for a num-
ber of species, particularly Platanthera flava, P.
grandiflora, P. psycodes, Spiranthes cernua and S.
lucida. These orchids thrive in the moist alluvial
soils with minimal competition. The Platantheras
also grow in adjacent riparian forests of Red Maple,
Silver Maple (Acer saccharinum), Red Ash
(Fraxinus pennsylvanica) and other trees.

Other Open Areas

Expanses of exposed bedrock maintain suitable
habitats for a number of orchids that grow in partial
to full sun and can tolerate some dryness.
Cypripedium arietinum, C. parviflorum var.
pubescens and Spiranthes lacera are characteristic
species of alvars, areas of thin soil over flat-lying
limestone and marble (Catling and Brownell 1995).
Over acidic rocks, Cypripedium acaule, Malaxis uni-
folia and Spiranthes lacera occur on Precambrian
knolls and are joined by Spiranthes casei and S. cer-
nua in areas of flat-lying, exposed sandstone. These
naturally-occurring openings are sometimes enlarged
by fires and blow-downs.

Ditches, borrow pits and abandoned fields are
human creations that in the past century have pro-
vided new, although often transient, habitats for
orchids that flourish in moist, exposed sites. Such
habitats are colonized, sometimes in large numbers,
by Liparis loeselii, Platanthera lacera, P. psycodes
and Spiranthes cernua. Other species occasionally
occurring are Malaxis unifolia, Platanthera clavel-
lata, P. grandiflora and S. casei (rarely). Mowing,
if done at the right time, maintains the habitat.
Sandy stretches beside bogs have supported, in
addition, Calopogon tuberosus and Pogonia
ophioglossoides.

1997

Local Distribution Patterns

Of the 44 species of Ottawa District orchids, all
but five have been recorded on both the Canadian
Shield and the St. Lawrence Lowlands within the
District. The five exceptions are species that are rare
or extirpated in the District: Listera auriculata and
Platanthera macrophylla (on the Shield) and
Aplectrum hyemale, Listera australis and
Platanthera leucophaea (on the Lowlands). Eight
other orchids occur predominantly on the Canadian
Shield, while seven additional species are found
mostly on the Lowlands.

The distributions of some orchids in the Ottawa
District correlate with one or more of calcareous
bedrock (marble and limestone), sandstone bedrock
and sand deposits.

Marble is exposed or close to the surface especially
in the western half of the Shield north of the Ottawa
River and in the west of the Study Area, while lime-
stone is close to or at the surface in the Lowlands on
the limestone plain in the southwest and in various
other locations. Amerorchis rotundifolia, Arethusa

Continental Distribution Patterns

Continental distribution patterns of Ottawa
District orchids fit roughly into the following North
American biomes: Montane, Boreal, Mixed,
(Eastern) Deciduous and Southeastern Coastal Plain
Forest Regions, and Prairie. These biomes are
mapped by Rowe (1972) for Canada and by Sheviak
(1983) for the United States, with some inconsisten-
cies between them.

The Ottawa District is in the Mixed Forest Region
and, as might be expected, the majority of orchids
(29 of 44) recorded here have their primary distribu-
tion in this Region.

Southern Affinities

A further five species are distributed both in the
Mixed Forest Region and in the Deciduous Forest
Region to the south. Four other species are predomi-
nantly inhabitants of the Deciduous Forest Region
and they are near or at their northern limits here:
Aplectrum hyemale, Galearis spectabilis,
Platanthera flava var. herbiola and Spiranthes cer-
nua. The distributions of four species, Calopogon
tuberosus, Corallorhiza maculata, C. striata and
Malaxis unifolia, reach various parts of Mexico,
Central America and the West Indies (Luer 1975;
Homoya 1993).

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT ies

bulbosa, Calypso bulbosa, Corallorhiza striata,
Cypripedium arietinum, C. parviflorum, C. reginae,
Epipactis helleborine, Malaxis monophylla,
Platanthera dilatata, P. flava, P. huronensis, P. leu-
cophaea and Spiranthes lucida have distributions that
correlate with the occurrence of calcareous rock.

Thinly covered or exposed sandstone bedrock is
present at some borders of the Canadian Shield, such
as in the Stony Swamp Conservation Area, and also
along a portion of the Gloucester Fault near South
Gloucester. The distribution of Spiranthes casei cor-
relates with the occurrence of sandstone on the
Lowlands.

Important sand deposits occur in several areas of
the Shield (Hoffman, Miller, and Wicklund 1967;
Lajoie 1962, 1967; Bélanger and Harrison 1980) and
on the Lowlands in the southeast (Wicklund and
Richards 1962; Bélanger and Harrison 1980). Liparis
loeselii, Malaxis unifolia, Platanthera grandiflora,
P. lacera and Spiranthes cernua occur mainly on
sand in the Ottawa District.

Northern Affinities

Ten species of the Mixed Forest Region also occur
in the Boreal Forest Region to the north. Four addi-
tional orchids, Amerorchis rotundifolia, Calypso bul-
bosa var. americana, Listera auriculata and Plat-
anthera obtusata, are mainly boreal species and they
are near or at the southern edges of their distribu-
tions in the District.

Prairie and Coastal Affinities

Platanthera leucophaea and Spiranthes cernua
are two species that also occur in the Prairie Region,
while Listera australis is a representative of the
Southeastern Coastal Plain Forest Region. Calo-
pogon tuberosus, Pogonia ophioglossoides and Spi-
ranthes cernua are other local species with important
presences in the latter region.

European and Asian Affinities

Ten native Ottawa District species, most with
northern affinities, also occur in Europe and/or Asia
as the same or a different variety: Calypso bulbosa,
Coeloglossum viride, Corallorhiza trifida, Goodyera
repens, Liparis loeselii, Listera cordata, Malaxis
monophylla, Platanthera hyperborea, P. obtusata
and Spiranthes romanzoffiana (Luer 1975). In addi-
tion, Epipactis helleborine is a relatively recent
arrival from Europe.

12 THE CANADIAN FIELD-NATURALIST

Rare Species

Of the 44 orchid species that have occurred in the
Ottawa District, six have been designated as rare in
Ontario, 13 as rare in Quebec and two as rare in
Canada by The Rare and Endangered Plants Project
of the Botany Division of the National Museum of
Natural Sciences (Canadian Museum of Nature).

Ottawa District orchids designated rare in Ontario
(Argus and White 1982) are Aplectrum hyemale,
Listera australis, Platanthera blephariglottis, P.
grandiflora, P. leacophaea and P. macrophylla.

Ottawa District orchids designated rare in Quebec
(Bouchard et al. 1983) are Aplectrum hyemale,
Arethusa bulbosa, Corallorhiza striata, Cypripedium
arietinum, C. reginae, Galearis spectabilis, Good-
yera pubescens, Listera australis, Platanthera ble-
phariglottis, P. flava, P. macrophylla, Spiranthes
casei and S. lucida.

Ottawa District orchids designated rare in Canada
(Argus and Pryer 1990) are Aplectrum hyemale and
Platanthera leucophaea.

These lists have provided a basis for the two
provinces, Ontario and Quebec, to begin the process
of determining what species of plants will be consid-
ered for legal protection. This process is on-going,
but at present there is no specific legal protection for
any of the District’s vascular plants or their habitats

The Need For Protection and Management

Orchids are one of the components of global biodi-
versity that are threatened by human activities. Loss
of natural areas containing orchids causes the extirpa-
tion of populations and, bit by bit, could lead to the
extinction of these species. To counteract these loss-
es, we must ensure that large natural areas represent-
ing all significant local ecosystems are protected.

This paper contains many references to losses of
orchid populations. Large scale human disturbance
and destruction of the natural environment began in
the last years of the 18th century with the establish-
ment of the settlements beside the Ottawa River that
grew into the cities of Ottawa and Hull, and by log-
ging of the forests followed by conversion of suit-
able lands to agricultural use.

Most orchids have survived so far through the ran-
dom accidents of being in places not yet accessible
to or economically viable for destructive develop-
ment. The establishment of large publicly-owned
areas such as the Marlborough Forest, the Greenbelt
and Gatineau Park has played a significant role in
protecting some orchid populations. Although these
areas were not set up primarily for the maintenance
of biodiversity, their establishment was an essential
first step in that direction. If that goal is to be
achieved, it is imperative that protection of these
reserves continue and be improved.

Vol. 111

in either province. The species listed above for
Quebec are recognized as “espéces de la flore vascu-
laire menacées ou vulnérables susceptibles d’étre
ainsi [par la loi] désignées” (Gazette officielle du
Québec 1993; Brouillet 1994). Lists compiled by the
Natural Heritage Information Centre of the Ontario
Ministry of Natural Resources (Oldham 1996*) are
released from time to time with further refinements
of the status of species in Ontario.

Platanthera leucophaea is assigned the status of
vulnerable in the 1996 list of The Committee on the
Status of Endangered Wildlife in Canada
(COSEWIC). The current Government of Ontario
has removed from consideration those species desig-
nated as vulnerable.

In the Ottawa District, all plants are protected on
the property of the National Capital Commission
(notably Gatineau Park and the Greenbelt) by the
National Capital Act (Regulation #26).

Orchids that are rare or possibly extirpated in the
Ottawa District are Amerorchis rotundifolia,
Aplectrum hyemale, Calypso bulbosa, Listera auric-
ulata, L. australis, Platanthera blephariglottis, P.
leucophaea and P. macrophylla. All but P. macro-
phylla are at or near the limits of their distributions
here.

Current threats to orchids on private rural land
include ongoing urban expansion, estate lot develop-
ment (particularly in woodlands), limestone quarry-
ing (including in alvars), logging, drainage of wet-
lands in anticipation of development and flooding of
wetlands as a byproduct of building highways and
access roads. There has also been some deliberate
destruction of habitats by owners who do not want
their hopes for development impeded by rare plants.
Throughout the District, predation by White-tailed
Deer (Odocoileus virginianus) and disruption of hab-
itats by Beavers (Castor canadensis) have had some
impact on orchid populations. A more serious threat
is the encroachment of invasive plants on some of
the area’s most important habitats: wetlands are
threatened by Glossy Buckthorn (Rhamnus frangula)
and shorelines by Purple Loosestrife (Lythrum sali-
caria).

Because the interrelationships of the components
of ecosystems are complex, the best way to protect
orchids is to protect whole ecosystems. This requires
the establishment of areas large enough to ensure
that colonies lost to the natural processes of succes-
sion, fire and changes in drainage will be replaced in
other suitable habitats nearby.

Management specifically for orchids usually will
involve measures to maintain the habitat against

1997

natural succession and water level disruptions.
Managers will need as much information as possible
on the life cycle of the orchids and on their habitat
requirements. In addition, close monitoring of the
colony and its habitat will be required along with
cautious experimentation with remedial procedures
when problems are detected and before the colony
becomes too depleted. A remarkable local example
of personal initiative in approaching such problems
is Joe Purdon’s 50-year management of a fen to
enhance and maintain a population of Cypripedium
reginae against encroachment by Eastern White
Cedar and flooding by Beavers. This task has now
been taken over by the Mississippi Valley
Conservation Authority.

Colour Forms and Their Nomenclature

As with other characters, the intensities and hues
of flower colours exhibit ranges, sometimes quite
broad, that are characteristic of the species. Outside
these normal ranges, more dramatic variations arise
from specific genetic mutations. Some mutations
disrupt biosynthetic pathways so that the plant
becomes unable to produce an otherwise typical pig-
ment. Other mutations interfere with the pattern con-
trol mechanisms that determine the distribution of
pigment on the flower with the result that colour will
appear in atypical parts of the flower or disappear
from typical parts. A number of these variations
have been found in the District (A. H. Reddoch and
J. M. Reddoch 1987b).

It will be useful to review the general mechanisms
responsible for these forms in terms of flower pig-
ments and their colours. Most flower pigments
belong to one of three classes of chemical com-
pounds (Harborne 1982; Griesbach 1983; Harborne
and Turner 1984; Arditti 1992). Colours ranging
from purple through magenta to pink are usually pro-
duced by anthocyanins. Green is usually the result of
chlorophyll, the primary photosynthetic agent of the
leaves. Yellow may arise from a variety of classes of
compounds, including anthocyanins, but most often
from carotenoids. Carotenoids frequently occur with
chlorophyll and also have a photosynthetic function.
The presence of combinations of these pigments in a
given flower can be demonstrated readily by simple
chromatographic methods.

Within the typical colour range of a species, minor
and sometimes major variation in intensity occurs.
This is particularly noticeable for pink-flowered
species such as Arethusa bulbosa, Cypripedium
acaule and Platanthera grandiflora, but it can also
be seen in some green or yellow flowers.

When two or more classes of pigments are present,
intermediate colours arise that are characteristic of a
given species. These colours can vary in hue as well

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 13

Regardless of the nature of the management, there
will be a need for continuing monitoring by dedicat-
ed individuals. It is our experience that organizations
often have poor corporate memories; the original
information and objectives are sometimes lost.
Sometimes instructions and precautions do not sur-
vive the chain of command from the conservation
office to the operator of the cutter. The Ottawa Field-
Naturalists’ Club could take a role in such monitor-
ing, although, ultimately, it will be enthusiastic and
knowledgeable individuals who must do the job. It is
encouraging that agencies on both sides of the
Ottawa River employ competent and dedicated biol-
ogists to lay the groundwork for more enlightened
conservation measures.

as in intensity from plant to plant with variations in
the concentrations of these pigments. The resulting
colours can be understood by the relatively complex
theory of subtractive colour mixing (Billmeyer and
Saltzman 1981), but it will be adequate to follow the
artists’ experience of the mixing of pigment colours.
Mixing yellow and green produces a range of interme-
diate colours that occur in many of our greenish
orchids. Yellow and magenta can yield brownish hues
such as are seen in Corallorhiza maculata and some-
times in the sepals of Cypripedium acaule.

When equivalent amounts of the approximately
complementary chlorophyll green and anthocyanin
magenta are mixed, they yield a dark greyed colour of
an intermediate hue. When the magenta dominates the
mixture, the result is a darkened and greyed purplish
colour. Such a colour can be found on the lower part
of the lip of Cypripedium arietinum and frequently on
the petals and sepals of C. acaule. If green is the dom-
inant component, then the mixture will be a darker
green with a reddish cast such as is sometimes found
on the lower leaves and stem of Epipactis helleborine.

Absence of anthocyanin

This aberration is the most striking and familiar of
the colour forms in the Ottawa District. Floral antho-
cyanin is absent occasionally in a wide range of plant
species as a result of genetic mutations that disrupt the
biosynthetic pathway for production of this pigment
(Griesbach 1985). If no other pigments are normally
present, then the result is a pure white, such as the lip
of Cypripedium acaule f. albiflorum and the flowers
of Platanthera psycodes f. albiflora. However, when a
yellow pigment is also present, the loss of antho-
cyanin results in the yellow form seen in Corallorhiza
maculata f. flavida. When both yellow and green pig-
ments are also present, usually in moderate concentra-
tions, they, together with the anthocyanin, combine to
produce brown or greyish magenta. The absence of
the anthocyanin then reveals a yellow-green colour as

14 THE CANADIAN FIELD-NATURALIST

in the sepals and petals of Corallorhiza trifida var.
verna, Cypripedium acaule f. albiflorum and C. ariet-
inum f. albiflorum.

This mutation causes the complete absence of
anthocyanin so that the flower is pure white in the
absence of other pigments or shows the colours of
any other pigments present. A flower with a very
small amount of anthocyanin may appear nearly
white but evidently the plant’s biosynthetic pathway
for this pigment is intact. The pale colour lies near
the limit of the normal range permitted by the control
function. Thus the plant is not an albino. For exam-
ple, plants of Platanthera grandiflora and P. psy-
codes with flowers so pale that the pink colour can be
verified only by the boundary between the outer part
of the lip and the pure white base are not f. albiflora.

It is known that some orchid species contain more
than one anthocyanin pigment in their flowers
(Arditti 1992). In such cases more complicated
colour variations may arise if genetic mutations
occur that affect the formation of only one of the
pigments. It should be fairly straightforward to ana-
lyze the resulting colour changes.

Absence of chlorophyll

The absence of chlorophyll is another genetic
defect, but is much rarer than the absence of antho-
cyanin mainly because most plants cannot develop
to maturity without this essential photosynthetic
agent (Furman and Trappe 1971). A few species,
such as those of Monotropa and, among our orchids,
of Corallorhiza, can develop normally with the aid
of their associated mycorrhizae. In the District, two
orchids normally bearing chlorophyll, Epipactis
helleborine and Platanthera hyperborea, are known
to have produced a few achlorophyllous plants. The
colour of the plants was generally white although in
some cases some small amounts of yellow could be
seen in buds, leaves and stems. In E. helleborine,
anthocyanins could be seen in the lip and the base of
the stem, as is sometimes the case for normal plants.

Pattern variation

The occurrence of well-defined, reproducible
genetic changes in the colour patterns of flowers is
familiar in such well known plants as Snapdragon

Blooming Periods

Table 1 summarizes the blooming dates of the
orchids in the Ottawa District and Study Area. A
related table was published previously (A. H.
Reddoch and J. M. Reddoch 1987e). Similar charts
for other areas in the northeast include those of
Brown (1985) for the Bruce Peninsula, Ontario,
Whiting and Catling (1986) for Ontario, Keenan
(1987) for New England, Smith (1993) for
Minnesota, and Homoya (1993) for Indiana.

Apparently split blooming periods of
Corallorhiza maculata, Platanthera dilatata and P.

Vol. 111

(Antirrhinum majus) and Petunia (Petunia hybrida)
and has proved useful in the study of gene expres-
sion (Fincham 1987; Jorgensen 1995). Among the
orchids of the Ottawa District, Galearis spectabilis f.
willeyi appears to show this effect. In this form, the
magenta pigment of the sepals and petals extends to
a greater or lesser extent over the normally white lip.

Nomenclature

Many colour variations have been given formal
names, usually at the forma level. However, it is
questionable whether these names are really a good
method of communication. In the cases of distinc-
tive, genetically based variations, it is usually possi-
ble to apply the names to well-defined entities.
Nevertheless, it would be useful to refer to these
plants by the mechanism involved, for example,
acyanic, achlorophyllous or pattern variants, rather
than by often obscure honorific or somewhat random
descriptive names.

It is even more questionable to apply formal
names to variations in hue or intensity that are
within the normal variation of these characters.
Even colours near the limits of the range are part of
the normal expression of genetic or environmental
conditions of the species. There is no natural
boundary to separate such plants from their more
typical relatives. Hence the creation of a variety or
form for such plants requires an arbitrary designa-
tion of a boundary, a process that makes the variety
or form itself arbitrary. There is the additional com-
plication, in the case of colours, that the usual ver-
bal descriptions of colour are not sufficiently
unambiguous to define clearly such a boundary. A
single type specimen could not designate a bound-
ary even if its colour were stable. Formal names in
these situations suggest a level of precision that is
illusory; it would be more appropriate for most pur-
poses simply to provide a concise, precise descrip-
tion. In this work, we generally do not use names
that have been applied to plants with flower colours
within the normal ranges for their species. In
describing the flower colours of each species in the
District, we include the full normal range and dis-
cuss special cases separately.

hyperborea (sensu lato) are shown for the Bruce.
From our data for C. maculata in the Ottawa
District, it seems likely that the split is related to
the early and late blooming varieties, var. occiden-
talis and var. maculata respectively, although our
overall set of dates for the District does not show
the effect so clearly. On the other hand it seems
unlikely that the split for P. hyperborea can be
explained by the inclusion of P. huronensis, which
have essentially the same blooming period in this
region.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 15

Blooming Periods of Ottawa District Orchids

SPECIES MAY JUN JUL AUG SEP OCT ft

Amerorchis rotundifolia onl

Arethusa bulbosa —-

Calopogon tuberosus ————

Calypso bulbosa —

Coeloglossum viride ——

Corallorhiza maculata —

Corallorhiza striata —_———

Corallorhiza trifida —— 55

Cypripedium acaule ———

Cypripedium arietinum ————

Cypripedium parviflorum ——

Cypripedium reginae —_——

Epipactis helleborine ——

Galearis spectabilis “a

Goodyera pubescens a

Goodyera repens ———

Goodyera tesselata ———

Liparis loeselii a. eee

Listera auriculata a

Listera australis a

Listera cordata —————

Malaxis monophylla —_——

Malaxis unifolia ———

Platanthera blephariglottis =

Platanthera clavellata a

Platanthera dilatata —

Platanthera flava soo ERIE = 16

Platanthera grandiflora ———

Platanthera hookeri ——

Platanthera huronensis ——

Platanthera hyperborea oe 2 ae

Platanthera lacera i

Platanthera leucophaea —

Platanthera macrophylla ee

Platanthera obtusata ——

Platanthera orbiculata ee a

Platanthera psycodes serene OTR UTE ecm
omen MEY reins

Pogonia ophioglossoides 79
Spiranthes casei oxo 42
Spiranthes cernua Sia 3-1, aad 125
Spiranthes lacera aco MEI cress 82
Spiranthes lucida ea 19
Spiranthes romanzoffiana cra = 33

MAY ci JUN uinedUl os wAUG tr SEP). OCT.

TABLE |. All data came from the Ottawa District except those for Listera australis and Platanthera macrophylla, which
were supplemented by data from adjacent parts of eastern Ontario and western Quebec. The light bar shows the range and
the heavy bar shows one standard deviation about the mean.

16 THE CANADIAN FIELD-NATURALIST

Capsules and Seeds

Capsules

Capsules of some Ottawa District orchids collect-
ed after seed release are illustrated in Figures la - d.
These species were chosen to complement those
shown by Homoya (1993) for Indiana, which are
also representative of plants in the District except for
Spiranthes lacera. The specimen of Platanthera
hyperborea illustrated by Homoya is comparable to
the mesic forest plant of the Ottawa District.

Sizes

Among species, the smallest capsules are those of
Listera cordata at 0.3 x 0.2 cm and the largest are
those of Cypripedium reginae at 3.6 x 1.2 cm.

Colours

Generally, at the time of seed release, capsule
colours are in the range of grey-brown through light
to dark brown. Sometimes there is a tendency to
blackening. At the same stage, Epipactis helleborine
capsules may occasionally still be partially green. A
few species have quite light-coloured capsules:
Liparis loeselii, Malaxis monophylla, M. unifolia
and, most notably, Listera cordata. The latter species
is also notable for its early seed release, which can
occur even when the flower is still fresh-looking and
the capsule is still green.

Orientation

The capsules of most species are erect or ascend-
ing, but those of the Corallorhizas and sometimes of
Epipactis helleborine are pendent. Capsule orienta-
tion appears to be determined by gravity rather than
by the orientation of the inflorescence axis. We have
seen examples in the field of Pogonia ophioglos-
soides and Platanthera macrophylla where the stem
had been accidentally leaning at an angle during and
after capsule development. In these cases, the cap-
sule orientation was distinctly vertical rather than
erect (parallel to the stem). Presumably this can hap-
pen in other species also.

For most species, the dried-up remnants of the
flower remain attached to the end of the capsule,
sometimes conspicuously so as in the larger
Cypripediums. In Calopogon tuberosus and
Arethusa bulbosa, however, the column is the most
distinctive remnant. Often for Pogonia ophioglos-
soides, the entire flower disappears, apparently by
dehiscence.

Capsule Yields

The yield (the fraction of flowers producing cap-
sules) for individual plants tends to be highly vari-
able and the data somewhat limited, but patterns
emerge for some species. The uncommon orchid
Arethusa bulbosa has very low yields, of the order
of a few percent. At the other end of the scale,
Spiranthes cernua and S. casei, both reported to be

Volwidd

autogamous (Catling and Catling 1991), regularly
approach 100% yields. A third group of species
tends to have rather random yields but with an
appreciable number of plants approaching 100%
yields. This group includes Coeloglossum viride,
Epipactis helleborine, Platanthera clavellata and
P. hyperborea, all of which are reported to be auto-
gamous (Catling 1983a; Catling and Catling 1991),
at least in some parts of their ranges. Corallorhiza
maculata and Spiranthes lacera usually have
yields over 50%, the first of these also being
reported as being autogamous (Catling 1983a;
Catling and Catling 1991). Goodyera pubescens
seems unique in having either very high or very
low yields.

For Arethusa bulbosa and Listera cordata our lim-
ited results are similar to those reported elsewhere;
however, for Calopogon tuberosus, Cypripedium
acaule, Galearis spectabilis and Pogonia ophioglos-
soides, our estimates of Ottawa District yields are
appreciably higher.

Seeds

Seeds have much the same range of colours as the
capsules and there is some rough correlation between
the colours of the seeds and of the capsules they
come from. The darkest seeds tend to occur in some
of the Platanthera species, particularly in the section
Blephariglottis. The seed studies of Arditti,
Michaud, and Healey (1979, 1980) and Healey,
Michaud, and Arditti (1980) show a level of colour
variability comparable to what we find for capsule
colours and, with more limited data, for seeds. Their
seed colours are similar to ours in some cases, in oth-
ers they are somewhat darker. Some of these differ-
ences could reflect terminological differences
between their “subjective” colour names and the
Methuen Handbook of Colour names (Kornerup and
Wanscher 1978) used here. We find a moderate level
of correlation between our colour observations and
the levels of pigmentation reported by Stoutamire
(1983).

For most species, seed release begins between
September and mid October, after the capsules have
changed colour. Exceptional species that release
seeds earlier are Listera cordata, Epipactis helle-
borine, Platanthera dilatata and Spiranthes lacera.
The late blooming S. casei and S. cernua tend to
release their seed in the latter half of October.
These observations are consistent with those of
Stoutamire (1964) for species in the upper Great
Lakes region.

Knowledge of seed release times may be useful
for management purposes, for example, to avoid
mowing before most of the seeds have been
released.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT ty

Sy, i
Wiel (Has 2

FiGuRE la. Capsules after seed dispersal (left to right, upper row): Amerorchis rotundifolia, Corallorhiza striata and C. trifi-
da; (lower row): Cypripedium arietinum, Goodyera repens var. ophioides and G. tesselata; approximately life size.

18 THE CANADIAN FIELD-NATURALIST Voli he)

FiGuRE Ib. Capsules after seed dispersal (left to ri ght): Listera cordata var. cordata, Malaxis monophylla var. brachypoda,
Platanthera blephariglottis var. blephariglottis and P. dilatata var. dilatata, approximately life size.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

FiGuRE Ic. Capsules after seed dispersal (left to right): Platanthera grandiflora, P. huronensis and P. leucophaea
mately life size.

, approxi-

20 THE CANADIAN FIELD-NATURALIST Vol. 111

d\

(i |
Sle ; AW S

FiGuRE Id. Capsules after seed dispersal (left to right): Platanthera macrophylla (scale bar = 5 cm), P. obtusata, Spiranthes
casei and S. lacera var. lacera, the latter three approximately life size.

1997

Overwintering States

Orchids in the Ottawa District are dormant for half
the year, thus it is worthwhile to describe the state in
which they spend half their time. Most District
orchid plants re-create themselves every year.
Notable exceptions are the Cypripediums, which rise
from annual extensions of long-lived perennial rhi-
zomes, the Goodyeras, which produce flowering
stems from creeping rhizomes on a cycle several
years long, and the Corallorhizas, which produce
flowering stems annually or at intervals of several
years from coralloid root systems.

Most species with fibrous, fleshy or tuberous roots
begin producing the shoots for next year’s plants at
flowering time or even before. Shoots develop at the
bases of the current year’s stems or a few centimetres
away on horizontal roots or other structures. Except
for some late-blooming Spiranthes, new roots also

Longevity of Colonies

Some orchid plants and their colonies are short-
lived even in habitats that appear not to change.
Other orchid colonies are transient because their spe-
cialized habitats change. However, most species that
live in stable environments may persist many
decades, either because individual plants are long-
lived or because reproduction is consistent and abun-

Variability of Characters

All characters show a range of sizes or values
reflecting genetic or environmental influences. The
ratio of the standard deviation to the average pro-
vides a convenient measure of such variation. For
heights, this ratio is between 18% and 33% for most
of our species. However, Platanthera hyperborea is
notable for its ratio of 50%. Liparis loeselii and
Epipactis helleborine are the next most variable
species with ratios of 36% and 37% respectively. For
the first two species much of the variation may be

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 4) |

form and begin to elongate before dormancy. Most
plants overwinter with the green shoots projecting just
above the ground surface beside the brown remains of
the current year’s stems, while the shoots on some
species remain just below the ground or moss surface
until spring. Currah, Smreciu, and Hambleton (1990)
illustrate the seasonal development of mycorrhizal
roots and tubers of Coeloglossum viride, Platanthera
hyperborea (sensu lato), P. obtusata and P. orbiculata
in Alberta. Although plants produce new shoots, these
shoots may die in the spring, followed during the sum-
mer by the roots.

In orchids growing from corms, the new corms
begin to develop in the spring and mature during the
summer. These orchids overwinter as the mature
corms, from which they produce new stems the fol-
lowing year.

dant. In fact, the three decades during which we
have monitored colonies can be only a short segment
of the life of some orchid colonies.

In forests, colonies may be almost as old as their
forest habitats, typically to about 65 - 70 years in the
District, whereas in stable peatlands, colonies may
be several hundred years old.

attributed to adaptation to different habitats, which is
discussed in the individual species accounts follow-
ing. For E. helleborine no specific cause is evident,
although the species is often regarded as being quite
variable.

For the number of flowers, these three species
again show great variability with ratios of 69%, 61%
and 67% respectively. However the largest ratio,
70%, comes from Platanthera huronensis. For other
orchid species, the ratio runs from 25% to 56%.

pd THE CANADIAN FIELD-NATURALIST

Methodology

The species descriptions are based on our observa-
tions made in the field in the Ottawa District and the
surrounding Study Area since 1966 and on herbari-
um specimens from the same areas examined at the
following institutions (listed in Index Herbariorum
Part I (Eighth Edition)): Canadian Museum of
Nature (CAN), Carleton University (CCO),
Agriculture Canada (DAO), Field Museum of
Natural History (F), Royal Botanical Gardens
(HAM), Université de Montréal (MT), McGill
University (MTMG), U.S. National Arboretum
(NA), Canadian Forestry Service (OTF; now incor-
porated in CAN), Queen’s University at Kingston
(QK), Royal Ontario Museum (TRT) and
Smithsonian Institution (US). In a few cases infor-
mation from nearby surrounding regions is included
and identified as such. The descriptions also include
information from the following publications of The
Ottawa Field-Naturalists’ Club: Transactions of the
Ottawa Field-Naturalists’ Club (1879 - 1887), The
Ottawa Naturalist (1887 - 1919), The Canadian
Field-Naturalist (1919 - present) and Trail &
Landscape (1967 - present), as well as Transactions
of the Ottawa Natural History Society (1867 - 1869).

Our orchid specimens and photographs of rare
taxa made during the course of this study have been
deposited in CAN and DAO; mosses have been
deposited in CANM.

ENGLISH NAMES: These names come mainly from
Whiting and Catling (1986). We rejected names that
gave erroneous impressions, for example, Tall White
Bog-orchid for Platanthera dilatata because this
species does not grow in bogs, and any name for P.
macrophylla that implies, incorrectly, that this
species has exceptionally large leaves or leaves larg-
er than P. orbiculata.

FRENCH NAMES: We have taken the French names
from Sabourin (1993), Argus and White (1982) and
Fleurbec (1994°), transferring names from Habénaire
to Platanthére where necessary. For Platanthera
macrophylla we propose a name related to the English
name that satisfies the objectives stated above.

SYNONYMS: Names in this section are those used in
the (mostly older) Ottawa District and other litera-
ture and are not a complete synonymy. Some author-
ities cited in the older literature have been amended
to current usage.

INTRODUCTION: Each species is introduced with a
comment on notable features followed by field charac-
ters that distinguish it from other species in the same
genus or from similar orchids in the same habitat.

PLANT DESCRIPTIONS: Plant descriptions are con-
fined to heights, flower numbers and colours, num-
bers of leaves, overwintering states, capsules and

Vol. 111

seeds. These characters were chosen because they
allowed a more quantitative account or were other-
wise of interest to us. Other aspects are well covered
by recent publications (for example: Smith 1993;
Homoya 1993).

Heights: The heights are based on our measure-
ments of herbarium specimens and plants in the
field. The full range is reported along with the range
including one standard deviation about the mean.
See Numerical Results below.

Flowers: The numbers of flowers were obtained
from herbarium specimens and our field counts. For
multiflowered species the full range is reported
along with the range including one standard devia-
tion about the mean. (See Numerical Results below.)

To have a standardized terminology for the colour
names of flowers, leaves, capsules and seeds, we
have used those in the Methuen Handbook of Colour
(Kornerup and Wanscher 1978). This book has a rea-
sonably convenient format for field work and, with
1266 colours, it permits fairly subtle distinctions for
reporting colour variation through the use of its
alphanumeric notation. In this work, we are con-
cerned with broader ranges and hence report the
colours by the more general (and more familiar)
English names given in the Handbook. (Two areas
where these names are too broad are the rather
extensive ranges described as greyish green and
greyish magenta.) Standardized nomenclature is
especially helpful for colours in the pink-purple-rose
group and for reporting the darker greens of the
leaves of the Goodyeras. Although Goodyera leaves,
as well as those of Cypripedium arietinum, have
sometimes been described as bluish green, no exam-
ples that we have seen conform to the use of that
term in the Handbook, the Centroid colour system
(National Bureau of Standards s.d.; Kelly and Judd
(1976)) or The RHS Colour Chart (1966). Greyish
green or dark green are the Handbook names that we
have found appropriate.

The sense of smell seems to be both more com-
plex and less understood than the sense of colour
vision. Orchid fragrances may be the product of as
many as 50 different compounds in a wide range of
concentrations (Arditti 1992) while human sensitivi-
ty is extremely variable in its ability to detect these
compounds. The intensity and composition of these
fragrances, which presumably are used primarily to
attract and perhaps to guide insect pollinators, can be
strongly dependent on the time of day and probably
other factors such as temperature. Hence it is diffi-
cult to achieve precise and concise descriptions of
such fragrances.

The fragrances reported here are our subjective
description of our field experience. Joyce is notably

1997

more sensitive than Allan, who is consistently
unable to detect odours from some species such as
Corallorhiza maculata. Such failures are not includ-
ed in the accounts. Joyce’s mother, Florence
Dunston, provided an expert second opinion on
occasion.

Leaf or Leaves: The numbers of leaves of flowering
plants were obtained from herbarium specimens and
our field counts. For multi-leaved species the maxi-
mum range observed is reported. When this range
seems unusually large, we also report a smaller,
more typical range. Counts for non-flowering plants
are reported separately when available. In some
cases, other leaf characteristics such as orientation
and colour are recorded.

Overwintering State: We report the dormant condi-
tion of the shoot or bud as observed above ground or
within the moss substrate in the autumn, having
taken care not to injure the plants. We obtained some
evidence of the condition at anthesis from herbarium
specimens.

Capsules: We describe the condition of the capsules
in the autumn.

Sizes: We measured small samples of capsules for
each species and cite the averages as a general
impression of size.

Yields: We report yields derived from relatively
small samples taken on a casual basis at various
locations over the past 15 years. For single-flowered
plants the yield is the ratio of the number of capsules
to the number of flowers in the sample and for multi-
flowered plants it is the average of such ratios for the
individual plants in the sample. Where several sites
were studied, sometimes over several years, it was
possible to report on the variability of the yields.

Seeds: The colours of the seeds represent a small
sampling which may not adequately reveal the range
of variation in some species. The colours describe a
small pile of seeds rather than a single seed. Under a
microscope, it can be seen that the colour is some-
times determined by the embryo and sometimes by
the testa or a part thereof.

We derived the first dates of seed release from our
records of dates when one or more capsules were
observed to be open in various colonies and years.
These individual dates for each species are spread
over a period of a week to 10 days and hence, in the
species accounts, are summarized as being in the
early, mid or late part of the month. If our records
did not also include dates when no capsules were
open, then we report that capsules were open “by” a
particular time. Presumably various factors including
weather, habitat, genetics and the frequency of our
visits contribute to the spread of our data.

BLOOMING PERIOD: These results are based on
herbarium collections and our field records. The

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 23

only literature reference is for Arethusa bulbosa and
Listera australis, for which the early date of 28 May
reported by Fletcher (1896) is included. The results
are summarized in Table 1. The same ranges are
given in the species accounts. See Numerical Results
below.

CoLony SIZEs: These results are from the Native
Orchid Location Survey and our field records. A
colony is taken to include all plants seen at a given
UTM coordinate, that is, in an area 100 m by 100 m.
The minimum (usually one) and maximum numbers
of plants are given and, in some cases, a smaller, more
typical upper limit. See Numerical Results below.

CURRENT STATUS: This topic is included when a
species is rare in the Ottawa District, Quebec,
Ontario or Canada, or is very abundant. Gillett and
White (1978) included the status of Ottawa District
species based on herbarium specimens.

DISTRIBUTION: General North American distribu-
tions are included to give impressions of where the
Ottawa District lies within them. They come mainly
from Homoya (1993) but also from Smith (1993),
Luer (1975) and specialist papers cited in individual
species accounts. Distribution patterns in the District
are summarized briefly with regard to the Canadian
Shield and the St. Lawrence Lowlands and with
emphasis on significant correlations with relevant
bedrock and surficial geological features.

DISTRIBUTION Maps: The maps include all plottable
records from as early as 1860 to the end of the 1996
field season (up to 137 years). Thus they show the
cumulative occurrences of each species and not nec-
essarily its current status.

The distribution data derive from three sources:
herbarium specimens, additional sight records from
the Native Orchid Location Survey (1965 - 1996)
and literature references in the Transactions of the
Ottawa Field-Naturalists’ Club, The Ottawa
Naturalist and the typescripts of John Macoun (circa
1911°) and Charles Macnamara (circa 1940’).
Locations cited on herbarium specimens except
those from the last couple of decades are often
described rather generally and therefore cannot be
plotted very accurately. The many early specimens
from locations cited only as “Ottawa”, “Hull”,
“swamp, Ottawa’, “environs d’Ottawa” and similar
forms could not be plotted at all. By contrast, the
3500 records generated by the Native Orchid
Location Survey give the location of each colony to
within 100 m.

Major occurrences of calcareous bedrock (marble
on the Shield and limestone on the Lowlands), sand-
stone bedrock (Nepean and March Formations) and
sand deposits on the Lowlands are shown for those
species where there are strong correlations with
orchid occurrences. Although limestone is
widespread on the Lowlands, it is deeply covered by

24 THE CANADIAN FIELD-NATURALIST

overburden in many places, especially in the south-
east. The Canadian Shield is shown on most maps.
The information is derived from a number of local
geological maps (Geological Survey of Canada
1912, 1973, 1974, 1977; Wicklund and Richards
1962; Hogarth 1970; Bélanger and Harrison 1980).

The distribution maps were initiated by Ed
Greenwood and were updated by successive Native
Orchid Location Survey coordinators; Joyce
Reddoch and Marc Guertin produced the base map
and the geological overlays.

HABITATS: The descriptions are specific to the
Ottawa District and are derived from our own obser-
vations and from herbarium labels. Most names of
vascular plants and the order in which they are listed
are from Gillett and White (1978).

LONG-LIVED COLONIES: Under this heading we
report our observations of colonies that we have
monitored for from one to three decades. These
treatments contribute additional details of habitats
for specific sites.

EARLY History: The first known collection and
other early collections are emphasized; the complete
known history is related for recently-discovered
species. John Kerr McMorine’s collections, Braddish
Billings Jr.’s list (1867), James Fletcher’s lists
(1880, 1893), John Macoun’s list (circa 1911") and
Charles Macnamara’s illustrated typescript (circa
1940°) are referred to where relevant. (See also
Appendix 1.)

SPECIAL Topics: Topics treated include accounts of
colour forms, morphological variation with habitat,
aberrations, exceptional plants, seasonal develop-
ment, vegetative reproduction, dermatitis and further
diagnostic notes.

FIGURES: The illustrations were prepared by Susan
Laurie-Bourque from our photographs and drawings,

Voli 111

and from fresh flowers. They show living plants that
are typical of this area. All of the plants depicted
grew in the District or Study Area except Amer-
orchis rotundifolia (Frontenac County), Calypso bul-
bosa (Frontenac County) and Listera australis
(Alfred Bog). Habitat, location and date are given
for each subject.

NUMERICAL RESULTS: Dates and measurements are
reported in the form a (b - c) d, where a and d are the
upper and lower limits and b and c represent one
standard deviation below and above the average,
respectively. While various authors report only a and
d, these values refer only to the most unusual mem-
bers of the species. The range b - c describes the
more normal plants and, for a normal distribution,
includes 68% of the population. Some caution is
required in the more detailed application of the stan-
dard deviation because the data for blooming dates,
flower counts and plant heights frequently deviate
from the normal distribution. These deviations often
reflect a slight excess of late dates or large heights or
flower counts. In the case of the heights and counts,
such asymmetric distributions are not surprising for
quantities that have large fractional variation but
cannot be negative.

By contrast distributions of colony sizes are not
even approximately normal because most species
have colonies containing only a few plants while large
colonies can occur but are infrequent. For this reason
the full range is given and, when 15 or more records
are available, a statement “typically to” is included
indicating an upper limit that includes 95% of the
colonies. This value was obtained simply by exclud-
ing 5% of the colonies at the upper end of the range.

In Spiranthes cernua, where some results are
given mainly for purposes of comparison, an average
value is given followed by the standard deviation in
brackets.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 25

Amerorchis rotundifolia (Banks) Hultén
Small Round-leaved Orchis Orchis a feuille ronde
SYNONYMS: Orchis rotundifolia Banks, Habenaria rotundifolia (Banks) Richardson

Amerorchis rotundifolia is an attractive but rare fen orchid, apparently approaching extirpation in the District.

The only currently known colony produces few and often no flowering plants. This species can be recognized
by its single, basal, elliptical, somewhat pointed leaf and by its irregular inflorescence of purple and white flow-

ers with three-lobed lips.

DESCRIPTION

Height: 9 (15 - 24) 26 cm [25 plants].

Flowers: | (3 - 9) 17 [29 plants]; dorsal sepal pale
violet outside, white with similarly coloured mark-
ings inside; lateral sepals white, sometimes with a
pale violet stripe on both surfaces; petals purple, or
greyish magenta or light lilac edged with purple; lip
white with purple, purplish red or deep magenta
spots or larger markings (see Colour Pattern Variant
below) and yellowish green or paler colour at
entrance to spur; spur white; column similar in
colour to petals; ovary light green with colour simi-
lar to dorsal sepal distally as stripes or solid colour;
in a loose, irregular, somewhat one-sided inflores-
cence; no fragrance detected.

46°

Leaf: 1; 1 on non-flowering plants.

Overwintering State: a whitish green shoot about
1 cm long, within the moss substrate, rising from a
horizontal rhizome several centimetres away from
the current year’s stem, by early fall; herbarium
specimens show the new shoot and partially elongat-
ed roots present at anthesis.

Capsules: dark brown, narrowly ellipsoid, typi-
cally 1.2 x 0.4 cm, erect (see Figure 1a); yield highly
variable, averaging 45% [32 plants].

Seeds: brown, released in late September.

BLOOMING PERIOD: 10 June (19 June - 5 July) 12
July [12 records].

45°
76° 30

75°

Amerorchis rotundifolia: 1 = herbarium specimen. Major areas underlain by calcareous rock

(marble and limestone) are shaded.

26 THE CANADIAN FIELD-NATURALIST Voli 111

FIGURE 2. Amerorchis rotundifolia, treed fen habitat, Palmerston Township, Frontenac County, Ontario, | July 1972 (plant
and flowers), 19 June 1996 (column); for the column (front view), scale bar = 0.5 mm.

1997

CoLony SIZE: 0 - 12 flowering plants, between
1967 and the present, at the only extant colony in
the District; as loose groups of flowering and
non-flowering plants in larger colonies in Eastern
Ontario.

CURRENT STATUS: rare and local in the Ottawa
District; one colony known to be extant, but close to
extirpation.

DISTRIBUTION: The Ottawa District is near the
southern edge of the distribution of this orchid of the
Boreal Forest Region (Catling 1983b). Within the
District, only three calcareous fens are known to
have harboured this orchid (J. M. Reddoch and A. H.
Reddoch 1987a). The nearest known colonies out-
side the District are 100 km southwest of Ottawa in
Frontenac County and 100 km northwest near
Westmeath in Renfrew County. This species is con-
fined to treed and open fens in regions of calcareous
bedrock.

HABITATS: This orchid grows in fairly open treed
fens, open sedge fens and some swamps associated
with these fens.

In treed fens and swamps, Black Spruce,
Tamarack, Eastern White Cedar and Balsam Fir are
the dominant trees; saplings of the latter two species
constitute the understory. Labrador Tea (Ledum
groenlandicum) is the most common shrub, and
Carex paupercula, C. stricta, C. trisperma,
Eriophorum viride-carinatum, Three-leaved False
Solomon’s Seal (Smilacina trifolia), Cypripedium
parviflorum, C. reginae, Platanthera huronensis,
Pink Pyrola (Pyrola asarifolia) and Twinflower
(Linnaea borealis) are common herbaceous plants.
The peatland floors are extensively covered with
mosses, especially Sphagna. Within these treed
habitats, A. rotundifolia plants, flowering and non-
flowering, are grouped in well-defined open areas
where the canopy is thin or somewhat open and the
shrub layer is absent. They are usually growing in
mounds of wet Sphagna, including S. russowii (pH
6.3 in the treed fen near Poltimore (J. M. Reddoch
and A. H. Reddoch 1987a)). In 1996, we recorded a
temperature of 14.5C at the root level at anthesis in
the colonies near Poltimore and in Frontenac
County.

In a swamp dominated by Eastern White Cedar
near Westmeath, the herbaceous layer also included
Shining Clubmoss (Lycopodium lucidulum), Dwarf
Scouring-rush (Equisetum scirpoides), Rattlesnake
Fern (Botrychium virginianum), Carex leptalea,
Goldthread (Coptis trifolia), Viola spp., Spikenard
(Aralia racemosa), One-flowered Wintergreen
(Moneses uniflora) and Canada Fly-honeysuckle
(Lonicera canadensis), as well as the additional
orchids Corallorhiza trifida, Liparis loeselii, Listera
cordata, Malaxis monophylla and Platanthera
obtusata.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT og,

In sedge fens, plants of A. rotundifolia are scat-
tered across the open fen, both on mounds of fen
mosses and in the wet peat of the fen floor.

LONG-LIVED COLONIES: Ed Greenwood discovered
the only currently extant colony near Poltimore in
1966. The 12 plants were situated in a swamp habitat
that was near the outside edge of a peatland com-
posed mainly of a treed fen/sedge fen complex. From
time to time, naturalists continued to find a few plants
here and there in this peatland, usually in the treed
fen close to the sedge fen opening. In 1970 and 1971,
there were about a dozen plants scattered on the open
fen floor; they could not be found there again in 1972
(J. Donald Lafontaine, Sheila and Harry Thomson,
personal communications) or in any subsequent year.
Logging of the treed fen and swamp between 1983
and 1985 resulted in a vastly changed habitat and it
was not until 1989 that a few flowering plants were
encountered again, in a small cluster of uncut trees.
There were four flowering plants in 1994, 1995 and
1996. In 1996, there were two additional flowering
plants nearby in the sedge fen growing among flow-
ering plants of Arethusa bulbosa.

The colony in Frontenac County was flourishing
in 1959 when Otto Devitt (1961) described the habi-
tat as “rather open”. From 1969 until the early
1980s, we recorded many hundreds of flowering
plants in and around the frequent clearings. By 1996
the trees had grown considerably but there were
numerous A. rotundifolia still thriving in the remain-
ing openings. One or two to two dozen (rarely three
dozen) flowering plants were present in various
openings depending on their size.

The colony in Renfrew County near Westmeath
numbered close to 200 plants when Hue and Elva
MacKenzie searched the area in 1968 (records of the
Native Orchid Location Survey, personal communi-
cations). We found a similar number there in 1984,
but in 1989 we could find only a few flowering
plants.

These patterns of fluctuating numbers are similar
to Fred Case’s observations in Michigan (Case
LOS):

EARLY History: Dow’s Swamp was the first place
in the District where A. rotundifolia was discovered.
The earliest collection was probably the one by R. B.
Whyte on 16 June 1878 cited by John Macoun in his
undated typescript of about 1911°. The whereabouts
of that collection is unknown. Extant herbarium
specimens record James Fletcher’s collections there
in July of 1878, as well as in 1879 and 1882 [DAO
17027 and 17029, MTMG 47597, TRT 15747, US
27620]. Amerorchis rotundifolia occurred at Dow’s
Swamp “in large numbers” along with the similarly
abundant orchids, Cypripedium arietinum and
Malaxis monophylla (Whyte and Small 1883).
Dow’s Swamp was a calcareous treed fen and pond-
edge open fen located southeast of Dow’s Lake. It

28 THE CANADIAN FIELD-NATURALIST

lingered into the middle of the 20th century and was
better known in its latter days as a good birding spot
(Reddoch 1978b; Iman 1980).

In 1903, John Macoun (circa 1911*) encountered
A. rotundifolia “in the cedar swamp east of Stittsville
and a little north of the railway” and made several
collections [CAN 117000, DAO 17028, MTMG
8602]. This swamp, which was visited by naturalists
a number of times over the years (see, for example,
Eddy and Halkett (1913)), was destroyed in 1980 in
an expansion of the Amberwood subdivision. In that
year, we found Corallorhiza trifida, Cypripedium
arietinum, Cypripedium reginae, Malaxis monophyl-
la and Platanthera hyperborea, as well as Epipactis
helleborine, but no A. rotundifolia, in the rapidly
draining swamp.

COLOUR PATTERN VARIANT: Amerorchis rotundifo-
lia f. lineata (Mousley) Hultén was discovered at the
Frontenac County colony in 1959 (Devitt 1961). The
form differs from the typical by having two broad
purplish stripes on the lip (Mousley 1941).

This description is, however, an oversimplifica-
tion that does not reflect the highly variable pattern

Vol. 111

of lip coloration of the species. In this form, the
genetic control of the colour pattern formation
appears not to be very rigorous. In the typical form,
the small dots, while of fairly uniform size, are quite
irregular in their arrangement from one flower to
another in the same inflorescence and are also some-
what variable in size from plant to plant. The ideal-
ized f. lineata is only one aspect of the observed pat-
terns. These include spots that are fewer and larger
than typical, as well as mixtures of large spots on
one side of the lip and a single stripe on the other,
stripes broken into two segments and even a single
patch of colour covering the entire centre of the lip.
Usually, no two flowers on the same inflorescence
have the same patterns. However, the aberrant pat-
terns do not seem to coexist with the typical small
spots on the same plant.

The plants with aberrant lip patterns constituted
about 15% of the Frontenac County population in
1996. Contrary to the observations of Whiting and
Catling (1986), we found that the aberrant-lipped
plants grew among plants with typical lips, not in
separate groups.

[997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 29

Aplectrum hyemale (Muhlenberg ex Willdenow) Nuttall

Putty-root

Aplectrelle d’hiver

Aplectrum hyemale was reported in the District in the 1880s and in 1905, but no subsequent records are known.
Such erratic sightings seem to be characteristic of the species in this part of its range. Unfortunately, the only
reported collection can not be found, but the accounts of the discovery of the first colony are convincing enough
to accept A. hyemale as a historic member of the Ottawa flora.

CURRENT STATUS: rare in the Provinces of Quebec
(Bouchard et al. 1983) and Ontario (White et al.
1982a), very rare (S2) in Ontario (Active List,
Oldham 1996"), rare in Canada (Argus and Pryer
1990); apparently extirpated in the Ottawa District
(no plants reported since 1905).

DISTRIBUTION: The Ottawa District is at the northern
edge of the distribution of this orchid of the
Deciduous Forest Region.

History: Aplectrum hyemale was added to the
Flora Ottawaensis following its discovery in 1885
by Lieutenant-Colonel William White, the first pres-
ident of The Ottawa Field-Naturalists’ Club (Whyte,
Macoun, and Small 1887). The location was cited as
“McKay’s wood” (Fletcher 1887) or “Beechwood”
(Fletcher, Small, and Baptie 1887b, 1888; Fletcher

46°

1893). “The woods beyond Rideau Hall [were] pop-
ularly known as McKay’s Bush or Beechwood”
according to an anonymous account of Club excur-
sions in 1887. This area, stretching eastward from
Rideau Hall to present-day St. Laurent Blvd., was a
rich and diverse natural area frequented by natural-
ists for more than 70 years (Reddoch 1979b).

On 15 October 1887, Lieutenant-Colonel White
led a group of botanists to the site of his discovery
(Fletcher, Small, and Baptie 1887b). “After a short
search beneath fallen leaves Mr. Fred Magee suc-
ceeded in finding a fine patch of three plants — two
of these bore two leaves each.” This event was
described again by the same authors (1888) in their
Report of the Botanical Branch for the season of
1887. Here they related that the intent of the ex-
cursion was to collect “roots of the rare orchid

45°

76° 30

ra

Aplectrum hyemale: & = literature reference. The Canadian Shield is shaded.

30 THE CANADIAN FIELD-NATURALIST

Aplectrum hyemale” and that “a fine patch of seven
roots was found by Mr. Fred Magee, three of which
were removed for study under cultivation”.

There is no indication that a herbarium specimen
was prepared. The description of the plants in the
1887 account was minimal, saying, in addition to the
quotation above about the leaves, that, in reference
to the corms on two of the plants, “there are two of
these bulbs on a plant”. The account also included a
general description of the seasonal development of
the species that showed that the authors were well
informed on the appearance of the leaves (“a large
handsome ribbed leaf ... dark green above and purple
beneath”) and corms. These descriptions and the
presence of the leaves in mid-October make this a
credible report. In the 1970s, we and other botanists
(Darbyshire 1982) searched in vain for A. hyemale in
the remnant forests in Beechwood Cemetery and
across Hemlock Road on the east side of McKay
Lake, now mostly destroyed (Darbyshire 1981).

John Macoun (circa 1911°), who was familiar
with this orchid from other places in southern
Ontario (Macoun 1888), listed his own collection of
13 September 1905 from “cool rich woods Billing’s

Vol. 111

[sic] bush, Rideau park (15 plants)”. This location
likely was on the Billings property east of Billings
Bridge (Belden 1879). We have not been able to
locate this collection in any of the herbaria that we
visited or to which we wrote (British Museum
(Natural History) (BM), University of Notre Dame
(NDG), New York Botanical Garden (NY),
University of Calgary (UAC)). Macoun’s collections
are held in many herbaria throughout North America
and Europe.

Three other colonies have been reported some 50
to 80 km beyond the District. At Oka, Quebec, about
100 plants were studied from 1930 to 1942, but no
plants have been found there more recently (Lavoie
1994). In 1964, G. N. Gogo collected a plant [DAO
s.n.] north of Summerstown, Charlottenburg
Township, Glengarry County, Ontario. In May of
1988, Don Cuddy (personal communication) discov-
ered 15 plants in Frontenac Provincial Park,
Frontenac County, Ontario, while leading a Field
Botanists of Ontario outing. He noted that one of
these plants had a flowering stalk. We saw 13 plants
there in the autumn of 1989, none of which had
flowered that year.

1997

Arethusa bulbosa Linnaeus

Arethusa

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT Sil

Aréthuse bulbeuse

Arethusa bulbosa, regarded by many as one of our most beautiful orchids, is one of the three pink to purple fen
orchids in the District. It is the least abundant of the three. It can be distinguished from the other two, Pogonia
ophioglossoides and Calopogon tuberosus, by its essentially erect lateral sepals. It differs from the former by its
very small floral bracts and lack of an elliptical cauline leaf, and from the latter by its solitary flower.

DESCRIPTION

Height: 8 (13 - 24) 38 cm [172 plants].

Flowers: 1, rarely 2; usually purple, sometimes
purplish red, deep magenta, greyish magenta or light
lilac, occasionally purplish pink, purplish white or
lighter; lip white with distal border (and sometimes
sides) of same colour as sepals or lighter and with
stripes of more intense colour, with 3 rows of bristles
along centre line pale yellow or lighter; fragrance
moderate, sweet, floral, like violets or lilacs.

Leaf: 1, developing to maturity after anthesis, |
on non-flowering plants.

Overwintering State: as the corm of the current
year.

Capsules: greyish brown, ellipsoid to obovoid,
typically 2.5 x 0.6 cm, ascending to vertical; yield
very low, perhaps a few percent (Boland and Scott

46°)

(1991) report 16% in Newfoundland, Thien and
Marcks (1972) report 5% in Wisconsin).
Seeds: light brown, released by late September.

BLOOMING PERIOD: 28 May (10 June - 26 June) 3
July [42 records].

CoLony SIZEs: | - 300 flowering plants [10
records], as scattered individuals; the number of
plants counted on a single visit is about a quarter of
the plants flowering in a month because individual
flowers last only about a week (Ed Greenwood, per-
sonal communication 1987, quoted in J. M. Reddoch
and A. H. Reddoch (1987d)).

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

45°
76° 30!

75°

Arethusa bulbosa: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record. Major areas underlain by calcareous rock (marble and limestone) are shaded.

32 THE CANADIAN FIELD-NATURALIST

DISTRIBUTION: The Ottawa District is close to the
northern edge of the range of this Great Lakes — St.
Lawrence — Atlantic Coast orchid. It is a species of
the Mixed Forest Region and adjacent parts of the
Deciduous and Boreal Forest Regions. Within the
District, A. bulbosa occurs in a few peatlands on the
Canadian Shield and on the Lowlands, mostly in
areas of calcareous bedrock.

Hasirats: Arethusa bulbosa is confined to calcare-
ous treed and open fens (intermediate to rich fens)
and the minerotrophic parts of bogs (poor fens).
Most colonies occupy the semi-open treed fen zone
surrounding open sedge fens. Here the plants are
supported in deep, moist leaf humus or mounds of
Sphagna and other mosses on hummocks around the
bases of Eastern White Cedar, Tamarack and Black
Spruce. Common substrate mosses are Sphagnum
warnstorfii, S. teres, S. magellanicum and S. rus-
sowii. Some companion vascular plants are Three-
leaved False Solomon’s Seal (Smilacina trifolia),
Sweet Gale (Myrica gale), Pitcher-plant (Sarracenia
purpurea), Buckbean (Menyanthes trifoliata), Bog
Rosemary (Andromeda glaucophylla), Labrador Tea
(Ledum groenlandicum) and Small Cranberry
(Vaccinium oxycoccos). The corm of one flowering
plant of A. bulbosa that we discovered at White Lake
Fen was lodged in the bark of an Eastern White
Cedar tree about 5 cm above the hummock surface.

Occasionally, A. bulbosa is found in the open
among the sedges of the fen floor if it is not too wet,
and sometimes the odd plant turns up in an opening
in a cedar swamp adjacent to the treed fen zone con-
taining the main colony.

Arethusa bulbosa usually occurs in the same fens
as do Calopogon tuberosus and Pogonia ophioglos-
soides, but while the latter two orchids flourish on
the wet, open fen floor, A. bulbosa usually is found
in somewhat higher areas.

Vol. 111

LONG-LIVED COLONY: We have followed the
progress of the colony in the calcareous fen near
Poltimore (Reddoch 1989) since 1967. During the
1970s, the fen became increasingly wetter and the
plants scattered over the fen floor disappeared. At
the same time, dozens of new plants began to flower
on the hummocks surrounding the trees along the
edges of the fen and have continued to thrive there.
In the 1990s, flowering plants again appeared on the
fen floor close to those on the hummocks.

EARLY History: Arethusa bulbosa was added to the
Flora Ottawaensis in 1883 when it was discovered
in the Mer Bleue Bog growing among hundreds of
Pogonia ophioglossoides (UMTMG 47593], Fletcher
1884, Macoun, Whyte, and Fletcher 1884).
Subsequent visitors, apparently to the same locality
“near the gas springs” at the south edge of the peat-
land near Carlsbad Springs, reported large numbers
of plants (Whyte, Craig, and Cowley 1894; Fletcher
1896; Whyte, Craig, and Macoun 1897; Gibson
1908). Between 1883 and 1912, at least sixteen col-
lections were made in the Mer Bleue and distributed
to various Ontario and Quebec herbaria. There has
been only one recent report of A. bulbosa in the Mer
Bleue, that of Shirley Black (personal communica-
tion 1980) of the National Capital Commission, who
noticed a few plants east of the Borthwick Ridge in
the early 1970s.

ACYANIC Form: Arethusa bulbosa f. albiflora Rand
and Redfield has been reported from the fen near
Poltimore by Brunton (1985). While we have seen
several very pale flowers, there and elsewhere, as
described above, they still have some anthocyanin
and do not represent f. albiflora. This form has also
been reported from Alfred Bog, east of the Study
Area, by Emerson Whiting (personal communication
circa 1980).

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 33

aN

ern ANY
Se aan
FINS

Z| | Ps
V7
se Le

FiGurRE 3. Arethusa bulbosa, sedge fen habitat, Val-des-Monts Municipality (Wakefield Township, Gatineau County),
Quebec, 20 June 1976.

34 THE CANADIAN FIELD-NATURALIST

Calopogon tuberosus (Linnaeus) BSP.

Grass-pink

SYNONYM: Calopogon pulchellus (Salisbury) R. Brown

Vol. 111

Calopogon tubéreux

Calopogon tuberosus is the most conspicuous of the three purple to pink peatland orchids of the District with its
several brightly coloured flowers. It can be distinguished from the others, Arethusa bulbosa and Pogonia
ophioglossoides, by its several flowers with lips uppermost. It differs from the former by its spreading lateral

sepals and from the latter by the absence of a cauline leaf.

DESCRIPTION

Height: 18 (30 - 46) 64 cm [180 plants], at most
half the height of some plants in the southern part of
the range (Luer 1975).

Flowers: | (2 - 5) 10 [150 plants], at most half the
number of some plants in the southern part of the
range (Luer 1975); purple, occasionally light lilac or
greyish magenta; lip erect but lying on column after
fertilization, with 3 rows of white clavate hairs,
sometimes orange yellow or light yellow at the tips;
fragrance very faint floral or lacking.

Leaf: 1, 1 on non-flowering plants.

Overwintering State: as the corm of the current
year.

46°

Capsules: light brown, ellipsoid, typically 1.5 x
0.6 cm, erect or nearly so; yield highly variable,
averaging 55% [62 plants], appreciably larger than
reported for Newfoundland, 19% (Boland and Scott
1991), for Maine, 24% (Firmage and Cole 1988) and
Wisconsin, 16% (Thien and Marcks 1972).

Seeds: pale yellow to pale orange, released in
early September.

BLOOMING PERIOD: 12 June (24 June - 18 July) 6
August [85 records].

CoLony SIZEs: | - 400, typically to 165, flowering
plants, one exceptional colony with an estimated
3000 flowering plants on a lake-edge fen in the

45°
76° 30!

75°

Calopogon tuberosus: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 35

FIGURE 4. Calopogon tuberosus, bog habitat, Val-des-Monts Municipality (Wakefield Township, Gatineau County),
Quebec, 2 August 1980.

36 THE CANADIAN FIELD-NATURALIST

southwest (Adolf Vogg, personal communication
1988) [32 colonies], as scattered individuals.

DISTRIBUTION: The Ottawa District is in the northern
part of the distribution of this eastern orchid. It is a
species of the Deciduous, Mixed and Southeastern
Coastal Plain Forest Regions and adjacent edges of
the Boreal Forest. It is also found in subtropical
Florida and in Cuba. Within the District, C. tubero-
sus is confined to several peatlands and related areas
scattered across the Canadian Shield and the
Lowlands.

HasitaTs: Calopogon tuberosus is most commonly
encountered in the minerotrophic parts of bogs (poor
fens) and intermediate to rich fens, both floating and
consolidated, and sometimes in openings in treed
fens. The plants rise from corms deep in the wet peat
among typical peatland vegetation. Calopogon
tuberosus is almost always accompanied by Pogonia
ophioglossoides in these peatlands.

Occasionally, colonies appear from time to time
adjacent to large bogs on open, moist sands that have
been cleared temporarily of competing vegetation by
excavation (Mer Bleue Bog) or by fire (Morewood
Bog). These colonies are particularly susceptible to
lowering of water levels, disappearing entirely after
a drought.

Overburden stripping in the 1960s on the Dolman
Ridge beside the Mer Bleue Bog left open stretches of
mesic to wet-mesic sand beside the lagg of the bog.
The sand (pH 4.5 - 5.0) was quickly colonized by old-
field and bog plants, including Field Horsetail
(Equisetum arvense), Wood Horsetail (E. sylvaticum),
Poverty Grass (Danthonia spicata), Hudsonian Club-
rush (Scirpus hudsonianus), Wild Iris (Iris versicolor),
Blue-eyed Grass (Sisyrinchium angustifolium),
Buttercup (Ranunculus acris), Wild Strawberry
(Fragaria virginiana), Silvery Cinquefoil (Potentilla
argentea), Rough-fruited Cinquefoil (P. recta),
Dewberry (Rubus hispidus), Meadowsweet (Spiraea
latifolia), Steeplebush (S. tomentosa), Cow Vetch
(Vicia cracca), Evening-primrose (Oenothera bien-
nis), Sheep Laurel (Kalmia angustifolia), Labrador
Tea (Ledum groenlandicum), Blue Vervain (Verbena
hastata), fleabane (Erigeron sp.), and Brown-eyed
Susan (Rudbeckia hirta). In the early 1970s, five
orchids also grew in one such field: Calopogon
tuberosus, Liparis loeselii and Platanthera lacera in
both the mesic centre of the field and the wet-mesic
border beside the lagg, Malaxis unifolia in the mesic
centre of the field, and Spiranthes cernua beside the
lagg. Pogonia ophioglossoides appeared one year in
the centre of the field. The colony of C. tuberosus was
likely seeded from the 600 or so flowering plants
across the lagg in the bog. It built up to about 50 flow-
ering plants in 1973, but there was none at all in the
dry summer of 1975 and in subsequent years. The
field filled in rapidly with Spiraea and Speckled Alder

Vol. 111

(Alnus rugosa), and the orchids soon disappeared. In
1978, there was one flowering plant of C. tuberosus,
along with a few flowering plants of Malaxis unifolia.
In 1980, two Liparis loeselii were the only orchids
found.

Beside the Morewood Bog in the late 1970s, the
area of an old burn supported about 25 flowering
plants of Calopogon tuberosus, two Liparis loeselii,
six Spiranthes lacera and about 50 Malaxis unifolia.
The sand substrate was covered with patches of
lichens, Polytrichum and Sphagnum mosses, and
Staghorn Clubmoss (Lycopodium clavatum). Other
plants noted were White Beak-rush (Rhynchospora
alba), Round-leaved Sundew (Drosera rotundifolia),
Dewberry, Pink Pyrola (Pyrola asarifolia),
Leatherleaf (Chamaedaphne calyculata), Sheep
Laurel and Labrador Tea. Grey Birch was the domi-
nant tree nearby.

EARLY History: On 9 July 1860, Braddish Billings
Jr. collected both C. tuberosus [QK 12563] and
Pogonia ophioglossoides at Dow’s Swamp, presum-
ably on the sedge fen mat surrounding the small
pond shown in the 1925 air photo (Reddoch 1978b).
He described both orchids as rare. He also included
C. tuberosus among the plants he collected in the
vicinity of Ottawa during the summer of 1866
(Billings 1867). James Fletcher collected C. tubero-
sus at Dow’s Swamp in 1878 [DAO 17646] and stat-
ed in his Flora Ottawaensis of 1893 that it was rare
there. The species was collected there again in 1902
[CAN 227228].

Other early collections were those of John Kerr
McMorine in 1862 from Ramsay, Ontario [QK
66597, DAO 17651 (photograph of the latter in Ross
1984)].

Henry Ami’s 1879 specimens from the Mer Bleue
Bog near Carlsbad Springs [CAN 232431, MTMG
3442] were the first of many from that peatland.
James Fletcher (1893) stated that C. tuberosus
occurred “in enormous profusion in the Mer Bleue
and at Lake Flora, Hull”. Lake Flora was a small
pond on I’Ile de Hull (Nagy 1974) that is now Parc
Fontaine. This species, along with Pogonia
ophioglossoides and Spiranthes romanzoffiana (list-
ed as S. cernua) (Fletcher 1893), probably grew on a
sedge fen mat on the edge of the pond.

ACYANIC FORM: Calopogon tuberosus f. albiflorus
Britton has not been reported from the Study Area
but should be watched for since it has been seen in a
fen in Leeds County not far south of the District by
E. W. Greenwood (1965 letter to W. K. W. Baldwin,
at CAN).

ABERRATION: One unusual plant of C. tuberosus that
we collected [DAO 691503] on a sedge mat (poor
fen) near St.-Francois-de-Masham, Quebec, had three
flowering scapes and two small leaves arising from a
single corm. The scapes bore 4, 6 and 10 flowers.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT |

Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown) Luer

Calypso, Fairy Slipper

SYNONYM: Calypso borealis (Swartz) Salisbury

Calypso bulbeux

Calypso bulbosa is a rare plant of cedar swamps that is among the first orchids to bloom in the spring. It can be
recognized by its single, pleated, basal (often seemingly detached) leaf and its single pink to purple flower with
a saccate lip with white apron and twin points at the apex. Because of the scarcity of living plants in the area,
heights, colours, blooming period and colony sizes below are based, in part, on observations made at two now

extinct colonies in Eastern Ontario.

DESCRIPTION

Height: 6 (7 - 13) 15 cm [31 plants].

Flower: 1; sepals and petals purplish red, purplish
pink or reddish lilac; column similar but lighter in
colour, sometimes white; lip white, sometimes pur-
plish white at end of apron, light yellow on 3 rows of
hairs and on apron beneath them, slightly greyer yel-
low on twin tips at apex, with ruby spots on apron
and on hairs, sometimes symmetrically placed, and
somewhat darker ruby lines within pouch; fragrance
rose-like or lacking.

Scape: greyish magenta or reddish lilac.

Floral bract and sheaths: purplish grey.

Leaf: 1, 1 on non-flowering plants; formed in late
summer, Overwintering and dying after anthesis.

Overwintering State: as the corm of the year
subtending a leaf (see above).

Capsule: light brown, ellipsoid, typically 1.5 x
0.6 cm, erect; pedicel, about 0.6 cm long at anthesis,
continuing to grow to 4 - 6 cm long when capsule
mature (from herbarium specimens elsewhere in
Ontario and Quebec);

Seeds: colourless with orange white embryos
(from herbarium specimens elsewhere in Ontario and
Quebec).

BLOOMING PERIOD: 13 May (20 May - 7 June) 11
June [13 records].

COLONY SIZES: up to 25 flowering plants [5
colonies], generally as scattered individuals; occa-

46°

45°
76° 30

75°

Calypso bulbosa var. americana: @ = herbarium specimen. Major areas underlain by calcare-

ous rock (marble and limestone) are shaded.

38 THE CANADIAN FIELD-NATURALIST

sionally as clumps of up to 5 plants, all flowering or
a mixture of flowering and non-flowering plants.

CURRENT STATUS: rare if not extirpated in the
Ottawa District; one recently-discovered colony just
beyond the 50-km circle in Quebec; species declin-
ing throughout the southern part of its range in
Ontario (Whiting and Catling 1986).

DISTRIBUTION: The Ottawa District is near the
southern edge of the transcontinental distribution of
this orchid in North America. This variety is a plant
of the Boreal Forest Region and adjacent Mixed
Forest, as well as of the Montane Forest.

Within the District, the few localities known to
have harboured this orchid were on both the
Canadian Shield and the Lowlands. Outside the
District, the nearest known colonies in Ontario were
on the Shield 100 km west of Ottawa in Frontenac
and in Renfrew Counties. The former was discov-
ered by Elva MacKenzie in 1970 and the latter by
Sheila Thomson in 1979. Both colonies now appear
to be extinct. In Quebec, 70 km north of Ottawa,
Theresa Aniskowicz found and photographed a sin-
gle plant at Heney Lake in 1971. Calypso bulbosa is
confined to areas of calcareous bedrock.

HABITATS: Calypso bulbosa inhabits calcareous,
semi-mature to mature Eastern White Cedar
swamps. The plants grow in partially open places in
moist needle-mould on mounds around trees or on
the moist to wet swamp floor.

The colony just outside the 50-km circle near Wolf
Lake (see History below) is apparently of recent origin.
The plants appeared in an opening created by a survey
line cut through an ancient Eastern White Cedar -
Balsam Fir - Black Ash swamp where one of the cedar
stumps revealed 230 annual rings (Ray Chipeniuk and
Sonia Sawchuk, personal communication 1993). The
plants grew on mounds and in hollows with such other
species as Oak Fern (Gymnocarpium dryopteris), Wild
Lily-of-the-valley (Maianthemum canadense),
Platanthera obtusata, Foamflower (Tiarella cordifo-
lia), dewberry (Rubus sp.), Kidney-leaved Violet
(Viola renifolia), bedstraw (Galium sp.), Twinflower
(Linnaea borealis) and Red Maple shoots.

The Renfrew County site was a similarly ancient
cedar swamp. A tight clump of four flowering plants
grew on the moist swamp floor beside a large fallen
tree in 1979. Sheila and Harry Thomson did not see
them again, although they discovered two other
flowering plants nearby in two different years since
(Sheila Thomson, personal communication 1996).

The Frontenac County Eastern White Cedar -
White Spruce - Tamarack swamp was somewhat
younger. The trees were fairly closely spaced but were
free of live branches for three to five metres from the
ground. The C. bulbosa plants grew on mounds
around the cedars in small openings, not in the densest
part of the stand. In 1973, there were 25 flowering

Vol. H11

plants and many non-flowering plants, in 1975 and
1979, four flowering plants and a dozen or more non-
flowering plants. No plants could be found in the mid-
1980s (David White, personal communication).

At Heney Lake, the single flowering plant was in
a somewhat different habitat, a semi-mature conifer-
ous forest on a slope above the lake (Theresa Anis-
kowicz, personal communication 1981).

History: This orchid was rather less rare in earlier
times than it is now. It was first found at several local-
ities close to the new city, too close to survive for
long. In 1856, Elizabeth Keen White painted a plant
gathered from “Wet woods. Ottawa” by her husband,
William White (Dore 1965°). Stewart’s Bush, the
“damp, rich wood” where James Fletcher made the
first herbarium collections in 1877 [DAO 17808,
MTMG 47281], was already being cleared six years
later (Macoun, Whyte, and Fletcher 1884). In modern
terms, the bush was west of the Victoria Memorial
Museum between Bank Street and Bronson Avenue.

Fletcher (1893) also reported it from Billings
Bridge, where he found it abundant in 1878 but
could not find it at all after that. John Macoun (circa
1911°) noted it at several other localities close to the
city: “a few specimens occasionally found around
the swamp in front of Beechwood cemetery old
entrance gate” and “one specimen found near Rideau
Hall grounds in the spring of 1895 by young Mr.
Frechette”, as well as reporting R. B. Whyte’s col-
lections at “McKay’s Woods near Beechwood”,
1879, and “Beechwood”, 1888.

At the western edge of the District in Ontario,
Robert Campbell collected C. bulbosa at “Ramsay
near Almonte” in 1897 [DAO 204317, MTMG
20282, 20283], and Charles Macnamara found five
plants (and photographed one) in “thick mossy
woods” near Arnprior in 1915 (Macnamara circa
1940°; Reddoch 198 1c).

To the north, in Quebec, R. B. Whyte collected C.
bulbosa at High Falls on the Liévre River north of
Buckingham in 1892 (Fletcher, Scott, and Cowley
1892: Macoun circa 1911°). In 1898, John Macoun
discovered it at Chilcott’s Swamp, west of Alcove
[TRT 15300], and Faith Fyles found one flowering
plant there in 1911 ([DAO 200666], Fyles 1912). In
the 1950s, two specimens (at DAO) were collected
from Lac Gauvreau west of Wakefield.

Members of the Native Orchid Location Survey
did not turn up any colonies of C. bulbosa during
their years of intensive study of the Ottawa District
beginning in 1965. In 1993, Ray Chipeniuk and
Sonia Sawchuk discovered eight flowering plants on
their property northwest of Wolf Lake and just out-
side the District [photographs DAO 668349]. The
next year, we found nine flowering plants and nine
non-flowering plants there. In 1995, Ray and Sonia
(personal communication 1996) counted five flow-
ering plants, and, in 1996, one flowering plant.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 39

FiGuRE 5. Calypso bulbosa var. americana, cedar swamp habitat, Clarendon Township, Frontenac County, Ontario, 30
May 1973 and 2 June 1975.

40 THE CANADIAN FIELD-NATURALIST

Vol. 111

Coeloglossum viride (Linnaeus) Hartman var. virescens (Muhlenberg) Luer

Long-bracted Orchid

Orchis 4 longues bractées

SyNonyMs: Habenaria viridis (Linnaeus) R. Brown, Habenaria bracteata (Muhlenberg) R. Brown

Coeloglossum viride is a spring-blooming, green, forest orchid that occurs infrequently and does not seem to
survive for many seasons. It can be recognized by its cauline leaves, by the floral bracts which are as long as the
flowers at the top of the inflorescence and much longer at the bottom, and by the lip which is strap-like in shape
with a notch at the end and which is much longer than the shallow bulbous spur.

DESCRIPTION

Height: 14 (19 - 29) 40 cm [53 plants], apprecia-
bly shorter than the 50 or 60 cm or more cited by
various authors (for example, Luer 1975).

Flowers: 4 (5 - 16) 28 [41 plants]; greyish green,
occasionally somewhat more yellow; lip usually, but
not always, with coloration such as brownish red of
varying intensity and extent near base; fragrance
faintly spicy or apple blossom.

Leaves: 2 - 5, usually 4, less frequently 3; 2 on
non-flowering plants.

Overwintering State: a greyish green, broadly
conical shoot, | - 2 cm above ground beside the cur-
rent year’s stem, appearing there in late September;

Currah, Smreciu, and Hambleton (1990) describe the
underground seasonal development of C. viride in
Alberta: during dormancy, the new shoot is subtend-
ed by a new, fully elongated, tuberous rhizome and
four fleshy roots; a new plant begins development a
year before flowering.

Capsules: brown, ellipsoid, typically 0.8 x 0.35
cm, ascending to erect; yield generally fairly high,
sometimes approaching 100%, averaging 75% for 10
plants.

Seeds: light brown, released in mid September.

BLOOMING PERIOD: 10 May (25 May - 26 June) 9
July [45 records]; however, by mid-June the column
may be no longer functional although the sepals and

46° L

76° 30'

75°

Coeloglossum viride var. virescens: 1 = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. The Canadian Shield is shaded.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

FiGuRE 6. Coeloglossum viride var. virescens, deciduous forest habitat, Gatineau Park, Quebec, 4 June 1978.

41

42 THE CANADIAN FIELD-NATURALIST

petals remain in good form and may continue to be
so for another month. The persistence of flower parts
in good form may be the source of some records of
late-blooming plants.

COLONY SIZES: | - 27 flowering plants; 3 or fewer
in 70% of colonies [44 colonies], usually as separate
individuals, rarely as pairs (in one case, two stems
arising from one of the previous year).

DISTRIBUTION: The Ottawa District is close to the
northern regional limit of the transcontinental distri-
bution of this plant of the southern Boreal, Mixed
and northern Deciduous Forest Regions. Within the
District, C. viride is confined almost completely to
the Canadian Shield north of the Ottawa River. The
lower part of Gatineau Park is a favoured area.

HaBitaT: Coeloglossum viride inhabits semi-mature
to mature, deciduous and mixed forests of such trees
as Sugar Maple, Hop Hornbeam, Butternut (Juglans
cinerea), White Birch and White Pine. Plants gener-
ally grow in the partial shade of forest clearings and
edges or among widely-spaced trees. The leaf-cov-
ered soil is generally well-drained and often rocky.
Ground cover in forests is usually sparse or non-
existent, but in clearings and along trails may be a
mixture of forest and old-field species such as
Bladder Campion (Silene vulgaris), Hairy Sweet
Cicely (Osmorhiza claytonii), Fragrant Bedstraw

Vol. 111

(Galium triflorum), Yarrow (Achillea millefolium),
asters (Aster spp.) and some grasses.

LONGEVITY: It is easy to get the impression that C.
viride is quite transient. A plant will appear where
there was none before and then will not be there on
the next visit a few years later. The longest record
that we have for individual plants is four years after
we discovered them. Charles Macnamara (1911,
circa 1940") found the same plant for seven years
“under a moose-wood bush” in the Arnprior area. He
photographed the plant in 1906.

EARLY History: In June 1862, J. K. McMorine col-
lected C. viride at Ramsay, Ontario [QK 12943]. Also
on the western edge of the District, R. B. Whyte col-
lected this species on 25 July 1876 “near Arnprior”
(Macoun circa 1911"), probably near his family home
at Galetta. Like many of R. B. Whyte’s collections,
this one may no longer be extant. Other early collec-
tions came from places closer to the city, beginning
with James Fletcher’s from “below Beechwood
Cemetery” in 1878 [DAO 17299]. Macoun (circa
1911”) cites a collection by William Scott from King
Mountain in 1892, which was soon followed by oth-
ers from the Gatineau Valley and other parts of the
Canadian Shield north of the Ottawa River.

James Fletcher (1893) considered this species to
be “not common”, while Macoun (circa 1911’)
reported it to be “local and scarce”.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 43

Corallorhiza maculata (Rafinesque) Rafinesque var. maculata
Corallorhiza maculata (Rafinesque) Rafinesque var. occidentalis

(Lindley) Ames
Spotted Coral-root

SYNONYM: Corallorhiza multiflora Nuttall

Corallorhize maculée

Corallorhiza maculata is a widely distributed orchid of the forests of the Shield which displays considerable
variation in colour. It can be identified by its leafless habit, its overall colour in the range from magenta to
brown to yellow and its several flowers with white lips with, or rarely without, reddish spots.

DESCRIPTION

Height: 10 (18 - 33) 49 cm [169 plants].

Plant colours: stem ranging from violet brown,
brownish violet, greyish rose or greyish red to yellow-
ish brown or light brown to greyish yellow depending
on the relative amounts of red and yellow pigments,
sometimes with decreasing red toward top of stem;
sheath generally paler than stem, e.g., greyish red to
yellowish white with or without fine reddish lines on
veins; in f. flavida (see Acyanic Form below): entire
plant (except lip) yellow, or less frequently, light yel-
low, greyish yellow or greenish yellow.

Flowers: 4 (8 - 19) 31 [111 plants]; on typical
form: sepals greyish yellow, light yellow, yellowish
white, greenish white or occasionally brownish
orange; tips of sepals ruby or violet brown; lip white;
lip, column, petals and sepals with deep magenta,
ruby or dark purple spots; in f. flavida: lip white
without spots; fragrance of typical form sweet, like
apple blossoms.

Overwintering State: one or more pale green
shoots, several centimetres below ground, rising
from coralloid rhizomes near the current year’s
stem(s), by early fall.

46°

45° {=
76° 30'

75°

Corallorhiza maculata (both varieties): HM = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. The Canadian Shield is shaded.

44 THE CANADIAN FIELD-NATURALIST

Capsules: light brown, ellipsoid, typically 1.5 x
0.5 cm, pendent; yield usually 50% to 100%, averag-
ing 70% [58 plants] (often very low or none in
drought years or when attacked by weevils).

Seeds: pale yellow, released in mid September.

BLOOMING PERIOD: 3 June (30 June - 28 July) 2
August [56 records].

COoLony SIZEs: | - 37, typically to 15, flowering
stems [93 colonies]; as scattered individuals and in
clumps of up to a dozen or so stems.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of occurrence of this transcontinental
orchid. It is a species of the Montane, southern
Boreal, Mixed and northern Deciduous Forest
Regions in Canada and the United States. It is also
found in Mexico and adjacent Guatemala. Within the
District, C. maculata is confined mainly to the
Canadian Shield, where it is widespread.

HasitatTs: Corallorhiza maculata is exclusively a
woodland species. It inhabits mesic, semi-mature
and mature, deciduous and coniferous forests. The
deciduous tree communities are most commonly
Beech, Sugar Maple - Beech and Sugar Maple - Red
Oak. Eastern Hemlock is by far the most common
type of coniferous forest. Even in deciduous forests,
the orchids tend to be near hemlock trees if they are
present (as Michael Runtz (1984) has noted in the
Arnprior area). The most common companion
species on the shaded, leaf-covered forest floors is
Wild Lily-of-the-valley (Maianthemum canadense),
accompanied by tree seedlings in somewhat less
shaded locations. Sands and sandy loams are the
usual substrates.

There are enough observations of colonies on
forested slopes at the edges of ponds, swamps and
peatlands, and sometimes in the humus-rich soil of
the wetland perimeters, to consider these as special
microhabitats within the broader forest descriptions.

LONG-LIVED COLONY: Since 1975, we have fol-
lowed a colony of C. maculata in lower Gatineau
Park that Anne Hanes discovered and photographed
in 1965. The colony consists of both the typical
form and f. flavida. John Freudenstein annotated our
photographs of both forms from this colony [DAO
468578, 478803] as var. maculata. The plants are
scattered over both level and sloped areas under a
canopy of Beech with a sprinkling of Eastern
Hemlock, Sugar Maple and Hop Hornbeam. Since
1975, we have seen a total of 85 flowering stems.
The number of flowering stems appearing in a year
has varied rather randomly from 0 to 11 between
1975 and 1985 and from | to 6 between 1985 and
1996. Many of these stems occurred as isolated indi-
viduals, but clusters of 3 or 5 stems were noted.
Often the individual stems did not reappear the fol-

Vol. 111

lowing year, but some members of a five-plant clus-
ter did. (See also Acyanic Form below.)

EARLY History: In September 1861, John Kerr
McMorine collected C. maculata at Ramsay, Ontario
[QK 12620]. We have encountered three other nine-
teenth century specimens: a previous year’s stalk
collected by James Fletcher on 15 May 1879, from
“Swamp, Ottawa, Ont.” [DAO 17734]; two stems
rising from a single rhizome collected by W.
Macoun in July 1886 from “Ottawa, Ont.” [MTMG
2376]; and a Quebec specimen collected by W. Scott
on 25 June 1891 from North Wakefield (= Alcove)
[CAN 17165, 100563]. Fletcher (1893) describes
this orchid as rare and lists Beechwood, Clark’s
Wood, Chelsea and Kingsmere as locations. John
Macoun (circa 1911°) adds four localities of his
own, all in Quebec. Three are in the lower Gatineau
Valley and one is near Quyon.

MORPHOLOGICAL VARIANT: John Freudenstein
(1987) published a preliminary morphological analy-
sis of C. maculata showing several differences
between early and late blooming plants. These dif-
ferences include greater lip dilation, floral bract
length and number of flowers as well as a broader
sepal sinus shape in the early blooming plants. In his
forth-coming monograph, Freudenstein (personal
communication 1996) will apply the name
Corallorhiza maculata var. occidentalis (Lindley)
Ames to these plants. He annotated 20 collections
from the District at DAO and CAN, six as var. occi-
dentalis and the rest as var. maculata. The blooming
period for the specimens of the former variety is 3
June to 3 July, while for the latter variety it is 1 July
to 28 July. The date of 3 June is two weeks earlier
than any other Ottawa District record.

ACYANIC ForRM: The long-lived colony discussed
above has frequently had a few plants of C. maculata
var. maculata f. flavida (C. H. Peck) Farwell since
Anne Hanes first told us about them (A. H. Reddoch
and J. M. Reddoch 1987b). These plants are much
easier to find and count than the typical form
because of their bright and distinctive colour (see
flower description above). They have usually
occurred as isolated individuals and have rarely
reappeared close to the site of a previous occurrence.
Sometimes no plants of this form were seen for one
or two years. Over the two decades, 20 of the 85
stems were f. flavida. There have been no other
reports of this form in the District.

Forma flavida is interesting in that it reveals the
yellow pigment that is also present in the typical form.
In the typical form, the yellow is mixed with a reddish
pigment in varying proportions to give the reds,
browns and yellows described above. This reddish
pigment can be seen in its pure form in the spots on
the white background of the lip of the typical form.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 45

FiGuRE 7. Corallorhiza maculata, deciduous forest habitat, Gatineau Park, Quebec, 22 July 1973 (plant) and 22 July 1972
(flowers).

46 THE CANADIAN FIELD-NATURALIST

Corallorhiza striata Lindley
Striped Coral-root

SYNONYM: Corallorhiza macraei A. Gray

Vol. tf

Corallorhize striée

Corallorhiza striata is the least abundant of the three Corallorhizas. This spring-flowering orchid is widely but
thinly distributed in the calcareous areas of the District. It can be recognized by its leafless stem that is not
green but some colour in the range pale yellow to orange to ruby and by its striped flowers with boat-shaped,

ruby lips.

DESCRIPTION

Height: 4 (12 - 23) 35 cm [115 plants].

Plant colours: stem ranging from greyish ruby to
greyish rose to very pale, e.g., greyish orange (flesh),
depending on the amounts of red and yellow pig-
ments, sometimes with decreasing red toward top of
stem; sheaths, 1 or 2 above ground, generally paler
than stem, e.g., yellowish white with or without fine
reddish lines on veins.

Flowers: 2 (8 - 17) 25 [90 plants]; pedicellate
ovary and bract light or pale yellow rarely with faint
reddish component; sepals and petals with pale yel-
low to translucent grey or occasionally pale orange

background and dark magenta lines along veins; lip
similar but with broader lines of more intense colour,
sometimes coalescing to a single mass of colour on
outer half, variously ruby, greyish ruby, violet brown
or brownish violet; column light yellow or pale yel-
low usually with a conspicuous reddish spot on
underside; fragrance faint, sweet or daisy-like.

Overwintering State: whitish shoots developing
below ground at nodes of the coralloid rhizome with-
in a half metre or so of the current year’s or recent
years’ stems.

Capsules: light to medium brown, ellipsoid, typi-
cally 2 x 0.6 cm, pendent (see Figure 1a); yield very

76° 30

re

75°

Corallorhiza striata: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, & = literature reference. Major areas underlain by calcareous rock (marble and lime-

stone) are shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT AT

aa ro 3 TP. “
ee ily
aa Uy, AEG fi

CLP,
of Ai UV) ie
tld Yl La

ND

FiGuRE 8. Corallorhiza striata, mixed forest habitat, Gatineau Park, Quebec, 31 May 1969 (plant) and 4 June 1978
(flowers).

48 THE CANADIAN FIELD-NATURALIST

low to none (see Long-lived Colony below),
although on one occasion in our experience, 100%.
Seeds: pale yellow.

BLOOMING PERIOD: 23 May (27 May - 12 June) 22
June [29 records].

COLONY SIZEs: | - 155, typically to 30, flowering
stems; 3 or fewer in 65% of colonies [18 colonies];
as scattered individuals or in clumps of up to about a
dozen stems.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District is within the
eastern, Mixed Forest, distribution of this Great
Lakes - St. Lawrence and western montane species.
Within the District, C. striata is thinly scattered in
areas of calcareous bedrock on the Shield and on the
Lowlands near the Ottawa River.

Hasirats: Corallorhiza striata occurs in a variety
of mesic, coniferous and mixed forests, as well as in
a few cedar swamps.

It grows in partial sun among widely-spaced
clumps of Eastern White Cedar on the limestone
plains of the southwest and under the deeper shade
of Balsam Fir and Eastern Hemlock on the Shield to
the north. It is also found in dappled to deep shade in
semi-mature and mature mixed forests of such trees
as Eastern White Pine, Balsam Fir, Sugar Maple,
Beech and Eastern White Cedar. In one forest of just
this mix of trees in the upper Gatineau Valley,
Cypripedium parviflorum var. pubescens, Epipactis
helleborine, Goodyera tesselata, and Platanthera
hookeri were also present. Under a covering of leaf
and/or needle humus, the substrate is usually sand,
but occasionally it is clay.

A few colonies of C. striata have been encoun-
tered growing among the mosses or in the wet organ-
ic soil of Eastern White Cedar - Balsam Fir swamps.

LONG-LIVED CoLony: In 1968, John Pinder-Moss of
Carleton University told us about a colony of C. stri-
ata that he had just discovered in the southern part of
Gatineau Park [CCO 15486]; we have followed its
progress since then. (Photographs of plants in this
colony were used to prepare the Figure.)

The colony is spread over an area 25 m x 50 m in
a maturing Eastern White Pine - Sugar Maple forest
that is part of a larger Sugar Maple - Beech forest.
Within the colony, the pines average 23 m in height,
dbh 40 - 80 cm, and the maples, 19 m in height,
average dbh 20 cm. Herbaceous plants at scattered
locations on the mesic, sandy forest floor are Jack-
in-the-pulpit (Arisaema triphyllum), Lady Fern
(Athyrium filix-femina), Trout Lily (Erythronium
americanum), Wild Lily-of-the-valley (Maianthe-
mum canadense), Red Trillium (Trillium erectum),
White Trillium (7. grandiflorum), White Baneberry
(Actaea pachypoda), Red Baneberry (A. rubra),

Vol. 111

Wild Ginger (Asarum canadense), Blue Cohosh
(Caulophyllum thalictroides), Poison Ivy (Rhus radi-
cans) and lettuce (Lactuca sp). There are a few
plants of Cornus alternifolia and a few saplings of
Sugar Maple, Black Cherry and Basswood.

There were 60 stems of C. striata in 1968 and 155
in 1969. Since then, the number of stems/year has
varied between 32 and zero (zero in four non-con-
secutive years), with an average of seven stems/year.

More detailed mapping in the past 11 years has
revealed that most of the stems appearing during that
time are confined to five nodes. Three of the nodes
are about a metre in diameter and two others mea-
sure about 3 m x 7 m. We have found a total of only
ten stems outside these nodes (and their positions
may define additional nodes).

Within the nodes, stems emerge at intervals of one
to five years. The most prolific rhizome has been
producing stems for four consecutive years (1993 to
1996). The stems within a node tend to have a simi-
lar amount of greyish ruby colour from year to year,
while the amount of the colour varies from node to
node. From these observations, we infer that the
stems at each node derive from a single rhizome
system.

This colony is heavily attacked by weevils
(Stethobaris ovata (LeConte) (Howden 1995)) that
cause the stems to dry up soon after flowering; thus
the colony rarely produces capsules. This observa-
tion leads us to conclude that since so little seed has
been produced during the past three decades, the
stems appearing during that time likely have come
from long-lived rhizomes. Insect or slug activity
occurs below ground as well; sheaths on one subter-
ranean shoot examined already showed the holes that
are seen on mature stems.

EARLY History: In 1861 and 1862, John Kerr
McMorine collected this orchid at Ramsay, Ontario
[QK 12654, 66619, 79687]. Closer to the city of
Ottawa, C. striata was added to the Flora
Ottawaensis in 1885 when Miss C. L. Hanington dis-
covered it at New Edinburgh (Whyte, Macoun, and
Small 1887; Fletcher 1887). The July 1885 collec-
tion by Mrs. Chamberlin at DAO [17709] labelled
“Beechwood, Ottawa” presumably records this
event. (Mrs. Agnes D. (FitzGibbon) Chamberlin, the
daughter of Suzannah Moodie and the niece of
Catharine Parr Traill, prepared illustrations of
Canadian wildflowers that subsequently appeared in
two books with her aunt’s text. She was a member of
The Ottawa Field-Naturalists’ Club from 1880 to
1894.) In 1894, James Fletcher reported that “several
plants of this beautiful orchid were collected last
spring at Beechwood, and in the woods at the back
of Rideau Hall. The first specimen was found by the
Hon. Archie Gordon. At the meeting of the Ottawa
Electoral District Agricultural Society, on June Sth, a
bunch of these flowers was exhibited.”

1997

Other nineteenth century collections, in fact all
other herbarium collections that we have examined
to the present, came from Quebec. James Fletcher
(1893) reported “Kingsmere” as an additional loca-
tion for this “very rare” orchid. John Macoun (circa
1911°*) listed two additional collections that we have
not seen: “Snell Lake, June 15, 1889” by Richard
Lee and “one specimen found in the woods east of
McKay Lake, June 1902” by Henry St. Jacques. We
have not been able to locate Snell Lake.

In 1967, W. G. Dore remembered that participants
on an Ottawa Field-Naturalists’ Club excursion in
May of some year between 1930 and 1933 discov-
ered a colony of C. striata “on rocky land under
cedars” on the northeast side of Fairy Lake. No spec-
imen was taken. His note describing the event is
attached to an unnumbered herbarium sheet at DAO.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 49

Charles Macnamara (circa 1940*) stated that he
had “not found it so very uncommon around
Arnprior”’.

MISCELLANEOUS: On a number of occasions over a
period of years at two sites, we have noticed a black
wasp about 2.5 cm long with orange legs visiting C.
striata inflorescences. We took a wasp to the
Biosystematics Research Institute of the Central
Experimental Farm where it was identified as
Pimpla pedalis Cresson (or Coccygomimus pedalis
(Cresson) Townes and Townes). The wasp has been
seen both on plants in bud and in flower, but we do
not know what it was doing on these visits.
Although it mainly visited the inflorescence, we
could see no evidence that it was functioning as a
pollinator.

50 THE CANADIAN FIELD-NATURALIST

Corallorhiza trifida Chatelain
Early Coral-root

SYNONYM: Corallorhiza innata R. Brown

Volott

Corallorhize trifide

Corallorhiza trifida is a widely distributed member of the spring flora in the Ottawa District. It can be recog-
nized as a leafless, early-flowering orchid with greenish sheaths, stem and flowers, the latter with white lips

with or without spots.

DESCRIPTION

Height: 4 (11 - 21) 32 cm [189 plants].

Plant: stem greyish green; sheaths | or occasion-
ally 2, similar in colour to stem, often with brownish
tips in typical variety. Its lack of leaves in mature
plants and its coralloid root are usually regarded as
evidence of a dependence on mycorrhiza; however,
spectroscopic studies have shown the presence of
some chlorophyll, and physiological studies have
shown that some photosynthesis also occurs (as sum-
marized by Freudenstein and Doyle 1994).

Flowers: 2 (4 - 11) 15 [101 plants]; sepals and
petals greyish green similar to stem and sheaths or
sometimes more yellowish, lip white; in the typical
variety: column, upper and lower surfaces of lip and

46°

45°
76° 30

of petals with deep magenta or ruby spots, sepals
often with brownish tips and petals with lighter
brown tips; occasionally, brownish colour absent
from tips of sepals and petals, and spots on lip small
and restricted to base, a variation also reported by
Case (1984) from the north shore of Lake Superior;
in var. verna (see Acyanic Form below): red and
brown colorations absent; odour faint and acrid or
richly floral, or lacking.

Overwintering State: in swamps: several or many
pale green shoots, 4 mm in diameter, 1.5 - 2.5 cm
tall, about 4 cm down in the moss substrate, rising
from nodes of coralloid rhizomes several centimetres
from the current year’s stems, by early fall; in forests:
shoots likely in a similar state below ground.

75°

Corallorhiza trifida (both varieties): HH = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 51

Ninny
/80

FiGurRE 9. Corallorhiza trifida var. verna (plant): deciduous forest habitat, Gatineau Park, Quebec, 19 May 1968; C. trifi-
da var. trifida (flowers), cedar swamp habitat, West Carleton Township, Regional Municipality of Ottawa-
Carleton, Ontario, 24 May 1975.

52 THE CANADIAN FIELD-NATURALIST

Capsules: brown, ellipsoid, typically 1 x 0.5 cm,
pendent (see Figure la); yield depending on habitat:
clumps in swamps approaching 100%, scattered
individuals in woodlands often very low; previous
year’s stems persisting well into the next season.

Seeds: brownish orange, released in late
September.

BLOOMING PERIOD: 12 May (25 May - 14 June) 23
June [55 records].

CoLony SIZEs: | - 200, typically to 50, flowering
stems; 10 or fewer in 69% of colonies [139
colonies]; the large colonies occur in swamps: one
exceptional colony with 2500 flowering stems in
West Carleton Township cedar swamp in 1969 (Hue
MacKenzie, personal communication); as clumps of
typically 20 stems in swamps and as scattered indi-
viduals in woodlands.

DISTRIBUTION: The Ottawa District is well within
the North American range of this orchid. It is a
transcontinental species of the Boreal and Mixed
Forest Regions and adjacent regions of the Montane
and Deciduous Forests and Tundra. Within the
District, C. trifida is widely distributed except in the
southeast.

HABITATS: Various swamps, treed fens and mesic
forests harbour this orchid in the Ottawa District.
These habitats provide adequate moisture and dap-
pled to deep shade.

Corallorhiza trifida is frequent in swamps, espe-
cially those dominated by Eastern White Cedar and
accompanied by a selection of Black Spruce, White
Spruce, Tamarack, Black Ash and Yellow Birch. In
the open or sometimes under ferns or the lower
branches of conifers, the orchids grow in moist or
wet organic soil or among mosses. Typical compan-
ion species in more open swamps are Cinnamon
Fern (Osmunda cinnamomea), Jack-in-the-pulpit
(Arisaema triphyllum), Bluebead-lily (Clintonia
borealis), Cypripedium reginae, Malaxis
monophylla, Platanthera hyperborea, Naked
Mitrewort (Mitella nuda), Foamflower (Tiarella
cordifolia) and One-flowered Wintergreen (Moneses
uniflora). Treed fens and stream edges are other wet
environments inhabited by this orchid.

Some deciduous, mixed and coniferous forests are
also suitable habitats for C. trifida. Sugar Maple -
Beech, Sugar Maple - Beech - Eastern Hemlock -
Eastern White Pine, Eastern White Pine - Red Maple,
Eastern Hemlock - Balsam Fir and Eastern Hemlock
- Eastern White Cedar are some of the common
mesic forest types. Dense, dryish Eastern White
Cedar thickets sometimes also shelter this orchid.
The substrates are usually sands and sandy loams.

LONG-LIVED COLONY: A colony that we have
noticed from time to time since 1968 in Gatineau

Voh Ti

Park is at the edge of a seasonal pond under a
canopy of Yellow Birch, Eastern Hemlock and Red
Maple. The plants grow in wet, partially-decayed
leaf-litter close to the high water line. In 1968 there
were a dozen flowering stems. The number of flow-
ering stems has varied over the years from 15 (in
1996) to none in some years. There was one flower-
ing stem in 1995. It is possible that coralloid rhi-
zomes are long-lived and put up flowering stems
under favourable conditions for many decades.

EARLY History: James Fletcher collected C. trifida
at “Billings Bridge, Nr. Ottawa” on 21 May 1877
and 12 May 1878 from “damp meadow at edge of
wood” [both on one sheet, DAO 17733]. Other nine-
teenth century collections came from Dow’s Swamp
[at CAN, MTMG and TRT] and Beaver Meadow
near Hull [at MTMG and TRT]. Among the speci-
mens cited by John Macoun (circa 1911°) is a
Robert B. Whyte collection from “Swamp at Lake
Flora. Hull” on 24 May 1878 and one of his own “in
woods at Britannia 1903”. The whereabouts of these
collections are unknown. James Fletcher (1893) con-
sidered this orchid to be “rather rare” in the District.

This species and Galearis spectabilis were the two
representatives of the orchid family gathered during
an Ottawa Field-Naturalists’ Club excursion to
Ironsides, Quebec, on Saturday, 23 May 1909
(Halkett 1909).

ACYANIC FORM: Corallorhiza trifida var. verna
(Nuttall) Fernald differs most notably from the typi-
cal plant in lacking the red and brown colorations
(see descriptions of plant and flowers above). The
whole plant except the lip is greyish green or some-
times more yellowish; the lip is white without spots.

This variant is more common in the southern part
of the range of the species (Luer 1975). The two
variants are about equally abundant in the District
and sometimes grow adjacent to each other in
swamps. Verna is the variant generally found in
mesic forest habitats.

Nuttall (1823) and Fernald (1946) each describe
morphological differences in the sepals, petals and
lip between var. verna and the typical plant, in addi-
tion to the colour differences. Some recent authors
ignore the morphological differences, but John
Freudenstein (personal communication 1996)
believes that they are real although not yet well
understood. In the District, we do not find these mor-
phological differences to be significant or to corre-
late with the colour differences.

If the only difference were the obvious colour dif-
ference, it would be appropriate to reduce the rank of
var. verna to that of a form. However, it would be
premature to do so until the nature and significance
of the morphological variation throughout the range
of the species are established.

[O97

Cypripedium acaule Aiton

Pink Lady’s-slipper
Stemless Lady’s-slipper

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 53

Cypripéde acaule
Sabot de la vierge

This species is a showy and abundant orchid that inhabits a variety of shaded, acidic habitats, both wet and dry.
It is easily recognized by its mostly pinkish lip with a longitudinal slit and its pair of basal, ascending, strongly

ribbed leaves.

DESCRIPTION

Height: 10 (18 - 36) 59 cm [231 plants].

Flower: one, often greyish magenta but varying
from pink to ruby to brownish violet, the colour
usually more intense on the veins; sepals and petals
usually darker than lip, greyish ruby but occasionally
light or reddish brown; lateral sepals fused but
sometimes with a slight double tip; petals and interi-
or of lip near column with long dense pubescence
that is mostly colourless but pinkish or reddish near
central areas; staminode white, green and brown in
varying proportions; in f. albiflorum (see Acyanic
Form below): lip and staminode white, sepals and
petals yellowish green; fragrance usually pleasantly
floral, varying in intensity from quite strong to none,
sometimes with sharp notes.

Leaves: 2 on flowering plants, 1 or 2 on non-
flowering plants.

Overwintering State: one or more, greyish green,
broadly conical shoots, 1 - 2 cm above ground beside
the current year’s stem, appearing in late September.

Capsule: brown, ellipsoid, typically 3 x 1 cm,
somewhat ascending, persisting to following year
with conspicuous remains of flower (unfertilised
flowers dehiscent); yields from 10% to 40%, occa-
sionally higher, averaging 30% [474 plants]; these
yields, from eight counts at five different sites over a
number of years, are notably higher than the percent-
ages reported elsewhere for pollen mass removal,
flower pollination and fruit set, which are generally
less than about 12% (Plowright, Thomson, and Thaler
1980; Davis 1986; Primack and Hall 1990; Gill 1996).

46° =

45° |.
76° 30'

as

75°

Cypripedium acaule: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded; areas underlain by sand and

sandstone have been omitted for clarity.

54 THE CANADIAN FIELD-NATURALIST

Seeds: greyish yellow, released in early to mid
October.

BLOOMING PERIOD: 18 May (29 May - 16 June) 4
July [105 records].

CoLony SIZEs: | - 1500, typically to 250, flowering
and non-flowering plants [273 colonies], mostly as
scattered individuals but with some clumps of up to
about a dozen flowering stems, in all habitats.

CURRENT STATUS: one of the most abundant orchids
in the District.

DISTRIBUTION: The Ottawa District is well within
the range of this mainly Great Lakes - Appalachian -
Atlantic Coast species of the Mixed Forest Region
and adjacent Deciduous and Boreal Forests. Within
the District, C. acaule is widespread across the
Canadian Shield and the Lowlands. It occurs in wet-
lands and in areas of acidic rocks and of sands left
by the Laurentide ice sheet, the Champlain Sea and
subsequent rivers.

HABITATS: Many wet, moist and dry sites provide
suitable habitats for this orchid, as long as they are
acidic and at least partially shaded.

Old bogs and swamps are the usual wet-mesic
habitats. Colonies of C. acaule grow in Sphagnum
moss in old bogs. The plants occur among erica-
ceous shrubs or with little other ground cover, in bog
forests of Black Spruce and Tamarack or near groves
of these trees in more open bogs (and among scat-
tered Grey Birch east of the Rideau River). In areas
of calcareous bedrock, plants are found occasionally
in semi-open Eastern White Cedar - Black Spruce -
Tamarack fens and swamps. Here they live above
the wetland floor in the more acidic conditions of
hummocks and wetland margins. Rarely there are a
few plants in seepage areas in otherwise mesic
forests.

Mesic coniferous, mixed and deciduous forests on
deep sand and in shallow soil over acidic rocks fur-
nish suitable habitats for C. acaule. Red Pine planta-
tions are common habitats as are forests of Eastern
White Pine, in pure stands or mixed with other trees
such as Red Maple, Sugar Maple, Red Oak,
Trembling Aspen and Grey Birch. Familiar species
growing with C. acaule in the needle or leaf humus
are Bristly Clubmoss (Lycopodium annotinum),
Bracken (Pteridium aquilinum), Bluebead-lily
(Clintonia borealis), Wild Lily-of-the-valley
(Maianthemum canadense), Starflower (Trientalis
borealis) and Twinflower (Linnaea borealis). A Red
Oak - Trembling Aspen - Jack Pine - Red Pine forest
on old dunes beside the Ottawa River at Constance
Bay also shelters C. acaule.

Precambrian knolls and hilltops are typical dry
sites. Plants grow in shallow soil and humus
among lichens, Polytrichum mosses and Bracken,
rarely in grasses, around trees and in open woods.

Vol. 111

Comparable dry conditions are provided by sand-
stone flats on the Lowlands.

LONG-LIVED COLONIES: Individual plants of C.
acaule may live for many decades and colonies may
last for even longer.

In the Larose Forest, the large colonies already
well established in 1969 in 40-year-old Red and
Scots Pine plantations (see also Dore 1969), remain
to the present (1996). Numbers have dwindled where
a picnic ground was installed and where the trees
have been so well trimmed that there is more light
and, consequently, increased ground cover. Plants of
Malaxis unifolia and Spiranthes lacera often accom-
pany C. acaule on the needle-strewn plantation floor.

The Eastern White Pine forest and its colony of C.
acaule at Blueberry Point near Aylmer, Quebec,
were presumably already of considerable age when
first visited by naturalists at the turn of the century.
Herbarium collections of C. acaule record the exis-
tence of the colony between 1905 [CAN 116907]
and 1948 [Breitung 6421 at DAO].

EARLY History: The earliest known collection of
C. acaule was made in 1862 by John Kerr McMorine
at Ramsay, Ontario [QK 2681]. Around the turn of
the century, this species was known from many loca-
tions, in both wetlands (Fletcher 1893; Whyte, Craig,
and Macoun 1897; Fyles 1912) and forest habitats.

Plants were regularly sought on Ottawa Field-
Naturalists’ Club excursions to Gilmour’s Grove on
the Gatineau River at Chelsea, Quebec (Ami 1896;
Clarke 1904, 1908; Reddoch 1979b) and evidently
were collected enthusiastically by many of the 200
naturalists and student teachers who attended. In
1908 Clarke reported that “Dr. Fletcher ... deprecat-
ed the digging up of the roots of [C. acaule] which
could not be cultivated [like the other species men-
tioned]. Through this useless destruction the plant is
now very rare here”.

Earlier, Whyte, Craig and Macoun (1897) urged
the preservation of our wild flowers because many
populations were disappearing, including “C. acaule
[which] is not to be found in Dow’s Swamp, where it
was formerly abundant ...”.

ACYANIC FORM: Cypripedium acaule f. albiflorum
Rand and Redfield, which is quite common in New
Brunswick (Hinds 1986), Nova Scotia (Roland and
Smith 1969), New Hampshire (Brackley 1985) and
Quebec (Ed Greenwood, personal communication
1996) but uncommon elsewhere, has been collected,
photographed or reported at least nine times in the
Ottawa District (A. H. Reddoch and J. M. Reddoch
1987b). From one to a dozen such plants occur,
either as individuals or in clumps, within larger
colonies of typical plants. The white-flowered plants
persist for several years. One colony, in Huntley
Township west of Stittsville growing in an area 10 m
across on sand under Eastern White Pine, had sever-

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 55

FicurE 10. Cypripedium acaule, mixed forest on sandstone, Stony Swamp Conservation Area, Regional Municipality of
Ottawa-Carleton, Ontario, 15 June 1980.

56 THE CANADIAN FIELD-NATURALIST

al of these flowers, along with typical ones, for at
least 13 years. In the Larose Forest, we have seen
one or a few white flowers at various times over a
period of 30 years, but these random sightings have
involved different plants.

These striking flowers, lacking the magenta antho-
cyanin, have a lip that is pure white, except for pale
green outside near the ovary, and a variable mixture
of green and yellow on the sepals, petals and stamin-
ode. Close inspection may show a faint reddish blush
on the staminode, ovary and adjacent parts that aris-
es from a pigment on the tips of the glandular hairs.
As discussed in the Introduction, the green and yel-
low colours seen in the acyanic form combine with
anthocyanin to produce the characteristic brown or
greyish magenta hues of the normal flower.

ABERRATIONS: C. acaule seems to have more struc-
tural aberrations than the other Lady’s-slippers. In
one plant that might be called scapeless, the lip,
tepals and floral bract all pointed upwards at ground
level from between a pair of leaves no larger than
these floral parts. Another short plant, in which the
flower had aborted at an early stage, produced a
scape about two cm long, so that the green floral
bract was leaning on one of the leaves.

One collection from the Ottawa District [Senn
1463 at NA] is of a plant with two flowers on sepa-
rate scapes. A different type of two-flowered plant,
seen in the field, had the flowers on a single scape,
the lips being joined back-to-back in a manner simi-
lar to that reported from New Hampshire (Brackley
1985).

Vol. 111

We observed a different type of aberration in a
plant bearing one extra tepal. This was outside a
petal and perhaps partly fused to it. It was boat
shaped, about the size of the lip, pink veined on the
outside and mostly white on the inside, looking
somewhat like a half lip. On another plant, a more
deformed flower with a possibly related aberration
had only half a lip and it was fused to the opposite
petal. It also had one normal petal and normal sepals.
The column was outside the twisted lip-petal and had
only the stigmatic surface and one pollinium, lacking
the staminode.

DEVELOPMENT: As the buds begin to open, the scape
is still arched over with the flowers pendent. The lips
are almost white, suggesting f. albiflorum, but the
colour intensifies as the flowers develop.

In large colonies, some 20% to 35% of the plants
are flowering, 35% to 65% are non-flowering with
two leaves and 15% to 40% are non-flowering with
only one leaf.

Some of the variation in the height of flowering
plants arises from the continuing growth of the
scapes during anthesis, about five mm per day. This
growth can be seen in the way that the previous
year’s capsules tower over the flowering plants until
near the end of the flowering period.

DERMATITIS: The floral bract and perhaps other
parts of this Cypripedium occasionally cause allergic
reactions in susceptible people (Reddoch and
Reddoch 1984), but the effects are not as severe as
from C. parviflorum and C. reginae.

1997

Cypripedium arietinum R. Brown

Ram’s-head Lady’s-slipper

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 57

Cypripéde téte-de-bélier

Cypripedium arietinum is the smallest, least conspicuous, least common and hence least familiar of our Lady’s-
slippers. When in bloom, it is easily identified by its small, purplish flower with its uniquely-shaped lip. In con-
trast to our other Lady’s-slippers, the lateral sepals are not united. The leaves differ from those of the other
species in being relatively small and narrow and in being arranged in a rough spiral about the stem.

DESCRIPTION

Height: 11 (16 - 26) 34 cm [224 plants].

Flower: 1, rarely 2; lip a small conical cup, the
opening formed by a circular inrolled edge sur-
rounded by numerous white bristles, with several
transparent windows in sides, essentially white with
pale yellowish green or greyish magenta at the apex
and greyish magenta, deep magenta or greyish ruby
as stripes centred on a set of vertical veins and irreg-
ular cross veins on the front and sides, and as nar-
row, vertical, parallel, equally spaced lines on the
back; staminode convex above with a marked cen-
tral, vertical groove; lateral sepals and petals nar-
row, reddish or violet brown, occasionally as stripes
on a yellowish green background; in f. albiflorum
(see Acyanic Form below): lip white with yellowish

46°

green on basal apex, sepals and petals yellowish
green; fragrance sweet, floral or vanilla-like, of
quite variable intensity.

Leaves: 3 - 4, deep green (but not, as described
by some authors, bluish green); 3 - 4 on non-flower-
ing plants.

Overwintering State: one or more, pale green,
conical shoots, appearing at ground level beside the
current year’s stem in early October.

Capsule: brown, ellipsoid, typically 1.7 x 0.7 cm,
ascending (see Figure 1a; unfertilized flowers dehis-
cent); yields from 30 to 90%, the higher values
associated with more open habitats [760 plants];
some capsules attacked by weevils (Stethobaris
ovata (LeConte) (Howden 1995)) before seed
release.

45°
76° 30'

75°

Cypripedium arietinum: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. Major areas underlain by calcareous rock (marble and lime-

stone) are shaded.

58 THE CANADIAN FIELD-NATURALIST

Seeds: greyish orange, beginning to release in
early to mid October.

BLOOMING PERIOD: 16 May (24 May - 7 June) 16
June [48 records].

CoLony S1zEs: | - 1000 plants, typically to 120,
flowering and non-flowering plants [45 colonies],
occurring as scattered individuals and in clumps of
up to 6 stems; wetland colonies restricted to a few
scattered individuals; about 50% of plants bearing
flowers [800 plants].

CURRENT STATUS: rare to uncommon (S3) in
Ontario (Active List, Oldham 1996’), rare in the
Province of Quebec (Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the range of this mainly Great Lakes - St.
Lawrence orchid of the Mixed Forest Region.

In the District, this species is confined to regions
underlain by limestone on the Lowlands and by mar-
ble on the Shield. Cypripedium arietinum is known
only west of the Rideau and Gatineau Rivers, with
particular concentrations in the southwest where the
limestone of the Smith Falls Limestone Plain
(Chapman and Putnam 1984) is close to the surface.

HABITATS: Cypripedium arietinum occurs in two
different types of habitats, one dry to moist and the
other moist to wet.

The first habitat is provided by maturing conifer-
ous forests in areas of shallow soil over flat-lying
calcareous rocks, including alvars (Catling and
Brownell 1995). Eastern White Cedar is usually the
dominant tree, often accompanied by White Spruce
and Balsam Fir. Trembling Aspen and other decidu-
ous trees are sometimes present. Plants grow in the
dappled to partial shade of small to large openings in
the forest canopy. Ground cover is usually sparse
and often includes Poison Ivy (Rhus radicans) and
Gaywings (Polygala paucifolia), the latter in flower
at the same time as the orchid. Cypripedium parvi-
florum var. pubescens is a constant companion in
more open sites.

A few scattered plants occasionally are found in
the moist to wet conditions of semi-open Eastern
White Cedar - Black Spruce - Tamarack fens and the
edges of cedar swamps. Plants sometimes are
drowned by rising water levels in the latter habitat.

LONG-LIVED COLONIES: Some locations are known
to have provided stable habitats for C. arietinum for
many decades.

Chilcott’s Swamp, near Alcove, Quebec (Fyles
1912), has been a known location for C. arietinum at
least since John Macoun collected it there in 1898
[TRT 15391]. In 1992, André Sabourin (personal
communication) found 22 plants there.

Near Braeside, Ontario, the dry, sandy, shallow
soil under White Spruce and Eastern White Cedar
supports thousands of plants over a wide area of the

Vol: ti

limestone plain (Adolf Vogg, personal communica-
tion 1994). Charles Macnamara (circa 1940°)
described the orchids and their habitat there in the
early years of this century. This spectacular alvar
population is currently threatened by estate lot devel-
opment.

In The Burnt Lands Alvar, several hundred plants
of C. arietinum and about half as many of C. parvi-
florum var. pubescens have flourished since at least
1968. The plants grow in shallow, clayey soil (pH
7.5) over limestone bedrock in open, partially shaded
clearings among widely-spaced Eastern White
Cedar, White Spruce, Balsam Fir, Trembling Aspen
and Balsam Poplar. The diverse but sparse ground
cover includes some grasses and Wood Lily (Lilium
philadelphicum), Wild Lily-of-the-valley (Maian-
themum canadense), Star-flowered False Solomon’s
Seal (Smilacina stellata), Malaxis unifolia,
Spiranthes lacera, Barren-strawberry (Waldsteinia
fragarioides), Cooper’s Milk-vetch (Astragalus
neglectus), Gaywings, Poison Ivy, Bearberry
(Arctostaphylos uva-ursi), Spurred-gentian (Halenia
deflexa) and Large-leaved Aster (Aster
macrophyllus).

The Marlborough Forest shelters several large
colonies of C. arietinum. In one colony that we first
encountered in 1981, the plants are clustered in scat-
tered glades of dappled sun beneath mature Eastern
White Cedar with White Spruce and Balsam Fir.
Ivory Sedge (Carex eburnea) and Wild Lily-of-the-
valley often accompany C. arietinum in the black,
moist needle-mould of this more shaded habitat.
Clear-cutting along the southern edge of this colony
in 1995 wiped out all of the plants in that section. In
1996, Don Cuddy (personal communication) and
members of the Ontario Ministry of Natural
Resources’ Environmental Youth Corps Program
sampled the population and estimated a concentra-
tion of 1375 plants per hectare in the vicinity of this
colony. They estimated that there are about 750 ha of
suitable habitat on public land in the forest. This may
be the highest concentration of this species on public
land in Ontario.

EARLY HIStory: Cypripedium arietinum was added
to the Flora Ottawaensis in 1882 when it was dis-
covered in Dow’s Swamp “in great profusion within
a limited area” (Fletcher 1883; Whyte and Small
1883). James Fletcher made collections on 12 June
of that year [MTMG 25532 and US 27620].
Although recognized as very rare in the District
(Fletcher 1896), this orchid was collected a number
of times at the same place between 1883 and 1906
(see, for example, Fletcher, Scott, and Cowley
1892).

Until the 1950s, this species was known only from
Dow’s Swamp, Chilcott’s Swamp and from near
Aylmer, Quebec, as well as at Braeside (see Long-
lived Colonies above). In 1952, several colonies

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 59

FiGurE 11. Cypripedium arietinum, mesic cedar forest habitat, City of Nepean, Regional Municipality of Ottawa-Carleton,
Ontario, 31 May 1971.

60 THE CANADIAN FIELD-NATURALIST

were found in open, mixed woods and adjacent cedar
woods near Fitzroy Harbour, Ontario (Calder 1952).

ACYANIC ForM: One plant of Cypripedium ariet-
inum f. albiflorum House was photographed by
Michael Runtz (1994; personal communication
1995) near Braeside, Ontario, in the Study Area
beyond the 50-km circle. The description above is
based on this photograph.

Vol. 111

ABERRATIONS: Only one two-flowered plant has
been seen in the District. One other plant had a
“Siamese twin” flower that had two fused lips, one
column, four sepals (9 mm long), three petals (2, 4
and 6 mm long) and a large floral bract (6 cm long).

DERMATITIS: Contrary to our expectations (Reddoch
and Reddoch 1984), we have found no evidence for
dermatitis from this species.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 61

Cypripedium parviflorum Salisbury var. pubescens (Willdenow) Knight
Cypripedium parviflorum Salisbury var. makasin (Farwell) Sheviak

Yellow Lady’s-slipper Cypripede jaune
SYNONYMS: Cypripedium calceolus Linnaeus var. pubescens (Willdenow) Correll, C. calceolus Linnaeus var.
parviflorum (Salisbury) Fernald, C. pubescens Willdenow.

This is one of our most distinctive and abundant orchids, occurring in a variety of calcareous habitats and show-
ing considerable variation in flower size and colour. It can be recognized in flower by its yellow lip and other-
wise by its cauline leaves that are two-ranked rather than spiralled about the stem.

TAXONOMY: We follow the concepts of Sheviak (1993, 1994, 1995), who recognizes the North American C.
parviflorum as a species distinct from the European C. calceolus. He proposes two varieties in the northern
part of the range, the highly variable var. pubescens (Willdenow) Knight and the small-flowered var. makasin
(Farwell) Sheviak. He distinguishes them by differences in flower size, colour distribution on sepals and
petals, fragrance and amount of pubescence on the upper sheathing bract (Sheviak 1994).

The majority of the plants in the District are referable to var. pubescens and it is these plants that we
describe below. Some of the plants in a few cedar swamps and fen edges that have small flowers with very

dark, uniformly coloured sepals and petals, and intense, sweet fragrance may be referable to var. makasin.

DESCRIPTION

Height: 12 (23 - 41) 57 cm [328 records].

Flowers: 1, occasionally 2 (one above the other
with the lower having a larger floral bract); lip 2.6 -
4.5 cm long, yellow (sometimes light, vivid or dark
yellow) with red or brownish red spots in rows with-
in and sometimes irregularly along edge of orifice;
sepals and petals ranging from greyish or yellowish
green with narrow reddish brown lines along veins,
through reddish, dark or violet brown as rows of dots
or lines of variable width to a nearly uniform field of
these colours; lateral sepals united with distinctly
split apex; petals usually corkscrewed with 3 - 5 half
turns, less commonly 0 - 2 or with a wavy edge; sta-
minode similar in colour to lip or slightly more
intense, with reddish spots usually in two rows near
edges but occasionally irregular or as lines, and often
with a green or deep green line along centre line; fra-
grance variable in character and intensity: rose, rasp-
berry, strawberry, lilac or vanilla.

Leaves: 3 - 4, less commonly 5, two-ranked, the
lowest often the smallest; 3 - 4 on non-flowering
plants.

Overwintering State: one or more, greyish green,
broadly conical shoots, 0 - 2 cm above ground beside
the current year’s stem, appearing in late September,
or pale green, remaining below ground level until
spring.

Capsules: brownish orange to light brown, ellip-
soid, typically 2.6 x 1 cm to 3.7 x 2 cm, ascending;
yield highly variable from colony to colony and year
to year from very small to about 50%, rarely to 80%;
32% and 9% in 1990 and 1991, respectively, in a
Gatineau Park study (Tremblay 1994);

Seeds: light brown, released in early to mid
October.

BLOOMING PERIOD: 9 May (27 May - 18 June) 6
July [142 records].

COLony SIZEs: | - 835, typically to 125, flowering
and non-flowering plants [257 colonies]; singly and
in clumps of up to about 20 stems; one clump just
south of Stony Swamp Conservation Area about 80
cm across with about 75 flowering stems in 1969,
declining to about 30 flowering stems four years
later.

CURRENT STATUS: one of the most abundant orchids
in the District.

DISTRIBUTION: The Ottawa District is in the north-
eastern part of the continental distribution. This is an
orchid of the Montane, Boreal, Mixed and
Deciduous Forest Regions and adjacent northern
Prairie.

Within the District, C. parviflorum is widely scat-
tered across the Canadian Shield and more concen-
trated on the Lowlands west of the Rideau River. It
occurs in areas underlain by Precambrian marble and
other calcareous rocks.

HABITATS: Cypripedium parviflorum occupies a
wide range of habitats in calcareous regions. In wet-
lands, it commonly occurs in wet-mesic, partially-
shaded openings and edges of coniferous and mixed
swamps. These swamps are treed with various com-
binations of Eastern White Cedar, Tamarack, White
Spruce, Black Spruce, Balsam Fir, Trembling Aspen,
Black Ash and Red Maple. The herbaceous layer
often includes Bulblet Fern (Cystopteris bulbifera),
Crested Wood Fern (Dryopteris cristata), Bluebead-
lily (Clintonia borealis), Wild Sarsaparilla (Aralia
nudicaulis), Bunchberry (Cornus canadensis) and
Starflower (Trientalis borealis).

62 THE CANADIAN FIELD-NATURALIST Voloht1

spa AAS Sogo plate CNT 5
a

FIGURE 12. Cypripedium parviflorum var. pubescens, alvar habitat, West Carleton Township, Regional Municipality of
Ottawa-Carleton, Ontario, 27 May 1979.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 63

46°

45°

76° 30

75°

Cypripedium parviflorum (both varieties): Hi = herbarium specimen, @ = Native Orchid
Location Survey sight record, A = literature reference. Major areas underlain by calcareous

rock (marble and limestone) are shaded.

Cypripedium parviflorum is also found at the
semi-open transition zone between open fen and
swamp among Eastern White Cedar, Black Spruce
and Tamarack and accompanied by typical fen vege-
tation. The sedge peat mat is usually consolidated;
rarely, it is floating (Adolf Vogg, personal communi-
cation 1988).

Mesic to dry woodlands are other common habi-
tats for C. parviflorum. These woodlands may be
coniferous, mixed or deciduous with almost any
combination of trees, but they have in common that
they are usually relatively open, either because they
are young to semi-mature or because they contain
large clearings. The soil may be rich with humus
(sometimes in rocky crevices) or shallow and clayey
over flat-lying bedrock. The herbaceous layer in rich
deciduous woods often includes Rattlesnake Fern
(Botrychium virginianum), Maidenhair Fern
(Adiantum pedatum), Wild Onion (Allium
canadense), Trout Lily (Erythronium americanum),
White Trillium (Trillium grandiflorum) and Bellwort
(Uvularia grandiflora).

Limestone plains and alvars are also typical habi-
tats for C. parviflorum. Plants grow in shallow soil

in the open sun or in the partial shade of conifers.
See The Burnt Lands Alvar in the Long-lived
Colonies section of the C. arietinum account for fur-
ther details of this habitat.

LONG-LIVED COLONIES: Cypripedium parviflorum
is a long-lived plant. Its colonies can persist for
decades wherever the habitat is stable. For example,
the colonies in Chilcott’s Swamp first recorded in
1911 (Fyles 1912, collections at DAO) were still
extant in 1992 (André Sabourin, personal communi-
cation 1995). In our own experience, colonies
observed in the late 1960s in The Burnt Lands Alvar
(see C. arietinum account) and in various swamps
continue to thrive.

EARLY History: The earliest known collections of
C. parviflorum were the three made in 1861 and
1863 at Ramsay, Ontario, by John Kerr McMorine
[QK 12719, 12726, 66622]. Another early collection
was that of Braddish Billings (1867) in 1866; its cur-
rent whereabouts is unknown. In 1870, Elizabeth
Keen White painted a plant gathered from “low
woods. Ottawa” by her husband, William White
(Dore 1965").

64 THE CANADIAN FIELD-NATURALIST

Collections in local herbaria show that in the early
days C. parviflorum was found close to the city in a
variety of rocky woods and swamps on both sides of
the Ottawa River. James Fletcher (1893) described it
as local but abundant. Fairy Lake and Beaver
Meadow (Fletcher, Small, and Baptie 1887a) on the
Quebec side, and Dow’s Swamp, MacKay’s Grove
and Billings Bridge on the Ontario side were well-
known localities. Lemieux Island and other rocky
areas above and around Chaudiére Falls supported a
large population and some plants persisted on

Vol. 111

Lemieux Island until at least 1925 [DAO 627979]. In
1897, Whyte, Craig and Macoun noted that many
local populations were disappearing, the inevitable
result of the growth of the city.

DERMATITIS: This species can produce dermatitis
similar to that from Cypripedium reginae. (For a
review, see Hausen (1984).) In our trials, all pubes-
cent parts, flowers, leaves, stems and new shoots,
caused irritation within a few days (Reddoch and
Reddoch 1984).

1997

Cypripedium reginae Walter
Showy Lady’s-slipper

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 65

Cypripéde royal

SYNONYMS: Cypripedium spectabile Salisbury, C. hirsutum Miller

Cypripedium reginae is indeed a showy plant with its large, bright flowers and its frequently large colonies.
However, while its image is familiar, the living plant is perhaps less so because it tends to occur in relatively
inaccessible wetlands. It is readily recognized by its magenta and white-lipped flowers and large, pubescent,

cauline leaves.

DESCRIPTION

Height: 33 (40 - 62) 81 cm [40 plants].

Flowers: | - 2; large, lip white with greyish
magenta or purplish red of variable intensity, occa-
sionally very faint, on front, sides and top, often as
diffuse vertical bands separated by white lines
marking the veins, and rows of similarly coloured
spots on interior; staminode white with broad yel-
low bands along edges toward apex, with spots or
irregular lines, above and below, of orange brown or
darker, depending on the anthocyanin concentration,
superimposed on the yellow bands; sepals and petals

white, with some long pubescence at base of petals;
fragrance delicate, sweet, floral, rose or raspberry-
like.

Leaves: 4 - 7, large, pubescent, strongly ribbed, in
a spiral around the stem; 4 - 7 on non-flowering
plants.

Overwintering State: one or more greyish green
shoots, 8 mm in diameter, about 6 cm down in the
moss substrate beside the current year’s stem, by
early fall (see illustration of rhizome and buds in
Whitlow (1983)).

Capsules: light brown, ellipsoid, typically 3.6 x

46°

45°
76° 30!

75°

Cypripedium reginae: M@ = herbarium specimen, @ = Native Orchid Location Survey sight
record, & = literature reference. Major areas underlain by calcareous rock (marble and lime-

stone) are shaded.

66 THE CANADIAN FIELD-NATURALIST

1.2 cm, ascending and erect, often with remains of
flower, persisting for one or sometimes two years;
yield relatively low, averaging 25% [78 plants].

Seeds: greyish orange, released in mid to late
September, some three weeks earlier than our other
Cypripediums.

BLOOMING PERIOD: 27 May (14 June - 4 July) 14
July [64 records], the latest of our Cypripediums.

Colony Sizes: | - 1000, typically to 250, flowering
and non-flowering plants [128 colonies]; occurring
singly and in clumps.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District lies near the
northern boundary of the distribution of this essen-
tially Great Lakes - St. Lawrence - Atlantic Coast
orchid. It is a species of the Mixed Forest Region
and adjacent regions. Within the District, colonies
are scattered across those parts of the Lowlands and
the Shield where limestone and marble bedrocks are
close to the surface.

HABITATS: Calcareous fens and swamps are the most
common habitats for C. reginae. In these wetlands,
plants are scattered in and around openings and
edges where there is no more than partial shade.
Occasionally, a few plants live for a few years in
mesic forests adjacent to the wetlands. Cypripedium
parviflorum is a frequent companion in all habitats.

In fens, colonies grow in wet peat among Eastern
White Cedar, Black Spruce and Tamarack with the
usual fen mosses and vascular plants, most frequent-
ly with various sedges and Marsh Fern (Thelypteris
palustris) and Labrador Tea (Ledum groen-
landicum).

Swamps range in type from mostly coniferous:
Eastern White Cedar, White Spruce, Balsam Fir and
Tamarack, through various mixtures to mostly decid-
uous: Black Ash, White Elm and Red Maple.
Characteristic ground cover sharing the wet humus
or peat includes various calcareous swamp mosses,
Rattlesnake Fern (Botrychium virginianum),
Cinnamon Fern (Osmunda cinnamomea), Bulblet
Fern (Cystopteris bulbifera), Crested Wood Fern
(Dryopteris cristata), Oak Fern (Gymnocarpium dry-
opteris), Sensitive Fern (Onoclea sensibilis) and
Poison Ivy (Rhus radicans).

Ditches along railway lines through calcareous
swamps sometimes provide favourable environ-
ments for hundreds of C. reginae while the adja-
cent, more shaded swamps may contain only a few
plants.

LONG-LIVED COLONIES: Under favourable condi-
tions, colonies continue to bloom year after year, but
they disappear quickly when they become too shad-
ed or if the water regime changes.

Vol. 111

Purdon’s Fen, just outside the Study Area near
McDonald’s Corners, Ontario, is a well-known
place to enjoy thousands of Showy Lady’s-slippers
every June. The wetland property was purchased by
the Mississippi Valley Conservation Authority in
1984. Joe Purdon told us that in the 1930s he had
noticed that many new orchids appeared in a part of
the fen that had been logged a few years before,
while they were disappearing from more shaded
areas. After experimenting in a small area, he
adopted the practice of selective thinning of the
Eastern White Cedar trees and at the same time
controlling beaver activities to maintain a constant
water level. From a few dozen plants, the fen now
shelters an estimated 16 000 (Mosquin 1986°;
White 1988). This is the only place where we have
seen a three-flowered plant.

The colony at Chilcott’s Swamp described in 1893
by Whyte, Craig and Cowley (1893) was still extant
in 1992 (André Sabourin, personal communication
1995). (See also below.)

EARLY History: The first Ottawa District collec-
tion was likely made in 1866 by Braddish Billings
Jr. (1867). The whereabouts of this collection is
unknown, but he could have made it in Dow’s
Swamp, less than 2 km west of the family home at
Billings Bridge. A year later, John Kerr McMorine
collected this orchid “near Clayton, Ramsay” in
Lanark County at the western edge of the Study
Area [DAO 25692]. There are several extant collec-
tions taken from Dow’s Swamp between 1879 and
1909.

Chilcott’s Swamp was discovered in late 1892 and
the next year naturalists found it “to contain large
numbers of the Showy Ladies-slipper (Cypripedium
spectabile). The display of these lovely plants was
even more extensive than had been expected, and
charmed and delighted the party, who returned laden
with spoil” (Whyte, Craig, and Cowley 1893). In
1912, Faith Fyles related that “Lady Grey paid a visit
to the swamp when the Showy Lady’s Slippers were
in full bloom, and she sat [on a stump] to rest in the
midst of thousands of these beautiful pink and white
orchids”. (Alice, Lady Grey, was the wife of Albert
Edward, 4th Earl Grey, who was Canadian Governor
General from 1904 to 1911 (Hubbard 1977) and
patron of The Ottawa Field-Naturalists’ Club.)

James Fletcher (1893) considered this orchid to be
common. “This is probably our most beautiful wild
flower, but [it] is gradually disappearing from ruth-
less digging up of the roots”.

ABERRATION: The number of flowers examined is
fairly small and only one aberration has been noted
in our plants. This is a two-flowered plant recorded
in photographs (at DAO) by Beatrice Treichler near
White Lake in which one flower was normal while
the other lacked a lip and had a row of magenta spots
along the lower edge of one petal.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

vibes

ftir the
are

SS LOT oe Ie

3

poh iry
kan

a dice
IRN

aie

FIGURE 13. Cypripedium reginae, cedar swamp habitat, McNab Township, Renfrew County, Ontario, 12 June 1979.

67

68 THE CANADIAN FIELD-NATURALIST

DERMATITIS: Contact dermatitis caused by C. regi-
nae, although perhaps not widely known, has been
reported for over a century (see Reddoch and
Reddoch 1984; Hausen 1984). Each of us has had
quite marked reactions comparable to that from
Poison Ivy within two days of contact, accidentally

Vol. 111

as a result of measuring living plants and deliberate-
ly in tests to verify the source of the affliction. On
the other hand, Joe Purdon (see above), who handled
the plants on his property frequently over a period of
at least 50 years, told us that he had never had any
reaction to them.

1997

Epipactis helleborine (Linnaeus) Crantz

Broad-leaved Helleborine

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 69

Epipactis petit-hellébore

Epipactis helleborine, an Old World species, was collected in Ottawa half a century after its discovery at
Syracuse, New York, the earliest North American record. In the following 65 years, it has become the most
common orchid in the District. As in much of its range, it shows great variation in stature and flower colour. It
can be recognized by its several cauline leaves, the lowest of which is oval and much shorter than the one above
it, by its stem that is sometimes greyish magenta toward the base and by its green and greyish magenta flowers

with an open, cup-shaped hypochile (basal part of lip).

DESCRIPTION

Height: 10 (28 - 60) 102 cm [232 plants]; the rela-
tive variation of the height (the ratio of the standard
deviation of the height to the average height is 0.37)
is exceeded only by that of Platanthera hyperborea,
which has two habitat-dependent forms.

Flowers: 2 (6 - 32) 53 [103 plants]; colours show-
ing considerable variation from plant to plant,
reflecting differing amounts of chlorophyll and
anthocyanin, the latter also increasing as flowers
age; sepals greyish green or deep green sometimes
with some red; petals often greyish magenta, but also
pink, dark purple, etc. as well as white, violet white
and pale green; epichile (apical part of lip) similar to

petals but usually somewhat lighter, e.g., white,
pink, greyish ruby; interior of hypochile cup usually
darker than petals, e.g., greyish magenta, dark ruby,
reddish brown, violet brown, but occasionally very
light, e.g., pale red (peach); often in a one-sided
inflorescence as flowers turn toward the light; droop-
ing with age; fragrance slightly sweet or lacking.

Leaves: 3 - 14 leaves and bracts, typically 3 - 8,
5 - 6 in 60% of plants [105 plants]; 2 - 5, rarely 1, on
non-flowering plants (Marilyn Light, personal com-
munication 1996).

Overwintering State: one to several white to
pinkish buds, 5 to 16 cm or more below ground,
rarely at ground level, beside the current year’s stem;

75°

Epipactis helleborine: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record. Major areas underlain by calcareous rock (marble and limestone) are shaded.

70 THE CANADIAN FIELD-NATURALIST Voll tt

FiGurE 14. Epipactis helleborine, edge of mixed forest, Gatineau Park, Quebec, 9 August 1980.

1997

Light and MacConaill (1991) observed at their study
site in lower Gatineau Park that the perennating bud
and flower primordia form one or more years before
they emerge; the shoots of the year appear above
ground in late May.

Capsules: brownish orange, ellipsoid to spheroid,
typically 1.1 x 0.7 cm to 1.9 x 1 cm, descending;
yield highly variable often approaching 100%, aver-
aging 80% [63 plants].

Seeds: light yellow to greyish orange, released in
late August to early October.

BLOOMING PERIOD: 18 June (15 July - 12 August)
29 August and 15 September [80 records].

Co.Lony SIZEs: | - 1000, typically to 300, flowering
plants [191 colonies], as scattered individuals and
occasionally in clumps of up to a dozen stems.

CURRENT STATUS: the only orchid that can be con-
sidered common in the District, with many more
locations than the distribution map shows; this
species has become so abundant that people no
longer record its presence.

DISTRIBUTION: The Ottawa District is in the northern
part of the North American range of this introduced
orchid. It is spreading slowly through the Mixed
Forest Region and in other, scattered parts of the
continent (Soper and Murray 1985; Homoya 1993).
Epipactis helleborine is widespread and abundant
throughout the District, especially in areas of cal-
careous bedrock.

HABITATs: This orchid is found most often in the
moderate to deep shade of mesic, deciduous and
mixed forests; however, it occurs in most types of
forests from young to mature and from deciduous to
coniferous. Occasionally, plants also grow in the
wet-mesic soils of swamps and forested stream
edges. These treed habitats are usually relatively
open at ground level with little other vegetation near
the plants except mosses.

More open environments for E. helleborine are
forest edges and tracks, and, increasingly in recent
years, old-fields, suburban lawns and gardens. (See
Dore (1968, 1977, 1986) for garden occurrences and
persistence.)

The plants are rooted in soils that range in compo-
sition from pure clays and sands to the corresponding
loams; in swamps, the substrate is usually organic.

Loca History: Epipactis helleborine has been in
the Ottawa District since at least 1930 when Hoyes
Lloyd collected it on his property in Rockcliffe Park
Village [TRT 202651] (Malte 1933). This 0.6 ha
property at 285 Mariposa Avenue supported a mesic,
mature Sugar Maple forest with a rich spring flora
(Elizabeth Lloyd, personal communication 1981)
until it was subdivided after Mr. Lloyd’s death in
1978. Remnants remain to the present.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 71

Herbarium collections and field observations sug-
gest that E. helleborine did not begin to be common
in the District until the 1960s (John Arnold, personal
communication 1977; Dore 1968, 1977). The popu-
lation seems to have established itself in two stages,
the first being the appearance of a few individuals
here and there on both the Lowlands and the Shield,
and the second being the significant increase in the
number of plants at sites already colonized, as well
as the rapid emergence of new colonies. This latter
process is continuing.

It is possible that the Ottawa population came up
the Ottawa Valley from Montreal, 160 km to the
east, where the species was first collected in 1892
and which is sometimes considered one point of
introduction of the species in North America (Doyen
and Cayouette 1966). However, it should be noted
that Syracuse, New York, where the species was first
discovered on this continent in 1879 (Correll 1950;
Brackley 1985), is only 250 km to the south of
Ottawa.

ACHLOROPHYLLOUS FORM: Two small plants of E.
helleborine f. monotropoides (Mousley) Scoggan,
were discovered by Bob Bracken in Stony Swamp in
1979 (A. H. Reddoch and J. M. Reddoch 1987b).
The plants were about 15 cm tall and had 3 and 4
buds. We could see no evidence of chlorophyll in
them; the leaves were pale grey. The lower parts of
the stems were reddish lilac, showing the presence of
anthocyanins that sometimes occurs in normal plant
stems. The buds were light yellow. When the flowers
opened, the lips were white except for the inner sur-
face of the cup, which was light brown. These two
plants withered during the flowering period and did
not reappear the following year. They may have
been under stress since they grew in quite thin soil
over sandstone. Salmia (1986) reported that 70% of
a colony of achlorophyllous plants in Finland
appeared for only one year, while 27% appeared for
two years and 4% for three years. This pattern is
similar to that reported by Light and MacConaill for
normal plants. (See Population Studies below.)

ABERRATION: One flower on an otherwise normal
plant was partially doubled while retaining bilateral
symmetry. It had four sepals, the extra one being
opposite the dorsal sepal. It had two petals and a sin-
gle broadened column, below which were two com-
plete lips radiating away from the column.

INSECT PREDATION: We have seen a few examples
of E. helleborine infested with weevils, identified by
Anne Howden (personal communication 1995) as
Stethobaris ovata (LeConte), an insect found on sev-
eral local orchid species (Howden 1995). They were
clustered in the inflorescence feeding on the rachis
and inhibiting the development of the inflorescence.

POPULATION STUDIES: For more than a decade,
Marilyn Light and Michael MacConaill (1990, 1991,

72 THE CANADIAN FIELD-NATURALIST

1994) have studied a large colony of E. helleborine
in Gatineau Park and have reported on patterns of
appearance and correlations with variations of cli-
mate (as well as below-ground changes; see
Overwintering State above). Over a six year period,
they found that only two plants appeared every year,
while two-thirds appeared only once in that time.
The rhizomes of half of the dormant plants remained
alive and three plants reappeared after a three-year
absence. They also determined that the appearance
of plants is strongly correlated with the amount of
rainfall in the previous August and September and
that plants growing in the open are more likely to
reappear than those in shaded areas.

MyYcorRuiZa: In 1979, W. I. Illman collected a
plant from beneath an old Red Oak on his lawn in

Vol. 111

southeast Ottawa [DAO 124498]. He appended a
discussion of a fungus surrounding the roots and of
fungal hyphae that had invaded some cortical cells,
implying a mycorrhizal association. The fungus had
been identified by J. Bissett as Trichoderma hama-
tum (Bon) Bain. Because this common and readily
apparent soil fungus is not known to form mycor-
rhizae, it seems likely that the hyphae in the root
cells came from some other, less easily detected
fungal species (Jim Ginns, personal communication
1996). W. G. Dore (1986) refers to the above collec-
tion and to a plant on his own lawn, noting that fine
roots from nearby trees entered the mass of the soil
fungus surrounding the E. helleborine roots but that
they did not seem to have any physiological connec-
tion. Thus the nature of mycorrhizal activity in
mature plants in the District remains unclear.

107

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 73

Galearis spectabilis (Linnaeus) Rafinesque

Showy Orchis

SYNONYM: Orchis spectabilis Linnaeus

Galéaris remarquable

Galearis spectabilis, as part of the spring woodland flora, is one of the earliest orchids to bloom in the District.
It can occur in sizable patches and, although not particularly showy from a distance, has attractive flowers. It
can be recognized by the two oval basal leaves and the inflorescence of several lilac-coloured flowers with usu-

ally whitish lips.

DESCRIPTION

Height: 5 (9 - 15) 21 cm [269 plants]; appreciably
shorter than the 35 cm reported (Correll 1950, Luer
1975) for plants presumably from the southern part
of the range, but quite comparable to heights from
the northern part (Case 1987; Smith 1993).

Flowers: | (2 - 5) 7 [303 plants], about half as
many as reported on southern plants (Luer 1975);
sepals and petals typically light lilac, ranging from
violet white and pale violet to lilac; lip white, yel-
lowish white or yellowish grey or like sepals (see
Colour Pattern Variant below); column darker and
sometimes redder than sepals, thus purple, greyish
magenta or greyish rose; spur usually translucent

OG: aan

45° 4
76° 30'

white but occasionally with traces of lilac; lateral
sepals connivent with petals and dorsal sepal to form
a galea, but rarely free and spreading; flowers in a
somewhat one-sided and irregular inflorescence; fra-
grance moderate, floral or sometimes undetectable.

Leaves: 2, basal, with occasionally one large
cauline bract on flowering plants; 1 or, more com-
monly, 2 leaves on non-flowering plants.

Overwintering State: one or more greyish green,
broadly conical shoots, about 1 cm above ground,
appearing beside the current year’s stem in late
September; herbarium specimens show the new
shoot bud and partially elongated roots already pre-
sent at anthesis.

75°

Galearis spectabilis: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

74 THE CANADIAN FIELD-NATURALIST

Capsules: light brown, ellipsoid, typically 1.9 x
0.5 cm, erect or nearly so; yield usually less than
60%, averaging 45% [32 plants], appreciably higher
than the 5% yield reported for Ohio (Deringer 1982).

Seeds: dark brown, released in early October.

BLOOMING PERIOD: 15 May (23 May - 6 June) 15
June [72 records].

CoLony SIZEs: 1 - 278, typically to 100, flowering
and non-flowering plants [55 colonies], generally as
small to large patches; non-flowering plants some-
what more abundant than flowering plants.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District is close to the
northern limit of distribution of this eastern orchid.
It is a species of the Deciduous Forest Region and
adjacent Mixed Forest Region.

Within the District, it occurs in scattered locations
up to the southern part of the Canadian Shield north
of the Ottawa River. In addition to the colonies
mapped, Bob Bracken (personal communication
1989) encountered additional colonies in a number
of farm woodlots on the Lowlands during a bird pop-
ulation survey.

Hasirats: This orchid is a plant of relatively open,
deciduous or sometimes mixed forests that are semi-
mature or mature. Sugar Maple is almost always the
dominant tree, accompanied by a variety of other
species. Ground cover is usually sparse but may
include other spring wildflowers such as White
Trillium (Trillium grandiflorum), as well as
Maidenhair Fern (Adiantum pedatum), Christmas
Fern (Polystichum acrostichoides), Wild Onion
(Allium canadense) and Sugar Maple shoots. In the
forest, the orchids grow in mesic or wet-mesic sandy
loam or clay loam on the level floor or in depres-
sions, seasonal runoff areas and at the bases of
slopes. This species occurs over various bedrock
types.

At two localities, we have seen a few plants in
more open, drier situations beyond woodland edges,
close to and under clumps of Eastern White Cedar at
one site and of Juniper (Juniperus communis) at the
other.

LONG-LIVED COLONIES: Colonies of G. spectabilis
can persist and multiply for many decades if the site
does not become heavily shaded or overgrown with
vegetation such as maple shoots.

We have followed a colony in lower Gatineau
Park from which W. G. Dore made a collection in
1969 [DAO 267276]. The number of plants has
increased from 65 in 1973 to 173 by 1994. In 1973,
40% of the plants flowered, while in 1994, 46%
flowered. About one-third of the plants produce
flowers with coloured lips. (See Colour Pattern
Varient below.)

Vol. 111

The colony is situated in wet-mesic clay loam at
the base of a steep, rocky, southwest-facing slope
under a high, almost closed canopy. Sugar Maple is
the dominant tree, accompanied by single trees of
Yellow Birch, White Birch, Basswood, White Ash
and Red Oak, and single saplings of Eastern
Hemlock, Balsam Fir and Beech. The sparse herba-
ceous layer consists of a few plants of White Trillium
and Early Meadow-rue (Thalictrum dioicum), as well
as many Sugar Maple seedlings. The bedrock, much
in evidence on the slope above, is marble.

Galearis spectabilis persisted in at least two sub-
urban woodlots until two decades ago. We last saw a
few plants in Pleasant Park Woods in Alta Vista
(Reddoch 1972) in 1971 and 18 plants in Niven’s
Woods in Rothwell Heights (Reddoch 1980, [DAO
691465]) in 1980. The former colony probably died
out naturally with increased shade, but the latter,
while decreasing, was destroyed by housing devel-
opment. It had been followed by Anne Hanes since
the early 1960s. Both these woodlots were seven
hectares in area.

EARLY History: Galearis spectabilis is one of the
seven orchids that Braddish Billings Jr. (1867)
included in his list of species collected in the sum-
mer of 1866, presumably in the vicinity of his family
home at Billings Bridge. In 1869, Elizabeth Keen
White painted a plant provided by her husband,
William White, from “Rich woods. Ottawa” (Dore
1965’).

Macoun (circa 1911") cited an 1878 collection by
R. B. Whyte from “woods east of Bank St. road near
Ottawa”. James Fletcher collected G. spectabilis in
1878 and 1879 at Beaver Meadow, Hull [DAO
267272], the Chelsea Mountains (now Gatineau
Park) [DAO 267273] and Patterson Creek Wood (in
the Glebe in Ottawa) [DAO 17040]. In 1879, Henry
M. Ami collected this orchid at MacKay’s Grove
[CAN 232401].

Galearis spectabilis was specifically mentioned in
accounts of Ottawa Field-Naturalists’ Club excur-
sions to New Edinburgh and Hemlock (McKay)
Lake (Fletcher, Small, and Baptie 1887b; Fletcher
1898), Beaver Meadow, Hull (Cowley, Macoun, and
Whyte 1892; Eifrig 1909), Ironsides, Quebec
(Halkett 1909) and Gilmour’s Grove (Clarke 1904,
1908). Eifrig (1909) remarked that the west bank of
Beaver Meadow was “perhaps the most prolific
place near Ottawa for the botanist, rare plants like
the showy orchis (Orchis spectabilis) being found
there, as well as other kinds of commoner ones in
profusion...”. By 1913, Beaver Meadow was being
destroyed by residential development (Anonymous
1913).

For his Beaver Meadow and Patterson Creek
Wood collections, James Fletcher noted that the
flowers were entirely the same colour, light mauve at
the first site and deep purple at the second. Similarly,

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT Wis)

FicurE 15. Galearis spectabilis, mixed forest habitat, Gatineau Park, Quebec, 10 June 1970 (plant) and 25 May 1979
(column); for the column (front view), scale bar = 0.5 mm.

76 THE CANADIAN FIELD-NATURALIST

Clarke (1904) described the plants found on an
excursion at Gilmour’s Grove as having completely
purple flowers. (See Colour Pattern Variant below.)

The photograph of this species that Charles
Macnamara took in the Arnprior area on 7 June 1913
is shown in his undated manuscript (circa 1940")
and in Reddoch (1981c). This photograph, credited
only to the American Orchid Society, is included in
the book, A History of the Orchid (Reinikka 1972).
Galearis spectabilis is considered to be rare in the
Arnprior area (Runtz 1984).

COLOUR PATTERN VARIANT: Galearis spectabilis f.
willeyi (Seymour) P. M. Brown has been seen in a
. number of colonies in both the Ontario and Quebec
parts of the District (A. H. Reddoch and J. M.
Reddoch 1987b). Some colonies are more prone

Vol. 111

than others to produce plants with coloured lips.
Although Seymour (1970) reported that Willey had
observed the colours to be constant for several years,
Marilyn Light (personal communication 1996) has
determined that the effect is not reproducible from
year to year. The lip colour of this putative form is
similar to that of the sepals but can be variable in
intensity and irregular in distribution. It is often lilac
but can be a darker shade, such as purple, greyish
violet or deep violet, or a fairly light one, such as
light lilac or pale violet. The colour may be uniform
across the lip or, when very light, merely a fringe
near the edge of the lip. This form may be an exam-
ple of the colour pattern variation arising from
genetic mutation (Fincham 1987; Jorgensen 1995) or
it may arise from environmental factors.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 77

Goodyera pubescens (Willdenow) R. Brown in Aiton

Downy Rattlesnake-plantain

Goodyérie pubescente

Goodyera pubescens is the largest of the three Goodyeras in the District. It is a species of mature, mesic forests
and swamps that can form large patches by vegetative reproduction. These patches can be one or two metres
across and can contain 100 to 200 plants. It can be recognized by its basal rosette of tessellated leaves with
broad light green to nearly white lines bordering the median vein. It can be distinguished from G. repens by its
cylindrical inflorescence. The most definitive character in separating G. repens and G. tesselata, the rostellar
beak on the column, is essentially absent in this species (Whiting and Catling 1986).

DESCRIPTION

Height: 14 (19 - 27) 36 cm [164 plants].

Flowers: 7 (15 - 36) 64 [104 plants]; white, fre-
quently with light green along veins of sepals and
sometimes on tip of dorsal sepal; fragrance faintly
sweet or spicy.

Leaves: 2 - 15, typically 2 - 11 [99 plants]; 2 - 4.5
cm long, 1.2 - 2.2 cm wide [98 leaves]; greyish
green or dark green with light green to nearly white
markings on upper surface only (see Figure).

Overwintering State: as a rosette of leaves (see
Vegetative Reproduction and Flowering Frequency
below).

Capsules: light brown, spheroid, typically 0.6 x

46°

45° |
76°.3

0.4 cm, ascending; yield often either 0% or ap-
proaching 100%, averaging 45% [57 plants].
Seeds: dark brown, released in mid October.

BLOOMING PERIOD: 26 July (4 August - 1
September) 20 September and 20 October [20
records].

Co.ony SIZEs: 2 - 321, typically to 250, flowering
and non-flowering rosettes [21 colonies], as small
groups or as patches of up to 200 rosettes.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District is close to the

75°

Goodyera pubescens: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

78 THE CANADIAN FIELD-NATURALIST

northern edge of the distribution of this eastern
species of the Mixed and Deciduous Forest Regions.
Within the District, this orchid is thinly scattered
across the Shield over a variety of bedrock types. It
is represented on the Lowlands only by collections
from 1903 and 1945.

HABITATS: Goodyera pubescens is a plant of fairly
mature mesic forests and sometimes of swamps. In
all habitats, it inhabits areas of dappled to deep shade
with little other ground cover except mosses.

This orchid is most often found in deciduous or
mixed forests and only occasionally in dominantly
coniferous forests. It thrives in forests of Sugar
Maple with Beech or with conifers such as Eastern
White Pine, Eastern Hemlock, Balsam Fir and
Eastern White Cedar. (See Long-lived Colonies
below for additional details.) The plants are as likely
to be on a flat forest floor as on a slope beside a
stream or pond. There is typically little other vegeta-
tion sharing the usually deep, but sometimes shal-
low, sandy or sandy loam substrate. Some compan-
ion species are Bristly Clubmoss (Lycopodium
annotinum), Bluebead-lily (Clintonia borealis), Wild
Lily-of-the-valley (Maianthemum canadense),
Goldthread (Coptis trifolia) and Wild Sarsaparilla
(Aralia nudicaulis).

In the typical Ottawa District cedar swamps,
rosettes of G. pubescens occasionally line mossy,
rotting logs or grow on peaty mounds close to trees.
Eastern White Cedar is accompanied by such trees as
Black Ash, Yellow Birch, White Elm, Balsam Fir,
Red Maple and Largetooth Aspen.

LoNG-LIVED COLONIES: Since 1975, we have fol-
lowed a flourishing colony of G. pubescens in lower
Gatineau Park. (There is a herbarium specimen taken
by D. Erskine from this locality in 1968 [DAO
96646].) There are eight separate groups of plants
scattered over a distance of 200 metres and ranging
in size from a dense patch of 200 rosettes to a small
group of eight. The large groups were well-estab-
lished in 1975 while the smallest one is fairly recent.
The orchids grow in shallow or deep sand, on slopes
or on flat land, over bedrock of marble, calc gneiss
and granite pegmatite (Hogarth 1970). The largest
patches are in a dominantly Eastern Hemlock stand
with Red Oak and Beech, while other groups are
under Sugar Maple with White Pine, Eastern
Hemlock and Hop Hornbeam. Most plants are in
dappled shade, but it is clear at one location that
those plants in a small opening, and thus receiving
more light, are larger. However, another group in a
somewhat sunny location disappeared during the
course of our study. Burial of the plants by heavy
layers of fallen leaves compacted by winter snow
and deer browsing of the inflorescences are hazards
in this location.

Five km to the northwest, there is another colony
of G. pubescens that we have followed and pho-

Vol. 111

tographed from time to time since 1975. (The plant
illustrated in the Figure grew in this colony.) It is on
a north-facing slope of a small stream valley under a
canopy of Beech and Sugar Maple. The bedrock
below the sandy substrate is porphyroidal gneiss
(Hogarth 1970). The 15 rosettes present in 1975 had
multiplied to 66 in 1987 and currently (1996) num-
ber 86. The 65 cm x 40 cm area of the colony in
1975 has expanded to 135 cm x 100 cm in 1996.

Because the pattern on each leaf is different, we
have been able to trace the season by season devel-
opment of some of the rosettes in these colonies. Our
findings are reported under Vegetative Reproduction
and Flowering Frequency below.

EARLY History: Most early collections of G.
pubescens came from the lower Gatineau valley.
William Scott discovered the first colony in the
District in 1891 near Ironsides, Quebec (Fletcher,
Scott, and Cowley 1892). He collected several
specimens there in July and, with John Macoun, in
September of that year [DAO 96648, TRT 15483;
TRT 15481, 15488, CAN 116928]. Macoun (circa
1911°) described the locality as “in the pine woods
towards Chelsea beyond Ironsides”. He listed two
other collections of his, one “by the mountain on
the old mine road to Old Chelsea” in 1902 (current
whereabouts unknown) and “in woods near
Wakefield” in 1903 [TRT 15502]. Botanists found
additional colonies in the lower Gatineau valley
near Chelsea in 1906 and near Ironsides in 1918
(specimens at CAN). James Fletcher (1893) also
included Hull and Kingsmere as localities for this
species.

John Macoun collected both G. pubescens and G.
tesselata at Leonard, Ontario, in 1903 (specimens at
CAN). In his circa 1911° list of the Ottawa flora,
Macoun stated that he made these collections, as
well as one of G. pubescens in 1911 (current where-
abouts unknown), “in thick woods at Leonard sta-
tion, C.P:R: Short Line’. He also cited an R. B:
Whyte collection of unknown date from “Eastman
Springs” (= Carlsbad Springs, Ontario).

Charles Macnamara (circa 1940°) included G.
pubescens in his treatment of the orchids of the
Arnprior area; however, the photograph that he used
to illustrate this species shows clearly that the plants
he described were G. tesselata. Goodyera pubescens
has not been found in the Arnprior area in more
recent years either (Runtz 1984, personal communi-
cation 1995).

VEGETATIVE REPRODUCTION AND FLOWERING
FREQUENCY: A plant, as the term is used in this sec-
tion, consists of a rhizome that is or has been
sheathed with leaves along its length and that termi-
nates in a rosette of leaves and a growing tip. (The
rhizome originated from a parent rhizome the year
after the parent plant flowered.) As the rhizome
grows, it puts down a few roots and the old leaves

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

4 Ss ea

aa i

FiuRE 16. Goodyera pubescens, deciduous forest habitat, Gatineau Park, Quebec, 25 August i OTeR

oe

80 THE CANADIAN FIELD-NATURALIST

die in turn, leaving rings around the rhizome to mark
where they were attached. These rings or leaf scars
are from 0.5 cm to 2.2 cm apart. The rhizome is usu-
ally on or just below the surface of the humus or soil
substrate; it may be straight or with bends where the
growing tip changed direction. When a plant flow-
ers, the inflorescence rises from the centre of the
rosette, first appearing in early June.

The year after a plant flowers, one to three, rarely
four, new growths emerge from the rhizome among
the older leaves of the plant and extend distances of
three to five cm from the parent. The parent plant
dies that year or the next. The weakest of the
growths often dies within a year. Each new growth
produces two leaves the first year; the maturing plant
generates three to five new leaves each year until the
year it flowers. (It does not produce any leaves the
year it flowers other than sometimes one or a few
narrow, basal-cauline leaves.) Each leaf lasts two to
four years, the lifetime being shorter for leaves heav-
ily covered with fallen tree leaves. (Keenan (1990)
reported a lifetime of four years in New Hampshire.)
By the time a plant flowers, after a minimum of
three years and typically five to eight years, it has
generated a total of 21 - 30 leaves, 8 - 15 of which
are still present at flowering. The growing tip is now
14 - 18 cm from the parent. The progeny of the same
parent do not all bloom the same year; the most
robust plant (generated more leaves, travelled far-
ther) flowers first and some plants do not flower at

Voktit

all. The qualitative features described here are simi-
lar to those reported by Ackerman (1975) for
Goodyera oblongifolia in California.

Usually only a few of the plants in a colony bloom
each year; the yearly average over 20 years for the
colonies described above under Long-lived Colonies
was 11%. However, the percentage of plants flower-
ing from year to year is not at all uniform; in some
years as many as 66% of the plants produce flowers,
while in other years none do. We have found that
there is a tendency for the colonies described above
to behave similarly, having simultaneous large, mod-
est or zero flowerings. There were large flowerings
in 1982, 1986, 1989, 1994 and 1996, and slight or
zero flowerings in 1977, 1983, 1985, 1987, 1990,
199, 1995, and 1395:

Of the four groups in the first colony described
above for which we have extensive records, three
showed intergroup correlations of annual flower pro-
duction that are statistically significant at the 1%
level or better. The fourth group, followed for 11
years, did not show high correlations with the others.
(Its growth cycle was frequently interrupted by bur-
ial in fallen leaves borne by drainage on a steep
slope.) These groups also correlated with the colony
5 km away with a significance level of 5% or better.
Such correlated, but irregular, flowering suggests the
influence of external factors, presumably the annual
weather patterns, perhaps with some local variation
in sensitivity to these factors.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 81

Goodyera repens (Linnaeus) R. Brown in Aiton var. ophioides Fernald

Lesser Rattlesnake-plantain

Goodyérie rampante

Goodyera repens is the smallest of the three Goodyeras in the District. It is a species of dense, shady forests and
swamps that can form small, compact patches by vegetative reproduction. It can be recognized by its basal
rosette of tessellated leaves lacking light lines along the median vein and can be distinguished from G. pubes-
cens by its essentially one-sided inflorescence. The most definitive, but not most convenient, character is the
rostellar beak length, which is less than 0.5 mm (Kallunki 1976).

DESCRIPTION

Height: 4 (7 - 14) 19 cm [67 plants].

Flowers: 1 (7 - 16) 28 [62 plants]; white, some-
times with a slight greenish tinge on the veins on the
outside of the sepals of the upper flowers; tending to
face the brightest available light in a one-sided inflo-
rescence.

Leaves: 2 - 7 [28 plants]; 1.4 - 2.8 cm long, 0.8 -
1.5 cm wide [17 leaves]; greyish green or dark green
with light green markings bordering the veins except
the median vein (see Figure); occasionally the lowest
cauline bract enlarged and leaf-like.

Overwintering State: as a rosette of leaves (veg-
etative reproduction apparently similar qualitatively
to G. tesselata).

Capsules: light brown, spheroid, typically 0.5 x

46°

0.3 cm, ascending (see Figure 1a); yield highly vari-
able but usually low, averaging 30% [44 plants].
Seeds: released in early October.

BLOOMING PERIOD: 7 July (20 July - 15 August) 27
August [19 records].

CoLony SIZEs: | - 208, typically to 70, flowering
and non-flowering rosettes [37 colonies], as scat-
tered individuals and small groups, occasionally in
patches of as many as 90 rosettes (see Long-lived
Colonies below).

DISTRIBUTION: The Ottawa District is well within
the range of this mainly eastern variety of G. repens
(Kallunki 1976). It occupies primarily the Mixed
Forest Region and adjacent Boreal Forest Region.
Within the District, this orchid is thinly scattered

45°
76° 30

fo:

Goodyera repens var. ophioides: HM = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. The Canadian Shield is shaded.

§2 THE CANADIAN FIELD-NATURALIST

across the Shield and rarely present on the
Lowlands.

HABITATS: Goodyera repens is a plant of dense,
shady, coniferous forests and swamps. It also occurs
occasionally in mixed forests and on forested slopes
adjacent to wetter areas.

The mesic coniferous forests that harbour G.
repens are composed of trees such as Eastern
Hemlock, Balsam Fir, Eastern White Cedar and
White Spruce. The plants grow among mosses or in
leaf mould on the otherwise relatively bare forest
floor. Sometimes they are accompanied by such
herbs as Goldthread (Coptis trifolia), Wild
Sarsaparilla (Aralia nudicaulis) and Bunchberry
(Cornus canadensis). Occasionally, Goodyera tesse-
lata is also present.

The coniferous swamps are generally in areas of
calcareous bedrock. They are usually dominated by
fairly mature Eastern White Cedar, accompanied by
a selection of other trees such as Black Spruce,
White Spruce, Tamarack and Black Ash. The trees
are often well-spaced with little under-story.
Goodyera repens plants are rooted in the moist to
saturated humus of the swamp floor and on mounds
around trees, among leaf litter or mosses.
Accompanying herbs may include Corallorhiza trifi-
da, Cypripedium acaule, Malaxis monophylla,
Platanthera hyperborea, P. obtusata, P. orbiculata,
Foamflower (Tiarella cordifolia), Wild Sarsaparilla
and One-flowered Wintergreen (Moneses uniflora).

LonG-LIVED COLONIEs: In 1977, Clarence and Enid
Frankton showed us a colony of G. repens that they
had discovered in 1972 in the Stony Swamp
Conservation Area [DAO 627112]. There were
about 90 rosettes, 80 of them in a dense patch 30 cm
square and the rest less than a metre away. They
were under a White Spruce in an Eastern White
Cedar - White Spruce forest growing in a thin layer
of leaf litter over Nepean sandstone. We measured a
pH of 4.0 at the colony. Although the colony likely

Vol. 111

had been there for a long time, it disappeared in the
1980s. Perhaps its location so closely in contact with
the bedrock far from soil humidity made it particu-
larly vulnerable to environmental stresses.

Near Poltimore, Quebec, a north-facing sandy
slope above a fen supported several dozen scattered
plants of G. tesselata and of G. repens in 1978.
Logging removed much of the dominantly Balsam
Fir forest cover in the early 1980s. By the mid-
1990s, the decade-old growth of Trembling Aspen,
Balsam Fir and Eastern Hemlock sheltered a similar
mixed colony of these two orchids.

EARLY History: John Kerr McMorine collected G.
repens in August 1862 at Ramsay, Ontario [DAO
96772, QK 12922]. James Fletcher made the only
other known 19th century collection at “Stewart’s
Bush, Ottawa, Carleton Co.” in 1879 [DAO 17595].
Elizabeth Keen White painted a plant provided by
James Fletcher from “Rich Woods. Ottawa” in 1877
(Dore 1965"), possibly also Stewart’s Bush.

John Macoun (circa 1911°) included in his list for
G. repens three collections of his own, one currently
at CAN that is referable to G. repens as well as two
that we have not seen. He also included James
Fletcher’s specimen of G. tesselata that is currently at
DAO (see G. tesselata). Given this mixed message as
to Macoun’s knowledge of G. repens, we have not
mapped the two collections that we have not seen. It is
interesting that the three Macoun specimens were col-
lected near railway stations, the G. repens at CAN
[116935] at “Wakefield, Quebec, in woods north of
railway station, August 10, 1903”, while the latter two
unknown Goodyeras were collected “in the Mer Bleue
at Blackburn Station, May 27, 1903” and “at Navan
station, C.P.R. Short Line, September 5, 1911”.

Hysrips: Kallunki (1976) has annotated two possible
hybrids between G. repens and G. tesselata. Both are
from Quebec, one from Gatineau Park [DAO 96754]
and one from Papineau County [DAO 16902]. Such
plants are extremely difficult to identify.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 83

sea ae eS
alas 5 ae Ea sy
A LETTS
‘ Ni :
—<<e \ Re
LUM \ y
u

FiGuRE 17. Goodyera repens var. ophioides, coniferous forest habitat, Val-des-Monts Municipality (Wakefield Township,
Gatineau County), Quebec, 6 August 1978.

84 THE CANADIAN FIELD-NATURALIST

Goodyera tesselata Loddiges

Tessellated Rattlesnake-plantain

Vol. 111

Goodyérie panachée

Goodyera tesselata is a species of shaded forests and swamps, particularly on the Shield, that can form small,
dense patches by vegetative reproduction. It can be recognized by its basal rosette of tessellated leaves with nar-
row greenish white lines bordering the median vein. The most definitive, but not most convenient, character is
the rostellar beak length, which lies between 0.5 mm and 2 mm (Kallunki 1976).

DESCRIPTION

Height: 12 (16 - 26) 31 cm [114 plants].

Flowers: 8 (16 - 29) 35 [65 plants]; white, tending
to face the brightest available light in a somewhat
one-sided inflorescence; fragrance faintly spicy.

Leaves: 2 - 5 on flowering and non-flowering
plants, occasionally to 10 and rarely to 15 on non-
flowering plants [48 plants]; 1.8 - 4 cm long, 0.9 -
1.8 cm wide [18 leaves]; greyish green (not bluish
green as is sometimes reported), with greenish white
markings on upper surface only (see Figure), mark-
ings occasionally broadened to cover most of leaf
except directly over veins; occasionally 2 erect
leaves at base of stem, occasionally lowest cauline
bract expanded to form a small spatulate leaf.

Overwintering State: as a rosette of leaves (see

Vegetative Reproduction and Flowering Frequency
below).

Capsules: light brown, spheroid, typically 0.4 x
0.35 cm, ascending (see Figure la); yield highly
variable but usually high, averaging 70% [14 plants].

Seeds: brownish orange, released in early
September, a month earlier than our other
Goodyeras.

BLOOMING PERIOD: 9 July (20 July - 15 August) 3
September [36 records].

CoLony SIZEs: | - 400, typically to 100, flowering
and non-flowering rosettes [57 colonies], as scat-
tered individuals, small clumps and patches of sever-
al dozen rosettes.

DISTRIBUTION: The Ottawa District is well within

46°

45°
76° 30'

75°

Goodyera tesselata: @ = herbarium specimen, @ = Native Orchid Location Survey sight

record. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

_ FIGURE 18. Goodyera tesselata, hemlock - maple - birch forest habitat, Gatineau Park, Quebec, 26 July 1975.

85

86 THE CANADIAN FIELD-NATURALIST

the range of this eastern orchid. It occurs predomi-
nantly in the Mixed Forest Region and adjacent
Boreal Forest Region (Kallunki 1976). Within the
District, G. tesselata is confined largely to the
Canadian Shield, especially in areas of calcareous
bedrock and of sand. On the Lowlands, the mixed
sandstone and calcareous rocks of the Stony Swamp
Conservation Area support an exceptionally large
concentration of colonies.

HABITATS: Mature, mesic forests and various
swamps are the two typical habitats for G. tesselata
in the District. Forested slopes adjacent to swamps
appear to be especially favoured localities. These
habitats provide heavy shade, adequate moisture and
little competition from other herbaceous plants.

Deciduous, mixed and coniferous forests impor-
tant to this orchid are composed of various combina-
tions of the following species (in order of decreasing
frequency): Sugar Maple, Eastern Hemlock, Eastern
White Pine, Eastern White Cedar, Yellow Birch,
Balsam Fir, Beech and Largetooth Aspen. The leaf-
covered sandy humus of the forest floor is some-
times shared, at a distance, with such species as
Ground-cedar (Lycopodium complanatum), Shining
Clubmoss (L. lucidulum), Wild Lily-of-the-valley
(Maianthemum canadense) and Wild Sarsaparilla
(Aralia nudicaulis). Occasionally, Goodyera repens
and, rarely, Corallorhiza striata, C. trifida,
Cypripedium parviflorum var. pubescens, Epipactis
helleborine and Platanthera hookeri also occur.
Rarely, G. tesselata is found at the interfaces
between pine plantations and mixed forests. We have
recorded a pH of 5.0 at two forest locations.

In swamps of Eastern White Cedar, Black Ash,
Yellow Birch and other trees, G. tesselata plants
grow on rotting logs and on the roots of trees among
various swamp mosses.

LONG-LIVED COLONIEs: Since 1975, we have moni-
tored a colony of about a dozen rosettes of G. tesse-
lata in Gatineau Park (Reddoch and Reddoch 1989).
(A plant in this colony is illustrated in the Figure.)
We discovered that the pattern on every leaf is dif-
ferent and thus we were able to follow the leaves in
photographs of the colony from year to year. On this
basis, we obtained the results described below under
Vegetative Reproduction and Flowering Frequency.
From computer simulations of the growth of this
colony, based on an exponential growth model and
on a random walk model, we estimated that the
colony likely originated from a single rosette in the
1940s.

Until 1980, the colony thrived in heavy shade
under a canopy of Eastern Hemlock, Sugar Maple,
Yellow Birch and White Birch. In that year, some
low-lying large trees shading the colony fell over as
a result of a new beaver dam, thus exposing the
colony to full afternoon sun and heat, and reducing
ambient humidity and soil moisture. As a conse-

Vol. 111

quence of these stresses, several changes occurred.
The number of rosettes dropped from 21 to 5 in
1980. The colony persisted but with reduced vigour.
After 1980, leaf lengths and inflorescence heights
were two-thirds of their former sizes. The distance of
new growths from the parent stem halved and the
further distance travelled before flowering was also
much less. On the other hand, neither the average
number of leaves on a non-flowering rosette nor the
number of new growths produced by a flowering
rosette changed very much. Significantly, the time
between the flowering of a parent and of its off-
spring decreased from a minimum of four years
before 1980 to three years between 1980 and 1986,
the last time that a plant flowered.

In 1989, after several hot, dry summers, there
were four rosettes left. Continued dryness resulted in
only one rosette remaining in 1990; it still survives
to the present (1996). Our records show that this
rosette appeared in the colony in 1983 and has never
flowered, while continuing to produce an average of
two new leaves per year.

Another long-lived colony of G. tesselata coexist-
ing with G. repens and described under that species,
has survived from at least 1978 to the present despite
logging of the forest habitat.

EARLY History: Although G. tesselata was
described in 1824, it was not until the turn of the
20th century that its status as a separate taxon was
clarified (Kallunki 1976; Morris and Eames 1929).
Thus it is not surprising that when early local
botanists first collected it, they treated it as G.
repens. Goodyera tesselata was not included in any
of the early lists of the Ottawa Flora (Billings 1867;
Fletcher 1880, 1893; Macoun circa 1911°). Macoun
listed the DAO collection cited below under G.
repens.

The first Ottawa District collections of this orchid
were made in the late 1870s. James Fletcher collect-
ed it on 4 August 1877, from “Hull P.Q. near
Ottawa” [DAO 96807] and on 8 August 1879, from
“Stewart’s Bush, Ottawa, Ont.” [MTMG 47563]
(with “Ottawa, Ont.” of the same date [TRT 15499]
a likely duplicate). In July of 1879, H. M. Ami col-
lected this species at “Dom. Springs” (= Dominion
Springs = Carlsbad Springs, Ontario; [MTMG
3419]), probably on the Ottawa Field-Naturalists’
Club excursion there on 19 July.

In 1974, we examined a specimen of G. tesselata
at the Canadian Forestry Service herbarium (OTF)
at the Central Research Forest in Gloucester. M. L.
Anderson had collected a flowering plant [Anderson
869] on 1 August 1974, from a group of two flower-
ing and eight non-flowering rosettes. He directed us
to the colony, located near the herbarium on the
Dolman Ridge beside the Mer Bleue Bog in a heavi-
ly shaded hemlock forest on damp sand. The herbar-
ium was moved to Petawawa and, subsequently

1997

(1995), its contents were incorporated into the
National Herbarium (CAN). At that time, this speci-
men could not be found (Albert Dugal, personal
communication). The colony has not been seen in
recent years.

VEGETATIVE REPRODUCTION AND FLOWERING
FREQUENCY: The year after flowering, a plant dies,
but not before producing one or two, or occasionally
three, new growths at an average distance of 2.3 cm
from the parent. Each new growth generates two or
three new leaves each year until the year it flowers.
Each leaf lasts about two years. By the time a rosette
reaches flowering size after a minimum of four
years, it has at least three to five leaves and is now

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 87

about 3.5 cm from its parent. It does not produce any
leaves the year it flowers. If a flowering stem is bro-
ken off at the base during flowering, the plant may
not create any new growths. Occasionally a new
rosette appears where there have been no recent
flowering plants; it may be either a seedling or a
growth from a non-flowering plant. (See Reddoch
and Reddoch (1989) for further details on this
colony.) On average, 24% of the rosettes flower in a
given year. This is consistent with the four-year
blooming cycle discussed above.

Hysrips: Kallunki (1976) has annotated two possi-
ble hybrids between G. repens and G. tesselata (see
the account of the former species).

88 THE CANADIAN FIELD-NATURALIST

Vol fT

Liparis loeselii (Linnaeus) L. C. M. Richard

Loesel’s Twayblade

Liparis de Loesel

In recent times L. /oeselii has been widespread in many habitats, especially in wetlands and on moist sands. It
shows considerable variation of stature according to its habitat. It can be recognized by its two basal, erect or
arching, elliptical to lanceolate leaves and its several greyish green or greenish yellow flowers.

DESCRIPTION

Height: 4 (8 - 17) 29 cm [219 plants] (see
Morphological Variation below regarding variation
with habitat).

Flowers: | (3 - 10) 24 [164 plants]; greyish green
in shady habitats to greenish yellow in sunny habi-
tats; no odour detected.

Leaves: 2, rarely 3 [DAO 691473]; 2 on non-
flowering plants; greyish green, usually long and
arching in shade; yellowish green, short and erect in
sun.

Overwintering State: as the corm of the current
year.

Capsules: pale yellow to yellowish white, obovoid,
typically 1.2 x 0.5 cm, erect or nearly so; yield usually
greater than 40%, averaging 70% [89 plants].

Seeds: brownish orange, released in late
September to late October.

76° 30

BLOOMING PERIOD: 9 June (18 June - 10 July) 2
August [72 records].

CoLony SIZEs: | - 430, typically to 150, flowering
and non-flowering plants [112 colonies], as scattered
individuals and sometimes in small clumps.

DISTRIBUTION: The Ottawa District is at the northern
edge of the range of this eastern species. It occurs
predominantly in the Mixed Forest Region. Within
the District, this orchid is thinly scattered on the
Shield but is rather more common on the Lowlands,
especially in areas of sandstone and of sand deposit-
ed by the Champlain Sea and subsequent rivers.

Hasirats: Liparis loeselii grows in many mesic to
wet-mesic, semi-shaded to open habitats over a vari-
ety of bedrock types.

Relatively natural habitats include willow swales,
alder thickets, coniferous and deciduous swamps,

75°

Liparis loeselii: 1 = herbarium specimen, @ = Native Orchid Location Survey sight record,
A = literature reference. Areas of major sand deposits on the Lowlands are shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 89

FicureE 19. Liparis loeselii, left plant and flowers (mesic, open habit): height 13 cm, sandy borrow pit, Borthwick Ridge,
Mer Bleue Conservation Area, Regional Municipality of Ottawa-Carleton, Ontario, 23 June 1970; right plant (wet-
mesic shade habit): height 20 cm, cedar-ash swamp, Gatineau Park, Quebec, 19 July 1970. (The two plants are
drawn to the same scale.)

90 THE CANADIAN FIELD-NATURALIST

forest stream edges and moist patches on forest
floors. Open sedge fens and floating sedge mats are
other naturally-occurring locations. In fens, the
corms are lodged in peat among sedges and mosses,
while in the other habitats they are supported in leaf-
mould or among mosses on the ground or on rotting
logs. In these latter habitats, there is generally little
other vegetation except some mosses.

Like Pogonia ophioglossoides, L. loeselti occasion-
ally grows on shoreline rotting logs that have fallen
into the water (see Local History below). Both species
also used to grow on floating boom logs on the
Gatineau River (Bayly 1972, personal communication
1977, [|CCO 3038]). (For additional details on these
two habitats, see the P. ophioglossoides account.)

The disturbed habitats of open roadside ditches,
old-fields and borrow pits harbour the majority of
the District’s colonies of this orchid. These habitats
have in common that they are sandy and mesic to
wet-mesic, with sparse vegetation and little shade.
Most of the locations plotted on the distribution map
in the Lowlands are such disturbed habitats. (See the
accounts of Calopogon tuberosus and Platanthera
clavellata for additional details of open sandy fields
near the Mer Bleue Bog.)

LONG-LIVED COLONY: Wetlands and forests can be
relatively stable and colonies of L. loeselii can per-
sist in them for many decades. For instance, the plant
of the wet-mesic shaded form illustrated grew near
the edge of an Eastern White Cedar - Black Ash
swamp in a colony that has continued from at least
the late 1960s to the present.

LocaL History: Braddish Billings Jr. collected L.
loeselii on 9 July 1860 in Dow’s Swamp and noted
that it was rare there [QK 13217]. He did not include
it in his list of plants collected in the summer of
1866 (Billings 1867). A number of other collections
were made in and adjacent to Dow’s Swamp
between 1879 and 1953.

James Fletcher (1893) cited stumps and floating
logs in the Rideau Canal as a habitat for L. loeselii.
He made two collections on the edge of the canal at
Patterson’s Creek, in 1878 and 1879 [DAO 17766,
17769]. At that time it is likely that the banks of the
65-year-old canal were still relatively unimproved.
Patterson’s Creek is now an elongated pond running
from the west bank of the canal almost to Bank
Street, south of the Queensway.

Morphological Variation with Habitat

average (range) [sample size]

Height
Leaf Length
Number of flowers

mesic, open
9.2 (5 - 19) cm [87]
5.7 (3.5 - 8.7) cm [6]
8 (3-17) [18]

Vol ttl

Both James Fletcher (1893) and John Macoun
(circa 1911") knew of this orchid only from relative-
ly natural habitats, except for one 1911 collection
that Macoun listed from the roadside near the new
entrance to Beechwood Cemetery. Fletcher (1893)
considered this orchid to be “not uncommon”.

Near Arnprior, Charles Macnamara (circa 1940")
described it as “scarce and local” in his caption of a
1913 photograph. In recent years, Runtz (1984)
regarded it as common in the larger Arnprior area.

In 1953, W. G. Dore noted on his collection from
the railway right-of-way adjacent to Dow’s Swamp
that L. loeselii was “scarce in the District”. A dozen
years later, members of the Native Orchid Location
Survey began turning up colonies with some fre-
quency, especially on the Lowlands in disturbed,
open areas on sands and sandstone. It is likely that
many of these habitats were created only in the
1960s with regrading of ditches, removal of sand for
roadbuilding and abandonment of farming.

MORPHOLOGICAL VARIATION: We have observed,
as have various authors, systematic variation in the
stature of this orchid depending on its habitat. To
study this variation more quantitatively, we classi-
fied the habitats as wet-mesic and shady (swamps),
wet-mesic and open (fens), mesic and open (early
succession, sandy, old-fields), and mesic and shady
(transitional, sandy, old-fields). Of the two plants
illustrated in the Figure, the larger is from a wet-
mesic, shady habitat, while the smaller is from a
mesic, open habitat.

The characters examined were plant heights, leaf
lengths and numbers of flowers. We found that the
variations in the numbers of flowers did not have a
significant correlation with the habitats. However,
the plant heights and the leaf lengths, although form-
ing a continuum, did correlate significantly with the
habitat type. The averages of these two characters
were largest in the wet-mesic, shady sites and small-
est in the mesic, open sites. For mesic, shady sites,
the corresponding averages were intermediate
between, and significantly different (5% level) from,
those of the two former habitats. For wet-mesic,
open sites, the averages suggested an intermediate
position, but insufficient data prevented a more
definitive account. The leaf length was nearly pro-
portional to the plant height in all habitats and so the
ratio of these two measurements did not vary greatly
or significantly among habitats.

wet-mesic, shaded
15.2 (5 - 29) cm [73]
10.3 (3.5 - 23) cm [59]
6 (1 - 14) [67]

mesic, shaded
11.6 (8 - 18) cm [18]
8.3 (5.5 - 10.3) cm [16]
6 (1 - 13) [16]

1997

Plants in a colony can change in stature, apparently
with changing light conditions. During the 1970s, we
followed a colony in a borrow pit on the Borthwick
Ridge beside the Mer Bleue Bog. At the beginning of
the decade, the many plants grew on the open, mesic
sand among sparse vegetation. These plants were of
the mesic, open form. (One of the plants from this
colony is shown in the Figure and the photograph of
a 1969 clump accompanied an article on the Mer
Bleue Bog (Dunston 1970).) By the end of the
decade, the site was becoming overgrown with alders
and other vegetation. The plants were now of the
mesic, shaded form. (With increasingly complete
shade, the colony disappeared within a few years.)

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 91

In conclusion, both the plant height and the leaf
length are larger in the presence of more moisture
and less light, but the flower number is not
strongly dependent on these conditions. The mor-
phology of plants in a colony changes with chang-
ing light conditions and likely with changing mois-
ture regimes.

ABERRATION: A plant at the mesic, open Borthwick
Ridge habitat described above had three flowers, the
top two arising from between two floral bracts, side
by side, and having fused pedicels and ovaries, but
otherwise being complete. The flowers were erect
with their lips touching each other.

92 THE CANADIAN FIELD-NATURALIST

Listera auriculata Wiegand

Auricled Twayblade

Vol. 111

Listere auriculée

Listera auriculata, a small green-flowered plant of shaded stream banks, is an ephemeral member of the Ottawa
flora, the only known colony having been discovered in 1967 and having survived for only a decade. It can be
recognized by its two sub-opposite, sessile, cauline leaves and its several greenish flowers. The lip has essen-
tially parallel sides and its apex has a small notch between two rounded lobes.

DESCRIPTION

Height: 8 - 12 cm [4 plants].

Flowers: 4 (6 - 12) 15 [18 plants]; greyish green
at the centre of the lip, tending to a more greyish,
translucent colour at the edges and on the petals and
sepals.

Leaves: 2; 2 on non-flowering plants.

Overwintering State: a shoot | - 2 cm high at the
base of the current year’s stem below ground;
herbarium specimens from elsewhere in Ontario and
Quebec show the new shoot present at anthesis.

Capsules: light brown, ellipsoid to spheroid, typi-
cally 0.5 x 0.3 cm, ascending.

Seeds: orange white (from herbarium specimens
elsewhere in Ontario and Quebec).

BLOOMING PERIOD: 14 June (16 June - 4 July) 9
July [9 records].

46°

COLONY SIZES: to 162 flowering and non-flowering
plants [1 colony], as scattered individuals and occa-
sionally as pairs and small groups.

CURRENT STATUS: rare to uncommon (S3) in Ontario
(Active List, Oldham 1996"); rare or extirpated in the
Ottawa District (no extant colonies known).

DISTRIBUTION: The Ottawa District is near the
southern edge of the range of this largely eastern
Boreal Forest Region species. Within the District,
one colony was located in Gatineau Park on the
Canadian Shield in an area of porphyroidal gneiss
bedrock (Hogarth 1970).

HABITAT: The plants were situated on the mesic to
wet-mesic floodplain of a small stream in the dap-
pled shade of fairly mature Eastern Hemlock,
Yellow Birch and Sugar Maple. They grew in leaf-

45°
76° 30'

75°

Listera auriculata: #@ = herbarium specimen. The Canadian Shield is shaded.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

FIGURE 20. Listera auriculata, mixed forest - flood plain habitat, Gatineau Park, Quebec, 27 June 1970.

93

94 THE CANADIAN FIELD-NATURALIST

covered, sandy, alluvial soil and on rotting logs,
stumps and tree roots among mosses; Stinging Nettle
(Urtica dioica), Touch-me-not (/mpatiens capensis)
and Sugar Maple shoots were present at the edges of
the colony.

LocaL History: Hue MacKenzie discovered the
first plant of L. auriculata on 16 September 1967, on
an Ottawa Field-Naturalists’ Club outing, which we
also attended. The plant, with conspicuous capsules,
was on a rotting log beside a rivulet running into
Fortune Creek. The next June, Hue and members of
the Native Orchid Location Survey confirmed the
identity of the group of two flowering plants and one
non-flowering plant (Greenwood 1968b; MacKenzie
and Greenwood 1969). The photograph by Gary
Hanes in the former reference shows the 1968 flow-

Vol. 111

ering plant as well as its old flowering stems from
the two previous years. From one to four plants
appeared each year until 1973, except for 1971 when
the terrain was flooded and no plants were visible.

In 1970, we encountered 162 plants on the flood-
plain of Fortune Creek about 100 m from the first
group [DAO 600828, 691478]. About two-thirds of
the plants produced flowers. This group declined to
11 plants in 1972 and was last seen in 1977. That
year, there were three non-flowering plants along
the stream edge about 50 cm above the stream bed.
They were growing in a 30 cm-wide border of the
moss Atrichum undulatum. Brunton and Crins
(1975) reported several hundred plants in the general
area of this colony in 1972. The relationship of this
report to the plants described above is not clear.

Loo

Listera australis Lindley

Southern Twayblade

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 95

Listére australe

Listera australis is a tiny green and reddish orchid that grows in peatlands among mosses of similar colours. It
was found in the Mer Bleue Bog at the end of the nineteenth century, but after about a decade was not seen
again, there or elsewhere in the District. It can be recognized by its two sub-opposite, sessile, ascending, cauline
leaves, its brown stem, and the reddish colour of the flowers. The lip has a very deep notch separating two long,
pointed lobes. The sepals and petals are about 1/4 of the length of the lip and are strongly reflexed.

DESCRIPTION

Height: 14 - 21 cm [10 plants].

Flowers: 7 - 16 [10 plants]; red to brown with
green depending on the proportions and concentra-
tions of the pigments: lip brownish red to greyish
ruby with greyish green or greyish yellow translu-
cent centre, sepals and petals greyish green or grey-
ish yellow with traces of red (colours based on
Alfred Bog plants).

Leaves: 2.

Overwintering State: herbarium specimens show
a shoot at the base of the current year’s stem in the
moss substrate; the new shoot is present at anthesis.

BLOOMING PERIOD: 28 May (3 June - 19 June) 21
June [7 records, including 4 from Alfred Bog].

CoLony Sizes: in Alfred Bog, over 40 flowering

46°

45°

plants in one part of the bog (Whiting and Bobbette
1974), as scattered individuals and in loose groups;
difficult even to estimate given the inconspicuous
nature of the plants and their widely scattered occur-
rence.

CURRENT STATUS: rare in the Provinces of Quebec
(Bouchard et al. 1983) and Ontario (Catling, White
et al. 1982), very rare (S2) in Ontario (Active List,
Oldham 1996"), rare in Canada (Argus and Pryer
1990); apparently extirpated in the Ottawa District
(no plants reported since 1902).

DISTRIBUTION: The Ottawa District is at the northern
edge of the distribution of this orchid of southern
Ontario and Quebec, New York State and coastal
plain areas from Nova Scotia to the Gulf states
(Catling, White et al. 1982). It is a species predomi-

76° 30

75°

Listera australis: 1 = herbarium specimen. The Canadian Shield is shaded.

96 THE CANADIAN FIELD-NATURALIST Vol. 111

FIGURE 21. Listera australis, poor fen habitat, Alfred Bog, Caledonia Township, United Counties of Prescott and Russell,
Ontario, 14 June 1974.

1997

nantly of the Southeastern Coastal Plain Forest
Region.

Within the District, there were two colonies of L.
australis in the Mer Bleue Bog a century ago.
Elsewhere in the Ottawa Valley, there is a thriving
population in various parts of the 4000 ha Alfred
Bog, 20 km east of the edge of the Study Area.

HasiratT: In Alfred Bog, plants of L. australis are
scattered through the sedge - shrub fen openings of
the patterned bog community and in openings in the
Black Spruce - Tamarack bog forest. (See Cuddy
(1983) for a vegetation map of Alfred Bog and a
description of the vegetation communities.)

The fen mats in the patterned bog are at least par-
tially floating and there are many areas of open
water. The orchids grow above the water level in wet
Sphagnum magellanicum and S. nemoreum, on the
fen floor and on the sides of hummocks around trees.
There is little competing shrub cover where the
orchid occurs; Three-leaved False Solomon’s Seal
(Smilacina trifolia) is almost the only other species
sharing the patches of Sphagna, sometimes accom-
panied by Pitcher-plant (Sarracenia purpurea),
Small Cranberry (Vaccinium oxycoccos) and
Buckbean (Menyanthes trifoliata). Farther away,
Pale Laurel (Kalmia polifolia) and Labrador Tea
(Ledum groenlandicum) are common. (See Whiting
and Bobbette (1974) for the report of the discovery
of this orchid in Alfred Bog and for photographs and
a description of the fen habitat. See Whiting (1974)
and Reddoch (1983a) for additional background and
for our photographs of this habitat.)

In the bog forest, L. australis plants grow on the
flat Sphagna-covered floor among scattered sedges.

EARLY History: On 21 June 1893, James Fletcher

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 97

discovered L. australis for the first time in the
Ottawa District and for the first time in Canada. He
found “a bed of this rare little orchid” in the southern
part of the Mer Bleue Bog, north of the Poplar Ridge
(Poplar Island) near Eastman’s Springs (Carlsbad
Springs) (Whyte, Craig, and Cowley 1894, [CAN
116993, TRT 15703]). Ottawa Field-Naturalists’
Club members again visited the colony on 28 May
1896 and found about a dozen plants in full flower
(Fletcher 1896; Whyte, Craig, and Macoun 1897).
The latter authors reported that specimens were col-
lected; however, we have not been able to find a rel-
evant collection in the herbaria that we visited. The
naturalists also visited the colony in 1898 (Campbell,
Macoun, and Whyte 1899).

On 20 June 1902, John Macoun (circa 1911°) col-
lected L. australis from the northwest corner of the
Mer Bleue Bog “amongst spruce trees ... about a
mile east of Blackburn Station and half a mile south
of the railway” [TRT 15701, MTMG 8596].

There have been no further reports of L. australis
from the bog despite periodic searches. It is probable
that fires and other disturbances such as various
drainage schemes (Ashley 1979°) caused drastic
changes in the habitats. Much of the burned area is
now an expanse of ericaceous shrubs with Black
Spruce and Tamarack making a comeback in some
places.

ABERRATIONS: A plant with nine flowers that we
collected at the Alfred Bog [DAO 691479] had two
otherwise normal flowers side by side arising from a
single broadened pedicel.

Another plant that we saw there had three leaves,
one of them quite small and opposite a normal-sized
leaf. A second normal-sized leaf grew about a cen-
timetre below the small leaf.

98 THE CANADIAN FIELD-NATURALIST

Vol. 111

Listera cordata (Linnaeus) R. Brown var. cordata

Heart-leaved Twayblade

Listere cordée

Listera cordata, a tiny, inconspicuous plant found in a few swamps, is the only Listera species currently known
in the District. It can be recognized by its pair of spreading, cauline leaves and its flowers that have spreading
sepals and petals about half as long as the lip. The narrow lip has a deep notch between two long, narrow, point-

ed, diverging lobes.

DESCRIPTION

Height: 7 (9 - 17) 22 cm [132 plants].

Flowers: 4 (6 - 12) 22 [123 plants]; sepals greyish
green, lip and petals greyish green to brown; no
odour detected.

Leaves: 2; 2 on non-flowering plants.

Overwintering State: a greyish green shoot, | -
1.5 cm high, at the base of the current year’s stem in
the moss substrate; herbarium specimens show the
new shoot present at anthesis.

Capsules: greyish yellow, spheroid to ellipsoid,
typically 0.3 x 0.2 cm, ascending (see Figure 1b);
yield averaging 60% [11 plants], similar to the 70%
yield reported from California (Ackerman and
Mesler 1979).

Seeds: yellowish white, released in late June to
early July while the flower parts are still fresh; this is

by far the earliest release of seeds of any Ottawa
District orchid.

BLOOMING PERIOD: 4 June (10 June - 28 June) 13
July [14 records].

COLONY SIZEs: | - 300 flowering plants [10
colonies], as scattered individuals and in loose
groups.

DISTRIBUTION: The Ottawa District is well within
the North American range of this transcontinental
orchid. It is a species of the Montane, Boreal and
Mixed Forest Regions. Within the District, L. corda-
ta is thinly scattered across the Shield and in the
Lowlands of the southwest.

HABITAT: Listera cordata is confined to semi-
mature and mature coniferous swamps, usually in
areas of calcareous bedrock. Plants occur in the

46°

45°

76° 30! -)

for

Listera cordata var. cordata: @ = herbarium specimen, @ = Native Orchid Location Survey

sight record. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 99

FIGURE 22. Listera cordata var. cordata, coniferous swamp habitat, Val-des-Monts Municipality (Wakefield Township,
Gatineau County), Quebec, 11 June 1975 (plant), 11 June 1978 (flowers).

100

shade of Eastern White Cedar and other trees such as
Balsam Fir, Eastern White Pine, White Spruce,
Black Spruce, Yellow Birch and Speckled Alder.
They grow on mesic or wet-mesic swamp floors in
leaf mould and in various mosses, including
Sphagnum magellanicum, S. fallax and Hylocomium
splendens, that may enclose the plants up to their
leaves. Herbs and shrubs associated with this orchid
include Bristle-stalked Sedge (Carex leptalea),
Three-leaved False Solomon’s Seal (Smilacina trifo-
lia), Goldthread (Coptis trifolia), Labrador Tea
(Ledum groenlandicum), Starflower (Trientalis bore-
alis) and Twinflower (Linnaea borealis).

EARLY History: This species is not included in
James Fletcher’s Flora Ottawaensis of 1893 or in

THE CANADIAN FIELD-NATURALIST

Vol. 111

any earlier list. John Macoun (circa 1911") cited as
his only record for L. cordata a collection that he
made on 2 June 1898, “in a sphagnum swamp 3
miles west of North Wakefield”. This description fits
Chilcott’s Swamp, the calcareous peatland at the
edge of Johnston Lake, Quebec, first visited by
botanists in the fall of 1892 (Whyte, Craig, and
Cowley 1893). Although we have been unable to
find this specimen, we found three others from this
locality in local herbaria, collected in 1940, 1945 and
1960 [DAO 138637, 138638, CAN 386175].

ABERRATIONS: Rarely there is a small cauline bract
between the leaves and the inflorescence. On one
plant, in a swamp near Poltimore, Quebec, this bract
was the size of a normal leaf.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

101

Malaxis monophylla (Linnaeus) Swartz var. brachypoda (Gray)

Morris & Eames

White Adder’s-mouth

Malaxis 4 pédicelles courts

SYNONYMS: Microstylis monophyllos (Linnaeus) Lindley, Malaxis brachypoda (Gray) Fernald

Malaxis monophylla is a widely distributed, but not abundant, midsummer swamp orchid. It is a small and
inconspicuous plant that becomes more noticeable after the light-coloured capsules have formed. It can be iden-
tified by its single, seemingly cauline leaf (which, in fact, sheaths the stem to the base) and by its numerous,
very small flowers with pointed lips. In contrast to M. unifolia, the pedicels are no longer than the height of the
flowers, resulting in a sparse, narrow inflorescence about 7 mm in diameter.

DESCRIPTION

Height: 8 (10 - 17) 25 cm [105 plants].

Flowers: 9 (18 - 34) 51 [41 plants]; yellowish
white or pale white, sometimes light green or pale
green; fragrance, none detected.

Leaf: 1, rarely with a second smaller, lower,
opposite leaf, 1 on non-flowering plants.

Overwintering State: as the corm of the current
year.

Capsules: pale orange, greyish orange or orange
white, spheroid, typically 0.5 x 0.3 cm, ascending
(see Figure 1b); yield highly variable, averaging

76° 30'

40% [29 plants].
Seeds: orange white, released in early October.

BLOOMING PERIOD: 8 June (23 June - 15 July) 29
July [38 records].

COLONY SIZEs: | - 275, typically to 50, flowering
plants [117 colonies], generally as scattered individ-
uals.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the distribution of this mainly eastern
orchid. It is a species of the Mixed Forest Region
and adjacent Boreal Forest Region. It is widespread

75°

Malaxis monophylla var. brachypoda: @ = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. Major areas underlain by calcareous rock

(marble and limestone) are shaded.

102 THE CANADIAN FIELD-NATURALIST Vol. 111

FIGURE 23. Malaxis monophylla var. brachypoda, plant: cedar - ash swamp habitat, Gatineau Park, Quebec, | July 1975;
flowers: cedar swamp, Goulbourn Township, Regional Municipality of Ottawa-Carleton, Ontario, 21 June 1970.

1997

in the calcareous regions of the District, chiefly in
the southwest and in the Gatineau Valley.

HasiraTs: Malaxis monophylla usually grows in
partially to deeply shaded, moist to wet swamps and
low-lying woods. At least a few plants are present in
most calcareous swamps in the District, especially
swamps dominated by Eastern White Cedar and
Black Ash. Rarely, this species also occurs in Silver
Maple swamps.

In swamps, the orchid corms are lodged in leaf
mould or in moist to wet organic soil, often among
mosses. Where the swamp floor is very wet, the
plants grow on mounds around trees and over tree
roots, as well as on rotting logs. There may be no
other companion plants or there may be such typical
swamp vegetation as Cinnamon Fern (Osmunda cin-
namomea), Oak Fern (Gymnocarpium dryopteris),
Jack-in-the-pulpit (Arisaema triphyllum), Coral-
lorhiza trifida, Cypripedium reginae, Platanthera
hyperborea, Naked Mitrewort (Mitella nuda),
Foamflower (Tiarella cordifolia), Water Avens
(Geum rivale), Wood-sorrel (Oxalis acetosella),

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

103

One-flowered Wintergreen (Moneses uniflora), One-
sided Pyrola (Orthilia secunda) and Twinflower
(Linnaea borealis).

Once in a while, a few plants are encountered in
moist to wet situations in willow swales, alder thick-
ets and stream edges, as well as on somewhat drier
ground in moist coniferous and mixed forests. Rarely
a few plants are found in fens.

Where M. monophylla and M. unifolia occur
together, the former species is usually in the lower
and wetter parts of the moisture gradient.

EARLY History: Early naturalists knew of M.
monophylla at only a few locations and considered it
rare in the District (Fletcher, Scott, and Cowley
1892; Fletcher 1893). James Fletcher made the first
collections in Dow’s Swamp in 1878 [DAO 17822,
MTMG 47282], where it grew “in large numbers”
along with two other rare orchids, Amerorchis rotun-
difolia and Cypripedium arietinum (Whyte and
Small 1883; Fletcher, Scott, and Cowley 1892). John
Macoun (circa 1911") was not acquainted with this
species in the District.

104

Malaxis unifolia Michaux

Green Adder’s-mouth

THE CANADIAN FIELD-NATURALIST

Vol. 111

Malaxis unifolié

SYNONYM: Microstylis ophioglossoides (Muhlenberg) Nuttall

Malaxis unifolia is a widespread and relatively common, if inconspicuous, orchid found in a wide variety of
habitats. It can be recognized by its single, sheathing, cauline leaf and its numerous tiny greenish flowers. In
contrast to M. monophylla, the pedicels are notably longer than the height of the flowers. As a result, the flat-
topped, dense inflorescence is at least 1 cm in diameter. The lip has a distinct notch at the apex, often containing

a small point.

DESCRIPTION

Height: 6 (10 - 17) 23 cm [250 plants] (see also
Notable Plant below).

Flowers: 10 (16 - 49) 80 [58 plants]; greyish
green like the rest of the plant or greenish yellow
with centre of lip greyish green; fragrance lacking or
mild and sweet; flower orientation somewhat irregu-
lar, especially for the lower flowers.

Leaf: 1, rarely 2, one clasping the other at mid-
stem, | on non-flowering plants.

Overwintering State: as the corm of the current
year.

Capsules: pale orange, obovoid, typically 0.5

76° 30'

x 0.25 cm, horizontal to ascending; yield usually less
than 40%, averaging 15% [83 plants].

Seeds: greyish orange, released in mid to late
September.

BLOOMING PERIOD: 16 June (2 July - 30 July) 19
August [105 records].

COoLony SIZEs: | - 150, typically to 60, flowering
plants; 10 or fewer in 74% of colonies; one excep-
tional colony in the Constance Creek valley, West
Carleton Township, Ontario, with 1100 plants in
1968 (Hue MacKenzie, records of the Native Orchid
Location Survey) [141 colonies]; generally as scat-

ia

Malaxis unifolia: @ = herbarium specimen, @ = Native Orchid Location Survey sight record,
A = literature reference. The Canadian Shield is shaded; areas underlain by sand and sand-

stone are omitted for clarity.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 105

FiGuRE 24. Malaxis unifolia, plant: pine plantation habitat, Larose Forest, Cambridge Township, United Counties of
Russell and Prescott, Ontario, 26 July 1969; single flower and top view of inflorescence: sandy old-field habitat,
Mer Blue Conservation Area, Regional Municipality of Ottawa-Carleton, Ontario, 23 June 1970.

106

tered individuals and occasionally in clumps of up to
a dozen flowering and non-flowering plants.

CURRENT STATUS: A century ago this species was
considered to be rare in the District (Fletcher 1893).
Perhaps its abundance has been enhanced in the lat-
ter half of this century by the increased number of
disturbed sites on sand.

DISTRIBUTION: The Ottawa District is in the northern
part of the range of this eastern species. In North
America it inhabits the Mixed and Deciduous Forest
Regions. It is also found in parts of Mexico, Central
America and the Caribbean (Catling 1991). This
orchid is widely scattered across the District, espe-
cially in areas of acidic rock and of sand left by the
Champlain Sea and subsequent rivers.

HaBitaTs: Malaxis unifolia occurs in a variety of
habitats, from dry hilltops to moist swamps, under
open sun to deep shade. The most common soil sub-
strate is sand or sandy loam.

Plants are perhaps most often encountered grow-
ing in thin sandy soil on open or partially shaded
Precambrian knolls or on expanses of flat-lying
sandstone. They are also frequent in recently dis-
turbed, open areas of mesic sand amongst sparse
vegetation (see description of Dolman Ridge habitat
of Calopogon tuberosus).

Colonies also become established on the bare
ground of heavily-shaded coniferous forests and pine
plantations (see description of Larose Forest colony
of Cypripedium acaule). Shady deciduous forests
and more-open forest edges also support colonies on
occasion. Sometimes there are a few plants in semi-
open forests at the shores of rivers and small lakes.

THE CANADIAN FIELD-NATURALIST

Vol. 111

In the wet-mesic to wet conditions of cedar
swamps, plants tend to stay above wet swamp floors
on rotting logs and on mounds around the bases of
trees (see account of M. monophylla above).

Although most habitats occupied by M. unifolia
are sandy and acidic, there is the occasional site that
is neither (see description of The Burnt Lands
colony of Cypripedium arietinum in clay soil over
limestone).

EARLY History: Braddish Billings Jr. collected M.
unifolia “on the nearly perpendicular slope of King’s
Mountain” in what is now Gatineau Park on 10 July
1860 [QK 13271]. Another early collection at
Queen’s University is that of John Kerr McMorine
in 1862 from Ramsay, Ontario [QK 66701].

James Fletcher made collections near Dominion
Springs (now Carlsbad Springs) in 1879 [DAO
17835, MTMG 47291, TRT 15728] and from Dow’s
Swamp in 1880 [DAO 17835]. At Dow’s Swamp, R.
B. Whyte, and H. B. Small (1883) reported only one
plant in 1882 and thought it curious that in previous
years it had been as plentiful there as M.
monophylla. By 1893 James Fletcher knew of it only
from Dominion Springs, Dow’s Swamp and Aylmer,
and regarded it as rare.

NOTABLE PLANT: In August of 1995 we found in the
Larose Forest a plant 35 cm tall growing out from
between two Red Pine logs lying on the ground. Of
that height, 13 cm of the stem was hidden from view,
passing between the logs to the corm sitting on the
sandy ground beneath. The previous year’s flowering
stem was present beside it. A month later the logs had
been removed and there was no evidence of the plant.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

107

Platanthera blephariglottis (Willdenow) Lindley var. blephariglottis

(Northern) White Fringed-orchid

Platanthére 4 gorge frangée

SyNonyM: Habenaria blephariglottis (Willdenow) Hooker

This orchid is one of our rarest, having been recorded only from three very wet peatlands in the District. It is
fairly conspicuous, however, with its clusters of white flowers standing just above the sedges and other wet-
land plants. It is easy to identify by its habitat and by its brilliant white flowers with short-fringed single-lobed

lips.

DESCRIPTION

Height: 11 (26 - 39) 50 cm [148 plants], signifi-
cantly shorter than the 100 cm reported by Case
_ (1987) for Michigan, southwestern Ontario and

Ohio.

Flowers: 3 (7 - 20) 35 [48 plants]; white on all
parts, sometimes with pale yellow at end of spur;
column white with pale yellow from the pollinia
showing through the anther sacs; pedicel often white
at top, blending into pale yellow or pale green at
base; inflorescences with few flowers appearing
globular or irregular, those with more flowers
becoming more cylindrical; fragrance none or faintly
spicy during day and early evening.

Leaves: 2, sometimes 3, grading into cauline

ee

45° fa
76° 30'

bracts for a total of 3 - 5 leaves and bracts; one leaf
on non-flowering plants.

Overwintering State: a greyish green shoot
beside the current year’s stem in the moss substrate,
rising from the stem or from a horizontal root up to 1
cm from the base; herbarium specimens show the
new shoot and partially elongated roots present at
anthesis.

Capsules: light to dark brown, ellipsoid, typically
1.2 x 0.3 cm, ascending (see Figure 1b); yield
approaching 100% on 5 plants.

Seeds: light brown to brown, released in late
September to early October.

BLOOMING PERIOD: 12 July (18 July - 5 August) 16
August [21 records].

fo

Platanthera blephariglottis var. blephariglottis: Hi = herbarium specimen, @ = Native Orchid
Location Survey sight record. The Canadian Shield is shaded.

108 THE CANADIAN FIELD-NATURALIST Vol. 111

FIGURE 25. Platanthera blephariglottis var. blephariglottis, poor fen habitat, Mer Bleue Conservation Area, Regional
Municipality of Ottawa-Carleton, Ontario, 19 July 1973.

1997

CoLony SIZEs: 3 - 100 flowering plants [9 records
from two peatlands], as scattered individuals or
occasionally in groups of two or three.

CURRENT STATUS: rare in the Provinces of Quebec
(Bouchard et al. 1983) and Ontario (White et al.
1982b), rare to uncommon (S3) to common (S4) in
Ontario (Watch List, Oldham 1996"); rare in the
Ottawa District.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the distribution of this northern variety
of P. blephariglottis. This variety occurs primarily in
the Mixed Forest Region (Catling 1983b). Within the
District, P. blephariglottis has been found in several
parts of the Mer Bleue Bog and in two small peat-
lands on the Canadian Shield. Elsewhere in the
Ottawa Valley, a colony occurs in the patterned bog
- fen complex of Alfred Bog, 20 km beyond the east-
ern edge of the Study Area (Cuddy 1983).

HABITAT: This species is confined to minerotrophic
areas in certain bogs that can be categorized as poor
fens. The largest colonies thrive in sunny openings
near scattered Black Spruce and Tamarack where the
Sphagnum-covered fen floor is very wet and loosely
consolidated. Shrubs are thinly scattered or absent
and sedges are the most common herbs. Large patch-
es of Bog Rosemary (Andromeda glaucophylla) and
numerous plants of Three-leaved False Solomon’s
Seal (Smilacina trifolia) are frequent. Small
Cranberry (Vaccinium oxycoccos) is often present
and Sphagnum magellanicum is the most abundant
moss. Leatherleaf (Chamaedaphne calyculata) and
Labrador Tea (Ledum groenlandicum) are occasional
shrubs, especially close to the trees.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

109

We observed that as one site in the Mer Bleue
became somewhat drier, shrubbier and more bog-
like, it supported fewer and fewer plants. A colony
of 95 flowering plants in 1973 had decreased to three
flowering plants in 1994. A few dozen flowering
plants of Cypripedium acaule continue to persist
there.

LocaL History: Ottawa Field-Naturalists’ Club
botanists began to explore the Mer Bleue in July of
1879 and P. blephariglottis was one of the first of
many rarities to be discovered there. It was collected
that July by Henry M. Ami [DAO 17057, MTMG
3388, TRT 15508] and by James Fletcher [CAN
116937], near Eastman’s (Carlsbad) Springs
(Macoun circa 1911°). Sporadic collections followed
from the southern and western edges of the bog. In
1971, Don Lafontaine and David White discovered
87 flowering plants out in the middle of the peatland
(Lafontaine 1971).

In 1963, W. G. Dore collected a plant, one of a
small number, from a “sphagnous bog mat around a
small lake” near Danford Lake west of Kazabazua,
Quebec [Dore 20359 at DAO]. Somewhat closer to
Gatineau Park, Monty and Grace Wood (personal
communications 1976, 1996) discovered this orchid
on their newly purchased property near St.-Francois-
de-Masham, west of Wakefield, in 1970. Several
hundred flowering plants were scattered on a partial-
ly floating, poor fen bordering a pond
[DAO 691514]. Similar numbers continue to flourish
there (Monty Wood, personal communication 1996).
This habitat is illustrated on the front cover of the
fourth-quarter issues of Trail & Landscape (Reddoch
1988).

110

THE CANADIAN FIELD-NATURALIST

Vol. 111

Platanthera clavellata (Michaux) Luer var. clavellata

Club-spur Orchid

Platanthére claviforme

SyNonyMs: Habenaria clavellata (Michaux) Sprengel, Habenaria tridentata (Muhlenberg) Hooker

Platanthera clavellata is neither abundant nor conspicuous. It can be identified by the single cauline leaf at or
somewhat below the centre of the narrow, ridged stem, the rather irregular arrangement of the flowers and the
three rounded teeth at the end of the otherwise undivided and unfringed lip.

DESCRIPTION

Height: 10 (18 - 27) 36 cm and 46 cm (see
Notable Plant below) [311 plants].

Flowers: | (3 - 10) 26 [180 plants]; pale green,
pale yellow or white, sometimes greyish green along
centre line on outside of dorsal sepal and at end of
spur; irregularly arranged as a result of incomplete
rotation about the pedicellate ovary, appearing to lie
on their sides, often facing in a common direction
toward the stronger light; when there are more flow-
ers, the inflorescence becomes more regular; fra-
grance mild and sweet or none.

Leaves: 1, occasionally 2 on robust plants, the
upper one much reduced, with 2 small bracts above;
| leaf on non-flowering plants.

Overwintering State: a greyish green shoot, 1.5 -

2 cm tall, 1.5 mm diameter, the tip just below ground
or moss substrate level in early fall, rising from a
horizontal root about 2 cm away from the current
year’s stem; herbarium specimens show the new
shoot present at anthesis.

Capsules: brown, ellipsoid, typically 0.8 x 0.4
cm, ascending irregularly; yield approaching 100%
for three-quarters of the plants, highly variable for
the remainder, averaging 85% [44 plants], plants
susceptible to predation by White-tailed Deer, espe-
cially in Gatineau Park.

Seeds: light brown, released in mid September to
early October.

BLOOMING PERIODs: 5 July (11 July - 6 August) 17
August [28 records].

46°

45°
76° 30

75°

Platanthera clavellata var. clavellata: @ = herbarium specimen, @ = Native Orchid Location
Survey sight record. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 111

FIGURE 26. Platanthera clavellata var. clavellata, old-field habitat beside Black Lake, Gatineau Park, Quebec, 18 July 1970.

112

CoLony SIZES: 2 - 77, typically to 50, flowering
plants [17 colonies], as scattered individuals and
occasionally in groups of from two to as many as
nine flowering stems.

DISTRIBUTION: The Ottawa District is in the northern
part of the range of this eastern orchid, which is pre-
dominantly in the Mixed and Deciduous Forest
Regions. Within the District, P. clavellata is thinly
spread across the Canadian Shield and the Lowlands.

HABITATS: Two types of habitats provide suitable
environments for P. clavellata: moist, sandy, old-
fields and openings, and the borders of peatlands,
swamps and lakes. Old-fields, openings and
lakeshores generally provide short term habitats,
while peatlands and swamps are more likely to offer
stable conditions for decades or longer.

In old-fields and openings, plants favour light to
medium shade adjacent to (and sometimes well
under) trees and shrubs. Depending on the amount of
shade, plants may be accompanied by any mixture of
old-field herbs, including Field Horsetail (Equisetum
arvense), Marsh Fern (Thelypteris palustris), Liparis
loeselii, Wild Strawberry (Fragaria virginiana),
Dewberry (Rubus hispidus), Heal-all (Prunella vul-
garis), Indian Tobacco (Lobelia inflata), Yarrow
(Achillea millefolium), goldenrods (Solidago spp.)
and various grasses. Occasionally, Royal Fern
(Osmunda regalis), Malaxis unifolia and Platanthera
lacera, as well as patches of Sphagnum and
Polytrichum mosses are present. A pH of 5.0 was
measured in two colonies near the Mer Bleue.

In peatlands, usually fens, colonies are sometimes
encountered on the open sedge mat near the border
with the adjacent swamp, growing in Sphagnum
magellanicum and other minerotrophic Sphagna in
the light shade of Tamarack and Labrador Tea
(Ledum groenlandicum). A pH of 4.5 was measured
at the roots in the Sphagna of one fen.

LONG-LIVED COLONY: A colony of a few dozen
plants that we have followed since 1968 flourishes
along the moist edge of a White Cedar - Black Ash
swamp over marble bedrock in Gatineau Park. The
plants grow on mossy hummocks (pH 5.3) in
dappled shade along with Bristly Clubmoss

THE CANADIAN FIELD-NATURALIST

Vol. 111

(Lycopodium annotinum), Cinnamon Fern (Osmunda
cinnamomea), Royal Fern (O. regalis), Bluebead-lily
(Clintonia borealis), Goldthread (Coptis trifolia),
Wild Sarsaparilla (Aralia nudicaulis), Bunchberry
(Cornus canadensis), Starflower (Trientalis
borealis), Red Maple seedlings and other swamp
species. Some plants grow equally well a metre
away from the swamp edge in the mesic leaf mould
of the adjacent mixed forest with Rattlesnake Fern
(Botrychium virginianum), Bluebead-lily, Wild Lily-
of-the-valley (Maianthemum canadense), Wild
Sarsaparilla, Sugar Maple seedlings and other mesic
forest inhabitants.

EARLY History: James Fletcher collected P. clavel-
lata in a “tiny bog” (Fletcher 1893) at Black Lake in
what is now Gatineau Park on 8 July 1878 [DAO
267278, MTMG 47571]. He was referring, perhaps,
to the small, partially-floating, poor-fen mat that per-
sisted at the southeast corner of the lake until recent
times. (The mat was mostly flooded by Beavers in
1983 (J. M. Reddoch and A. H. Reddoch 1987d)).
He collected Pogonia ophioglossoides there on the
same day.

We have not found P. clavellata on the fen mat,
but in the late 1960s and early 1970s we did
encounter some 50 flowering plants on the north
shore of the lake, about 100 m away from the mat, in
moist, old-field conditions. They are no longer there.
One of these plants is depicted in the Figure.

From 1878 until 1950, no additional collections
seem to have been made in the District.

NOTABLE PLANT: In 1970, we encountered and pho-
tographed a strikingly large plant that grew in the
Eastern White Cedar - Black Ash Swamp described
above. One of a group of eight medium to large
plants, it was 46 cm tall with two leaves and 26
flowers, 30% taller and with 50% more flowers than
the next largest plants in the District. The inflores-
cence was cylindrical in shape. Unlike most plants,
its flowers, except for the lowermost ones, were
arranged in four nearly vertical ranks with lips
downward, giving the appearance of a small P.
orbiculata. We have not seen a comparable plant at
that site or at any other.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

113

Platanthera dilatata (Pursh) Lindley ex Beck var. dilatata

Fragrant White Orchid

SynonyM: Habenaria dilatata (Pursh) Hooker

Platanthere dilaté

An inhabitant of some of our fens, but not of our bogs, Platanthera dilatata is a striking, tall, white-flowered
plant that justifies the second part of its other common name, Bog Candles. It can be recognized by its narrow,
dense inflorescence of pure white, fragrant flowers with narrow lips and short spurs.

DESCRIPTION

Height: 27 (38 - 61) 77 cm [167 plants].

Flowers: 9 (11 - 40) 65 [24 plants]; white, some-
times becoming greyish green toward the tip of the
spur and at bases of sepals; arranged in dense or
loose inflorescences sometimes showing evidence of
spiral structure; fragrance spicy, often strong; Luer
(1975) included a photograph (Plate 60, #1) of an
inflorescence in the open sedge fen near Poltimore,
Quebec, to illustrate this species.

Leaves: 5 to 8 leaves and bracts.

Overwintering State: a greyish green, narrowly
conical shoot, appearing above ground beside the
current year’s stem in late September; herbarium
specimens show the new shoot and partially elongat-
ed roots present at anthesis.

46°

Capsules: light brown, ellipsoid, typically 0.8 x
0.35 cm, ascending to erect (see Figure 1b); yield
variable, averaging 60% [9 plants].

Seeds: light brown, released in mid to late August.

BLOOMING PERIOD: 8 June (20 June - 22 July) 10
August [40 records].

CoLony SIZEs: | - 190 flowering plants [12
colonies], as scattered individuals and, rarely, in
small groups.

DISTRIBUTION: The Ottawa District is in the south-
ern part of the eastern range of this primarily north-
eastern and western species. In the east, it is mainly a
plant of the Boreal and Mixed Forest Regions.
Within the District, P. dilatata is confined to fens in
areas of calcareous bedrock.

75°

Platanthera dilatata var. dilatata: @ = herbarium specimen, @ = Native Orchid Location Survey
sight record. Major areas underlain by calcareous rock (marble and limestone) are shaded.

114 THE CANADIAN FIELD-NATURALIST Vol. 111

FIGURE 27. Platanthera dilatata var. dilatata, sedge fen habitat, Val-des-Monts Municipality (Wakefield Township,
Gatineau County), Quebec, 1 July 1974.

1997

HasitatT: Platanthera dilatata occurs only in open
sedge fens and in a few adjacent treed fens. It never
grows in the more acidic environments of bogs. In
sedge fens, plants are scattered across the fen floor
in full sun, accompanied here and there by Pogonia
ophioglossoides and Calopogon tuberosus. It
avoids the wettest parts of fens, that is, the centres
of basin fens and the outer edges of floating fens.
(Our photograph in Baird (1983) shows a plant of
P. dilatata in typical sedge fen habitat near
Poltimore, Quebec; see also Appendix 2.) There are
sometimes suitable conditions for a few plants in
sedgy openings among Eastern White Cedar, Black
Spruce and Tamarack where the central open fen
gives way to treed fen. Populations have been pre-
sent in these fens for 30 years and likely have been
there for a century or longer.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

115

EARLY History: Braddish Billings Jr. made the
first two Ottawa District collections at Dow’s
Swamp, less than 2 km west of the family home at
Billings Bridge, on 9 July 1860 [QK 66646] and 8
July 1861 [CAN 227237]. He made a third collec-
tion in 1866 (Billings 1867), its current location
unknown. These collections likely came from the
floating sedge fen on the edge of the central pond
shown in the 1925 air photo (Reddoch 1978b). He
collected Pogonia ophioglossoides there on the
same dates and Calopogon tuberosus there on the
first date. James Fletcher collected P. dilatata at
Dow’s Swamp in 1878 [DAO 17071]. Collections
in this century began fifty-four years later with Bill
Dore’s 1932 specimen from Glenfarne (= Danford
Lake, Pontiac County), Quebec.

116 THE CANADIAN FIELD-NATURALIST Vok fii

Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer

Northern Tubercled Orchid Platantheére a gorge tuberculée var. petite-herbe
SYNONYMS: Habenaria flava (Linnaeus) R. Brown var. herbiola (R. Brown) Ames and Correll, Habenaria
virescens Sprengel, Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown) Luer f. /utea (Boivin)
Whiting and Catling

Platanthera flava is a rather rare orchid in the District, but, being green and inconspicuous and inhabiting wet,
weedy or woody shores and flood plains, it may be somewhat overlooked. It is a plant with cauline leaves and a
cylindrical inflorescence of yellowish green flowers with the spurs somewhat longer than the lips. However the
most diagnostic feature of the flower is the mound or tubercle rising from the base of the lip. The tubercle is

most clearly seen from the side.

DESCRIPTION

Height: 18 (24 - 35) 40 cm [52 plants].

Flowers: 13 (18 - 33) 50 [40 plants]; bicoloured
with greyish green sepals and pale yellow, some-
times to pale green, petals, lip and spur; in a moder-
ately dense cylindrical inflorescence of somewhat
irregular appearance with flowers rotating in both
directions on a single plant; fragrance moderately
strong, sweet and very pleasant.

Leaves: 3, occasionally 4, with 2 or 3 cauline
bracts; 2 on non-flowering plants.

46°

Overwintering State: a shoot rising from a hori-
zontal root several cm away from the current year’s
stem; herbarium specimens show the new shoot pre-
sent at anthesis.

Capsules: brown, ellipsoid, typically 0.6 x 0.3
cm, ascending.

Seeds: dark brown, released in early to mid
October.

BLOOMING PERIOD: | July (8 July - 1 August) 6
August [16 records].

75°

Platanthera flava var. herbiola: @ = herbarium specimen, @ = Native Orchid Location
Survey sight record. Major areas underlain by calcareous rock (marble and limestone) are

shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 117

Cae
&
i
2
ie
z
e
ye
£
:

FIGURE 28. Platanthera flava var. herbiola, Ottawa River shore among shrubs, Britannia, City of Ottawa, Regional
Municipality of Ottawa-Carleton, Ontario, 3 July 1975.

118

CoLony SIZEs: | - 300 flowering and non-flowering
plants [13 colonies], as scattered individuals and as
patches of flowering and non-flowering stems.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983); rare to uncommon (S3) in
Ontario (Watch List, Oldham 1996'); rare in the
Ottawa District. Most of the colonies mapped no
longer survive.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the range of this eastern orchid of the
Deciduous and Mixed Forest Regions. Within the
District, most discoveries of P. flava have taken
place on the north and south shores of the Ottawa
River upstream from Ottawa, but some colonies have
been found in a few other locations in Quebec. The
Native Orchid Location Survey contains a record
from the Mississippi River just south of the Study
Area. Colonies generally occur in areas of calcareous
bedrock.

HasirTaTs: Moist, open or partly shaded areas on the
flood plains of the Ottawa River and some of its trib-
utaries provide favourable habitats for this orchid.
On these flood plains, several zones are occupied:
open, grassy and weedy beaches not far above the
summer water-line, shrub zone edges facing the
water, clearings in willow thickets, and forest floors
under Red Maple, Silver Maple (and their hybrids)
and Red Ash. Soils are sandy or silty. Purple
Loosestrife (Lythrum salicaria) is an almost constant
companion in these four situations and may be a
threat to the survival of these habitats. In more
exposed locations, Spiranthes lucida occasionally
shares the same habitat.

One 1947 collection [DAO 17093] from the
Britannia area came, not from the river edge, but
from “along the railway tracks” that ran parallel to
the Ottawa River a short distance away. Here, the
plants grew in “rich black loam”. Colonies have
been known on the river shore in the Britannia area
for several decades.

EARLY History: The only collection that Macoun
(circa 1911°) cited was one made by Robert B.
Whyte in August 1879 in a “wet place, bank of river
near Ottawa”. Very few of Whyte’s collections are
held in public herbaria, and this is not one of them.
James Fletcher (1880) did not have this species in
his first Flora Ottawaensis but included it as an
addition to the list a year later (sub Habenaria
virescens, Spreng.; Fletcher 1881), perhaps on learn-
ing of Whyte’s collection. Curiously, a decade later,
James Fletcher, Robert B. Whyte and William Scott
(1891) and Fletcher (1891) reported Scott’s collec-

THE CANADIAN FIELD-NATURALIST

Vol. 111

tion of P. flava at Thurso on 6 August 1890 [DAO
267279, TRT 15571] as a new species for the
District. Fletcher described only this latter collection
in his Flora Ottawaensis of 1893.

On 5 July 1906, W. Hague Harrington found P.
flava “in some abundance” on the north shore of the
Ottawa River “near the Country Club” [DAO
267281]. The habitat was “a marshy river-front”
where the plants were dispersed among sedges and
other plants. When he visited the colony the next
year, he found it much reduced as a result of being
“badly trampled by cattle which seek the river either
to drink or stand in the water, and destroy much of
the littoral vegetation” (Harrington 1917, [CAN
116954, 116955)).

COLOUR PATTERN VARIANT: Boivin typified f. lutea
(Boivin) Whiting and Catling (Catling 1982a), an
obscure name attributed to Louis-Marie. He chose
for the type a 1966 collection of L. C. Sherk [547 at
DAO] and E. W. Greenwood from the now extirpat-
ed colony at Remic Rapids, and described it as
floribus luteis.

The collectors did not report the flower colour but
Ed Greenwood has told us (personal communication
1996) that they chose the plant as representative of
the colony. When we saw and photographed that
colony in 1969, the flowers were generally
bicoloured with greyish green sepals and pale yellow
lips and petals. In 1996, the bicoloured character of
the type was still evident in spite of some fading.
The sepals were greyish green to the same degree as
the leaves, while the petals and lips were light brown
and showed no sign of green. This description
applies to most of the P. flava of the District. Such
bicoloured patterns are typical of a number of
Ottawa District orchids.

A small sample of collections and published
colour photographs from other parts of the range of
P. flava var. herbiola suggests that such bicoloured
flowers may be the norm. Some photographs (e.g.,
Luer 1975) do show yellowish green in the lips and
petals of both var. herbiola and var. flava; however,
Homoya (1993) reported green to be the flower
colour of var. herbiola as distinguished from var.
flava. These variations in lip and petal colour could
be the result of explicit colour pattern variation or be
within the normal range of control of chlorophyll
formation.

These observations suggest to us that f. /utea may
not differ from the type of var. herbiola. Although
we have not seen the approximately 200 year-old
type specimen, it may be, however, that the original
colours have disappeared.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

119

Platanthera grandiflora (Bigelow) Lindley

Large Purple Fringed-orchid

SyNONYM: Habenaria fimbriata (Dryander) R. Brown

Platantheére grandiflore

Platanthera grandiflora is one of our most spectacular orchids, being both large and colourful. It is similar to
the more common P. psycodes (Stoutamire 1974) with which it sometimes grows. These two species can be
easily recognized by their numerous purple fringed flowers on a leafy stem. Platanthera grandiflora is distin-
guished from P. psycodes by having a large round entrance to the spur rather than a somewhat flattened one and
by having anther sacs that diverge downward instead of being close and parallel. The former species also tends

to have flowers nearly twice as large as the latter.

DESCRIPTION

Height: 33 (44 - 73) 97 cm [71 plants].

Flowers: | (12 - 34) 55 [87 plants], on average
about three-quarters as many as on P. psycodes; gen-
erally light lilac, pale violet or purple, occasionally
deep violet or so pale as to be almost indistinguish-
able from white (Petrie (1981) illustrates the latter
colour on an Ottawa District plant), with white at
base of lip and on column; spur translucent, often
becoming pale violet sometimes with a trace of
green toward tip; in a fairly dense, regular inflores-

cence; fragrance light, sweet, daisy-like.

46°

45°
76° 3

Leaves: 3 - 5, with 2 - 5 bracts, 1 - 3 on non-flow-
ering plants.

Overwintering State: a greyish green, broadly to
narrowly conical shoot, | - 4 cm above ground beside
the current year’s stem, appearing there in late
September; herbarium specimens show the new shoot
and partially elongated roots present at anthesis.

Capsules: light brown, ellipsoid to oblong, typi-
cally 1.3 x 0.35 cm, ascending to erect (see Figure
Ic); yield highly variable, averaging 50% [10 plants].

Seeds: dark brown, released in mid to late
September.

75°

Platanthera grandiflora: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. Areas of major sand deposits on the Lowlands are shaded, as

is the Canadian Shield.

120 THE CANADIAN FIELD-NATURALIST Vol. 111

di

FIGURE 29. Platanthera grandiflora, deciduous forest/stream flood plain habitat, Gatineau Park, Quebec, 3 July 1976
(plant), 15 July 1980 (flowers).

1997

BLOOMING PERIOD: 16 June (27 June - 13 July) 23
July [55 records], of shorter duration and two to
three weeks earlier than P. psycodes.

Co.Lony SIZEs: | - 15, typically to 8, flowering
plants [14 colonies] (sometimes to 50 flowering
plants (Paul Catling, personal communication
1996)), as scattered individuals, rarely in clumps of
several flowering and non-flowering stems.

CURRENT STATUS: rare in the Province of Ontario
(Catling, Whiting et al. 1982), extremely rare (S1) in
Ontario (Active List, Oldham 1996’).

DISTRIBUTION: The Ottawa District is at the north-
western limit of the distribution of this Appalachian -
Atlantic Coast species of the Mixed Forest Region
(Stoutamire 1974). It has not been found farther up
the Ottawa Valley (Runtz 1984, personal communi-
cation 1996; Whiting and Catling 1986). Within the
District, this orchid is scattered across the Canadian
Shield, generally in areas of acidic bedrock or deep
sand. On the Lowlands, it is localized on the sands
deposited by the Champlain Sea and subsequent
rivers.

HABITATS: The most common places for P. grandi-
flora on the Canadian Shield are moist to wet
depressions, stream edges and flood plains under the
high canopies of deciduous forests, usually with
Sugar Maple and Beech dominating.

The leaf-carpeted sandy loam or muck is often
completely saturated and sometimes plants are stand-
ing in shallow water. Companion species include
Lady Fern (Athyrium filix-femina), Ostrich Fern
(Matteucia struthiopteris), Sensitive Fern (Onoclea
sensibilis), Stinging Nettle (Urtica dioica) and
Touch-me-not (Jmpatiens capensis), as well as
Platanthera psycodes.

On the Lowlands, P. grandiflora usually grows in
the dappled shade of poorly-drained Red Maple
forests. One site that was more mesic than usual was
a shaded, second growth forest of White Pine, Sugar
Maple, Largetooth Aspen and other trees. The sandy
substrates in forests on the Lowlands are moist to
wet owing to the relatively impervious layer of clay
underneath but are susceptible to drying out during
periods of drought.

Both on the Shield and in the Lowlands, colonies
are found only rarely in the open, moist, sedgy
patches that are a common habitat for P. psycodes.

LONG-LIVED COLONIES: The mixed colony of P.
grandiflora and P. psycodes in Gatineau Park that
we discovered in 1969 (Reddoch 1976; J. M.
Reddoch and A. H. Reddoch 1987a) continues to
thrive. Numbers, never large, vary from year to year
as plants appear, flower for a few years and then dis-
appear. The habitat is a wide stream valley that is
kept saturated by steady seepage from a beaver pond
upstream. The formerly closed canopy of Sugar
Maple, Beech, White Elm, White Birch and Yellow

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

121

Birch is now more open with the demise and decom-
position of the large elms. White-tailed Deer are a
menace in this colony because they eat off the tops
of flowering plants and occasionally trample shoots.

The mixed colony that Ed Greenwood recorded in
1968 on the Lowlands in Cumberland Township (J.
M. Reddoch and A. H. Reddoch 1987a) survives
with only P. grandiflora remaining. The P. psycodes
plants in the shallow roadside ditch disappeared a
couple of years after we saw them, apparently the
victim of spraying; however, a few P. grandiflora
continue to appear in the ditch from time to time.
They occur as well in the adjacent, young Red Maple
- Trembling Aspen - Yellow Birch forest. The forest
colony may serve as a seed source for restocking the
ditch after spraying, or more recently, cutting. Both
habitats are moist to wet with substrates of silty
humus and sandy clay. Sensitive Fern, Cinnamon
Fern (Osmunda cinnamomea) and Marsh Fern
(Thelypteris palustris) accompany the orchids at this
site, along with many roadside weeds in the ditch.

Mixed colonies of P. grandiflora and P. psycodes
are known at a few other places in Gatineau Park and
on the floodplain of the Picanoc River at the north-
ern edge of the Study Area (Reddoch 1976).

EARLY History: James Fletcher added this species
to his 1880 Flora Ottawaensis in 1881, probably on
the basis of Henry M. Ami’s specimen collected in
1880 in the “Laurentian Hills, Blanche River, East
Templeton, P.Q.” [DAO 84103]. Although only the
top half has been preserved, it is enough to show that
the plant was exceptionally robust. By 1893
(Fletcher 1893), this orchid had been found at two
other sites, Eastman’s Springs (Macoun, Whyte, and
Fletcher 1884) and Kingsmere. It was collected at
Eastman’s Springs (= Carlsbad Springs) also in 1887
and 1908 (([TRT 15561, DAO 84105], Gibson 1908).
Both Fletcher (1893, 1896) and W. Hague
Harrington (1917) concluded that the species was
rare in the District.

Harrington (1917) made collections on the edge of
the railway ditch near Kirk’s Ferry in 1906 and near
Chelsea in 1906 and 1907 [CAN 116949 - 116952].
The best locality known to him was “a small area of
springy ground near a cedar swamp in the deep
woods north of Chelsea, but this habitat will proba-
bly have been destroyed by the fires which followed
the cutting down of the beautiful forest. Instead of
grateful shade and lovely woodland vistas, there are
left the crumbing rocks denuded of soil and desolate
with the blackened trunks and stumps of the forest
monarchs”.

ACYANIC FORM: Two herbarium collections from
the Ottawa District may be referable to the white-
flowered f. albiflora (Rand and Redfield) Catling.
One of Harrington’s 1906 collections from Chelsea
[CAN 116951] includes one plant labelled “albino”.
This identification is credible but cannot be verified,

P22

given the present brown colour of the specimen. A
1943 collection by Gaston Lamarre from Thurso
[DAO 84111] was annotated “Habenaria psycodes
var. grandiflora f. leuacophaeopsis” by B. Boivin in
1966, although the collector did not describe the
flower colour. The specimen currently is uniformly
brown with no anthocyanin evident. If the plant had
initially had only a small amount of pigment, this
could have faded in the intervening 23 years before
Boivin examined it.

The palest flowers that we have seen, on a plant in

THE CANADIAN FIELD-NATURALIST

Vol. 111

the long-lived Gatineau Park colony discussed
above, could easily have been described as albino.
However, the presence of anthocyanins was con-
firmed by a clear boundary between the white base
of the lip and the faint violet white of the rest of the
lip. If a complete absence of anthocyanin, that is, a
genetic mutation preventing its formation, is
required in the definition of f. albiflora, then this
plant does not qualify. Similarly, the 1943 collection
discussed above could also have had pale, rather
than pure white, flowers.

1997

Platanthera hookeri (Torrey) Lindley
Hooker’s Orchid

SYNONYM: Habenaria hookeri Torrey

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

123

Platanthere de Hooker

Platanthera hookeri is widely but thinly distributed in the forests of the District. Although it is one of the green
orchids, its spiky, open inflorescence is often conspicuous above the bare forest floor. Like P. orbiculata and P.
macrophylla, it has two conspicuous, round, basal leaves. It can be distinguished from these two species by its
flowers, which are green or yellowish green instead of whitish. Its spurs taper to points and its lips are upturned.
In addition, with rare exceptions, P. hookeri lacks cauline bracts.

DESCRIPTION

Height: 14 (22 - 33) 44 cm [129 plants].

Flowers: 2 (8 - 16) 25 [138 plants]; sepals deep
green, petals, lip and spur greyish green or yellowish
green sometimes tending to greenish yellow; fra-
grance light, floral, by day.

Leaves: 2, basal; 1 - 2 on non-flowering plants;
either ascending or lying on the ground at anthesis,
plants apparently maintaining the same orientation
from year to year; dull to slightly lustrous; rarely 1
cauline bract, seen twice in the small colony at Mud
Pond (see below) but nowhere else in the District.

Overwintering State: a greyish green, broadly
conical shoot, 1 - 2 cm above ground beside the cur-

46°

rent year’s stem, appearing there in late September;
herbarium specimens show the new shoot and par-
tially elongated root usually present at anthesis.
Capsules: greyish orange, brownish orange or
dark brown, ellipsoid to obovoid, typically 1.4 x 0.5
cm, erect; yield usually less than 60%, averaging
30% [30 plants]; pollinators not known (Catling and
Catling 1991) and often not very effective, especially
in the Gatineau Park colony that we have monitored
(see below); on two occasions we have found similar
lepidopteran bristles and scales on the stigmatic sur-
faces of flowers at different sites. In one of the flow-
ers, both hemipollinaria had been removed; in the
other, neither had been disturbed but there was

45°
76° 30

15°

Platanthera hookeri: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

124

pollen on the stigmatic surface. These observations
together with the flower colour suggest that the polli-
nators are moths.

Seeds: brownish orange to brown, released in
early to mid October.

BLOOMINg PERIOD: 8 May (1 June - 29 June) 16
July [67 records].

CoLony SIZEs: | - 143, typically to 45, flowering
and non-flowering plants; 3 or fewer in 50% of
colonies [126 colonies], generally as scattered indi-
viduals.

DISTRIBUTION: The Ottawa District is near the north-
ern limit of the range of this Mixed Forest Region
species. Within the District, P. hookeri is almost
completely confined to the Canadian Shield and
some adjacent Lowland areas.

HABITATS: This woodland species is most often
found in the medium shade of moist, fairly mature,
Sugar Maple - Beech, Sugar Maple - Eastern
Hemlock, Eastern White Pine and Eastern White
Cedar forests. It is also found in forests just above
the edges of swamps or other wet areas. Occasional
plants are encountered in pine plantations and in par-
tially shaded Trembling Aspen and other young
deciduous or mixed woods.

Generally, P. hookeri grows where there is little
other vegetation. Soils are shallow to deep sands,
sandy loams or gravels over many bedrock types.
Cypripedium acaule is sometimes a companion in
coniferous forests on sand.

Rarely, a few plants grow in the moist humus of
calcareous peatlands.

LONG-LIVED COLONIES: Platanthera hookeri is a
long-lived plant and colonies can thrive for many
years in undisturbed surroundings. The patterns of
development that colonies follow are fairly individ-
ual and may depend on conditions related to their
specific habitats.

In 1984 we began to monitor a well-established
colony of P. hookeri near Kingsmere in Gatineau
Park that we had known since 1978. The plants are
scattered across an area about 30 m x 15 m under a
fairly dense canopy of Sugar Maple with some
Eastern Hemlock, Eastern White Pine and Hop
Hornbeam. Young hemlocks form the understory
and many of the plants are under the edges of the
hemlock boughs. The sandy loam is heavily covered
with deciduous leaf litter and fairly devoid of other
vegetation, except for a few patches of Wild Lily-of-
the-valley (Maianthemum canadense). The underly-
ing bedrock is granite pegmatite (Hogarth 1970).

A total of 22 plants has been followed between
1984 and 1996. Nine plants have been present for the
whole 13 years and all of these have flowered at least
once. Three additional plants that were present in
1984 have disappeared. Ten new plants have been
found during the course of this study, three of which

THE CANADIAN FIELD-NATURALIST

Vol. 111

have since disappeared, one after only one year.
(Some of the plants that have disappeared may have
been smothered by the heavy annual blanket of fallen
tree leaves; in June we have had to lift off the matted
leaves to locate a number of plants over the years.)

Of the 22 plants, 14 have flowered at least once.
Of these, 4 flowered once, 4 twice, | three times, |
four times, 2 five times, and | each seven and eight
times. The plant blooming most frequently flowered
7 years out of 8 with a break in the 6th year and then
flowered again (in 1995) after a break of 3 years. (It
also had flowered in 1981.) It did not set seed in any
of the years that it flowered.

In 1991, we found two additional plants near the
original colony. Between 1991 and 1996, one has
flowered four times and the other twice.

For all 24 plants, the average percentage of plants
flowering per year between 1984 and 1996 is 24%,
with a range from 0% to 55%. There were four years
when the average was between 0% and 7%, and four
other years when it was 46% to 55%. The relatively
high frequency of years with few flowers or with a
comparatively large number of flowers is inconsis-
tent with a model of independent random flowering,
as can be shown with a chi-squared analysis.
Evidently there is a tendency to coordinated flower-
ing, possibly the result of annual climatic influences,
although other factors may be involved as well.

Capsule production in the colony as a whole has
been very small. Eighty-three percent of the flower-
ing plants examined since 1981 did not produce cap-
sules. In the exceptional year of 1981, six of eight
flowering plants produced capsules with a yield per
inflorescence of 17% to 57%.

Most of this colony consisted of two-leaved plants
from the first time they were recorded. Only two
plants declined to one leaf for a couple of years
before disappearing.

Recently we began to monitor another colony of
P. hookeri in Gatineau Park, this one near Black
Lake. It is in a relatively open, moist, mostly decidu-
ous forest of Sugar Maple, Red Oak, Beech and
Eastern Hemlock over calc gneiss bedrock. Ground
cover is dominantly Wood-betony (Pedicularis
canadensis). After only three years, we can see that
this colony presents a somewhat different picture of
flowering frequency and capsule production than the
colony described above. All seven plants have flow-
ered each year and the capsule yield ranges from 5%
to 95% with an average of 40%.

Across the Ottawa River in Ottawa-Carleton, Ed
Greenwood discovered a colony of P. hookeri near
Mud Pond in 1962. (The plant in the Figure was a
member of this colony.) The plants were growing in
dappled shade at the edge of a cedar swamp. We fol-
lowed this colony of up to six plants from 1967 to
1977. During this time, one of the plants flowered
for six consecutive years and another flowered for
five consecutive years. None of the other plants

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 425

FiGuRE 30. Platanthera hookeri, plant: cedar swamp habitat near Mud Pond, City of Kanata, Regional Municipality of
Ottawa-Carleton, Ontario, 14 June 1969; flowers: mixed forest habitat, Gatineau Park, Quebec, 6 June 1978.

126

flowered during that period. In 1977, we found that
the colony had been flooded out by rising water lev-
els; it did not appear again.

EARLY History: Platanthera hookeri was one of
the 1867 additions to Braddish Billings Jr.’s (1867)
list of plants collected in 1866. In 1878, James
Fletcher collected the species in various rich and
rocky woods in the “Chelsea Mountains” and at
“Hull”, Quebec [MTMG 48465, DAO 267282,

THE CANADIAN FIELD-NATURALIST

Vol. 111

267285] at the beginning of concerted botanical
exploration north of the Ottawa River. Ottawa Field-
Naturalists’ Club excursions to Gilmour’s Grove,
near Chelsea, Quebec, in 1904 and 1908 found P.
hookeri along with Cypripedium acaule and
Galearis spectabilis (Clarke 1904, 1908). Gilmour’s
Grove consisted of a stand of ancient Eastern White
Pines, a hemlock grove and numerous deciduous
trees (Reddoch 1979b).

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

127

Platanthera huronensis (Nuttall) Lindley

Fragrant Green Orchid

Platanthére de la Huronie

SyNONYMS: Platanthera hyperborea (Linnaeus) Lindley var. huronensis (Nuttall) Luer, Platanthera x media
(Rydberg) Luer, Habenaria hyperborea (Linnaeus) R. Brown var. huronensis (Nuttall) Farwell

The taxon referred to here as Platanthera huronensis has been considered by Luer (1975) and others to be a puta-
tive hybrid between P. hyperborea and P. dilatata. Schrenk (1978) suggested that the hybrid may be, in fact, a
hybridogenic new species. Plants referable to P. huronensis as described by Charles Sheviak (personal communi-
cation 1996) occur in several Ottawa District fens, and it is these plants that we describe below. Both P. huronen-
sis and P. hyperborea can be distinguished from our other Platantheras by their several cauline leaves and numer-
ous small, greenish flowers (not pure white as in P. dilatata) with entire lips comparable in size to the spurs. In
the Ottawa District, P. huronensis can be distinguished from P. hyperborea by its open sedge fen habitat (rarely
in swamps), and its larger, light green flowers with a distinct rose-like fragrance. The two species can also be
separated by differences in the orientations of the anther sacs in the column (Paul Catling, personal communica-
tion 1994). See Further Diagnostic Notes below and the P. hyperborea account following.

DESCRIPTION

Height: 27 (39 - 70) 91 cm [56 plants].

Flowers: 12 - 105 [24 plants]; light green (but not
white) in the field; under the microscope, colours
darker but not as dark as P. hyperborea; dorsal sepa-
Is green or yellowish green with some readings of
light green or greyish green, lip, spur and petals
greyish green, usually lighter than the sepals, edges
of petals white; (lighter measurements in the field

46°

may be the result of surface light-scattering effects);
arranged in moderately dense to dense inflores-
cences; fragrance rose-like.

Leaves: 4 - 11 leaves and bracts [23 plants]; 1 - 4
leaves on non-flowering plants.

Overwintering State: a greyish green, conical
shoot within the moss substrate at the base of the
current year’s stem, in late September.

Capsules: light brown to brown, oblong, typically

75°

Platanthera huronensis: @ = herbarium specimen, ® = our sight record and photographs.
Major areas underlain by calcareous rock (marble and limestone) are shaded.

128 THE CANADIAN FIELD-NATURALIST Volo 1M

AY i
be /ag ~ BN

FIGURE 31. Platanthera huronensis, plant and column: sedge fen, Val-des-Monts Municipality (Wakefield Township,
Gatineau County), Quebec, 28 June 1996; flowers: cedar - ash swamp, Gatineau Park, Quebec, 6 July 1969; for the
column (front view showing self-pollination), scale bar = 0.5 mm.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 129

1.0 x 0.4 cm to 1.5 x 0.6 cm, nearly erect (see Figure
Ic); yield 90% - 100% [8 plants].

Seeds: light brown to brown, released in early
September to early October, several weeks earlier
than P. hyperborea.

BLOOMING PERIOD: 9 June (20 June - 14 July) 23
July and 14 August [16 records].

CoLony SIZES: 2 - 13 flowering plants [8 records];
as scattered individuals, sometimes in small clumps.

CURRENT STATUS: Our experience suggests that this
species may be rare in the Ottawa District.

DISTRIBUTION: The Ottawa District is apparently
well within the range of this transcontinental species
of the Montane, Boreal and Mixed Forest Regions
(Schrenk 1978; Charles Sheviak, personal communi-
cation 1996). Within the District, this orchid is
apparently restricted to calcareous wetlands.

Since 1968 we have identified and photographed
four colonies of the taxon currently recognized as P.
huronensis, three in fens and one in a calcareous
swamp. These are the colonies plotted on the distribu-
tion map along with a 1911 herbarium collection. A
small number of additional swamp colonies of P.
huronensis may have been recorded as P. hyperborea
in the Native Orchid Location Survey and mapped as
such; however, the vast majority of swamp colonies
are of P. hyperborea. The general distribution pattern
shown for the abundant P. hyperborea would not be
significantly altered by the inclusion of a few colonies
of P. huronensis.

Hasirats: Platanthera huronensis is known to us so
far only from open sedge fens and clearings in adja-
cent treed fens as well as in an Eastern White Cedar
- Black Ash swamp. In fens, the plants occur in the
open but close to the margins. They are not as far
away from the edges as P. dilatata, which invariably
grows nearby. Platanthera hyperborea is usually
present elsewhere in the wetland complex, generally
in the swamp component.

P. huronensis

LONG-LIVED COLONY: Since 1968 we have followed
the colony in the sedge fen near Poltimore and it is
from this colony that most of our knowledge about
this orchid comes. Although the colony has persisted
for many decades (and possibly centuries), individu-
al plants do not appear to last longer than two or
three years.

LocaL History: We have seen only one herbarium
specimen that is referable to P. huronensis. It was col-
lected by Faith Fyles in June 1911 from
“Northwakefield, P.Q. Swamp” [Fyles 2243 at DAO].
North Wakefield, present-day Alcove, was the loca-
tion often given by Fyles and other botanists for
Chilcott’s Swamp, the calcareous peatland at the edge
of Johnston Lake (Fyles 1912). Both P.dilatata and P.
hyperborea were collected from the same peatland in
the 1940s [DAO]. In 1979, we collected a specimen
of P. huronensis from the sedge fen near Poltimore,
Quebec [DAO 691524].

FURTHER DIAGNostTic NOTES: We provide below a
comparative list of characters observed for plants
referable to P. huronensis and P. hyperborea. Note
that these results are restricted to the Ottawa District
and the sample of P. huronensis is rather small, no
more than 10 measurements of floral parts.

In our local populations, the flowers are at the
small end of the range described by Sheviak and
they differ from populations observed elsewhere
(Sheviak, personal communication 1996) by being
self-pollinating. We deduce that the flowers are self-
pollinating because the hemipollinaria rotate for-
ward and bring the pollinia into contact with the
stigmatic surface (see Figure 31), where the pollen
loses its colour. This action can be seen when the lip
is up or down. We have done no experiments to
exclude pollinators, and it may be that both external
pollination and self-pollination takes place. The
high capsule yields reported above also suggest self-
pollination.

P. hyperborea

Habit: semi-stout to stout slender to stout
Height: to 91 cm to 87 cm
Leaves: usually ascending usually arcuate-spreading

Flowers: pale green yellowish, greyish or deep green
fragrant no odour
sepals: spreading to somewhat reflexed reflexed
lip: moderately or slightly dilated lanceolate or slightly dilated
length: 5.0 - 6.2 mm 3.5-5.5 mm
width: 2.0 - 2.5 mm 0.6 - 1.5 mm
spur length: 5.5 - 7.0 mm 2.7 - 5.0 mm

anther sacs:

separated at top,
diverging slightly (+ 30°) downward

viscidia: elliptic

viscidium spacing: +2.5 mm
rostellum: moderately arched (90° - 120°)
self-pollinating: yes

touching at top,
diverging widely (+ 90°) downward
orbicular
+ 1.1mm
slightly arched (+ 150°)
yes

130

THE CANADIAN FIELD-NATURALIST

Vol. 111

Platanthera hyperborea (Linnaeus) Lindley var. hyperborea

Northern Green Orchid

SYNONYM: Habenaria hyperborea (Linnaeus) R. Brown

Platanthere hyperboréal

Platanthera hyperborea is widely distributed in the District, where it is notable for the variation in its habit and
habitat. Both P. hyperborea and P. huronensis can be distinguished from our other Platantheras by their several
cauline leaves and numerous small, greenish flowers (not pure white as in P. dilatata) with entire lips compara-
ble in size to the spurs. Platanthera hyperborea can be distinguished from P. huronensis by its smaller, darker
green flowers without fragrance. (See the preceding account of P. huronensis for additional details, including

the column structure.)

DESCRIPTION

Height: 6 (16 - 47) 87 cm [204 plants].

Flowers: 2 (4 - 23) 60 [131 plants]; sepals green
or yellowish green, occasionally greyish green or
deep green, lip, spur and petals yellowish green to
greyish green, usually lighter and yellower than the
sepals; arranged in dense to lax inflorescences, occa-
sionally in 3 or 4 spiral ranks; no detectable odour.

Leaves: 2 - 8, grading to an additional 1 - 3
bracts; 1 - 4 on non-flowering plants.

Overwintering State: a greenish white or greyish
green, conical shoot, appearing at ground level
beside the current year’s stem, in October.

46°

e 9 oe

45° £

Capsules: brown to dark brown, oblong, typically
0.8 x 0.3 cm on small plants to 1.6 x 0.6 on large
ones, nearly erect; yield usually greater than 50%,
half of the plants approaching 100%, averaging 80%
[58 plants].

Seeds: light brown to dark brown, released in late
October (rarely September).

BLOOMING PERIOD: 14 June (24 June - 16 July) 15
August [85 records].

CoLony SIZEs: | - 690, typically to 140, flowering
and non-flowering plants [204 colonies], mostly as
individuals, but also in twos and threes.

76° 30!

75°

Platanthera hyperborea var hyperborea: @ = herbarium specimen, ®@ = Native Orchid
Location Survey sight record, A = literature reference. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 131

FIGURE 32. Platanthera hyperborea var. hyperborea, plant: swamp habitat, Gatineau Park, Quebec, 5 July 1980; flowers
and column: mesic, mixed forest, Gatineau Park, Quebec, 17 July 1996; for the column (front view showing self-
pollination), scale bar = 0.5 mm.

132

CURRENT STATUS: one of the most abundant and
widespread orchids in the District.

DISTRIBUTION: The Ottawa District is well within
the range of this transcontinental species of the
Boreal, Montane and Mixed Forest Regions. This
orchid is distributed throughout much of the District
especially in areas of calcareous bedrock.

HasiratTs: Platanthera hyperborea thrives in some-
what shaded, moist to wet places such as swamps,
low-lying woods, seeps and stream edges. Colonies
are also found in moist deciduous forests and cedar
woods. This orchid grows in both mineral and organ-
ic soils over a variety of bedrock types. It does not
grow in bogs.

Almost any type of swamp provides suitable habi-
tat for P. hyperborea, from Eastern White Cedar -
Black Ash swamps to mature Eastern White Cedar
swamps with widely-spaced, large trees. Ground
cover may be abundant, or almost absent; plants are
rooted in mosses or organic soils.

Plants also grow in running water around springs,
in the rich muck of seasonal stream beds and at the
edges of streams and ponds. Usually these places are
at least partially shaded, but occasionally colonies
occur in the open in wet spots in meadows, gravel
pits and old-fields.

Mesic to dryish cedar woods and fairly mature
deciduous forests provide other habitats for this
orchid. Plants come up through moist leaf mould
under more or less closed canopies of Eastern White
Cedar or of Beech, Sugar Maple, Largetooth Aspen
and other trees. Ground cover is usually sparse.

Sandy road or track edges through swamps or
forests sometimes support a few plants.

EARLY History: John Kerr McMorine’s two 1862
collections from Ramsay, Ontario, [QK 13017,
66657] are the earliest from the Ottawa District. P.
hyperborea was included in Braddish Billings Jr.’s
(1867) list of species that he collected in 1866, but
specimens supporting this list have not been located.
If the plant came from the fen at Dow’s Swamp
(Reddoch 1978b), it could have been referable to P.
huronensis. Henry M. Ami collected P. hyperborea
in “MacKay’s Grove” in July 1879 [CAN 23407].
(MacKay’s Grove was likely on the MacKay Estate,

Morphological Variation with Habitat

average (range) [sample size]

Habit

Height

Inflorescence Height
Number of Leaves
Number of Flowers
Number of Flowers / cm

THE CANADIAN FIELD-NATURALIST

Mesic Forest Habitats

Vol. 111

the property that extended across the centre of what
is now the Village of Rockcliffe Park, Ontario
(Belden 1879).) Across the Ottawa River in Quebec,
the east bank of Beaver Meadow, the outflow valley
of Fairy Lake west of Hull, was considered “a splen-
did locality” for this orchid (Eifrig 1909).

MORPHOLOGICAL VARIATION: Platanthera hyper-
borea is the most variable species in the District; for
example, the ratio of standard deviation to average
height is 0.5, appreciably greater than the ratios of
0.16 to 0.37 for the other species. This variation cor-
relates with habitat. Plants growing in mesic forests
have a restricted range of habit partly overlapping
the broader range of wetland plants. On average,
mesic forest plants are shorter, have fewer leaves
and flowers, and have somewhat shorter and more
lax inflorescences. They are generally slender, while
plants of wet habitats range from slender to stout.
The averages given below all differ between the
habitats at a statistically very significant level. The
blooming periods of the two groups are very similar.

ACHLOROPHYLLOUS FORM: The form P. hyperborea
(Linnaeus) Lindley var. hyperborea f. alba Light
was described in Light and MacConaill (1989) from
plants in Gatineau Park. In addition to these plants,
they also reported striped plants with green pigmen-
tation in part. Some of the achlorophyllous plants
flowered for at least two consecutive years.

On a subsequent visit to the site, we found some of
the achlorophyllous plants to be pale yellow in the
Methuen terminology (Kornerup and Wanscher
1978). The yellow colour, suggesting residual
carotenoid pigments, was visible in the absence of
chlorophyll. The plants were of the mesic forest habit
and were fairly small, although not much smaller
than chlorophyllous plants in the same colony.

Marilyn Light (personal communication 1996) has
been aware of achlorophyllous plants in this colony
for at least a decade. She has found individual plants
to be short-lived. She observed that whereas chloro-
plasts in the normal green leaves stained positively
for starch with iodine, those same structures in f.
alba did not. This observation provides an indepen-
dent demonstration of the absence of photosynthetic
activity, consistent with the lack of chlorophyll, in f.
alba.

Wet Habitats

slender slender to stout
23 (6-47)cm_ [115] 41 (20-87)cm [59]
6(2-12)cm [64] 9(3-19)cm_=_ [31]
3 (1 -7) [89] 7 (4- 13) [43]
10 (2 - 30) [87] 21 (5 - 60) [39]
1.3 (0.5 - 3.6) [61] Zil,t=3) [29]

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

133

Platanthera lacera (Michaux) G. Don in Sweet var. lacera

Ragged Fringed-orchid

SYNONYM: Habenaria lacera (Michaux) R. Brown

Platanthére lacéré

One of the more recent orchids to be discovered in the District, P. lacera is, on close inspection, an attractive
plant, but from a distance it is easily lost among the grasses and weeds with which it grows. It is distinguished
by its greenish flowers with long-fringed, tripartite lips. The spur is about | to 1.5 times as long as the lip.

DESCRIPTION

Height: 11 (30 - 53) 74 cm [127 plants].

Flowers: 4 (9 - 31) 56 [61 plants]; light green,
pale green, pale yellow, greenish white or yellowish
white; lip white at base and occasionally overall,
often with sepals more green than lip and petals;
flowers generally lighter than surrounding vegeta-
tion; laceration of lip quite variable in length and
regularity, occasionally almost lacking on central
lobe while present on side lobes; in a moderately
dense cylindrical inflorescence of some irregularity,
but in one case approximately five-ranked; fragrance
sweet and floral, becoming much stronger at sunset.

Leaves: | - 5, grading into an additional 2 - 4
bracts for a total of 4 - 7 leaves and bracts.

46°

45°
76° 30'

Overwintering State: a greyish green, broadly
conical shoot, appearing above ground beside the cur-
rent year’s stem, in late September; herbarium speci-
mens show the new shoot and partially elongated root
often present at anthesis. We have found that the pres-
ence of a green shoot does not necessarily mean that a
plant will grow from it the next year. The shoot may
disappear by early summer even while the roots are
still crisp and alive-looking. By September, however,
all of these roots will have disappeared.

Capsules: dark brown, ellipsoid to oblong, typi-
cally 1.4 x 0.4 cm, nearly erect; yield variable, aver-
aging 50% [11 plants].

Seeds: brownish orange to brown, released in late
September to early October.

‘

fo

Platanthera lacera var. lacera: @ = herbarium specimen, @ = Native Orchid Location
Survey sight record, A = literature reference. Areas of major sand deposits on the Lowlands

are shaded.

Vol. 111

THE CANADIAN FIELD-NATURALIST

134

y old-field habitat, Greenbelt, City of Gloucester, Regional Municipality of

FIGURE 33. Platanthera lacera var. lacera, sand

Ottawa-Carleton, Ontario, 6 July 1977.

1997

BLOOMING PERIOD: 28 June (2 July - 24 July) 8
August [31 records].

CoLony SIZEs: | - 123, typically to 60, flowering
plants; 10 or fewer in 70% of colonies [47 colonies];
as scattered individuals.

CURRENT STATUS: Platanthera lacera is generally
rare in Eastern Ontario (Whiting and Catling 1986)
and “occasionelle” in Western Quebec (Sabourin
1993) but is capable of becoming locally common
when ideal habitat is available. Interestingly, this is
more likely to happen as a result of human activity in
removing vegetation from sandy areas than by natur-
al means. The distribution map reflects the coinci-
dence of the work of the Native Orchid Location
Survey and the “population explosion” of this
species in the 1960s and 1970s.

DISTRIBUTION: The Ottawa District is at the regional
northern limit of the distribution of this orchid of the
Mixed and Deciduous Forest Regions. Within the
District, P. lacera is localized on sands deposited by
the Champlain Sea and subsequent rivers. In the
southeast the sands are mainly deltaic and estuarine
deposits of the Ottawa River, while those at other
sites in Stony Swamp - Bridlewood and Gatineau
Park are beaches and shallow deposits of the
Champlain Sea.

HABITATS: Two types of sand-based habitats support
colonies of this orchid, one in the open and the other
in shrubby or forested areas.

Platanthera lacera has been principally a species
of open, disturbed sites on moist, acidic sand
(pH 4.5 - 5.0) where vegetation is still sparse and sod
has not yet developed. In the 1960s, such sites were
abundant as a result of the formation of the
Greenbelt that left many abandoned fields and bor-
row pits. Further east, in the Larose Forest, there
were many broad, open, sandy roadsides that also
provided ideal habitat. But by the 1970s, the old-
fields were becoming covered with heavy shrub
growth, pine plantations were shading out the Larose
Forest, and roadsides in all parts of the District south
of the Ottawa River were being maintained by herbi-
cide spraying, leaving little opportunity for plants
other than stress-tolerant grasses to survive. Only for
a couple of decades was abundant suitable habitat
readily available, and now much of that habitat has
disappeared.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

135

Typical species accompanying P. lacera are Field
Horsetail (Equisetum arvense), Sensitive Fern
(Onoclea sensibilis), Marsh Fern (Thelypteris palus-
tris), Bladder Campion (Silene vulgaris), Dewberry
(Rubus hispidus), Meadowsweet (Spiraea latifolia),
Cow Vetch (Vicia cracca), Common Milkweed
(Asclepias syriaca), Heal-all (Prunella vulgaris),
Brown-eyed Susan (Rudbeckia hirta), goldenrods
(Solidago spp.) and, occasionally, Dwarf Grape
Fern (Botrychium simplex), Adder’s-tongue Fern
(Ophioglossum vulgatum), rushes (Juncus spp.),
Liparis loeselii and Platanthera clavellata.

Rarely, a few plants are found along mesic wood-
land edges and in clearings among Eastern White
Cedar or White Spruce in the Stony Swamp -
Bridlewood area. Also in this area, a few plants
sometimes occur in clearings in willow and alder
thickets. In the southeast, some plants grow in the
shade of moist to wet Red Maple forests with Lady
Fern (Athyrium filix-femina) and Sensitive Fern.

LocaL History: Platanthera lacera does not ap-
pear on any early plant lists for the District (Billings
1867; Fletcher 1880, 1893; Macoun circa 1911’).
Among the few herbarium specimens, the earliest
was collected in 1941 from a “mixed wood” near
Old Chelsea in Gatineau Park by H. A. Senn, W. A.
Minshall and M. N. Zinck [DAO 17175]. The next
was collected 27 years later from a roadside near
Ramsayville, City of Gloucester, by W. J. Cody
[18110 at DAO]. These and other Department of
Agriculture botanists had been botanizing the Ottawa
District seriously since the 1940s, but such was the
rarity of this orchid at that time that it was collected
only twice. Between 1965 and 1975, members of the
Native Orchid Location Survey searched the District
intensively and discovered 41 colonies, mainly in the
southeast.

ABERRATION: In 1970, we found two plants in a
colony on the Dolman Ridge beside the Mer Bleue
Bog that totally lacked fringes and lobes on the lips,
leaving the lips resembling those of P. hyperborea.
But all other flower parts, including the column as
well as the leaves and bracts, were characteristic of
P. lacera, and so we had to conclude that some
mutation had occurred to produce two plants with
fringeless lips. (Photograph in J. M. Reddoch and A.
H. Reddoch 1987c.) We did not see these two plants
again, nor have we observed this aberration since.

136

THE CANADIAN FIELD-NATURALIST

Vol. 111

Platanthera leucophaea (Nuttall) Lindley

Eastern Prairie Fringed-orchid

SYNONYM: Habenaria leucophaea (Nuttall) A. Gray

Platantheéere blanchatre

Platanthera leucophaea is the most recently discovered orchid in the District. It is a rare species that is restrict-
ed to fens, a habitat that is also becoming rare. It is one of our largest and most fragrant orchids. This species
can be identified by its large white flowers with tripartite, fringed lips and long spurs, as well as by its fen habi-

tat. The spur is about 2 to 2.5 times as long as the lip.

DESCRIPTION

Height: 28 (47 - 73) 93 cm [127 plants].

Flowers: 3 (6 - 17) 29 [60 plants]; sepals light
yellow, pale yellow or greyish green with darker
veins; lip and petals white often with pale green or
greenish white at base of lip, spur greyish yellow; in
an open, irregular inflorescence; fragrance in day-
time sweet and penetrating, like Common Milkweed
(Asclepias syriaca) or Dogbane (Apocynum
androsaemifolium), becoming much stronger in
evening.

Leaves: | - 5, typically 2 - 4, grading to bracts for
a total of 5 - 9 leaves and bracts.

46°

45°
76° 30

Overwintering State: a pale green shoot more
than 6 cm tall, 4 mm diameter, the greyish green,
acicular tip at the moss surface beside the current
year’s stem, in October; herbarium specimens show
the new shoot usually present at anthesis.

Capsules: brown, oblong, typically 1.7 x 0.4 cm,
ascending to erect (see Figure Ic); yield usually
greater than 50%, averaging 85% [8 plants].

Seeds: dark brown, released in mid to late
September.

BLOOMING PERIOD: 26 June (4 July - 22 July) 22
July [10 records].

75°

Platanthera leucophaea: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record. Major areas underlain by calcareous rock (marble and limestone) are shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 137

FIGURE 34. Platanthera leucophaea, sedge fen habitat, Goulbourn Township, Regional Municipality of Ottawa-Carleton,
Ontario, 10 July 1977.

138

COoLony SIZEs: | - 68 flowering plants [7 records],
as scattered (sometimes very widely scattered) indi-
viduals; up to an estimated 1000 in one wetlands
complex (see below); the number of flowering plants
varying widely from year to year.

CURRENT STATUS: rare in the Province of Ontario
(White et al. 1982c), very rare (S2) in Ontario
(Active List, Oldham 1996°); rare in Canada (Argus
and Pryer 1990); vulnerable (1996 list of The
Committee on the Status of Endangered Wildlife in
Canada (COSEWIC)); rare in the Ottawa District;
not known in Quebec (Sheviak and Bowles 1986;
Sabourin 1993).

DISTRIBUTION: The Ottawa District is part of a
northeastern disjunction of this orchid of the
Prairie, northern Deciduous Forest Region and
adjacent Mixed Forest Region. There is growing
realization that much of the present-day population
of the species is in southern Ontario (Sheviak and
Bowles 1986). Within the District, P. leucophaea is
known only in fen habitats in the Richmond
Wetlands Complex in the southwest. The next clos-
est extant colony (Greenwood 1968a) is in Leeds
County.

HABITAT: Calcareous sedge fens are the exclusive
habitat of P. leucophaea in Eastern Ontario. The
plants grow in the wettest parts of the fen, which
means they may be in ankle deep water in wet years
and in peat that is damp rather than surface-dry in
dry years. The peat is up to two metres deep and pHs
are in the range of 6.2 - 6.6. This orchid is accompa-
nied by typical fen species, including the co-domi-
nant sedges, Carex lasiocarpa and C. livida. At
some spots in the two local fens, as well as in the
Leeds County fen, Calopogon tuberosus, Liparis
loeselii and Pogonia ophioglossoides are also pre-
sent. Where it was first discovered at the edge of the
Richmond Fen, other wet habitat species were pre-
sent, such as Swamp Milkweed (Asclepias
incarnata), Northern Bugleweed (Lycopus uni-

THE CANADIAN FIELD-NATURALIST

Vol. 111

florus), Common Skullcap (Scutellaria galericulata),
Marsh Bedstraw (Galium palustre) and Spotted Joe-
Pye-Weed (Eupatorium maculatum).

LocAL History: Ewen Todd discovered P. leu-
cophaea in the Richmond Wetlands Complex in
1976. The colony consisted of several dozen flower-
ing plants at various places in the fen (Reddoch
1977a). No one knows how long P. leucophaea
has been in the Richmond Wetlands Complex, but
we do know that over a century ago the fen was
under water for a period of time when the Jock
(Goodwood) River was dammed (Belden 1879).

Two years after the initial discovery, we encoun-
tered a second colony in another fen in the wetlands
complex. The next year we found more plants in
another part of this large fen [DAO 691529]. Local
residents told us that this fen is known as “The
Burn” because area farmers in the early days used to
burn the fen each fall to drive out the deer.

In 1996, Don Cuddy (personal communication)
and members of the Ontario Ministry of Natural
Resources’ Environmental Youth Corps Program
studied P. leucophaea in the Richmond Wetlands
Complex. The results of their sampling led them to
estimate the population at between 800 and 1000
flowering plants.

About 1900, Frank Morris found this orchid “on
the margin of a mud lake near Smith’s Falls...”
(Morris 1920), but it had disappeared before 1929
(Morris and Eames 1929). There are several “Mud
Lakes” with shoreline fens near Smiths Falls, but P.
leucophaea is not currently known from any of them
(Don Cuddy, personal communication 1986). Smiths
Falls is just beyond the Study Area to the southwest.

The earliest evidence for the existence of the
Leeds County colony is a collection by W. J. Cody,
W. G. Dore and J. H. Soper in 1956 [DAO 17187].
Luer (1975) included illustrations of an inflorescence
and a flowering plant in this fen as photographs #3
and #4 on Plate 47.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 139

Platanthera macrophylla (Goldie) P. M. Brown

Goldie’s Round-leaved Orchid

SyNONYM: Habenaria macrophylla Goldie

Platanthere orbiculaire de Goldie

Platanthera macrophylla can be an impressive sight with its bright and sturdy inflorescence floating above two
large green leaves in the shade of the forest where little else grows. However, only one plant has been seen in
the District. Platanthera macrophylla forms a species pair with P. orbiculata and both share the distinguishing
features of two large roundish leaves flat on the ground, a stem with a few cauline bracts and an inflorescence
of long-spurred, greyish green and white flowers. Platanthera macrophylla can be separated most reliably from
P. orbiculata by the average length of its spurs, which is 28 mm or more, and also by the average length of its
hemipollinaria, which is 4.6 mm or more. The two species do not differ significantly in height or, in spite of the
epithet macrophylla, in leaf size. For more details, see the P. orbiculata account and Reddoch and Reddoch
(1993). Given the small number of plants found in or near the District, additional herbarium records from an
area bounded by latitudes 45° and 47°, and longitudes 73° and 77°, were included for the heights, flower num-
bers and blooming dates given below. Most of these specimens were from north or south of Montreal and

included the type specimen from the Island of Montreal (K!).

DESCRIPTION

Height: 34 (35 - 51) 63 cm [18 plants].

Flowers: 9 (9 - 21) 33 [16 plants]; sepals greyish
green often with white margin, lateral sepals white at
base; petals and lip white, sometimes greyish green
toward the tips; spur translucent, white, greyish
green distally; in a lax, regular to irregular inflores-
cence; fragrance not detected by day.

46°

45°
76° 30'

Leaves: 2, basal, shiny, deep green, silvery
below.

Overwintering State: likely similar to P. orbicu-
lata; herbarium specimens show the new shoot pre-
sent at anthesis.

Capsules: light brown to brown, ellipsoid to
oblong, typically 2.1 x 0.5 cm, vertical even on a
sloping rachis (see Figure 1d).

,

75°

Platanthera macrophylla: ® = our sight record and photographs (see text). The Canadian

Shield is shaded.

140 THE CANADIAN FIELD-NATURALIST Vol! 1411

CTA
NN:

aint
= sti
Cy MINN

FIGURE 35. Platanthera macrophylla, mixed forest habitat, Gatineau Park, Quebec, 15 July 1973; the only record known
for the Ottawa District.

1997

Seeds: brown to dark brown, released in mid
October.

BLOOMING PERIOD: 2 July (5 July - 25 July) 6
August [16 records].

CoLony SIZEs: usually single plants, rarely more.

CURRENT STATUS: rare in the Provinces of Quebec
(Bouchard et al. 1983) and Ontario (Reddoch et al.
1982), very rare (S2) in Ontario (Watch List,
Oldham 1996"); rare or extirpated in the Ottawa
District (no extant colony known).

DISTRIBUTION: The Ottawa District is near the mid-
dle of the range of this species of the Mixed Forest
Region (Reddoch and Reddoch 1993). Within the
District, the only recorded location of P. macrophyl-
la is on the Canadian Shield.

HABITAT: The Ottawa District habitat was similar to
the typical habitat for the species across its range. It
was a fairly mature forest of Sugar Maple and
Beech, with a few trees of Eastern Hemlock and
Largetooth Aspen, on a low slope beside a beaver
pond. There was little herbaceous cover and the
mesic forest floor was under a thick layer of leaf
mould. This type of habitat is very common on the
Shield but one can walk through many kilometres of
forest without finding any P. macrophylla.

LocaL History: As far as we can determine, only

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

141

one plant of P. macrophylla has been reported in the
District. We discovered this plant in 1972 as a pair
of leaves on the forest floor near the Champlain
Lookout in Gatineau Park, about 25 m from a group
of P. orbiculata that we were monitoring (see P.
orbiculata account). In 1973, the plant produced a
scape 36.5 cm tall with 11 flowers; it is shown in
the Figure. The following year, the plant again pro-
duced 11 flowers, this time on a scape 42 cm high.
In both 1973 and 1974, the leaves measured 12.5 cm
across. In 1975, the plant produced two healthy
leaves but did not flower. After that, we could not
find the plant again.

A few kilometres east of the Study Area near
Montebello, Quebec, William Scott collected a plant
in 1890 [TRT 189735].

Patricia Rothschild discovered and photographed
a plant several kilometres southwest of the Study
Area in Lanark County in the early 1980s. In 1983
and 1984 we examined one flowering plant and three
non-flowering plants at the edge of the Eastern
White Cedar - Tamarack - Black Ash swamp. The
plants were growing on a hummock with the moss
Hylocomium splendens about 30 cm above the water
level (photographs at DAO). Patricia told us in 1989
that no plants bloomed in 1985 or 1986 and the
colony disappeared soon after, likely the result of the
site being logged.

142

THE CANADIAN FIELD-NATURALIST

Vol 111

Platanthera obtusata (Banks ex Pursh) Lindley

Blunt-leaf Orchid

Platanthere a feuille obtuse

SYNONYM: Habenaria obtusata (Banks ex Pursh) Richardson

Platanthera obtusata is a small green orchid, widespread in the District but restricted to swamps. The single
blunt, basal leaf, with or without a cauline bract, and the relatively small, green and white flowers with short

tapering spurs serve to identify this species.

DESCRIPTION

Height: 6 (10 - 17) 25 cm [122 plants].

Flowers: 2 (4 - 12) 20 [116 plants]; sepals light
green or greyish green with colourless edges, petals
white often with light green along upper edges and
veins, lip and spur greyish green with white at base;
in an irregular inflorescence; fragrance lacking or
faint, sweet or sharp.

Leaf: single, basal and blunt, 10 - 15% of plants
with a smaller, narrow cauline bract, in one plant the
bract being only about 2 cm above the basal leaf and
larger than usual suggesting a second, smaller leaf.

Overwintering State: a greyish green, conical
shoot appearing above ground beside the current
year’s stem by October; Currah, Smreciu, and
Hambleton (1990) describe the underground season-

46°

al development of P. obtusata in Alberta: at dorman-
cy, the new shoot is subtended by a new, fully elon-
gated, tuberous root and two fleshy roots; a new
plant begins development a year before flowering.

Capsules: light brown, ellipsoid, typically 0.6 x
0.3 cm, ascending (see Figure Id); yield highly vari-
able, averaging 55% [16 plants], appreciably greater
than the 14% average reported from Wisconsin for
326 plants (Thien and Utech 1970).

Seeds: greyish orange, released by early October.

BLOOMING PERIOD: 10 June (20 June - 14 July) 25
July [36 records].

CoLony SIZEs: 1| - 2000, typically to 200, flowering
plants [77 colonies], as scattered individuals and
sometimes in clumps of four or five.

45°
76° 30'

75°

Platanthera obtusata: @ = herbarium specimen, ®@ = Native Orchid Location Survey sight

record. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 143

FIGURE 36. Platanthera obtusata, mixed swamp habitat, La Péche Municipality (Low Township, Gatineau County),
Quebec, | July 1980.

144

DISTRIBUTION: The Ottawa District is near the
southern boundary of the North American distribu-
tion of this transcontinental orchid. It is a species of
the Montane, Boreal and Mixed Forest Regions, as
well as the Tundra. Within the District, P. obtusata
is scattered across the Shield and more sparsely on
the Lowlands.

HasiratT: Platanthera obtusata frequents the Eastern
White Cedar - White Spruce - Black Ash swamps that
are common in the District especially in areas of cal-
careous bedrock. These swamps range in character
from fairly closed and dense to high-canopied and
open. Plants on the shaded, moist to saturated, swamp
floors grow among swamp mosses or in coniferous
and deciduous leaf mould. Companion species may
include Oak Fern (Gymnocarpium dryopteris), Jack-

THE CANADIAN FIELD-NATURALIST

Vol. 111

in-the-pulpit (Arisaema triphyllum), Wild Lily-of-the-
valley (Maianthemum canadense), Malaxis mono-
phylla, Platanthera hyperborea, Foamflower (Tiarella
cordifolia), Dewberry (Rubus hispidus), Wood-sorrel
(Oxalis acetosella), Kidney-leaved Violet (Viola reni-
folia), Wild Sarsaparilla (Aralia nudicaulis),
Bunchberry (Cornus canadensis), One-flowered
Wintergreen (Moneses uniflora), Twinflower
(Linnaea borealis) and tree seedlings.

EARLY History: The earliest reference to P.
obtusata comes from John Macoun’s circa 1911°
manuscript. He cited only one collection - his own,
which he made in 1903 in a swamp near Wakefield,
Quebec [Macoun 61032 at CAN]. This swamp may
be Chilcott’s Swamp, where other collections of
this species were made in 1940, 1959 and 1960.

17

Platanthera orbiculata (Pursh) Lindley
Large Round-leaved Orchid

SyNnonyM: Habenaria orbiculata (Pursh) Torrey

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

145

Platanthére a feuilles orbiculaires

Platanthera orbiculata can be quite distinctive with its greenish white inflorescence rising from between two
large green leaves. While it has been recorded widely on the Shield, it is not a common sight. Platanthera
orbiculata forms a species pair with P. macrophylla and both share the distinguishing features of two large
roundish leaves flat on the ground, a stem with a few cauline bracts and an inflorescence of long-spurred, green
and white flowers. Platanthera orbiculata can be separated from P. macrophylla by the differences in the aver-
age lengths of their spurs and hemipollinaria. In P. orbiculata, the average spur length is less than 28 mm, and
the average hemipollinarium length is less than 4.6 mm. The two species do not differ significantly in height or,
in spite of the epithet macrophylla, in leaf size. For more details, see Further Diagnostic Notes below and

Reddoch and Reddoch (1993).

DESCRIPTION

Height: 18 (24 - 38) 58 cm [82 plants].

Flowers: 3 (7 - 17) 27 [68 plants]; sepals greyish
green often with white margin, lateral sepals white at
base; petals and lip white, sometimes greyish green
toward the tips; spur translucent, white, greyish
green distally, often horizontal but sometimes
descending; in a lax to dense, regular inflorescence;
fragrance often undetectable but occasionally faint
_and sweet in late afternoon, flowers collected and

46°

45° 4
76° 3

kept in water moderately fragrant after dark although
not at dusk.

Leaves: 2, basal; 1 - 2 on non-flowering plants;
shiny, deep green, silvery below, approximately cir-
cular, but ratio of length-to-width ranging from 0.6
to 2.5; bracts 1 - 4, typically 3 - 4.

Overwintering State: a greyish green, broadly
conical shoot, 1 - 2 cm above ground beside the cur-
rent year’s stem, appearing above ground in late
September; Currah, Smreciu, and Hambleton (1990)

75°

Platanthera orbiculata: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

146 THE CANADIAN FIELD-NATURALIST Vol 111

FIGURE 37. Platanthera orbiculata, plant: coniferous forest habitat, La Péeche Municipality (Masham Township, Gatineau
County), Quebec, 14 July 1979; flowers: mixed forest, Gatineau Park, Quebec, 9 July 1970.

1997

describe the underground seasonal development of
P. orbiculata in Alberta: at dormancy, the new shoot
is subtended by a new, fully elongated, tuberous root
and 5 - 6 fleshy roots; a new plant begins develop-
ment a year before flowering.

Capsules: light brown to brown, ellipsoid to
oblong, typically 1.5 x 0.45 cm, erect; yield usually
less than 50%, averaging 30% [23 plants].

Seeds: dark brown, released in early October.

BLOOMING PERIOD: 30 June (6 July - 22 July) 10
August [41 records].

CoLony SIZEs: | - 190 flowering and non-flowering
plants; 10 or fewer plants in 79% of colonies [52
colonies]; usually as scattered individuals but some-
times in pairs and small groups; non-flowering
plants typically 1 - 3, but occasionally 5 - 10, times
as numerous as flowering plants.

DISTRIBUTION: The Ottawa District is well within
the distribution of this transcontinental orchid. It is a
species of the Montane, Boreal and Mixed Forest
Regions (Reddoch and Reddoch 1993). Within the
District, P. orbiculata is confined almost completely
to the Canadian Shield.

HABITATS: Platanthera orbiculata is a plant of
coniferous swamps and of a few coniferous and
mixed forests. The largest colonies occur in swamps;
usually only small numbers are found in forests.

The coniferous swamps are usually of some con-
siderable age. Widely spaced clumps of large
Eastern White Cedars are accompanied by a few
White Spruce, Tamarack, Black Ash and other trees.
The humus of the swamp floor is usually moist
rather than wet with the water table below the sur-
face. Only sometimes is the soil saturated. In places,
the floor is covered with patches of swamp mosses
and in others is bare except for layers of fallen
leaves. The herbaceous layer is scant or is luxuriant
with such species as Bulblet Fern (Cystopteris frag-
ilis), Crested Wood Fern (Dryopteris cristata), Oak
Fern (Gymnocarpium dryopteris), Bluebead-lily
(Clintonia borealis), Platanthera obtusata,
Foamflower (Tiarella cordifolia), Wood-sorrel
(Oxalis acetosella), Wild Sarsaparilla (Aralia nudi-
caulis), Bunchberry (Cornus canadensis) and One-
flowered Wintergreen (Moneses uniflora).
Platanthera orbiculata plants grow on mounds at the
bases of trees as well as on the swamp floor.

A few fairly mature forests of Sugar Maple and
Beech with scattered conifers or of mainly conifers,
particularly Eastern White Cedar, occasionally shel-
ter up to a couple of dozen P. orbiculata plants. The
plants are rooted in moist leaf mould and sandy loam
where there is little other vegetation.

LONG-LIVED COLONIEs: In 1969 we discovered and
began monitoring a group of P. orbiculata plants in a
mixed forest near the Champlain Lookout in

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

147

Gatineau Park. The plants, ultimately numbering
five, grew close together in a roughly curved line
separated from each other by 3 to 7 cm. Initially
there were two plants, one flowering and one not.

The already vigorous non-flowering plant pro-
duced two leaves in 1969 and in the following 10
years. In that decade, it flowered in six of the next
eight years, including four consecutive years. It did
not appear in the twelfth year and then had one leaf
for three more years (until 1983), after which it was
not seen again.

The plant that flowered in 1969 continued as a
two-leaved plant for six more years, blooming once
more, in 1973, and remained as a one-leafed plant
for four more years (until 1979). It did not come up
after that. Its decline may have been hastened by
being often under the large leaves of the adjacent,
more vigorous plant described above.

A third plant appeared in 1971 as one leaf beside
the preceding plant. It did not come up the next year,
put up one leaf for the next three years and then was
two-leaved for one year before attempting to flower
in the seventh year as a 2.5 cm stunted scape. It
flowered again the year after and, after a one-year
absence, for the next two years. Sometime during the
last flowering, the scape and one leaf were ripped
off, likely by a deer. The plant lasted as two leaves
for four of the next six years (until 1987) and then
disappeared.

A fourth plant came up in 1978 beside the third
plant, as one leaf for five of the next seven years,
with two absences, and the last two years with two
small leaves (until 1987). It never flowered.

A fifth plant produced one leaf each year from
1981 to 1983 and did not flower.

The apparently sudden disappearance of the third
and fourth plants at the two-leaf stage in 1987 may
have been the result of tree removal by Beavers that
exposed the colony to sunlight. We have seen no
plants in the area since.

In summary, we can conclude from this study that
plants can live for 15 and probably at least 20 years
or for as few as three years. The general pattern of
development is to produce one small leaf each year
for several years, move to two large leaves each year
during the plant’s prime (which can last at least as
long as 11 years), and then end with one small leaf
each year for several more years. In their prime,
plants can flower at least six times in eight years
with four of those years in succession. Once or twice
at some random time in even a vigorous plant’s life,
a plant will fail to come up at all for one year. Leaf
sizes, scape heights and flower numbers are at their
greatest in the middle of the plant’s life.

In 1966, Ed Greenwood (records of the Native
Orchid Location Survey) discovered a very large
colony of P. orbiculata in an Eastern White Cedar -
Tamarack - White Spruce swamp that is part of the

148

Manion Corners Long Swamp in Ottawa-Carleton.
This colony declined drastically in the 1970s after
this part of the swamp was logged in reaction to the
designation of the wetland as a Natural Environment
Area in the Region’s Official Plan. Several dozen
plants survive to the present in the nearby unlogged
portions of the swamp. Other colonies, totalling sev-
eral hundred plants, have been noted elsewhere in
this peatland over the years.

EARLY History: Platanthera orbiculata was one of
the orchids collected by John Kerr McMorine at
Ramsay, Ontario, in 1862 [QK 13099, 13100].
Another early collection was James Fletcher’s 1878
specimen from the “Chelsea Mountains, P.Q.”, now
Gatineau Park [DAO 200660]. Fletcher (1893) cited
Rockcliffe and Beechwood as other locations for this
species, but we have found no supporting collec-
tions. Early botanists considered this species to be
rare in the District (Fletcher 1893; Macoun circa
1911°).

FURTHER DIAGNOSTIC NOTEs: A better separation of
P. orbiculata and P. macrophylla can be made
through a discriminant function, in this case the sum

THE CANADIAN FIELD-NATURALIST

Vol. 111

of the average spur length and twice the average
hemipollinarium length. This sum will be less than
38 mm for P. orbiculata. Numerical studies
(Reddoch and Reddoch 1993) show that none of the
measured characters permit a perfect separation of
these species, the discriminant function being about
99.7% accurate, while the average spur length alone
is about 99.0% accurate.

The discriminant function can be useful for deal-
ing with certain plants growing in swamps. Swamp
plants tend to have fairly large flowers, although still
within the norms of the species, and to have relative-
ly long spurs, but a few of them have average spur
lengths exceeding the critical value of 28 mm by | or
2 mm. For most of these long-spurred plants, the dis-
criminant function will be less than 38 mm and they
will key out to P. orbiculata; however, for a few, it
will be 38 mm or greater. While these marginally
long-spurred plants, in isolation, key out as P.
macrophylla, it is best to assign them to the same
species as the rest of the colony. They lie in the
small area of intrinsic overlap in the characters of the
two species that causes the less than 100% accuracy
noted above.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

149

Platanthera psycodes (Linnaeus) Lindley

Small Purple Fringed-orchid

SyNONYM: Habenaria psycodes (Linnaeus) Sprengel

Platanthére papillon

Platanthera psycodes is one of our most distinctive orchids and among our tallest. It is widespread in the
District, but it is not as abundant as the map suggests because many colonies are rather short-lived. Platanthera
psycodes closely resembles P. grandiflora, with which it occasionally grows. These two species can be easily
recognized by their numerous purple fringed flowers on a leafy stem. Platanthera psycodes is distinguished
from P. grandiflora by the shape of the entrance to the spur and by the appearance of the anther sacs. The
entrance to the spur in P. psycodes is flattened, sometimes almost divided into two smaller entrances, rather
than being large and round. The anther sacs are close and parallel instead of diverging downward. Platanthera
psycodes also tends to have flowers only about half as large as P. grandiflora.

DESCRIPTION

Height: 24 (42 - 74) 101 cm [194 plants].

Flowers: 7 (14 - 46) 100 [111 plants], on average
about one third more than P. grandiflora; commonly
lilac or light lilac but also violet white, pale violet or
purple; pure white except for pollinia in f. albiflora
(see Acyanic Form below); base of lip pure white,
spurs translucent becoming similar to lip colour near
the tip, occasionally darker presumably from small
concentration of chlorophyll; fragrance usually
daisy-like, sometimes rich, rose-like, but occasional-

46°

ly musty or even like a cat’s spray, the latter perhaps
related to Brackley’s (1985) experience.

Leaves: 2 - 5 (typically 3 - 4) with 2 - 8 (typically
3 - 6) bracts; 2 - 3 on non-flowering plants.

Overwintering State: a greyish green, broadly con-
ical shoot, 0.5 - 2 cm above ground beside the current
year’s stem, appearing above ground in late
September; herbarium specimens show the new shoot
and partially elongated roots present at anthesis.

Capsules: brown, oblong, typically 1 x 0.25 cm,
ascending to erect; vertical or nearly so even on bent

45°
76° 30

75°

Platanthera psycodes: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, A = literature reference. The Canadian Shield is shaded.

150 THE CANADIAN FIELD-NATURALIST Vol. 111

FIGURE 38. Platanthera psycodes, plant: old-field/stream edge habitat, Aylmer Municipality, Quebec, 4 August 1980;
flowers: deciduous forest/stream flood plain habitat, Gatineau Park, Quebec, 22 July 1972.

Foo?

or fallen stems; yield highly variable, averaging 65%
[22 plants].

Seeds: dark brown, released in mid to late
September.

BLOOMING PERIOD: 23 June (11 July - 2 August) 18
August [121 records], more extended and two to
three weeks later than for P. grandiflora.

COLONY SIZEs: | - 118, typically to 70, flowering
plants; 10 or fewer in 71% of colonies [250 colonies],
primarily as scattered individuals.

DISTRIBUTION: The Ottawa District is in the northern
part of the range of this eastern orchid. It is a species
of the Mixed Forest Region and adjacent Boreal and
Deciduous Forest Regions (Stoutamire 1974).
Within the District, this species is found most com-
monly on the Canadian Shield and in the southeast
on the sands deposited by the Champlain Sea and
subsequent rivers.

The number of sight records shown on the distrib-
ution map is over-represented compared with the
other orchids in the District. Many sightings were
the byproduct of Ross Layberry’s intensive searches
of roadside sedge patches for rare butterflies in the
early 1980s.

HABITATS: Colonies are encountered most often
among grasses and sedges in sunny, moist or wet
roadside ditches, in old-fields and at the margins of
ponds, streams and rivers. They also occur among
ferns in partially-shaded stream valleys and depres-
sions in fairly mature deciduous forests. (See
description of the latter habitat under P. grandiflo-
ra.) Occasionally, there are plants in alder thickets
and in the edges of swamps. The soils are usually
moderately acidic sands or sandy humus, but some-
times, on the Lowlands, the soil is Leda clay. In
forests and swamps, the substrate is often highly
organic.

Open field plants tend to be shorter and stouter
and to have more flowers and paler, yellower leaves
than plants growing in wetter, more shaded sites.

EARLY History: John Kerr McMorine collected this
species in 1862 at Ramsay, Ontario [QK 13119]. A
few years later, Braddish Billings Jr. (1867) included
it in his list of species collected during the summer
of 1866, but we have not found a voucher for verifi-
cation. Other early collections were made by Robert
B. Whyte in 1875 (Macoun circa 1911°), Henry M.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

151

Ami in 1879 [MTMG 3383, 26283] and James
Fletcher in 1881 [MTMG 25529].

In 1917, W. Hague Harrington had this to say:
“The tall purplish spikes of bloom appear some sea-
sons in great abundance and beautify the low mead-
ows and roadsides which are their usual habitat. The
Beaver Meadow, Hull, P.Q., was often richly
adorned in July with these charming plants but, alas!
the devastation and ravages of the extensions of Hull
are fast destroying the beautiful scenes to which the
Field-Naturalists’ Club frequently resorted to study
the rich fauna and flora”.

ACYANIC Form: The white-flowered P. psycodes f.
albiflora (R. Hoffman) Whiting and Catling has
turned up at least four times in the last century, each
time in the southern Quebec part of the District. The
first report was in 1888: “A beautiful albino form of
Habenaria psycodes, Gray, [sic] was collected this
past summer near Ironsides by Mr. H. M. Ami. It
was growing amongst a number of plants of the ordi-
nary colour and was of vigorous growth.” (Fletcher,
Small, and Baptie 1888). The whereabouts of this
collection is not known at present.

Ed Greenwood observed two such plants near
Buckingham in 1962 (personal communication
1977). Some 20 years later, the site had become
overgrown due to succession and we could find no
P. psycodes there. In 1980, Ross Layberry discov-
ered the albino form in a colony of about 35 typical
plants along a stream edge in a farm field south of
Mountain Road near lower Gatineau Park. At this
colony we saw two plants with pure white flowers
and buds (A. H. Reddoch and J. M. Reddoch 1987b,
[DAO 691538]) and one other plant with white flow-
ers and buds but with a very pale pinkish colour on
the uppermost flowers and the outsides of the buds.
The site of the colony, by a stream at the edge of a
field, had evidently been cut and ploughed in previ-
ous years and was ploughed again a few years later.
In 1982, Clarie Frankton (personal communication
1987) found two albinos with a number of typical
plants in the southern part of Gatineau Park.

ABERRATION: On some plants that we examined in
1976 in Cumberland Township, the flower petals
were appreciably lighter than the sepals, giving a
two tone effect. At that time, the roadside ditches
where these plants grew were being sprayed for
weed control. (See Long-lived Colonies under P.
grandiflora for a further description of this colony.)

152

THE CANADIAN FIELD-NATURALIST

Vol. 111

Pogonia ophioglossoides (Linnaeus) Jussieu

Rose Pogonia

Pogonie langue-de-serpent

Pogonia ophioglossoides is one of the three pink or purple orchids to be found in the fens of the District. It can
be separated from the other two, Calopogon tuberosus and Arethusa bulbosa, by the elliptical cauline leaf, the
similar but somewhat smaller floral bract, the lip in the usual lower position, in contrast to that of C. tuberosus
and the spreading lateral sepals, in contrast to the erect ones of A. bulbosa.

DESCRIPTION

Height: 10 (18 - 32) 62 cm [264 plants].

Flowers: 1, occasionally 2; sepals and petals usu-
ally purplish pink, sometimes more or less intense:
purplish red, greyish magenta, pink or pinkish white;
margin of lip similar with more intense lines; centre
of lip with light yellow or lighter papillae near the
end and finer fibres of similar or greener colour near
the base; fragrance moderate, sweet, strawberry-like,
or lacking; Luer (1975) displays a photograph (Plate
65, #4) of a typical flower in the open sedge fen near
Poltimore, Quebec.

Leaf: 1, cauline, rarely also with 1 basal, petio-
late, lanceolate leaf; 1 or 2, petiolate, lanceolate, on
non-flowering plants.

Overwintering State: a greyish green shoot, | -
1.5 cm high, at the base of the current year’s stem

about 4 cm deep in the moss substrate, in September;
herbarium specimens show the new shoot and par-
tially elongated roots present at anthesis.

Capsules: brownish orange to light brown, ellip-
soid to oblong, typically 2.1 x 0.5 cm, vertical even
on a sloping rachis, flowers frequently dehiscent
from capsule; yield usually high, 50% or more [60
plants] (10% - 100% in Wisconsin (Thien and
Marcks 1972)).

Seeds: brown, released in late September.

BLOOMING PERIOD: 8 June (23 June - 17 July) 14
August [79 records].

COLony SIZEs: | - 2500, typically to 1000, flower-
ing plants [40 colonies], as scattered individuals.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the range of this eastern orchid. It is a

46°

45°
76° 30!

75°

Pogonia ophioglossoides: @ = herbarium specimen, @ = Native Orchid Location Survey
sight record, & = literature reference. The Canadian Shield is shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT bS3

FIGURE 39. Pogonia ophioglossoides, left plant: moist, sandy borrow pit, Borthwick Ridge, Mer Bleue Conservation Area,
Ontario, 26 June 1973; right plant: sedge fen, Val-des-Monts Municipality (Wakefield Township, Gatineau
County), Quebec, 16 July 1977; flower: sedge mat, Black Lake, Gatineau Park, Quebec, 9 July 1969.

154

species of the Mixed, Deciduous and Southeastern
Coastal Plain Forest Regions. Within the District, P.
ophioglossoides is confined to several peatlands and
related areas scattered across the Canadian Shield
and the Lowlands.

HaBITATS: Open sedge fens are the most common
habitat for P. ophioglossoides and the habitat where
it occurs in largest numbers. Both calcareous sedge
fens (intermediate to rich fens) and the minerotroph-
ic edges of bogs (poor fens) are occupied, as are
floating pond-edge fens in both categories. The
orchids grow on the wet fen floor among such moss-
es as Sphagnum warnstorfii and S. magellanicum.
Calopogon tuberosus almost always accompanies P.
ophioglossoides in these habitats. (For more details
on accompanying calcareous sedge fen vegetation,
see Appendix 2.)

Certain borrow pits, shore-edge fallen logs and
floating boom-logs also have provided suitable habi-
tats for P. ophioglossoides.

During the 1970s, a colony flourished on the moist
to wet floor of a borrow pit on the Borthwick Ridge
beside the Mer Bleue Bog. Removal of overburden a
decade earlier left a layer of bare sand over imperme-
able clay that was then colonized by both peatland
and old-field plants. A patch of P. ophioglossoides
developed in one of the more low-lying areas (pH
5.0), along with saplings of Tamarack, Trembling
Aspen, Grey Birch and Speckled Alder. Prominent
herbaceous plants included Field Horsetail
(Equisetum arvense), Round-leaved Sundew
(Drosera rotundifolia), Meadowsweet (Spiraea
alba), Red Clover (Trifolium pratense), Sheep Laurel
(Kalmia angustifolia), Labrador Tea (Ledum groen-
landicum) and Small Cranberry (Vaccinium oxycoc-
cos). Polytrichum commune was the common moss.
The colony expanded to more than a hundred flower-
ing plants and many more non-flowering plants in the
years 1977 to 1980. During that time, Rose Chafers
(Macrodactylus subspinosus) infested the colony and
ate the flowers. By 1987, the area was so overgrown
that it was difficult to recognize and there was only
one flowering plant left.

Meanwhile, in a similar, but slightly drier, habitat
on the Dolman Ridge beside the Mer Bleue Bog, 11
flowering plants of P. ophioglossoides appeared for
one year (1973), growing along with Calopogon
tuberosus and other orchids. Although there were
hundreds of flowering plants of P. ophioglossoides
in poor fen habitat across the lagg to provide a con-
tinual source of new seed, subsequent dry summers
and increased growth of shrubs and trees quickly
made this site unsuitable for P. ophioglossoides, and,
ultimately, for the rest of the orchids. (See
Calopogon tuberosus account.)

Somewhat waterlogged and decayed fallen trees
along undisturbed lake edges can provide suitable
habitat for P. ophioglossoides. At Lac Bonin, in

THE CANADIAN FIELD-NATURALIST

Vol. 111

Quebec cottage country, two dead, partially floating
Eastern White Cedar trees supported a small colony
of this orchid for at least two decades, from 1973 to
1993 (Ed Dodson, personal communication 1996).
Although the habitat did not change, no flowering
plants were present in 1994 and 1995 (idem). The
trees remained anchored to a shore lined with Sweet
Gale (Myrica gale) and Leatherleaf (Chamaedaphne
calyculata) while vegetation developed at two places
on them over the years. In 1983, when Ed Dodson
showed the site to us, we counted 10 flowering
plants of P. ophioglossoides and two of Liparis loe-
selii. They were accompanied by Royal Fern
(Osmunda regalis) (< 20 cm high), Wild Iris (/ris
versicolor), Sweet Gale, Pitcher-plant (Sarracenia
purpurea), Round-leaved Sundew, Marsh St. John’s-
wort (Triadenum virginicum), Leatherleaf (< 40 cm
high), Cut-leaved Bugleweed (Lycopus americanus)
seedlings and Red Maple seedlings. Campylium stel-
latum, Sphagnum contortum and S. turgescens were
the mosses present.

Partially waterlogged and decayed boom-logs on
the Gatineau River between Chelsea and Cascades
have provided habitat for many local plants, includ-
ing Liparis loeselii (Bayly 1972) and P. ophioglos-
soides (Isabel Bayly, personal communication 1977).
Boom-logs were large squared timbers of Douglas-
fir, Western Hemlock or Western Red Cedar (Bayly
1972) that were chained together to restrain cut logs
en route to the mills on the Ottawa River. Most
plants living in this harsh environment of glaring sun
and low fertility remained dwarfed in size, even
while becoming fully mature (Bayly 1972); howey-
er, the P. ophioglossoides plant collected by Martha
Camfield in 1959 at Ramsay’s Crossing (between
Burnet and Cascades) was a relatively normal 23 cm
high [CCO 3482]. About a decade ago, when trucks
became the sole means of transporting cut logs, the
boom-logs were removed (Isabel Bayly, personal
communication 1996).

LONG-LIVED COLONY: On 8 July 1878, James
Fletcher [1791 at DAO] collected P. ophioglossoides
at Black Lake in what is now Gatineau Park. He prob-
ably made the collection from the partially floating,
poor fen mat (pH 5.0) at the southeast end of the lake.
The colony of P. ophioglossoides is still extant. We
counted 80 flowering plants there in 1969; however,
Beavers dammed the outlet stream in 1983, raising the
lake level half a metre and inundating much of the
mat (J. M. Reddoch and A. H. Reddoch 1987d). From
the shore in 1995, we spotted 37 flowering plants on
the remnant mat; in 1996 we were able to see 19.
Accompanying vegetation in 1996 included Marsh
Fern (Thelypteris palustris), Water Arum (Calla
palustris), sedges (Carex spp.), Wild Iris, Marsh St.
John’s-wort, Leatherleaf and Sphagnum spp.

EARLY History: Braddish Billings Jr. collected P.
ophioglossoides at Dow’s Swamp on 9 July 1860

1997

[QK 13410] and on 8 June 1861 [QK 13412, -13,
CAN 227241], likely on the sedge mat surrounding
the small pond shown in the 1925 air photo
(Reddoch 1978b). He collected Calopogon tuberosus
there on the former date and noted that both species
were rare. He did not include P. ophioglossoides in
his list of plants collected during the summer of
1866 (Billings 1867), although he did have C.
tuberosus. There appear to be no other collections
from Dow’s Swamp.

In 1862, J. K. McMorine collected both P.
ophioglossoides and C. tuberosus from Ramsay,

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

155

Ontario. The date cited for the former species is 9
July [QK 66723, 66727].

Robert B. Whyte’s 1879 collection from the Mer
Bleue Bog cited by John Macoun (circa 1911°)
appears to be the earliest of many collections made
over the years from the west end of the peatland
between Blackburn Hamlet on the north and
Carlsbad Springs on the south. Several early
authors commented on the great abundance of P.
ophioglossoides in the Mer Bleue (Macoun, Whyte,
and Fletcher 1884; Fletcher 1893; Macoun circa
1911°).

156

THE CANADIAN FIELD-NATURALIST

Vol. 111

Spiranthes casei Catling and Cruise var. casei

Case’s Ladies’-tresses

Spiranthe de Case

One of the latest orchids to bloom, S. casei was first collected here only a half century ago and was not recog-
nized as a separate species until two decades ago (Catling and Cruise 1974). Spiranthes casei is distinguished
from the more abundant and widespread S. cernua by its off-white rather than pure white flowers that are less
than 8 mm long and are arranged in a single, corkscrew-like spiral.

DESCRIPTION

Height: 14 (21 - 32) 45 cm [211 plants], on aver-
age about 4 cm taller than S. cernua.

Flowers: 4 (13 - 26) 50 [178 plants]; yellowish
white, rarely almost white, with a greyish yellow
patch at the centre of the lip visible from above and
below, usually with greyish green to deep green at
the base of the tepals; more or less horizontal; in a
single corkscrew spiral of up to 8 turns, rising to the
left or right, with 3 - 8, typically 5 - 6 flowers per
turn; no odour detectable, but a strong sour odour
when bruised or collected.

Leaves: 3 - 4, basal, withering to leave 0 - 3 at
anthesis; and 1, cauline, near the base, sometimes
bract-like, starting to wither at anthesis.

Overwintering State: a lanceoloid shoot or a pair

of leaves | - 2 cm in length surrounding a central
bud, appearing above ground beside the current
year’s stem at anthesis or shortly after; shoots
appearing earlier and developing further on decapi-
tated and non-flowering plants; Mousley (1924b, sub
S. cernua var. ochroleuca) reported that the new
shoot appeared at the base of the flowering stem at
anthesis and the 3 - 4 fleshy roots made their appear-
ance the following spring, near Hatley, Quebec.

Capsules: light brown, ellipsoid, typically 0.6 x
0.3 cm, ascending (see Figure 1d); yield approach-
ing 100%.

Seeds: greyish orange, released in late October.

BLOOMING PERIOD: 10 August (21 August - 12
September) 23 September [42 records], about the
same duration as S. cernua but a week earlier.

76° 30

75°

Spiranthes casei: @ = herbarium specimen, @ = Native Orchid Location Survey sight record.

Areas underlain by sandstone are shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 1IS7

FIGURE 40. Spiranthes casei, old-field habitat, Stony Swamp Conservation Area, Regional Municipality of Ottawa-
Carleton, Ontario, 24 August 1980.

158

CoLony SIZEs: | - 100, typically to 50, flowering
plants, 10 or fewer in two-thirds of the colonies [32
colonies]; as scattered individuals or occasionally
as pairs.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983).

DISTRIBUTION: The Ottawa District is at the northern
edge of the range of this orchid of the Mixed Forest
Region (Catling 1982b; New York Flora Association
1990). Within the District, S. casei is restricted in the
Lowlands to regions of sandstone bedrock of the
Nepean and March Formations (shaded areas on
map). The major concentration of colonies has been
recorded in the Stony Swamp Conservation Area and
adjacent Bridlewood. On the Canadian Shield, S.
casei is found in scattered locations on sand deposits
or over acidic Precambrian rocks (J. M. Reddoch and
A. H. Reddoch 1987a).

HABITAT: Spiranthes casei favours old-fields and
woodland clearings in areas of acidic rock where
weeds and grasses are neither high nor dense.
Poverty Grass (Danthonia spicata) is usually the
dominant vegetation. Here, plants grow in deep or
shallow sand or sandy loam that is moist or some-
what drier. Where there are expanses of exposed
bedrock, plants are scattered along soil-filled
crevices and on adjacent open ground. Some associ-
ated species are fruticose lichens, Bracken
(Pteridium aquilinum), Meadowsweet (Spiraea lati-
folia), Blueberry (Vaccinium angustifolium), Slender
Gerardia (Gerardia (Agalinis) tenuifolia), Gray
Goldenrod (Solidago nemoralis) and, increasingly in
Stony Swamp, Stiff Eyebright (Euphrasia stricta).

Near White Lake, the unusually high concentra-
tion of 39 flowering plants was observed in a 1.4 m?
area of a sandy rural lawn (Adolf Vogg, personal
communication 1994).

Spiranthes cernua is almost always a companion
plant. Where sites are relatively flat-lying, as most of
them are, the two Spiranthes are intermingled.
Where the sites are sloped, S. casei usually is situat-
ed on the higher, drier ground, while S. cernua
inhabits the moister end of the gradient. Spiranthes
lacera occasionally accompanies S. casei on drier

THE CANADIAN FIELD-NATURALIST

Vol. 111

sites. Colonies on dry sites are particularly affected
by drought. In some years, very few or no plants
flower.

LONG-LIVED COLONIES: Many colonies of S. casei
last for many years because the soil is too shallow or
too infertile to support dense vegetation and woody
plants. In a few locations, annual mowing is the
agent that controls competing growth.

In Stony Swamp, the colony recorded by a 1972
collection continues to thrive. Several dozen plants
are scattered in the sand-filled cracks (pH 4.5) of
flat-lying exposures of sandstone in a large clearing
surrounded by second-growth mixed forest. A simi-
lar number of S. cernua plants run along the lower
edge of the opening.

At Kingsmere in Gatineau Park, a colony of up to
55 plants of S. casei and a few S. cernua has sur-
vived since at least 1981 because the field they grow
in is mowed annually to keep down woody plants.
Here, the gneiss bedrock (Hogarth 1970) is partially
exposed but generally covered thinly to deeply with
sand. It has been the usual practice for the National
Capital Commission to mow Gatineau Park fields in
September to avoid disturbing ground-nesting birds.
But since mowing at this time sometimes cuts down
flowering S. casei and it is N.C.C. policy to protect
provincially-rare flora, the staff is trying to arrange
for mowing of that field to be done either later in the
fall or very early in the spring.

LocAL History: We have found no record of S.
casei in the Ottawa area before the late 1940s. In
1947, W. J. Cody, J. A. Calder and H. L. Mitchell
made a collection [DAO 89678] near Jarnac,
Quebec, just outside the 50-km circle to the north-
east (Reddoch and Reddoch 1974). In 1946, 1947
and 1948, L. Mcl. Terrill collected S. casei north of
Merrickville, about 5 km south of the District [CAN
338026, -27, -28]. S. casei was first collected in
Gatineau Park in 1968 and in Stony Swamp in 1972.

During the late 1980s and the 1990s, housing con-
struction in Bridlewood destroyed all of the colonies
there, almost half of the population recorded a
decade earlier in the combined Stony Swamp -
Bridlewood area.

L997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

159

Spiranthes cernua (Linnaeus) L. C. M. Richard

Nodding Ladies’-tresses

Spiranthe penchée

The last orchid of the season, Spiranthes cernua is one of the most abundant, although it did not become so until
comparatively recently. In the field, it can be distinguished from S. casei by its pure white flowers greater than 8
mm in length, often with a faint yellowish colour in the throat, from S. romanzoffiana because its flowers are
tubular and more or less horizontal rather than hooded and ascending, and from the other Spiranthes by its late
blooming period, September, and lack of strong yellow or green in the throat.

DESCRIPTION

Height: 6 (16 - 28) 43 cm [1327 plants].

Flowers: | (8 - 21) 40 [492 plants]; white, often
faint yellowish white or yellowish grey near the cen-
tre of the lip, often with a narrow band of faint grey-
ish green across the bases of the tepals; in 1, 3 or 4
ranks (see Spirality below); fragrance usually rich,
sweet, coumarin-like, rarely, lacking.

Leaves: 4 - 5, basal, starting to wither a month
before anthesis, leaving 0 - 2 by the time the last
flowers open; and 1, cauline, near the base, some-
times bract-like, starting to wither at anthesis or
shortly after; 2 - 4 on non-flowering plants.

Overwintering State: a lanceoloid shoot or a pair
of leaves 1 - 2 cm in length surrounding a central
bud, appearing above ground beside the current

46°

year’s stem at anthesis or shortly after; shoots appear-
ing earlier and developing further on decapitated and
non-flowering plants (see Seasonal Development
below); Mousley (1924a, b) in describing the under-
ground development of S. cernua near Hatley,
Quebec, reported that the new shoot appeared at the
base of the flowering stem at anthesis and that the
first pair of root buds was evident in the early spring
of the following year; by late May, one or two addi-
tional fleshy roots had been produced.

Capsules: light brown, ellipsoid; typically 0.7 x
0.3 cm, ascending; yield approaching 100%.

Seeds: light brown, released in mid to late
October.

BLOOMING PERIOD: 15 August (28 August - 21
September) 16 October [125 records].

45°
76° 30'

ke

Spiranthes cernua: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record. Areas of major sand deposits on the Lowlands are shaded.

160

CoLony SIZEs: | - 5000, typically to 500, flowering
plants [235 colonies]; as scattered individuals, occa-
sionally in twos and threes, and rarely up to groups
of five.

CURRENT STATUS: one of the most abundant orchids
in the District.

DISTRIBUTION: The Ottawa District is near the north-
ern limit of the range of this eastern species. The
range encompasses much of the Mixed, Deciduous
and Southeastern Coastal Plain Forest Regions and
the adjacent American Prairie. Within the District,
most colonies are located on sands deposited by the
Laurentian ice sheet, the Champlain Sea and subse-
quent rivers. These sands are mainly in the
Lowlands, especially in the southeast, but they also
occur in scattered locations on the Shield.

HaABITATs: The principal habitat in terms of number
and size of colonies is open, wet to moist expanses
of acidic sand with sparse plant cover. Human activi-
ties have created these conditions in broad, shallow
roadside ditches, abandoned sand pits and scrapes,
and fallow fields and pastures. Common companion
plants are Variegated Horsetail (Equisetum variega-
tum), Slender Gerardia (Gerardia (Agalinis) tenuifo-
lia), Meadowsweet (Spiraea latifolia) and other
roadside weeds. On the ridges adjacent to the Mer
Bleue Bog, additional companion plants include
Round-leaved Sundew (Drosera rotundifolia),
Labrador Tea (Ledum groenlandicum) and Sheep
Laurel (Kalmia angustifolia).

With increasing shade and competition from other
herbs, colonies dwindle in size and eventually may
disappear. However, individuals or small numbers
may persist among moderately dense, knee-high
grasses, in young, semi-open shrub thickets and
woodlands, and in residual openings and along trails
in more mature second growth forest.

A few sizeable colonies are also encountered in
naturally disturbed settings on the wet to moist
shores of some small Shield lakes and along the
Ottawa River west of the city. Plants have been
found on both sandy and clayey beaches, in exposed
soil or among rocks, and, rarely, in shallow water.

Some colonies thrive in thin, dryish to moist soil,
usually sand but sometimes clay, over flat-lying
bedrock. Bedrock under S. cernua includes sand-
stone and gneiss (pH 4.0), and limestone and marble
(pH 7.5).

Occasionally, the substrate has a moderate compo-
nent of humus, but there is no evidence in the
District for a peatland habitat.

LONG-LIVED COLONIES: Colonies can continue to
flourish for decades under conditions that maintain
suitable habitat, for example, mowing, re-exposure
of bare substrate and the presence of soil too shallow
or infertile for competing vegetation to thrive. A gra-
dient within a habitat that allows the colony to sur-

THE CANADIAN FIELD-NATURALIST

Voli

vive in one part after the rest of the habitat has been
altered can be important.

There has been a colony of several hundred S. cer-
nua in a Shallow, sandy ditch (pH 5.0) beside
Anderson Road in Gloucester since at least 1973.
The habitat is maintained by mowing. Installation of
a gas pipeline in the mid-1980s obliterated most of
the colony except for a few plants along the fence-
line. Within a few years, several dozen plants
appeared on the newly-exposed sand in the ditch and
numbers quickly increased to their previous level.
(See also Seasonal Development below.)

Up to 500 plants of S$. cernua have flowered on
the Borthwick Ridge adjacent to the Mer Bleue Bog
since at least 1966. For most of that time, they grew
in wet to moist sand (pH 4.5 - 5.0) in a seasonally-
flooded scrape that increasingly was being shaded by
willows. Companion plants included the species list-
ed above for the sand habitat, as well as Pogonia
ophioglossoides, Bog Clubmoss (Lycopodium inun-
datum) and Marsh Fern (Thelypteris palustris). The
S. cernua plants persisted into fairly deep shade
before disappearing along with all other herbaceous
plants. However, several dozen plants still can be
found adjacent to the scrape in moist sand at slightly
higher elevations in the dappled shade of second
growth Trembling Aspen, Grey Birch, Tamarack,
Red Maple and Speckled Alder. The sparse ground
cover is dominated by Polytrichum mosses and
includes Ground-cedar (Lycopodium complanatum)
and Labrador Tea. Throughout the known history of
this colony, four-ranked plants have been dominant,
although ranking was less well-defined in the treed
habitat (see Seasonal Development below for a
description of the effects of shade).

Patches of shallow sand on exposed flat-lying
sandstone (pH 4.0) in the Stony Swamp Conser-
vation Area have supported a mixed colony of sever-
al hundred S. cernua and several dozen S. casei since
at least 1977. Conditions do not support a dense
growth of vegetation and Poverty Grass (Danthonia
spicata) is the most common herb.

Whether a colony is allowed to bloom depends on
the timing of the operation that maintains the habitat.
A striking example was the appearance in 1980 of
several hundred flowering plants in a rocky pasture
on the Hendrick farm at Old Chelsea, Quebec. The
cattle and horses are moved from field to field
throughout the season. Apparently only that year was
a field free from livestock from July to September.
We do not know for how many years the plants were
present in the field before or after 1980.

See also this heading under S. casei.

EARLY History: The earliest records of S$. cernua in
the Ottawa District appear to be collections made by
F. P. Clappison in a “beaver meadow” at Kemptville
in 1898 [TRT 15859, 15865] and by John Macoun
“in the old gravel pit” at Britannia in 1903 [MTMG

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 161

FiGuRE 41. Spiranthes cernua, right inflorescence: moist scrape, Dolman Ridge, Mer Bleue Conservation Area, 2
September 1973; Centre inflorescence: moist scrape, Borthwick Ridge, Mer Bleue Conservation Area, 2 September
1980; whole plant: moist old-field habitat, Stony Swamp Conservation Area, 13 September 1980. The two inflores-
cences are drawn to the same scale.

162

8601]. References to S. cernua in the Flora
Ottawaensis (Fletcher 1880, 1893) were likely based
on James Fletcher’s collection of $. romanzoffiana at
“Take Flora, Hull, P.Q.” in 1877 that he mislabelled
S. cernua [DAO 89679] (see S. romanzoffiana
account).

That this species was not collected earlier suggests
that it was very rare here before 1900. Indeed, only
one additional site was reported up to 1917 (in 1909
[CAN 117012], Harrington 1917). It is likely that
human alteration of the landscape in the 19th century
enabled S. cernua to expand its range northward into
this area and/or increase its numbers dramatically.

In the Arnprior area at the western edge of the
District, S. cernua was unknown to Charles
Macnamara (circa 1940°) in the first part of the 20th
century. Photographs that Macnamara labelled S.
cernua are Clearly S. romanzoffiana. Spiranthes cer-
nua is currently considered uncommon in the
Arnprior area (Michael Runtz 1984, personal com-
munication 1995).

SEASONAL DEVELOPMENT: In late 1994, we exam-
ined overwintering shoots in the Anderson Road
colony described above. In 1995, we marked 15 ran-
domly selected plants in this colony and observed
them throughout the growing season, usually once a
week during the more rapid stages of growth. These
plants grew in the sun on the east-facing slope of the
shallow ditch below a wire fence where they would
be safe from mowing. We made additional observa-
tions on the other plants in the colony at the same
times.

The above-ground seasonal development of the
mature plant of S. cernua can be considered in terms
of several distinct, consecutive processes with some
temporal overlap, i.e., growth and decay of the basal
leaves, growth of the flowering stem and cauline
leaf, development of the inflorescence, appearance
of new overwintering buds, and seed production and
release.

Development of Basal Leaves

An unusual mid March thaw melted the snow suf-
ficiently on the lip of the ditch to reveal the overwin-
tering buds. They were about | cm high, some still a
lanceoloid bud, others with two leaves opened to
reveal a minute growing tip. By mid May, the two
leaves were 2.5 - 3.5 cm long (almost their maximum
length) and the central shoot had elongated slightly.
By 4 June, two new leaves were growing rapidly
from the centre and had become 5 - 10 cm long. On
robust plants, there was an additional leaf about 1 cm
long in the centre. Leaf growth had effectively
stopped by 2 July when the two new leaves were 11 -
13 cm long and the inner leaf was 5 - 7 cm long. The
original two leaves were yellow or brown. By 22
July, the two large leaves were turning yellow or
brown on their outer halves or losing their tips, and
the original two leaves had disappeared.

THE CANADIAN FIELD-NATURALIST

Vol mit

Development of the Flowering Stem
and the Inflorescence

In the second week of July, the flowering stem
and clasping cauline leaf appeared in the centre of
the basal leaves and was 1.5 - 5 cm tall by 22 July.
The flowering stem achieved its most rapid growth
over the next two weeks, reaching 4 - 15 cm by 6
August. The basal leaves continued to decay. The
cauline leaf near the base of the stem had grown to 2
- 6 cm, nearly its maximum length. The stem
reached 7 - 17 cm on 13 August and its maximum of
13 - 19 cm on 20 August. The cauline leaf had
achieved its greatest length of 3 - 7 cm by 28 August
and on some plants was turning yellow.

On 20 August, the floral buds of most plants were
quite distinct and arranged in three nearly vertical
ranks independent of the rank of the ultimate mature
inflorescence. About three-quarters of the flowers
were open by 28 August and almost all by 3
September. By 13 September, half of the plants had
brown flowers. The individual flowers lasted about
two weeks or a little longer. Most inflorescences
were four-ranked and the rest were three-ranked.

For the 156 flowering plants in the colony, the
average height (for a 25 plant sample) was 20.4 (4.8)
cm and the average number of flowers was 15 (5.7).

Overwintering Buds, Capsules and Seed Release

New overwintering buds were appearing beside
the flowering plants by 13 September. Almost all
flowering plants produced capsules, which expanded
during the next month and then released seed in
October.

Non-flowering Plants

Of the 15 plants originally selected, four did not
produce flowers. The initial development of the non-
flowering plants was similar to that of the flowering
plants except that the leaves were shorter, their maxi-
mum lengths being 6.5 - 7.5 cm. There was usually a
total of four leaves on the more robust or mature non-
flowering plants and two or three on smaller plants.
Most of the non-flowering plants had produced new
overwintering buds by 3 September, at least 10 days
before the flowering plants. Similar early bud appear-
ance was also noted on a flowering plant that had
only one flower and on another that had been acci-
dentally decapitated between 6 August and 13
August, probably as a result of mowing nearby.

The Effects of Mowing

The broad, flat part of the ditch was mowed in late
June and again in the first week of August. The 17
flowering plants in this part of the ditch were unusu-
ally short, 10.8 (2.3) cm, with few flowers, 7.3 (4.6).
Although the flowers opened at the same time as the
rest of the colony, only a few had brownish flowers
by 13 September compared with about 50% for the
marked plants discussed above. The stems of some
of these small plants were deformed, in some cases
to the extent that they were horizontal on the ground.

1997

The Effects of Shade

On a section of the east-facing slope that was in
the open but lightly shaded by nearby bushes for
most of the day, there were three flowering and sev-
eral non-flowering plants. The flowering plants
developed more slowly, flowered later, were taller,
more slender, and had longer, narrower leaves and
fewer, more widely-spaced flowers than those of the
rest of the colony. The flowers were arranged in a
single rank. The plants flowered between 13
September and | October, about two weeks later
than the rest. (Some of their lower flowers were
brown by the latter date.) They were 21, 26 and 27
cm tall and had 9, 10 and 12 flowers.

Similar effects of shade were expressed in the
recent plants on the Borthwick Ridge in the aspen-
birch-tamarack-maple-alder habitat (see Long-lived
Colonies above). In 1994, the average height of 33
plants there was 28.3 (4.3) cm and the average num-
ber of flowers was 14.4 (5.7). Half of the inflores-
cences were four-ranked and half were single-
ranked.

The Effects of a Cooler Season

The summer of 1995 was one of the hottest and
driest on record (with very little rain after a record-
breaking deluge on 2 June). As a result, seasonal
development of the leaves was earlier than usual. By
comparison, the spring of 1996 was cooler than nor-
mal and the summer until August was slightly cooler
than normal. August temperatures were closer to
normal. Overall precipitation was slightly below nor-
mal. Development of the leaves in 1996 was slower
than in 1995. For example, in late July the two (or
three) longer basal leaves were still green with very
little dying-off at the tips. Some of the original pairs
of basal leaves remained green, while others were
yellow or had turned brown. At anthesis, the original
two basal leaves had disappeared but the two or
three more recent leaves, as well as the cauline leaf,
were still green or at most tinged with yellow. By the
time the flowers were brown, these leaves remained
mostly green but the yellowish component was
stronger and the ends were brown or had died off.
The cauline leaf remained green. Timing of the
emergence and development of the flowering stalk
was similar to that in 1995.

In 1996, there were 622 flowering plants with an
average height (for a 50 plant sample) of 19.5
(5.7) cm.

The Amount of Repeat Flowering
from One Year to the Next

Of the 16 plants marked and followed in 1995,
five in the open sun flowered in both 1995 and 1996
as did one of the three in the shade. However, all of
the monitored flowering stalks were bitten off in mid
August before the flowers opened. Perhaps our han-
dling of the plants had left scents that attracted the
selective attentions of a herbivore.

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

163

Throughout the summer, new seedlings appeared
where there were none before. It is our impression
that individual plants have a life span of only a few
years and the colony is maintained by seeding.

The Effects of Not Mowing

In 1996, the broad flat section of the ditch was not
mowed in August at our request and with the gra-
cious cooperation of Regional Municipality of
Ottawa-Carleton staff (Craig Huff, Urban Forester;
Richard Scott, Supervisor; and Bob McNeil). The
plants in the ditch followed the normal patterns of
development of the rest of the colony and did not
show any distortion of form. The average height of
these plants was consistent with that of the rest of the
colony. Instead of the 17 flowering plants in 1995,
there were many hundreds. The difference in the
total count of the colony, from 156 flowering plants
in 1995 to 622 in 1996, was largely due to the
greater number in the ditch.

SPIRALITY: Spiranthes species are often classified
as single-ranked or multi-ranked. For example, S.
casei is single-ranked and S. cernua is usually
multi-ranked. However, these terms refer to differ-
ent concepts that are not always mutually exclusive,
sometimes causing some confusion in the descrip-
tion of S. cernua. One concept, single-ranking,
describes the order in which the flowers developed
on the rachis, while the other, multi-ranking,
describes apparent patterns of flowers that the
human eye perceives.

In Spiranthes inflorescences there is always a sin-
gle fundamental spiral marking the positions of suc-
cessively formed flowers. (This spiral is equally like-
ly to rise to the left or to the right with the flowers
facing to the right or to the left, respectively.) If the
spiral is not too compressed, it is easily seen and the
plant is classified as single-ranked. Otherwise, the
eye is more likely to discern a set of parallel ranks of
flowers crossing the fundamental spiral. These ranks
may be vertical or may spiral up the inflorescence at
a moderate angle to the axis. Plants with this appear-
ance are termed multi-ranked. However, the funda-
mental spiral is still present and may be apparent in
four-ranked plants.

Sheviak (1982) has argued that the use of rank as
a character has resulted in considerable confusion in
the determination of some S. cernua plants, partly
because it recognizes only the extreme limits of sin-
gle- and multi-ranking instead of the continuous
range of possible arrangements. In addition, he finds
that this character can be inconstant. He proposes
that the phyllotaxy should be used instead of the
rank. The phyllotaxy is the ratio of the number of
turns of the fundamental spiral to the number of
flowers encountered along that spiral, counting from
a given flower to the next one directly above it.

However, we believe that the phyllotaxy also pre-
sents problems in its application to real plants. It still

164

does not adequately describe the continuous char-
acter of possible spiral behaviour from one plant to
another because it uses the idealization of integral
numbers of ranks, at least as it is often used. Fur-
ther, it is applied to the structure that the human eye
perceives rather than to the actual structure of the
inflorescence.

For a more detailed analysis and understanding, it
is useful to consider the concept of the divergence,
which is simply the angle between two successive
flowers as seen along the axis of the inflorescence.
The divergence is closely related to the phyllotaxy
but the former permits an explicit account of the
continuous character of the spirality and does not
depend on perception. It can thus be used to describe
some of the more subtle effects that can arise.

In the Ottawa District, S. cernua is commonly
three- or four-ranked. In the four-ranked plants, the
ranks are usually quite vertical, reflecting divergence
angles of about 88° - 92° (middle inflorescence in
Figure). Some other four-ranked plants have diver-
gences up to 103°, in which case the ranks twist in
the same direction as the fundamental spiral. A few
plants have divergences around 86° and the ranks
twist in the opposite direction. The four-ranked
inflorescences have 13 - 19 flowers with an average
spacing along the rachis of 3.8 (0.3) mm. If the fun-
damental spiral is lax, the inflorescence may appear
to be simultaneously single-ranked and four-ranked.

Plants with divergences larger than 103° appear
three-ranked. A few S. cernua have a divergence of
120° and display three vertical ranks. Most, howev-
er, have angles in the range of about 103° - 115°, in
which case the ranks spiral in the opposite direction
to the fundamental spiral (plant on right in Figure).

THE CANADIAN FIELD-NATURALIST

Vol. 111

In the occasional plant with a divergence greater
than 120°, the two types of spirals turn in the same
direction. Three-ranked plants have about 15 - 40
flowers with a spacing of 3.3 (0.1) mm.

Inflorescences with a divergence of about 107°
can show four ranks simultaneously with the usual
three ranks. However, the four ranks involve flowers
in every other turn of the fundamental spiral instead
of in adjacent turns.

A few of the S. cernua colonies on the open sands
of the southeast contain some single-ranked plants.
They have divergences of about 60° - 80°, corre-
sponding to six or five flowers, respectively, per turn
of the fundamental spiral (left inflorescence in
Figure and photograph in Reddoch and Reddoch
1974). Such plants have 9 - 14 flowers with an aver-
age spacing of 4.4 (0.1) mm. These plants may be
part of single-ranked triploid populations centred in
the Eastern Townships of Quebec, New England and
adjacent New York (Catling 1980; Sheviak 1982,
Figure 18a-d).

Often, flowers near the top or bottom of an in-
florescence are not as regularly arranged as those
near the centre. Some inflorescences are irregular
throughout.

Sheviak (1982) has noted that single-ranked plants
tend to have fewer flowers than others. The flower
counts given above support this result and show a
highly significant correlation, r = 0.62 for 57 plants,
between divergence and flower number. The single-
ranked plants have the fewest flowers while the
three-ranked plants have the most.

A sampling of several local colonies has shown
that the buds develop in a three-ranked formation
regardless of the ranking of the open flowers.

1997

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

165

Spiranthes lacera (Rafinesque) Rafinesque var. lacera

Northern Slender Ladies’-tresses

SYNONYM: Spiranthes gracilis (Bigelow) Beck (in part)

Spiranthe découpée

This is an inconspicuous orchid, easily mistaken at first glance for a grass. On closer inspection, it is clearly an
orchid and may be distinguished from our other Spiranthes by the greenish lip and the small, ovate, basal

leaves.

DESCRIPTION

Height: 12 (21 - 35) 46 cm [206 plants].

Flowers: 9 (13 - 25) 35 [77 plants]; white with
centre of lip yellowish green, light green or greyish
green inside and outside or mainly along the veins;
in a single rank with almost no spiral or up to 3 loose
turns, occasionally to 5 turns, rising to the right or
left; fragrance usually marked, sweet, floral, occa-
sionally spicy.

Leaves: 3 - 5 in a basal rosette with 0 - 2 remain-
ing at anthesis; and 1, cauline, near base; non-flow-
ering plants with | - 5 leaves.

Overwintering State: a rosette of leaves surround-
ing a central bud (rarely two or three rosettes), appear-
ing above ground beside the current year’s stem in
late August to mid September shortly after anthesis

and developing to almost ultimate size and number of
leaves before winter; new rosettes appearing earlier
and developing further on decapitated plants.

Capsules: brown, ellipsoid to spheroid, typically
0.3 x 0.2 cm, slightly ascending (see Figure 1d);
yield generally over 50%, averaging 85% [26
plants].

Seeds: light brown, released in early to mid
August, about two weeks after the flowers have
turned brown.

BLOOMING PERIOD: | July (13 July - 12 August) 7
September [82 records].

CoLony SIZEs: | - 200 flowering plants, fewer than
30 in 87% of colonies; one exceptional colony with
1721 flowering plants (see Long-lived Colonies

46°

45°
76° 30

75°

Spiranthes lacera var. lacera: @ = herbarium specimen, @ = Native Orchid Location Survey

sight record. The Canadian Shield is shaded.

166 THE CANADIAN FIELD-NATURALIST

sy)
qi
y)
7
Ww
—*
N
Bi
.
rN
VSS
SS
[=
as
EN
bas
ey
ase
AS
EH:
ia
|
t

FIGURE 42. Spiranthes lacera var. lacera, open mixed woods, Gatineau Park, Quebec, 9 August 1980.

Vol. 111

1997

below) [137 colonies]; flowering plants occurring
mostly as scattered individuals, occasionally 2 or 3
together.

DISTRIBUTION: The Ottawa District is near the north-
ern edge of the range of S. lacera var. lacera. This
variety extends over much of the Mixed Forest
Region and the northern part of the Deciduous Forest
Region (Luer 1975). Within the District, this species
is widespread, with concentrations along the
Eardley-Masham Road in Gatineau Park, in the
Stony Swamp Conservation Area and in The Burnt
Lands Alvar.

HaBITATs: This orchid frequents areas of dry to
moist, shallow to deep sand over almost any rock
type, acidic or basic. Colonies are found in open,
sandy or rocky, deciduous or coniferous woods, in
pine plantations, in sandy or rocky old-fields and on
open Precambrian knolls. Rarely, a plant or two
becomes established in a coniferous swamp or on a
semi-shaded river shore. Plants generally occur close
to trees and shrubs, even in old-fields. In most situa-
tions, there is little or no competing vegetation.

LonG-LIVED COLONIES: Where habitats are relative-
ly stable, colonies of S. lacera can endure for several
decades at least.

In 1968, 140 flowering plants were scattered
around outcrops of calc gneiss bedrock (Hogarth
1970) among young White Birch, Sugar Maple, Red
Oak and Eastern White Pine in Gatineau Park. The
colony continues to carry on, although in reduced
numbers, where the canopy is still fairly open.

In the Larose Forest, there were 1721 flowering
plants of S. lacera in 1969, growing in deep sand
under the high canopy of a well-trimmed, 40-year-
old Scots Pine plantation. The ground cover was
mainly a thick layer of pine needles with a few

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

167

patches of Polytrichum mosses under Bracken
(Pteridium aquilinum). Now, a few hundred flower-
ing plants remain in the areas that have not become
overgrown nor been cleared for a picnic site.
Cypripedium acaule is a constant companion here.

EARLY History: Spiranthes lacera was first record-
ed in the Ottawa District in 1887 (Fletcher 1888) and
was described as “very rare” in 1893 (Fletcher
1893). Until the second decade of this century,
Blueberry Point near Aylmer, Quebec, was one of
the few known localities for this species. There, it
was collected from sandstone openings among
Eastern White Pines [CAN 117018, -19, -20] and on
a hummock at the edge of a swamp (Fletcher 1893).
Cypripedium acaule was also collected at Blueberry
Point. The construction of the Wychwood subdivi-
sion destroyed much of the natural habitat and put an
end to Ottawa Field-Naturalists’ Club excursions
there (Groh 1910; Reddoch 1979b).

SEASONAL DEVELOPMENT - SUMMARY: Before and
during anthesis, the basal leaves of the current year’s
plant die. Shortly after the flowers die, one, two or
three (rarely zero) new shoots appear above ground
beside the plant. Usually only one shoot survives; it
develops into a rosette of almost full-size leaves sur-
rounding a central bud before winter. Non-flowering
plants produce new shoots according to a similar
schedule.

In the spring, the last leaf unfurls and the leaves
complete their growth. Some of the outer leaves
begin to turn yellow in June.

The plant may produce a flowering stalk or remain
as a non-flowering plant. If it is going to flower, the
flowering stalk rises from the centre of the rosette in
June and expands throughout June and July until
flowering. In the two weeks after the flowers die, the
capsules ripen and release their seed.

168

Spiranthes lucida (H. H. Eaton) Ames

Shining Ladies’-tresses

THE CANADIAN FIELD-NATURALIST

Vol. 111

Spiranthe lustrée

This is the smallest, the earliest blooming and the least abundant of our Spiranthes. It can be recognized by the
bright yellow of the lip and the late June to early July blooming period.

DESCRIPTION:

Height: 11 (14 - 20) 28 cm [66 plants].

Flowers: 5 (8 - 16) 21 [56 plants]; white with lip
yellow to vivid yellow, above and below, often with
greyish green to dark green as a wash in centre or as
three broad parallel stripes or, when faint, following
the veins; nodding or horizontal, occasionally
ascending; in 3 ranks or occasionally 4; fragrance
rich and sweet or spicy or undetectable.

Leaves: 3 - 6, typically 4 - 5, basal, shiny, mostly
present at anthesis; | - 5 on non-flowering plants.

Overwintering State: a rosette of 3 greyish green
leaves, appearing above ground beside the current
year’s plant in early summer before anthesis (as
shown in Plate 80 of Correll (1950)); in October,
leaves | - 3 cm long, surrounding a narrowly conical
bud 0.5 cm tall, usually accompanied by a last fading
leaf of the current year’s plant.

Capsules: light brown to brown, ellipsoid, typi-
cally 0.8 x 0.35 cm, ascending.
Seeds: greyish brown.

BLOOMING PERIOD: 19 June (25 June - 7 July) 11
July [19 records].

COLONY SIZES: 3 - 90 flowering plants [13
colonies], as scattered individuals or in clumps of | -
3 flowering plants, accompanied at anthesis by new
shoots (see Overwintering State above) and occa-
sionally non-flowering plants.

CURRENT STATUS: rare in the Province of Quebec
(Bouchard et al. 1983), rare in the Ottawa District.

DISTRIBUTION: The Ottawa District is at the northern
limit of the range of this lower Great Lakes -
Atlantic Coast species. It is mainly a plant of the
Mixed Forest Region and the adjacent Deciduous

76° 30'

75°

Spiranthes lucida: @ = herbarium specimen, @ = Native Orchid Location Survey sight
record, & = literature reference. Major areas underlain by calcareous rock (marble and lime-

stone) are shaded.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 169

FiGurE 43. Spiranthes lucida, Ottawa River shore, Innis Point, City of Kanata, Regional Municipality of Ottawa-Carleton,
Ontario, 27 June 1978.

170

Forest Region. Within the District, S. /ucida is essen-
tially restricted to the shores of the Ottawa River and
some of its tributaries in areas of limestone bedrock
on the Lowlands and marble on the Shield. Whiting
and Catling (1986) have mapped two additional col-
lections in the District from the Madawaska and
Mississippi Rivers.

HaBitaT: Colonies are confined to rocky or sandy
shores that are scrubbed annually by ice and flood
waters to a distance inland of up to 15 m. Plants usu-
ally are within a few metres of the water’s edge,
among low sedges, rushes and grasses, in the wet
sand that has lodged around stones and boulders or
in crevices in the bedrock pavement. Some years,
high water covers the plants from time to time.
Additional accompanying plants include Meadow
Spikemoss (Selaginella apoda), Marsh Fern
(Thelypteris palustris), Wild Iris Uris versicolor),
Silverweed (Potentilla anserina), Shrubby
Cinquefoil (P. fruticosa), Kalm’s St. John’s-wort
(Hypericum kalmianum), Purple Loosestrife
(Lythrum salicaria), Indian Hemp (Apocynum
cannabinum) and, occasionally, Platanthera flava.

In 1912, Charles Macnamara discovered and pho-
tographed two flowering plants of S. Jucida in a field
close to the Ottawa River near Arnprior, Ontario.
Here, the plants grew in tall grasses and sedges on a
wet bank beside a small stream. He followed the
colony for at least three years (Macnamara circa
1940°; Reddoch 1981c).

Along the Mississippi River just south of the
Study Area, S. lucida occupies the eroded rocky
or sandy bank adjacent to the aquatic zone.
Companion species there include Meadow
Spikemoss, Arrow-grass (Triglochin maritimum),
sedges (Carex spp.), Golden Sedge (Carex aurea),
Twig-rush (Cladium mariscoides), Wild Iris, Tall
Meadow-rue (Thalictrum polygamum), Grass-of-
Parnassus (Parnassia glauca), Purple Loosestrife,
Marsh Bellflower (Campanula aparinoides),

THE CANADIAN FIELD-NATURALIST

Vol. 111

Cardinal-flower (Lobelia cardinalis) and Kalm’s
Lobelia (L. kalmii) (Patricia Rothschild, personal
communication 1984; Lemky and Keddy 1988).
Purple Loosestrife is currently being kept under con-
trol by hand weeding (Patricia Rothschild, personal
communication 1996).

LocaL History: Spiranthes lucida was first noted
in the Ottawa District in 1905. A single plant was
discovered and collected on the Ottawa River shore
below the Victoria Hotel at Aylmer, Quebec, by W.
Hague Harrington ({CAN 117016], Harrington
1917). In 1908, a second plant was found “by the
brookside near Old Chelsea” (Harrington 1917).
These two plants are the only records from the
Quebec part of the District.

Colonies at Shirley’s Bay - Innis Point in the
western Greenbelt have been known for almost 50
years. They are scattered along the Ottawa River
from the sandy beaches of Shirley’s Bay to the lime-
stone pavements of Innis Point and beyond, a dis-
tance of several kilometres. Along this shore, the
colonies vary in size and location from year to year.
Access to this property is restricted by the
Department of National Defence, thus affording
some protection to S. /ucida and the other rare plants
of the Innis Point shoreline (White 1979).

SEASONAL DEVELOPMENT - SUMMARY

The shoot for the next year’s plant appears above
ground in late June beside the current year’s plant. It
comprises three leaves and a growing tip which con-
tinue to expand through the growing season. At the
end of the season, the leaves are as long as 3 cm and
the growing tip has become a partially unfurled leaf
up to 1.6 cm long. At this time, the leaves of the cur-
rent year’s plant, some of which remained green into
September, are now brown and some have disap-
peared.

The next spring, the new leaves continue to expand
to full size. The flowering stalk extends from the cen-
tre of the basal leaves during the month of June.

1997

Spiranthes romanzoffiana Chamisso

Hooded Ladies’-tresses

REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT

Led

Spiranthe de Romanzoff

This Spiranthes is encountered infrequently in the District. Its colonies are usually small and often short-lived.
Spiranthes romanzoffiana is sometimes mistaken for S. cernua but can be recognized by its ascending, hooded,
creamy white flowers and by its lip, which has a constriction where it emerges from the hood.

DESCRIPTION

Height: 9 (18 - 33) 53 cm [64 plants].

Flowers: 4 (9 - 29) 45 [58 plants]; white to yel-
lowish white with light green at the base of the sep-
als, lip yellowish white, occasionally with greenish
white at its centre; usually arranged in 3 slightly spi-
ral ranks to right or left, occasionally in 4 ranks or
irregular; fragrance strong and rich to light vanilla.

Leaves: 3 - 4, basal, withering to leave 2 - 3 at
anthesis; and 1, cauline near base, sometimes bract-
like; 4 on non-flowering plants.

Overwintering State: a greyish green, lanceoloid
shoot, 1 cm above ground beside the current year’s
stem, appearing by early September; from near
Hatley, Quebec, Mousley (1924a, b) reported that
the new shoot was present at the base of the flower-
ing stem by early August and the first pair of root

buds appeared in early October; sometimes there
were two shoots and, on one occasion, three, each
with two root buds; 1 - 2 and sometimes 4 or more
additional fleshy roots were produced the following
spring.

Capsules: light brown to brown, ellipsoid, typi-
cally 0.8 x 0.4 cm, ascending.

Seeds: light brown, released by mid September.

BLOOMING PERIOD: 4 July (20 July - 19 August) 15
September [33 records].

COLony SIZEs: | - 15 flowering plants, 3 or fewer in
60% of colonies [24 colonies], as scattered individu-
als or rarely as a loose group.

DISTRIBUTION: The Ottawa District is in the south-
eastern part of the North American range of this
transcontinental species. It occurs in the Montane,

76° 30!

75°

Spiranthes romanzoffiana: #1 = herbarium specimen, ®@ = Native Orchid Location Survey
sight record, A = literature reference. The Canadian Shield is shaded.

BIZ

Boreal and Mixed Forest Regions. Spiranthes
romanzoffiana is scattered thinly across the District.

HasiraTs: This species has been found in a variety
of habitats, some wet and some moist.

Small colonies have been encountered on the wet
floors of almost every calcareous sedge fen and in
some semi-open swamps. Other plants occasionally
turn up in open or partially shaded, moist, sandy
seepage areas and the edges of lakes, streams, beaver
ponds, fens and swamps.

In addition to the above typical habitats, S. roman-
zoffiana plants appear from time to time in moist, par-
tially shaded, sandy forest edges. In 1973 and 1974, in
the Larose Forest, a clump of seven flowering plants
grew in well-drained sand several tree-rows in from
the edge of a 45-year-old Red Pine plantation.
Another flowering plant was in a similar situation
across the road. In another part of the forest two
decades later, a single plant was in bloom at the edge
of a Trembling Aspen - White Birch woodland beside
a forest track adjacent to another Red Pine plantation.
In the former location, S. romanzoffiana was accom-
panied by a large colony of Malaxis unifolia on the
pine-needle covered ground among a few stalks of
Bracken (Pteridium aquilinum). At the latter site,
Bracken, Wild Lily-of-the-valley (Maianthemum
canadense), Dewberry (Rubus hispidus), One-flow-
ered Wintergreen (Moneses uniflora) and Polytrichum
mosses were prominent accompanying plants.

At the Stony Swamp Conservation Area in the
western Greenbelt in the early 1980s, a few plants of
S. romanzoffiana were scattered among Bracken on a
low ridge at the edge of an Eastern White Pine - Red
Maple woodland. (One of these plants is the subject
of the Figure.) They were adjacent to an open, moist

THE CANADIAN FIELD-NATURALIST

Vol. 111

area of exposed sandstone bedrock that is home to a
long-established mixed colony of S. casei and S. cer-
nua. In a drier part of the same opening are scattered
plants of S. casei and S. lacera.

Spiranthes romanzoffiana does not form large
colonies as does S. cernua.

EARLY History: Spiranthes romanzoffiana was first
collected in the District in 1877 at “Lake Flora, Hull,
P.Q.” by James Fletcher. This collection of two
plants [DAO 89679] was identified by the collector
as S. cernua and recorded as such in both versions of
his Flora Ottawaensis (1880, 1893). It was annotated
by Paul Catling as S. romanzoffiana in 1973. Lake
Flora was a small pond on I’Ile de Hull (Nagy 1974)
that is now Parc Fontaine. Spiranthes romanzoffiana,
along with Calopogon tuberosus and Pogonia
ophioglossoides (Fletcher 1893), likely grew on a
sedge fen mat surrounding the pond. Other orchids in
this calcareous peatland were Corallorhiza trifida
(Macoun circa 1911°), Cypripedium acaule and C.
parviflorum (Fletcher 1893).

The first recognized discovery of S. romanzoffi-
ana came in 1890 (Fletcher 1891). William Scott
collected this species at East Templeton, Quebec
(Fletcher, Whyte, and Scott 1891; Fletcher, Scott,
and Cowley 1892; collections at CAN, DAO, TRT).
The 1892 authors reported that “W. Scott found
Spiranthes romanzoffiana in profusion in a hayfield
close to the East Templeton wharf, and between the
wharf and the lightkeeper’s house”. The 1891
authors also mentioned that Robert B. Whyte had
found the same species a short while later on Kettle
Island, nearby in the Ottawa River; however,
Fletcher (1893) included only the first location in his
Flora Ottawaensis.

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 173

FiGurE 44. Spiranthes romanzoffiana, edge of mixed forest, Stony Swamp Conservation Area, Regional Municipality of
Ottawa-Carleton, Ontario, 22 July 1979.

174

Possible Additional Species and Hybrids

Three orchid species have been discovered in
recent years not far from the Ottawa District. Thus
these species in particular should be watched for.

Corallorhiza odontorhiza: This species of the
Deciduous Forest Region has been discovered 150
km southwest of the District near Trenton, Ontario
(Brownell et al. 1994) and 80 km east of the District
near the mouth of the Ottawa River (lac des Deux-
Montagnes) in Quebec (Sabourin 1993).

Listera convallarioides: The closest known locations
for this eastern Boreal Forest, Mixed Forest and
Montane Forest species are both 90 km from the
Ottawa District: northwest in the Ottawa Valley near
Petawawa, Ontario (Whiting and Catling 1986) and
northeast in the Laurentian Mountains north of
Montreal, Quebec (Sabourin 1993). The Petawawa
colony consists of a hundred or so plants but only a
few plants flower each year (Grant Bickel, personal
communication 1996).

Acknowledgments

We are indeed grateful to the following people
whom we have had the pleasure of interacting with
over the years: members of the Councils and
Publications Committees of The Ottawa Field-
Naturalists’ Club for their interest and for financial
support in publishing this article, including a grant
in commemoration of the Club’s centennial in
1979; Charles Gruchy and John M. Gillett of the
Canadian Museum of Nature for research associate-
ships; curators, curatorial assistants and staffs,
especially of the National Herbarium (Canadian
Museum of Nature) and Agriculture Canada, for
many different kinds of assistance; Robert R.
Ireland for identifying the mosses cited; Donald D.
Hogarth for answering questions about geology;
John H. Day for the loan of soil maps; Donald G.
Cuddy for providing copies of Ontario Ministry of
Natural Resources reports and other information;
André Sabourin, Roger Perreault and Jacques
Cayouette for obtaining copies of obscure references;
J. Donald Lafontaine for early discussions on habi-
tats; Edward W. and Mary Greenwood for reviewing

THE CANADIAN FIELD-NATURALIST

Vol. 111

Spiranthes ochroleuca: This species of the Decid-
uous Forest has been found 90 km southwest of the
Ottawa District in Frontenac County, the only record
in Eastern Ontario (Whiting and Catling 1986). It
has not been recorded in Quebec (Sabourin 1993,
personal communication 1995).

Homoya (1993) included a disjunct dot in the
Ottawa District on his distribution map for this
species; however, he informed us (personal commu-
nication 1995) that this dot was the result of a pro-
duction error.

Hybrids: There are several species in the Ottawa
District that have produced hybrids elsewhere in their
ranges. These hybrids are Platanthera x vossii Case
(P. blephariglottis var. blephariglottis x P. clavellata
var. clavellata), P. x keenanii P. M. Brown (P. lacera
x P. grandiflora), P. x andrewsii (Niles) Luer (P. lac-
era x P. psycodes) and Spiranthes x simpsonii Catling
and Sheviak (S. lacera var. lacera x S. romanzoffiana).

much of the manuscript; Charles J. Sheviak, Marilyn
H.S. Light and John V. Freudenstein for helpful
information on various genera; Paul M. Catling and
Karen L. McIntosh for examining the species illus-
trations; two reviewers for valuable comments on
the entire manuscript; Florence Dunston for assis-
tance with field work; several property owners for
granting us permission to search for and study
orchids on their properties; and the many people
listed as contributors to the Native Orchid Location
Survey or cited for personal communications for
sharing their finds and knowledge. We particularly
recognize the contributions of Ed Greenwood, who
founded the Native Orchid Location Survey and
continually prodded us to complete the present
account, and Paul Catling for providing useful
information and stimulating discussions. We
enjoyed working with Susan Laurie-Bourque, who
prepared the final drawings of plants, flowers and
capsules, and with Marcel Jomphe, who generated
the computer versions of the introductory maps and
Table.

1997

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Received 3 October 1996
Accepted 10 December 1996

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 181

Appendix 1: Summary of Lists of Orchids of the Ottawa District and Adjacent Areas

1861 =herbarium specimens of John Kerr McMorine at QK and DAO (dated 1861 — 1867)

1867 =List of Plants collected by Mr. B. Billings in the vicinity of the City of Ottawa, during the Summer of 1866
(Billings 1867)

1880 = Flora Ottawaensis (Fletcher 1880)

1893 = Flora Ottawaensis (Fletcher 1893)

1911 = The Ottawa Flora (Macoun circa 1911)

1958 = Checklist of Plants of the Ottawa District (Gillett 1958)

1977 = Native Orchid Location Survey (Reddoch 1977b); Checklist of Vascular Plants of the Ottawa-Hull Region, Canada
(Gillett and White 1978)

1940 = Some Orchids of Eastern North America Photographed by Charles Macnamara (Macnamara circa 1940")

1984 = Orchids of the Arnprior Area (Runtz 1984)

OTTAWA DISTRICT ARNPRIOR

1861 1867 1880 1893 1911 1958 1977 1940 1984

Amerorchis rotundifolia - - x
Aplectrum hyemale - - -
Arethusa bulbosa - -
Calopogon tuberosus X 8
Calypso bulbosa =
Coeloglossum viride
Corallorhiza maculata
Corallorhiza striata
Corallorhiza trifida
Cypripedium acaule X
Cypripedium arietinum =
Cypripedium parviflorum x
Xx

1 PO;
!
1s pe) pe 1 pd. id ds ed
a a oo oo
PS OP OPK OPK OP OP OS OP OP OP ODO

Cypripedium reginae
Epipactis helleborine
Galearis spectabilis . X
Goodyera pubescens . - . X
Goodyera repens X . x X
Goodyera tesselata - : - :
Liparis loeselii - . X X X
Listera auriculata - - - - >
Listera australis - = : 2
Listera cordata - = ~ -
Malaxis monophylla - - x
Malaxis unifolia X . X
Platanthera blephariglottis - - X
x
x

ec a os

1
1 > >

>

Bd) pel 54
Dd ibd) Sei SeeBibd! Ube) isda bd Sd) ibdll Sei dic 154!) dl isd Sei bai sa)
x ®m

Co eo

Platanthera clavellata - -
Platanthera dilatata - x
Platanthera flava - - -
Platanthera grandiflora - - -
Platanthera hookeri - x x
Platanthera huronensis - - . -
Platanthera hyperborea X x X X
Platanthera lacera - - - - -
Platanthera leucophaea - - - - -
Platanthera macrophylla - . - - -
Platanthera obtusata - . - -
Platanthera orbiculata x x
Platanthera psycodes X x X
Xx xX

~ > oh OOO
~ > > pd md OO

*
1 PS PK Pt KOK OK OK OK KK OOK OK KOK OK OK OKO OKO OKO OKO KO OK OK

Pogonia ophioglossoides
Spiranthes casei - -
Spiranthes cernua - - - - X
Spiranthes lacera - - - x X
Spiranthes lucida . - - . -
Spiranthes romanzoffiana : - ‘4 X X

Total species: 13 8 23 31 35
@ discussed as Goodyera pubescens but photograph shows G. tesselata

#presumably a mistaken identification
*erroneously identified as S. cernua (see species account).

a eo oo =
a a eo a

!

1

oe)
oo
TS
Ne)
i)
Ns)
os)
jan)

182 THE CANADIAN FIELD-NATURALIST Vol. 111

Appendix 2: Significant Vegetation of Six Calcareous Sedge Fens

Two of the Lowland fens, the Richmond Fen and the Phragmites Fen, are components of the 20 km? palustrine wetlands
complex adjacent to the Jock River. A third fen, the Manion Corners Long Swamp Fen, is a patterned fen of about 20 ha
area in a 15 km? peatland (Reddoch 1978a). It is a patterned fen that has evolved in a saddle between the Mississippi River
and Rideau River watersheds. The pattern is an array of sedge fen lenses aligned across the direction of water flow and sep-
arated from one another by treed peat ridges. The fourth fen is the Mud Pond Fen, a partially floating fen behind an ice-
pushup ridge or ancient beach at the edge of a small, marl-bottomed pond surrounded by swamp.

Two fens on the Shield have formed in long, narrow bays of small lakes, the Quebec fen near Poltimore, and the White
Lake Fen adjacent to the lake in Renfrew County named for its marl-covered bottom.

This list is an update of previously published lists (Reddoch 1979a, 1984, 1989).

RICH = Richmond Fen, PHRA = Phragmites Fen, L.S. = Manion Corners Long Swamp Fen, M.P. = Mud Pond Fen, POLT
= Fen near Poltimore, W.L. = White Lake Fen.

RICH PHRA LS. M.P. POLT W.L.
Vascular Plants*
Scheuchzeria palustris X i : : - -
Triglochin maritimum X X X - - x
Triglochin palustre® - - - - - x
Bromus ciliatus X X X . . -
Calamagrostis stricta ssp. stricta® - - - - -
Muhlenbergia glomerata
Phragmites communis
Carex chordorrhiza - -
Carex diandra X -
Carex disperma - %
Carex exilis
Carex lasiocarpa
Carex limosa
Carex livida
Carex pauciflora - - - -
Carex paupercula = C. magellanica - - X
Carex prairea - - x
Carex tenuiflora - - X -
Carex trisperma - - X
Carex vaginata - - X
Cladium mariscoides x Xx - X - X
Dulichium arundinaceum x
Eleocharis elliptica® - -
Eriophorum viridi-carinatum - x
Rhynchospora alba X X
Rhynchospora capillacea - - - - - Xx
Scirpus acutus - - - - - X
Scirpus hudsonianus X X X
Smilacina trifolia - ¥ x
Amerorchis rotundifolia - - - - X -
Arethusa bulbosa - - x X
Calopogon tuberosus - X X x
Cypripedium arietinum - - x
Liparis loeselii X - - - - -
Platanthera clavellata - - - -
Platanthera dilatata - X
Platanthera huronensis = -
Platanthera leucophaea x
Pogonia ophioglossoides -
Spiranthes romanzoffiana X
Salix candida x
Salix pedicellaris x
Salix serissima : . X X - =
Betula pumila var. glandulifera
Sarracenia purpurea Xx
Potentilla fruticosa - - - X -
Potentilla palustris % - - - X -
Rosa palustris - - . X - -
Viola nephrophylla - X - - - -
Epilobium leptophyllum x X X x X -

1997 REDDOCH AND REDDOCH: THE ORCHIDS IN THE OTTAWA DISTRICT 183

RICH PHRA L.S. M.P. POLT W.L.
Vascular Plants*
Epilobium strictum - -
Andromeda glaucophylla - 2
Chamaedaphne calyculata - 4
Kalmia angustifolia -
Ledum groenlandicum -
Vaccinium macrocarpon . -
Vaccinium oxycoccos - x
Menyanthes trifoliata s§ X
Utricularia cornuta = 2
Utricularia intermedia X
Utricularia minor x
Galium labradoricum X
Lonicera oblongifolia x
Campanula aparinoides X
Lobelia kalmii Xx
Aster borealis X
Aster umbellatus -
Solidago uliginosa

Pad

MP KK OK OOOO OOO
I
1

>
~ <> OM
x «>»

1
x « «>

#designated rare, sparse and uncommon in Gillett and White (1978)
@ Adolf Vogg (personal communication 1994)

& Brunton (1990*)

*Paul Catling (personal communication 1991)

Mosses**

Aulacomnium palustre Xx X x X » -
Calliergon giganteum X X - - x -
Calliergon trifarium - - - - - x
Calliergonella cuspidata x x
Campylium chrysophyllum - x
Campylium stellatum a X
Cinclidium stygium - X
Drepanocladus vernicosus X 8 X
Fissidens adianthoides % - X - - -
Meesia triquetra - - X Xe
Paludella squarrosa . - - - - Xx
Plagiomnium ciliare - - - - . X
Rhizomnium pseudopunctatum - - Xx - - -
Scorpidium scorpioides - X X X
Sphagnum riparium - - - - > x
Sphagnum russowii - - - - : Xx
Sphagnum squarrosum - - . X - -
Sphagnum teres - - - -
Sphagnum warnstorfii - - Xx
Thuidium delicatulum Xx - « - - -
Thuidium recognitum - X X

Tomenthypnum falcifolium - - : - - Xx
Tomenthypnum nitens - - X - X X

**collections by Joyce Reddoch, except most at Manion Corners Long Swamp Fen by Anne Hanes and Cinclidium stygium
at Phragmites Fen by Gilda Trucco; all mosses identified by Robert R. Ireland.

184

About the Authors

Joyce and Allan Reddoch’s interests in studying
native orchids arose from a number of different
directions. They have always been interested in
nature and in hiking and they are always curious
about what they encounter along the way. Their
hobby of photography led them to taking pictures of
wildflowers and, after joining the Native Orchid
Location Survey, more especially of orchids. Their
scientific backgrounds prompted them to think in
terms of quantitative and statistical techniques when
studying various aspects of orchid ecology and tax-
onomy. Their work has resulted in numerous
descriptive articles in Trail & Landscape (The
Ottawa Field-Naturalists’ Club’s local natural histo-
ry journal) and other publications, as well as a taxo-
nomic paper on Platanthera orbiculata and P.
macrophylla in Lindleyana. They have given many
illustrated talks and lectures to audiences in the
Ottawa valley and beyond. With other members of
the Club’s Conservation Committee, they explored
and documented natural areas in the Ottawa District
and developed recommendations for the Ottawa-
Carleton Regional Official Plan and other land-use
planning exercises in Ontario and Quebec. These
activities led them to a more complete understanding
of the orchids and their habitats in the District as
well as the threats to them.

Joyce Marilyn Dunston was born in Mimico
(Toronto), Ontario on 27 December 1938, a first
generation Canadian of English parents Margery
Florence Green and George Vincent Dunston. Joyce
grew up in Oakville, Ontario, and earned degrees in
chemistry from the University of Toronto (B. Sc.
1961, M. A. 1962, Ph. D. 1966). She did postdoctor-
al work in organic chemistry and photochemistry at
Cornell University and, after moving to Ottawa in
the fall of 1966, at Carleton University and the
National Research Council. Briefly she was an
Assistant Professor in the Chemistry Department at
Carleton University.

Since joining The Ottawa Field-Naturalists’ Club
in 1966, Joyce has been a member of the Council

THE CANADIAN FIELD-NATURALIST

Vol. 111

and of most of its committees. From 1971 to 1979
she was an Associate Editor of Trail & Landscape
and from 1980 to 1989 she was the Editor. For her
contributions to the Club, she received the Member
of the Year Award for 1981 and the Service Award
of 1989.

In the 1970s, Joyce was the chairman of the
Gloucester Environmental Advisory Committee
established by the Gloucester Council to advise on
environmental matters, a member of the South Nation
Conservation Authority representing the Regional
Municipality of Ottawa-Carleton and a member of the
Club’s Conservation Committee, among other conser-
vation activities. The Ottawa Fish and Game
Conservation Association awarded Joyce the Murray
Boegel Waterfowl Conservation Trophy in 1978 for
her work toward protecting wetlands and other signifi-
cant natural areas in the Region.

Allan Harvey Reddoch was born in Montreal,
Quebec on 19 January 1931 to Scottish parents Mary
Love Harvey and Allan Reddoch. He attended
school in Montreal and in Ottawa where the family
arrived early in the Second World War. He then
studied chemistry at Queen’s University (B. Sc.
1953, M. Sc. 1955) and chemical physics at the
University of California at Berkeley (Ph. D. 1960).
Returning to Ottawa as a Postdoctoral Fellow at the
National Research Council, he subsequently became
a Research Officer there until his retirement in 1991.
His work involved the application of electron spin
resonance spectroscopy to a variety of studies in
physical chemistry and solid state physics.

Allan joined The Ottawa Field-Naturalist’s Club
in 1967 and in the early 1970s became a member of
the Club’s Council and its Corresponding Secretary.
During that time, he was also Chairman of its
Conservation Committee when it was involved in the
evaluation of candidate natural areas for the Ottawa-
Carleton Regional Official Plan and of natural areas
in the lands of the National Capital Commission in
Ontario and Quebec.

Index to Special Topics

COLOUR FORMS

Achlorophyllous Forms
MRIS PEST) ROP UE ir, Sette seh rates ve tows oes Vast ihamaas Sadie eakh aac cones aps das wea biowuannecsa bebsitienenawisnyoehs bane! FA
MMT EMRT UCR ILE, fone Sec cocsec-ynsccen ss Onan onan cist Aeon te teams BU AEA eas calld e aea Sadan LAS To Lino dswnrtcnn 132
Acyanic Forms
IN Et tn segine <meta 95 8 Roos wee seceine stein ok debs faa aneeninnong Saarnnained aalemaninaandesagnas evans ews
TN Te SPTTRIBE IEE DCP Ices I occa tra ccc ahs dae oles raha ya cien Ie ccbne canine saa napnnnnameb aes sepedee 44
RMR EPIL TOG Cree eee gnc cose nat eres nanan seen ena Bah icbepectines te shaeneepeess JuateGhe ouch iivaiadadins chased batdeonshingneaatanwiy 52
EE CLE EI cose cae rc sas cx ces neannngnen oie one isu tac URE eae a oe ASUS Sep be Secs dace Rat cso nS Ad dh bats 54
RIPE RISC IED 3 ooo cn ewe pasipeignenca goo «ete ea ae ee the hea aA a ncto bes See ap abo eA 60
EERE) ES 9 a a ne eee co nA Are ee ee 121
I RIS Re sic snaagsisouspncdentsb Sebel tte een mee eter Mace eaten ae etn octet ween nen sonmetanoseebuors 151
Colour Pattern Variants
aN NOTE PE CIE HI BCE. Sp 258 se ocd Re ct wc Raa eee Sua a tite, sh Sande cde eee AME na sows oda 28
SELL geal a RA id aia eee ine ak i soe I< Ge sf RE a DEBE re Ret Seasons orn, bee ee ae ee 76
5 Tl inl aaag a inlaid deaeeistsoeiar ta: Nara eA EEL nc ht Ls aA ere Ee a Uo ee AE PE 118
DERMATITIS
ENTRIES PETE ee Oe ee Pen ee ee ooo 8 ce aehsnnnnarchansdendsnisebiwuanaidanes divided vedenesnadinaddécuannsvonnnes 56
(2) RSS LEZ LEE ei lteter nA eR AOE PE MRSS 5 Se 60
MUIR? TEREST TINE oc sManc Recta a neet ene cet ace sea sean Re aves wae coun ows eae aR Che ea a eee mcbcneet ys 64
SNe re oar oat ae ceenetodcninsan on senenontenginapPacarmrccncnmesnrsaanspnenneStes 68

MORPHOLOGICAL VARIATION WITH HABITAT

NIUE 521 2 ree 098, AC) AE eee OEE... EOE Sal, IN es SI, WA, Sa 90
MTT OE OC Petra. octet sina Rees oe oaaacatncnl ote eccaucendetstpercnaes wlecdscometontetespdssbiccasoueae 12

POPULATION STUDIES (LONG-LIVED COLONIES)

ea RAL IU TRERE ATC = 1D CHL CUTE eo ete ere Fe OI sn antsuinvn cute Caer aSana go deeb sa dae ede dab d evade nb eut Meee ne 44
SNES ON TCU TE ULECI Sa te eT, Pe EE tA cod, AT iSisea sane dante ee a as es 48
IE ACER Tete Pere ee Ne MEY ics Seoncn Sade cr ces stot cau sc ontabe ct ste dashenasnnadentgecdaantszenecdeetaeu secs 78
NEES 1 NTI ree ees ene ee aoe once econo conc Jose eames tawadaseesorenreneivarnaenntonineanunincn ties 86
RE FOIVIPIEE OPEN EE es I ane Re Oo nls os sat, Os hae wotiininascinmwannansahinwasuie est wdsne Kae eae 124
EDIE EI OTE CEE ET ie te Se Pe RE SO EE accu de nocd Vanes anninsigSevigdeisaesnopbie wa phn seehealaaie 147
IPERS RELL oe ce eres eee eae 2 ae Aine Se cia acter napigcbce tuitl Sun sabeanavmeh Wee neta otek 160

SEASONAL DEVELOPMENT AND VEGETATIVE REPRODUCTION

SUNT TREAEOURELESEZIETINS Foner, , Oa Sept ens Jom ee Eek Oa oc A Tah a cera sales acanpaaerabo bs ben aR iten lade sas abasess ckanans 56
8 DE ELPA BST a7 ei ae te RMS IE peel cscs bb Ie Re MOEN Oe ME Cefn NIE en 78
MNES MEER MO TCREER I basco eh bn Satna d ganna onus ons ais dae ems ed ines Sh aes ask he Neel eet ws AE etd ok ain So Soee 87
DALE LL DOE EL TL 7 EE a OEM pe LT ee eee Lc we Ree ae eit rine et ee ee eee 162
ND or 8 si we cdma edbarpn sigan suc ARAER RO RO ES in a OO a AS 167
MINOR AIL A LCBO he Pies Soe sc ci rote: «A es Sa Aas ois nnn 1c NS as elt da Me one 170
Pete BEN EN SPIRANTHES INFLORESCENCES 2002.20. 0....00.-.050.c0secsecceoseeseseesrsetnteetacorsuacsacsstecssscntentecteneateeevese 163

185

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186

THE ORCHIDS OF THE OTTAWA DISTRICT

Species Accounts

DEAT IR RENE pe eo en So Seyccg cn ecnenteceoonemre eote ea dene shen evabed ony onnovswanobieniwenerevesnes 25
CAR AARNE ee Eo cp ncn cnnd bd Ao cana ER Gans -odioa nates lana cnbtatnadienneaeedniassiniesses 29
se a ae og) a ne eee Pen ewe ac cn ee 31
NOUR A MORI Le tne Be tah oh gs ke hace das aap aba Rem oreen Donte oad oLadRe ca Ren datnanaiitnwcus 34
I NaN NIC IRE kk arey oan aes iasSeeo nt cna oe mreteinecinc east bertese nce OAaananaadlve dees enaeainnnayincnse ag
RIUM EIER TAT ME MELE SEIN 355550 sncos Secs ocsirnnncd creado siodacs shh naetaunbe Sobindav ondetNalsnnbeedsuethtvainiehecseneses 40
nen erco Mmachiata Val. MaCculata and Val. OCCIGENIATIS. .........0.ccsnssnnececnvensesonescnneneccosesnescneceereesess 43
SITE A NEE ge J oe nm ee RP PME Wr cae PR Ce Ce Se Oe eee 46
TE EEN TUES ke Sg RE Rape etn ct Ce ener ich a eee aeeie te Seis onc? Sneret ee oe ee 50
SI Rs eS gn eg Da gS ce na EON ear a ttinn soa: Coen dave pan aceeelrt reese wants 53
LEE RIEE 5G he sti ce cou «nena na oa seie ds wae eels traatee Sues c oa deen reece de aleedoncacse ade teas teareeniicee af
PeetOeAiUIN PATVIFIOTUM Var. PUDESCENS ANG VAL. MAKASIN ......1.0.1.cccccsonensseoeeseesessneneoeceecensnsssccoonccees 61
NS SORES 8 P08 pe oe SY sd Sie ale ete i te ad eaten Ram se Moa uneeTntesadwaninennnonnnie 65
OTS AER SE Se ae oe Seller en de 28 OP wae ee SRN arte PLS OE EP ON ae ea 69
IRIE URLS Nc cere Fee Ne ha oe estes eae ones dabee Sant dawnueaaesionaabmadetnanereedanss 73
INO MRI prt ee ei allan ndlsd we vn ce dba puin cae ddavignubs yuaaambbuseedbwasn vans i
Sa le aE AR PITS oe aac se se evs aay vaca ooeuasnnd Onc cetundndynabadacadaveeseusdabaneatinibievsunsdeanae 81
ITN UP er cr Saige op nlecd dvniranb bl eoatbivntepE ban enncainnale oncom 84
PN ce aaa Ie ea ea ac Ses dA Se tildes solr aitsdacentda dai Dadueesanicd uanene 88
NU 2 reo Fo tahoe wg A eae was deasnine sa eMMa we xnndabeeddivietngevnnnaiianane 92
a UNUUNINNTII ceee ons ee sense ac eae chase wads acinepesbnddapualasdedgn sancevanediesnan 95
SES CR FED co oa he uF vail ads eaeicacn six egdnncdenirvedlavgddbedudetvuiisasedvendntnindsadudcestanseenes 98
EURMRNE MIE AR DVICHY DOUG. ooo sce eeciaten-senedabenatisusodessdsededssnsnernvéansdinsvedemondactevensacsndosacsesneeeeuas 101
NSN ae dst ne a ba eb ce Gani ds danicanic pine sceonnnPavondi namnaddadbornandoes 104
REE Fe PCDNGTISIONNS Val. DICPHOTIGIONIS <....<cc0cec-sosesncacssnecnnseaneconsennsenen aeons cnaecsnnacessesacacdvatecnenees 107
RE IEEE VAN CVV. oor ca cexsencc ci ncdcvvintstovacavunineaneb insecdarsenisanenccarvetusersvasndavanindussdeceonsusa 110
RINE SPECT GM OTN OIED 5a con ae oe coe sc enecrsg sts badcng’ssvia waned capanabstinvaenesastndviaceoiaiacdoavnecctonsie 113
EN RT IE TANIA EE oe ra oy ca bps teenie nx caovk dala desde i Ganan viinsnddciecndadenanenuinsnbdsviexdvaunaseaes 116
EASELS) RICE ce ee ce aa ead S sick cs dnnd aah cana od ween nwa aecghedbanudnastnaanbecsed 119
AO et aod Ss 9 och waar andes esinchn dees damnnnsudesdanxensioiadoonssavaccastease 123
I Ee a gree eS Sa els hoes wren Miavcewlaaann dss dalsetvaniiandacatahden enonddvdiews 127
ESE ROCIO VAL. PCF DOVE soci icscpecnn sie seine a asacgaeweusietunn cursdevevdscnssdsveneecunioysiensinenvncssinir’ 130
EET? Es 21 St ene ene SO cn 133
I Ce re Pt 9 i Sheol Lelia ah aN aon vilane ho ob aespaaatpinleb dna ea arin be 136
PNR AE TY Oe te 8 JLo nr catia ahivualasvnsnracmadieususlssiacdaudaisy evashndaiivaniosvodenveaieideds 139
NE RAMI PN 8 Ly kde shirt cca cused pneute adds wncanelps ok daseaaes rssh alate sgenueaiotat caabecxtaes 142
EGE iN ose psn epee are avsnteas du oss unsaid athe onan Was eierdxnahz 145
i ID EOS EN oda ls ca ctw vlna oh accailed a vpn ouy voce bnlontnoniaornes Dldaass sn ialan piurias 149
NORTON BRE TDG AGE Ce ah a i. fas Pisce Sid sore oe aia cre aavasee eve eawnanadiow ince aan Nvuaiokaied 152
UN Me UE PR EON 8 soca awe scs cod Lptunuies il cbd adbeeei eos tas eu oe phleoendie Uc SRabw ka aanics 156
MRA OS ee asa b dasinsudn sakes Coo a eae os auideehapcensennbuientacan 159
NE NIN BIA oy ot cc she vn Ceccerses es pcshcck oehgtadite deeneece eau sasbivobekindianwivnnwianmes 165
LE aoa Se eS Sa a0 Se Sen ae one a ee ene, Ree ne \ cc er 168
ONE IO a Soc ace deca ec ec ea ok teas wach cezsnildsvaeirsiouen 171

ere er OMA SPCCICS ANC EAVES. .:..05-.0i.200ncenacse icesanconenxactavsceseccvosecenenaessivnveindbessssuecvesnveninsesues 174

THE CANADIAN FIELD-NATURALIST Volume 111, Number 1

The Orchids in the Ottawa District: Floristics, Phytogeography,
Population Studies and Historical Review

JOYCE M. REDDOCH and ALLAN H. REDDOCH

Introduction and Discussion

The Ottawa District

History of Orchid Collecting and Recording

The Orchids of the Ottawa District

Climate, Geology and Vegetation

Principal Orchid Habitats

Local Distribution Patterns

Continental Distribution Patterns

Rare Species

The Need for Protection and Management

Colour Forms and Their Nomenclature

Blooming Periods

Capsules and Seeds

Overwintering States

Longevity of Colonies

Variability of Characters
Methodology
The Species Accounts (see inside back cover for species list)
Acknowledgments
Documents Cited
Literature Cited
Appendices:

1. Summary of Lists of Orchids of the Ottawa District and Adjacent Areas

2. Significant Vegetation of Six Calcareous Sedge Fens
About the Authors

Index to Special Topics
Advice to Contributors

Mailing date of the previous issue 110(4): 2 April 1997

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lume 111, Number 2 April-June 1997

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B3H 4J1. See status report by Reeves and Whitehead, pages 293-307.

The Canadian Field-Natusalist

Volume 111, Number 2 rT , , ¥ pril—June 1997

The Decline and Current Status of the Dune Raee-etDwarf Cherry,
Prunus pumila, var. pumila, on the Canadian Shores of the
Lower Great Lakes

PAuL M. CATLING,! and BRENDON M. H. LARSON?

'Biological Resources Division, Agriculture Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6
"Department of Botany, University of Toronto, 25 Willcocks Avenue, Toronto, Ontario MSS 3B2

Catling, Paul M., and Brendon M. H. Larson. 1997. The decline and current status of the dune race of Dwarf Cherry, Prunus
pumila var. pumila, on the Canadian shores of the lower Great Lakes. Canadian Field-Naturalist 111(2): 187-193.

Based on a survey of the floristic literature, herbarium collections, and recent field study, the dune race of Dwarf Cherry,
Prunus pumila var. pumila, has declined at most of 13 regions of occurrence on the Canadian shores of the lower Great
Lakes and is considered threatened. The disappearances or declines at Lake Ontario sites, and at Long Point National
Wildlife Area on Lake Erie, and at Pinery Provincial Park on Lake Huron, all followed very high populations of White-
tailed Deer (Odocoileus virginianus), and current absence or rarity of the shrub at some other locations can also be associ-
ated with high deer populations in recent times. Deer have had major impacts on herbs and shrubs in some of the shoreline
parks, and were observed to selectively browse Dwarf Cherry. Other factors have probably also played a role in the decline
of Dwarf Cherry at some sites, including erosion due to high water levels and shoreline modifications associated with
human use. Accurate monitoring of significant natural resources and more effective management of deer populations
appears necessary to maintain biodiversity, not only for posterity and science, but also for potential utilization of genetic
resources by people.

Key Words: Prunus pumila var. pumila, Dwarf Cherry, dune race, Odocoileus virginianus, White-tailed Deer, conserva-

tion, biodiversity, germplasm, Lake Erie, Lake Ontario, Lake Huron, Ontario.

The dune race of Dwarf Cherry (Prunus pumila L.
var. pumila), and other races within the Dwarf
Cherry complex, were selected and cultivated for
fruit production in the late 1800s (Bailey 1892;
Wight 1915). Although little exploited since that
time, they are currently important with respect to the
development of new Canadian crops and as breeding
stock and sources of resistance genes for improve-
ment of existing cherry crops (Lezzoni et al. 1991).
Recently, Small (1995) has noted the importance of
native fruits as diversification crops in Canada and
Davidson (1995) has stressed the need for a concert-
ed effort to determine which taxa are in need of pro-
tection. Regardless of previously disputed taxonomic
rank (see below), the patterns of geographic varia-
tion represented by the four currently recognized
varieties of Dwarf Cherry would provide a useful
basis for germplasm protection and selection. In
many cases the status of native germplasm is
unclear, and this applies particularly to native berry
plants. However, some species or races are definitely
rare or threatened in Canada. In 1993 the most
restricted race of Dwarf Cherry, P. pumila var. pumi-
la, could not be found at one of the best known and
accessible stations in Presqu’ile Park, eastern Lake

Ontario. This race of Dwarf Cherry was recently
highlighted as one of a number of priorities for the
protection of Canadian native germplasm of eco-
nomically important plants (Catling and Porebski
1996). Its apparent decline not only required expla-
nation, but also raised the question of its history and
current status elsewhere. In order to contribute to the
protection of the dune race of Dwarf Cherry we pre-
sent information relating to decline and current
status on the Canadian shores of Lake Erie, Lake
Ontario and southern Lake Huron.

Taxonomic status of Dwarf Cherry

Fernald (1923) established Prunus susquehanae
Willd. as the correct name for the plant of acid sandy
or rocky barrens known as P. cuneata Raf., and dis-
tinguished the prostrate P. depressa Pursh of river
gravels from the erect P. pumila L. of Great Lakes
sand dunes. In his 8th edition of Gray’s manual
(Fernald 1950) he maintained the three eastern
species and included in the group the western P.
besseyi Bailey, which was reported from Manitoba
and Kansas west to Wyoming and Colorado. More
recent workers have tended to recognize only one
species in the Dwarf Cherry group in eastern North

187

188

America, referring all the eastern plants in the group
to P. pumila (e.g. Boivin 1966; Soper and
Heimburger 1982; Voss 1985). Groh and Senn
(1940) were also “unable to follow Fernald” and
noted that the “entities show a complex series of
intergradations”, but they did report patterns of
occurrence of the taxa in Ontario similar to those
outlined by Fernald (1923). Recently, Gleason and
Cronquist (1991) recognized the four species as vari-
eties of Prunus pumila, the var. pumila being the
erect, narrow-leaved shrub of dunes along the shores
of the Great Lakes. Subsequent reference here to
“Dwarf Cherry” refers to the erect, narrow-leaved
plant of the Great Lakes dunes.

Methods

Field surveys were conducted at Point Pelee
National Park and Rondeau Provincial Park on Lake
Erie, and Presqu’ile Provincial Park, Outlet Beach
Provincial Park, and Sandbanks Provincial Park on
Lake Ontario. Appropriate habitats were systemati-
cally searched in 10 m wide paths and the numbers
of shrubs seen as well as any evidence of browsing
was recorded. Specimens at CAN, DAO, HAM,
LKHD, OAC, QK, TRT, TRTE, UWO, and WAT
(acronyms from Holmgren et al. 1990) were exam-
ined to determine when P. pumila var. pumila was
present and what its status was in the past.
Specimens were identified using Fernald’s (1950)
key, but admitting to var. pumila plants with pits 5-8
mm wide instead of 6-8 mm wide. The concept of P.
pumila var. pumila applied here thus involves a nar-
row-leaved, erect plant with a broad range of pit
widths. Locations of collections were mapped and
regions of occurrence, i.e.; areas where plants were
found from | to 200 acres in extent, up to 7 km in
length, were indicated. Field botanists and park offi-
cials were contacted for information on current sta-
tus, and available reports concerning vegetation
management and monitoring were examined.
Relevant botanical literature and crown survey
records were also searched for information on the
occurrence and status of P. pumila var. pumila on
the Canadian shores of the lower Great Lakes.

Results
Distribution pattern

Specimens exist to substantiate reports from all of
the sites listed in Table 1 and shown in Figure 1.
Specimens examined were referable to Prunus pumila
var. pumila, the only exceptions being some collec-
tions from the Inverhuron shore on Lake Huron, and
Point Abino on Lake Erie (Figure 1, Table 1), some of
which were referable to var. depressa (Pursh) Bean.
The only specimens collected near the shore referable
to P. pumila var. cuneata (Raf.) L. H. Bailey (=P.
susquehanae) were those from sandy oak woodland
and prairie-like scrub one to five km distant from the

THE CANADIAN FIELD-NATURALIST

Vol. 111

Lake Erie shore north of Long Point as well as that
reported from Lambton, 2 km north of Lake Ontario
in the Humber valley of Toronto (Scott 1913), where
a similar habitat once existed. Since 1882 Dwarf
Cherry has been found at 13 shoreline regions in the
eastern lower Great Lakes (Figure 1, Table 1).

Lake Ontario

In 1901 Mrs. Burnham Greeley reported a
“species of bilberry which was greatly prized by the
early settlers” in western Prince Edward County
(Carlisle et al. 1973). Since there are no ericaceous
plants in the area, there seems to be little doubt that
this was a reference to Prunus pumila, which is often
referred to locally (and incorrectly) as bilberry or
huckleberry.

Dwarf Cherry was reported from Sandbanks
(Figure | - region 1) by Klugh in 1911. It was com-
mon there until the early 1970s (personal observa-
tion, specimen labels at DAO — Table 1; Carlisle et
al. 1973*; Neugebauer 1974*). In 1994 there were
still more than 30 patches in the more remote central
portions of the bar, but it had become very scarce at
either end, where once abundant. The remaining
population, however, appears to be the largest along
the Canadian shores of Lakes Erie and Ontario. In
the early 1970s, there were still at least seven large
clumps on the foredune at Outlet Beach (personal
observation), but none could be found in 1993.

Carlisle et al. (1973*) indicated that Dwarf Cherry
was one of the dominant species along the front and
top of Huycks Bay Bar (Table 1, Figure 1 — region
2). It was frequent at North Beach in 1990 (S.
Blaney, personal communication). No additional
information is available for either of these sites.

At Presqu’ile Provincial Park (Figure 1 — region
3), Dwarf Cherry was frequent on the foredunes in
the late 1960s and early 1970s, when there were at
least 15 large patches at the south end of the beach
opposite Bluff Island (personal observation). It could
not be found in 1993 (personal observation), but a
single stem was located in 1996 (S. Blaney, personal
communication).

Lake Erie

Zenkert (1934) reported Dwarf Cherry as rare but
locally abundant at intervals along the shore from
Point Abino to Port Colborne (Figure | — region 1).
It was collected on Point Abino in 1941, 1948 and
1975, and seen by Yaki and MacDonald in 1988
(MacDonald 1990*). Current status in this area is
unclear.

In 1793* William Chewett, then senior surveyor
for Upper Canada, noted “innumerable quantities of
Dwarf Cherry” eight miles from the eastern extremity

*Document, see Documents Cited between Acknowledg-

ments and Literature Cited.

oF

440

430

420

CATLING AND LARSON: DECLINE OF DWARF CHERRY

189

80° 79° 78°

FicurE 1. Distribution of locations where Prunus pumila var. pumila has been found on the Canadian shores of the lower
Great Lakes (dots) and corresponding regions (numbers). For the basis of the map and key to numbers, see Table 1.

of Long Point (Figure 1 — region 5). James Black’s
1854* survey included a “shrub community” extend-
ing from the tip of the point 12 km to the west [see
also Heffernan (1978*) and Heffernan and Nelson
(1979)]. Dwarf Cherry was one of the species fre-
quently noted in this zone on Black’s transects.
Dwarf Cherry was probably not listed in subsequent
early surveys of the flora of Norfolk County because
it had already become rare. Dwarf Cherry was pre-
sent, but very rare on the outer portions of Long Point
and particularly in the Gravelly Bay area in the late
1960s (personal observation). Heffernan and Nelson
(1979) noted that no Dwarf Cherry occurred in the
areas where it had been reported by Chewett (1793*)
or Black (1854*), and that there were only a few,
very small clumps in the Provincial Park at the base
of Long Point. Dwarf Cherry was not seen on the
outer portions of the point in the 1980s by Reznicek
and Catling (1989), who also reported it only from
the park at the base of the point.

There is a specimen of Dwarf Cherry from
Rondeau Park (Figure 1 — region 6) collected in
1905 (Table 1), but there are no other records from
this location, nor was any reference to it at this loca-
tion found in any of the early surveyor’s notes. It
could not be located there in 1994 during a survey of

the entire east beach, although there is extensive
sandy shore habitat.

Dodge (1914) noted that Dwarf Cherry was “fre-
quent at Point Pelee (Figure | — region 7) in sandy
open ground and on upper beach.” Some later refer-
ences with almost identical wording (e.g., Mosquin
1988*) are evidently based on Dodge’s report.
Specimens were collected here between 1882 and
the early 1960s (Table 1). W. Botham, who visited
Point Pelee frequently between 1950 and 1980 lists
no flowering dates, yet he made field notes exten-
sively. Thus he likely either did not observe it or it
was very infrequently observed. The only reference
he makes to it is a specimen collected by Frith in
1961 in the collection at the park (Botham 1981).
The park herbarium contains specimens collected by
Frith on the west beach in 1961 and on the east
beach in 1962. A specimen collected by Neal in
1948 was from near the tip thus demonstrating that
the plants were widespread in the park. Maycock et
al. (1978*) found it “restricted and rare” along the
west beach in the late 1970s (see also Battin 1975*).
Failure to locate Dwarf Cherry in the park in 1994
following a search of both the eastern and western
beaches, provides further evidence of a decline at
Pelee since Dodge’s time (1914).

190

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE |. Locations of Prunus pumila var. pumila on the Canadian shores of Lakes Erie and Ontario. For locations in liter-
ature for which no justifying specimens were found, at least one literature report or personal communication is included.
Herbarium acronyms are from Holmgren et al. (1990).

Location

Lake Ontario
1. Sandbanks and Outlet

Outlet Beach

2. Huyck’s Bay and North Beach

North Beach
3. Presqu’ile Park

Lake Erie

4. Point Abino & Port Colbourne

5. Long Point, base

Long Point, middle
Long Point, tip
6. Rondeau Park
7. Point Pelee
South Point
near tip
east & west beach

8. Fish Point, Pelee Island

Southern Lake Huron
9. Walpole Island
10. Sarnia, Caratara Park
Point Edward
11. Grand Bend to Kettle Point
Grand Bend

Pinery Park

Port Franks

Ipperwash

Kettle Point
12. Wright Point
13. Inverhuron shore

Latitude/Longitude

44 56 30 77 19 30
43 54 00 77 16 00

43 54 00 77 17 00
43 55 20 77 18 00
43 55 00 77 17 00
44 53 3077 13 30
44 56 00 77 29 45
43 57 30 79 31 30
44 00 00 77 44 00

42 50 00 79 05 00

42 53 00 79 06 00
42 34 00 80 09 00

42 33 00 80 06 00
42 33 00 80 03 00
42 19 00 81 51 00
41 45 00 82 40 00
41 57 00 82 31 00
41 54 00 82 31 00
41 45 00 82 40 00
42 30 00 82 33 00
43 00 20 82 24 20
43 00 00 82 25 10
43 19 00 81 45 00

43 16 00 81 50 00

43 13 00 81 54 00

43 13 00 81 58 00

43 13 00 81 57 00
43 13 00 82 01 00
43 48 00 81 44 00
44 17 00 81 35 00

There are reports of Dwarf Cherry from Fish Point
(Figure 1 — region 8), on Pelee Island (e.g.; Jones
1912; Campbell 1976*), but, interestingly, some
important regional sources do not list it (e.g., Dodge

year of collection/collector/herbarium or reference

1993 Catling (DAO)

1947 Gillett (DAO), 1948 Soper (TRT, DAO, OAC),
1952 Calder (DAO), 1960 Hainault (QK),

1964 Lubke (UWO), 1965 Garwood (QK),

1967 Garwood et al. (QK), 1967 Baum (DAO),

1969 Catling (LKHD, QK, TRT), 1969 Saifi (CAN,
LKHD, TRT), 1977 Conley (OAC), 1979 Crowder (QK)
1981 Ball (TRTE)

1993 Catling (DAO)

1973 Gad et al. (LKHD, TRT), 1980 Bezdek (TRTE)
1973 Carlisle et al.

1973 Carlisle et al.

1990 Blaney, personal communication

1969 Catling (TRT)

1941 Soper (DAO, OAC, TRT), 1951 Landon (OAC),
1975 Catling & McKay (DAO, TRT)

1975 Catling & McKay (DAO, TRT)

1951 Falls & Klawe (TRT, WAT), 1980 Pringle (HAM),
1980 Reznicek & Catling (DAO, MICH)

Chewett (1793)

Black (1954)

1905 Fall (TRT)

1892 Macoun (CAN)

(CAN)

1961-62 Frith (Pelee Park herbarium)

1882 Burgess (DAO, WAT), 1948 Neal (DAO)
1892 Macoun (CAN)

1957 Gillett & Senn (DAO)
1962 Stewart (DAO, UWO)
(CAN)

1936 Sutton (UWO), 1940 Soper (DAO, TRT),
Brown (TRT)

1940 Gaiser (HAM), 1960 Scoggan (CAN),

1962 Shaw (UWO), 1969 Cody (DAO),

1974 Montgomery (WAT), 1974 Helley (OAC)

1891 Dearness (DAO), 1919 Tripp (OAC),

1934 Wood (DAO), 1941 Fox (UWO),

1941 Colgrove (UWO), 1957 Gaiser (OAC),

1959 Lamarre (QK), 1983 Tiedje (UWO), Soper (TRT),
Crabe (TRTE)

1906 Tripp (OAC), 1940 Soper et al. (DAO, GH, HAM, TRT);

1946 Gaiser (HAM), 1955 Tamsalu (HAM),

1957 Yang (OAC), 1958 Lamarre (DAO),

1982 Aboud et al. (UWO)

1960 Stewart (DAO, UWO)

1934 Gaiser (DAO, HAM)

1990 Dalpe (DAO)

1963 Stewart (DAO)

1914; Core 1948; Botham 1981). There is, however,
a specimen collected in 1892 (Macoun 5608, CAN).
Dwarf Cherry has not been observed in several com-
prehensive field surveys over the past seven years. It

LOOT

may never have been common on the limited sandy
habitats on the island.

Lake Huron and Lake St. Clair

There are old collections of Dwarf Cherry from
Walpole Island Indian Reserve (Figure | — region 9)
and it occurred on the north end of Squirrel Island
until 1969 (personal observation). Extensive recent
botanical surveys of this valuable natural area over
the past 10 years have failed to find Dwarf Cherry
(G. Allen and R. Brown, personal communication).
There is limited habitat at this site and it may never
have been common.

There are old collections of Dwarf Cherry from
Sarnia (Figure | — region 10, Table 1), but it was not
listed for Sarnia by Tiedje and Tiedje (1989). Since
botanists have been fairly active in the area (M. J.
Oldham, personal observation), it seems likely that
Dwarf Cherry is no longer present.

Until seven or eight years ago Dwarf Cherry was
abundant or frequent on the first dune ridge extending
from Grand Bend to Kettle Point (Figure | — region
11) , a strip of shore 25 km in length (personal obser-
vation; Sparling 1965). It is estimated that Dwarf
Cherry has been reduced to 10% of its former cover
over the past 10 years along this shore (M. A. Maun,
personal communication; Maun and Crabe, in press).
Nevertheless, this and the population at Sandbanks on
Lake Ontario are the largest remaining populations on
the Canadian shores of the lower Great Lakes.

Dwarf Cherry has been collected at several loca-
tions along the Lake Huron shore near Inverhuron
(Figure 1 — regions 12 and 13, Table 1), but no addi-
tional information is available on its status in this
region or at the nearby Farm Point Provincial Park.
Some of the collections from this area are recent in
contrast to most Lake Erie regions of occurrence.

Possible causes of decline
Evidence for excessive browsing by White-tailed

Deer as a major cause

Recent declines at all of the Lake Ontario sites
(regions 1, 2, and 3) as well as the Grand Bend —
Kettle Point dunes on Lake Huron (region 11) have
been coincident with major increase in populations
of White-tailed Deer, Odocoileus virginianus, over
the past 10 years (D. Tyerman, personal communica-
tion; Gedge and Maun 1992). In the latter regions
deer have increased to a very high population level
over the past 10 years, and the present level is four
times that at which deer and browsed vegetation can
coexist in balance (Gedge and Maun 1992). The deer
have had a devastating impact on low woody vegeta-
tion (Landplan Collaborative 1992*; Pearl et al.
1995*; Bakowsky 1996*).

The relatively well documented declines on Long
Point prior to 1920, were also coincident with the
development of very dense deer population. Shrubs
were generally scarce on Long Point in the late

CATLING AND LARSON: DECLINE OF DWARF CHERRY

191

1920s, the scarcity at that time being attributed to
“overstocked” deer (Snyder 1931a, b). Snyder
included photos of forest and dunes lacking shrubs,
and in this respect, comparable to photos published
58 years later (Reznicek and Catling 1989). Deer
were introduced to Long Point on several occasions
after 1870 during a period when they were extirpat-
ed on the mainland (Snyder 1931a, b), and they
have continued to profoundly influence the vegeta-
tion until very recently (McCullough and Robinson
1988*). Reznicek and Catling (1989) suggested that
the absence of shrubs like Dwarf Cherry over large
parts of Long Point was probably attributable to the
large deer population. They similarly reported
(1989, page 157) Dwarf Cherry as “rare, a few
shrubs on low dunes near the base of the Point in
Long Point Provincial Park’, one of the few regions
of the point where heavy human traffic leads to rel-
atively fewer deer.

Declines at Pelee and Rondeau may have slightly
preceded the deer population explosion in those loca-
tions, but may be accounted for through selective
browsing by a steadily growing deer population. The
current rarity of Dwarf Cherry at these two sites can
be associated with extremely dense populations of
deer in recent times. The White-tailed Deer popula-
tion in Rondeau was very high between 1912 and
1942 and most of the shrubs were destroyed by
browsing during this period (Bartlett 1958*). In the
late 1960s, at about the time that Dwarf Cherry was
last observed in the park, the deer population in Pelee
was below the maximum sustainable population level
(Theberge and Oosenburg 1975*), but by the early
1990s the deer population had increased six times and
was far beyond this level (Maun and Crabe 1995).

In addition to coincidence of the decline of Dwarf
Cherry with the increase in deer and well-document-
ed impacts of a few other observations also suggest
deer as a major cause of Dwarf Cherry decline.
Species of Prunus, including Dwarf Cherry, are
among the plant species most browsed by deer where
they are available (Miller et al. 1992; Strole and
Anderson 1992). Dwarf Cherry was selectively
browsed by deer at Sandbanks Provincial Park (per-
sonal observation) and at Pinery Provincial Park (M.
A. Maun, personal communication), and at both
locations it was preferred over a number of other
woody species. Finally, Dwarf Cherry was least
impacted at Long Point and the Grand Bend — Kettle
Point shore in built up areas where density of deer
was lower (personal observation).

Other factors

The first suggestion of a decline on the Outlet
Beach bar was that of Carlisle et al. (1973*), who
noted that Beach Grass (Ammophila breviligulata
Fern.) and Dwarf Cherry were not as common at
Outlet as at Sandbanks probably due to increased
human traffic at the former site. At Presqu’ile, in

192

addition to human traffic, prevention of dune forma-
tion due to scraping of the beach to keep it clean and
level, probably contributed to the decline of Dwarf
Cherry. Nutrient loading of sand due to use by peo-
ple and Ring-billed Gulls (Larus delawarensis Ord)
may also have contributed to the decline at this loca-
tion. The gulls nest nearby and the beach is some-
times used as a resting area by thousands of birds.

On Long Point changes in sand deposition due to
high water levels and shoreline modification have
reduced dune habitat in some areas, but as noted by
Reznicek and Catling (1989), sand is still accumulat-
ing in places and the major shoreline changes are
recent, whereas Dwarf Cherry declined on Long
Point prior to 1920.

In 1986, high water levels (1.5 m higher than nor-
mal) rapidly eroded half of the first dune along the
Grand Bend — Kettle Point shore and when the water
levels dropped, rapid sand accumulation buried
many Dwarf Cherry plants (Gedge and Maun 1992).
Recovery from such events is often rapid, however,
leading to the conclusion that this was not the most
important factor in recent decline.

Conclusions

Considering the preceding observations, Dwarf
Cherry has declined in most of 13 regions of occur-
rence on the Canadian shores of the lower Great
Lakes, including all major stations. Severely browsed
Dwarf Cherry could be easily overlooked, so it may
be premature to say that it has been extirpated in
some areas, but regardless of whether it has declined
or totally gone, it is threatened on the Canadian
shores of the lower Great Lakes. Since the dune race
is essentially confined to the Great Lakes shores, and
is also rare along much of the United States shore-
lines of the lower Great Lakes (Bonanno 1993;
personal observation), this means that it is at least
seriously threatened globally. The two regions where
it is still fairly common, Sandbanks and Grand Bend,
are particularly important for conservation.

There is compelling evidence that browsing by
deer has been an important factor in the decline, but
other factors have probably also played a role. A
number of the sites where Dwarf Cherry has
declined are parks that have had management plans
for many years, but to adequately protect significant
germplasm and rare plants in situ, a focussed pro-
gram of monitoring and management, including
management of deer populations, is necessary. Very
extensive data on the impact of excessive browsing
by deer on the natural ecosystem and on endangered
species has been insufficient to produce effective
management. Impact on humans through loss of
potentially valuable genetic resources should result
in a more determined effort. At some locations
where the shrubs have only recently been severely
browsed, a substantial recovery from root crowns

THE CANADIAN FIELD-NATURALIST

Vol. 111

and aerial stem bases may be possible, but this pos-
sibility diminishes as heavy browsing continues.

Acknowledgments

V.R. Catling assisted with the field survey. W. J.
Cody provided useful comments on the manuscript.
Other field biologists providing valuable information
included S. Blaney, J. Bowles, M. Gartshore, M. A.
Maun, D. Metsger, G. Moulin, M. J. Oldham, J.
Robinson, D. Tyerman, G. Waldron, and A.
Woodliffe. S. Porebski assisted with mapping. A.
Day of the Crown Land Survey Records Section of
the Ontario Ministry of Natural Resources
Information Branch kindly searched the field notes
of early surveyors for references to occurrence and
status of Dwarf Cherry in the Lake Erie region.

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Black, J. 1854. Field notes and diary: survey of Long
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Campbell, C. A. 1976. Preliminary ecological report on
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Received | May 1995
Accepted 23 October 1996

A Taxonomic Study of the Grass Genus Glyceria (Mannagrass) in

British Columbia

ROSARIO RUIZ DE ESPARZA and JACK MAZE!

Department of Botany, University of British Columbia, Vancouver, British Columbia V6K 1Z4

‘Author for correspondence.

de Esparza, Rosario Ruiz, and Jack Maze. 1997. A taxonomic study of the grass genus Glyceria (mannagrass) in British

Columbia. Canadian Field-Naturalist 111(2): 194-199.

The relationships among the nine species of Glyceria occurring in British Columbia were evaluated using principal compo-
nents analysis of morphological features. The analysis resolved five distinct species and two groups, each consisting of a
pair of similar species. Independent analyses of these two groups indicated both consisted of different, albeit close, species.
The interpretation that quantitatively similar species are distinct is corroborated by qualitative differences among those

same species. A key to the nine species is included.

Key Words: Glyceria, mannagrass, principal components analysis, British Columbia.

The grass genus Glyceria is composed of about 25
species, mostly of temperate to subarctic regions of
the Northern Hemisphere. The species are all tender
and readily eaten, but furnish comparatively little
forage since they occur in wet areas, usually in limit-
ed quantity (Hitchcock et al. 1969).

The status of the genus in the Province of British
Columbia is not a matter on which there is agree-
ment (Table 1). An objective of this study was to
clarify what species occur in the province, offer
insight into potential taxonomic problems that have
surfaced as the new Manual of Grasses is being writ-
ten; in particular the relationship between G. striata
and G. elata (Barkworth, personal communication)
and to quantify the relationships among the species
occurring in British Columbia as a first step in deter-
mining taxonomic problems requiring more detailed
study.

As a Starting point, we used the species which had
been listed for British Columbia by different authori-
ties (Table 1), but not all the species listed were
included in further analyses. Glyceria declinata and
G. fluitans were not found in the herbaria used and
were thus excluded from this study and the species
G. pauciflora was earlier transferred to other genera,
Torreyochloa (Church 1949) or Puccinellia
(Hitchcock et al. 1969).

In order to quantitatively describe the relation-
ships among the species of Glyceria we used
Principal Components Analysis (PCA) of a correla-
tion matrix to assay for distinct groups and deter-
mine the features which best distinguish among
those groups should they coincide with species des-
ignations. PCA has been used, with some success, to
estimate relationships among other grass species by
Allred (1984), Barkworth (1970) and Rapson and
Maze (1994). Discriminant function analysis or

canonical variates analysis was not used since they
assume the existence of groups whereas we wished
to test for the existence of such.

The most complete key to the Glyceria of British
Columbia is in the Vascular Plants of British
Columbia edited by Douglas et al. (1994). A final
objective of this study was to update the key of
Douglas et al. (1994) by including our results on the
nine species analyzed here.

Materials and Methods
Material studied:

Collections of G. borealis (Nash) Batch., G.
canadensis (Michx) Trin., G. elata (Nash) M. E.
Jones, G. grandis S. Wats. ex A. Gray, G. lep-
tostachya Buckl., G. maxima (Hartm.) Holmb., G.
occidentalis (Piper) J. C. Nels., G. pulchella (Nash)
K. M. S. and G. striata (Lam.) A. S. Hitch. were
obtained from UBC, UAC and ALTA. A list of the
specimens used in the analysis can be obtained from
the senior author on request.

Characters:

The first data set for PCA comprised 16 characters
assessed for each of 195 specimens (OTUs). Two
criteria were used in choosing the variables. They
reflected the form of the grasses and were readily
measurable. These variables are: Measurements
taken in the Flag leaf: Length of the sheath
(SHEATH), blade (BLADEL), ligule (LIGULE) and
the width at the mid point of the blade (BLADEW).
Inflorescence: Overall length to the tip of the termi-
nal spikelet (INF); overall length to the tip of the ter-
minating spikelet of the lower most branch
(BRANCH) and length from the inflorescence axis
to the first node of the lower most branch (NODE).
The spikelets measured were chosen from the middle

194

1997

DE ESPARZA AND MAZE: GLYCERIA IN BRITISH COLUMBIA

H9

TABLE 1. List of the species recognized as growing in British Columbia by four authorities. +, species recog-
nized; -, species not recognized or beyond the geographic range of the work; ASH, Hitchcock and Chase
(1951); CLH, Hitchcock et al. (1969); RLT, Taylor and MacBryde (1977); GWD, Douglas et al., 1994.

Species

G. borealis +
G. canadensis -
G. declinata

G. elata

G. fluitans

G. grandis

G. leptostachya

G. maxima

G. occidentalis

G. pauciflora

G. pulchella

G. striata

Total number of species recognized

Ntr tee +e $i

of the inflorescence, the measurements taken were:
Spikelet width at the middle (SPIKW) and length
(SPIKL); and the length of the first and second
glumes (GLI and GL2). In the same spikelet we
measured in the lowest floret the length of the:
Lemma (LEMMA), palea (PALEA), anther
(ANTHER) and scarious portion at apex (SAPEX).
The number of anthers per floret were counted
(NUANT). The variables are listed in Table 2.

TABLE 2. Characters and character states used in principal
components analysis.

Variable Acronym
Flag leaf
Sheath length SHEATH
Blade length BLADEL
Blade width at the midpoint BLADEW
Ligule length at the middle LIGULE
Inflorescence
Overall length from lowest node to
tip of terminal spikelet INF
Length of the lower most branch to
tip terminating spikelet BRANCH
Length of lower most branch from the
inflorescence axis to the first node NODE
Spikelet (chosen from middle of inflorescence)
Spikelet width at the middle SPIKW
Spikelet length SPIKL
First glume length GLI
Second glume length GL2
Lowest floret of measured spikelet
Lemma length LEMMA
Palea length PALEA
Number of anthers NUANT
Anther length ANTHER
Length of scarious portion of
apex measured at midvein SAPEX

Authorities
CLH RLT. GWD

+ + +
- Ae 2
= ak =
+ + +
= + z=
+ + +
+ + +
- - +
+ + +
= + ~
+ +

"/ 10 8

Analyses:

A PCA on the data for all species analyzed. For
for those instances where species were not clearly
resolved in the initial analyses, a PCA on those
species alone was done to see if the resolution could
be improved by excluding those species that strongly
polarize the data.

The final analyses were not always based on the
full set of variables. For all comparisons made, an
initial PCA was done on all variables. The variables
were then ranked by the square of their eigenvector
loadings adjusted by the eigenvalue and summed
over all axes. Those sums were then analyzed for
natural breaks, variables separated by large values,
and all variables with summed values above that nat-
ural break were used in the next PCA. This approach
was used by Maze et al. (1992) and is designed to

TABLE 3. PCA of all species. Variable acronyms as in
Table 2.

PCA axes
| 2

Eigenvalues 6.54 1.23
Percent variation

accounted for 65.38 12.26
Eigenvectors

SHEATH 0.780 0.462

LIGULE 0.754 0.518

INF O:771 0.338

SPIKL 0.904 0.125

GLI 751 -0.482

GL2 0.848 -0.324

LEMMA 0.880 -0.212

PALEA 0.912 -0.184

NUANT 0.763 0.204

ANTHER 0.700 -0.408

196

Factor Two
_ Oo
9 eR
No
es

Factor One

FIGURE |. 70% confidence ellipses for all nine species.
Factor one, first PCA axis; Factor two, second PCA
axis; 0, Glyceria occidentalis; b, G. borealis; m, G.
maxima; |, G. leptostachya; g, G. grandis; p, G. pul-
chella: c, G. canadensis; e, G. elata; s, G. striata.

eliminate those variables whose vectors do not close-
ly parallel major axes of variation; it is an approach
designed to eliminate “noise” in the data. Three
PCAs were performed, one for all species, one com-
paring G. canadensis and G. pulchella and one com-
paring G. elata and G. striata.

The results of the PCA, the PCA axis scores,
were used to produce scatter plots, summarized as
confidence ellipses, and subjected to analysis of
variance (ANOVA) to offer a quantitative summary
of the scatter plots. For this we relied on Tukey's
multiple range test. In all the comparisons using
PCA axis scores we used only the first two axes as
these are the ones that account for most the varia-
tion in the data.

These and subsequent analytical comparisons
were done using SYSTAT Network Version 5.04
Wilkinson (1991).

Results
Analysis of all species:

The first PCA, done with all the variables for all
species, resulted in eliminating six variables,
BLADEL, BLADEW, BRANCH, NODE, SPIKW
and SAPEX. In the PCA on the remaining variables
(Table 3), the first axis of the PCA explained
65.38% of the data variation. The characters load-
ing heaviest on this axis pertained to the spikelet
and floret characteristics; the four most important
were: PALEA, SPIKL, LEMMA and GL2.
However all variables have fairly high loading and
the same sign indicating the first axis most likely

THE CANADIAN FIELD-NATURALIST

Vol T#]

reflects size differences. The second axis explained
12.26% of the data variation and the most impor-
tant characteristics were LIGULE, GL1, SHEATH
and ANTHER. The presence of both positive and
negative signs in the loading on the second axis
indicate that differences along that axis represent
differences in shape.

Distribution of the study samples along compo-
nents | and 2 is illustrated in the Figure 1. Seven
groups can be recognized. Two consist of pairs of
closely related species, G. canadensis — G. pul-
chella and G. elata — G. striata, and five species
standing apart from each other. Three species, G.
grandis, G. leptostachya and G. maxima form a
linear series of two related pairs along the first
axis, G. grandis and G. leptostachya and G. lep-
tostachya and G. maxima. However these three
species were not subjected to further analyses.
The main axis of variation for G. leptostachya, G.
grandis and G. maxima is orthogonal, or nearly so,
to the main axes of variation of all the species.
Thus when analyzed alone they would be separated
along their main axes of variation. The same rea-
soning was used in not subjecting G. borealis and
G. maxima, which overlap on the first axis, to fur-
ther analyses.

In the Tukey analysis (p > 0.05), G. canadensis
and G. pulchella were not significantly different on
components | and 2, and in six SHEATH, LIGULE,
INF, SPIKL, GL1 and PALEA, characteristics; G.
elata and G. striata were not different on factor 2
and seven characteristics; SHEATH, LIGULE,
SPIKL, GL1, GL2, LEMMA, and PALEA. Because
of the similarities between first two pairs of species
it was necessary to include each pair in further anal-

TABLE 4. PCA of G. canadensis and G. pulchella. Variable
acronyms as in Table 2.

PCA axes
1 ys
Eigenvalues 4.06 3225
Percent variation
accounted for 33.83 og ae
Eigenvectors
SHEATH 0.640 0.469
BLADEL 0.720 0.319
BLADEW 0.595 0.443
LIGULE 0.332 0.683
INF 0.812 0.166
BRANCH 0.548 0.470
RACHIS 0.169 0.653
SPIKW 0.462 -0.695
SPIKL 0.760 -0.440
GL2 0.416 -0.622
LEMMA 0.666 -0.631
PALEA 0.521 -0.363

Foo 7,

Factor Two

G. canadensis

-3 -2 -1 0 1 2 3 4
Factor One

FIGURE 2. 70% confidence ellipses for Glyceria elata and
G. striata. Factor one, first PCA axis; factor two,
second PCA axis.

yses, following the same criteria established for all
the species.

Analysis of G. canadensis and G. pulchella:

An initial PCA of only G. canadensis and G.
pulchella using all variables indicated that
BRANCH, GL1, NUANT, ANTHER and SAPEX
contributed little to the main axes of variation.
Those variables were eliminated and a second PCA
was done. The results of that PCA are presented in
Table 4. The first axis has all positive loadings
with the largest being those for INF, SPIKL,

TABLE 5. PCA of G. elata and G. striata. Variable acro-
nym as in Table 2.

PCA axes
1 o)
Eigenvalues 5.04 2.60
Percent variation
accounted for 38.81 20.01
Eigenvectors
SHEATH 0.634 O57
BLADEL 0.582 0.480
BLADEW 0.646 0.204
INF 0.796 0.387
BRANCH 0.772 0.395
RACHIS 0.622 0.336
SPIKL _ 0.463 -0.576
GLI 0.629 -0.423
GL2 0.607 -0.502
LEMMA 0.344 -0.722
PALEA 0.529 -0.529
NUANT 0.502 -0.440
ANTHER 0.801 0.160

DE ESPARZA AND MAZE: GLYCERIA IN BRITISH COLUMBIA

LOW

BLADEL, LEMMA and SHEATH; it accounts for
33.83% of the variation in data; the second axis
accounts for 27.11% of the variation in the data. It
is a shape axis with two groups of variables being
recognized, those for the spikelet variables and
those for the vegetative variables and the inflores-
cence. The variables making the strongest contri-
bution to the second axis are SPIKW, LIGULE,
RACHIS, LEMMA and GL2.

The distribution of the study samples along com-
ponents | and 2 is illustrated in the Figure 2. The
two species occur in different parts of the ordination
and are separated on both axes with the greatest sep-
aration being on the second. In the Tukey analysis
both species were significantly different on compo-
nents | and 2. The combination of the loadings and
position on the ordination indicate G. pulchella is
smaller than G. canadensis.

Analysis of G. elata and G. striata:

The initial analysis, using all the variables,
showed that LIGULE, SPIKW and SAPEX, were
not major contributors to the variation and were not
considered in the second analysis. The result of the
second PCA is shown the Table 5. The factor load-
ings indicate that component | revealed mainly dif-
ferences in ANTHER, INF, BRANCH, BLADEW
and SHEATH, explaining 38.81% of the data varia-
tion, whereas component 2 revealed differences in
LEMMA, SPIKL, PALEA, GL2 and BLADEL,
explaining 20.01% of the data variation. As in the
analysis of G. canadensis and G. pulchella, the first
axis is likely a size axis and the second shape. Also,
like G. canadensis and G. pulchella, the second axis

Factor Two

-3 -2 -1 0 1 2 3 4
Factor One

FIGURE 3. 70 % confidence ellipses for Glyceria canaden-
sis and G. pulchella. Factor one, first PCA axis; fac-
tor two, second PCA axis.

198

is bipolar with the main separation being between
spikelet and non-spikelet characters, with the
exception of ANTHER. Figure 3 presents the distri-
bution of the G. striata and G. elata along compo-
nents | and 2.

The two species are separated along the first, but
not the second axis. This relationship is reflected in
the Tukey analysis wherein the two species are sig-
nificantly different on the first but not the second
axis. The combination of loadings on the first two
PCA axes and the loadings on the first PCA axis
indicate that G. striata is the smaller of the pair.

Discussion

The PCA found a certain number of groupings
which coincide most closely with those in Douglas
et al. (1994), quantified the distinctiveness of the
groups and identified character variation of value
for identification of the groups. In those instances
where there was apparently some overlap when all
species were analyzed, analyses using only the
closely related pairs of species, G. striata — G.
elata and G. canadensis — G. pulchella, indicated
the species occupied separate parts of the
ordination.

Although the differences among the species are
not strong, we feel that we are dealing with distinct
species. The reason for this is that differences
among the species are often represented in charac-
ters that are difficult to quantify without establish-
ing artificial categories; such characters are texture,
degree of prominence of lemma nerves, the form of
apices of appendages and the shape of structures.
The combination of these variables, along with the
analyses done, allow the recognition of nine
species, and this opinion is presented in a key to the
nine species that is presented at the end of this
discussion.

The analyses of the pairs of G. canadensis — G.
pulchella and G. striata — G. elata implies that,
once the effect of size has been eliminated, there are
different growth responses with the spikelet features
showing a different response from those of the veg-
etative parts of the plant. That such should occur is
not surprising. The parts with the different respons-
es are derived from in different physiological states,
the vegetative features and inflorescence variables
are formed by an induced meristem while the parts
of the spikelet are the product of a spikelet meris-
tem. This relationship is, obviously, descriptive.
Without differently designed studies, we cannot
ascribe any sort of functional or developmental rela-
tionship that goes beyond that descriptive. But what
is indicated is that a more detailed study directly
addressing the development of these species might
give some insight into the relationship between the
developmental events in parts of different ontoge-
netic origin.

THE CANADIAN FIELD-NATURALIST

Vol. 111

Key to the species
1. Spikelets 10 - 15 mm long, linear in outline

2. Lowest lemma 5 - 6 mm long; anthers
1.1 - 1.6 mm long G. occidentalis

2. Lowest lemma 3 - 4 mm long; anthers 0.5 - 1.0
mm long

3. Blades papillose on the adaxial surface;
Lemma scabridulous only on the nerves,
the internerves usually glabrous G. borealis

3. Blades scabridulous on the adaxial surface;
lemma scabridulous on both nerves and
internerves G. leptostachya

1. Spikelets 2.5 - 6.5 mm long, ovate to oblong in
outline

4. Nerves of the lemma not prominently raised;

Spikelets 3 - 5 mm wide; palea often exposed

G. canadensis

4. Nerves of the lemma prominently raised;

Spikelets at most about 2.5 mm wide; palea
concealed by lemma

5. First glume 1.3 - 3.5 mm long; palea apices
with wide, V-shaped notches

6. Lemma with broad scarious, margins
distinctly erose at top; stamens 2;
glumes commonly obtuse. G. pulchella

6. Lemma nearly entire at apex, without or
with only inconspicuous scarious mar-
gins; stamens 3, rarely 2; glumes acute.

7. Lowest lemma 2.0 - 2.5 mm long;
anthers 0.5 - 0.8 mm long; palea 1.9 -
3 mm long. G. grandis

7. Lowest lemma 2.5 3.5 mm long;
anthers 1.0 - 2.0 mm long; palea 2.7 -
3.5 mm long, rugose G. maxima

5. First glume 0.6 - 1, rarely to 1.5 mm long;
palea apices with narrow slits.

8. Ligules of the lower leaves usually
closed in front, mostly 1 - 3 mm long;
blades 2 - 5, occasionally to 6 mm wide;
anthers 0.4 - 0.6 mm long; glumes ovate,
rounded to obtuse, first glume 0.5 - 0.7,
or even to | mm long; lemma barely 2
mm long; firm; leaf blades mostly 2 - 4
mm wide; plants 2, but usually 3 - 8 dm
tall. G. striata

8. Ligules of the lower leaves usually open
in front, mostly 3 - 6 mm long; blades
broad, 6 - 10, or even 15 mm broad;
anthers 0.7 - 0.8 mm long; glumes lance-
olate-ovate, mostly acute, first glume
mostly | but rarely 0.7 to 1.2 mm long;
lemma mostly 2 - 2.2 mm long; leaf
blades usually 4 - 10 mm wide; plants
mostly 10 - 15 dm tall. G. elata

L997

Acknowledgments

This research was supported by a scholarship from
the Secretaria de Relationes Exteriores de Mexico to
Rosario Ruiz de Esparza and by the Department of
Botany at the University of British Columbia. Olivia
Lee and Alan Reid were extremely kind and helpful
in bringing this study to fruition.

Literature Cited

Allred, K. W. 1984. Morphologic variation and classifica-
tion of the North American Aristida purpurea complex
(Gramineae). Brittonia 36: 382-395.

Barkworth, M. E. 1970. A Taxonomic study of the large-
glumed species of Stipa (Gramineae) occurring in
Canada. Canadian Journal of Botany 56: 606-625.

Barkworth, M. E. 1994. Glyceria. In The Vascular Plants
of British Columbia. Part 4 Monocotyledons. Edited by
G. W. Douglas, G. B. Straley, and D. Meidinger.
Ministry of Forests Research Program, Victoria, British
Columbia, Canada.

Church, G. L. 1949. A cytotaxonomic study of Glyceria
and Puccinellia. American Journal of Botany 36:
155-165.

Douglas, G. W., G. B. Straley, and D. Meidinger.
Editors. 1994. The Vascular Plants of British Columbia.
Part 4 Monocotyledons. Ministry of Forest Research
Program, Victoria, British Columbia.

DE ESPARZA AND MAZE: GLYCERIA IN BRITISH COLUMBIA

199

Hitchcock, A. S., and A. Chase. 1951. Manual of the
Grasses of the United States, second edition. United
States Department of Agriculture, Miscellaneous
Publication Number 200.

Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1969. Vascular Plants of the Pacific
Northwest. Part 1, Vascular Cryptograms, Gymno-
sperms and Monocotyledons. University of Washington
Press. Seattle, Washington.

Maze, J., S. Banerjee, Y. A. EL-Kassaby, and L. R.
Bohm. 1992. Quantitative genetics of integration in
Douglas fir. International Journal of Plant Science 153:
333-340.

Rapson, G. L., and J. Maze. 1994. Variation and inte-
gration in the rare grass Achnatherum (Oryzopsis)
hendersonii: phenotypic comparison with parapatric
common congeners. Canadian Journal of Botany 72:
693-700.

Taylor, R. L., and B. MacBryde. 1977. Vascular Plants of
British Columbia: A descriptive Resource Inventory.
Technical Bulletin Number 4 The Botanical Garden.
University of British Columbia. Vancouver, British
Columbia.

Wilkinson, L. 1991. SYSTAT: The System for Statistics.
Evanston, Illinois.

Received 29 December 1995
Accepted 22 October 1996

Coyote, Canis latrans, Visitations to Scent Stations in Southeastern

Alberta

SYBILLE WOELFL and MANFRED WOELFL

Canadian Wildlife Service, 4999-98 Avenue, Edmonton, Alberta 2X3 T6B
Present address: Hainerbachstrasse 4, 83052 Bruckmuehl, Germany

Woelfl, Sybille, and Manfred Woelfl. 1997. Coyote, Canis latrans, visitations to scent stations in southeastern Alberta.

Canadian Field-Naturalist 111(2): 200-203.

During a scent-station survey from July until September 1994 in southeastern Alberta, we found no differences in Coyote,
Canis latrans, response to the standard attractant FAS (Fatty Acid Scent) and Sardines, Sardina pilchardus. However, total
September visitation rate was significantly higher than in July or August. This increase was mainly caused by the increas-
ing mobility of pups. Using the scent-station method, we succeeded in documenting Coyote presence in each ecosection.
Differences in visitation rates among ecosections existed. There was a negative correlation between the number of visits

and the relief of the different ecosections.

Key Words: Coyote, Canis latrans, abundance indices, attractant, habitat, scent stations, Alberta.

The scent-station technique was developed to
obtain indices of relative abundance of carnivore
populations and was originally used for Red Fox
(Vulpes vulpes) and Gray Fox (Urocyon cinereoar-
genteus) (Wood 1959). It has now been used on
Coyotes (Canis latrans — Linhart and Knowlton
1975), Bobcats (Lynx rufus — Brady 1981;
Knowlton and Tzilkowski 1981; Diefenbach et al.
1994), Raccoons (Procyon lotor — Smith et al.
1994), Opossums (Didelphis virginiana — Sumner
and Hill 1980; Conner et al. 1983), Black Bears
(Ursus americanus — Lindzey et al. 1977), and vari-
ous small mammals (Barrett 1982; Clark and
Campbell 1983). These studies have focused on rela-
tionships between population size and scent-station
indices (Conner et al. 1983; Diefenbach et al. 1994,
Smith et al. 1994), survey design (Roughton and
Sweeny 1982; Hatcher and Shaw 1981) and efficien-
cy of odor attractants (Turkowski et al. 1983;
Roughton 1982). Debates about type and presenta-
tion of attractant, the number of days survey lines
should be operated, distances between scent-stations,
and the timing of surveys are ongoing.

The purpose of this study was to compare the effi-
cency of a standard attractant (FAS) with an easily
manageable alternative (Sardines, Sardina
pilchardus) and to evaluate the effect of habitat and
month on the visitation rate.

Study Area

The study was conducted in the Canadian Forces
Base Suffield in southeastern Alberta (50° 32'N,
110° 32’E). The 420-km2 area stretches north of
Medicine Hat along the west side of the South
Saskatchewan River and is the Dry Mixed Grass
Ecoregion comprised of three broad physiographic
units: Eastern Alberta Uplands (dominated by glacial

fluvial outwash plains: 24% of the study area),

Eastern Alberta Plains (sand hills 36%; moraine

deposits 21%) and the South Saskatchewan River
Valley (consisting of terraces, ravines and interven-
ing steep slopes; 19%) (Usher and Strong 1994). The
Suffield area lies in one of the driest regions in
Alberta. The combination of warm summer tempera-
tures (maximum 35-40°C) and low precipitation
(summer average 40 mm) produces potential mois-
ture deficits exceeding 100 mm. Winter temperatures
average -7.2°C with thin snow cover and relatively
few days with continuous snow cover. The Needle
Grass-Blue Grama (Stipa comata-Bouteloua
gracilis) community-type is the most common plant
community in the Southern part of the study area and
occurs on a variety of landforms. Plant communities
with a significant component of sagebrush
(Artemisia cana) dominate the vegetation in the
northern portion, with sagebrush-sand grass
(Calamovilfa longifolia)/needle grass and sage-
brush/needle grass community-types being the most
abundant.

Methods

Habitat classification: Usher and Strong (1994)
defined 10 different ecosections for the Suffield
National Wildlife Area. Basically following their
“Ecological Land Classification”, we distinguished
nine ecosections for our carnivore survey. We con-
sidered major landform components based on phys-
iographic and/or geologic patterns (eolian, fluvial,
glacial fluvial landforms, and morainal deposits),
vegetation (herb/shrub dominant), slope, and
drainage. We clustered similar sites and excluded
ecosites < 500 ha (Table 1).

Survey Design: Twenty-seven transect lines with
10 scent-stations each were established and con-

200

shee I

TABLE 1. Ecosections ( > 500 ha, similar vegetation and
slope), based on Usher and Strong (1994), southeastern
Alberta.

Ecosection Description % Slope % Shrub Area (ha)

E1 Sand Dunes 2-30 99.8 16433

Fl Ravines >45 42.1 665

P2 River Valley >30 89.1 1452

Gl Channel Banks 15-30 28.5 846

G2A—- Fluvial Terraces 0-5 80.1 729
type (Shrub dominant)

G 2B - Fluvial Terraces 0-9 19.9 2239
type (Herb dominant)

G3 Fluvial Outwash 0-15 4.4 11079

M1 Ground Morain 5-30 - 6987

M2 Morainal Plain 5-15 51.0 1770

trolled once a month during July, August and
September 1994. Depending on accessability the sta-
tions were checked by truck, ATV or on foot. We
placed three transects in each of the nine ecotypes,
each transect consisting of 10 stations spaced at
0.3 km intervals (Roughton and Sweeny 1982).
Distances between the transects varied according to
ecosite distribution, but were at least 1500 m apart.
At each station, the vegetation was removed, the sur-
face cleared of rocky material, leveled, and
smoothed with sifted soil within an area of 1 m2. The
contrast of foot or paw imprints was improved by
using a thin layer of flour. In the center of the circle
we placed a wooden stake 0.20 m in height above
ground and attached the attractants.

We used two attractants, fatty acid scent (FAS, see
Roughton and Sweeny 1982) and Sardines (in oil).
The liquid FAS was applied to a cottonball and
placed in perforated plastic capsules. The sardines
were wrapped in short pieces of cotton bandages
and, like the FAS capsules, nailed on top of the
wooden stake. We alternately activated the stations
with FAS or sardines on the first day of the survey
and then checked the stations for the next two con-
secutive days. During the final check of the lines we
removed the capsules or sardines from each station.
We recorded daily weather conditions, number of
inoperable stations (due to unreadable tracks because
of weather/livestock disturbance), and species of car-
nivores and other animals that visited each operable
station. While we were checking the lines, tracks
were recorded and erased, the ground was smoothed,
and a thin layer of flour added. Missing attractants
were replaced with fresh capsules or sardines as
needed.

Data analyses: We used the Chi-square goodness-
of-fit test to determine whether Coyote visitations to
the scent-stations differed between type of attractant
and month of use. Data were pooled over month to
provide a meaningful sample size. Differences
among habitat-types were analysed by a least-

WOELFL AND WOELFL: COYOTE VISITATIONS TO SCENT STATIONS

201

squares analysis of variance (ANOVA — SYSTAT,
Inc. 1992), and a Tukey Test to determine where dif-
ferences among the ecosections existed. The visita-
tion rate was calculated according to Linhart and
Knowlton (1975) (total Coyote visits/ total operable
station nights * 1000).

Results and Discussion

Of 1601 operable station nights, 91 stations were
visited by Coyotes and the visitation rate, as calcu-
lated by Linhart and Knowlton (1975), was 57.
Beside mere sniffing (n = 47) Coyotes responded to
the scent by carrying the capsule/sardine away (n =
33), defecating/urinating (n = 7) on the station,
scratching (nm = 3), and biting at the capsule (n = 1).
Scratching occured only at stations scented with
FAS.

There was no difference in visitation rate between
stations scented with FAS versus Sardines (Table 2;
x2 = 2.5, 1 df, P > 0.26). FAS is a commonly used
superior attractant that can be used as a standard to
which other attractants can be compared (Roughton
and Sweeny 1982). Our results suggest that sardines
were as effective as FAS.

Sardines are readily available and cost 10% less.
However, sardines needed more replacement due to
insect and weather related deterioration. FAS needs
to be ordered and its acidity demands precautions
during handling (Turkowski et al. 1983). Visitation
rates to FAS appeared comparable with other studies
(e.g., Sumner and Hill 1980; Roughton and Sweeny
1982; Turkowski et al. 1983). However, the basic
comparability of scent-station studies is discussed
controversially (Diefenbach et al. 1994; Smith et al.
1994).

Number of visits per site was significantly higher
in September (x? = 9.5, 2 df, P > 0.01) than in July or
August (Table 2). Roughton and Sweeney (1982)
suggested that timing of a survey was very important
and recommended a period when juveniles were dis-
persing and the population was at its annual peak.
Several studies compared the effect of seasons.
Stolzenburg and Howard (1989) found monthly
peaks in visitation rate of Coyotes during January and
September. Martin and Fagre (1988) reported sum-

TABLE 2. Coyote visits to scent-stations with two different
attractants during July, August, and September 1994, in
southeastern Alberta. ~

Attractant
Month Visits FAS Sardines
July 23 14 9
August 24 11 13
September 44 28 16
TOTAL 9] 53 38

Number of operable station nights = 1601

202

mer (July-September) and fall (October-December)
as the seasons with the highest visitation rates.

Smaller tracks at scent-stations and several sight-
ings of Coyote pups near transect lines indicated that
the higher visitation rate during September in our
study was caused by increased mobility of pups. The
number of transects visited by Coyotes increased
from 11 in July, to 13 in August, to 16 in September.

Period of highest response to scent-stations coin-
cides with time of juvenile Coyote dispersal
(Stolzenburg and Howard 1989; Martin and Fagre
1988). When the ultimate goal of scent-station sur-
veys is to assess trends in furbearer abundance, we
suggest conducting surveys during dispersal periods.
Nevertheless the advantage of obtaining higher visi-
tation rates during dispersal and therefore detecting
changes in relative abundance more easily
(Roughton and Sweeny 1982) was questioned by
Diefenbach et al. (1994). They found that increased
visitation rates increased the variance; 1.e., multiple
visits from individuals became more likely, and thus
the relationship between population size and scent-
station index changed.

There were significant differences in the number
of visits among the nine ecotypes (Table 3;
F =5.456;18.df:-P =:0.002): Differences, occurred
between ecosections M2 and El, Fl, F2, G1 and M1
(P,,, = 0.006, P,., = 0.001, P,., = 0.004, P,, = 0.040,
Py; = 9.015, respectively). However, these results
should be treated with caution because ecosection
M2 consisted of only one, relatively small unit.
Therefore, the three replicates were relatively close
together (about 1 km) and not separated by other
ecosections. This configuration was very susceptible
to multiple visits.

A conspicuous difference throughout the study
area was the varying percentage of cover (=sage-
brush-sand grass/needle grass plant community-
type) per ecosection. Stolzenburg and Howard
(1989) explained low visitation rate on one of their
two study areas in New Mexico by suggesting that
Coyotes in the more open area of the Plains of San

TABLE 3. Coyote visits to scent-stations per transect and
ecosection! during July-September 1994, southeastern
Alberta.

Ecosections

Replicate

transect Fl... FL, .B2. Gl GA G2B. Gs. Wil az
# | 0 0 1 2 7 2 a 2 8
#2 l l Do G2 162 6 2 ava
#3 3 0: ;O 4 4 1 5 | Ss

Total ee Ts Pa, Be Re. 7 DAO a AG, tee

'For definition of ecosections see Table 1.
*Excluded from analysis because transect was situated near
a Coyote den.

THE CANADIAN FIELD-NATURALIST

Vok fi

Augustin were more reluctant to investigate the
attractants than those on the brushy Jornada del
Muerto. In our study (with a non-hunted Coyote
population) the ecosections with the highest percent-
age of shrub vegetation (E1: 99.8%, Fl: 42.2%, F2:
89.1%) had the lowest visitation rates. However,
there was no correlation between number of visits at
the scent-stations and the shrub-percentage per eco-
section (r, = -0.183). Cover apparently was not an
important factor influencing the visitation rate at the
scent-stations.

We found a significant correlation between relief
(% slope) of ecosections and number of scent-station
visits (7, = -0.817, P < 0.01). Landforms with high
relief (>15%) had less visits than landforms with low
relief (<15%). The steeper and rougher the terrain,
the fewer Coyotes visited the scent-stations.

Field observations (tracks, sightings and howling
activity) often contradicted the low response to the
scent-stations and suggested Coyote presence also in
areas with no or very few visits to scent-stations.
Along the river shore (F2) or on sandy roads close to
the transects (El) we regularly found fresh Coyote
tracks. Along the river shore where Coyote move-
ment was often restricted to a travel corridor <5 m
wide between the river and the steep eroded slopes,
Coyotes passed the stations very close (< 2m), with-
out altering their travel direction or pace, apparently
ignoring the scent-stations. Smith et al. (1994)
reported that Raccoons although their travel path
was in line with an established station, ignored or
even avoided the station by veering to one side. One
explanation for failure of individual animals to visit
stations is that their prior investigation of stations,
either on the previous day or elsewhere on the tran-
sect, satisfied the animal’s curiosity.

Acknowledgments

This study was funded by the Canadian Wildlife
Service and the Canadian Forces Base Suffield. We
thank the CWS staff, especially L. Carbyn and G.
Trottier for their support during the study. D. Davies
and B. Bennett of the CFBS safely guided us
through unknown terrain. We are grateful to B. Scott
from the Alberta Energy Company for his help and
friendliness. Special thanks to E. Klausz and L.
Carbyn who got us started and shared hard labour
during the initial setup of the transects. S. Barry
helped us with the statistical analyses, and G. Mowat
commented on earlier drafts of the manuscript.

Literature Cited

Barrett, R. H. 1982. Smoked aluminum track plots for
determining furbearer distribution and relative abun-
dance. California Fish and Game: 188-190.

Brady, J. R. 1981. Preliminary results of Bobcat scent
station transects in Florida. Pages 101—103 in Pro-
ceedings of the Bobcat Research Conference 1979, Front
Royal, Virginia.

1007

Clark, T. W., and T. M. Campbell, III. 1983. A small
carnivore technique. Great Basin Naturalist 43:
438-440.

Conner, M. C., R. F. Labisky, and D. R. Progulske.
1983. Scent-station indices as measures of population
abundance for bobcats, raccoons, gray foxes, and opos-
sums. Wildlife Society Bulletin 11: 146-152.

Diefenbach, D. R., M. J. Conroy, R. J. Warren, W. E.
James, L. A. Baker. 1994. A test of the scent-station
survey technique for bobcats. Journal of Wildlife
Management 58: 10-17.

Hatcher, R. T., and J. H. Shaw. 1981. A comparison of
three indices to furbearer populations. Wildlife Society
Bulletin 9: 153-156.

Knowlton, F. F., and W. M. Tzilkowski. 1981. Trends in
Bobcat visitations to scent-station survey lines in west-
ern United States, 1972-1978. Pages 8-12 in Bobcat
Research Conference Proceedings: Current research on
Biology and Management of Lynx rufus. National
Wildlife Federation Science and Technology Series 6.

Lindzey, F. G., S. K. Thompson, and J. I. Hodges. 1977.
Scent-station index of Black Bear abundance. Journal of
Wildlife Management 41: 151-153.

Linhart, S. B., and F. F. Knowlton. 1975. Determining
the relative abundance of Coyotes by scent-station lines.
Wildlife Society Bulletin 3: 119-124.

Martin, D. J., and D. B. Fagre. 1988. Field evaluation of
a synthetic Coyote attractant. Wildlife Society Bulletin
16: 390-396.

WOELFL AND WOELFL: COYOTE VISITATIONS TO SCENT STATIONS

203

Roughton, R. D. 1982. A synthetical alternative to fer-
mented egg as a canid attractant. Journal of Wildlife
Management 46: 230-234.

Roughton, R. D., and M. W. Sweeney. 1982. Refine-
ments in scent-station methodology for assessing trends
in carnivore populations. Journal of Wildlife
Management 46: 217-229.

Smith, W. P., D. L. Borden, and K. M. Enders. 1994.
Scent-station visits as an index to abundance of rac-
coons: an experimental manipulation. Journal of
Mammalogy 75: 637-647.

Stolzenburg, H. W., and V. W. Howard. 1989. Acti-
vation of liquid bait devices by Coyotes in southern New
Mexico. Wildlife Society Bulletin 17: 306-312.

Sumner, P. W., and E. P. Hill. 1980. Scent-stations as
indices of abundance in some furbearers of Alabama.
Proceedings Annual Conference Southeastern
Association of Fish and Wildlife Agencies 34: 572-583.

Turkowski, F. J., M. L. Popelka, and R. W. Bullard.
1983. Efficacy of odor lures and baits for Coyotes.
Wildlife Society Bulletin 11: 136-145.

Usher, R. G., and W. L. Strong. 1994. Ecological land
classification for the proposed Suffield National Wildlife
Area. Environment Canada Conservation and Protection,
35 pages.

Wood, J. E. 1959. Relative estimates of fox population
levels. Journal of Wildlife Management 23: 53-63.

Received 5 January 1996
Accepted 11 October 1996

Beaver, Castor canadensis, Home Range Size and Patterns of Use in
the Taiga of Southeastern Manitoba: I. Seasonal Variation

MICHELLE WHEATLEY

Taiga Biological Station, Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
Current address: Sahtu Renewable Resources Board, P.O. Box 490, Norman Wells, Northwest Territories XOE 0VO

Wheatley, Michelle. 1997. Beaver, Castor canadensis, home range size and patterns of use in the taiga of southeastern
Manitoba: I. Seasonal variation. Canadian Field-Naturalist 111(2): 204—210.

Seasonal variation in Beaver (Castor canadensis) home ranges in the taiga of southeastern Manitoba was monitored by
radiotelemetry and visual observations between 1986 and 1992. The results demonstrated that summer home ranges were
larger than fall home ranges (p < 0.001, n = 24). Both summer and fall home ranges were larger than winter home ranges
(p < 0.001, n= 19 and p< 0.001, n=22) which were restricted to the vicinity of the lodge. All Beavers had core areas in
both summer and fall (p < 0.05) but no core areas were present in winter. Core areas were larger in summer than in fall
(p < 0.001, n = 24). However, there was no significant difference in the proportion of the home range made up by the core
area in summer (27.2%) and fall (31.7%) (p > 0.05, n = 24), nor in the proportion of all activity which occurred in the core
areas in summer (73.7%) and fall (76.1%) (p > 0.05, n = 24). Summer and fall home range sizes were positively correlated
(p < 0.001) as were summer and fall core area sizes (p < 0.001).

Key Words: Beaver, Castor canadensis, home range, taiga, Manitoba, season.

An animal’s home range consists of the area that
animal uses during its normal activities of feeding,
breeding and raising young (Burt 1943). But animals
do not use all areas of their home range equally,
resulting in areas of concentrated use or core areas
(Springer 1982; Samuel et al. 1985).

Seasonal variation in activity and behaviour has
been well documented for Beavers, Castor canaden-
sis, in Manitoba (Green 1936), Alberta and the
Northwest Territories (Novakowski 1967), Quebec
(Potvin and Bovet 1975), Ohio (Brady and Svendsen
1981), California (Busher and Jenkins 1985) and
Minnesota (Buech 1995). Similarly the species of
tree eaten (Jenkins 1979) and the specific parts of
plants eaten (Aleksiuk 1970; Jenkins 1981) have
been found to vary seasonally.

Seasonal variation should be most noticeable in
regions with the greatest climatic variation among
seasons, and especially in regions where Beavers
must cope with extended periods of ice cover
(Novakowski 1967). In northern regions, Beaver kits
are born in late May or early June and much of the
summer is devoted to raising these offspring
(Banfield 1974). In the north, summer is also a time
for rapid growth of kits, yearlings and two-year old
Beavers (Aleksiuk and Cowan 1969b). In fall,
Beavers build food caches and repair or construct
lodges and often dams. Winter is spent almost entire-
ly in the lodge and under the ice, feeding on stored
food (Green 1936; Novakowski 1967; Aleksiuk and
Cowan 1969a; Potvin and Bovet 1975). Growth is
rare in winter, except in kits and sometimes year-
lings, and weight loss may often occur (Novakowski
1967; Aleksiuk and Cowan 1969a, 1969b; Aleksiuk

1970). Despite this acknowledged seasonal variation
in activities, only Davis (1984) has compared sea-
sonal movement patterns for Beavers, in South
Carolina, recording the maximum movement in fall
or winter and minimum movement in spring or sum-
mer. The taiga accounts for a large proportion of the
North American distribution of Beavers. However,
most studies of Beavers have been undertaken in
more southern regions of the Beavers’ distribution,
and no quantification of the seasonal variation in
Beaver home range size has been made for the taiga
regions. The objectives of this study were to quantify
the seasonal differences in Beaver home range size,
and to compare seasonal patterns of home range use
by Beavers in the taiga.

Study Area

This study was based at Taiga Biological Station
(TBS), 51°02'40"N, 95°20'40" W, 250 km northeast
of Winnipeg, Manitoba. The area was burnt during
the 1980 Wallace Lake fire. There are irregular
patches of unburned land in the area, some bordering
both rivers and lakes. The bogs, and also some
ridges where erosion has not been too extensive,
support a dense growth of Trembling Aspen
(Populus tremuloides), Alder (Alnus crispa), Black
Spruce (Picea mariana) and Jack Pine (Pinus
banksiana).

The climate of the study area is typical for mid-
continent boreal forest. Temperatures normally range
from -40°C in winter to +35°C in summer (Taiga
Biological Station Records). Freeze-up occurs in late
October or very early November and breakup in late
April or early May. Ice cover is continuous through-

204

1997

out the period except in areas of fast flowing water.
February ice thicknesses range from 50 cm to 100
cm depending upon thickness of the snow cover and
air temperatures.

Methods

Between September, 1986 and May, 1992, I live-
trapped 60 different Beavers using Hancock traps
(Hancock Trap Co., Custer, South Dakota) baited
with a piece of Trembling Aspen 30 cm long and a
scent consisting of ground up Beaver castor, anise
extract and glycerine (Aleksiuk 1968). I measured
total length, tail length, ear from notch, hind foot
length, and the weight of each animal, and deter-
mined the sex by palpating externally for the bacu-
lum. To facilitate visual identification of individuals,
I attached uniquely numbered and coloured ear tags
(Monel Number 3) to both ears (Miller 1964). I
installed radio-transmitters using two methods:
intraperitoneal implantation (42 transmitters, Austec
Electronics, Edmonton, Alberta); and tail collars (3
transmitters, Wildlife Materials Inc., Carbondale,
Illinois) (Wheatley 1989, 1994). Beavers were
released at the trap site 24 to 48 hours after surgery.
All work was approved by the University of
Manitoba Animal Care Committee and animals were
cared for in accordance with guidelines of the
Canadian Council on Animal Care (1980).

I tracked Beavers with an H-shaped antenna and a
24-channel receiver (Wildlife Materials Inc.,
Carbondale, Illinois). Those animals residing on the
river or lake were generally tracked and observed
from a canoe, while those living in ponds were
observed from a high ridge or similar good vantage
point. The telemetry equipment was used to locate
the Beaver and then, as much as possible, I contin-
ued with visual observations of activities and move-
ments. After dark, I also relied on telemetry to locate
animals, but additionally used sounds of chewing or
tree felling, or visual observation of water ripples to
determine activity. In winter, all movements were
monitored by radiotelemetry locations.

I recorded information about the location, time,
type of activity and movement during the period of
observation. On most nights, I visited each Beaver
activity area in turn, but the order of the visits varied
so that the same Beaver or group of Beavers was not
observed at the same time each night. Observation
periods in each activity area varied in length from

TABLE 1. Summary of number of observations per Beaver
in each season.

Season Mean SE Minimum Maximum N
Summer P24:8 ) »120:8 TW 2934 34
Fall 203.7 40.2 45 1134 27
Winter 1008.7. 102.1 471 1931 22

TABLE 2. Distribution of Beavers by habitat, season, sex and age class. Habitat designation is based on the location of the primary lodge for each Beaver. Ponds are areas of still

water, created and maintained at a relatively constant level by a Beaver dam; lakes are larger areas of non-flowing water, without the presence of a dam and rivers are areas of flow-

ing water, without the presence of any Beaver dams.

Total

River

Lake

Pond

WHEATLEY: BEAVER SEASONAL VARIATION

Fall Winter Summer Fall Winter Summer Fall Winter Summer Fall Winter

Summer

Age and sex

Adult male

Adult female

Yearling male

Yearling female
Juvenile male

Juvenile female

Kit male

Kit female

De

pA|

34

9

12

1] 13

6

14

Total

205

206

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 3. Home range size (ha) for Beavers for all animals in the study and the subset of the data used for pairwise tests
between the seasons are given. Median values followed by different letters (within each group) are significantly different

from each other (Wilcoxon Signed Rank Test p < 0.05).

Group Season Mean (ha) SE
All Summer 10.34 1.379
Fall 3.07 0.356
Winter 0.25 0.000
Summer - fall Summer 10.50 1.835
Fall 3:28 0.380
Summer - winter Summer 10.29 2.065
Winter 0.25 0.000
Fall - winter Fall 285 0.320
Winter 0.25 0.000

0.1 to 13 hours, depending upon weather conditions
and the order and number of the activity areas visit-
ed. Most observation periods lasted from 0.5 to 4
hours. In open water seasons, most observations
were made between 1800 and 2400 h. Earlier
research (Wheatley 1989) had shown that most
movement about the home range occurred in the first
part of the night, while more continuous feeding
bouts occurred during the second part of the night
(2400 to 0700 h). More visits to the lodge did occur
in the first part of the night (Wheatley 1989), so core
area estimates may emphasize the lodge area.
However, the lodge is used as a rest site for 10 to 12
hours per day, so is clearly going to be an important
part of the home range.

I transferred each Beaver’s movement data onto
maps with grid overlays equivalent to a 50 m by 50
m grid (0.25 ha) on the ground. For this I considered
1 minute as | observation because a Beaver can
swim approximately 50 m in one minute (personal
observation). This does not allow for true indepen-
dence of observations (Swihart and Slade 1985a;
1985b); however, Beavers move throughout their
home range too much to be able to determine home
range accurately using less frequent observations.

I defined a year as having three seasons: (1) sum-
mer, the period from breakup of ice cover to the
beginning of food cache construction; (2) fall, the
period from the beginning of food cache construc-
tion until freeze-up; and (3) winter, the ice-covered
period. I tallied the number of observations in each
grid cell and the total number of observations for
each Beaver for each season.

Samuel et al. (1985) described a method of deter-
mining core areas of home ranges based on the num-
ber of observations in each grid cell. Their method,
based on the Kolmogoroyv-Smirnov one-tailed good-
ness-of-fit procedure, was found to be unreliable and
the chi-square goodness-of-fit procedure was recom-
mended in its place (Samuel and Green 1988). To
determine the core area of each individual’s home
range, I compared the observed values in each cell

Median Minimum Maximum N
Tas 2:25 AD:T5 34
2.750 1.00 8.00 =o
0.250 0.25 0:25 22
GH Poi 2:25 42.75 24
2.750° 1.00 8.00 24
Lt DOF 225 42.75 19
0.2504 0.25 0.25 19
21 30F 1.00 6.75 22
0.250! 0.25 0.25 pe,

with the hypothesized uniform distribution for each
Beaver using the chi-square goodness-of-fit proce-
dure (Fowler and Cohen 1986). If this test showed
that a core area was present, I classed all cells with a
greater proportion of observations than uniform as
core areas (Samuel et al. 1985). I then transferred to
the map the outline of the home range and core areas
within the home range. I did this for each Beaver in
each season. While other methods of home range
and core area analysis are available, my examination
of several of these methods showed the grid method
to be most suitable for analysis of the Beaver home
ranges (Wheatley 1994).

I calculated home range size and core area size by
totalling the number of grid cells in each. I calculat-
ed the percent of home range that was included in
the core area for each Beaver, and refer to this as
core percent. I calculated core activity as the percent
of all observations that occurred within the core area.

All statistical analyses were made using non-para-
metric statistics because the data did not meet the
normality assumptions of parametric statistics. | used
the Wilcoxon Matched Pairs Test to test differences
in the variables between each of the seasons, and
Spearman Rank Correlation to determine correla-
tions between the same variables for summer and
fall, and between variables within each season
(Fowler and Cohen 1986). Although data for all
Beavers are presented, all statistical comparisons
between seasons are made only for those Beavers
where paired tests are possible in order to limit the

TABLE 4. Spearman Rank Correlation Coefficients between
summer and fall variables.

Variable r, (Summer-Fall) N P
Home Range 0.665 24 <0.001
Core Area 0.619 24 <0.005
Core Percent -0.016 24 NS
Core Activity 0.071 24 NS

1997

WHEATLEY: BEAVER SEASONAL VARIATION

207

TABLE 5. Core area size (ha) for Beavers for all animals and the subset of the data used for pairwise statistical analysis.
Median values followed by different letters are significantly different from each other (Wilcoxon Signed Rank Test,

p < 0.05)

Group Season Mean (ha) SE

All Summer 2.79 0.442
Fall 0.94 0.130

Paired data Summer 2.93 0.612
Fall 1.03 0.669

influence of other factors such as family status, sex,
age class and habitat, which are examined separately
(Wheatley 1997a, 1997b).

Results

Home range size was determined for 34 Beavers
in summer, 27 in fall and 22 in winter. A summary
of the number of observations on which these home
range estimates were based is given in Table 1.
Twenty-four of these Beavers were matched sum-
mer-fall pairs, 22 fall-winter pairs and 19 summer-
winter pairs. The breakdown of these Beavers by
sex, age and habitat is given in Table 2. Core areas
were documented for all Beavers in both summer
(chi-square test, p < 0.01) and fall (p < 0.05). In win-
ter, activity was restricted to one grid square and
therefore there were no core areas. Home ranges
usually followed shorelines and core areas only
occasionally included areas of open water, but these
were usually associated with food piles. Usually,
several unconnected areas constituted the core area
(multiple core areas). Main lodges were usually to
one side or one end of the home range, and rarely
towards the centre. Beavers used almost all of their
home range each night.

All Beavers had different summer and fall home
ranges based on differences in location, size, shape
and intensity of use. Fall home ranges generally con-
sisted of a portion of the summer home range but
some new areas were often included in fall.
Likewise, most Beavers used the same lodge year-
round, but five beavers (one family and one individ-
ual) built new lodges in the fall. Changes in home
range use coincided with changes in location of food
harvesting sites. In winter Beavers usually remained
outside the lodge only for a few minutes at a time,

Median Minimum Maximum N
2.125 Ob) [425 34
0.750 0.25 ZS Og |
2.0002 0.75 14.25 24
0.750° 0.50 pA hs) 24

but Beavers spent extended periods outside the lodge
in winter if the water level under the ice dropped,
creating air spaces. Even when these conditions
occurred, they never appeared to travel much beyond
the food pile.

Home ranges were larger in summer than in fall
(Table 3), and for the matched summer-fall pairs,
this difference was significant (Wilcoxon Signed
Rank Test, T = 1.0, p< 0.001, n= 24) and summer
and fall home ranges were significantly positively
correlated (Table 4). Winter home ranges were sig-
nificantly smaller than matched summer home
ranges (Tf = 0, p< 0.001, n=19) and fall home
ranges, (T= 0. pa0001,. 1 =22) (lable) Core
areas were also larger in summer than fall (Table 5)
and within the matched group the difference was sig-
nificant (T = 6.5, p< 0.001, n= 24) and also posi-
tively correlated (Table 4).

Core percent did not differ between summer and
fall (Table 6) (T = 89, p> 0.05, n= 24) and values
for the two seasons were not significantly correlated
(Table 4). Neither did core activity differ between
summer-and: fall (Table?) (1 =123 49 >0.05;
n = 24) and there was no significant correlation
(Table 4).

In summer, there was a significant positive corre-
lation between home range size and core area size
(Table 8). There was no significant correlation
between any of the other summer variables (Table
8). In fall, home range size and core area were also
significantly positively correlated as were core area
and core percent (Table 9). There was a significant
negative correlation between home range size and
core activity in the fall (Table 9). There was no sig-
nificant correlation between any of the other fall
variables (Table 9).

TABLE 6. Percent of home range included in the core area (core percent) for all animals and the subset of data used for
paired statistical analysis. Median values in the paired group are not significantly different from each other (Wilcoxon

Signed Rank Test, p > 0.05).

Group Season Mean (%) SE

All Summer Oe 0.963
Fall 31.68 2.622

Paired data Summer 27.62 1.095
Fall 33.41 2.748

Median Minimum Maximum N
26.201 16.36 37.50 34
30.000 16.67 75.00 oiafl
26.794 16.36 37.04 24
30.385 18.18 75.00 24

208

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 7. Percent of all activity occurring in the core area (core activity) for all Beavers and for the subset of the data used
for the pairwise statistical test. Median values in the paired group are not significantly different from each other (Wilcoxon

Matched Pairs Test, p > 0.05).

Group Season Mean (%) SE
All Summer 73.68 1.159
Fall 76.14 1.874
Paired data Summer 73.69 1.450
Fall 75.64 2.085
Discussion

Seasonal variations in home range size have been
described for many animals (including Mountain
Sheep (Ovis spp.) (Geist 1971), Muskrat (Danell
1978; MacArthur 1978; Proulx and Gilbert 1983),
Coyotes (Canis latrans) (Springer 1982) and
Porcupines (Erethizon dorsatum) (Craig and Keller
1986)) and are consistent with seasonal differences
in activities and food availability. The limited previ-
ous studies of Beaver home range size have focussed
primarily on the summer. Busher (1975) reported
home ranges 200 m to 800 m long for Beavers in
California and Gillespie (1977) found home range
sizes of 0.4 to 22 ha for Beavers in southern Ontario.
Davis (1984) measured the nightly distance between
extreme points in South Carolina as 84 m to 1863 m
for the whole year. While in general slightly lower
than the summer home range sizes of 2.25 to 42.75
ha in this present study, these estimates are none the
less comparable.

Only Davis (1984) has measured seasonal differ-
ences in patterns of home range use. His work
showed that total daily movement was greatest in fall
or winter and least in spring and summer, while the
distances between extremes showed maxima in
spring or fall and minima in summer. Data in the
present study showed that taiga Beaver home ranges
also vary seasonally in size, but the results differ
sharply from Davis’ results in that home ranges were
largest in summer, smallest in winter and intermedi-
ate in fall. In South Carolina, winter ice cover was
not present, and since nights are longer in winter
than in summer, Beavers have more time to travel
farther afield in search of food or for exploration.
Beavers in South Carolina also do not have to build
food caches in the fall for winter use, allowing
greater freedom of movement in the fall. In the taiga,

TABLE 8. Spearman Rank Correlation Coefficients between
summer variables, n = 34.

Home Range Core Area Core Percent

Core Area 0.939*** 1.00
Core Percent ~0.252"5 0.036NS 1.00
Core Activity 0.108NS 0.250NS 0.228NS5

*** = p < 0.001, NS = not significant.

Median Minimum Maximum N
F527 60.89 84.40 34
7720 59090 95.40 pe |
75.365 60.89 84.40 24
76.630 35250 95.40 24

Beavers must build a food cache in fall and rely on
this cache for up to six months of ice cover. This is
an important difference in the obligate seasonal
activities between the two regions. Similar to Davis’
findings, Busher and Jenkins (1985) found that much
more time was spent outside the rest site (lodge) in
fall than in summer. However, time outside the rest
site does not necessarily imply a larger home range.
Jenkins (1981) reported a seasonal change in feed-
ing site, possibly as a result of a seasonal preference
for different species of trees, or because of seasonal
variation in site accessibility. Similar seasonal
changes in food choice occurred in my study area
(Wheatley 1994). But in contrast to other studies
(Green 1936; Tevis 1950; Townsend 1953; Brenner
1962; Northcott 1971; Gillespie 1977; Svendsen
1980; Jenkins 1981; Belovsky 1984), the Beavers in
my study area did not rely heavily on aquatic vegeta-
tion (Wheatley 1994). Fall home ranges usually con-
sisted of a portion of the summer home range, but in
some cases contained new area not included in the
summer range. Fall food harvesting sites were usual-
ly situated within summer home ranges in areas close
to the lodge, but were not used as sources of food in
the summer. This could reflect a seasonal difference
in food preference, a method of reducing energy
expenditure in the fall by reducing the distance that
food must be transported to the food cache, or a
means of extending the time period over which a par-
ticular lodge site is usable. MacArthur (1978) pro-
posed a similar explanation for the reduction in size
of Muskrat foraging areas as winter advanced.
Semyonoff (1953) (working in the Russian taiga),
Lancia (1979), and Lancia et al. (1982) (working in
Massachusetts) reported more winter movement than
I found in Beavers in the Manitoba taiga. However,
Semyonoff’s Beavers used sushinetz (air spaces

TABLE 9. Spearman Rank Correlation Coefficients between
fall variables, n = 27.

Home Range Core Area Core Percent

Core Area 0.796*** 1.00
Core Percent -0.108NS 0.468** 1.00
Core Activity -0.482** -0.2185 0.320NS

1997

under the ice) extensively and Lancia’s Beavers were
active fairly frequently above the ice. Smith and
Peterson (1991) found behaviour similar to
Semyonoff when water levels under the ice dropped
and speculated that there was a thermal advantage to
staying in the air spaces under the ice. Beavers in the
present study did not move extensively using the
sushinetz, when they occurred. This is most likely
due to the hindrance imposed by the irregular shape
of shorelines and frequent location of lodges on
islands.

Core areas were larger in summer than fall, con-
sistent with the seasonal differences in home range
size but core activity and the percent of the home
range included in core areas did not differ between
summer and fall. No other studies have quantified
core area size, but Davis (1984) reported areas of
concentrated use when food was clumped, but not
when food was regularly distributed. Knowledge
about core areas and the pattern of use of home
ranges (as reflected by data for core activity and the
core percent) are important in understanding how a
species uses its environment. No other studies have
examined these variables for Beavers, so a compari-
son to some other species is necessitated. Beavers in
the taiga appear to have some inherent pattern of use
of their home range such that about 75% of their
activity takes place in about 27% of their home
range. This value for core activity is similar to
Springer’s (1982) value of 82.9% Coyotes (Canis
latrans) and Samuel et al.’s (1985) value of 76% for
Black Bears (Ursus americanus). For core percent,
the percent of the home range included in the core
area, Springer’s (1982) value of 6.9% for Coyotes is
much lower than my value for Beavers, while
Samuel et al.’s (1985) value of 34% for Black Bears
is much more similar to my value for Beavers. As
with the Beavers in this study, both the Black Bears
and Coyotes had multiple core areas (Springer 1982;
Samuel et al. 1985). For the Beavers, these patterns
of home range use may be related to food availabili-
ty in the area, to habitat types, or to other factors that
we cannot detect. The correlation between summer
and fall home range size may likewise indicate that
the home range size reflects the habitat, or possibly
quality of the habitat, in which the lodge is situated.

There are many other factors which also affect
home range size. These include the age and the sex
of the Beaver and whether they are part of a family
group (Wheatley 1997a), the habitat in which the
Beavers live (Wheatley 1997b), and the shape and
size of the body of water on which they live
(Gillespie 1975; Wheatley 1994). Statistical analysis
of the interactions between these different factors is
difficult to achieve due to the large numbers of ani-
mals that would be necessary. But by using the
matched pairs statistical procedures, it is possible
make the comparison between seasons, without hav-
ing to include consideration of these other factors.

WHEATLEY: BEAVER SEASONAL VARIATION

209

In this study I have shown that, in the taiga,
Beavers have home ranges that are largest in sum-
mer, smallest in winter and intermediate in fall.
While summer values of home range size are compa-
rable to data from other regions, there are major dif-
ferences between my findings for seasonal variation
in home range size for taiga Beaver and the findings
for Beavers in more temperate regions. The correla-
tions between summer and fall home range sizes and
core sizes, and the similarity in core percent and core
activity between the two seasons indicates that there
may be some inherent pattern of home range use
possibly related to food harvest sites or food avail-
ability. The differences from other studies reflect the
differences in seasonal activities necessitated by the
extended winter period and preparations for this
period. These results must be considered to reflect
Beaver behaviour in the mid-continental taiga, and
not necessarily other regions of North America.

Acknowledgments

This study was funded by grants from the Taiga
Biological Station Research Trust; Wildlife Branch,
Manitoba Department of Natural Resources; Sigma
Xi; Manitoba Chapter, The Wildlife Society and the
Manitoba Wildlife Federation and a University of
Manitoba Graduate Fellowship. W. O. Pruitt, Jr.,
provided advice, support and assistance throughout
this study and read many earlier drafts of this paper.
Numerous field assistants made this study possible.
S. C. Jay, S. H. Jenkins, R. Riewe and K. Stewart
and two anonymous reviewers improved earlier ver-
sions of this manuscript.

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Received 29 January 1996
Accepted 24 October 1996

Beaver, Castor canadensis, Home Range Size and Patterns of Use in
the Taiga of Southeastern Manitoba: II. Sex, Age, and Family Status

MICHELLE WHEATLEY

Taiga Biological Station, Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
‘Current address: Sahtu Renewable Resources Board, P.O. Box 490, Norman Wells, Northwest Territories XOE 0VO

Wheatley, Michelle. 1997. Beaver, Castor canadensis, home range size and patterns of use in the taiga of southeastern
Manitoba: II. Sex, age, and family status. Canadian Field-Naturalist 111(2): 211-216.

The differences in home range size and patterns of use relating to age, sex and family status for 25 Beavers living in family
groups, and 9 non-family Beavers in summer and 23 family and 4 non-family Beavers in fall, were examined in southeast-
ern Manitoba. Home ranges were larger for non-family Beavers (x =17.56 + 3.8 ha, n = 9) than for those living in family (x
=7.74 + 0.87 ha, n = 25) groups in summer (T = 233.5, p < 0.005) but not in fall (T = 78.5, p > 0.1). Core areas were also
larger for non-family animals in summer (T = 211.5, p < 0.05) but not in fall (T = 63, p > 0.1). The percent of home range
area included in the core area and the percent of activity within the core area did not differ significantly between family
and non-family Beavers in either season (p > 0.1). Within families, adult females had smaller home ranges than other fami-
ly members in summer (p < 0.05) but not in fall (p > 0.1). Home ranges did not differ significantly in size among all other
family members in both summer and fall (p > 0.05). Core area size, percent of home range area included in the core area

and percent of activity within the core area did not differ among the sex and age classes within family groups (p > 0.1).

Key Words: Beaver, Castor canadensis, home range, boreal forest, Manitoba.

Beavers are generally considered to be a monoga-
mous species (Jenkins and Busher 1979) having a
prolonged term of parental care. Beaver families
spend much of the summer raising kits that were
born in late May or early June (Banfield 1974). In
northern regions, summer is also the primary time of
year for growth and fat deposition (Aleksiuk and
Cowan 1969a, 1969b). In fall, Beavers build food
caches and repair or construct lodges and dams.
Winter is spent almost entirely in the lodge and
under the ice, feeding on the stored food
(Novakowski 1967). Growth is rare in winter, except
in kits and yearlings, and weight loss may occur
(Novakowski 1967; Aleksiuk and Cowan 1969a,
1969b). Parental care usually continues through the
second year of life, but yearlings also assist in the
care of the kits. Subadults generally leave their natal
lodge at two years of age, but occasionally leave at
their first birthday or stay beyond their second birth-
day (Jenkins and Busher 1979).

Hodgdon and Larson (1973) and Busher and
Jenkins (1985) reported differences in behavioural
patterns between adult male and female Beavers,
while Buech (1995) cited differences in the time spent
on different types of activities by adult males and
females, and Townsend (1953) reported differences in
movement patterns between adult males and females.
In addition, Busher and Jenkins (1985) noted that
yearling female behaviour was more like that of adult
females, while yearling male behaviour was more like
that of kits. Brady and Svendsen (1981) found sex
and age class differences in the interactions between
animals. Davis (1984) assumed that no such differ-
ences in movement or behaviour patterns existed. No

authors appear to have examined variations in home
range size among Beavers in different sex and age
classes. The objective of this study was to examine
patterns of use of home range by Beavers according
to sex and age classes both for Beavers living in fami-
ly groups and for non-family Beavers.

Methods

The study area, methods of capturing, marking
and tracking Beavers, and data manipulation are
described in Wheatley (1997). I used Buckley and
Libby’s (1955) classification system to separate the
different age classes. “Kits” were animals less than
12 months of age, “yearlings” referred to Beavers
in their second year of life and “juveniles”, to those
in their third year of life. “Adult” referred to any
Beaver over 36 months. Beavers were classed as
being part of a “family” if they were living in a
lodge where there was a breeding pair, and there-
fore kits. All “non-family” Beavers were either
solitary, or if living with another beaver, were not
breeding or resident in a lodge with a breeding pair
and kits.

Since earlier analysis had shown significant sea-
sonal differences in home range size for Beavers
(Wheatley 1997), all analysis of differences
between family and non-family Beavers and
between sex and age classes of family Beavers was
carried out within each season (summer and fall). I
calculated home range size and core area size by
totalling the number of grid cells in each (Wheatley
1997). I calculated the percent of home range that
was included in the core area for each Beaver, and
refer to this as core percent. I calculated core activi-

a |

242 THE CANADIAN FIELD-NATURALIST Vol. 111

TABLE |. Summer home range size (ha) for Beavers comparing all family and non-family animals (Mann Whitney U test),
different age classes and sexes within family groups (Kruskal-Wallis Analysis of Variance) and males and females in the
non-family group (Mann Whitney U test). Median values followed by different letters are significantly different from each
other (p < 0.05) (within each group — all Beavers, family only and non-family only). All other comparisons within each
group are not significantly different (p > 0.05).

Family Age Class and

Status Sex Mean (ha) SE Median Minimum Maximum N

Family All 7.74 0.869 6.750 2.25 20.50 2

Non-family All 17.56 3.804 14.000° 4.75 42.75 9

Family Adult Male 10.89 5.789 LISS I25 20.50 *
Adult Female 4.39 0.946 4.0054 2.23) 9.50 7
Yearling Male S25 2.449 T25@ Bete) LS 5
Yearling Female 7.69 1.804 7.000 4.25 12.50 4
Juvenile Female Vide 0.750 7.250 6.50 8.00 2

Non-family Male 13.33 3.282 13.875 4.75 20a 4
Female 20.95 6.258 14.500 8.25 42.75 5

ty as the percent of all observations that occurred
within the core area.

The data were analysed by sex and age class and
family status. I tested for differences between fami-
ly and non-family Beavers using the Mann-
Whitney U Test (Fowler and Cohen 1986). Within
known family Beavers I tested for differences due
to age and sex using the Kruskal-Wallis One Way
Analysis of Variance, with subsequent multiple
comparisons if the Kruskal-Wallis Test Statistic
indicated a significant difference among the groups
(Conover 1980). The group of non-family Beavers
did not have sufficient numbers in the different sex
and age classes to allow for statistical comparison
when separated into these classes, so data are sum-
marised by sex for the non-family group.

Results
I captured and ear tagged 60 different Beavers,
and installed 42 radiotransmitters. I was able to

identify home ranges of 25 animals living in family
groups, and nine non-family animals in summer
and 23 family and four non-family Beavers in fall.
Non-family Beavers included five juvenile females,
one adult male and three yearling males in summer
and three juvenile females and one adult male in
fall.

While all Beavers living within the same family
group customarily used the same general area for
their home range, the different family members did
not usually associate together at any particular
feeding site or other location. When kits were pre-
sent in the lodge, one of the family members was
usually present in or near the lodge at all times.
During the observation periods, I observed all fami-
ly members taking food into the lodge, apparently
for the kits. In fall, kits were active outside the
lodge, but remained in the vicinity of the lodge.

Summer home ranges for Beavers living in fami-
lies were significantly smaller than those of non-

TABLE 2. Fall home range size (ha) for Beavers comparing all family and non-family animals (Mann Whitney U test), dif-
ferent age classes and sexes within family groups (Kruskal-Wallis Analysis of Variance) and males and females in the non-
family group (no statistical test, sample size too small). Median values are not significantly different from each other
(p > 0.05) (within each group — all Beavers and family only).

Family Age Class and

Status Sex Mean (ha) SE Median Minimum Maximum N

Family All 2.87 0.378 2.500# 1.00 8.00 23

Non-family All 4.25 0.919 3.875 pee fe) 6.50 4

Family Adult Male 3.96 0.935 2ISOF 1.00 8.00 Es
Adult Female 1.90 0.341 2.000° 1.00 PM 5
Yearling Male 33 0.260 Dvlney 2.50 fs) 4
Yearling Female BM es 1.010 1.500° 1.00 4.25 3
Juvenile Female Jie) 0.000 Dae IU: Salo aud) l
Kits 1.42 0.083 1.500° La 1.50 |

Non-family Male veh 0.000 2.750 Oe fs OE be l
Female 4.75 1.090 5.000 ale 6.50 3

1997 WHEATLEY: BEAVER SEX, AGE AND FAMILY STATUS DANS

TABLE 3. Summer core area size (ha) for Beavers comparing all family and non-family animals (Mann Whitney U test),
different age classes and sexes within family groups (Kruskal-Wallis Analysis of Variance) and males and females in the
non-family group (Mann Whitney U test). Median values followed by different letters are significantly different from each
other (p < 0.05) (within each group — all Beavers, family only and non-family only). All other comparisons within each
group are not significantly different (p > 0.05).

Family Age Class and

Status Sex Mean (ha) SE Median Minimum Maximum N

Family All 2.10 0.204 A750? 0.75 4.50 25

Non-family All 4.72 1.440 3.000° 1.00 135225 9

Family Adult Male 2.82 0.422 2.500 Ld 4.50 fe
Adult Female 1:32 0.335 1.250 0.75 3.25 a,
Yearling Male 2.10 232 2.000 1.50 £109 fe 5
Yearling Female Zil3 0.924 2.000 1.25 5a +
Juvenile Female 2.25 0.750 2.250 1.50 3.00 2

Non-family Male pAE 0.700 3.000 1.00 4.00 4
Female 6.30 2.402 3.000 2.00 14.25 5

TABLE 4. Fall core area size (ha) for Beavers comparing all family and non-family animals (Mann Whitney U test), differ-
ent age classes and sexes within family groups (Kruskal-Wallis Analysis of Variance) and males and females in the non-
family group (no statistical test, sample size too small). Median values are not significantly different from each other
(p > 0.05) (within each group — all Beavers and family only).

Family Age Class and

Status Sex Mean (ha) SE Median Minimum Maximum N

Family All 0.90 0.133 0.750 0.25 BAD 23

Non-family All 1.19 0.472 0.875 0.50 2350) 4

Family Adult Male 132 0.317 1.250 0.50 pa i Ui
Adult Female 0.55 0.050 0.500 0.50 OFS 5
Yearling Male 0.88 0.072 0.875 0.75 1.00 4
Yearling Female 0.83 0.220 0.750 0.50 PAs) 3
Juvenile Female 2.00 0.000 2.000 2.00 2.00 1
Kits 0.25 0.000 0.250 0.25 O.25 3

Non-family Male 0.50 0.000 0.500 0.50 0.50 1
Female 1.42 0.583 1.250 0.50 2.50 3

TABLE 5. Percent of home range area included in the core area (core percent) in summer for Beavers, comparing all family
and non-family animals (Mann Whitney U test), different age classes and sexes within family groups (Kruskal-Wallis
Analysis of Variance) and males and females in the non-family group (Mann Whitney U test). Median values are not sig-
nificantly different from each other (p > 0.05) (within each group — all Beavers, family only and non-family only).

Family Age Class and

Status Sex Mean (%) SE Median Minimum Maximum N

Family All 28.22 0.984 DiczAS 19.51 37.50 25

Non-family All 24.46 BOS 24.000 16.36 35.51 9

Family Adult Male 27.78 2.073 28.125 19.51 33.33 if
Adult Female 29.63 1.340 30.000 25.00 34.21 1
Yearling Male 26.24 2.845 24.138 2052 37.04 5
Yearling Female 27.95 1.116 27.749 26.00 30.30 4
Juvenile Female 30.29 TAZ 30.288 23.08 5150 2

Non-family Male ZEOZ 2.699 19.562 16.36 28.57 4
Female Zhe S11 24.242 18.97 a5 a

214 THE CANADIAN FIELD-NATURALIST Vol 141

TABLE 6. Percent of home range area included in the core area (core percent) in fall for Beavers, comparing all family and
non-family animals (Mann Whitney U test), different age classes and sexes within family groups (Kruskal-Wallis Analysis
of Variance) and males and females in the non-family group (no statistical test, sample size too small). Median values are
not significantly different from each other (p > 0.05) (within each group — all Beavers and family only).

Family Age Class and

Status Sex Mean (%) SE Median Minimum Maximum N

Family All 32.85 2.929 30.000 16.67 75.00 23

Non-family All 24.96 4.780 21.591 18.18 38.46 4

Family Adult Male 35.45 4.702 34.375 18.18 50.00 Z
Adult Female 33.14 6.110 37.500 18.18 50.00 5
Yearling Male 28.11 1.366 28.333 25.00 30.77 4
Yearling Female 45.92 14.586 33333 29.41 75.00 a
Juvenile Female 38.10 0.000 38.100 38.10 38.10 ]
Kits 17.78 1a ia 16.667 16.67 20.00 3

Non-family Male 18.18 0.000 18.182 18.18 18.18 1
Female pH PA 5.958 25.000 18.18 25.00 3

TABLE 7. Percent of activity occurring in the core area (core activity) in summer for Beavers, comparing all family and
non-family animals (Mann Whitney U test), different age classes and sexes within family groups (Kruskal-Wallis Analysis
of Variance) and males and females in the non-family group (Mann Whitney U test). Median values are not significantly
different from each other (p > 0.05) (within each group — all Beavers, family only and non-family only).

Family Age Class and

Status Sex Mean (%) SE Median Minimum Maximum N

Family All 74.68 1.411 77.200 60.89 84.40 25

Non-family All 70.92 1.741 68.000 63.90 78.40 9

Family Adult Male T4A2 2.877 77.800 64.50 81.40 7
Adult Female 69.84 2GST 72.800 60.89 79.00 i;
Yearling Male 76.70 312 78.500 66.30 84.40 5
Yearling Female 78.95 0.821 78.905 77.20 80.79 +
Juvenile Female 79.96 4.125 79.955 75.83 84.08 2

Non-family Male 70.64 3.146 71.150 63.90 76.34 4
Female Tis 2.224 68.000 67.33 78.40 5

TABLE 8. Percent of activity occurring in the core area (core activity) in fall for Beavers, comparing all family and non-
family animals (Mann Whitney U test), different age classes and sexes within family groups (Kruskal-Wallis Analysis of
Variance) and males and females in the non-family group (no statistical test, sample size too small). Median values are not
significantly different from each other (p > 0.05) (within each group — all Beavers and family only).

Family Age Class and

Status Sex Mean (%) SE Median Minimum Maximum N

Family All 76.97 2.098 78.500 53.50 95.40 24

Non-family All yg Ne 5 i We} 71.080 65.50 77.80 4

Family Adult Male 78.14 1.668 75.600 73.10 84.90 7
Adult Female 72.80 6.322 64.400 62.50 95.20 5
Yearling Male 75.38 3.056 77.050 66.60 80.80 4
Yearling Female 79.43 13.082 89.400 53.50 95.40 3
Juvenile Female 79.20 0.000 79.200 79.20 79.20 1
Kits 80.07 L517 78.600 78.50 83.10 3

Non-family Male 76.36 0.000 76.360 76.36 76.36 l
Female 69.70 4.051 65.800 65.50 77.80 3

1997

family animals (Mann-Whitney U test, T = 233.5,
p < 0.005) (Table 1), but in fall there was no signifi-
cant difference in size of family and non-family
home ranges (T = 78.5, p > 0.1) (Table 2). Likewise,
in summer, core areas of family Beavers were sig-
nificantly smaller than those of non-family animals
(T = 211.5, p < 0.05) (Table 3) , but in fall the core
area sizes did not differ between family and non-
family animals (T = 63, p>0.1) (Table 4). Core
percent did not differ between family and non-fami-
ly Beavers in summer (T = 115.5, p > 0.1) (Table 5)
nor in fall (T = 40.5, p > 0.1) (Table 6). Neither did
core activity differ between family and non-family
animals in summer (T = 120, p>0.1) (Table 7) or
fat 1. — 35, p > 0.1) (Table 8).

In summer there was a significant difference in
home range size among the sex and age classes with-
in the family group (Kruskal-Wallis ANOVA,
H = 9.559, df = 4, p < 0.05) (Table 1). Multiple pair-
wise comparisons showed adult female home ranges
to be significantly smaller than those of adult males
(Dunn’s Method, p < 0.05), but no other significant
differences among the adults and yearling males,
yearling females and juvenile females (Table 1). In
fall there was no significant difference in home
range size among the sex and age classes within the
family group (H = 8.618, df=5, p>0.1) (Table 2).
Core area size was similar for all sex and age classes
in summer (H = 9.388, df =4, p > 0.05) (Table 3).
Although the Kruskal-Wallis ANOVA showed a sig-
nificant difference in core area size in fall among the
sex and age classes (H = 13.445, df =5, p < 0.05),
the subsequent Dunn’s Multiple Comparison did not
reveal any significant differences in the pairwise
tests of the different sex and age classes (Table 4),
possibly due to the small sample sizes. Core percent
did not differ among the sex and age classes in both
summer (H = 5.824, df =4, p>0.1) (Table 5) and
fall (H = 2.822, df =5, p>0.1) (Table 6) and core
activity did not differ among the sex and age classes
in either summer (H = 2.27, df =4, p >0.1) (Table
7) or fall (H = 7.803, df =5, p> 0.1) (Table 8) with-
in the family group.

Discussion

The family Beavers in the Wallace Lake study
area demonstrated smaller home ranges and core
areas than non-family Beavers in summer, but there
were no differences between the groups in the fall.
The primary difference between Beavers living in
family groups and those not living in families is the
presence of kits in the family groups. Kits require
nursing and the provision of solid food from outside
the lodge, especially in the summer. This ties mem-
bers of family groups to the lodge, and may restrict
the degree to which they may roam freely from the
lodge during the night. Non-family Beavers do not
have this requirement of returning to the lodge dur-
ing their nightly activities, and are therefore freer to

WHEATLEY: BEAVER SEX, AGE AND FAMILY STATUS

215

travel greater distances from the lodge during the
night. In the fall, the obligate activities of family and
non- family Beavers are more similar resulting in
more similar home range sizes. Alternatively, or
simultaneously, it may be that the better habitats are
occupied by the family groups, and therefore the
individual Beavers are left to use poorer habitat,
which requires more movement in order to find food.
Unattached Beavers may also be travelling further
afield in search of a mate and suitable habitat for
establishing a lodge and raising a family.

Within family groups, the adult female’s home
range was smaller than that of other family members
in the summer, but otherwise there were no significant
differences in home range size or patterns of use of
the home range among the members of family groups.
While I observed all the members of family groups
apparently provisioning the lodge with food for the
kits, the additional need for nursing, which only the
adult female is capable, best accounts for the smaller
adult female home ranges in summer. In contrast, in a
comparison between adult male and adult female
Beavers, Buech (1995) cited the adult male as spend-
ing more time in the lodge than the adult female in the
summer, but also cited the adult male as spending
more time travelling, while the adult female spent
more time feeding. As in the present study, Buech
(1995) found a greater similarity in behaviours
between adult males and females in fall. Other studies
have found adult males to have a great deal of sum-
mer movement (Tevis 1950; Townsend 1953). The
role attributed to the adult male Beaver in raising the
kits varies greatly, from little or no participation
(Seton 1909; Bradt 1938; Busher 1975) to some or
full participation (Brady and Svendsen 1981; Busher
and Jenkins 1985; Buech 1995).

Most of the studies of Beaver behaviour have
focussed on the adult members of family groups, so
there is very little in the literature on the role and
behaviours of juveniles, yearlings and kits. However,
as I found for members of family groups, Tevis
(1950) and Brady and Svendsen (1981) found that
Beavers were independent of each other during the
night. Busher and Jenkins (1985) showed that the
behaviour of yearling females was more similar to
that of adult females while that of yearling males was
more similar to kits. This distinction between sexes
among yearlings was not evident in the present study,
but similarity of home range sizes and behaviours are
not necessarily synonymous. As in the present study,
Busher and Jenkins (1985) found that all family
members except the kits had similar behaviour pat-
terns in the fall. They also report that much more
time was spent outside the rest site in fall (>80% of
active time) than in summer (Busher and Jenkins
1985). The kits in this present study (fall data only),
while not differing significantly from other family
members in home range parameters, did tend to
remain close to the lodge. As the smallest family

216

members, these kits are most susceptible to predation,
and therefore likely remain closer to the lodge.

In summary, the greatest difference in Beaver
home ranges is between those Beavers living in fam-
ily groups, and those Beavers not residing in these
family groups. These differences may reflect differ-
ences in obligate activities, habitat quality or behav-
iour. Within family groups, there is little difference
in home range between the family members except
for the smaller home range for adult females in sum-
mer. This probably relates to the different role that
the adult female must play in raising the young.

Acknowledgments

This study was funded by grants from the Taiga
Biological Station Research Trust; Wildlife Branch,
Manitoba Department of Natural Resources; Sigma
Xi; Manitoba Chapter, The Wildlife Society and the
Manitoba Wildlife Federation and a University of
Manitoba Fellowship. W. O. Pruitt, Jr., provided
advice, support and assistance throughout this study
and read many earlier versions of this paper.
Numerous field assistants made this study possible.
S. C. Jay, S. H. Jenkins, R. Riewe and K. Stewart
and two anonymous reviewers improved earlier
drafts of this manuscript.

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Aleksiuk, M., and I. Mc T. Cowan. 1969b. The winter
metabolic depression in arctic beavers (Castor canaden-
sis Kuhl) with comparisons to California beavers.
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Banfield, A. W. F. 1974. The mammals of Canada.
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Bradt, G. W. 1938. A study of beaver colonies in
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Davis, J. R. 1984. Movement and Behavior Patterns of
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Wheatley, M. 1997. Beaver, Castor canadensis, home
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Naturalist 111: 204-210.

Received 29 January 1996
Accepted 23 October 1996

Beaver, Castor canadensis, Home Range Size and Patterns of Use in
the Taiga of Southeastern Manitoba: HI. Habitat Variation

MICHELLE WHEATLEY

Taiga Biological Station, Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
Current address: Sahtu Renewable Resources Board, P.O. Box 490, Norman Wells, Northwest Territories XOE OVO

Wheatley, Michelle. 1997. Beaver, Castor canadensis, home range size and patterns of use in the taiga of southeastern
Manitoba: III. Habitat variation. Canadian Field-Naturalist 111(2): 217-222.

Differences in home range size among Beavers living in ponds, lakes and rivers were examined in southeastern Manitoba.
Beavers living in river habitat had larger summer home ranges than those living in ponds (p < 0.05), but there were no dif-
ferences between river and lake or pond and lake dwelling beavers (p >0.05). However, when non-family Beavers were
excluded from the analysis, there was no significant difference in home range size among the habitats (p > 0.05). Summer
core areas were significantly smaller for pond Beavers than for river animals (p < 0.05) when non-family Beavers were
included, but not when they were excluded from the analysis (p > 0.05), and there were no differences between pond and
lake or river and lake animals (p > 0.05). In fall there were no differences in home range or core area size among the habi-
tats (p > 0.05). The percent of the home range included in the core area did not differ among the habitats in summer or fall
(p > 0.05). The percent of activity occurring in the core area did not differ among the habitats in summer, but in fall core
activity was greater for lake than river Beavers regardless of whether non-family Beavers were included in the analysis

(p < 0.05), but there were no differences between pond and river or pond and lake (p > 0.05).

Key Words: Beaver, Castor canadensis, habitat, boreal forest, home range, Manitoba.

Little is known about the home range size of
Beavers, and especially how home range size relates
to habitat. Much early work on Beavers relied on
examining evidence of Beaver activity: lodges;
dams; food piles; and cutting areas (Morgan 1868 as
cited in Morgan 1986; Warren 1927). Only with the
advent of radio-telemetry has the study of the actual
movements of this nocturnal, semi-aquatic mammal
been possible.

Davis (1984) found minima and maxima of move-
ment to be different for stream and lake Beavers in
South Carolina. Busher (1975) quantified daily
movements of Beaver living in a stream habitat.
Neither author delineated actual home range areas.
Likewise, Brady and Svendsen (1981), Bergerud and
Miller (1977) and Green (1936) all described Beaver
activities without quantifying movements or home
range size. Gillespie (1975) described smaller home
range sizes for Beavers living in a pond as compared
to lake habitat.

The traditional view of the Beaver is an animal
living in a pond created by a Beaver dam which
maintains a constant water level. In reality, many
Beavers also live on free-flowing rivers and on lake
shores. Beavers living in lake and river habitats are
potentially subject to greater fluctuations in water
level than those living in pond habitats, while those
living in pond habitats have a requirement of greater
effort on maintenance activity than river or lake
Beavers. Therefore, the expectation was that there
would be differences in the home range size and pat-
terns of use among Beavers living in these three dif-

ferent habitats. In this paper I examine the differ-
ences in home range size and other home range char-
acteristics demonstrated by Beavers living in pond,
lake and river habitats within the boreal forest of
eastern Manitoba.

Methods

The study area, methods of capturing, marking
and tracking Beavers, and data manipulation are
described in Wheatley (1997a). I classified Beavers
as pond, lake or river Beavers based on the location
of their primary residence, the lodge or bank burrow
used most often, during each season. I defined ponds
as areas of still water, created and maintained at a
relatively constant level by a Beaver dam; lakes as
larger areas of non-flowing water, without a dam;
and rivers as areas of flowing water, without the
presence of any Beaver dams.

I calculated home range size and core area size by
totalling the number of grid cells in each (Wheatley
1997a). I calculated the percent of home range that
was included in the core area for each Beaver, and
refer to this as core percent. I calculated core activity
as the percent of all observations that occurred with-
in the core area. Family Beavers are those residing in
a group with a breeding pair and kits, while non-
family animals are solitary animals or if paired, not
living in a lodge with a breeding pair and kits.

I analysed the data for all Beavers by habitat type.
Since earlier analysis showed that season affected
home range size and core area size (Wheatley
1997a), I analysed data only within each season.

217

218

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE |. Summer home range size (ha) for Beavers living in pond, lake and river habitats for all Beavers and for family
Beavers only. Median values followed by different letters are significantly different from each other (p < 0.05) (within each
group — all Beavers and family only). All other comparisons within each group are not significantly different (p > 0.05).

Family Status Habitat Mean (ha) SE

All Beavers Pond 6.54 0.642
Lake 77 1.663
River 16.94 4.004

Family only Pond 6.19 0.513
Lake 8.78 1.883
River 10.06 2.923

Earlier analysis also showed that family status
affected home range size (Wheatley 1997b), so I
analysed habitat differences for all Beavers regard-
less of family status, and then separately for just
family Beavers. I tested for differences between the
habitats using the Kruskal-Wallis One Way Analysis
of Variance, with subsequent multiple comparisons
(Dunn’s Method) if the Kruskal-Wallis Test Statistic
indicated a significant difference among the habitats
(Conover 1980).

Results

I determined summer home range size for 34
Beavers (14 river, 11 lake and 9 pond) and fall home
range size for 27 Beavers (6 river, 13 lake and 8
pond). All Beaver home ranges tended to follow
shorelines, with movement across open water limited
to travel directly from one shore to another. In sum-
mer, five of the river Beavers occupied home ranges
which included lake areas. Three of these Beavers
were non-family Beavers and include the two largest
home ranges in the data set (42.75 ha and 26.75 ha).
The other two were an adult breeding pair with a
lodge near the river mouth. Beavers living in ponds
did not necessarily remain within the boundaries of
the pond, but frequently moved both downstream
over their dam and upstream, crossing other dams
above their home pond.

When all Beavers were considered, summer home
range size was larger for Beavers living in rivers as
compared to those living in ponds (p < 0.05) but
there was no difference between pond and lake

Median Minimum Maximum N
6.6252 2.50 12.50 14
9.75 2D 20.50 11

1375" 2225 42.75 9
6.625 2.50 8.25 12
6.00 25 20.50 9

11.00 225 16.00 4

(p > 0.05) or lake and river home ranges (p > 0.05)
(Table 1). When non-family Beavers were excluded
from the analysis, there was no significant difference
in summer home range size among the Beavers liv-
ing in the different habitats (p > 0.05) (Table 1).
Similarly, when all Beavers were considered, sum-
mer core areas were larger for river Beavers than for
those living in ponds (p < 0.05), but there was no
difference between pond and lake or lake and river
Beavers (p > 0.05) (Table 2). Again, when non-fami-
ly Beavers were excluded from the analysis, there
was no significant difference in summer core area
among Beavers in the different habitats (p > 0.05)
(Table 2). The percent of the home range area
included in the core area did not differ among the
Beavers (p > 0.05) (Table 3) in summer and neither
did the percent of activity occurring in the core area
differ among the Beavers from different habitats
(p > 0.05) (Table 4).

In fall, there was no significant difference among
the Beavers living in different habitats for home
range size (p > 0.05) (Table 5), core area size
(p > 0.05) (Table 6) or percent of home range includ-
ed in the core area (p > 0.05) (Table 7) whether all
Beavers were included or when only family Beavers
were included in the analysis. Fall core activity was
greater for lake Beavers than for river Beavers
(p < 0.05) regardless of whether non-family Beavers
were included in the analysis (Table 8). There was
no significant difference in core activity between
pond and river (p > 0.05) animals or pond and lake
animals (p > 0.05) (Table 8).

TABLE 2. Summer core area size (ha) for Beavers living in pond, lake and river habitats for all Beavers and for family
Beavers only. Median values followed by different letters are significantly different from each other (p < 0.05) (within each
group — all Beavers and family only). All other comparisons within each group are not significantly different (p > 0.05).

Family Status Habitat Mean (ha) SE

All Beavers Pond 1.86 0.194
Lake 9 Os 0337
River 4.83 1.434

Family only Pond 1.83 O.191
Lake 2.08 0.354
River 2.94 0.786

Median Minimum Maximum N
Lis OW5 3.00 14
2.00 0.75 4.00 |
3.25° O75 14.25 9
| BY fe 0.75 3.00 12
2.00 0.75 4.00 9
a5 O:75 4.50 4

1997 WHEATLEY: BEAVER HABITAT VARIATION 219

TABLE 3. Percent of the home range area included in the core area (core percent) in summer for Beavers living in pond,
lake and river habitats for all Beavers and for family Beavers only. Median values are not significantly different from each
other (p > 0.05) (within each group — all Beavers and family only).

Family Status Habitat Mean (7%) SE Median Minimum Maximum N
All Beavers Pond 28.55 1.418 28.16 2105 37.50 14
Lake 2a-03 12339 25.00 18.96 35.33 1]
River 27.69 2.366 28.13 16.36 ees 9
Family only Pond Bare os) 1.448 30.15 23.08 3750 12
Lake 25.45 1.426 25.00 1951 35.35 9
River 30.42 1.993 30°75 26.00 34.21 4

TABLE 4. Percent of all activity occurring in the core area in summer (core activity) for Beavers living in pond, lake and
river habitats for all Beavers and for family Beavers only. Median values are not significantly different from each other
(p > 0.05) (within each group — all Beavers and family only).

Family Status Habitat Mean (%) SE Median Minimum Maximum N

All Beavers Pond 74.54 2.067 76.52 60.90 84.40 14
Lake 74.34 2.006 76.34 60.89 80.67 11
River FPLDS 1790 68.00 66.66 80.79 Sy
Family only Pond bales | 2201 76.52 60.90 84.40 12
Lake 74.12 2.467 78.50 60.89 80.67 5
River 74.65 2.980 72.30 67.20 80.79 4

TABLE 5. Fall home range size (ha) for Beavers living in pond, lake and river habitats for all Beavers and for family
Beavers only. Median values are not significantly different from each other (p > 0.05) (within each group — all Beavers
and family only).

Family Status Habitat Mean (ha) SE Median Minimum Maximum N
All Beavers Pond 2.69 0.523 2.50 1.00 Sez5 8
Lake 2565 0.457 2.50 213 20.50 13
River 4.50 0.922 3.50 Pe bs) 8.00 6
Family only Pond 2.69 0.523 2.50 1.00 23 8
Lake 2.46 0.449 2.00 1.00 6.50 12
River 5.00 1.561 2.00 Aaa be) 8.00 3

TABLE 6. Fall core area size (ha) for Beavers living in pond, lake and river habitats for all Beavers and for family Beavers
only. Median values are not significantly different from each other (p > 0.05) (within each group — all Beavers and family
only).

Family Status Habitat Mean (ha) SE Median Minimum Maximum N
All Beavers Pond 1.00 O22 0.405 0.50 PI pe 8
Lake O:73 0.118 0.500 0.25 1.50 13
River 1.33 0.427 0.875 0.50 OE be 6
Family only Pond 1.00 0299. . 0.405 0.50 OE bes 8
Lake 0.69 0.120 0.500 0.25 2.00 12

River 1.50 0.661 1.250 0.50 ao 3

220

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 7. Percent of home range area included in the core area in fall for Beavers living in pond, lake and river habitats for
all Beavers and for family Beavers only. Median values are not significantly different from each other (p > 0.05) (within

each group — all Beavers and family only).

Family Status Habitat Mean (%) SE
All Beavers Pond 40.33 6.137
Lake 28.91 00
River 26.13 3.743
Family only Pond 40.33 6.137
Lake 29.24 3.254
River 2732 4.790
Discussion

Beaver home ranges are not uniform in size and
do not conform to any regular shape. Rather, the
home ranges follow the convoluted shorelines of
lakes, the meandering routes of rivers, or the natural,
irregular outlines of ponds. Shorelines are the prima-
ry source of Beaver food, either the leaves and bark
of terrestrial trees or the aquatic plants growing in
shallow water near the shore (Jenkins and Busher
1979):

When moving between feeding sites and lodges,
Beavers tended to stick close to shore. Tevis (1950)
also reported frequent movement by Beavers along
shorelines. The eyesight of Beavers is poor com-
pared to their senses of smell and hearing (Jenkins
and Busher 1979). By travelling near the shore they
may be better able to track their location by smell.
Similar movement patterns, in which the animals
stay close to the shore and avoid deep water, except
to cross, have been reported for other semi-aquatic
species such as Nile Crocodiles (Crocodilus niloti-
cus) (Hutton 1989) and Snapping Turtles (Chelydra
serpentina) (Galbraith et al. 1987). While tracking
their location by smell might be expected on the
river, it seems unusual on the lake or ponds, but by
remaining near the shore the Beavers are able to
maintain a reference point for as long as possible.

In summer, river home ranges were larger than
pond home ranges, and river core areas were larger
than pond core areas. However, this habitat differ-
ence disappeared when non-family Beavers were

Median Minimum Maximum N
37.80 20.00 75.00 8
26.67 16.67 50.00 13
23.80 18.18 38.46 6
37.80 20.00 75.00 8
28.33 16.67 50.00 12
29.41 18.18 34.38 3

excluded from the analysis. Earlier research had
shown a significant difference between the size of
family and non-family home ranges and core areas
(Wheatley 1997b). Whether this difference in home
range and core area sizes is directly due to the preva-
lence of non-family Beavers in river habitat, or
whether, instead, it is due to characteristics of the
habitat occupied by these non-family Beavers is diffi-
cult to assess. River habitat in the study area was bor-
dered by numerous bogs containing Black Spruce
(Picea mariana), Jack Pine (Pinus banksiana), Alder
(Alnus crispa) and Tamarack (Larix laricina), with a
few ridges containing Jack Pine and Trembling
Aspen (Populus tremuloides). Lakes and ponds tend-
ed to have more numerous ridges and fewer bogs
along their shores. Trembling Aspen is a favoured
food of Beavers (Jenkins and Busher 1979), and there
was a greater abundance of Trembling Aspen along
lake and pond shores than along river banks. Any
better quality habitat along the river was likely
already occupied by established Beaver families. This
leaves the poorer quality habitat for the less settled or
potentially transient non-family Beavers. If these
Beavers then had to travel farther in search of food,
this may account for the larger home ranges. But, this
poorer quality habitat in terms of food, also provides
the softer banks necessary for bank burrows. The
rockier lake shores afford little opportunity to build
bank burrows. Most of the non-family Beavers were
juveniles (Wheatley 1997b), in the process of seeking
a mate and a site for establishing their own family

TABLE 8. Percent of all activity occurring in the core area (core activity) in fall for Beavers living in pond, lake and river
habitats for all Beavers and for family Beavers only. Median values followed by different letters are significantly different
from each other (p < 0.05) (within each group — all Beavers and family only). All other comparisons within each group

are not significantly different (p > 0.05).

Family Status Habitat Mean (%) SE

All Beavers Pond 75.44 3.941
Lake 81.12 1 32
River 66.28 3.269

Family only Pond 75.44 3.941
Lake 81.39 1.638
River 63.33 5.658

Median Minimum Maximum N
Tis 62.50 95.40 8
78.608 75.60 95.20 13
65.65° 53.50 76.36 6
hy 2 62.50 95.40 8
79.70° 75.60 95.20 12
63.40! 53.50 FaAD 2

1997

base. A bank burrow, many of which already existed
along the river, provides a dwelling that uses less
energy to establish and maintain, especially if the
likelihood of establishing permanent residence at the
site is limited. So while not providing a high quality
habitat in terms of food, rivers may provide an opti-
mum habitat when both food and shelter are consid-
ered. At the same time, the regular movements of
these Beavers might be expected to range more wide-
ly in search of a suitable site for colonization. The
larger home ranges in this group of Beavers likely are
a combination of this searching behaviour of juvenile
animals and the characteristics and quality of the
habitat available to them.

While the river in this study may generally be
classed as providing poorer quality habitat, it would
be expected that Beaver ponds, which require a
greater energy input for maintenance of the dam,
would provide better quality habitat. I hypothesize
this because there would seem to be a need for a cer-
tain quality of habitat in order to justify the extra
effort necessary for building and/or maintaining a
dam. If this higher quality of pond habitats is reflect-
ed by a greater abundance of food (including aquatic
plants which may be eaten by Beavers (Jenkins
1979)), then pond Beavers should be able to obtain
sufficient food while occupying smaller home
ranges. Similarly if food is more concentrated, core
areas should be smaller. However, home ranges of
family Beavers living in ponds in this study were not
different from lake or river Beavers. In contrast with
my findings, Gillespie (1975) reports much smaller
home ranges for the pond Beavers she observed as
compared to the lake Beavers. Gillespie’s (1975)
estimates of home range size for the period of June
to October were much smaller than my estimates for
pond Beavers (1.76 ha) but larger than my estimates
for lake Beavers (15.0 ha).

The quality of the habitat may not just be reflected
in the abundance of food. Ponds may also provide
better protection from predators and a more secure
habitat in terms of water levels. While a pond has an
area that is more definitively delineated than the
river or lake, the pond Beavers in this present study
did not always remain within the boundaries of their
own pond. In contrast, Tevis (1950) found Beavers
were restricted to their own pond. This may be
dependent on the density of Beavers in the area.
Because there were no other Beavers living in the
immediate vicinity of the pond Beavers I studied,
there were no potential territorial limits from neigh-
bouring Beavers. The ponds also had a long history
of Beaver occupation (over 20 years, Bill Conley,
personal communication). This may have reduced
the quality of the habitat in terms of food availabili-
ty, resulting in the need to travel farther to find food,
while maintaining the other benefits of pond resi-
dence. So, while initially ponds might be expected to
be associated with smaller home ranges and core

WHEATLEY: BEAVER HABITAT VARIATION

221

areas, this relationship does not seem to extend to
older ponds if food availability decreases.

Similarly to the ponds, many of the lodges on the
river had been occupied for many years. In contrast to
the river and pond Beavers, lake Beavers generally
occupied newly constructed lodges, in areas where
there was little evidence of previous Beaver activity.
Because the areas had not been subject to previous
harvesting by Beavers, food was likely denser in these
areas, resulting in more concentrated activity and
explaining the higher value of fall core activity for
lake Beavers. Buech (1995) has also hypothesized that
differences between habitats in the time spent on dif-
ferent types of behaviour may account for differences
in home range size and apparent patterns of use.

Although there are some small differences
between the habitats in patterns of home range use,
the relative similarities between the three habitats
were surprising. One final explanation for these sim-
ilarities may be the forest fire history of the area.
The entire study area was burnt in May, 1980, six
years before the start of this study. While the intensi-
ty of the forest fire varied across the study area, all
the locations where I studied Beavers were affected
to some extent. The short period between the time of
the fire and the study may mean that there was little
time for differentiation between the habitats, which
were all still in the early successional stages through-
out this study. As succession proceeds at different
rates in different locations throughout the region,
habitats may become more distinct and differences
in Beaver home ranges more apparent. But, because
forest fires are a natural part of succession in the
boreal forest, it may be that the lack of differentia-
tion between habitats is the more common scenario
within the boreal regions.

Acknowledgments

Funding for this study was provided by the Taiga
Biological Station Research Trust; Wildlife Branch,
Manitoba Department of Natural Resources; Sigma
Xi; Manitoba Chapter, The Wildlife Society and the
Manitoba Wildlife Federation and a University of
Manitoba Fellowship. W. O. Pruitt provided advice,
support and assistance throughout this study and read
many earlier drafts of this paper. Numerous field assis-
tants made this study possible. S. C. Jay, S. H. Jenkins,
R. Riewe and K. Stewart and two anonymous review-
ers improved earlier versions of this manuscript.

Literature Cited

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dynamics of Newfoundland Beaver. Canadian Journal of
Zoology 55: 1480-1492.

Brady, C. A., and G. E. Svendsen. 1981. Social behaviour
in a family of Beaver, Castor canadensis. Biology of
Behavior 6: 99-114.

Buech, R. R. 1995. Sex differences in behavior of Beavers
living in near-boreal lake habitat. Canadian Journal of
Zoology 73: 2133-2143.

222

Busher, P. E. 1975. Movements and activities of Beavers,
Castor canadensis, on Sagehen Creek, California. M.A.
thesis, San Francisco State University, California. 86
pages.

Conover, W. J. 1980. Practical non parametric statistics.
John Wiley and Sons, New York.

Davis, J. R. 1984. Movement and behavior patterns of
beaver in the Piedmont of South Carolina. M.Sc. thesis,
Clemson University, Clemson, South Carolina.
83 pages.

Galbraith, D. A., M. W. Chandler, and R. J. Brooks.
1987. The fine structure of home ranges of male
Chelydra serpentina: are snapping turtles territorial?
Canadian Journal of Zoology 65: 2623-2629.

Gillespie, P. S. 1975. Summer activities, home range and
habitat use of beaver. M.Sc. thesis, University of
Toronto, Toronto, Ontario. 149 pages.

Green, H. U. 1936. The beaver of the Riding Mountain,
Manitoba: An ecological study and commentary.
Canadian Field-Naturalist 50: 1-8, 21-23, 36-50,
61-67, 85-92.

Hutton, J. 1989. Movements, home range, dispersal and
the separation of size classes in Nile crocodiles.
American Zoologist 29: 1033-1049.

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Jenkins, S. H. 1979. Seasonal and year-to-year differences
in food selection by Beavers. Oecologia 44: 112-116.
Jenkins, S. H., and P. E. Busher. 1979. Castor

canadensis. Mammalian Species 120: 1-8.

Morgan, L. H. 1986. The American beaver: A classic of
natural history and ecology. Dover Publications Inc.,
New York.

Tevis, L., Jr. 1950. Summer behavior of a family of
Beavers in New York State. Journal of Mammalogy 31:
40-65.

Warren, E. R. 1927. The beaver its works and its ways.
American Society of Mammalogists Monographs 2,
Baltimore, The Williams and Wilkins Company.

Wheatley, M. 1997a. Beaver, Castor canadensis, home
range size and patterns of use in the taiga of southeastern
Manitoba: I. Seasonal variation. Canadian Field-
Naturalist 111: 204-210.

Wheatley, M. 1997b. Beaver, Castor canadensis, home
range size and patterns of use in the taiga of southeastern
Manitoba: II. Sex, age and family status. Canadian
Field- Naturalist 111: 211-216.

Received 29 January 1996
Accepted 23 October 1996

Food Preference and ad libitum Intake of Wild-captured Sitka Mice,
Peromyscus keeni sitkensis

Ep O. REESE!, JEFFREY C. BARNARD, and THOMAS A. HANLEY2

U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, P.O. Box 20909, Juneau, Alaska
99802-0909

‘Current address: Hamngatan 39B, 185 32 Vaxholm, Sweden

2Author to whom correspondence should be addressed.

Reese, Ed O., Jeffrey C. Barnard, and Thomas A. Hanley. 1997. Food preference and ad libitum intake of wild-captured
Sitka Mice, Peromyscus keeni sitkensis. Canadian Field-Naturalist 111(2): 223-226.

Food preferences of Sitka Mouse (Peromyscus keeni sitkensis) were studied in four experiments involving fruits from five
major shrub species and seeds from three tree and four shrub species. Each experiment consisted of individual animals
being offered mixed-species assortments of foods fed over a four-day period. Ad libitum intake trials were conducted with
the same foods used in the preference experiments but with only one food per trial. Intake trials also included beetles
(Seaphinotus and Pterostichus spp.) and Pink Salmon (Oncorhynchus gorbuscha). Salmonberry (Rubus spectabilis) and
Stink Current (Ribes bracteosum) fruit and seeds emerged as highly preferred and valuable foods in terms of both relative
preference and ad libitum intake. Devilsclub (Oplopanax horridus) and Elderberry (Sambucus racemosa) fruits were least
preferred in both preference and intake, although their seeds were relatively palatable and nutritious (in terms of weight
loss). Blueberry (Vaccinium ovalifolium) fruit was intermediate in both preference and intake. Tree seed palatability and
intake ranked in order of Sitka Spruce (Picea sitchensis) (highest), Western Hemlock (Tsuga heterophylla), and Red Alder
(Alnus rubra) (lowest). Ad libitum intake of beetles was low, and that of Pink Salmon was near zero. These results should
be helpful in interpreting diet composition studies of Sitka Mouse throughout its range and possibly other Peromyscus
species in wet coastal forests of the Pacific Northwest and Alaska.

Key Words: Sitka Mouse, Peromyscus keeni sitkensis, diet choice, diet selection, voluntary intake, food habits, Alexander

Archipelago, Alaska.

The Sitka Mouse (Peromyscus keeni sitkensis) 1s
endemic to the outer-coast islands of the Alexander
Archipelago of southeastern Alaska and to the
Queen Charlotte Islands, British Columbia (Swarth
1911; Nagorsen 1990). Until recently, it was consid-
ered a separate species (P. sitkensis), Hogan et al.
(1993), however, have proposed that it and the Deer
Mouse (P. maniculatus) in southeastern Alaska and
northern British Columbia coast as well as the
Columbian Mouse (P. oreas) of northern Vancouver
Island and vicinity are all subspecies of Keen’s
Mouse (P. keeni). The Sitka Mouse inhabits temper-
ate, coniferous rain forest and associated habitats
within the Western Hemlock-Sitka Spruce (Tsuga
heterophylla — Picea sitchensis) forest type (Ruth
and Harris 1979). Its distribution and population
ecology have seldom been studied, and even less is
known of its feeding ecology. Like others of the
Peromyscus genus (Van Horne 1982), however, its
diet is one of mostly seeds, supplemented with
insects and other invertebrates (T. A. Hanley, J. C.
Barnard, and M. Ben-David, U.S. Forest Service and
University of Alaska, unpublished data).

Food preference experiments, where differential
availability of foods is not a factor, have never been
reported for the Sitka Mouse or any other
Peromyscus species living in similar habitats.
Therefore, we studied food preferences and ad libi-

tum intake of major summer foods of the Sitka
Mouse under controlled conditions.

Methods

The study was conducted during July and August,
1995, at the Kadashan River field station on
Chichagof Island, southeastern Alaska (57° N, 135°
W). Animals were captured as needed, using
Longworth traps baited with rolled oats. Traplines
were in Sitka Spruce floodplain, mature Red Alder
(Alnus rubra) floodplain, and old-growth Western
Hemlock-Sitka Spruce upland forests (Hanley and
Hoel 1996). All juveniles (based on pelage col-
oration) and pregnant or lactating females were
released; we used only nonbreeding adults. Mice
were maintained individually in_ separate
40 X 38 X 30-cm plywood boxes with a screen top
and an aluminum bottom liner to facilitate cleaning.
Each box contained a water bottle, steel nest box
(9 xX 14 X 8 cm) filled with polyester bedding, and
set of feeding stations. Mice were fed a millet-based
bird seed mixture during non-trial periods and were
allowed a minimum of one week to adjust to captivi-
ty before being used in a feeding trial. Feeding
stations constructed of plywood with sheet alu-
minum dividers and a raised floor of hardware cloth
allowed simultaneous presentation of 1-4 foods in
1/2-pint (240-ml) wide-mouth Mason jars while min-

223

224

imizing spillage and mixing. Fresh food was present-
ed and the orts collected each day for four consecu-
tive days. Dry matter was determined for each food
(100° C for 24 hrs, measured with a Mettler balance
to 0.01 g) and multiplied by wet weight to determine
dry weight offered. Feces were removed, and orts
were dried also. Animals were weighed daily (with
an Avinet spring scale to 0.5 g), and a trial for any
given animal was discontinued if that animal lost
more than 25 percent of its initial body weight. We
followed the American Society of Mammalogists'
(1987) care protocol.

Relative preferences were analyzed in four sepa-
rate experiments of two to four foods each. All 12
foods could not be fed simultaneously for the fol-
lowing reasons: (1) their availability changed
throughout the summer, so all were not simultane-
ously available; (2) we could not harvest sufficient
quantities of more than three or four foods at a time;
and (3) the number of mice needed for statistical
analysis was estimated to be 3 X the number of
foods in an experiment (Roa 1992). We used a total
of 17 mice (6 males, 11 females). Some mice were
used in several experiments, others were used in
only one. Mice were selected on the basis of appear-
ing healthy and in good body condition. Each exper-
iment was conducted and analyzed independently.
Therefore, relative preferences could be determined
within experiments but not between experiments.

Selection of offered foods was based on the tim-
ing of their local availability and our ability to col-
lect sufficient quantities. Foods were grouped in
similar-kind trials (fruits, tree seeds, fruit seeds)

THE CANADIAN FIELD-NATURALIST

Vol. 111

accordingly. Fresh fruit was collected daily for fruit
trials. Seeds of Sitka Spruce, Western Hemlock, and
Red Alder were separated from their cones by a
mechanical separator. Fruit seeds were obtained by
crushing, screening, and washing ripe fruits.

Nine mice were used in Experiment 1, which
involved fresh fruits: Salmonberry (Rubus
spectabilis), Blueberry (Vaccinium ovalifolium), and
Elderberry (Sambucus racemosa) (14-17 July). Six
mice were used in Experiment 2, which also
involved fresh fruits: Stink Current (Ribes bracteo-
sum) and Devilsclub (Oplopanax horridus) (28-31
July). Nine mice were used in Experiment 3, which
involved tree seeds: Sitka Spruce, Western Hemlock,
and Red Alder (4-8 August). And 12 mice were used
in Experiment 4, which involved seed cleaned from
fruit: Salmonberry, Elderberry, Stink Current, and
Devilsclub (25-28 August) (Blueberry seeds were
too small to include).

Ad libitum intake trials were conducted with three
or four mice (depending on our ability to harvest
food) for each food during the week immediately
following the respective preference experiment.
Each food was fed separately. Additionally, intake
trials were conducted with fresh and partially
decayed Pink Salmon (Oncorhynchus gorbuscha)
and various species of live beetles captured in pitfall
traps (predominantly Seaphinotus angusticollis with
a few Pterostichus crenicollis and P. castaneus).

The preference experiments were analyzed with
multivariate analysis of variance as suggested by
Roa (1992). Total dry-matter intake of each food by
each mouse over the 4-day trial was divided by the

TABLE 1. Mean dry-matter intake (total four-day intake divided by mean body weight, for each mouse) and mean change
in body weight (grams) in four preference experiments. N = Number of mice. SD = Standard devation. P = Probability of

null hypothesis, multivariate analysis of variance.

Mean dry-

Experiment/Food matter intake

Experiment | (N = 9)

Salmonberry fruit 0.380

Blueberry fruit 0.069

Elderberry fruit 0.007
Experiment 2 (N = 4)

Stink Current fruit 0.161

Devilsclub fruit 0.077
Experiment 3 (N = 9)

Spruce seed 0223

Hemlock seed 0.094

Alder seed 0.012
Experiment 4 (N = 12)

Devilsclub seed 0.214

Salmonberry seed@ 0.098

Elderberry seed@ 0.078

Stink Current seed 0.011

Mean body
SD P weight change SD
0.000 -3.67 2.70
0.061
0.038
0.044
0.009 -5.75 3.16
0.035
0.015
0.000 -0.94 2.86
0.044
0.051
0.028
0.000 La 3.47
0.065
0.043
0.048
0.013

“Foods with same superscript do not differ from one another at alpha = 0.05.

£997,

REESE, BARNARD, AND HANLEY: FOOD OF SITKA MICE

225

TABLE 2. Mean ad libitum dry-matter intake (grams per day divided by mean body weight, for each mouse) and daily body
weight change (grams) while on single foods during three- to four-day trials. N = number of mice in trial. Mean intake
values with same superscript do not differ at the alpha level of 0.05, analysis of variance.

Food N

Salmonberry fruit
Stink Current fruit
Salmonberry seed
Stink Current seed
Elderberry seed
Spruce seed
Hemlock seed
Alder seed!
Devilsclub seed
Blueberry fruit!
Beetles

Salmon (rotten)
Elderberry fruit
Devilsclub fruit
Salmon (fresh)?

G0 Gd) (GoGo) SS SG Is

Mean dry-matter intake Mean weight change

0.138? -0.70
0.1082" -1.82
0.09225 -1.13
0.0902>«< -1.92
0.0892>«< -0.46
0.084>< -0.47
0.078>¢ -0.41
0.064°«-4 -2.72
0.061>«4 -0.67
01057268 -2.50
0.050°-4¢ pis}
0.0214¢ -2.47
0.017%¢ -1.56
0.015% -1.63
0.000° -3.07

'One animal with only two days in trial is included in the alder seed trial and in the blueberry fruit trial.
?Two animals with only two days in trial are included in the fresh salmon trial.

individual’s average body mass to standardize for
differences in body mass. MANOVA and Tukey’s
test statistic (Wilkinson et al. 1992) were used to
determine statistically significant differences in
means. Results of ad libitum intake trials were com-
pared using ANOVA and Tukey’s test. An alpha
level of 0.05 was used for all statistical tests and
comparisons. All mice were released upon comple-
tion of the study.

Results and Discussion

Mean pretrial body weight of all mice in all pref-
erence experiments and intake trials was 32.1 g (SD
= 4.2). Most animals (73 of 90) lost weight during
all experiments and trials, which is common when
captive animals are fed single- or few-species rations
of natural foods. Two of the six mice in Experiment
2 lost > 25% body weight and were dropped from
the four-day trial. All other mice completed their
four-day trials in each of the other experiments.
Significant differences in mean intakes (MANOVA)
occurred in all four preference experiments, and all
foods differed significantly from each other (Tukey’s
test) in all experiments except Experiment 4 where
Salmonberry seed and Elderberry seed did not differ
(Table 1). Among fruits, Salmonberry was clearly
most preferred and Elderberry least preferred.
Interestingly, however, Devilsclub was the most
highly preferred seed and Stink Current the least pre-
ferred seed (Experiment 4), which was exactly the
opposite order of their preferences as fruits
(Experiment 2). Among tree seeds, Sitka Spruce was
most highly preferred and Red Alder least preferred
(Experiment 3).

Mice were restricted to only one food in the intake

trials and lost weight much more quickly than in the
preference experiments (Table 2). In 16 instances,
intake trials were terminated after only three days,
and in four instances after two days. Rather than dis-
card these data, results were expressed as mean daily
intake rather than total trial intake, and all mice with
22 days of data were included. Food intakes were
significantly different (P < 0.05; Table 2). Greatest
daily intakes were of Salmonberry and Stink Current
fruit and seed, while lowest intakes were of Pink
Salmon, Devilsclub fruit, Elderberry fruit, and bee-
tles. Ad libitum intake is a function of at least two
counteracting variables: palatability and nutritional
quality (Robbins 1993). High palatability should
result in high intake, but high intake could also result
from low nutritional value as animals are forced to
eat more food to maintain nutritional balance. It is
impossible to determine cause-and-effect in such
simple trials.

Salmonberry and Stink Current fruits were highly
preferred in the preference experiments and were
consumed in greatest quantity in the intake trials.
Salmonberry, however, appears to be nutritionally
superior to Stink Current in terms of mice weight
loss (Tables 1 and 2). Seeds resulted in less weight
loss than did fruits in the other species. Both
Devilsclub and Elderberry, especially, appeared to
be both more highly palatable and nutritious as
seeds than as fruits in both preference experiments
and intake trials. Red Alder seeds were the least
preferred of the tree seeds, and animals lost sub-
stantial weight when forced to eat them in the
intake trials.

Although Pink Salmon were not used in prefer-
ence experiments, the extremely low intake of Pink

226

Salmon and high weight losses of mice on Pink
Salmon in the intake trials indicate that Pink Salmon
probably are not eaten by mice in the field. That
also is consistent with stable isotope analysis (!°C
and !°N) of riparian food webs in our study area (M.
Ben-David, University of Alaska, unpublished
data). Pink Salmon carcasses scattered along the
streams and throughout the riparian forests by bears,
eagles, and other animals are readily available to
mice throughout much of each summer. We had
expected that they might provide a protein-rich food
source for the generalist-feeding Sitka Mouse at that
time.

Studies that quantify diet composition in terms of
broad categories such as fruit and seeds, insects,
etc., do not take into account the wide variation in
palatability, voluntary intake, and nutritional value
of food types and species. Field studies, regardless
of detail, also are subject to differential availability
of foods. Our study helps to identify the relative pri-
orities of major foods in terms of feeding behavior
by the Sitka Mouse. Although our preference exper-
iments and intake trials provide insight into only a
small part of the feeding ecology of the Sitka
Mouse, they should be useful in interpreting field
data from other studies throughout the species’
range and possibly also that of other Peromyscus
species in similar habitats.

Acknowledgments

We thank our field assistants in this study and all
other work of the 1995 season: Nicklas Fronth,
Lillian Lundin, Aashild Odden, Christian Riser,
Anna Rooth, Peter Rosen, and Anja Wannag.

THE CANADIAN FIELD-NATURALIST

Vel HI

Literature Cited

American Society of Mammalogists. 1987. Acceptable
field methods in mammalogy: preliminary guidelines
approved by the American Society of Mammalogists.
Journal of Mammalogy 68 (Supplement 4): 1-18.

Hanley, T. A., and T. Hoel. 1996. Species composition of
old-growth and riparian Sitka spruce-western hemlock
forests in southeastern Alaska. Canadian Journal of
Forest Research 26: 1703-1708.

Hogan, K., M. C. Hedin, H. S. Koh, S. K. Davis, and I. F.
Greenbaum. 1993. Systematic and taxonomic implica-
tions of karyotypic, electrophoretic, and mitochondrial-
DNA variation in Peromyscus from the Pacific
Northwest. Journal of Mammalogy 74: 819-831.

Nagorsen, D. W. 1990. The mammals of British Columbia:
A taxonomic catalogue. Memoir Number 4, Royal
British Columbia Museum, Vancouver. 140 pages.

Roa, R. 1992. Design and analysis of multiple-choice feed-
ing-preference experiments. Oecologia 89: 509-515.

Robbins, C. T. 1993. Wildlife feeding and nutrition, second
edition. Academic Press, New York, New York. 352 pages.

Ruth, R. H., and A. S. Harris. 1979. Management of west-
ern hemlock-Sitka spruce forests for timber production.
General Technical Report PNW-88. U.S. Department of
Agriculture, Forest Service, Pacific Northwest Research
Station, Portland, Oregon. 197 pages.

Swarth, H. S. 1911. Birds and mammals of the 1909
Alexander Expedition. University of California
Publications in Zoology 7: 9-172.

Van Horne, B. 1982. Niches of adult and juvenile deer
mice (Peromyscus maniculatus) in seral stages of conif-
erous forest. Ecology 63: 992-1003.

Wilkinson, L., M. Hill, J. P. Welna, and G. K.
Birkenbeuel. 1992. SYSTAT for Windows: Statistics,
version 5 edition. SYSTAT, Evanston, Illinois. 750 pages.

Received 5 February 1996
Accepted 12 October 1996

Summer Food Habits and Population Density of Coyotes,
Canis latrans, in Boreal Forests of Southeastern Québec

CLAUDE SAMSON! and MICHEL CRETE2

'\Département de biologie, Pavillon A.-Vachon, Université Laval, Sainte-Foy, Québec G1K 7P4

2Service de la faune terrestre, Ministére de l'Environnement et de la Faune, 150 boul. René-Lévesque Est, Québec,
Québec GIR 4Y1, and Centre d'études nordiques, Pavillon J.-C. Bonenfant, Université Laval, Sainte-Foy, Québec
G1K 7P4 [Author to whom all correspondence should be sent. ]

Samson, Claude, and Michel Créte. 1997. Summer food habits and population density of Coyotes, Canis latrans, in boreal
forests of southeastern Québec. Canadian Field-Naturalist 111(2): 227-233.

The Coyote (Canis latrans) arrived on the Gaspé peninsula in the mid 1970s. The population increased through 1990 fol-
lowed by a general decline. We compared summer food habits of the Coyote in 1988 and 1991 in central Gaspé peninsula,
a forested area dominated by conifers. We analyzed 231 scats in 1988 and 435 scats in 1991 to determine Coyote food
habits. In 1991, we also marked seven Coyotes with radio-collars and a radioactive isotope (Zn) to estimate population
density in the study area. The density estimate of 0.2 - 0.3 Coyote/10 km? (+ 35 - 36%) was among the lowest ever record-
ed. Woodchuck (Marmota monax) and White-tailed Deer (Odocoileus virginianus) dominated the diet in May-June 1988
but were replaced by Moose (Alces alces), Snowshoe Hares (Lepus americanus) and Beavers (Castor canadensis) in 1991.
Berries were preferred in mid-summer and comprised 56% and 80% of the volume of scats in August of 1988 and 1991,
respectively. The use of small mammals remained relatively constant during both years; 1.e., 9 - 19% of scat volume.
Results are related to levels of prey abundance on the Gaspé peninsula.

Les coyotes (Canis latrans) sont arrivés dans la péninsule gaspésienne au milieu des années 1970. La population a connu
une expansion jusqu’en 1990, suivi d’un déclin général. Nous avons comparé le régime alimentaire estival du Coyote en
1988 et 1991 dans la région centrale de la péninsule gaspésienne qui est dominée par les foréts conifériennes. Pour ce faire,
nous avons analysé le contenu de 231 féces récoltées en 1988 et de 435 féces en 1991. En 1991, nous avons aussi marqués
7 coyotes avec un collier-émetteur et un isotope radioactif (°°Zn) pour estimer la densité de la population de I’aire d’ étude.
La densité de cette population, estimée a 0,2 - 0,3 coyote/10 km? (+ 35-36%), était parmi les plus faibles jamais observées.
Les principales proies utilisées en mai-juin 1988 étaient la marmotte (Marmota monax) et le cerf de Virginie (Odocoileus
virginianus), mais ces proies furent remplacées par |’ orignal (Alces alces), le li¢vre d’ Amérique (Lepus americanus) et le
castor (Castor canadensis) en 1991. Les petits fruits étaient préférés au milieu de |’ été, représentant jusqu’a 56% du vol-
ume des féces en aoa 1988 et 80% en aout 1991. L’utilisation des petits mammiféres est restée relativement stable durant
les deux années, variant entre 9 et 19% du volume. Ces résultats sont discutés en relation avec l’abondance des proies dans
la péninsule gaspésienne.

Key Words: Eastern Coyote, Canis latrans, summer diet, population density, Québec.

The Coyote (Canis latrans) is as an adaptable car- The decline of White-tailed Deer (Odocoileus vir-

nivore that changes its diet in response to seasonal
food availability (Todd et al. 1981; Van Vuren and
Thompson 1982; Barrett 1983; Bowyer et al. 1983;
Harrisson and Harrisson 1984; Parker 1986; Andelt
et al. 1987; MacCracken and Hansen 1987; Dibello
et al. 1990). This flexible behaviour has helped the
Coyote expand its range following European settle-
ment in North America. Coyotes arrived on the
Gaspé peninsula in southeastern Québec in mid
1970s and were well established in the mid 1980s
(Créte and Lemieux 1994*). Eastern Coyotes are
larger than other conspecifics and may hunt during
winter in family unit, enabling them to kill larger
prey (Harrisson 1992; Lariviére and Créte 1992).
Consequently, two species of ungulates on the Gaspé
peninsula have been seriously impacted by Coyotes.

*see Documents Section preceding Literature Cited.

ginianus) in the Gaspé peninsula between 1986 and
1991 was attributed to a series of harsh winters.
Mortality from starvation was nevertheless enhanced
by Coyote predation (Poulle et al. 1993; Créte and
Lemieux 1994*). Most large winter deer yards were
unoccupied by 1992 and the hunting season was
closed (Créte and Lemieux 1994*). After the deer
crash, Coyotes also declined on the Gaspé peninsula
(Créte and Lemieux 1994*).

Coyotes also threatened the last population of
Caribou (Rangifer tarandus) which remained on the
mainland east of the Fleuve Saint-Laurent: the
Gaspésie Park Caribou herd (Créte et al. 1990; Créte
and Desrosiers 1995). From 1984 to 1988, summer
mortality of calves approached 90% and most mor-
talities were from Coyote predation (Créte and
Desrosiers 1995). Black Bears (Ursus americanus)
also killed calves, but most bears in the area appar-
ently avoided the alpine tundra occupied by Caribou
(Boileau et al. 1994). Dumont (1993), however,

227

228

observed that the presence of hikers in alpine tundra
of the Gaspésie Park disturbed females with calves
and forced them to enter forested habitats where
they were more vulnerable to predation. The
removal of Coyotes and bears after 1990 and the
control of visitor access since 1992, have helped to
increase the survivorship of calves (Créte and
Desrosiers 1995).

Although many aspects of Coyote ecology and
population dynamics in northeastern North America
have been studied (see Boer 1992 and Parker 1995),
little is known of its ecology in eastern boreal
forests. The boreal forest is probably a marginal
habitat for Coyotes and White-tailed Deer, as both
species are at the northern limits of their range. More
knowledge on the ecology of Coyotes is necessary to
understand Coyote-prey relationships in that biome.
Poulle et al. (1995) examined the fecundity and the
seasonal variations in body reserves of Coyotes from
the Gaspé peninsula. In this study, we report on the
summer food habits of Coyotes in 1988, when num-
bers peaked on the Gaspé peninsula, and in 1991,
after Coyotes began to decline. We also estimated
the density of Coyotes in boreal forests in the centre
of the Gaspé peninsula during 1991.

Study Area

The study area included parts of Gaspésie
Conservation Park (GCP) and the Chic-Chocs
Game Reserve (CCGR), in the centre of the Gaspé
peninsula (=49°00’N, 66°00’W). Topography is
rugged, with the McGerrigle Mountains forming an
elevated plateau dominated by the Mont Jacques-
Cartier (1270 m). At low elevation, forests are dom-
inated by Balsam Fir (Abies balsamea), Paper Birch
(Betula papyrifera) and Black Spruce (Picea mari-
ana). At higher elevation in GCP, alpine tundra
occupies wind-exposed sites and Black Spruce and
Balsam Fir krummohlz grow in protected areas. No
logging, hunting or trapping has occurred in GCP
since 1977. These activities persist in CCGR. A net-
work of gravel roads covers most of GCP and
CCGR at low elevation, but only a few hiking trails
provide access to the alpine areas. The climate is
continental with a mean yearly temperature below
0°C (Gagnon 1970*). Annual precipitation averages
1663 mm on Mont Logan (1128 m altitude), includ-
ing a snow fraction of 33%; snow cover persists
from early October to early June. Near sea level,
precipitation is 854 mm with a snow fraction of
32%; snow covers the ground from late November
to early May.

Methods
Population density

The density of Coyotes was estimated in a
660 km? capture area in CCGR (Figure 1). We
avoid biased estimates by maintaining sufficient dis-

THE CANADIAN FIELD-NATURALIST

Vol. 111

tance (~10-20 km) from where Coyotes were
removed in GCP to reduce predation on Caribou.
Coyotes were captured in May-June 1991 using
modified foot-hold traps. Seven Coyotes were fitted
with radiocollars and injected intramuscularly with
2.2 MBg radioactive solution of ®Zn. In July 1991,
scats were collected on roadsides in the capture area.
The proportion of radioactive scats was determined
using a Nal detector with a multichannel analyzer
(see Jolicoeur et al. 1993). The number of marked
coyotes in the capture area was determined by locat-
ing the radiocollared animals from a helicopter in
late June. Of the seven marked Coyotes, five were
located in the capture area. We could not determine
if the two remaining individuals had left the area or
if their transmitters failed. Therefore, two estimates
of population density were made, one which
assumed that all marked Coyotes were still in the
capture area in July, and the other that assumed five
of seven marked Coyotes were in the area. The size
of the Coyote population was estimated from the
proportion of radioactive scats (Lancia et al. 1994)
by using a Petersen Index (modified by Bailey
(1951) in Caughley (1977)).

Food habits

Coyote scats were searched along gravel roads
once every two or three weeks from June to August
1988 and from May to October 1991. Other scats
were collected opportunistically around trapping
sites in both CCGM and GCP. No effort was made
to collect scats on the plateau used by caribou due to
lack of access. Scats were oven-dried at 70°C for 24
hours to kill parasites and then washed through a 2
mm sieve. Non-digested remains were collected, dif-
ferent items were separated, identified and their vol-
ume in each scat was visually estimated. Mammal
hairs were identified using the pigment patterns of
the medulla and the morphology of cuticular scale
(Adorjan and Kolenosky 1969). A reference collec-
tion served to identify hairs, bones, berries and vege-
tation. Comparison between 1988 and 1991 was per-
formed on a monthly basis with the Mann-Whitney
U test on the mean volume per scat.

Results
Population density

In July 1991, 62 scats were collected in the cap-
ture area of which 19 contained the radioactive
marker. Assuming that all marked Coyotes occupied
the area, population size was estimated at 22 + 8
individuals (+ 95% confidence intervals); i.e., 0.3
Coyote/10 km2. If only five of the seven marked
Coyotes stayed in the capture area, population size
was estimated at 16 + 6 individuals or 0.2
Coyote/10 km2.

Food habits
Monthly scat collection was not uniform. In 1988,

1997 SAMSON AND CRETE:

Gaspésie Park
______ Chic-Chocs Game Reserve
2 Density estimate area
Paved Roads
Gravel Roads (permanent)
Gravel Roads (temporary)

FOOD AND DENSITY OF COYOTES

229

FiGurE 1. Location of Gaspésie Conservation Park and Chic-Chocs Game Reserve in southeastern Québec, showing the

area where Coyote population density was estimated.

15 scats were collected in June, 50 in July and 166 in
August. In 1991, scats were collected in May (n=73),
June (n=29); July (n=125), August (n=74),
September (n=125) and October (n=19).

The use of Woodchuck (Marmota monax) and
deer declined from June to August 1988, passing
from 34% and 16% to 10% and 0.5% of volume
respectively (Figure 2). During the same period, the
use of berries (mostly Vaccinium sp., Amelanchier
sp., and Aralia nudicaulis) increased (2% to 56% of
volume) whereas the use of small mammals (17% to
14% of volume) and vegetation (11% to 13%)
remained stable (Figure 2). In 1991, the feeding
habits of Coyotes shifted from predominantly Moose
(Alces alces) (28%-33% of volume) and Snowshoe
Hare (Lepus americanus) (49%-32% of volume) in
May-June, to predominantly berries in July-October
(41%-80% of volume) (Figure 2). The use of small
mammals remained more constant, fluctuating from
8% to 23% between May and October (Figure 2).

Food habits differed significantly between 1988
and 1991. In June 1988, White-tailed Deer,
Woodchuck and vegetation (grasses and sedges)
comprised up to 61% of the volume of scats, but
they were mostly replaced by Moose, Snowshoe
Hares and Beavers (Castor canadensis) in 1991
(total volume = 73%; Table 1). In July, the trend
remained the same for the Deer and Woodchuck,
whereas berries increased markedly from 1988 to
1991 (Table 1). Woodchuck consumption was higher
in August 1988 compared to 1991, while the intake

of berries was lower (Table 1). During the three
month period, the use of vegetation was higher in
1988 than in 1991.

Discussion
Population density

Precision of our population density estimates
(+35-36%) was limited by the small number of
Coyotes marked. Also, the size of this population
may have been overestimated as two of the seven
radiocollared Coyotes may have left the capture area.
The assumption of a closed population, required for
the Petersen index, is therefore violated, causing an
underestimation of the recapture/capture ratio.
Nevertheless we believe that this estimate is repre-
sentative of the low carrying capacity of eastern
boreal forests. Indeed, our estimates of density were
among the lowest ever recorded for a Coyote popula-
tion. Comparisons of population density between
studies are often difficult due to differences in tech-
niques of estimation. However, the summer density
of Coyotes seems much clearly higher in southern
United States and western Canada, varying from 3 to
20 Coyotes/10 km? (Bowen 1982; Pyrah 1984;
Andelt 1985; Hein and Andelt 1995). In northeastern
North America reported densities also vary: |
Coyote/10 km? in Ontario (Voigt and Berg 1987),
1.2 Coyotes/10 km2 in southern Québec (Messier et
al. 1986) and 2 Coyotes/10 km? in Minnesota (Berg
and Kuehn 1986). Fortin (1995) estimated a mini-
mum density of 0.7 Coyote/10 km? in the mixed

230

1988
100%

80%

Berries

60% Small mammals

YJ Woodchuck

Volume (%)

40%

20%

August Sept. Oct.

n=166

1991

Ea Other

Berries

Small mammals
E41 Hare

[_] Moose

Volume (%)

Sept. Oct.
n=125 n=19

August
n=74

May June July
n=73 n=29 n=125
FIGURE 2. Average volume (%) of food items in Coyote
scats during summers of 1988 and 1991 in the cen-
tre of the Gaspé peninsula (Québec). Sample size

was 231 scats in 1988 and 435 in 1991.

forests of Forillon National Park, 110 km east of our
study area. Fewer density estimates are available for
boreal forests. In Alberta, winter density averaged 3
Coyotes/10 km? (Todd et al. 1981). Density, howev-
er, fluctuated from 1.4-2.5 Coyotes/10 km’, when
Snowshoe Hares were scarce, to 4.4 Coyotes/10
km’, when Hares became abundant. As the number
of Coyotes in our study area was most likely near
their peak following colonization of the Gaspé
Peninsula, our low density estimate suggests a con-
sequent low carrying capacity of their environment,
especially since the White-tailed Deer had declined
(Poulle et al. 1993).

THE CANADIAN FIELD-NATURALIST

Vol. 111

Food habits

Differences in the diet between 1988 and 1991
may have been caused by a change in the availability
of certain prey types. If so, the decrease in the con-
sumption of deer and Woodchuck in 1991 could be
related to a decline in the abundance of these prey
types. Indeed, harvest of deer on the Gaspé peninsu-
la, which may be used as an index of population
trends, decreased from 620 individuals in 1988 to
143 in 1991. The consumption of Moose, Snowshoe
Hares and Beavers was greater in 1991 than 1988,
possibly due to a greater abundance of these species.
However, while Moose harvest increased from 1988
(n=590) to 1991 (n=679) (Bouchard 1990*;
Lamontagne et al. 1992*), the number of harvested
Hares declined (1988: 63.3 Snowshoe Hare/100
km2; 1991: 24.2 Hare/100 km2) (R. Lafond,
Ministére de |’Environnement et de la Faune, unpub-
lished data). In the CCGR, the harvest of Beavers
declined from 67 during the winter of 1987-1988,
to only 11 during winter 1990-1991 (R. Lafond,
Ministére de |’Environnement et de la Faune, unpub-
lished data). We hypothesize that the number of
Woodchucks and Deer may have been so depres-
sed in 1991 that Coyotes were forced to switch to
Moose, Beavers and Snowshoe Hares, even though
the two latter species were less abundant in 1991
than in 1998.

The decline of deer on the Gaspé peninsula
was due, in part, to Coyote predation (Créte and
Lemieux 1994*; Poulle et al. 1993). However, there
is no evidence in the literature that Coyotes may
significantly reduce Woodchuck numbers. Adult
Woodchucks may weight 5 kg (Lee and Funderbury
1982) and represent a profitable and reliable prey
for eastern Coyote. Open woodlands and clearings
are its prime habitat (Lee and Funderburg 1982),
but Woodchucks are confined to recent clearcuts
and roadsides in boreal forests. Although popula-
tions of this species are generally resilient (Davis
1962; Davis et al. 1964), the restricted distribution
of this rodent may have contributed to a decline in
Woodchuck numbers when Coyotes arrived in the
area. Woodchucks also were important in the sum-
mer diet of Coyotes when they colonized New
Brunswick (Parker 1986). Unfortunately, no data on
Woodchuck abundance in our study area were
available to test this hypothesis further.

The importance of Coyote predation to the de-
clines of Snowshoe Hare and Beaver in the study
area remains uncertain. Snowshoe Hare populations
do experience periodic fluctuation and predation has
been implicated in these demographic changes
(Keith et al. 1981; Keith 1990).

Monthly differences in the food habits of Coyotes
generally reflect changes in the food availability
(Van Vuren and Thompson 1982; Andelt et al. 1987;
MacCracken and Hansen 1987; Cypher et al. 1993).

1997

SAMSON AND CRETE: FOOD AND DENSITY OF COYOTES

231

TABLE 1. Comparison of the summer diet of Coyotes in boreal forests of the Gaspé peninsula (Québec) when the popula-
tion was expanding (1988), and declining following colonization (1991). Diet was estimated as percent volume (%) of
food items in scats. Monthly comparisons were tested with the Mann-Whitney U test; n = the number of scats examined.

June
1988 1991
(n=15) (n=29)
Moose 0.0** 28.0**
Deer 16.3* 1.8*
Snowshoe Hare 13.3 52-3
Woodchuck 35.75* ed aks
Beaver 0.0* | LP ARS fit
Small mammals 17.0 18.0
Other mammals 0.0 0.0
Birds 6.7 pF
Berries 1.7 0.0
Vegetation Pied OY ai

NO, **P < 0.05.

Moose and Deer were probably consumed as carrion
in May-June, as all hairs found in the scats were
from adults. Remains of large mammals that died
during winter are available to, and often used by,
Coyotes in spring (Hawthorne 1972; Parker 1986;
Reichel 1991). Parker (1986) hypothesized that
Coyotes have less difficulty catching Woodchucks in
May-June compared to other seasons because
Woodchucks are particularly vulnerable when forag-
ing for new vegetation growth; our results for 1988
support this view. The same may also apply to
Snowshoe Hares. Indeed, Snowshoe Hares may be
more easily found by predators in May because their
white winter coat is not fully replaced and the snow
has almost completey melted (Bittner and Rongstad
1982). Also, young Woodchucks begin to emerge

July August
1988 1991 1988 199]
(n=50) (n=125) (n=166) (n=74)
6.3 a8) i) L9
ISP a OOs> 0.5 0.0
ole 10.5 2.4 pe
21 .0F UES? a6" 0.4*
0.0 0.0 0.0 0.0
16.8 19.1 14.2 941
a7 3.7 0.0 L2
1.6 2.4 3d 2.4
14.6** 49.5** Sy le IDS**
12.8* 6.4* L2S IRS ha

from their natal dens in early summer when young
hares become available (Lee and Funderburg 1982;
Bittner and Rongstad 1982), and their inexperience
leaves them vulnerable to predation.

In our study area, berries were most available
from late July to early October, a time when Coyotes
made a large use of that food source. In fact, the use
of berries by Coyotes was more pronounced in our
study area than in other regions of North America
(Table 2). Although study periods differed in the
comparisons, each always included at least the peak
of berry ripening in July-August. In our study, the
volume of berries in scats averaged 24% during June
to August 1988, and 40% during May to October
1991. In comparison, berries represented less than
1% of the volume in most other studies (Table 2),

TABLE 2. Food habits (% volume) of Coyotes during summer in various regions of North America.

Items
Area Habitat type Animal Berries Period Source
Québec Boreal forests 64 24 June-August 1988 Present study
58 40 May-October 1991

Québec Mixed northern hardwood- 95 <1 May-October Fortin (1995)

coniferous forests*
Québec Mixed northern hardwood- 78 14 July-August Messier et al. (1986)

coniferous forests*
Alberta Boreal forests? 84 1 June-September Todd et al. (1981)
California Chapparal 80 a July-September Hawthorne (1972)
California‘ Chapparal-grassland = ie) l Summer‘ Ferrel et al. (1953)
Idaho Grassland 93 1 July-October Johnson and Hansen (1979)
Idaho Grassland 71 - July-August MacCraken and Hansen (1987)
Oklahoma Grassland-woodland 65 - July-August Holle (1978)
Central and Mostly grassland =| 4 July-August Young and Jackson (1951)

Western U.S.A.°

“with farmlands

>not specified but apparently not used.
“stomach contents.

4No precise date specified.

Ze

except in southern Québec where volume averaged
14% in July-August (Messier et al. 1986). Other
studies also concluded that berries were an important
part of the Coyote diet during summer (Barett 1983;
Harrisson and Harrisson 1984; Parker 1986; Andelt
et al. 1987; Litvaitis and Harrisson 1989; Dibello et
al. 1990; Riechel 1991; McClure et al. 1995).
However, those studies used the frequency of occur-
rence rather than volume to describe Coyote food
habits. The use of small food items, such as berries,
are often overestimated by frequency of occurrence
(Weaver and Hoffman 1979).

The high use of berries, especially in 1991, may
have results from a high availability of this food
source. Many scats were collected along logging
roads in clear-cuts where berries were common.
However, as White-tailed Deer, Snowshoe Hare and
possibly Woodchuck and Beavers were less abun-
dant in 1991, it is also possible that Coyotes con-
sumed more berries to compensate for the low num-
ber of animal prey. Indeed, some authors suggest
that rodents and lagomorphs are preferred by
Coyotes and that other items are used only when
those prey are not available (Todd et al. 1981;
Barrett 1983; Cypher et al. 1993; Hernandez and
Delibes 1994).

If food habits and low population density were
due to a scarcity of animal prey in our study area,
then the low abundance of prey may also explain
why the fecundity of female Coyotes is exceptional-
ly low on the Gaspé peninsula (Poulle et al. 1995).
In Alberta, fecundity of Coyotes was correlated
with abundance of hares (Todd et al. 1981). The
fecundity of females Red Fox (Vulpes vulpes), also
appeared to be influenced largely by the availability
of prey (Lindstr6ém 1988). If such is true for
Coyotes, it might suggest that a high consumption
of berries, containing mostly carbohydrates, may
not compensate for required lipids and proteins that
Coyotes obtain from animal prey, thus influencing
reproduction. This hypothesis is based on the
assumption that foraging activities in summer and
early fall influence female reproduction during the
next spring.

Acknowledgments

We are particularly indebted to R. Lemieux who
was responsible for the field work and scat analysis.
R. Faubert, S. Lariviére and B. Picard participated in
the field work. Personnel of the Ministére de
Environnement et de la Faune in Sainte-Anne-des-
Monts always provided quick assistance with the
logistics. N. Lemay performed some of the statistical
analyses on Coyote diet as part of an undergraduate
course at Université du Québec a Rimouski. D.
Murray, G. R. Parker and an anonymous person
kindly reviewed an earlier draft of this manuscript.
This study was supported by the Ministére de
Environnement et de la Faune du Québec.

THE CANADIAN FIELD-NATURALIST

Vol. 111

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Received 26 February 1996
Accepted 4 October 1996

Egg Composition and Post-DDT Eggshell Thickness of the American
White Pelican, Pelecanus erythrorhyncos

SHAWN C. BUGDEN and ROGER M. EVANS

Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2

Bugden, Shawn C., and Roger M. Evans. 1997. Egg composition and post-DDT eggshell thickness of the American White
Pelican, Pelecanus erythrorhyncos. Canadian Field-Naturalist 1 11(2): 234-237.

Eggs of the American White Pelican (Pelecanus erythrorhyncus) were collected from a breeding colony at East Shoal
Lake, Manitoba. Egg contents were 17.5 + 0.4 percent yolk, had an energetic density of 4.77 + 0.06 kJ/g and contained
82.8 + 0.2 percent water. These results suggest a typically altricial composition for the eggs of the American White Pelican
and strengthen the evidence that this is true of the family Pelecanidae in general. After thinning significantly in the prairie
region during the late 1960s, eggshell thickness of the American White Pelican in Manitoba has recovered. Eggshell thick-
ness (0.685 + 0.018mm) and eggshell mass (16.54 + 0.55 g) were not significantly different from pre-DDT (prior to 1947)
measurements reported in the literature (0.683 mm and 16.56 g).

Key Words: American White Pelican, Pelecanus erythrorhynchos, egg composition, eggshell thickness, DDT.

The relative yolk content and energetic density
of eggs correlate well with the position of a species
on the altricial to precocial continuum of develop-
mental maturity at hatch (Nice 1962; Carey et al.
1980). Pelagic-feeding Procellariiformes, offshore-
feeding terns with their long incubation periods
(Carey et al. 1980; Sotherland and Rahn 1987), and
some Columbidae may represent significant excep-
tions to the general trends (Ricklefs 1977; Carey et
al. 1980). Two members of the altricial family
Pelecanidae have been studied. The Great White
Pelican (Pelecanus onocrotalus) produces eggs that
are typical of most altricial species (Jones 1979;
Williams et al. 1982). Eggs of the Brown Pelican
(Pelecanus occidentalis) (Lawrence and Schreiber
1974) were found to deviate from the typical com-
position of an altricial egg with energetic densities
similar to semi-precocial species (Ricklefs 1977;
Carey et al. 1980). However, some of the Brown
Pelican eggs studied (3 of 6) had been incubated for
up to three weeks (Lawrence and Schrieber 1974),
which may have significantly influenced the results
(Ricklefs 1977; Jones 1979). To assess further egg
composition in the family Pelecanidae, we exam-
ined a third species, the American White Pelican
(Pelecanus erythrorhyncos).

Prior to the widespread use of DDT (pre-1947),
eggshells of American White Pelicans collected in
the interior of North America had an average thick-
ness of 0.683 mm with 95% confidence limits of
0.673 - 0.693 mm and an average dry eggshell mass
of 16.56 g (Anderson and Hickey 1972). By 1952,
eggs collected in Manitoba showed some evidence
of thinning with an average thickness of 0.654 mm
with 95% confidence limits of 0.628-0.680
(Anderson and Hickey 1972). Eggs collected in the
prairie states and provinces in 1965 were significant-
ly thinner than pre-DDT eggshells with an average

thickness of 0.655 mm with 95% confidence limits
of 0.643 - 0.667 mm (Anderson et al. 1969). Another
sample of pelican eggs from the interior of North
America collected in 1965 to 1968 showed a signifi-
cant 5% decline in eggshell thickness with an aver-
age of 0.647 mm and 95% confidence intervals of
0.621 - 0.673 mm (Anderson and Hickey 1972). The
dry eggshell mass of these eggs also showed a
decline, with a 9% drop from pre-DDT measure-
ments (Anderson and Hickey 1972).

Eggshell thinning was even more pronounced in
eggs collected from other areas of North America
(Anderson and Hickey 1972). Eggs collected in the
late 1960s in California and British Columbia
showed declines of 8 and 13 percent from pre-1947
measurements. In 1972 an average eggshell thick-
ness of 0.620 + 0.0063 mm was found in eggs col-
lected from Gunnison Island, Great Salt Lake, Utah
(Knopf and Street 1974). Eggshells in 1965 and
1972 were significantly thinner than the pre-1940
samples and inversely related to the level of the
DDT metabolite DDE found in pelicans (Anderson
et al. 1969; Knopf and Street 1974). The widespread
usage of DDT from 1943 to 1971 and resulting
bioaccumulation in fish-eating birds like pelicans is
believed to have contributed to eggshell thinning
(Hickey and Anderson 1968; Anderson et al. 1969).
The only reported post-DDT (post-1971) measure-
ments of eggshell thickness in the American White
Pelican were from eggs collected in the Klamath
region in California (Boellstorff et al. 1985; Evans
and Knopf 1993). These eggs showed an increase in
shell thickness from 1969 (15% below pre-1947
thickness) but were still 10% below pre-1947 values
in 1977 and 1981 (Boellstorff et al. 1985). We
obtained current eggshell measurements for compar-
ison with measurements taken prior to and during the
widespread usage of DDT.

234

Loo7

Methods

Ten freshly laid eggs were collected in the spring
of 1994 from an American White Pelican breeding
colony at East Shoal Lake, Manitoba. Eggs were
removed from a single subcolony (Knopf 1979) to
minimize disturbance to the rest of the colony
(Boellstorff et al. 1988; Evans and Knopf 1993).
Prior observations from a protected tunnel blind
made it possible to identify and take eggs of known
chronology during a brief (10-min) visit to a single
edge subcolony, thereby reducing the amount of dis-
turbance. Egg length and breadth were measured
with vernier calipers to the nearest 0.1 mm. Egg vol-
ume was calculated using the formula, V = KLB?
where L is the egg length, B is the egg breadth and K
is the species constant of 0.500 (O’ Malley and Evans
1980). Egg volume can also be estimated by the dif-
ference between the mass of the egg in air (W,) and
its mass when submerged in water (W,). The density
of water was assumed to be 1.00 g/cm’. Initial
weights of the whole egg were made in both air and
water to the nearest 0.1 g. Specific gravity was cal-
culated from the formula, (W, / (W, - W,) (Evans
1969). The eggs were then boiled for 10 minutes,
allowed to cool to room temperature, placed in air
tight containers and then frozen for later analysis.

Eggs were thawed, separated into components, and
weighed to the nearest 0.01 g using a Mettler bal-
ance. Any loss in mass from the boiling and freezing
process was attributed to albumen water loss and cor-
rected by adding the difference between the fresh
whole egg and the thawed egg to the wet albumen
mass (Williams et al. 1982; Meathrel et al. 1987).
The components were dried for 72 hours at 60°C and
then weighed for dry mass. Shell thickness, including
membranes, was measured (+0.001 mm) using a
modified Starrett Model 1010M micrometer. Mean
shell-thickness was obtained by averaging eight lat-
eral readings, taken at the egg’s equator, midway
between the caps (Knopf and Street 1974).

Dried yolk and albumen were ground to a uniform
powder. Samples of the albumen and the yolk of
each egg were burned in a bomb calorimeter (Parr
1241 Calorimeter, Parr Instrument Co., Moline, III.)
to determine their energy content. Two samples each

BUGDEN AND EVANS: EGG COMPOSITION AND EGGSHELL THICKNESS

200

of yolk and albumen were tested and the average
result was taken. Additional aliquots of yolk and
albumen were used to determine the lipid content.
Lipid content was determined by Soxhlet extraction
using petroleum ether (Arnold 1992) for eight hours
for yolk samples and four hours for albumen sam-
ples. Samples were run in duplicate with the average
result taken. The energetic content of the egg was
then calculated indirectly assuming 40 kJ/g for lipids
and 23.5 kJ/g for the non-lipid fraction of the egg
contents (Sotherland and Rahn 1987). All means are
presented + 1 SE.

Results

Fresh pelican eggs had a mass of 156.2 + 3.8 g
and had a length of 87.8 + 1.1 mm and breadth of
56.9 + 0.7 mm. Calculation of egg volume based on
length and breadth gave a value of 142.4 + 3.6 cm?
which was not significantly different from egg vol-
ume based on (W, - W,) of 140.5 + 3.2 cm? (paired
t= 1.84, P> 0.05, df= 9). Specific gravity was 1.110
+ (0.005. The wet and dry mass of egg components
are shown in Table 1. The fraction of the yolk in the
contents was 0.175 + 0.004. Egg shell thickness was
0.685 + 0.018 mm.

The yolk made up 17.5 + 0.4 percent of the egg
contents. Dry yolks contained 61.7 + 0.5% lipid and
were shown by bomb calorimetry to contain 32.9 +
0.1 kJ/g (dry mass). Dry albumen contained 0.3 +
0.06% lipid and had an energetic content of 21.4 +
0.04 kJ/g (dry mass). The overall energy of the dry
egg contents was 27.7 + 0.1 kJ/g. The energy of the
wet contents was compared with other members of
the family Pelecanidae (Table 2). The energy of the
wet contents was also estimated based on the per-
centage of lipid in the contents (5.1 + 0.1) and litera-
ture values for the energy of lipid and non-lipid frac-
tions of egg contents (Sotherland and Rahn 1987).
This value, 4.73 + 0.06 kJ/g was very similar to the
energetic value based on bomb calorimetry (Table 2).

Discussion

Eggs did not differ significantly in size (length,
breadth and mass) from fresh eggs measured in previ-
ous studies in Manitoba (P > 0.50) (Evans 1969;

TABLE 1. Mass of fresh American White Pelican egg components.

Wet Mass (g)

Component

Albumen 114.14 + 3.16

Yolk 24.09 + 0.46

Contents 138.23 + 3.41

Shell 17.92 + 0.62

Total egg 156.2 + 3.7 100

Mean + SE %

donk
15.4

88.5
ELS

Water! Dry Mass (g)

% Mean + SE %
89.2 12.38 + 0.45 30.7
52.6 11.41 + 0.69 28.3
82.8 23.79 + 0.58 59.0

7.6 16.54 + 0.55 41.0
74.2 40.33 + 0.91 100

'Water as a percentage of wet mass of the component; other percentages are in relation to total

egg mass.

236

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 2. Energy and composition of egg contents in the family Pelecanidae.

Species kJ/g
Pelecanus erythrorhynchos 4.77
Pelecanus occidentalis 6.22
Pelecanus onocrotalus 5.16
Pelecanus onocrotalus 4.49

Yolk/Albumen

Ratio N Ref.
0.21 10 l
0.40 6 2
0.26 54 3
O17 1 4

References: 1. This study; 2. Lawrence and Schrieber (1974);
3. Jones (1979); 4. Williams et al. (1982).

“Energy value was based on bomb calorimetry.

O’Malley and Evans 1980). As eggs lose water at a
relatively constant rate during incubation, the specific
gravity of eggs can be used to estimate the time
elapsed since laying (Westerkov 1950). Specific
gravity was not significantly different than that of 21
freshly laid eggs in a previous study (t = 0.65,
P > 0.50, df = 29) (O’ Malley and Evans 1980).

The water content of the fresh bird egg typically
correlates with the state of hatching maturity of the
chick (Sotherland and Rahn 1987). The fraction of
water in the contents of altricial American White
Pelican eggs (Table 1) is similar to the altricial
species average of 84.3% (Carey et al. 1980). The
fraction of water in the albumen (Table 1) is relative-
ly uniform (85-90%) among all species of birds
(Ricklefs 1977; Sotherland and Rahn 1987). The
water content of American White Pelican egg yolk
(Table 1) fell within, but near the low end of, the
altricial range (52.4% to 65.5%) (Sotherland and
Rahn 1987).

Calculations of energetic density based on bomb
calorimetry and lipid extraction produced very simi-
lar results. The similarity of the results supports the
use of either bomb calorimetry or lipid extraction for
estimation of the energy content of eggs. The energy
of the dry contents of eggs is relatively uniform in all
species of birds (Carey et al. 1980; Ar et al. 1987).
Our results put the American White Pelican (27.7
kJ/g) within one standard deviation of Sotherland and
Rahn’s (1987) average energy density of 29.1 +
1.5(SD) for 99 species of birds.

The energy of the wet contents of avian eggs typi-
cally correlates with the hatchling’s maturity (Ar et
al. 1987; Sotherland and Rahn 1987). The energetic
density of the contents of the eggs of the American
White Pelican and the Great White Pelican (Table 2)
fall within the 95% confidence interval (4.771 -
5.193) of the average (4.982 kJ/g) reported for other
altricial species (Ar et al. 1987). The energetic densi-
ty previously reported for Brown Pelican eggs (Table
2) falls well outside of the normal range for altricial
species. The high energetic density and the low frac-
tion of water in egg contents (76.7%) reported for the
Brown Pelican probably reflects the effects of embry-
onic development and water loss associated with eggs

that had been incubated for up to three weeks
(Lawrence and Schreiber 1974; Jones 1979; Williams
et al. 1982). Analyses of eggs of the Great White
Pelican (Jones 1979; Williams et al. 1982) and the
American White Pelican (this study) support the con-
clusion that birds of the family Pelecanidae produce
eggs typical of altricial birds.

Eggshell thickness significantly increased from
measurements taken in 1972 (Utah), at the end of
almost 30 years of DDT usage (t = 3.16, P< 0.01,
df = 108) (Knopf and Street 1974). Our eggshell
thickness measurements were also significantly
greater (t= 3.54, P< 0.001, df = 79) than measure-
ments taken in 1981 (California; Boellstorff et al.
1985). Western populations were more affected by
DDT than interior populations (Anderson and Hickey
1972). Although prairie populations experienced a
smaller decline (5% in late 1960s; Anderson and
Hickey 1972), our results indicate that eggshell thick-
ness in this population returned to pre-DDT levels.
Present day post-DDT eggshell thickness (t = 0.12,
P>0.50, df = 100) and eggshell mass (t = 0.04,
P>0.50, df = 102) were not significantly different
from measurements taken from prairie populations in
the pre-DDT period prior to 1947 (Anderson and
Hickey 1972). Although a direct link between
eggshell thinning and reproductive success in
American White Pelicans has not been established
(Anderson et al. 1969; McEwen and Stephenson
1979) it is comforting to report that with the recovery
of eggshell thickness in the sampled population this
potential problem has been avoided.

Acknowledgments

We thank K. L. Campbell, R. A. MacArthur and
J.P. Ryder for their advice and assistance. D. W.
Anderson and P. R. Sotherland provided helpful com-
ments on the manuscript. This study was supported
financially by an operating grant to R. M. E. from the
Natural Sciences and Engineering Research Council,
Ottawa, Canada.

Literature Cited
Anderson, D. W., J. J. Hickey, R. W. Risebrough, D. F.
Hughes, and R. E. Christensen. 1969. Significance of

1997

chlorinated hydrocarbon residues to breeding pelicans
and cormorants. Canadian Field-Naturalist 83: 91-112.

Anderson, D. W., and J. J. Hickey. 1972. Eggshell
changes in certain North American birds. Pages 514-540
in Proceedings of the XVth International Ornithological
Congress. Edited by K. H. Voous. E. J. Brill, Leiden,
The Netherlands.

Ar, A., B. Arieli, A. Belinsky, and Y. Yom-Tov. 1987.
Energy in avian eggs and hatchlings: Utilization and
transfer. Journal of Experimental Zoology Suppplement
1: 151-164.

Arnold, T. W. 1992. Variation in laying date, clutch size,
egg size, and egg composition of yellow-headed black-
birds (Xanthocephalus xanthocephalus): a supplemental
feeding experiment. Canadian Journal of Zoology 70:
1904-1911.

Boellstorff, D. E., H. J. Ohlendorf, D. W. Anderson,
E. J. O'Neill, J. O. Keith, and R. M. Prouty. 1985.
Organochlorine chemical residues in white pelican and
western grebes from the Klamath Basin, California.
Archives of Environmental Contamination and
Toxicology 14: 485-493.

Boellstorff, D. E., D. W. Anderson, H. M. Ohlendorf,
and E. J. O’Neill. 1988. Reproductive effects of nest-
marking studies in an American White Pelican colony.
Colonial Waterbirds 11: 215-219.

Carey, C., H. Rahn, and P. Parisi. 1980. Calories, water,
lipid and yolk in avian eggs. Condor 82: 335-343.

Evans, R. M. 1969. Specific gravity of White Pelican eggs.
Auk 86: 560-561.

Evans, R. M., and F. L. Knopf. 1993. American White
Pelican (Pelecanus erythrorhynchos). In The Birds of
North America, Number 57. Edited by A. Poole and F.
Gill. The Academy of Natural Sciences, Philadelphia;
The American Ornithologists’ Union, Washington,
Die.

Hickey, J. J.. and D. W. Anderson. 1968. Chlorinated
hydrocarbons and eggshell changes in raptorial and fish-
eating birds. Science 162: 271-273.

BUGDEN AND EVANS: EGG COMPOSITION AND EGGSHELL THICKNESS

26

Jones, P. J. 1979. Variability of egg size and composition
in the Great White Pelican (Pelecanus onocrotalus).
Auk 96: 407-408.

Knopf, F. L. 1979. Spatial and temporal aspects of colonial
nesting of White Pelicans. Condor 81: 353-363.

Knopf, F. L., and J. C. Street. 1974. Insecticide residues
in White Pelican eggs from Utah. Wilson Bulletin 86:
428-434.

Lawrence, J. M., and R. W. Schreiber. 1974. Organic
material and calories in the egg of the Brown Pelican,
Pelecanus occidentalis. Comparative Biochemistry and
Physiology 47A: 435-440.

McEwen, F. L., and G. R. Stephenson. 1979. The use and
significance of pesticides in the environment. John
Wiley and Sons, New York.

Meathrel, C. E., J. P. Ryder, and B. M. Termaat. 1987.
Size and composition of Herring Gull eggs: Relationship
to position in the laying sequence and the body condition
of females. Colonial Waterbirds 10: 55-63.

Nice, M. M. 1962. Development of behavior in precocial
birds. Transactions of the Linnaean Society of New
York 8: 1-211.

O’Malley, J. B. E., and R. M. Evans. 1980. Variations in
the measurements among white pelican eggs and their
use as a hatch date predictor. Canadian Journal of
Zoology 58: 603-608.

Ricklefs, R. E. 1977. Composition of eggs of several bird
species. Auk 94: 350-356.

Sotherland, P. R., and H. Rahn. 1987. On the composition
of bird eggs. Condor 89: 48-65.

Westerkov, K. 1950. Methods of determining the age of
game bird eggs. Journal of Wildlife Management 14:
56-67.

Williams, A. J., W. R. Siegfried, and J. Cooper. 1982.
Egg composition and hatchling precocity in seabirds.
Ibis 124: 456-470.

Received 15 April 1996
Accepted 16 October 1996

Population Growth of Moose, Alces alces, in Labrador

Tony E. Cuupss! and JAMES A. SCHAEFER2

'Department of National Defence, 5 Wing Goose Bay, Station A, Happy Valley-Goose Bay, Labrador, Newfoundland
AOP 1SO

"Department of Forest Resources and Agrifoods, Newfoundland and Labrador Wildlife Division, Box 3014, Station B,
Happy Valley-Goose Bay, Labrador, Newfoundland AOP 1EO

Chubbs, Tony E., and James A. Schaefer. 1997. Population growth of Moose, Alces alces, in Labrador. Canadian Field-
Naturalist 111(2): 238-242.

Moose (Alces alces), although not historically resident in Labrador, have increased in numbers and range over the past four
decades. Densities of Moose remain comparatively low (0.013 - 0.168 animals/km7), but several indicators reveal an
expanding population. Successive surveys of two Moose management areas in south-central Labrador, conducted in 1986
and again in 1994, indicated an average finite rate of increase of 1.10. Similarly, calf:cow ratios are relatively high, ranging
from 0.30 to 0.67. Twinning rates were estimated at less than 11% of the total calf crop. During late winter animals
appeared to be non-gregarious with mean group size of 1.4. Moose hunting in Labrador commenced in 1977; hunter suc-
cess, as indicated by annual license returns, increased substantially during the period 1977-1993. Incidental observations

have indicated northward range expansion of Moose to the treeline in Labrador.

Key Words: Moose, Alces alces, population growth, density, winter range, Labrador.

In Labrador, the first Moose sightings occurred
during the early 1950’s (Harper 1961) in the south-
west near Ashuanipi Lake and are independent of
the introduction attempts in southeastern Labrador
near St. Lewis River in 1953 (Figure 1). By 1953,
Moose were already present in southwestern
Labrador (Mercer and Kitchen 1968). Range expan-
sion into Labrador appears to have occurred
through natural dispersion from Québec in a north-
easterly direction along river valleys (Mercer and
Kitchen 1968). By 1977, Moose had populated
south-central Labrador in numbers sufficient to
warrant the establishment of 13 management areas
and an annual sport hunt. More recently, in western
Labrador, an additional management area has been
established. Very low Moose densities and large
management areas precluded reliable systematic
counts prior to 1986.

Although anecdotal observations of Moose in
Labrador are strongly suggestive of a growing popu-
lation, there has been little formal analysis of the
extent and magnitude of this trend. Here we use a
combination of sources to document demographic
changes in Moose. First, we compared successive
surveys in two Moose Management Areas (MMAs),
conducted in 1986 and repeated in 1994. For region-
al comparisons, we summarized density estimates of
four other MMA’s. Second, we used hunter statis-
tics as an indirect index on population structure and
trends (Fryxell et al. 1988). The harvest of Moose in
Labrador has been well documented through licence
returns and telephone inquiries since the initial open
season in 1977. Finally, we collated information on
Moose sightings to determine the current northern
limit of the species in the region.

Study Area

Successive aerial surveys were conducted in two
MMaAs within the High Boreal Forest-Lake Melville
ecoregion (Lopoukhine et al. 1977; Meades 1990*),
consisting of taiga, subalpine tundra, and the transi-
tional vegetation types. Stands of Black Spruce
(Picea mariana) and Balsam Fir (Abies balsamea)
occur on shallow upland soils whereas open spruce
lichen forest can be found on upland till and out-
wash soils. White Birch (Betula papyrifera) and
Trembling Aspen (Populus tremuloides) occur on
well-drained alluvium soil. Mosses of the genus
Sphagnum dominate bogs. Common associates
included Labrador Tea (Ledum groenlandicum),
Sheep Laurel (Kalmia angustifolia), Bunchberry
(Cornus canadensis), Partridgeberry (Vaccinium
vitis-idea) and Low Sweet Blueberry (Vaccinium
angustifolium). Forest harvesting has continued in
the Muskrat Falls area since the 1950s, increasing
human access and expanding areas of successional
browse species preferred by Moose including Alder
(Alnus rugosa), White Birch, Willow (Salix sp) and
Balsam Popular (Populus balsamea). Approximately
27% of the Muskrat Falls area (Area 53; Figure 1) is
composed of deciduous species in various stages of
regeneration. The Grand Lake area (Area 54; Figure
1) has some commercial forestry harvest and signifi-
cant burned areas resulting in nearly 7% deciduous
regeneration.

This region has long, severe winters with heavy
snow accumulation and short summers. Snow nor-

*See Documents Cited between Acknowledgements and
Literature Cited.

238

1997

CHUBBS AND SCHAEFER: POPULATION GROWTH OF MOOSE

239

;

Sear \

=

__--77” “Approximate northern ay (

~Okak Bay

ie

—

====—5 ae Hebron Fiord
core o i

<->

0 100 km

x Ne Area 54
0.085/km
ya
— saan (1994) et
rea > Lee
0.077/km _/ Lake ‘a
(1995) Melville ee
Area's4 Area 53
0.017/km* 0.168/km
(1986) Area 52 5 (1994) prea 58 5
@ fee an ann St. Lewis
Ashwanipi : \ : |
Lake ee | 4
Québec cat Sa,
yes We =

FIGURE 1. Moose Management Areas in central Labrador indicating densities and year of
most recent survey. Extreme northern sightings of Moose or their sign are indicated

by open squares.

mally begins in October and may persist into June.
Mean daily temperatures range from -14 to -18°C in
February and 13°C in July. Average annual precipi-
tation is 1100 mm with an average snowfall accumu-
lation of 4 m.

Besides man, potential predators of Moose in the
study area include Wolves (Canis lupus), Black
Bears (Ursus americanus), and possibly Lynx (Lynx
canadensis).

Methods

During 1994 and 1995, population estimates were
conducted using stratified random sampling
(Gasaway et al. 1986). To estimate population
growth, we compared Moose densities in two MMAs
surveyed in 1986 and again in 1994 (Areas 53 and
54, Figure 1). We delineated all possible subunits on
1:50 000 scale topographic maps of the management
areas. Subunits (4 km?) were defined by Universal
Transverse Mercator grid lines on topographic maps.
The survey area was then stratified into “high” and
“low” density strata using Forestry Inventory Maps

and incidental Moose observations during prelimi-
nary flights to determine preferred habitat. Flights
were conducted using a Bell 206L helicopter at alti-
tudes ranging from 30 to 50 m agl at 50 to 100 km/h.
Calf:cow ratios and population sex and age structure
were obtained during aerial counts. Sex was deter-
mined by the presence or absence of a vulva patch
(Mitchell 1970), antlers or a calf.

During 8 to 12 March 1994, we conducted a strati-
fied random block survey to estimate Moose popula-
tion parameters in MMA’s 53 (Muskrat Falls) and 54
(Grand Lake). We estimated Moose populations and
quantified the rate of increase since the last census.
Both MMA’s share an adjoining boundary which has
changed slightly since 1986 but both encompass the
same area censused in 1986. To enable between sur-
vey comparisons, we examined changes in Moose
density and sex and age structure.

Areas of high Moose density generally included
riverine habitat, regenerated clear cuts and deciduous
forest. The low density stratum comprised areas of
high elevation, closed canopy mature Black Spruce

240

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE |. Moose Sex and Age Composition within two Moose Management Areas (MMA’s) in Labrador dur-

ing 11-12 March, 1994.

Adult sex ratio

MMA n bulls:cows
Grand Lake a 47:53
Muskrat Falls 79 40:60

forests, recent burns and barrens. Both strata were
assigned identification codes and individually num-
bered subunits were randomly selected for sam-
pling. A sightability correction factor (SCF) was
calculated using five high density subunits in the
Grand Lake MMA. We performed a standard search
effort (2-4 min/km7) to locate Moose followed by an
intensive search (5 to 8 min/km7?) the following day.
We determined a SCF of 1.6 for high density strata.
This is slightly lower than correction factor of 2.3
used on the west coast of Newfoundland (Albright
and Keith 1987) where Moose densities were rela-
tively high and cover dense. We believe that all
Moose were located in low density strata due to the
openness of the terrain and thus no correction factor
was applied. Since both management areas were
similar in strata composition, we used the same
sightability correction factor and sampling variance
for both areas.

The Grand Lake MMA has a total area of 3797
km’, of which 3732 km? was considered possible
Moose habitat. We delineated 264 km? (7%) as
high density stratum and 3468 km? (93%) as low
density stratum. Our survey encompassed 26 (39%)
of 66 subunits of the high density stratum and 39
(4%) of 975 low density subunits. The Muskrat
Falls MMA has a total area of 4021 km? of which
3872 km? was considered possible Moose habitat.
We delineated 800 km? (21%) as high density stra-
tum and 3072 km? (79%) as low density stratum.
We surveyed 43 (22%) of 200 subunits of the high
density stratum and 15 (2%) of 768 subunits in the
low density stratum.

Excellent weather conditions prevailed throughout
the entire survey period. Heavy snowfall in late
February until 7 March effectively covered old
tracks, enabling an efficient search effort. Days were
sunny and bright with temperatures ranging from -2°
to -15°C and winds were generally northwesterly
ranging from 0 to 15 km/hr. With constant weather
conditions, movement of Moose between strata
appeared minimal during the survey.

Hunter success rate was determined annually
using licence and mandible returns complemented by
intensive telephone inquiries. Outcomes of all
licences were accounted for each year. As an addi-
tional indication of population growth we used
regression analysis to correlate hunter success with
time during 1977-1993.

Percent of population

bulls cows calves unknown
35 38 12 15
25 38 25 ]

In addition, we summarized Moose densities from
the results of occasional aerial surveys conducted
between 1991 to 1995 (Newfoundland and Labrador
Wildlife Division, unpublished data) in six MMAs
(Figure 1). Prior to 1994, MMAs were surveyed
using stratified line transect surveys.

Results and Discussion
Distribution

Historically, Moose have been distributed more
southerly along the Labrador Ungava Peninsula.
Northern limits in Québec appeared to be along 53°E
latitude in the early 1950s (Brassard et al. 1974)
which corresponds to the northernmost observations
of Moose in the Lake Melville region of Labrador
(Mercer and Kitchen 1968). Range expansion in
Québec reached the 57° in the late 1970s (Brassard
et al. 1974). Recent observations in Labrador con-
firm a similar northward range extension north of
58° (Figure 1).

By 1980, Moose had been observed as far north as
Nain (F. Phillips, personal communication). Most
recently, independent sightings of two Moose during
fall 1995 (1 adult male; | unknown sex) have been
made as far north as Ikinet River (57°24’; 62°32’),
west of Okak Bay (J. Ennis, personal communication).
Evidence of browsing on willows (Salix spp.) and
fecal droppings were observed in October 1995, south
of the Ikarut River (58°04'; 63°22’) northwest of
Hebron Fiord (G. Baikie, personal communication).

This indicates a rate of spread of 8 km/yr since
1980, slightly lower than the 10 km/yr spread rate
initially reported for Moose in Labrador (Mercer and
Kitchen 1968). Similarly, Pimlott (1953) reported
that Moose introduced into Newfoundland spread at
a rate of 11 km/yr. Nevertheless, because the current
extension has now reached the treeline, it appears
that Moose may have reached their northern most
geographic distribution in Labrador. Although
Moose occur in treeless, shrub-dominated, alpine
habitat in Alaska (Gasaway et al. 1986), the expan-
sion of Moose into the subarctic tundra of Labrador
is unlikely (Ballard et al. 1991; Doerr 1983).

Our observations revealed that Moose in the
region tended not to aggregate during winter (1.4
Moose/group) compared to other ranges where
groups ranged from 1.8 to 2.3 Moose/group (Rolley
and Keith 1980; Mytton and Keith 1981). Similarly,
Dalton (1986) recorded a mean group size of 1.3.

1997
90 r =0.20
p = 0.082
80
® e e
oy od e
=
© 70
72)
9)
®
8
60
”
=
®
€ 50
=5
a
40
30
aera So al Poe:
1980 1984 1988 1992
Year

FIGURE 2. Hunter success for Moose in Labrador, 1977-
1993.

Yarding was nonexistent (largest group was four ani-
mals), even though the population had increased sig-
nificantly, probably due to the low densities of
Moose and lack of winter browsing areas.

Population Growth

During 1994, in the Grand Lake MMA, we count-
ed 46 and 6 Moose in the high and low density strata
respectively. Our estimate indicated a Moose popu-
lation of 321 + 218 (90% CI) or 0.085 Moose/km?.
Dalton (1986) estimated the population at 245 or
0.043 Moose/km? (no CI provided). This indicates
an increasing population, growing at a rate of \ =
1.09 over 8 years. In the Muskrat Falls MMA, we
counted a total of 77 and 2 Moose in the high and
low density strata respectively. We estimated a
Moose population of 677 + 359 (90% CI) or 0.168
Moose/km?. Dalton (1986) estimated the population
at 398 or 0.074 Moose/km? (no CI provided). This
population had increased over 8 years with \ = 1.11.

Moose densities in southern Labrador appear low
(0.013 - 0.168/km?) but comparable to analogous
ranges elsewhere in the north. Similar habitats in
north-central Québec produce 0.06 to 0.11
Moose/km? with densities exceeding 1.8 Moose/km?
in the most productive southern regions (Brassard et
al. 1974). Stenhouse et al. (1995) reported densities
of 0.155 Moose/km? in the Northwest Territories.
There is some evidence that the scope for further
increases in the Moose population are limited in
Labrador, possibly due to illegal harvests, wolf pre-
dation (Trimper et al. 1996), marginal habitat, or
some combination.

Population Structure
In the Grand Lake area, adult bull:cow ratio was

CHUBBS AND SCHAEFER: POPULATION GROWTH OF MOOSE

241

47:53, whereas in Muskrat Falls, where only male
licenses have been issued, the ratio was slightly
more in favour of females at 40:60 (Table 1).
Dalton’s (1986) bull:cow ratios of 43:57 and 42:58
for Muskrat Falls and Grand Lake respectively were
similar to those obtained in 1994. Dalton (1986) esti-
mated calves comprised 22% and 12% of the total
population in Grand Lake and Muskrat Falls respec-
tively. In 1994, calves comprised 12% and 25% of
the total population for Grand Lake and Muskrat
Falls respectively (Table 1). During the 1994 census
no evidence of twinning was observed in the Grand
Lake area while Muskrat Falls area had a low twin-
ning rate of 11%. These rates are much lower than
observed in the Northwest Territories (31%) where
Moose densities are similar (Stenhouse et al. 1995).
Our productivity is also much lower than that of
Moose populations across North America where
twinning rates average 33% (Boer 1992). Calf:cow
ratios were 0.48 and 0.24 in 1986 compared to 0.30
and 0.67 in 1994, for Grand Lake and Muskrat Falls
areas respectively. These results further imply that
continued growth of Labrador Moose populations is
likely limited.

Hunter Returns

Abundance estimates such as kills per day and
Moose seen per day have been shown to be valid
indices of Moose abundance provided that changes
in hunting pressure are also accounted for (Fryxell et
al. 1988). We used annual hunter success (kills per
licence) from 1977 to 1993 as an indicator of Moose
abundance. Hunter success rates ranged from a low
of 37% in 1977 to a high of 77% in 1993 (Figure 2).
Regression analysis indicated a moderate increase in
annual hunter success (7 = 0-20, p = 0.082).
Although licence quotas have increased slightly, the
increased success rate indicates that the Moose pop-
ulation has increased.

Acknowledgments

We would like to extend thanks to Conservation
Officers Wallace Lyall and Douglas Blake for their
participation as observers during the census, identi-
fying areas of winter Moose habitat and in the prepa-
ration of survey maps. J. Ennis and G. Baikie of
Universal Helicopters Ltd. provided detailed descrip-
tions of Moose sightings and browsing areas. Also,
we thank two anonymous reviewers for helpful com-
ments and suggestions. Funding for this project was
provided by the Monitoring and Mitigation Section
of the Department of National Defence and the
Newfoundland and Labrador Wildlife Division.

Documents Cited (marked * after date in text)

Meades, W. J. 1990. Natural regions of Newfoundland and
Labrador. Unpublished report submitted to the Protected
Areas Association, St. John’s, Newfoundland. 103

pages.

242

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Mitchell, H. B. 1970. Rapid aerial sexing of antlerless
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Moose populations near Rochester, Alberta, 1975-1978.
Canadian Field-Naturalist 95: 39-49.

Pimlott, D. H. 1953. Newfoundland Moose. Transactions
of the North American Wildlife Conference 18:
563-581.

Rolley, R. E., and L. B. Keith. 1980. Moose population
dynamics and winter habitat use at Rochester, Alberta,
1965-1979. Canadian Field-Naturalist 94: 9-18.

Stenhouse, G. B., P. B. Latour, L. Kutny, N. Maclean,
and G. Blover. 1995. Productivity, survival, and move-
ments of female Moose in a low-density population,
Northwest Territories, Canada. Arctic 48: 57-62.

Trimper, P. G., E. A. Young, and T. E. Chubbs. 1996.
Distribution of wintering Moose (Alces alces) in river
valleys of south-central Labrador and Northeastern
Québec. Alces 32: 1-9.

Received 22 April 1996
Accepted 20 October 1996

Resurgence of Breeding Merlins, Falco columbarius richardsonii,

in Saskatchewan Grasslands

C. STUART HousTon!, and KeIrH A. Hopson?

'863 University Drive, Saskatoon, Saskatchewan S7N 0J8
7#10, Site 16, RR #1, Telkwa, British Columbia VOJ 2X0

Houston, C. Stuart, and Keith A. Hodson. 1997. Resurgence of breeding Merlins, Falco columbarius richardsonii,
in Saskatchewan grasslands. Canadian Field-Naturalist 111(2): 243-248.

The Richardson’s Merlin (Falco columbarius richardsonii) almost disappeared from Saskatchewan grasslands in the early
1960s, coincident with extensive use of dieldrin for grasshopper control in adjacent grainfields. Merlins reappeared in
small numbers in 1975 and 1976 and gradually increased. Unlike the Merlins that have colonized cities such as Saskatoon
and prefer crow nests in conifers, the rural Merlins use magpie nests in deciduous trees. Productivity of four young per suc-

cessful nest is as high as reported anywhere.

Key Words: Merlin, Falco columbarius richardsonii, grasslands, Saskatchewan, dieldrin.

Richardson’s Merlin, Falco columbarius richard-
sonii, is distinct from other Merlins, since it alone
has five very wide white tail bands, paler plumage,
a longer wing chord, and a longer tail (Temple
1972). This subspecies suffered a drastic decline in
numbers across the prairies and virtually disap-
peared from the Kindersley area of Saskatchewan,
1962-1975 (Fox 1971). It is fortunate that subse-
quent Buteo studies have also been centered on the
Kindersley area, as this is the only locality on the
prairies where the decline and resurgence of the
Merlin are not purely anecdotal.

Since their resurgence, we have compared and
contrasted Merlins nesting in their “native” grass-
land habitat in thinly-populated rural areas, with the
new population of “city Merlins” studied so care-
fully since 1974 by Oliphant and associates in
Saskatoon.

When Bison (Bison bison) roamed the plains,
Merlins nested in partially wooded river valleys
and coulees of rough country, including the mar-
gins of the Cypress Hills and Wood Mountain,
Saskatchewan (Houston and Schmidt 1981;
Oliphant and Thompson 1979). In the first half of
this century, Merlins had large clutch sizes. W. J.
Brown at Lethbridge, Alberta, between 7 May and
5 June 1904 found eight Merlin nests; he took
seven sets of five eggs, and one of six eggs (Bent
1938). Fox (1971) reported that a small sample of
Merlins on the Great Plains prior to 1950 had 4.7
eggs per nest (n=10) and 4.3 young per nest
(m3).

*Dieldrin is the endo-exo isomer of 1,2,3,4,10,10-hex-
achloro-6,7-epoxy-1,4,4a,5,6,7,8,8a,octahydro-1,4-5,8-
dimethanonaphthalene (HEOD).

Many birders remarked that Merlins declined
noticeably in Saskatchewan grasslands in the early
1960s, a decline that was more severe than in adja-
cent Alberta. The decline was coincident with
extensive use of DDT across the continent and with
local use of dieldrin* for grasshopper control in
adjacent grainfields (Fyfe et al. 1976), but it also
coincided with breaking of much previous
grassland.

Decline and disappearance, 1958-1962

In grassland habitat near Kindersley, Saskatche-
wan, Merlin numbers seemed stable with six sites
occupied, some for two or three years, in 1958-
1960. In eight nests near Kindersley and two near
Battleford, followed through the season, during
what in retrospect was a period of declining pro-
ductivity, 45 eggs were laid (4.5 per nest), 27
hatched and 26 were fledged (2.6 young per suc-
cessful nest) (Fox 1964). By 1962, five of the six
previously occupied sites near Kindersley had been
deserted; these were in no-longer-occupied farm
shelterbelts where the majority of the trees had died
and there was a decrease in foliage of surviving
trees (Fox 1971). The sole surviving Merlin pair
near Kindersley in 1962 was the last to be recorded
for ten years. Hodson (1976) in 1972 found a pair
of Merlins with a failed nesting attempt 3 km north
of Marengo, 42 km west of Kindersley; egg analy-
sis showed persisting high levels (2.78 and 3.24
ppm) of dieldrin. That year he found another occu-
pied Merlin nest in a tributary of the Red Deer
River valley, barely within Saskatchewan, a mile
north-east of Empress, Alberta.

One of the avowed aims of Hodson’s thesis study
(1976) was “to probe the cause of the disappear-
ance of Merlins from the Kindersley area.” All for-
mer Merlin sites near Kindersley were checked by
Hodson (1976) in 1972 and by Fox and Hodson in

243

244

1974; these and other potentially suitable sites in
the area were also checked intermittently by
Richard Fyfe through 1975 (personal communica-
tion). All three observers failed to find a single
nesting pair (Hodson 1976).

In Alberta, with intensive search in 1971-1974,
Hodson’s active nests at Hanna numbered 19, 29,
18, and 25 per year, and in the river valleys north
and west of Medicine Hat, they numbered 9, 16, 10,
and 30, respectively; 98 of 709 eggs were taken for
analysis of pesticide levels. Hatching success for
the remaining eggs was only 57.8%; 84.4% of the
hatched young survived to fledging.

Quantification of pesticide use, 1958 - 1965

There are three complementary sources of infor-
mation about dieldrin usage. First, Robert V. Folker
(1970), wildlife ecologist with the Saskatchewan
Department of Natural Resources, did a study in
1959 in the Gravelbourg area of south-western
Saskatchewan, and sent a circular to 296 municipal-
ities of which 268 or 91% replied. That year 43
Gravelbourg farmers purchased 288 gallons of
dieldrin, an average of 6.7 gallons per farmer, and
sprayed 8205 acres with 260 gallons, very close to
the recommendation of | gallon per 32 acres (13
hectares). Of this spraying, 81% was done in June;
20% of the farmers sprayed grassland pasture as
well as crops. Province-wide, that year, 12 573
farmers purchased 69 584 gallons of dieldrin, about
5.5 gallons per farmer. [Gallons and acres are cited
unconverted as they were the units in use at the
time. Dieldrin in 1960 was delivered and sold in
five-gallon pails. Land in Saskatchewan is still
bought and sold in acres. ]

The second source is Hodson’s thesis (Hodson
1976), where Table 12 quantified the dieldrin pur-
chased in five villages and towns (Kindersley, Brock,
Smiley, Marengo and Glidden) on the western half of
the 1:250 000 Kindersley, Saskatchewan, map sheet
(Figure 1). Hodson postulated that “the heavy and

THE CANADIAN FIELD-NATURALIST

Vol. 111

widespread use of dieldrin is ... the factor which
rendered the coup de grace to the Merlins of the
Kindersley area.” In Alberta, Hodson found a high-
ly significant relationship between hatching failure
and levels of both DDE and dieldrin; unhatched
eggs from successful nests had 0.54 ppm of dield-
rin, whereas eggs from unsuccessful nests had 0.83
ppm (p < 0.02).

The annual reports of the Saskatchewan
Department of Agriculture from 1951 through 1965,
the third source, listed dieldrin gallonage supplied
to each rural municipality throughout one grass-
hopper outbreak and for three years thereafter
(Table 1). A small quantity, 500 gallons, of the new
chemical was used for grasshopper control in
Saskatchewan for the first time in 1951, near
Davidson, where the infestation was most severe.
For the next six years, grasshopper infestations
were minimal throughout the province, and dieldrin
was not required. Then, in 1958, 50 000 gallons
of dieldrin were used. In 1959, almost 70 000 gal-
lons were sprayed on about two million acres, with
residual chemical used the succeeding year. In
1961, the outbreak was unexpectedly severe,
requiring airlifts of dieldrin from Toronto; the total
distribution was 214 000 gallons. In 1962, about
100 000 gallons of dieldrin were used, estimated to
kill 95 to 98% of the grasshoppers. These govern-
ment figures of dieldrin usage, by municipalities,
correspond reasonably closely with Hodson’s fig-
ures of sales in the main village of each municipali-
ty. The department reported there were no deaths to
humans or to wildlife linked to this usage.

In 1962, sevin was available as an alternate treat-
ment, and in 1963, dimethoate as well. In 1965,
dieldrin was restricted to specified localities, so
very little was used. Milk and meat could not be
sold if livestock had been fed crops sprayed with
dieldrin. Dieldrin was banned in 1966.

In addition to dieldrin, Hodson incriminated

TABLE 1. Sales of dieldrin in five rural municipalities near Kindersley, 1958-1962 (in gallons). From Annual Reports,

Saskatchewan Department of Agriculture.

Price/. Kindersley _Prairiedale Hillsburgh Milton Newcombe Entire
gallon RM #290 RM #321 RM #289 RM#292 RM #260 Total Province
1958 $5.00 464 12 20 496 51 160
1959 $4.50 300 56 48 3352 300 1056 70 000
1960 $4.50 928 360 16 1196 120 2620 12 500 *
1961 $6.00 3195 980 594 2195 1650 8614 214 000
1962 $5.00 1365 600 550 1220 580 4315 100 000
1963 $6.00 22 360
1964 $6.00 10 281
1965 $6.00 740 restricted
Total 6252 1996 1208 4975 2670 17101

*= assuming equal quantities of dieldrin and heptachlor.
A gallon is 4.546 litres.

1997
l
o G
~+,s
o | 2
2
Bake
some (IS
~”
Cy
| op)
#321
Smiley
| #292 #290 #289
arengo| Kindersley} ack
Glidden
i S
#260
sr |
I t
110° 109°

HOUSTON AND HODSON: RESURGENCE OF MERLINS IN SASKATCHEWAN

Biggar e

245

Saskatoon

e Rosetown

FiGuRE 1. Map of Kindersley area, courtesy of Donald S. Houston.

DDT in the productivity decline. The least aggres-
sive adult Merlins had decreased productivity and
their eggs had thinner shells (Hodson 1976). A fol-
low-up study in Alberta by Fox and Donald (1980)
again showed that eggs with large DDE (the break-
down product of DDT) burdens had significantly
thinner shells, disappeared more often, and hatched
less frequently than normal.

Habitat

Fox (1971) thought the sudden decline in Merlin
numbers might have been accelerated by deteriora-
tion and disappearance of trees used as nest sites.
Hodson (1976) found no crow, magpie, or hawk
nests available for Merlin nest use in 7 of 14 previ-
ous Merlin sites near Kindersley. To quantify the
disappearance of grasslands, Hodson (1976) com-
pared air photos near Kindersley between 1956 and
1971. In 1956, ten of the 15 historic Merlin territo-
ries included more than 50% grassland, whereas by
1971, only 4 of the 15 retained this amount of
grassland. Put another way, near Kindersley 52% of
the Merlin hunting territories present in the 1940s
had been cultivated by 1971. Hodson calculated
that 58% of the land within 1.6 km of Merlin nests
was under cultivation in 1971, as compared to only
22% at Hanna, Alberta, where Merlins persisted as
a breeding species.

Colonization of urban habitat

In the earliest phase of their recovery, Merlins
followed the American Crow (Corvus brachyrhyn-
chos) and Black-billed Magpie (Pica pica) into the
cities, where corvid nests became available for city
Merlins to use. By 1971 Merlins were regular resi-
dents in Saskatoon (Oliphant and McTaggart 1977;
Sodhi et al. 1992). Since 1974 intensive studies of
both summering and wintering Merlins have been
carried out by Oliphant, James, Warkentin, Sodhi,
and colleagues (citations in Sodhi et al. 1993), with
a continued slow increase in the number of breed-
ing pairs. There are now over 30 pairs resident in
Saskatoon each year.

Methods
Post-pesticide productivity at Kindersley

Houston’s studies of the Merlins near Kindersley
were a by-product of more intense studies of
Swainson’s (Buteo swainsoni) and Ferruginous
hawks (Buteo regalis) since 1969 in the grasslands
within all but the northeast quarter of the 1:250
000 Kindersley map sheet, from 51° to 52° North
and from 108° to 110° West. Each known nest of
both buteos is visited at least once each year, the
Ferruginous in late June and the Swainson’s in late
July. Over the years, with increased effort, the

246

TABLE 2. Merlins banded per successful rural nest,
1975-1995.

Nests with # nests Total young % young

| young 5 5 1.0%

2 young 6 12 2.4%

3 young 20 60 11.8%

4 young 39 156 30.7%

5 young 55 275 54.1%
Total 125 508 100.0%

number of buteo nests under surveillance has grad-
ually increased (Houston and Schmutz 1995).
Early visits included government pastures where
Merlins were completely absent in the first six
years but appeared subsequently. Visits were timed
for the two buteo species; sometimes the Merlins
were too small for banding at the Ferruginous
banding visit but had fledged by the time of the
Swainson’s visit.

Results

During Houston’s first six years of Kindersley area
studies, 1969-1974 inclusive, no nesting Merlins
were encountered. One nesting pair was encountered
in each of 1975 and 1976. Since then, Merlins have
gradually re-established in the Kindersley area.
Since 1975, 508 Merlin nestlings have been banded
in 125 rural nests, with productivity of 4.06 young
per successful nest. The most common number in a
nest at banding time was five young (Table 2). The
highest number of rural nests encountered (12) and
the highest number of young (51), occurred in 1995,
when Swainson’s and Ferruginous hawk productivi-
ty were at all-time lows (Houston and Schmutz
1995a, 1995b).

Unlike the Saskatoon Merlins, which prefer previ-
ous crow nests in conifers, the Merlins in the
Kindersley area seemed to use preferentially the
cover provided by previous, roofed magpie nests in
deciduous trees (Table 3). However, two Merlin
pairs nested on top of the magpie nest, rather than
inside the covered nest cup.

Whereas in the city of Saskatoon, males return to
the same nest area in a subsequent year 61% of the
time, and females 29% of the time (Warkentin et al.
1991), near Kindersley Merlins rarely occupied the
same shelterbelt in consecutive years. For example,
a deserted farm shelterbelt in the Antelope Park
Pasture near Hoosier, always with at least two mag-
pie nests and one crow nest in good condition, had
Merlins nesting successfully in 1980, 1981, 1986,
1993, and 1995. Only one other known site in this
pasture was used once, in 1988. In the 63.5 km?
Kindersley-Elna Pasture, only four of nine aban-
doned farm groves have been used, and only in
1986 were two groves occupied simultaneously. The

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 3. Merlin nest trees (rural), 1975-1995.

Former occupant (builder of nest)

Tree BBMA AMCR SWHA _ Total
Maple 54 11 1 66
Aspen 15 6 l 22
Green Ash 1] 2 13
Willow sp. 11 11
Elm sp. - l 5
Cottonwood 2 2
Saskatoon 2 1 3
Chokecherry l 1
Spruce 2 2
102 21 2 125

BBMA - Black-billed Magpie, Pica pica
AMCR - American Crow, Corvus brachyrhynchos
SWHA - Swainson’s Hawk, Buteo swainsoni

first grove was used in 1975 with three successful
raptor species quite close together, a Ferruginous
Hawk, a Swainson’s Hawk, and a Merlin, and had
Merlins again in 1984. A smaller L-shaped grove
was used consecutively in 1985 and 1986. Another
grove was used once in 1986. The most popular
grove was used in 1976, 1982, 1983, 1987, 1988
and finally in 1989 when on 24 June a bigamous
male defended both a successful female with well-
developed young and another female nearby with
newly-hatched, downy young, but the second nest
failed before a repeat visit. A fifth site, in trees
along the shore of a tiny lake just outside the south
edge of the pasture, was used only in 1992. This last
nest was remarkable in that magpies were raised
inside the nest and Merlins simultaneously were
raised on top of the nest.

To date, there have been only seven recoveries
(0.8%) from 561 Merlins banded, all but one (after
2.5 years) within the first year after banding. Three
distant recoveries were from extreme southern
Saskatchewan, Nebraska, and Texas, at distances of
260, 1290, and 2005 km, respectively. The recovery
rate here is only one-third of that obtained from
banding in Alberta (Schmutz et al. 1991).

Discussion

Although this study relied on a small sample of
observations, it indicated that Merlins were virtually
absent from grasslands near Kindersley between
1962 and 1975. Disappearance in the 1960s was due
largely to pesticide toxicity, probably from both
dieldrin and DDT. Since 1975, Merlin populations
have gradually recouped in numbers and productivi-
ty near Kindersley at the same time as they invaded a
number of cities such as Saskatoon, Edmonton, and
Winnipeg and, to a lesser degree, Regina.

Grassland Merlins overwhelmingly use nests of
the Black-billed Magpie in deciduous trees, whereas

1997

in Saskatoon all 58 nests studied were in conifers
and 54 of the 58 had been built by crows
(Warkentin and James 1988). It is interesting that in
Montana, Merlins use magpie nests in stands of
Ponderosa Pine near grassland; Black-billed Magpie
nests combine easy access with maximum conceal-
ment (Sieg and Becker 1990).

Since 1977, the productivity per rural nest (aver-
age number of chicks reaching 15 days of age), has
been excellent (4.06), minimally better than the 3.8
reported by Sodhi et al (1992) for the city of
Saskatoon. The 44% (55/125) of nests raising five
young to banding age is a big improvement over the
early Kindersley figures (1958-1960), and from the
rural Alberta figures in the early 1970s. More than
half of the young in rural Saskatchewan are pro-
duced from nests with five young (Table 2).
Polygyny, considered rare in this species, has been
reported once by us and once in Saskatoon by Sodhi
(1989).

Why do the rural Merlins near Kindersley not
show the fidelity to nest areas demonstrated by
color-banded birds in the city of Saskatoon? One
doubts that the high mortality rate for this small fal-
con, much higher than that for larger hawks, is the
explanation; although the mortality in rural areas is
unknown, in Saskatoon the annual adult mortality
rate is estimated at 31% for males and 29% for
females (James et al. 1989). The Saskatoon studies
indicate greater dispersal of females, with recover-
ies of adult females in the breeding season at dis-
tances of 71, 123, 231, and 259 km (James et al.
1989), and a color-banded female sighted in
Edmonton in 1994 was believed to have originated
in Saskatoon (Rick Espie, personal communica-
tion). In England, early in this century, Merlin terri-
tories tended to be occupied annually, even after
removal of previous occupants: on the moors near
Skipton, Yorkshire, England, the late William
Rowan (1921) reported that on one territory, both
individuals of a pair of Merlins were killed by the
gamekeeper each year for 19 consecutive years, and
yet a pair inevitably replaced them the following
year.

Trees on Saskatchewan pastures have deteriorated
greatly since the 1960s, and even more grassland
has succumbed to the plough, yet in spite of these
adverse factors it has been heartening to observe the
resurgence of this attractive little falcon. It is indeed
fortunate that two raptor enthusiasts were suitably
situated to document the decline, confirmed by
additional surveys for a thesis project.

Addendum

Houston also banded young in single nests with
two and three young discovered incidentally in 1957
and 1958, before Merlin numbers declined, and four
young in a river coulee near Beechy, Saskatchewan,

HOUSTON AND HODSON: RESURGENCE OF MERLINS IN SASKATCHEWAN

247

in 1974. Another 43 nestlings in ten nests in cities
and large towns (Saskatoon and Nipawin) and in
mixed forest areas have been banded; all ten were
previous crow nests in spruce trees. Only one adult
has been banded.

Acknowledgments

We are grateful to Glen A. Fox and Richard W.
Fyfe, each of whom studied Merlins as a high
school student in the Kindersley area, and who criti-
cized an earlier version of this manuscript. Mary
Houston, Josef K. Schmutz, Lynn W. Oliphant,
Rick Espie, David J. Lieske, Margaret Belcher and
Frank Roy also offered constructive criticism. Jim
Hay and Paul W. Riegert assisted our search for
prices of dieldrin. This paper was presented at the
Raptor Research Foundation meeting at Duluth, 2-4
November 1995.

Literature Cited

Bent, A. C. 1938. Life Histories of North American Birds
of Prey, Part 2. [Pages 87-88.] Washington:
Smithsonian Institution.

Folker, R. V. 1960. A study of the effects of dieldrin
spraying on wildlife. Department of Natural Resources,
Regina, Saskatchewan. 10 pages.

Fox, G. A. 1964. Notes on the western race of the Pigeon
Hawk. Blue Jay 22(4): 140-147.

Fox, G. A. 1971. Recent changes in the reproductive suc-
cess of the pigeon hawk. Journal of Wildlife
Management 35(1): 122-128.

Fox, G. A. 1979. A simple method of predicting DDE con-
tamination and reproductive success of populations of
DDE-sensitive species. Journal of Applied Ecology 16:
737-741.

Fox, G. A., and T. Donald. 1980. Organochlorine pollu-
tants, nest-defense behavior and reproductive success in
Merlins. Condor 82(1): 81-84.

Fyfe, R. W., R. W. Risebrough, and W. Walker II. 1976.
Pollutant effects on the reproduction of the Prairie
Falcons and Merlins of the Canadian Prairies. Canadian
Field-Naturalist 90(3): 346-355.

Hodson, K. A. 1976. The ecology of Richardson’s Merlins
on the Canadian Prairies. M. Sc. thesis, University of
British Columbia.

Hodson, K. A. 1978. Prey utilized by merlins nesting in
shortgrass prairies of southern Alberta. Canadian Field-
Naturalist 92(1): 76-77.

Houston, C. S., and A. Schmidt. 1981. History of
Richardson’s Merlin in Saskatchewan. Blue Jay 39(1):
30-37.

Houston, C. S., and J. K. Schmutz. 1995a. Declining
reproduction among Swainson’s Hawks in prairie
Canada. Journal of Raptor Research 29(3): 198-201.

Houston, C. S., and J. K. Schmutz. 1995b. Changes in
Canadian grassland bird populations. Proceedings of
Conference, Conservation and Ecology of Grassland
Birds, Tulsa, Oklahoma, 26-26 October 1995
[Abstract].

James, P. C., I. G. Warkentin, and L. W. Oliphant.
1989. Turnover and dispersal in urban Merlins Falco
columbarius. Ibis 131(4): 426-429.

248

Oliphant, L. W., and W. J. P. Thompson. 1979. Recent
breeding success of Richardson’s Merlin in
Saskatchewan. Raptor Research 12: 35-39.

Oliphant, L. W., and S. McTaggart. 1977. Prey utilized
by urban Merlins. Canadian Field-Naturalist 91(2):
190-192.

Rowan, W. 1921. Observations on the breeding-habits of
the Merlin. I: General environment. British Birds 15(6):
122-129.

Saskatchewan Department of Agriculture. 1950 - 1966.
Annual reports for the twelve months ended March 31.
Queen’s Printer, Regina.

Schmutz, J. K., R. W. Fyfe, U. Banasch, and H. Arm-
bruster. 1991. Routes and timing of migration of fal-
cons banded in Canada. Wilson Bulletin 101(3):
44-58.

Sieg, C. H., and D. M. Becker. 1990. Nest-site habitat
selected by merlins in southeastern Montana. Condor
92(3): 688-694.

Sodhi, N. S. 1989. Attempted polygyny by a merlin.
Wilson Bulletin 101(3): 506—507.

THE CANADIAN FIELD-NATURALIST

Vol. 111

Sodhi, N. S., P. C. James, I. G. Warkentin, and L. W.
Oliphant. 1992. Breeding ecology of urban Merlins
(Falco columbarius). Canadian Journal of Zoology
70(4): 1477-1483.

Sodhi, N. S., L. W. Oliphant, P. C. James, and I. G.
Warkentin. 1993. Merlin (Falco columbarius) in The
Birds of North America, Number 44. Edited by A. Poole
and F. Gill. The Academy of Natural Sciences,
Philadelphia, and The American Ornithologists’ Union,
Washington, D.C.

Temple, S. A. 1972. Systematics and evolution of the
North American Merlins. Auk 89(2): 325-338.

Warkentin, I. G., and P. C. James. 1988. Nest-site selec-
tion by urban merlins. Condor 90(2): 734-738.

Warkentin, I. G., P. C. James, and L. W. Oliphant.
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108(2): 294-302.

Received 12 June 1996
Accepted 24 October 1996

Rare and Endangered Fishes and Marine Mammals of Canada:
COSEWIC Fish and Marine Mammal Subcommittee
Status Reports: XI

R. R. CAMPBELL
666 Plantation Drive, R.R. 2 Woodlawn, Ontario KOA 3M0O

Campbell, R.R. Editor. 1997. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine
Mammal Subcommittee Status Reports: XI. Canadian Field-Naturalist 111(2): 249-257.

Six status reports representing the 1996 fish and marine mammal status assignments have been prepared for publication.
Committee (COSEWIC) and Subcommittee (Fish and Marine Mammals) activities are briefly discussed as is proposed leg-
islation regarding a Canadian Endangered Species Act. Tabular lists of fish and marine mammal species assigned status to
April 1996, reports in progress, and species of possible interest are presented.

Six rapports de statut relativement aux poissons et aux mammiféres marins auxquels ont été attribués un statut en 1996 ont
été préparés pour publication. Les activitités du Comité (CSEMDC) et du sous-comité (des poissons et des mammiféres
marins) sont briévement discutées, de méme que la proposition législative quant a une Loi sur la protection des espéces en
péril au Canada. Les listes sous forme tabulaire des espéces de poisson et de mammifére marin qui ont recu un statut

jusqu'en avril 1996, des rapports en cours, et des espéces qui pourraient étre d'intérét sont présentées.

Key Words: Rare and endangered species, fish, marine mammals, COSEWIC.

As indicated in previous submissions (Campbell
1984 through 1996), the intent of the Subcommittee
on Fish and Marine Mammals is to publish the status
reports (on those species of fish and marine mam-
mals) which the Committee on the Status of
Endangered Wildlife in Canada (COSEWIC) has
reviewed, approved and used as a basis of assigning
status to species in jeopardy in Canada.* The group
of six reports presented herein represent the fish and
marine mammal component of those species
assigned status in 1996 (see Table 1). It is hoped that
we will be able to find continuing financial support
to offer, in succeeding volumes, those reports
reviewed in future years.

In addition to the six reports included here, a sev-
enth species was accorded status at the 1996 General
Meeting. The Eelgrass limpet, Lottia alveus alveus
was declared extinct based on a previously published
article by Carlton et al. (1991).

Progress
COSEWIC has undertaken to make available to
all Canadians supporting information on each

*Publications of Status Reports submitted to other subcom-
mittees, and subsequently approved by COSEWIC, includ-
ing some mammals, birds, amphibians and reptiles, and
plants have not had subcommittee publication support;
some have been published in The Canadian Field-
Naturalist on the individual initiative of their authors.
Full original Status Reports for all designated species are
available from the Committee on the Status of Endangered
Wildlife in Canada, Ottawa, Ontario, Canada K1A OH3.
Phone (819) 997-4991.

species classified (see Cook and Muir 1984). The
Fish and Marine Mammal Subcommittee has been
able to use this journal as one step in achieving the
goal. A series of reports have appeared in various
volumes and numbers from 1984 through 1996 [see
Canadian Field-Naturalist 98(1): 63-133; 99(3):
404-450; 101(2): 165-309; 102(1): 81-176 and
102(2): 270-398; 103(2): 147-220; 104(1): 1-145;
105(2): 151-250; 106(1): 1-72; 107(4): 395-546;
110(3): 454-532].

In April 1996 the Canadian Wildlife Federation
(CWF) discontinued support of COSEWIC report
preparation, with the exception of the Canadian
Wildlife Service contribution of $10 000, no funding
report remains at the time of writing. Negotiation
with other Canadian non-governmental agencies
such as World Wildlife Fund (Canada) will hopeful-
ly result in a matching funding agreement similar to
that enjoyed with CWE from 1993 to 1996 (see
Campbell 1996).

As of April 1996, COSEWIC has reviewed the
status of 86 fish species, two marine invertebrates,
and 41 marine mammals (Table 1). Of the 129
species investigated six are indeterminate (five fish,
one marine mammal), 54 (21 fish, 32 marine mam-
mals, one marine invertebrate) have been found not
to require status designation and another 44 (38 fish,
six Marine mammals) have been designated as vul-
nerable, mainly due to natural rarity, leaving 24
species of immediate concern (threatened and endan-
gered) which are of interest to the RENEW
(Recovery of Nationally Endangered Wildlife) orga-
nization which was established in 1990 to oversee
the development of recovery teams and plans for

249

250

THE CANADIAN FIELD-NATURALIST

TABLE |. Fish and Marine Mammal Species with Assigned COSEWIC Status to 19 April 1996.

Species

Fish

Lake Sturgeon
Bloater

Blueback Herring
Cutlips Minnow
Striped Shiner
Redfin Shiner
Hornyhead Chub
River Chub

Ghost Shiner
Blackchin Shiner
Leopard Dace
Mountain Sucker
Golden Redhorse
Least Darter
Tesselated Darter
River Darter
Green Sunfish
Longear Sunfish
Spoonhead Sculpin
Brook Silverside
Y -Prickleback
Darktail Lamprey
Bering Cisco
Flathead Catfish
Northern Madtom
Pixy Poacher
Lake Lamprey*
Chestnut Lamprey
Northern Brook Lamprey
Shortnose Sturgeon
Green Sturgeon
White Sturgeon
Spotted Gar
Spring Cisco
Squanga Whitefishe
Kiyi

Pacific Sardine
Redside Dace
Silver Chub
Pugnose Shiner
Bigmouth Shiner
Silver Shiner

Roseyface Shiner - Manitoba

Pugnose Minnow
Speckled Dace
Umatilla Dace
Central Stoneroller

Banded Killifish -Newfoundland

Blackstripe Topminnow
Lake Chubsucker
Bigmouth Buffalo
Black Buffalo

Spotted Sucker

River Redhorse
Greenside Darter
Brindled Madtom
Redbreast Sunfish
Warmouth
Orangespotted Sunfish

Fourhorn Sculpin - Arctic Islands

Scientific Name

Acipenser fulvescens
Coregonus hoyi

Alosa aestivalis
Exoglossum maxillingua
Luxilis chyrsocephalus
Lythrurus umbratilis
Nocomis biguttatus
Nocomis micropogon
Notropis buchanani
Notropis heterodon
Rhinichthys falcatus
Catostomus platyrhynchus
Moxostoma erythrurum
Etheostoma microperca
Etheostoma olmstedi
Percina shumardi
Lepomis cyanellus
Lepomis megalotis
Cottus ricei

Labidesthes sicculus

Allolumpenus hypochromus

Lethenteron alaskense
Coregonus laurettae
Pylodictis olivaris
Noturus stigmosus
Occella impi

Lampetra macrostoma
Ichthyomyzon castaneus
Ichthyomyzon fossor
Acipenser brevirostrum
Acipenser medirostris
Acipenser transmontanus
Lepisosteus oculatus
Coregonus sp.
Coregonus sp.
Coregonus kiyi
Sardinops sagax
Clinostomus elongatus
Macrhybopsis storeriana
Notropis anogenus
Notropis dorsalis
Notropis photogenis
Notropis rubellus
Opsopoeodus emiliae
Rhinichthys osculus
Rhinichthys umatilla
Campostoma anomalum
Fundulus diaphanus
Fundulus notatus
Erimyzon sucetta
Ictiobus cyprinellus
Ictiobus niger
Minytrema melanops
Moxostoma carinatum
Etheostoma blennioides
Notorus miurus
Lepomis auritus
Lepomis gulosus
Lepomis humilis

Myoxocephalus quadricornis

Status

NARa2
NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR

NAR
Indeterminate
Indeterminate
Indeterminate
Indeterminate
Indeterminate
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable
Vulnerable

Vol. 111

Date Assigned

April 1986
April 1988
April 1980
April 1994
April 1993
April 1988
April 1988
April 1988
April 1993
April 1994
April 1990
April 1991
April 1989
April 1989
April 1993
April 1989
April 1987
April 1987
April 1989
April 1989
April 1991
April 1990
April 1990
April 1993
April 1993
April 1991
April 1986
April 1991
April 1991
April 1980
April 1987
April 1990
April 19834
April 1992
April 1988
April 1987
April 1987
April 1987
April 1985
April 1985
April 1985
April 1983¢
April 1994
April 1985
April 19808
April 1988
April 1985
April 1989
April 1985
April 1994
April 1989
April 1989
April 19834
April 1983
April 1990
April 1985
April 1989
April 1994
April 1989
April 1989

1997 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA: STATUS REPORTS

TABLE |. Continued.
Species

Giant Stickleback
Unarmoured Stickleback¢
Blackline Prickleback
Bering Wolffish

Lake Simcoe Whitefish
Blackfin Cisco
Shortnose Cisco
Shortjaw Cisco

Black Redhorse

Copper Redhorse*
Eastern Sand Darter
Channel Darter
Margined Madtom
Enos Lake Stickleback
Shorthead Sculpin
Deepwater Sculpin - Great Lakes
Acadian Whitefish
Aurora Trout¢
Nooksack Dace

Salish Sucker
Paddlefish

Gravel Chub

Longjaw Cisco
Deepwater Cisco

Banff Longnose Dace‘
Blue Walleye

Marine Molluscs
Northern Abalone
Eelgrass Limpet

Marine Mammals
Sea Otter
Sea Mink
Northern Fur Seal
Hooded Seal
Bearded Seal
Northern Elephant Seal
Ringed Seal
Harbour Seal
Lac des Loups Marins
Steller Sea Lion
California Sea Lion
Atlantic Walrus
Eastern Arctic
Northwest Atlantic
Baird's Beaked Whale
Beluga
Beaufort Sea
Western and Southern Hudson Bay
High Arctic
Eastern Hudson Bay
St. Lawrence River
S.E. Baffin Island
Ungava Bay
Common Dolphin
Grey Whale
Northeast Pacific
Northwest Atlantic
Risso’s Dolphin
Short-finned Pilot Whale

Scientific Name

Gasterosteus sp.
Gasterosteus sp.
Acantholumpenus mackayi
Anarichus orientalis
Coregonus clupeaformis ssp.
Coregonus nigripinnis
Coregonus reighardi
Coregonus zenithicus
Moxostoma duquesnei
Moxostoma hubbsi
Ammocrypta pellucida
Percina copelandi

Noturus insignis
Gasterosteus sp.

Cottus confusus
Myoxocephalus thompsoni
Coregonus huntsmani
Salvelinus fontinalis timagamiensis
Rhinichthys cataractae ssp.
Catostomus sp.

Polyodon spathula
Erimystax x-punctata
Coregonus alpenae
Coregonus jJohannae
Rhinichthys cataractae smithi
Stizostedion vitreum glaucum

Haliotis kamtschatkana
Lottia alveus alveus

Enhydra lutris

Mustela macrodon
Callorhinus ursinus
Cystophora cristata
Erignathus barbatus
Mirounga angustirostris
Phoca hispida

Phoca vitulina mellonae

Eumetopias jubatus
Zalophus californianus
Odobenus rosmarus rosmarus

Berardius bairdi
Delphinapterus leucas

Delphinus delphis
Eschrichtius robustus

Grampus griseus
Globicephala macrohynchus

Status

Vulnerable
Vulnerable
Vulnerable
Vulnerable
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Threatened
Endangered
Endangered
Endangered
Endangered
Extirpated
Extirpated
Extinct
Extinct
Extinct
Extinct

N/Ab
Extinct

Threatened
Extinct
NAR

NAR

NAR

NAR

NAR
Vulnerable

NAR
NAR

NAR
Extirpated
NAR

NAR

NAR
Vulnerable
Threatened
Endangered
Endangered
Endangered
NAR

NAR
Extirpated
NAR
NAR

Date Assigned

April 1980
April 1983
April 1989
April 1989
April 1987
April 1988
April 1987
April 1987
April 1988
April 1987
April 1994
April 1993
April 1989
April 1988

November 1983

April 1987
April 1983
April 1987
April 1996
April 1986
April 1987
April 19872
April 1988
April 1988
April 1987
April 1985

April 1988
April 1996

April 1996#
April 1985
April 1996
April 1986
April 1994
April 1986
April 1989
April 1996

April 1987
April 1987

April 1987
April 1987
April 1992

April 1986
April 1993
April 1992
April 1988
April 1983
April 1990
April 1988
April 1991

April 1987
April 1987
April 1990
April 1993

232. THE CANADIAN FIELD-NATURALIST Vol. 111
TABLE |. Concluded.
Species Scientific Name Status Date Assigned
Long-finned Pilot Whale Globicephela malaena NAR April 1994
Northern Bottlenose Whale Hyperoodon ampullatus

Northwest Atlantic NAR April 1993

Gully Population Vulnerable April 1996
Pygmy Sperm Whale Kogia breviceps NAR April 1994
Atlantic White-sided Dolphin Lagenorhynchus acutus NAR April 1991
Pacific White-sided Dolphin Lagenorhynchus obliquidens NAR April 1990
Northern Right Whale Dolphin Lissodelphis borealis NAR April 1990
Hubbs’ Beaked Whale Mesoplodon carlhubbsi NAR April 1989
Blainville’s Beaked Whale Mesoplodon densirostris NAR April 1989
True’s Beaked Whale Mesoplodon mirus NAR April 1989
Stejneger’s Beaked Whale Mesoplodon stejnegeri NAR April 1989
Narwhal Monodon monoceros NAR April 1986¢
Dall's Porpoise Phocoenoides dalli NAR April 1989
False Killer Whale Pseudorca crassidens NAR April 1990
Sperm Whale Physeter macrocephalus NAR April 1996
Striped Dolphin Stenella coeruleoalba NAR April 1993
Bottlenose Dolphin Tursiops truncatus NAR April 1993
Cuvier's Beaked Whale Ziphius cavirostris NAR April 1990
Blue Whale Balaenoptera musculus Vulnerable April 1983!
Fin Whale Balaenoptera physalus Vulnerable April 1987
Sowerby's Beaked Whale Mesoplodon bidens Vulnerable April 1989
Harbour Porpoise Phocoena phonoeca

Northwest Pacific Indeterminate April 1991

Northwest Atlantic Threatened April 1990
Humpback Whale Megaptera novaeangliae

Northwest Atlantic Vulnerable April 1985

Northeast Pacific Threatened April 1982)
Bowhead Whale Balaena mysticetus Endangered April 1980!
Right Whale Eubalaena glacialis Endangered April 1980k

aNAR The use of NIAC (Not in Any Category) was discontinued in 1988 and replaced by RANSDR (Report Accepted No
Status Designation Required - not a category of risk). Species previously designated as NIAC were converted to RANS-
DR at the April 1989 meeting on the recommendation of the subcommittee. The Committee approved the use of a catego-
ry to be known as species Not At Risk (NAR) at the April 1994 meeting. All species previously designated RANSDR
were converted to NAR at the April 1995 meeting as recommended by the subcommittee (see Campbell 1996).
>Indeterminate — The use of a new list, “Report Accepted Insufficient Scientific Information For Status Designation”
(RAISIFSD), was approved at the 1990 General Meeting. The Committee agreed to establish this as the “Indeterminate”
(1) category at the April 1994 meeting. Species previously listed as RAISIFSD were converted to Indeterminate at the
April 1995 mmeting on the advice of the subcommittee (see Campbell 1996).

‘Endemic to Canada

dUpdated April 1994 no status change.

¢Updated April 1987 no status change.

fUpdated April 1984 no status change.

sUpdated April 1987 previous status of “Endangered” assigned April 1985.

5N/A Status Not Assigned. Until 1994 COSEWIC had no mandate for invertebrates. Report accepted and recommended
RANSDR Status agreed to, but not assigned. An updated report is in progress.

‘Endangered 1978 - updated April 1996 no status change.

iUpdated April 1985 North Atlantic stock downlisted to “Vulnerable”.

kKUpdated April 1985 and April 1990 no status change.

‘Updated April 1996 - downlisted to “Threatened”.

such species listed by COSEWIC (CWS 1993). To _ species (includes eight updates), one marine mollusc
date, recovery plans have been produced and imple- (update), and 11 on marine mammal species (three
mented only for the St. Lawrence River Beluga. updates) under review or in preparation (Table 2).
Recovery teams are in the process of development of Several of these will be presented to the Committee
recovery plans for the Gully population of the for status assignment in 1997. As well, some 78
Northern Bottlenose Whale and Harbour Porpoise. additional species of fish (plus eight to be updated),

There are currently 37 status reports on fish one marine mammal (plus one update) and 12

1997 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA: STATUS REPORTS 253

TABLE 2. Fish and Marine Mammal Species for which Status Reports are in preparation, or under review - to 19 April 1996.

Species

Fish

Shortnose Sturgeon*
Atlantic Sturgeon

Lake Sturgeon*

Atlantic Cod

Blueback Herring’

Red (Arctic) Char

Bull Trout

Mira Whitefish*
Opeongo Whitefish*
Lake Cisco

Lake Whitefish

Acadian Whitefish*”
Pygmy Whitefish

Round Whitefish

Pygmy Smelt

Pygmy Longfin Smelt*
Redfin Pickerel

Grass Pickerel

Chain Pickerel
Chiselmouth

Central Stoneroller’
Liard Hotspring Lake Chub*°
Western Silvery Minnow
Eastern Silvery Minnow
Weed Shiner

Bluntnose Minnow
Speckled Dace”

Jasper Longnose Sucker*
Striped Bass

Spinynose Sculpin
Cultus Pygmy Coastrange Sculpin*
Mottled Sculpin
Shorthead Sculpin°

Giant Stickleback*’
Unarmoured Stickleback*’
Texada Stickleback*
Bluefin Tuna

Marine Molluscs
Northern Abalone”

Marine Mammals
Minke Whale
Sei Whale
Bowhead Whale”
Blue Whale’
Beluga’
St. Lawrence River
Ungava Bay
Dwarf Sperm Whale
White-beaked Dolphin
Killer Whale
Grey Seal
Harp Seal

*Endemic to Canada ON = Ontario
PQ = Quebec

“Updated Status Report

marine invertebrates have been identified as being
worthy of consideration (Table 3). Many may be
found to not require status designation, but the pro-

Scientific Name

Acipenser brevirostrum
Acipenser oxyrhynchus
Acipenser fulvescens

Gadus morhua

Alosa aestivalis

Salvelinus alpinus ssp.
Salvelinus confluentus
Coregonus sp.

Coregonus sp.

Coregonus artedi

Coregonus clupeaformis
Coregonus huntsmani
Prosopium coulteri
Prosopium cylindraceum
Osmerus spectrum
Spirinichus thaleichthys
Esox americanus americanus
Esox americanus vermiculatus
Esox niger

Acrocheilus alutaceus
Campostoma anomalum
Couesius plumbeus ssp.
Hybognathus argyritis
Hybognathus nuchalis regius
Notropis texanus

Pimphales notatus
Rhinichthys osculus
Castostomus castostomus lacustris
Morone saxatilis
Asemichthys taylori

Cottus aleuticus

Cottus bairdi

Cottus confusus
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.

Thunnus thynnus

Haliotis kamtschatkana

Balaenoptera acutorostrata
Balaenoptera borealis
Balaena mysticetus
Balaenoptera musculus
Delphinapterus leucas

Kogia simus
Lagenorhynchus albirostris
Orcinus orca

Halichoerus grypus

Phoca groenlandica

BC = British Columbia
NS = Nova Scotia

Proposed Status

Vulnerable April 1980
iy

?

a
NAR April 1980

2

Vulnerable
Vulnerable
Threatened
Endangered
Threatened - Lake Erie, ON
Endangered April 1983
9

Vulnerable
Vulnerable
Vulnerable - BC
Vulnerable - Quebec
Vulnerable
Vulnerable - PQ, NB, NS
Vulnerable - BC
Vulnerable April 1985
Vulnerable - BC

9
Vulnerable
Vulnerable
Vulnerable
Vulnerable April 1980
Vulnerable
Endangered - PQ
Vulnerable - BC
Threatened - BC
Vulnerable - BC, Alberta
Vulnerable
Vulnerable April 1980
Vulnerable April 1983

Vulnerable
?
ue
2
?
Endangered
Vulnerable

Endangered April 1983
Endangered April 1988
Vulnerable

NB = New Brunswick

cess serves to bring together the information neces-
sary to make the appropriate determination and satis-
fy the need to fill those knowledge gaps. Although

254

THE CANADIAN FIELD-NATURALIST

Vol. 111

TABLE 3. Fish and Marine Mammal Species of Possible Interest to COSEWIC - April 1996 (Not listed by Priority).

Species

Species Updates

Fish

Lake Lamprey

Silver Chub

Bigmouth Shiner
Pugnose Minnow
Pugnose Shiner

Salish Sucker
Blackstripe Topminnow
Brindled Madtom

Marine Mammals

Humpback Whale
Northwest Atlantic
Northeast Pacific

Species Yet to be Considered
Fish

Spiny Dogfish

Smooth Skate

Thorny Skate

American Eel

American Shad

Atlantic Herring

Creek Chubsucker

Brassy Minnow

Pearl Dace

Emerald Shiner

Bridle Shiner

Spottail Shiner

Creek Chubsucker

Stonecat

Capelin

Broad Whitefish

Least Cisco

Giant Pygmy Whitefish

Pacific Salmonids

Atlantic Salmon

Monkfish

Tomcod

Haddock

Silver Hake

Pollock

Red Hake

Rock Grenadier

Balkwill Lake Benthic Stickleback
Balkwill Lake Limnetic Stickleback
Emily Lake Benthic Stickleback
Emily Lake Limnetic Stickleback
Hadley Lake Benthic Stickleback
Hadley Lake Limnetic Stickleback
Paxton Lake Benthic Stickleback
Paxton Lake Limnetic Stickleback
Priest Lake Benthic Stickleback
Priest Lake Limnetic Stickleback
Ninespine Stickleback

Pacific Ocean Perch

Aurora Rockfish

Redbanded Rockfish

Silvergray Rockfish

Copper Rockfish

Scientific Name

Lampeta macrostoma
Macrhybopsis storeriana
Notropis dorsalis
Opsopoedus emiliae
Notropis anogenus
Catostomus sp.
Fundulus notatus
Noturus miurus

Megaptera novaeangliae

Squalus acanthias
Malacoraja senta

Raja radiata

Anguilla rostrata
Alosa sapidissima
Clupea harengus
Erimyzon oblongus
Hybognathus hankinsoni
Margariscus margarita
Notropis atherinoides
Notropis bifrenatus
Notropis hudsonius
Erimyzon oblongatus
Noturus flavus
Mallotus villotus
Coregonus nasus
Coregonus sardinella
Prosopium sp.?
Oncorhynchus spp. (5 species)
Salmo salar

Lophius americanus
Microgadus tomcod
Melanogrammus aeglefinus
Merluccius bilinearis
Pollachius virens
Urophycis chuss
Coryphaenoides rupestris
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Gasterosteus sp.
Pungitius pungitius
Sebates alutus
Sebastes aurora
Sebastes babcocki
Sebastes brevispinis
Sebastes caurinus

Possible Status

Vulnerable April 1986
Vulnerable April 1985
Vulnerable April 1985
Vulnerable April 1985
Vulnerable April 1985
Endangered April 1986
Vulnerable April 1985

Vulnerable April 1985

Vulnerable April 1985
Threatened April 1982+

Vulnerable

9

9
Extirpated ?

9
Vulnerable - BC
Vulnerable - BC
Vulnerable
Vulnerable - BC

9
? - Alberta
9
Vulnerable - BC
Vulnerable - BC
Threatened - BC

9

a

ts
Vulnerable - Quebec

Vulnerable - BC
Vulnerable - BC
Vulnerable - BC
Vulnerable - BC
Vulnerable - BC
Vulnerable - BC
Threatened - BC
Threatened - BC
Threatened - BC
Threatened - BC
Vulnerable - BC

oD OD OD OD

1997 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA: STATUS REPORTS

TABLE 3. Concluded.

Species Scientific Name Proposed Status
Fish
Splitnose Rockfish Sebastes diploproa ?
Widow Rockfish Sebastes entomelas ?
Chilipepper Sebastes goodei is
Shortbelly Rockfish Sebastes jordani q
Quillback Rockfish Sebastes maliger q
Black Rockfish Sebastes melanops ?
Vermillion Rockfish Sebastes miniatus ?
Blue Rockfish Sebastes mystinus L
China Rockfish Sebastes nebulosus cy
Golden Rockfish Sebates norvegicus ?
Bocaccio Sebates paucispinis 2
Canary Rockfish Sebates pinniger ?
Yelloweye Rockfish Sebates ruberrimus ?
Stipetail Rockfish Sebates saxicola ?
Shortspine Thornyhead Sebastolobus alascanus ?
Longspine Thornyhead Sebastolobus altivelis ?
Lingcod Ophiodon elongatus 2
Rainbow Darter Etheostoma caeruleum u
Lumpfish Cyclopterus lumpus ?
Wolffish Anarhichas lupus z
Atlantic Mackeral Scomber scombrus ?
Swordfish Xiphias gladius ?
Summer Flounder Paralichthys dentatus ?
Witch Flounder Glyptocephalus cynoglossus ?
American Plaice Hippoglossoides platessoides z4
Atlantic Halibut Hippoglossus hippoglossus ?
Pacific Halibut Hippoglossus steolepis Z
Yellowtail Flounder Limanda ferrugineus i;
Winter Flounder Pseudopleuronectes americanus i
9

Greenland Halibut (Turbot)

Marine Mammals

Reinhardtius hippoglossoides

Harbour Seal Phoca vitulina ?
Marine Invertebrates
Crustaceans
Aesop Shrimp Pandalus montagui 2
Atlantic Rock Crab Cancer irroratus ?
Atlantic Lyre Crab Hyas araneus ?
King Crab Neolithodes grimaldi ?
Molluscs
Iceland Scallop Chlamys islandica 7
Blue Mussel Mytilus edulis ?
Softshell Clam Mya arenaria ?
Arctic Surfclam Mactromeris polynyma 2
Atlantic Surfclam Spisula solidissima ?
Ocean Quahog Arctica ilandica ?
Waved Whelk Buccinum undatum ?
Northern Shortfin Squid Illex illecebrosus ?

+Updated April 1985 North Atlantic stock downlisted to Vulnerable.

some of these may be of no immediate concern, the obtain updates on the status of selected species as
Subcommittee will, as opportunity allows, attempt to new information becomes available, or in the 10-
document these species to determine their status in year review process initiated in 1993 (Table 3) for
Canada. those species which had not already received fur-

In addition to soliciting further status reports on ther examination following their initial status
species of concern, the Subcommittee continues to assignment.

256

The Canadian Endangered Species
Protection Act: A Legislative Proposal

The Canadian Government is considering proposals
for the establishment of a Canadian Endangered
Species Protection Act which could be in place by
1997. The goal of the Act and the processes which
flow from it would be to prevent wild Canadian
species from becoming extinct (as a consequence of
human activities) and to recover species where possi-
ble and economically feasible. It is built around and
would embody the COSEWIC framework and pro-
vide a legal status to its mandate (see Environment
Canada 1995). However, the Act would apply only to
species within the federal mandate (In Canada, as a
result of the listing of federal and provincial powers in
the British North America Act and recapitulated in the
Constitution, wildlife is the property of the province
or territory in which it resides, except for those
species explicitly under the federal mandate; i.e., fish
and marine mammals and migratory birds.), and
would also exclude native peripheral species. Most
invertebrates, vascular plants, mosses, lichens, rep-
tiles, amphibians, non-migratory birds, and terrestrial
mammals do not fall under a federal mandate and
given the scope of the proposed Act, it is difficult to
see how these would ever get any national considera-
tion by COSEWIC in its new form, given that they
would only be of provincial interest.

With the current economic climate, the restraints
on government resources from within and without,
and the lack of provincial will in the national inter-
est, there is some fear that the enablement of the leg-
islation as proposed would set wildlife conservation
in Canada back forty years.

Concluding Remarks

The six reports included in the following series
are reports on the status of the respective species in
Canada. Status was assigned by consensus of the
COSEWIC Committee based on these reports which
are published under the name(s) of the original
author(s). The reports have undergone minor editing
to provide a brief introduction and some degree of
consistency in format and presentation.

As to the legislative proposal for a Canadian
Endangered Species Act, one should not be overly
pessimistic to such a colloquial Canadian bureau-
cratic approach to the situation. Canadians are great
infighters and when a job is to be done, a way will
be found to do it. Just look at COSEWIC and what it
has achieved in 20 years without any legislative
basis or legal mandate.

Acknowledgments

The members of COSEWIC and the Fish and
Marine Mammal Subcommittee would like to extend
their thanks to the various authors who have so gen-
erously contributed their time and talents in support

THE CANADIAN FIELD-NATURALIST

Vol. 111

of COSEWIC. The Committee also wishes to
acknowledge the members of the Subcommittee for
their unstinting efforts in reviewing the reports and
their helpful comments.

COSEWIC is grateful to the Canadian Wildlife
Federation, World Wildlife Fund Canada, the
Canadian Wildlife Service, the Canadian Museum
of Nature, and the Royal Ontario Museum for assis-
tance provided in cash and kind. A special mention
to The Canadian Field-Naturalist for assistance in
publication and editing and to all members of
COSEWIC for their dedication and interest in the
future of Canada’s flora and fauna. We gratefully
acknowledge the financial support provided by
Environment Canada, the Province of British
Columbia and the Canadian Nature Federation
which permitted the contracting of several new
reports.

Literature and Documents Cited

Campbell, R. R. 1984. Rare and endangered fishes of
Canada: The Committee on the Status of Endangered
Wildlife in Canada (COSEWIC) Fish and Marine
Mammal Subcommittee. Canadian Field-Naturalist
98(1): 71-74.

Campbell, R. R. 1985. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammals Subcommittee Status Reports: II.
Canadian Field-Naturalist 99(3): 404408.

Campbell, R. R. 1987. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: III.
Canadian Field-Naturalist 101(2): 165-170.

Campbell, R. R. 1988. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: IV.
Canadian Field-Naturalist 102(1): 81-86.

Campbell, R. R. 1989. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: V.
Canadian Field-Naturalist 103(2): 147-157.

Campbell, R. R. 1990. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: VI.
Canadian Field-Naturalist 104(1): 1-6.

Campbell, R. R. 1991. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: VII.
Canadian Field-Naturalist 105(2): 151-156.

Campbell, R. R. 1992. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: VIII.
Canadian Field-Naturalist 106(1): 1-6.

Campbell, R. R. 1993. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: IX.
Canadian Field-Naturalist 107(4): 395401.

Campbell, R. R. 1994a. An updated report on the status of
the Spotted Gar, Lepisosteus oculatus, in Canada.
Report to the Committee on the Status of Endangered
Wildlife in Canada (COSEWIC), Canadian Wildlife
Service, Ottawa, Ontario. 7 pages.

1997

Campbell, R. R. 1994b. An updated report on the status of
the Spotted Sucker, Minytrema melanops, in Canada.
Report to the Committee on the Status of Endangered
Wildlife in Canada (COSEWIC), Canadian Wildlife
Service, Ottawa, Ontario. 6 pages.

Campbell, R. R. 1996. Rare and endangered fishes and
marine mammals of Canada: COSEWIC Fish and
Marine Mammal Subcommittee Status Reports: X.
Canadian Field-Naturalist 110(3): 454461.

Carlton, J. T., G. J. Vermeij, D. R. Lindberg, D. A.
Carlton, and E. C. Dudley. 1991. The first historical
extinction of a marine invertebrate in an ocean basin: the
demise of the Eelgrass Limpet, Lottia alveus. Biological
Bulletin 180: 72-80.

Cook, F. R., and D. Muir. 1984. The Committee on the
Status of Endangered Wildlife in Canada (COSEWIC):
History and progress. Canadian Field-Naturalist 98(1):
63-70.

CWS. 1993. RENEW: Recovery of nationally endangered
wildlife. Report #3. Canadian Wildlife Service, Ottawa,
Ontario CW70-2/1993: 1-55.

CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA: STATUS REPORTS

257

CWS. 1995. The Canadian Endangered Species
Protection Act: A Legislative Proposal. Minister of
Supply and Services Canada. Ottawa, Ontario. CW 66-
143/1995.

Parker, B., P. McKee, B. Hindley, M. Brinkman, M. J.
Fenton, K. Rowan, J. Tanner, and C. Goddard.
1980. Rare, threatened and endangered fish species of
southern Ontario: Status reports. Beak Consultants
report to Department of Supply and Services,
Department of Fisheries and Oceans and the National
Museum of Natural Sciences, Ottawa, Ontario. 238
pages.

Parker, B., and P. McKee. 1984a. Status of the Spotted
Gar, Lepisosteus oculatus, in Canada. Canadian Field-
Naturalist 98(1): 80-85.

Parker, B., and P. McKee. 1984b. Status of the Spotted
Sucker, Minytrema melanops, in Canada. Canadian
Field-Naturalist 98(1): 104-109.

Accepted 8 July 1996

Status of the Nooksack Dace, Rhinichthys sp., in Canada*

J.D. McPHAIL
Biodiversity Centre and Department of Zoology, University of British Columbia, Vancouver, British Columbia V6S 1Z4

McPhail, J. D. 1997. Status of the Nooksack Dace, Rhinichthys sp., in Canada. Canadian Field-Naturalist.
111(2): 258-262.

The Nooksack Dace (Rhinichthys sp.) is a morphologically distinctive form derived from the common, widely distributed
Longnose Dace, Rhinichthys cataractae. Like the Salish Sucker (Catostomus sp.), it is a component of the Chehalis fauna.
This fauna survived the last glaciation in the ice-free area west of the Cascade Mountains and north of the Columbia
drainage system. In Canada, the Nooksack Dace is confined to four small streams tributary to the Nooksack River in the
Abbotsford, Aldergrove and Clearbrook areas of the lower Fraser Valley, southwestern British Columbia. The adult habitat
is riffles, typically with water velocities close to 0.25 m-sec and a loose, coarse gravel substrate. Young-of-the-year inhabit
shallow margins at the tail-ends of pools with mud/sand substrates, and typically recruit to the adult habitat after one year.
Although the Nooksack Dace is still moderately common, suitable habitat is deteriorating rapidly through explosive urban-
ization and gravel extraction in the headwaters of the streams. In western Washington state, the Nooksack Dace is
widespread and in no immediate danger but, in Canada, given its restricted distribution and deteriorating habitat it is endan-
gered.

Le naseux de la Nooksack (Rhinichthys sp.) est une forme morphologiquement distincte dérivée d’un poisson commun,
largement réparti, le naseux des rapides (Rhinichthys cataractae). Comme le meunier de Salish (Catostomus sp.), il fait
partie de la faune du Chehalis, faune qui a survécu a la derniére glaciation dans des espaces libres de glace, a l’ouest des
Cascade Mountains et au nord du bassin hydrographique du Columbia. Au Canada, le naseux de la Nooksack est confiné a
quatre ruisseaux tributaires de la riviére Nooksack, dans les secteurs d’ Abbotsford, d’ Aldergrove et de Clearbrook de la
partie inférieure de la vallée du Fraser, dans le sud-ouest de la Columbie Britannique. Comme habitat, les adultes adoptent
les rapides, particuli¢rement ceux qui ont une vélocité de prés de 0,25 m/s avec des substrats de gros gravier meuble. Les
jeunes de l’année habitent les recoins peu profonds, aux substrats de boue sablonneuse, en marge des étangs, et se retrou-
vent naturellement dans |’habitat des adultes aprés une année. Méme si le naseux de la Nooksack est relativement commun,
son habitat de prédilection se détériore rapidement a cause d’une urbanisation frénétique et de l’extraction de gravier a la
source des cours d’eau. Dans |’ ouest de |’état de Washington, le naseux de la Nooksack est trés répandu et ne court aucun
danger immédiat, mais au Canada, vu sa faible distribution et le déclin de son habitat, il est en danger de disparition.

Key Words: Nooksack Dace, naseaux de la Nooksack, Rhinichthys sp., cyprinids, endangered species, British Columbia.

The Nooksack Dace (Rhinichthys sp.) is a small
(up to about 105 mm in standard length) cyprinid
fish widely distributed in the clear, relatively rapid
rivers and streams of western Washington (McPhail
1967). In Canada, it is restricted to a few small trib-

conspicuous black mid-lateral stripe that extends
from the snout back to a diffuse dark spot at the
base of the tail.

Distribution

utaries of the Nooksack River in the lower Fraser
Valley, southwestern British Columbia (McPhail
and Lindsey 1986). It is a slim fish with a snout
that clearly overhangs the mouth, a streamlined
back and a flattened underside (Figure 1). In life,
adults are grey-green above with a dull brassy
stripe just above the lateral line. The sides below
the lateral line are dirty white grading into silver-
white on the underparts. Often there are scattered
dusky speckles on the sides below the lateral line
and a black stripe on the head in front of the eyes.
Viewed from above, there is a distinct pale mark at
the anterior and posterior base of the dorsal fin.
There is no striking sexual dimorphism in colour,
but males have conspicuously longer, and darker,
pectoral fins than females. In juveniles there is a

This distributional account is based on collec-
tions in the fish museums of the Department of
Zoology, University of British Columbia, and the
School of Fisheries, University of Washington. The
geographic distribution of the Nooksack Dace
forms a rough fish-hook around Puget Sound
(Figure 2) although, curiously, it is absent from
rivers on the west side of the Sound. On the west
side of the Olympic Peninsula it occurs from the
Queets system in the north to the Willipa River in
the south. On the east side of Puget Sound it
extends from the Pyuallup River in the south to the
Nooksack River in the north. The species is
widespread in the Chehalis system, but so far is
unrecorded from the Deschutes and Nisqually
rivers near Olympia, Washington. In Canada, the

*Reviewed and approved by COSEWIC 16 April 1996 —staus assigned: Endangered.

258

1997

McPHAIL: STATUS OF THE NOOKSACK DACE

259

FIGURE |. Nooksack Dace, Rhinichthys sp., 81 mm standard length.

Nooksack Dace is restricted to Nooksack tributaries
in the lower Fraser Valley of British Columbia;
Bertrand, Cave, Fishtrap and Pepin creeks (see inset
Figure 2).

Protection

No special protection is in place for the Nooksack
Dace; however, its Canadian distribution lies entirely
within the range of the Salish Sucker (Catostomus
sp.), a species assigned “Endangered” status in April
of 1986 by COSEWIC and in British Columbia, allo-
cated to the Ministry of Environment’s critically
imperiled category. Consequently, measures taken
by the B.C. Ministry of Environment, Lands and
Parks to protect the Salish Sucker also protect the
Nooksack Dace.

Population Sizes and Trends

Although the geographic distribution of the
Nooksack Dace in Canada is limited, museum
records from the 1960s suggest that the species once
was abundant within this restricted range.
Unfortunately, no formal estimates of past, or pre-
sent, numbers are available for the Nooksack Dace;
however, an October 1993 survey of five riffles in
Bertrand Creek revealed an average density of 1.4
(SE = + 0.24) adults m*. Furthermore, the popula-
tions in Bertrand, Cave, Fishtrap and Pepin creeks
appear healthy: a range of year classes is present in
each of the creeks, including substantial numbers of
young-of-the-year, although in 1993 the 1+ year
class was under represented both in our survey and
in a summer survey supported by the Habitat
Conservation Fund (S. Inglis, B.C. Ministry of
Environment, Lands and Parks, Victoria, British
Columbia; personal communication). Even though
the existing populations appear healthy, there is evi-
dence that suggests the British Columbia distribution
is shrinking. For example, Nooksack Dace are now
absent from some of the smaller tributaries (e.g.,
Howes Creek) and headwaters of both Bertrand and
Fishtrap creeks where they occurred in the 1960s. In

these creeks, rapid urbanization in the Aldergrove,
Clearbrook and Abbotsford regions has increased sil-
tation, pollution and fluctuations in discharge. The
result is deteriorating water quality and a loss of
habitat that is spreading downstream. Near the
United States border, the streams are in better condi-
tion. Here, they flow through a rural landscape, but
even in this area land clearing and gravel extraction
generate silt and decrease summer flows. By late
August most small tributaries are dry and, in drought
years, even the main creeks (Bertrand and Fishtrap)
are reduced to trickles. Since adult Nooksack Dace
depend on riffles with loose gravel substrate for for-
aging and breeding (see sections on Habitat and
General Biology), silt and low summer flows are
especially harmful to this species. Low summer
flows restrict adult habitat at the most productive
time of the year by reducing the riffle areas, and silt
tends to fill the interstices between rocks and cement
them into the substrate. This reduces both the cover
available to adults and their foraging opportunities.
Under low water conditions, adults will shift into
pools but they probably do not grow or survive as
well in this marginal habitat as they do in riffles.
Consequently, given that the demand for both hous-
ing and gravel in the lower Fraser Valley will contin-
ue to increase, habitat suitable for adult Nooksack
Dace will continue to decrease and the species prob-
ably will go extinct in Canada in the next one or two
decades.

Habitat

Adult Nooksack Dace are demersal and typically
occur in riffles with water velocities greater than
0.25 m** and a substrate of loose gravel (4-10 cm in
diameter), cobbles, or boulders. In Canada, the
species is associated with small to moderate sized
streams (1 to 4 m in width); however, this associa-
tion probably simply reflects the absence of large
Nooksack tributaries in Canada; whereas in
Washington, the species regularly occur in large
rivers. In contrast to adults, young-of-the-year are

260

TABLE 1. Habitat differences between adult and young-of-
year Nooksack Dace, Rhinichthys sp., Bertand Creek, 1

October 1993.
Adults Young-of-year

Average Water

Veocities 0.34 + 0.039 ms* 014+0.022 ms
Substrate fist-sized gravel, mud, sand,
cobbles and leaflitter

boulders

associated with slow water (0.14 + 0.022m‘* near the
downstream ends of pools (Table 1). Here, the
young occur in shallow water (about 10 to 20 cm
deep) but swim above the sand or mud substrate.
Thus, the species occupies two basic stream habitats:
adults in fast water over loose rock substrates, and
juveniles in quiet waters over sand or mud sub-
strates.

General Biology

In British Columbia, Nooksack Dace spawn in the
spring (April through May). Spawning apparently
occurs at night and the eggs are usually found near
the top ends of riffles. There is no evidence of the
spawning colours reported for Longnose Dace
(Rhinichthys cataractae) in Manitoba (Bartnik
1972). Both males and females mature at the end of
their second summer (1*) and breed in their third
spring (2*). To date, the oldest recorded individual (a
female 105 mm in standard length) was in her sixth
year (5*). As in most fish, egg number is a function
of female size, and in the Nooksack Dace fecundity
ranges from about 200 to over 2000 eggs. In sum-
mer, adults feed primarily on riffle dwelling insects
(e.g. nymphs of caddisflies and mayflies, dytiscid
beetle larvae and adult riffle beetles). In pools the
young feed primarily on chironomid pupae and
ostracods. Adults collected at mid-morning have
empty stomachs but packed hind guts. This suggests
nocturnal feeding. In the lower Fraser Valley, adults
appear to inhabit riffles throughout the year, but in
less benign climates they may shift to slower, deeper
water in the winter.

Limiting Factors

Habitat loss through human disturbance is the
greatest threat facing the Nooksack Dace in British
Columbia. Around Aldergrove, Clearbrook and
Abbotsford, housing developments, shopping malls
and industrial parks are replacing fields and wooded
areas at a dizzying pace. This accelerating urbaniza-
tion brings with it all the usual environmental prob-
lems, compounded in this case by the development
being in the headwaters of the streams. Thus, both
Bertrand and Fishtrap creeks and their tributaries are
vulnerable to the usual fate of urban streams (e.g.,

THE CANADIAN FIELD-NATURALIST

Vol. 111

straightening, siltation, industrial and domestic
chemical spills, and clandestine garbage disposal), as
well as attempts to “aesthetically improve” the
streams by creating parks and ponds that please the
human eye but destroy critical fish habitat. In the
past, accidental fish kills in the Canadian portion of
the Nooksack system would have been followed by
natural recolonization from the main river. This is no ,
longer the case. The U.S. portions of Bertrand and
Fishtrap creeks are ditched and silted. They no
longer contain either Nooksack Dace or habitat suit-
able for Nooksack Dace. Thus, the shrinking
Canadian populations are sandwiched between a
deteriorating environment upstream and unsuitable
habitat downstream.

Special Significance of the Species

The Nooksack Dace is a member of the Chehalis
fauna (McPhail 1967, 1987; McPhail and Lindsey
1986). This isolated fauna is derived from the
Columbia fauna and, with the exception of the
endemic Olympic Mudminnow (Novumbra hubbsi),
all the species are closely related to Columbia
species. These Chehalis isolates diverged from their
Columbia counterparts sometime before the last
(Fraser or Vashon) glaciation, and survived the ice-
sheets south of Puget Sound but north of.the
Columbia River. Consequently, their geographic dis-
tributions include the Chehalis River, the rivers on
the west side of the Olympic Peninsula and some
rivers on the east side of Puget Sound. In two cases,
the Salish Sucker and Nooksack Dace, Chehalis iso-
lates have dispersed postglacially as far north as the
lower Fraser Valley (Figure 2).

The Nooksack Dace is a typical Chehalis isolate: it
is related to, and presumably derived from, the west-
ern North American form of a widely distributed
species, the Longnose Dace (Rhinichthys cataractae).
It differs from this species in scale counts (McPhail
1967; Bisson and Reimers 1977), body shape and in
consistent sequence differences in both mitochondrial
and nuclear genes. For example, the mitochondrial
sequence differences between Nooksack and
Longnose Dace are comparable to the sequence dif-
ferences between such well established species as
Largescale Sucker (Catostomus macrocheilus) and
Longnose Suckers (Catostomus catostomus)
[McPhail and Taylor, in preparation]. The Nooksack
Dace also shows the characteristic distribution pat-
tern of a Chehalis isolate: scattered populations in the
Chehalis River and populations in rivers draining the
west side of the Olympic Peninsula and the east side
of Puget Sound (Figure 2).

Typically, the genetic separation between
Chehalis isolates and their Columbia relatives, as
measured by allozyme frequencies or gene
sequences, is greater than their morphological sepa-
ration (McPhail and Lindsey 1986; McPhail and

1997

ee

GEORGIA
STRAIT

OLYMPIC O

~
STRAIT
PENNINSULA Q

Quillayute R.

Copalis ey
Wa

Humptulips R
d

Chehalis R:

MCcPHAIL: STATUS OF THE NOOKSACK DACE

261

Nooksack Dace
@ - present
- absent

Fishtrap
Creek

CANADA

CANADA

— —- = ame

Nooksack Dace

@ - present
- absent from apparently
suitable habitat

FIGURE 2. Geographic range of the Nooksack Dace, Rhinichthys sp. inset depicts Canadian distribution.

Taylor, in preparation). Still, Chehalis isolates usual-
ly can be separated from their Columbia counterparts
by a combination of morphological traits. Because
the geographic distributions of most Chehalis iso-
lates do not overlap with their closest relatives, any
decisions regarding their taxonomic status (e.g.,
species or subspecies) are necessarily arbitrary; how-
ever their status as independant, divergent lineages

are not debatable, and for conservation purposes
they should be regarded as species. Certainly, their
distinctive morphologies, gene sequences and char-
acteristic geographic distributions argue that the
Nooksack Dace has not exchanged genes with the
Longnose Dace for a long time (i.e., since well
before the beginning of the last galciation and, per-
haps, since before the Pleistocene).

262

Evaluation

The Canadian distribution of the Nooksack Dace
probably will continue to shrink as long as the
Vancouver megalopolis continues to expand. To stop
the decline will require a concerted effort by all lev-
els of government to protect the remaining free-
flowing streams occupied by this species ... an
unlikely event. Even if the political will is there,
accidents are inevitable in urban streams, especially
with a drinking-water supply that for public health
reasons will require chlorination in less than a
decade. Once this happens, it will be a minor miracle
if, in Canada, the Nooksack Dace survives into the
next century.

Acknowledgments

The illustration of the Nooksack Dace was drawn in
“Freehand” on a Macintosh computer by Diana
McPhail. Marvin Rosenau, Juanita Ptolemy and Susan
Inglis (British Columbia Ministry of Environment,
Lands and Parks) helped in many ways. Over the
years, Ron Jones, Dave Greenfield, Clyde Murray and
Gordon Haas helped me outline the nature and distri-
bution of the Chehalis fauna. I am grateful for their

THE CANADIAN FIELD-NATURALIST

Vol. 111

support and enthusiasm. More recently, Mike Folkes
assisted in the field, Ruth Withler did an allozyme sur-
vey, and Claire Thompson and Ric Taylor did DNA
analyses. The Fisheries Branch, B.C. Ministry of
Environment, Lands and Parks, commissioned and
funded this report.

Literature Cited

Bartnik, V. 1972. Comparison of the breeding habits of
the two subspecies of Longnose dace, Rhinichthys
cataractae. Canadian Journal of Zoology 50 (1): 83-86.

Bisson, P. A., and P. E. Reimers. 1977. Geographic
variation among Pacific Northwest populations of long-
nose dace, Rhinichthys cataractae. Copeia 1977 (3):
518-522.

McPhail, J. D. 1967. Distribution of freshwater fishes in
western Washington. Northwest Science 41: 1-11.

McPhail, J. D., and C. C. Lindsey. 1986. Zoogeography
of the freshwater fishes of ascadia (the Columbia system
and rivers north to the Stikine). Pages 615-637 in The
Zoogeography of North American Freshwater Fishes.
Edited by C. H. Hocutt and E. O. Wiley. John Wiley and
Sons, New York. 866 pages.

Accepted 8 July 1996

Status of the Northern Fur Seal, Callorhinus ursinus, in Canada*

ROBIN W. BAIRD! and M. BRADLEY HANSON2

'Marine Mammal Research Group, Box 6244, Victoria, British Columbia V8P 5L5 and Biology Department, Dalhousie
University, Halifax, Nova Scotia B3H 4J1

?National Marine Mammal Laboratory, National Marine Fisheries Service, 7600 Sand Point Way N.E., Building 4, Seattle,
Washington 98115

Baird, Robin W., and M. Bradley Hanson. 1997. Status of the Northern Fur Seal, Callorhinus ursinus, in Canada.
Canadian Field-Naturalist 111(2): 263-269.

This report reviews the general biology, status, and management of the Northern Fur Seal (Callorhinus ursinus), with spe-
cial reference to its status in Canadian waters. While Northern Fur Seals do not breed within Canadian waters, they can be
found in large numbers in the waters offshore of British Columbia year-round, and occasional stragglers are found inshore.
Generally found only in small groups during the pelagic phase of their life, the largest numbers occur in British Columbia
waters from January through June. The eastern North Pacific population has declined significantly over the last 30 years,
but the cause is unknown.

Ce rapport examine en général la biologie, le statut et la gestion de |’otarie 4 fourrure, (Callorhinus ursinus) avec référence
particuliére 4 son statut dans les eaux canadiennes. Bien que les otaries a fourrure ne se reproduisent pas dans les eaux cana-
diennes, on les trouve quand méme toute |’année en grand nombre au large de la céte de la Colombie-Britannique et
quelques errants viennent parfois dans les eaux cétiéres. Bien que généralement ils ne forment que de petits groupes pendant
la phase pélagique de leur vie, c’est dans les eaux de la Colombie-Britannique, de janvier 4 juin, que |’on trouve les plus
grands regroupements. La population de |’est du Pacifique nord a connu un important déclin ces trente derniéres années,

mais la cause est inconnue.

Key Words: Northern Fur Seal, Callorhinus ursinus, Otarie a fourrure, otariid, status, British Columbia.

This review of the general biology, status, and
management of the Northern Fur Seal, Callorhinus
ursinus (Linnaeus 1758), was prepared on behalf of
the Committee on the Status of Endangered Wildlife
in Canada (COSEWIC). A compilation and assess-
ment of the available information on this species is
used to evaluate its status in Canadian waters. For
this species such a task is relatively easy in compari-
son to the many lesser known species of marine
mammals in Canadian waters, because of the exten-
Sive investigations undertaken by U.S., Russian, and
Canadian researchers.

The Northern Fur Seal, a member of the sea lion
family (Otariidae), is the second smallest pinniped
found on the west coast of Canada. Adult males
reach a length of 1.9 m and a weight of about 200
kg, while females are much smaller, reaching a
length of 1.3 m and a weight of 35 kg (Figure 1).
The pelage of adults is generally a brownish-grey
colour. The vibrissae colour varies with age, being
black in juveniles and white in fully grown adults.

Distribution

Northern Fur Seals range throughout the northern
Pacific from central Japan (latitude 36°N) and the
Sea of Japan north to the Bering Sea, and south along
the west coast of North America to the area of the

U.S.-Mexican border (latitude 32°N) [Figure 2]. Fur
seals can be found throughout this range in almost all
months of the year, but peak abundance varies sea-
sonally and geographically. Off the Canadian west
coast, females and subadult males are typically found
during the winter off the continental shelf (Bigg
1990). Occasional animals are seen in inshore waters
in British Columbia, and stragglers occasionally
come ashore, usually at sea lion haulouts (e.g., Race
Rocks, off southern Vancouver Island).

Three breeding colonies occur in Russia; at
Robben (Tyuleniy) Island and the central Kuril
Islands in the Sea of Okhotsk, and Commander
Island in the western Bering Sea. In the United
States, colonies occur at the Pribilof Islands (St.
George and St. Paul islands) in the eastern Bering
Sea, at Bogoslof Island in the southeast Bering Sea,
and at San Miguel Island and nearby Castle Rock off
southern California. Reeves et al. (1992) noted that a
few fur seals also haul out seasonally on Southeast
Farallon Island and occasionally on San Nicolas
Island, off California.

Protection
International

The Interim Convention on Conservation of North
Pacific Fur Seals lapsed in 1984, when the United

*Reviewed and approved by COSEWIC 16 April 1996 —staus assigned: Not at Risk (NAR).

263

264

States Senate failed to ratify a protocol for extension.
This international agreement protected the fur seal
from hunting at sea, but also allowed for the com-
mercial harvest of fur seals in the Pribilof Islands.
Attempts to establish a new treaty for the protection
of the fur seal have failed; consequently the species is
vulnerable to future hunting of animals at sea. Under
the terms of this agreement Canada received 15 per-
cent of the skins from harvests, and was also obliged
to undertake research on this species.

The Northern Fur Seal is not listed under the
Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES), thus
international trade is not monitored or regulated.

National

Canada: In Canada the Northern Fur Seal is pro-
tected under the 1993 Marine Mammal Regulations
of the Fisheries Act of Canada. These regulations
control the hunting of this species in Canadian
waters by all except aboriginals, who are required to
obtain a licence. Fees for such a licence are low ($5),
and issuance is at the discretion of the Minister of
Fisheries and Oceans. “Disturbance” is prohibited
through these regulations, except when hunting
under licence, as is the operation of aircraft within
600 m of any live seal on land.

United States: After the international convention
lapsed in 1984, management of this species in U.S.
waters became subject to the Fur Seal Act of 1966
and the Marine Mammal Protection Act of 1972.
The commercial harvest on the Pribilof Islands
ended because the National Marine Fisheries Service
(NMES) determined that such a harvest could not
take place under domestic laws. Current levels and
conditions for a subsistence harvest are regulated
under these Acts. In 1988 the U.S. population was
listed as depleted under the Marine Mammal
Protection Act. Under this Act, a conservation plan
was prepared for this species (National Marine
Fisheries Service 1993), outlining protective mea-
sures and research activities to be undertaken by the
U.S. National Marine Fisheries Service.

Population Size(s) and Trends

Several stocks have been generally recognized for
the Northern Fur Seal; however, tag returns suggest
that some exchange occurs between breeding
colonies. The Pribilof Islands population is the
largest, comprising three quarters of the world’s
total, with just less than one million individuals in
the early 1990s (National Marine Mammal
Laboratory, unpublished data). This population was
thought to number between two to three million indi-
viduals in the mid-1800s (National Marine Fisheries
Service 1993), and was estimated at approximately
2.1 million individuals in the 1940s (Kenyon et al.
1954; Lander and Kajimura 1982; Briggs and
Fowler 1984). Based on pup counts, the population

THE CANADIAN FIELD-NATURALIST

Vol. 111

on St. Paul Island (representing about 80% of the
Pribilof stock) has remained relatively stable since
the early 1980s, while the St. George stock has
undergone a significant decline since the late 1970s
(York and Fowler 1992). The Pribilof Island popula-
tion(s) appear to be well below the level of maxi-
mum net productivity (Ragen 1995). Northern Fur
Seals first began breeding at Bogoslof Island in the
southeast Bering Sea in 1980, and the population in
1988 comprised over 400 individuals, increasing at a
rate of 57% per year (Loughlin and Miller 1989).
Fur seals also began breeding at San Miguel Island
(California) in 1968 and at nearby Castle Rock in
1972. The San Miguel colony numbered approxi-
mately 5000 individuals by 1993 and was increasing
(DeLong et al. 1993).

Habitat

Terrestrial habitats for Northern Fur Seals are
generally limited to their rookeries, which are scat-
tered around the North Pacific rim in close proximi-
ty to the continental slope. Fur seals have strong
fidelity to traditional sites that are typically com-
posed of a rocky substrate (on San Miguel Island
they use a sand beach). Although all rookeries in the
U.S. are federally owned, the Pribilof Islands are
inhabited and non-rookery lands are owned by local
communities which are developing support facilities
for fishing industries. Consequently, several types
of habitat degradation are possible.

Ocean habitats appear to vary by the sex and age
group(s) of fur seals present. In the summer breeding
season, continental slope waters in the eastern Bering
Sea are the principal destination for adult females on
foraging trips. The use of continental shelf and slope
waters of British Columbia and the states of
Washington, Oregon, and California by adult females
during winter is well documented from pelagic seal-
ing data (Bigg 1990). Adult males from the Alaskan
populations appear to remain in Alaskan waters year-
round, some remaining in the Bering Sea and some
moving south into the Gulf of Alaska in mid-winter.
Subadults of both sexes use coastal waters of British
Columbia and Washington as well as offshore areas
of the North Pacific (Kajimura 1984; Bigg 1990).
The highest concentrations in the open ocean occur
along the continental shelf break and in association
with other major oceanographic frontal features, such
as canyons, sea mounts and valleys (Kajimura 1984).
Although water quality in these areas is unknown, it
appears to be suitable with the exception of areas
subjected to oil spills. Trends in the availability of
prey in these areas are unknown.

General Biology
Reproduction

Female Northern Fur Seals may produce their first
pup at four years of age, and the majority are pup-
ping by six years of age (York and Hartley 1981). A

1997

BAIRD AND HANSON: STATUS OF NORTHERN FUR SEAL

265

4

FiGurE 1. Northern Fur Seals. Photo by O.W. Otsen. The small animal at left is an adult female, the large animal is an

adult male.

single pup is born within two days of the arrival of a
female at the breeding colony. Parturition appears to
be stimulated by arrival at the breeding colony and
the presence of conspecifics (Bigg 1984). For the
first 8-10 days after birth, females remain ashore
until they come into estrous and mate. After that
time they regularly leave the colony for foraging
trips of 4-10 days. Insley (1992) demonstrated that
calls used between mother-offspring pairs of
Northern Fur Seals were highly variable between
individuals, but varied little for a particular individu-
al, allowing the mother and pup to reunite after sepa-
rating. Pups remain on the breeding islands until
they are weaned in late fall. Pup weight depends in
part on that of the mother and on sex, male pups are
larger at birth (Boltnev 1993). Adult males remain
ashore and fast while defending breeding territories.
Juvenile males also haul out during the breeding sea-
son and fast, typically losing about 20% of their
body mass during this time (Baker et al. 1993).
Diet

Knowledge of the diet of the Northern Fur Seals
comes primarily from the examination of stomach
contents of animals killed as part of a joint U.S.-
Canadian research program from 1958 to 1974

(Kajimura 1984; Perez and Bigg 1986), and in part
from scat analysis (Antonelis et al. 1990, 1993).
Primary prey species vary seasonally (Perez and
Bigg 1986). Differences in diet also occur through-
out their range, both on a large (Perez and Bigg
1986) and small scale (Antonelis et al. 1993). Small
schooling fishes are the primary food species in
terms of energy content. In British Columbia waters,
Pacific Herring (Clupea harengus) and various
species of squid comprise about 70% of the diet.
Walleye pollock, sablefish, rockfishes, whiting, and
salmonids form the remainder of the diet (Perez and
Bigg 1986). Historical evidence suggests that the
composition of the diet has fluctuated over time with
changes in fish stocks; sardines were once extremely
abundant in the eastern North Pacific but were over-
fished in the 1940s until the stocks collapsed. There
is some evidence that sardines were commonly eaten
by Northern Fur Seals off Vancouver Island in the
early 1930s (Clemens and Wilby 1933).

Movements

In general, Pribilof Island fur seals migrate south
to winter along the west coast of Canada and the
United States. However, patterns of movement of
this species are extensive and complex, with timing

266

and migratory routes depending on age, sex and
reproductive condition (Bigg 1990). Bigg (1990)
suggested that fur seal migration from the Bering
Sea after the breeding season facilitated both the
avoidance of low temperatures and access to
sources of prey. He also suggested that the age-
related differences in migratory timing and routes
likely results from a combination of the learning of
productive foraging areas and the need to return to
the breeding areas when animals reach reproductive
age. The origin of sex-related differences in migra-
tion may result in part from differences in the tim-
ing of arrival to and departure from the breeding
colonies. Adult females both arrive at and leave
colonies later than adult males. Males start to arrive
at colonies in May to establish territories and
females start arriving in mid-June. Males depart in
late summer while females remain until late fall
while they continue to nurse their young. The fall
movements of pups away from the rookeries is not
random; estimated minimum swimming speed of
pups between St. Paul Island and the Aleutians was
between 36 to 61 km/day (Ragen et al. 1993).
Fowler et al. (1993) demonstrated that male
Northern Fur Seals generally return to their natal
rookery, although individuals sometimes emigrate
to other rookeries or, rarely, form new colonies.

Short-distance movements around San Miguel
Island by females nursing pups were examined by
Antonelis et al. (1990), who found that females for-
aged primarily in oceanic waters over the continental
slope. These females departed the colony in greatest
numbers in mid-day, possibly a thermoregulatory
behaviour in response to increasing temperatures and
solar radiation.

Behaviour

Northern Fur Seals generally exhibit strong site
fidelity to their rookeries. Despite disturbance asso-
ciated with commercial harvesting by human inhabi-
tants on the Pribilof Islands from the late 1700s until
recently, seals continue to occupy nearly all the same
rookeries. This species appears to be tolerant to short
term disturbance associated with human activities
(Gentry and Gilman 1990). The proximity of these
islands to the continental slope likely contributes to
their continued use of these rookeries.

Limiting Factors

Commercial harvesting has affected the population
of Northern Fur Seals in the Pribilof Islands since
shortly after it was first discovered in 1786 (National
Marine Fisheries Service 1993). From 1786 to 1828
an average of a hundred thousand Northern Fur Seals
per year, primarily pups, were killed. Commercial
harvesting of this species was directly responsible for
the large reductions in population size in the late
1800s and early 1900s. The commercial harvest dur-
ing this early period included pregnant females; dur-

THE CANADIAN FIELD-NATURALIST

Vol. 111

ing the period of pelagic sealing, large numbers of
animals were taken off British Columbia and in the
Bering Sea (National Marine Fisheries Service 1993).
Hunts were reduced in size in the early 1900s, and
the population grew up to the 1940s. Approximately
300 000 females were killed between 1956 and 1968,
in an effort to move the population towards the level
where productivity would be maximized. The popu-
lation did not respond as expected however, and pup
production decreased (York and Hartley 1981).
When the commercial harvest of females ceased in
1968, pup production increased and the population
grew until 1976. While hunting continued during this
period, it is considered unlikely to be the cause of a
decline in the population size after 1976 (National
Marine Fisheries Service 1993). Commercial harvests
continued for this species up until the expiration of
the Interim Convention on Conservation of North
Pacific Fur Seals in 1984. Since that time, between
about 1200 and 3700 juvenile males have been killed
each year as part of a subsistence harvest. Such levels
are not thought to contribute to the lack of recovery
of the population (National Marine Fisheries Service
1993).

A variety of other natural and anthropogenic
sources of mortality for this species have been
observed. Several authors have examined causes
and levels of mortality in pups, both at breeding
colonies and during the winter. Calambokidis and
Gentry (1985) observed that pup survival from birth
to weaning was positively correlated with birth
weight, which in turn was correlated with the age of
the mother. Pups which weighed less than average
were more likely to die from trauma, parasitic infes-
tation and infectious disease, as well as “emaciation
syndrome”. Baker and Fowler (1992) examined pup
weight and overwintering survival, and found that
the overwintering survival of males increased with
pup weight. Their small female sample size likely
precluded a similar determination (Baker and
Fowler 1992). They suggested that larger over-
wintering animals were better able to withstand cold
temperatures. Larger animals are also able to dive
longer (Kooyman 1989 in Baker and Fowler 1992),
possibly increasing the ability to find prey.
Predation by sharks, foxes, Killer Whales (Orcinus
orca) and Steller Sea Lions (Eumetopias jubatus)
has been recorded (Bychkov 1967; Gentry and
Johnson 1980; Hanna 1922; Reeves et al. 1992;
National Marine Fisheries Service 1993). Three to
seven percent of fur seal neonates on St. George
Island were killed by sea lions in 1974 and 1975
(Gentry and Johnson 1980); no data on mortality
levels from Killer Whales or sharks are available.

Natural environmental fluctuations, such as the
periodic occurrence of El Nino, has negatively
affected the population breeding at San Miguel
Island, although conditions for the Pribilof Island

MIGRATORY ROUTES AND BREEDING AREAS ..
OF NORTHERN FUR SEALS

=
=>
Primary distribution ———

Migration route

~

June—October

q
XN
\

=~.
Commander Is.
i}

A =
Oy
/
su | }
ys ~/June—October
= =

y =z ¢ ne ow
4
Kuril Is. ?
4

\ “June—October

120 OOE 130 00 140 00 150 00 160 00 170 00 180 OOW

BAIRD AND HANSON: STATUS OF NORTHERN FUR SEAL

\UNK
ee

Ng ar
Bogoslof ea - June
I. a | K
~:*’
s

170 00

ALASKA

as 2s a ai ; of S Eig s a . aa AS

\;Pribilof Is.
\

me tA - =
y- ae May ~

¢ October 50 00

?

40 00
November— ‘
February

San Miguel |. 3
30 00

20 00

160 00 150 00 140 06 130 00 120 00 110 00

FiGurRE 2. Distribution of the Northern Fur Seal off the west coast of North America. Breeding islands are located in the
circles. Reproduced from Loughlin and Miller (1989) by permission.

population may be enhanced by increased sea sur-
face temperatures associated with these events (York
1991). The 1983 El Nifio occurred just prior to
implantation of embryos at San Miguel Island, and
resulted in an increase in pup mortality and a
decrease in pup weights. DeLong et al. (1993) sug-
gested that the later onset of the 1992 El Nifo affect-
ed this population less intensely. Competition with
fisheries has been suggested as a possible limiting
factor for this species, both in waters surrounding
breeding colonies, along migration routes, and dur-
ing the non-breeding season in the North Pacific.
The interactions between commercial harvesting of
prey species and fur seal movements, reproductive
rates or mortality is unclear, however (National
Marine Fisheries Service 1993).

Calambokidis and Peard (1982) examined levels
of chlorinated hydrocarbons from Northern Fur Seals
in Alaska, but found concentrations well below lev-
els thought to contribute to reproductive problems in
other populations of pinnipeds. Anas (1974) reported
levels of heavy metals in fur seals from Alaska and
Washington State, but no information is available on
potential impacts. An analysis of heavy metals by
Noda et al. (1995) revealed higher cadmium concen-
trations in northern fur seals than in other otartids.
Cadmium levels were higher, and mercury levels
lower, than those reported by Anas (1974); however,
heavy metal concentrations in Northern Fur Seals are
variable with age, location, and probably season,
making comparisons between studies difficult (Noda

et al. 1995). Entanglement in fishing gear is probably
a more significant problem. Northern Fur Seals were
the third most commonly caught species of marine
mammal recorded in an observer program of the
Japanese driftnet fishery for squid in 1989
(Anonymous 1990). Two animals were killed in an
experiment drift gillnet fishery for Neon Flying
Squid (Ommastrephes bartrami) in Canadian waters
in the mid-1980s (Jamieson and Heritage 1988).
Entanglement in marine debris is also a source of
mortality. Fowler (1987) suggested that mortality
due to entanglement in marine debris has contributed
significantly to the decline in the population on the
Pribilof Islands. Recent declines have also occurred
in the numbers of Steller Sea Lions (Loughlin et al.
1992) and Harbour Seals, Phoca vitulina (Pitcher
1990) in the central and eastern Gulf of Alaska.
While many possible causes have been identified
(reviewed above), the exact causes of these declines
continues to remain unclear.

Special Significance of the Species

This species is the only fur seal found in the tem-
perate waters of the north Pacific Ocean and is
endemic to this region. Alaskan natives on the
Pribilof Islands harvest approximately 2000 subadult
males annually for food.

Evaluation
The present world population of Northern Fur
Seals is substantially lower than historical levels,

268

and causes of the decline are unclear. Although the
market demand for furs is currently low, the lack
of any international regulatory body or agreement
on the management of the species means that
killing of this species at sea could be undertaken at
any time, and trade is not restricted or monitored
through any international agency. Rapid develop-
ment of fishing industry support services on the
Pribilof Islands has the potential to adversely
affect this population.

Acknowledgments

We thank the Canadian Wildlife Federation for
financial assistance for the preparation of this
report, Robert Campbell and COSEWIC for provid-
ing assistance and support, and several anonymous
reviewers for helpful comments on the manuscript.

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Bychkov, V. A. 1967. On killer whale attacks on fur seals
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Accepted 8 July 1996

Status of the Lacs des Loups Marins Harbour Seal,
Phoca vitulina mellonae, in Canada*

R. J. SMITH
Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1

Smith, R. J. 1997. Status of the Lacs des Loups Marins Harbour Seal, Phoca vitulina mellonae, in Canada. Canadian Field-
Naturalist 111(2): 270-276.

The Lacs des Loups Marins Harbour Seal (Phoca vitulina mellonae) is a subspecies that occurs in the area of Lacs des
Loups Marins (Lower Seal Lakes) (56°N, 73-74°W), 160 km east of Hudson Bay, on the Ungava peninsula of northern
Québec. With the possible exception of Lake Iliamna, Alaska, it is the only known Harbour Seal population that is resident
in freshwater year-round. Written references to the unique appearance and behaviour of this seal date back to 1818. The
subspecies was described primarily on the basis of its unique morphology and presumed long-time geographic isolation
from neighbouring oceanic Harbour Seals. Estimates of the population’s size are imprecise, and range from 100 to 600 ani-
mals. Little is known of the seals’ habitat requirements other than that they seem to feed exclusively in freshwater, and are
likely reliant on some specific environmental features such as under-ice air pockets to sustain them through the winter.
Pupping seems to take place substantially earlier (mid-April to mid-May) than in other Harbour Seal populations at similar
latitudes. The only known cause of human-induced mortality is occasional hunting by aboriginal peoples. Phoca vitulina
mellonae is potentially vulnerable because of its small population size, restricted range, and susceptibility to disturbance.

Les phoques communs des Lacs des Loups Marins (Phoca vitulina mellonae) sont une sous-espéce que |’on trouve dans la
région des Lacs des Loups Marins (Lower Seal Lakes) environ au 56-57°N, 73-74°0 a 160 km a l’est de la Baie d’ Hudson,
sur la péninsule d’Ungava, au nord du Québec. A |’exception, peut-étre, du Lac Iliamna en Alaska, c’est la seule popula-
tion de phoques communs connue qui reste dans les eaux douces toute l’année. On dispose de références écrites sur
l’apparence et le comportement uniques de ce phoque depuis 1818, et on a d’abord décrit cette sous-espéce a partir de sa
morphologie particuliére et selon l’hypothése qu’elle est depuis lontemps géographiquement isolée des phoques communs
océaniques avoisinants. Les estimations de leur population sont peu précises et donnent entre 100 et 600 individus. On en
connait peu sur les exigences des phoques quant a leur habitat a part le fait qu’ils semblent dépendre de certaines carac-
téristiques environnementales précises comme des poches d’air sous la glace pour qu’ils puissent passer V’hiver. La nais-
sance de petits semble subvenir sensiblement plus tot dans leur cas (mi-avril 4 mi-mai) que dans celui d’autres phoques a
des latitudes comparables. Les seules causes de mortalité connues sont quelques références, a des prises de chasse par des
Autochtones. Sa petite population, son aire restreint et sa vulnérabilité aux dérangements justifient qu’on accorde 4 cette
espeéce un statut de vulnérable.

Key Words: Harbour Seal, Phoca vitulina, Phocidae, pinniped, freshwater seals, phoque commun, Lacs des Loups Marins,
northern Québec, endangered species.

The Lacs des Loups Marins Harbour Seal, Phoca
vitulina mellonae (Doutt 1942) [Figure 1], is con-
fined to the area of Lacs des Loups Marins (Lower
Seal Lakes), approximately 160 km east of Hudson
Bay, on the Ungava peninsula of northern Québec
(Figure 2) [Doutt 1942; Anderson 1946; Scheffer
1958; Bigg 1981].

While there are numerous references to Harbour
Seals occurring in freshwater worldwide (Erlandson
1834; DeKay 1842; Allen 1880; Browne 1909;
Grenfell 1910; Prichard 1911; Strong 1930; Dunbar
1949; Fisher 1952; Wheeler 1953; Harper 1956,
1961; Beck et al. 1970; Paulbitski 1974; Roffe and
Mate 1984; Williamson 1988), Phoca vitulina mel-
lonae is the only known Harbour Seal population
resident in freshwater year-round (Atkinson 1818;
Clouston 1820; Hendry 1828; Finlayson 1830; Low
1898; Lewis 1904; Flaherty 1918; Twomey 1938;

Doutt 1942; Manning 1946; Doutt 1954; Graburn
1969; Power and Gregoire 1978; Smith and
Horonowitsch 1987; Consortium Gilles Shooner &
Associés et al. 1991). A freshwater population is
found in Lake Iliamna, Alaska, but it is the Pacific
subspecies Phoca vitulina richardsi, and it is not
certain whether it is landlocked (Everitt and
Braham 1980).

Written references to the unique appearance and
behaviour of Phoca vitulina mellonae date back to
Atkinson (1818). The subspecies was described pri-
marily on the basis of an unusually dark pelage and
an enlarged coronoid process on the mandible (Doutt
1942), with the presumption that the population had
been isolated for 3000 to 8000 years, trapped by the
Ungava peninsula’s isostatic rebound since the
retreat of the Laurentian ice sheet. Other authors dis-
puted this interpretation, however, arguing that the

*Reviewed and approved by COSEWIC 16 April 1996 —status assigned: Vulnerable.

270

1997

SMITH: STATUS OF THE LACS DES LOUPS MARINS HARBOUR SEAL

274

al ps3 f

~~ “7 !
in ane

FiGuRE |. One of the specimens of Lacs des Loups Marins Harbour Seal that were collected in March 1938, by
the Carnegie Museum of Natural History. The animal has just been shot by Daniel Petagumskum, one of
the expedition’s Cree guides (photograph by J. K. Doutt; reproduced with the permission of the
Carnegie Museum of Natural History, Pittsburgh, Pennsylvania).

supposed morphological anomalies of Phoca vitulina
mellonae are merely artifacts of a small sample size,
and that the seals are likely able to travel freely
between salt and freshwater (Mansfield and
McLaren 1958; Mansfield 1967; Smith and
Horonowitsch 1987; also see Honacki et al. 1982;
King 1983; Wiig 1989; Reeves et al. 1992). Other
work, the majority of it recent, strongly supports the
validity of a subspecific designation for Phoca vituli-
na mellonae (Davies 1958; Consortium Gilles
Shooner & Associés et al. 1991; Smith et al. 1994,
1996; Smith 1996).

Distribution

This is clearly a population with restricted dis-
tribution (Figure 2). There are historical references
to the presence of this seal in Lac Minto, at the
head of Riviere aux Feuilles (Flaherty 1918;
Manning 1947), Lac Beneta, situated in the basin
of Riviére aux Mélézes (Manning 1947), Petit Lac
des Loups Marins (Atkinson 1818; Clouston 1820;
Doutt 1942), and Lacs des Loups Marins (Hendry
1828; Finlayson 1830; Low 1898; Lewis 1904;
Doutt 1942; Doutt 1954; Power and Gregoire
1978; Berrouard 1984; Smith and Horonowitsch
1987). Several sightings have been made by
Hydro-Québec employees and contractors in the

Riviere aux Feuilles, Lac Melvin and Riviére
Delay (Consortium Gilles Shooner & Associés et
al. 1991). Inuit hunters, interviewed by Hydro-
Québec contractors, reported seeing or killing
freshwater seals in Lac Guillaume-Delisle, Riviére
Nastapoca, Riviere Boniface, Riviere Niagurnaq,
Riviere Kuunga, Riviere Longland, Lac Tasialuk,
and Lacs des Loups Marins (Archéotec Inc. 1990).
The Cree nation of Whapmagoostui considers the
range of Phoca vitulina mellonae to be Lacs des
Loups Marins, Petit Lac des Loups Marins, and
Lac Bourdel, with some reports of animals having
once been in Lac a 1’Eau Claire (Clearwater Lake)
(J. Petagumskum Sr., Whapmagoostui, Québec,
personal communication).

Hydro-Québec has recently compiled observa-
tions made of these freshwater seals between 1970
and 1990 (Consortium Gilles Shooner & Associés
et al. 1991). Though their survey efforts have been
concentrated largely in Lacs des Loups Marins,
Hydro-Québec’s data nevertheless indicate the
presence of seals in Riviere Nastapoca, Lacs des
Loups Marins, Petit Lac des Loups Marins, Lac
Bourdel, Lac a |’Eau Claire, and Petite Riviére de
la Baleine. In addition, evidence from recordings of
underwater vocalizations suggests the presence of
seals in Riviere aux Feuilles, Riviére aux Mélézes,

272

Riviére du Gué, Grande Riviére de la Baleine and
La Grande Riviére (Consortium Gilles Shooner &
Associés et al. 1991).

During the autumn of 1995, four seals were cap-
tured in Lacs des Loups Marins and affixed with
satellite-linked time-depth recorders (Wildlife
Computers, Redmond, Washington). All four tags
transmitted from early September to mid-
November, and during that time all four seals
remained within Lacs des Loups Marins or in the
immediate vicinity (R. J. Smith, unpublished data).

Protection

Because Canada possesses no specific marine
mammal or endangered species legislation, and
because it is unclear whether Phoca vitulina mel-
lonae, a marine mammal in freshwater, falls within
a provincial or federal jurisdiction, the population
currently has minimal legal protection. Freshwater
seals north of the 55th parallel are listed as a pro-
tected species under the James Bay and Northern
Québec Agreement (Québec 1976); however, this
protection does not have the force of law (J. Gunn,
Ministére de 1’Environnement et de la Faune,
Radisson, Québec, personal communication).
Phoca vitulina mellonae was recently listed by the
International Union for the Conservation of Nature
and Natural Resources (IUCN) as being “insuffi-
ciently known”, meaning that it is “suspected but
not definitely known to be endangered, vulnerable,
or rare due to a lack of reliable information”
(Reijnders et al. 1993). The government of Québec
has listed the population as “likely to be designated
as threatened or vulnerable” (Québec 1992a), and is
considering whether to give legal protection to a
portion of Phoca vitulina mellonae’s habitat
(Dubreuil 1983; Québec 1992b). This protection
should be a priority given that the proposed Grande
Baleine hydroelectric development could have an
adverse impact on a large portion of this popula-
tion’s range (Rosenthal and Beyea 1989; Rougerie
1990; Woodley et al. 1992; Smith et al. 1994).

Population Size And Trends

Estimates of the size of this small population are
imprecise. A maximum of 500 animals was the
“guess” of Doutt (1957), cited in Scheffer (1958).
Power and Gregoire (1978) estimated 200 and 600
animals by two different summations. The most
recent estimate by Consortium Gilles Shooner &
Associés et al. (1991) was approximately 100 ani-
mals, or 0.1 seals/km’, in Lacs des Loups Marins
and Lac Bourdel. Population trends over time obvi-
ously cannot be calculated.

Habitat
Little is known of the habitat and ecological
requirements of this subspecies. The few dead ani-

THE CANADIAN FIELD-NATURALIST

Vol. 111

mals that have been examined were found to have
salmonid (Salvelinus sp.) otoliths in their stomachs
(Consortium Gilles Shooner & Associés et al. 1991;
Smith et al. 1996). Comparisons of the stable-isotope
ratios and fatty acid profiles of the tissues of Phoca
vitulina mellonae and Harbour Seals collected from
oceanic locations indicate that, over a two year peri-
od, Phoca vitulina mellonae seemed to be feeding
exclusively in freshwater (Smith et al. 1996).

Recent investigations found no permanent
haulout sites on Lacs des Loups Marins and Lac
Bourdel (Consortium Gilles Shooner & Associés et
al. 1991). In winter, when the vast majority of the
lakes and rivers are covered in ice, the seals may
rely on several physical features for their sources of
air: areas that remain ice-free because of strong cur-
rents, fissures in the ice, and air pockets created by
the shoreline’s complicated geometry or by the
undulations in the bottom of the sheet ice on the
lake’s surface (Smith and Horonowitsch 1987;
Consortium Gilles Shooner & Associés et al. 1991;
Dean Consulting & Research Associates Inc. 1991).

None of the habitat of this population is protect-
ed. It is entirely on Crown land that could be
adversely affected by Hydro-Québec’s construction
of the proposed Grande Baleine hydroelectric pro-
ject (Woodley et al. 1992) which, though indefinite-
ly postponed by the current provincial government,
has not been cancelled altogether. One of the results
of the Grande Baleine environmental assessment
process has been that Hydro-Québec is now
required to evaluate properly the likely impacts of
the project on the population, prior to construction
(Review Bodies 1994). Some of these potential
impacts include the disappearance of ice-free areas
and under-ice shoreline shelters, upon which the
seals may rely in the winter, in water courses with
altered flows arising from hydroelectric develop-
ment. The Grande Baleine project may also affect
the distribution and abundance of the seals’ prey,
and contaminate the animals with methyl mercury
released from the flooded, decomposing vegetation
(Woodley et al. 1992). The negative effects of this
habitat destruction could lead to a decline in the seal
population and an impoverishment of its genetic
diversity (Alfonso and McAllister 1994).

General Biology

Reproduction probably occurs between mid-
April and mid-May in the Lacs des Loups Marins
area; substantially earlier than other Harbour Seal
populations at a similar latitude (Doutt 1942;
Archéotec inc. 1990; Consortium Gilles Shooner &
Associés et al. 1991; Temte et al. 1991; Smith et al.
1994). Since the lakes are still iced over at the time
of pupping, and no births have been observed on
the ice, several authors have postulated that pup-
ping takes place in under-ice shelters (Consortium

1997

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SMITH: STATUS OF THE LACS DES LouPS MARINS HARBOUR SEAL

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273

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R. AUX MELEZES R, DU GUE

L. BOURREL
.L. DES LOUPS

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FIGURE 2. Known range of Phoca vitulina mellonae in relation to Hydro-Québec’s proposed Grande Baleine

hydroelectric project.

Gilles Shooner & Associés et al. 1991), like those
of Ringed Seals (Phoca hispida) (Smith and
Stirling 1975).

The only known cause of human-induced mortali-
ty is occasional hunting of the seals by aboriginal
peoples (Clouston 1820; Low 1898; Flaherty 1918;
Doutt 1942; Doutt 1954; Consortium Gilles Shooner
& Associés et al. 1991; J. Petagumskum Sr., person-
al communication).

Seasonal movements of the population are poorly
known, though the sporadic observations of Gilles
Shooner & Associés et al. (1991) hint at seals spend-
ing the winter months in larger bodies of water like
Lacs des Loups Marins, Lac Bourdel, and Petit Lac
des Loups Marins, with some dispersal into outlying,
smaller bodies of water upon the melting of the ice.
These investigators report finding many worn trails
between bodies of water frequented by the seals,
some as long as 0.15 km, and on inclines as steep as
25°. There is no evidence that animals move between
the area of Lacs des Loups Marins and Hudson or
Ungava Bays. However, though there are a number
of impassable waterfalls on the Riviére Nastapoca,
some authors believe that if the seals could move
into the more placid rivers that flow north into
Ungava Bay, this would be a feasible avenue of
exchange between the fresh and saltwater popula-
tions (Mansfield 1967; SOGEAM 1985; Smith and
Horonowitsch 1987).

Preliminary evidence from DNA sequencing of
region I of the mitochondrial D-loop indicate that
Phoca vitulina mellonae has haplotypes that are
unique when compared to Harbour Seals in the
eastern Canadian arctic and Northwest Atlantic
(Smith 1996).

Animals hauled out in the spring months are usu-
ally in small groups, whereas at the end of the sum-
mer, they are usually hauled out singly or in pairs.
It has been suggested that this behaviour is related
to the moulting process (Consortium Gilles
Shooner & Associés et al. 1991).

Limiting Factors

The tendency of Harbour Seals to be distributed
in small local populations makes them vulnerable
to disturbance (Maine Seal 1994). There are a
number of examples of local Harbour Seal popula-
tions being extirpated, or their numbers drastically
reduced, by human activity. For example, a small
population that seemed to frequent Lake Ontario
was eliminated by the early 1800s (DeKay 1842;
Allen 1880); the population in Greenland is practi-
cally extirpated (Teilmann and Dietz 1993; R.
Dietz, personal communication), an important rea-
son being the intensity with which it has been
hunted and entangled in fishing gear; the popula-
tion in Hokkaido, Japan, is very small, with
removals from incidental catches in fishing gear

274

exceeding recruitment (Reijnders et al. 1993).
Given such evidence, the Lacs des Loups Marins
seal population is likely sensitive even to limited
disturbance by humans.

Special Significance of the Subspecies

This population of Harbour Seals is unique, in
that it is the object of reverence by the aboriginal
peoples of northern Québec (Archéotec Inc. 1990;
Richardson 1991; Posluns 1993; M. George,
Whapmagoostui, Québec, personal communica-
tion); it is the object of a wealth of historical refer-
ences, and seems to be unusual in a number of
ways, including aspects of its biology (Consortium
Gilles Shooner & Associés et al. 1991; Smith et al.
1994, 1996; Smith 1996). The population has also
acquired something of a public profile over the last
few years (e.g.: Dubreuil 1987).

Evaluation

It is essential for the future viability of this sub-
species that the potential impacts on the population
from the Grande Baleine hydroelectric project, and
any other future development in Québec’s north, be
eliminated or seriously mitigated. Because of the
inaccessibility of the population, there is no known
trade in the subspecies, legal or otherwise.

Acknowlegments

I thank D. M. Lavigne for his support. The com-
ments and criticisms of R. R. Campbell and two
other members of COSEWIC’s Fish and Marine
Mammal Subcommittee were also helpful. G.
Luste provided some of the more obscure refer-
ences. Funding for the production of this
manuscript was provided by the Natural Sciences
and Engineering Research Council of Canada and
the International Marine Mammal Association Inc.,
Guelph, Ontario.

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Jenkins Ltd., London. 151 pages.

Wheeler, E. P. 1953. Notes on pinnipedia. Journal of
Mammalogy 34: 253-255.

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Vol. 111

Wiig, O. 1989. A description of common seals, Phoca vit-
ulina L. 1758, from Svalbard. Marine Mammal Science
5: 149-158.

Williamson, G. R. 1988. Seals in Loch Ness. Scientific
Reports of the Whales Research Institute 39:
151-157.

Woodley, T. H., R. J. Smith, and D. M. Lavigne. 1992.
Potential impacts of hydroelectric development on
marine mammals of northern Quebec. IMMA Technical
Report Number 92-02. International Marine Mammal
Association Inc., Guelph, Ontario. 9 pages.

Accepted 8 July 1996

Updated Status of the Sea Otter, Enhydra lutris, in Canada*

JANE C. WATSON!, GRAEME M. ELLIS”, THOMAS G. SMITH?, and JOHN K. B. Forp?*

‘Malaspina University-College, 900 Fifth Street, Naniamo, British Columbia V9R 5S5
Pacific Biological Station, Department of Fisheries and Oceans, Nanaimo, British Columbia V9R 5K6
3Vancouver Aquarium, P.O. Box 3232, Vancouver, British Columbia V6B 3X8

Watson, Jane C., Graeme M. Ellis, Thomas G. Smith, and John K. B. Ford. 1997. Updated status of the Sea Otter,
Enhydra lutris, in Canada. Canadian Field-Naturalist 111(2): 277-286.

Sea Otters (Enhydra lutris) ranged historically across the north Pacific from the central Pacific coast of Baja California, to
northern Japan. An intensive fur hunt in the 18th and 19th centuries extirpated the Sea Otter from much of its range. The
species was protected in 1911, and now occupies approximately half of its historic range. Presently, the worldwide popula-
tion of Sea Otters is thought to be about 150 000 animals. The British Columbia Sea Otter population, originating from 89
animals reintroduced to Vancouver Island from 1969 to 1972, has increased at a finite rate of 18.6% per year, a rate similar
to reintroduced populations in Washington and southeast Alaska. In 1995, a minimum of 1522 Sea Otters were found in
British Columbia. Most of these animals occurred off Vancouver Island, but at least 135 Sea Otters were found near Goose
Island, 125 km north of Vancouver Island. The origin of this latter group is uncertain. Sea Otters are presently classed as
threatened in British Columbia and are protected under Federal and Provincial regulations.

A une certaine époque, les loutres de mer, Enhydra lutris, étaient repandues partout dans le Pacifique nord, de la céte
Pacifique centrale de Baja, en Californie, jusqu’au nord du Japon. Aux 18° et 19° siécles, une chasse intensive a la fourrure
a fait disparaitre la loutre d’une grande partie de son aire de répartition. On commenga a protéger l’espéce en 1911 et elle
est maintenant revenue dans prés de la moitié de son aire historique. On croit que la population mondiale de la loutre
s’éléve présentement 4 quelques 150 000 individus. La population de loutres de mer de la Colombie-Britannique, com-
mencée avec 89 animaux réintroduits a la fle de Vancouver de 1969 a 1972, a augmentée a un taux de croissance fixe de
18,6 % par année, un taux semblable a celui des populations réintroduites dans |’état de Washington et dans le sud-est de
l’ Alaska. En 1995, un minimum de 1522 loutres de mer ont été relevées en Colombie-Britannique. La plupart de ces ani-
maux ont été vus au large de l’ile de Vancouver, mais on en a trouvé au moins 135 prés de Goose Island a 125 km au nord
de l’ile de Vancouver. On n’est pas certain de l’origine de ce dernier groupe. Les loutres de mer ont présentement le statut
d’espéce menacé en Colombie-Britannique et elles sont protégées en vertu de reglements fédéraux et provinciaux.

Key Words: Sea Otter, Loutre de mer, Enhydra lutris, otters, keystone predator, mustelid, fur trade.

The Sea Otter, Enhydra lutris (Linnaeus, 1758), is Distribution

one of the largest members of the family Mustelidae,
but the smallest of marine mammals (Estes 1980).
An adult male Sea Otter (Figure 1) may weigh up to
45 kg and attain a length of about 1.4 m; females are
smaller (Kenyon 1969; Estes 1980). In contrast to
other marine mammals, which rely upon blubber for
insulation, Sea Otters depend upon their dense fur
for protection against chilly ocean water (Kenyon
1969). The coat is light to dark brown in colour,
often becoming grizzled with age, especially around
the head (Kenyon 1969; Garshelis 1984). Sea Otters
have a high metabolic rate consuming up to one
quarter of their body weight in food each day
(Riedman and Estes 1990). Sea Otters feed on benth-
ic invertebrates, often limiting the abundance of their
prey, which has wide-ranging ecological conse-
quences on community structure (Estes et al. 1989).
Previous COSEWIC (Committee on the Status of
Endangered Species of Wildlife in Canada) reports
on the Sea Otter population in Canada were pub-
lished by Monroe (1985) and MacAskie (1987).

Sea Otters ranged historically across the Pacific
rim, from northern Japan to central Baja California
(Figure 2) [Kenyon 1969; Riedman and Estes 1990].
An intensive fur hunt for Sea Otters, commencing in
the 1740s resulted in their near extinction (Kenyon
1969). Sea Otters were protected in 1911, when less
than 2000, animals in 13 relic groups, remained
(Figure 2) [Kenyon 1969].

Several of these remnant groups declined to
extinction, likely because of their small size. One
such group located off the Queen Charlotte Islands,
disappeared, and the last Sea Otter known to inhabit
British Columbian (Canadian) waters was shot in
1929 off Vancouver Island (Cowan and Guiguet
1960; Kenyon 1969; Monroe 1985).

The Sea Otter currently occupies approximately
half of its historical range (Figure 2), extending
from the northern Gulf of Alaska, westward through
the Aleutian archipelago to the Kamchatka
Peninsula and the Kuril archipelago (Estes 1990a).
Large areas to the south of the Gulf of Alaska

*Reviewed and approved by COSEWIC 16 April 1996, status assigned: Threatened (downlisted from Endangered).

277

278

remain uninhabited, except for recently reintroduced
populations in southeast Alaska, British Columbia,
Washington, and a remnant population in
California.

During the 1960s and early 1970s, Sea Otters
captured at Amchitka Island and Prince William
Sound were reintroduced to unoccupied habitat in
Alaska, British Columbia, Washington, and Oregon
(Riedman and Estes 1990; Estes 1990b). Eighty-
nine Sea Otters were reintroduced to Checleset Bay
off northwest Vancouver Island; 29 in 1969, 14 in
1970, and 46 in 1972. By 1972, Sea Otters had
been sighted from the Queen Charlotte Islands to
Barkley Sound, suggesting that some of the
translocated animals had dispersed (Bigg and
MacAskie 1978). In 1977, aerial surveys found Sea
Otters near the original transplant site in Checleset
Bay, and off Bajo Reef, Nootka Island, 75 km to
the southeast (Figure 3) [Bigg and MacAskie
1978]. Until 1991, the Vancouver Island Sea Otter
population was composed of two groups, one in
Checleset Bay and a second off Nootka Island
(MacAskie 1987), but by 1992 the population
ranged continuously from Estevan Point northwest
to Quatsino Sound (Figure 3) [Watson 1993].
Between 1977 and 1991 the range of the Vancouver
Island Sea Otter population expanded at a rate of
15.3% per year (Watson 1993).

In 1989 a group of Sea Otters was reported in the
Goose Island Group, 125 km northwest of
Vancouver Island (Figure 3). Single Sea Otters
were reported in the Goose Islands in 1976 (Bigg
and MacAskie 1978) and subsequently, but mothers
and pups were first sighted in 1989 (D. Powers,
Warden, Hakai Pass Recreation Area, c/o British
Columbia Parks, Caribou District, 540 Borland
Street, Williams Lake, B.C. V2G 1R8, personal
communication). The origin of these Sea Otters is
unknown.

Sea Otters were reintroduced to various sites in
southeast Alaska from 1965 -1969 (Jameson et al.
1982; Riedman and Estes 1990) and may account
for confirmed sightings off the north end of the
Queen Charlotte Islands. In 1969 and 1970, Sea
Otters were translocated to Washington State
(Jameson et al. 1986). Animals from this popula-
tion may account for some of the reports of Sea
Otters off southern Vancouver Island and in the
Strait of Georgia. The California Sea Otter popula-
tion, a relict group, extends from about Ano Nuevo
Point to Point Sal (Riedman and Estes 1990b; J.
Estes, National Biological Survey, University of
California, Santa Cruz, California 95064, personal
communication).

Protection
Sea Otters were protected in 1911 under the
International Fur Seal Treaty signed by the United

THE CANADIAN FIELD-NATURALIST

Vol. 111

States, Russia, Japan and Great Britain (for
Canada). In Canada, Sea Otters are protected under
the Federal Fisheries Act and the British Columbia
Wildlife Act and Regulations. Sea Otters were des-
ignated as an “endangered” species by COSEWIC
and under the British Columbia Wildlife Act in
1980. MacAskie (1987) retained this designation.
In the United States, Sea Otters are protected by the
Marine Mammal Protection Act of 1972, which
prohibits their take or harassment except for public
display or scientific research. Aboriginal peoples of
Alaska are permitted to take Sea Otters for tradi-
tional use. Sea Otters in California, which are con-
sidered a separate subspecies (Wilson et al. 1991),
are designated as “threatened” under the U.S.
Endangered Species Act of 1972 (Estes 1990b).

Population Size and Trends

Before commercial exploitation, the worldwide
population of Sea Otters was between 150 000 and
300 000 animals, although these estimates are high-
ly speculative, and some authors suggest this num-
ber may have been much larger (Kenyon 1969;
Johnson 1982). Since their protection the species
has recovered in many areas. Sea Otters from the
northern Gulf of Alaska, through the Aleutian
archipelago to the Kuril archipelago are probably at
or near equilibrium density throughout much of
their range (Estes 1990b). Recently reintroduced or
remnant populations in southeast Alaska, British
Columbia, Washington and California are increas-
ing in size.

The British Columbia Sea Otter population, now
well established with at least 1522 animals, origi-
nated from the 89 animals reintroduced from 1969
to 1972. Population surveys, conducted from
fixed-wing aircraft, helicopters and small vessels,
indicate the British Columbia Sea Otter population
has increased at a finite rate of 18.6% per year
from 70 animals in 1977 to 1522 animals in 1995
(Table 1, Figure 4, Watson 1993). This rate, which
may be near r,,. (Caughley 1977), is similar to that
of reintroduced populations in southeast Alaska
and Washington State, and is almost identical to
the instantaneous rate of 17.7% per year calculated
by Estes (1990a) using 1977, 1984, and 1987 data.
Estes (1990a) suggests that if this growth curve is
extrapolated back in time, the translocated popula-
tion had declined to 28 individuals by 1973. At
present, portions of the British Columbia Sea Otter
population may be at equilibrium density (Watson
1993),

Habitat

Sea Otters inhabit shallow coastal waters, seldom
ranging more than | to 2 km from shore, usually
foraging to depths of 30 m or less (Riedman and
Estes 1990). Sea Otters forage in areas with rocky

1997

Bae

FiGuRE 1. The Sea Otter, Enhydra lutris.

or soft substrate (Riedman and Estes 1990). Sea
Otters, especially mothers with pups, appear to pre-
fer areas with kelp canopies. Kelp, however, is not
an essential habitat requirement, because large pop-
ulations of Sea Otters occur in areas which lack
extensive kelp beds (Riedman and Estes 1990).
Habitat use varies with weather and sea state. Off
Vancouver Island, for example, Sea Otters move
offshore during extended periods of calm, and con-
gregate in sheltered inshore areas during storms
(Morris et al. 1981; Watson 1993).

In British Columbia, Checleset Bay Ecological
Reserve 109 was established in 1981 to protect Sea
Otter habitat. Sea Otters off Goose Island are locat-
ed within the boundaries of Hakai Pass Recreational
Area. However, these areas are under provincial
jurisdiction and are only partially exempt from com-
mercial shellfish harvest, which is under federal
jurisdiction.

General Biology
Reproductive Capability

Most female Sea Otters reach sexual maturity at
three years or older (Bodkin et al. 1993) and males
between 5 and 6 years of age (Riedman and Estes
1990). Sea Otters bear a single pup at approximately
one-year intervals (Bodkin et al. 1993). Pups are
usually born in the water and twinning is rare
(Kenyon 1969).

The number of large, dependent pups present off
Vancouver Island in July suggests a peak parturition
period from approximately April to July, as also
observed in Alaska (Payne and Jameson 1984;
Garshelis et al. 1984; Riedman and Estes 1990;

WATSON, ELLIS, SMITH, AND FORD: STATUS OF THE SEA OTTER

phe

Photo courtesy Vancouver Aquarium.

Watson 1993; Bodkin et al. 1993). However, pup-
ping can occur year round. The period of pup
dependency varies between individuals, but rarely
exceeds one year (Riedman and Estes 1990; Bodkin
et al. 1993).

In British Columbia, annual estimates of the popu-
lation birth rate (pups/adult) range from 0.12 to 0.33
(Watson 1993). Similar rates are reported for Alaska
(0.30 pups/adult) and California (0.20 pups/adult),
although the pupping period is more extended in
California, making it difficult to compare population
birth rates (J. Estes, personal communication). It
should be noted that pup abundance is easily over or
under-estimated if areas of high and low pup produc-
tion are not equally surveyed (Simon-Jackson 1986;
Jameson et al. 1986; Estes 1990a).

Species (Individual) Movement

Sea Otters are not migratory. Patterns of move-
ment and social organization in British Columbia
appear similar to those observed in Alaska and
California. Sea Otters segregate by sex and age,
although patterns vary somewhat with season. In
general, males and females rest in different loca-
tions. Adult males occupy territories that overlap
female areas especially during the autumn breeding
season. In California and Alaska large groups of
males are found at the edges of the sea-otter range.
These groups are composed of subadult and adult
males, with breeding males joining them outside of
breeding season. Males generally expand into new
areas first whereas females use areas which have
been occupied by Sea Otters for longer periods, and
expand into areas as they are vacated by male

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THE CANADIAN FIELD-NATURALIST

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WATSON, ELLIS, SMITH, AND FORD: STATUS OF THE SEA OTTER

281

% 1 CANADA

.
, .
f ‘e

PACIFIC OCEAN

Onginal Distnbution
Present Distnbution
Remnant Colonies

Surviving Translocation Populations

FiGuRE 2. Historic and present distribution of Sea Otters, with remnant colonies and translocated populations marked.

groups (Loughlin 1980; Ribic 1982a, 1982b;
Garshelis et al. 1984; Wendell et al. 1986; Pitcher
1989; reviewed in Riedman and Estes 1990).

Both male and female Sea Otters establish home
ranges (Jameson 1989), the size of which depends,
to a large degree, upon season. The home ranges of

Queen
Charlotte Islands

British Columbia sea otter
distribution

males are substantially larger than those of females,
which reflects seasonal movement of breeding
males. Females are sedentary by comparison. Daily
movement patterns vary with the territorial status of
the male and whether or not a female has a pup
(Jameson 1989, review in Riedman and Estes 1990).

» Wancouver *
Island es

FiGurE 3. The distribution of the British Columbia Sea Otter population.

282

Behaviour / Adaptability

Sea Otters in British Columbia are wary of
humans and easily disturbed by boat traffic. Females
with pups seem most sensitive to disturbance. As
these areas become more utilized by tourists, Sea
Otters may become less wary. Sea Otters adjacent to
inhabited areas habituate to the presence of humans
(e.g., Woolfenden 1985).

There is substantial variation in the diet of individ-
ual Sea Otters. Sea Otters are highly adaptable in
their diet, often exploiting seasonally abundant food
resources such as spawning squid (Riedman and
Estes 1990) or pelagic red crab during El Nino
events (in Riedman and Estes 1990). In one notable
example, a male otter learned to bite into discarded
aluminum cans to obtain octopus which hid inside
the cans (McCleneghan and Ames 1976). In
California, Sea Otter dietary preferences appear to be
maternally transmitted (Lyons 1989).

Sea Otters feed principally upon shellfish.
However, in parts of Alaska, where Sea Otters have
been long established, fish are also an important
food (Estes 1977). The high proportion of fish in the
diet of Sea Otters in these areas may be associated
with an increase in fish abundance and decline in
shellfish abundance (Estes et al. 1978). Estes
(1990a) speculated that inclusion of fish in the diet
of Sea Otters may increase the equilibrium density of
otter populations. Foraging on fish may be a feeding
strategy lost to many populations when Sea Otters
were reduced to a few remnant colonies at the end of
the fur hunting era (Estes et al. 1989). Sea Otters
have not been observed feeding on fish in British
Columbia.

Limiting Factors

Fur hunting in the 18th and 19th centuries was
responsible for the extirpation of Sea Otters through-
out most of their range. Today, the worldwide otter
population continues to rise, with further increases
expected as long as unoccupied habitat is available
(Estes 1990b).

Food limitation is probably the major cause of
death in Sea Otter populations at or near equilibrium
density (Riedman and Estes 1990). Other than
humans, predators include Bald Eagles, Haliaeetus
leucocephalus (Sherrod et al. 1975); Killer Whales,
Orcinus orca (in Riedman and Estes 1990); and
sharks (Ames and Morejohn 1980). Sources of Sea
Otter mortality have not been studied in British
Columbia. However, pup carcasses found at eagles’
nests, suggested that Bald Eagle predation on pups
may represent a substantial source of pup mortality.
Killer Whales, although probably an insignificant
source of mortality, have been seen pursuing and
consuming Sea Otters in Kyuquot Sound, British
Columbia (Watson 1993) and at Amchitka Island,
Alaska (J. Estes, personal communication). One

THE CANADIAN FIELD-NATURALIST

Vol. 111

otter carcass recovered near Kyuquot, British
Columbia, had injuries which could have been
caused by a boat propeller (G. Ellis, unpublished).

Human-related threats to Sea Otters are localized
and include environmental contamination, conflict
with commercial fisheries, incidental take, and the
possibility of the resumption of hunting by native
peoples.

Environmental Contamination

Perhaps the greatest threat to Sea Otters is that of
oil spills (Geraci and St. Aubin 1980; Geraci and
Williams 1990; Ralls and Siniff 1990). Sea Otters
are vulnerable to oil in several ways. Oil destroys
the water-repellent property of the fur, allowing
cold water to contact the skin, which may result in
hypothermia. Inhalation of volatile petroleum
vapors injures lungs and other organs. Lastly when
oil is ingested through grooming or foraging it
injures internal organs (Geraci and Williams 1990).
Furthermore, Sea Otters spend much of their time
in kelp beds which retain oil, and congregate in
groups, which means large numbers of Sea Otters
can be exposed to oil simultaneously (Ralls and
Siniff 1990).

Although methods for cleaning and rehabilitating
oiled otters have been developed (Costa and
Kooyman 1984; Davis et al. 1988; Geraci and
Williams 1990; Degange et al. 1995; Williams and
Davis 1995), their effectiveness, at a population
level, has been questioned (Estes 1991). On 24
March, 1989 the oil tanker Exxon Valdez ran
aground in Prince William Sound, spilling 10 mil-
lion gallons (37.8 million 1) of crude oil. At least
1013 Sea Otters were killed (Bayha and Kormendy
1990; Estes 1991; Ballachey et al. 1994); additional
carcasses undoubtedly sunk and were not recovered.
On 22 December 1988, 231 000 gallons (875 000 1)
of oil from the barge Nestucca spilled and spread
northwards from Grays Harbour, Washington. This
relatively small spill killed at least one Sea Otter in
Checleset Bay, British Columbia (Waldichuck
1989; Watson 1990, 1995). The continued trans-
portation of oil along the coast of British Columbia
poses a significant threat to British Columbia’s Sea
Otter population.

The effect of enviromnental toxins on Sea Otters
has not been well documented, although varying
levels of chlorinated-hydrocarbons and heavy met-
als have been found in sea-otter tissue (Riedman
and Estes 1990). The high rate of population growth
observed throughout much of the Sea Otters’ range,
suggests that environmental toxins are not at present
a significant threat to Sea Otters (Riedman and
Estes 1990), although recent evidence suggests that
locally high levels of environmental toxins may
affect pre-weaning mortality in Sea Otters (J. Estes,
personal communication; Bacon 1994).

1997

10000

1000

100

Population size

10

1975 1980 1990 1995 2000

1985

Year

FiGurE 4. Growth of the British Columbia Sea Otter popu-
lation from 1977-1995. The population has increased
at a rate of 18.6% per year (r°=0.969) since 1977.
Estimates of the Sea Otter population off of Goose
Island are not included, because the size of this group is
poorly known.

Fishery Conflicts

Sea Otters are capable of limiting the abundance
of many commercial species of shellfish (Johnson
1982; Estes and VanBlaricom 1985; Pitcher 1989).
This has resulted in considerable conflict between
commercial shellfish harvesters and Sea Otters
(Estes and VanBlaricom 1985). Sea Otters have had
a significant effect upon the Pismo clam fishery in
central California (Wendell et al. 1986), clam densi-
ty in Alaska (Kvitek et al. 1992; Kvitek and Oliver
1992) and sea urchin abundance in parts of British
Columbia (Morris et al. 1979, 1981; Breen et al.
1982; Watson 1993). Conflicts over Sea Otters and
shellfish stocks are not yet a problem in British
Columbia, but can be anticipated as the geographic
range of Sea Otters increases (Watson and Smith
1996).

Incidental Take of Sea Otters

Gill-net fisheries in California have probably lim-
ited or even reduced the southern Sea Otter popula-
tion (Wendell et al. 1985). Incidental take of Sea
Otters in net-fisheries in southeast Alaska and Prince
William Sound have been reported, but the impact
on the Sea Otter population is unknown (Matkin and
Fay 1980; Simon-Jackson 1986). Gill-nets are used

WATSON, ELLIS, SMITH, AND FORD: STATUS OF THE SEA OTTER

283

in both salmon and herring fisheries in British
Columbia, but incidental take of Sea Otters has not
been reported. The expansion of Sea Otters into
areas where gill-net fisheries, particularly for
salmon, occur could have local effects on Sea Otter
abundance.

Special Significance of the Species

Once on the verge of extinction, Sea Otters have
made a remarkable recovery, largely because of
protective legislation, an abundance of suitable
unoccupied habitat and successful reintroductions
(Mason and Macdonald 1990). The worldwide pop-
ulation of Sea Otters is thought to be about 150 000
animals, most of which are found from the Gulf of
Alaska and westward across the Aleutian archipela-
go (Estes 1980; Riedman and Estes 1990).

The "teddy-bear-like" appearance of Sea Otters,
their near brush with extinction, their important
role in nearshore community structure and their
vulnerability to oil spills has endeared them to the
public. Sea Otters do well in captivity and are
popular at zoos and oceanaria. Sea Otters use rocks
and other hard objects as tools to crack the
exoskeletons of their prey. With the exception of
primates, Sea Otters are the only mammal known
to use tools.

The Sea Otter, Enhydra lutris, is the sole member
of the genus Enhydra. Three subspecies are recog-
nized; Enhydra lutris lutris which is found from the
Kuril Islands to the Commander Islands; Enhydra
lutris kenyoni (Wilson et al. 1991) which historical-
ly occurred from the Aleutian Islands to Oregon,
and Enhydra lutris nereis or the southern Sea Otter,
which is presently found only in California. The his-
toric range of the southern Sea Otter is disputed
(Riedman and Estes 1990).

In the United States, the Marine Mammal
Protection Act allows aboriginal people in Alaska to
harvest Sea Otters for traditional use (Samuel and
Foin 1983; Simon-Jackson 1986). The number of
Sea Otters harvested in this fishery has increased
dramatically in recent years, from 55 animals in
1988 to 1248 in 1993 (J. Bodkin, National Biological
Survey, 1011 East Tudor Road, Anchorage, Alaska
99503, personal communication). The impact of this
harvest on local Sea Otter populations is unknown
(Estes 1990b).

Sea Otters exert a significant effect upon the struc-
ture of rocky nearshore communities (Estes and
Palrnisano 1974; Estes et al. 1978, 1989). By feeding
on herbivorous invertebrates, particularly sea
urchins, Sea Otters reduce grazing pressure and
increase the growth of fleshy algae, particularly
brown algae such as kelp (Estes et al. 1978; Breen et
al. 1982; Watson 1993). Consequently, rocky
nearshore communities in areas occupied by Sea
Otters are often characterized by an abundance of

284

fleshy algae, whereas areas without Sea Otters are
dominated by sea urchins and fleshy algae are rare
(Estes and Duggins 1995).

Evaluation

Although aboriginal hunting may have locally
extirpated Sea Otters prior to the arrival of
Europeans (Simenstad et al. 1978), Sea Otter popu-
lations were driven to the brink of extinction by an
intensive fur hunt commencing in the 1740s. Sea
Otters were protected in 1911, and have subse-
quently increased to historic levels throughout
much of their range. In British Columbia, the rein-
troduced Sea Otter population is increasing at near
maximum rate (Estes 1990a; Watson 1993).

Despite increases in the size and geographic
extent of Sea Otters in British Columbia, the popu-
lation is still relatively small in size and restricted
in distribution, making it vulnerable to environmen-
tal catastrophes such as oil spills. The species is no
longer in danger of imminent extirpation from
Canadian waters, but is still vulnerable and should
remain in a cateagory of risk which will ensure
continued protection and management. A downlist-
ing to "threatened" would retain their status on the
COSEWIC list as a species which could become
endangered if the factors limiting its distribution
and numbers are not controlled.

Acknowledgments

Financial support to produce this report was pro-
vided by the Canadian Wildlife Service. Field
research was supported by the Canadian
Department of Fisheries and Oceans, British
Columbia Ministry of Environment Lands and
Parks, United States Fish and Wildlife Service,
Vancouver Aquarium, University of California
Santa Cruz, Western Canadian Universities Marine
Biological Station, Archipelago Marine Research,
West Coast Whale Research Foundation, and
Friends of the Ecological Reserves. The British
Columbia Ministry of Environment Land and Parks
and the Department of Fisheries and Oceans made
unpublished data available. The manuscript was
improved by comments from James Estes, Ian
MacAskie, Ric Simmons and Andrew Trites.

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Accepted 8 July 1996

Status of the Northern Bottlenose Whale, Hyperoodon ampullatus,
in the Gully, Nova Scotia*

HAL WHITEHEAD, ANNICK FAUCHER, SHANNON GOWANS, and STEPHEN MCCARREY!

Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1
‘Department of Zoology, University of Tasmania, P.O. Box 252C, Hobart 7001, Australia

Whitehead, Hal, Annick Faucher, Shannon Gowans, and Stephen McCarrey. 1997. Status of the Northern Bottlenose
Whale, Hyperoodon ampullatus, in the Gully, Nova Scotia. Canadian Field-Naturalist 111(2): 287-292.

A population of approximately 230 Northern Bottlenose Whales, Hyperoodon ampullatus, uses the Gully, a prominent sub-
marine canyon on the edge of the Scotian Shelf. These animals use the Gully throughout the year. Approximately 57% of
the population reside in a 20 km X 8 km core area at the entrance of the canyon at any time. The Gully animals seem to be
largely or totally distinct from the population seen off northern Labrador: they are smaller and appear to breed at a different
time of year. Threats to the population include commercial shipping, fishing and oil and gas developments. One oil and gas
discovery of commercial interest, the Primrose Field, lies about 5 km from the core area of this population. The population
is vulnerable because of its small size, location at the extreme southern limit of the species’ range, and year-round depen-
dence on a small and unique sea area. It is threatened by plans for the development of the oil and gas fields close to the
Gully.

Une population d’ environ 230 baleines a bec communes, Hyperoodon ampullatus, se trouve dans le Gully, un canyon sous-
marin important situé sur le bord de la Plate-forme Néo-Ecossaise. Les animaux fréquentent le Gully a l’année longue.
Environ 57% de la population réside en tout temps dans une aire de 20 km X 8 km a l’entrée du canyon. Les animaux du
Gully sont en grande partie sinon totalement distincts de ceux des populations arctiques de cette espéce, étant plus petits et
semblant se reproduire a une période de |’année différente. Les facteurs menagant la population comprennent la navigation
commerciale, la péche et la présence de ressources pétroliéres et gaziéres potentiellement exploitables. L’une des décou-
vertes de pétrole et de gaz d’intérét commercial, le gisement Primrose, se situe 4 5 km de I’aire principale de cette popula-
tion. Cette derniére est vulnérable de par son petit nombre d’individus, sa situation a la limite extréme sud de |’étendue de
distribution de l’espéce et sa dépendance 4 |’année longue envers une petite zone marine unique. La population est men-
acée par des projets de développement de gisements de pétrole et de gaz a proximité du Gully.

Key Words: Northern Bottlenose Whale, baleine a bec commune, Hyperoodon ampullatus, Cetacea, Odontoceti, the Gully,
submarine canyon, Nova Scotia.

In this report we evaluate the status of the popula-
tion of Northern Bottlenose Whales, Hyperoodon
ampullatus (Forster 1770) whose members are found
in the Gully, a submarine canyon on the edge of the
Scotian Shelf. The Northern Bottlenose Whale
(Figure 1) is a 6- to 9-m member of the beaked
whale family (Ziphiidae) resident only in the north-
ern regions of the North Atlantic. Its closest relative
is the Southern Bottlenose Whale (Hyperoodon plan-
ifrons) of the Southern Oceans. Bottlenose whales
seen in the tropical Pacific may be Hyperoodon
planifrons or an unnamed species (Klinowska 1991).
The biology of the Northern Bottlenose Whale was
reviewed by Benjaminsen and Christensen (1979)
and Mead (1989). The general status of the species
in the North Atlantic was described by Reeves,
Mitchell and Whitehead (1993).

Distribution and Stock Identity
Northern Bottlenose Whales are present consis-
tently, throughout the year, ina 20km X 8 km “core

area,” at the entrance of the Gully, a submarine
canyon on the edge of the Scotian Shelf (Faucher and
Whitehead 1991; Reeves, Mitchell, and Whitehead
1993; Figure 2). They are also sighted, more rarely,
off the edge of the Scotian Shelf to the east and west
of the Gully (Figure 2), and there are very occasional
reports from the edge of the U.S. Shelf (Reeves,
Mitchell, and Whitehead 1993). During the Canadian
Patrol Frigate Shock Trials in November 1994,
Northern Bottlenose Whales were observed twice
and heard once near the detonation site (42° 05'N
61° 20’W), 110 km from the Shelf Break and 200 km
from the Gully (Parsons 1995; Figure 2).

The Gully is the southernmost area in the western
North Atlantic where Northern Bottlenose Whales
are found on anything other than on a very occa-
sional basis. The nearest other region where the
species may be consistently sighted is off northern
Labrador, 1400 km to the north.

Analysis of photographic identifications of indi-
vidual whales suggests that the animals in the Gully

*Reviewed and Approved by COSEWIC 16 April 1996 - status assigned: Vulnerable.

287

288

THE CANADIAN FIELD-NATURALIST

Vol. 111

agit a ee,

= Mbt PRs he,

FiGuRE |. Northern Bottlenose Whale photographed in the Gully showing signs of entangle-

ment in fishing gear.

at any time are about 57% of a population number-

ing about 230 animals (Whitehead et al. 1997;
H. Whitehead, unpublished). The geographic range
of this population is unknown but, based on the pat-
tern of sightings, we suspect that it is principally
the slope waters south of Nova Scotia. If the Gully
animals are an integral, freely-mixing part of the
population off northern Labrador, then the total
population numbers only about 230 animals —
unlikely given the numbers and geographical
spread of recent sightings in northern waters
(Reeves, Mitchell and Whitehead 1993).

The population analysis (Whitehead et al. 1997)
does not rule out occasional migrations of animals
between the northern (Labrador-Davis Strait) areas
and the Gully. The estimates of mortality + emigra-
tion + mark change are about 12% per year, and
have wide confidence limits. We have no objective
means of allocating the 12% between these three
possible causes. No estimates of mortality are avail-
able for Northern Bottlenose Whales, although mor-
tality is believed to be about 6% per year in the
Sperm Whale (Physeter macrocephalus), the species
most ecologically similar to the Northern Bottlenose
for which data are available (Rice 1989). Thus we
can only conclude from the population analysis that
emigration rates from the Gully population, and
immigration rates into it, are probably less than
about 10% per year and may be negligible.

Additional evidence that the Northern Bottlenose
Whales in the Gully are largely distinct from those
off Labrador comes from an examination of length
distributions (Whitehead et al. 1997). The photo-

graphically measured animals in the Gully are
about 0.7 m shorter than those caught off northern
Labrador. The same effect is true for the two sexes:
the Labrador population is 0.55 m larger for all
males, 0.94 m for females. Some of the difference
could be due to differential selection by the whalers
and our photographic measurement methods.
However, this cannot account for all the difference:
about 10% of the Labrador population were greater
than 8.5 m long, but animals this large were virtual-
ly absent from the Gully. It is possible that only
young animals visit the Gully. However, we do see
distinctive mature males as well as females with
calves. A 6.15 m male with five growth layers in its
teeth which stranded in the Bay of Fundy, and was
thus likely from the Gully population, lay below the
growth curve for animals caught in Labrador
(Mitchell and Kozicki 1975) and is further evidence
that individuals in the Gully population are smaller
than those from Labrador.

Our observations also indicate that the Bottlenose
Whales of the Gully may be on a different breeding
schedule to the Labrador population, which mates
and gives birth in April (Benjaminsen 1972). In the
Gully, we have five high-quality measurements
(probably representing two calves) of 3.0 to 3.3m
in August, and none between 3.3 to 4.0 m. Mead
(1989) suggests 3.5 m for the mean length at birth
of the populations studied in northern waters. Thus,
even allowing for smaller animals in the Gully, the
3.0 to 3.3 m calves observed in the Gully in August
were likely recently born. The gestation period of
Bottlenose Whales is about 12 months so that mat-

1997,

=]

Cohasset
Panuké’

WHITEHEAD, FAUCHER, GOWANS, AND MCCARREY: NORTHERN BOTTLENOSE WHALE

289

Scotian
Shelf

6 o Ps a @
Sane Primrose
° Sable Is
o oO

Poe

FIGURE 2. The Scotian Shelf, showing the Gully, oil and gas discoveries from Wade et al.
(1989) (circles, the Cohasset/Panuke and Primrose fields are filled), major shipping route
(dashed line), the core area of the Northern Bottlenose Whale population (shaded), and
additional sightings and catches of the species (m) from Reeves, Mitchell and Whitehead
(1993), the Sea Education Association and our own sightings. The site of the Canadian
Patrol Frigate Shock Trial at which Bottlenose Whales were sighted is marked on the inset

by a “+”.

ing and calving occur at the same time of year
(Benjaminsen and Christensen 1979). In the Gully,
the proportion of mixed groups of males and
females rises through June-August (proportion of
groups with both adult males and females: 10%
June; 17% July; 28% August), consistent with an
August mating/parturition season.

Protection

National legislation and international conven-
tions protecting Northern Bottlenose Whales from
hunting are described by Reeves, Mitchell and
Whitehead (1993). Neither now, nor in the near
future, is the population in the Gully likely to be
subject to whaling.

Several initiatives have been undertaken to pro-
tect the Gully population. In 1990, the oil company,
Lasmo, which was beginning exploitation the
Cohasset-Panuke field 110 km to the west of the
Gully, declared a “tanker exclusion zone” including
the Gully, so that shipping associated with the

development does not interfere with the whales.
The Department of Fisheries and Oceans has desig-
nated a “Whale Sanctuary” in the Gully for the
Northern Bottlenose Whales and, in the Canadian
Notices to Mariners Annual Edition (1994), pub-
lished guidelines for the behaviour of vessels with-
in the sanctuary. Shipping companies have been
asked (by letter) to avoid the area, and many have
agreed to do so. The establishment of a Marine
Protected Area in the Gully is being considered by
the Canadian Wildlife Service, the Department of
Fisheries and Oceans, Parks Canada, World
Wildlife Fund Canada and other organizations and
individuals (Amirault 1995).

Population Sizes and Trends

The population of Northern Bottlenose whales in
the Gully has been assessed using mark-recapture
methods on photographic individual identifications
collected between 1988 and 1995 (Whitehead et al.,
1997). This analysis suggests a population of about

290

230 animals (approximate 95% confidence interval
160-360). Data are insufficient to detect any trends
in population size.

Eighty-seven Northern Bottlenose Whales were
taken by whalers working from Blandford, Nova
Scotia between 1962-1967 (Reeves, Mitchell, and
Whitehead 1993). The great majority of these seem
to have been killed in, or near, the Gully (Reeves,
Mitchell, and Whitehead 1993). Assuming that the
population size is now somewhere between its lev-
els immediately before and after this whaling, a 29
to 41% reduction in numbers during the whaling
period is indicated.

Habitat

The habitat of the Bottlenose Whales south of
Nova Scotia is the waters near the edge of the con-
tinental shelf which are greater than 1000 m deep,
but the overwhelming focus of their distribution is
the 20 km x 8 km core area at the entrance of the
Gully. In terms of relief, and penetration into the
shelf, the Gully is the most prominent canyon in
the western North Atlantic. The whales are never
seen in waters less than about 800 m deep, even
though such depths are within a few km of their
core habitat.

Protection of this core habitat is likely to be
essential for the survival of the population.

General Biology

The general biology of Northern Bottlenose Whales
was described by Benjaminsen and Christensen
(1979), Mead (1989) and Reeves, Mitchell and
Whitehead (1993).

Limiting Factors

The size of the population that uses the Gully is
small, about 230 animals. This is well below the
2000 beneath which IUCN (International Union for
the Conservation of Nature) classifies cetacean
species as Endangered (Klinowska 1991).
However, the Gully Bottlenose Whales, although
apparently largely or totally distinct from the popu-
lations in more northern waters, are not (as far as
we know) a different species. It seems likely that
the population is naturally small, limited by avail-
able habitat in the area to the low hundreds.

The Gully population is at the extreme southern
limit of the species’ range in the western North
Atlantic. This likely limits the potential for alterna-
tive suitable habitat in nearby sea areas.

The proximate threats to these animals from
humans are principally:

Collisions with ships. Each year a number of whales
are found dead in the waters off Nova Scotia follow-
ing collisions with shipping (J. Conway, Department
of Fisheries and Oceans, Scotia-Fundy Region,
Halifax, Nova Scotia; personal communication),

THE CANADIAN FIELD-NATURALIST

Vol. 111

although there are no known reports of Bottlenose
Whale fatalities.

Acoustic pollution. Whales communicate and sense
their environment largely through the acoustic
channel. Noise affects the behaviour and movement
of whales; it has the potential to interfere with feed-
ing or mating, or cause physiological damage
(Richardson et al. 1991; Committee on low-fre-
quency sound and marine mammals 1994).

Fishing Gear. Entanglement in fishing gear (that in
active use as well as discarded, abandoned or lost
gear) is a major threat to many cetacean populations
(Cooke 1991). A number of the Bottlenose Whales
in the Gully show evidence of encounters with fish-
ing gear (e.g., Figure 1).

Marine Debris. Entanglement in floating debris,
such as plastic bags and discarded strapping, is
a source of mortality for many marine animals,
including cetaceans (Cooke 1991). The Gully has a
high level of such pollution (Dufault and
Whitehead 1994).

Chemical Pollution. The most obvious sources of
these dangers are commercial shipping, fishing
activity and petrochemical exploration and exploita-
tion. Explosions and loud underwater sounds made
for naval and scientific purposes may also pose a
threat (Richardson et al. 1991; Committee on low-
frequency sound and marine mammals 1994).

Ships may collide with the whales, are a source of
acoustic pollution, and can contribute to marine
debris and chemical pollution. The major east-west
trans-Atlantic shipping route lies about 30 km south
of the Gully core area (Figure 2). Commercial ships
(excluding fishing vessels) transit the core area of
the Bottlenose Whales about once per day (H.
Whitehead, unpublished data). This rate may have
decreased somewhat since the Department of
Fisheries and Oceans’ guidelines were published in
the Notices to Mariners, and requests for avoidance
were sent to shipping companies.

The shallow areas bordering the Gully were
heavily dragged for groundfish, while midwater
draggers take Redfish (Sebastes sp.) from within
the core area. Fishing vessels, especially while
dragging, are extremely noisy (H. Whitehead, per-
sonal observation). They may entangle whales in
their active, lost or discarded gear, and are sources
of other marine debris. The crash of groundfish
stocks during the last few years has considerably
reduced fishing activity in the area of the Gully.
Harpooning and long-lining for Swordfish (Xiphius
gladius) also takes place in the core area but this is
less obviously harmful.

Oil and gas have been found in commercially
exploitable quantities on the Scotian Shelf bordering
the Gully. One find, the “Primrose” Field, lies about
5 km from the core area of the Bottlenose Whales
(Figure 2). The only current exploitation in the area

1997

is at the “Cohasset/Panuke” condensate fields
(Figure 2). These are 110 km from the Gully, and
their exploitation probably poses little threat to the
bottlenose whales. In the next few years, a consor-
tium of oil companies led by Mobil Oil Canada
plans to exploit some of the gas discoveries around
Sable Island, the closest of which, the “Venture”
Field, is about 45 km from the core area of the
Bottlenose Whales in the Gully.

Oil and gas exploitation has the potential to harm
the Bottlenose Whales directly through the noise of
the drilling and other operations, spills and discard-
ed material, but also indirectly because of an
increase in shipping traffic. Noises associated with
offshore oil and gas production disturbed the
behaviour of Bowhead Whales (Balaena glacialis)
to ranges of about 3 to 11 km (Richardson, Wursig,
and Greene 1990). It is not known how sensitive
the Bottlenose Whale is to acoustic disturbance but
the most ecologically similar species for which
there are any data, the Sperm Whale, is especially
easily disturbed by sound (e.g., Watkins and
Schevill 1975; Watkins, Moore and Tyack 1985).
Bottlenose Whales have particularly weak social
sounds (Winn, Perkins, and Winn 1970) which
might suggest vulnerability to acoustic disturbance.

The most important limiting factor for the popula-
tion of Bottlenose Whales in the Gully is likely the
pattern and method of development of these oil and
gas fields.

Special Significance of the Population

Our research on the Northern Bottlenose Whales
in the Gully is unique: it is the only long-term study
of any beaked whale population, anywhere in the
world — these are the only living ziphiids to be
individually identified. There are no known loca-
tions at all comparable to the Gully in terms of the
potential for studies of beaked whale populations.
In the near future, most of what is known about the
natural behaviour of living beaked whales is likely
to come from the Gully. The population is not only
amenable to research: two film crews have visited
the Gully, and have successfully filmed the whales;
and local tourist operators are making plans to
bring limited numbers of whale watchers to the
Gully.

Unusual biological features of the Bottlenose
Whale include their maxillary crests, their deep and
prolonged dives, the types of sounds produced, and
pronounced curiosity towards boats (Mead 1989;
Reeves, Mitchell, and Whitehead 1993).

Evaluation

The population of Northern Bottlenose Whales in
the Gully is small, about 230 animals, at the south-
ern extreme of the species’ range, and largely or
totally distinct from the larger populations further

WHITEHEAD, FAUCHER, GOWANS, AND MCCARREY: NORTHERN BOTTLENOSE WHALE

291

north. The animals seem to be non-migratory,
spending an average of 57% of their time in a 20 km
x 8 km core area, which is bathometrically unique
in the western North Atlantic. These characteristics
make the population particularly sensitive to human
activities. Thus the population should be, at the
least, considered vulnerable.

The development and exploitation of the oil and
gas fields on the Scotian Shelf is approaching the
Gully, threatening the Bottlenose Whales and other
inhabitants of the Gully, which include exceptional
numbers of some other cetacean species (Gowans
and Whitehead 1995). With exploitation of the
Primrose Field, the core area of the Bottlenose
Whales in the Gully may be abandoned, endanger-
ing the population. The current plans for progres-
sive development of the oil and gas fields near the
Gully suggest that this population should be desig-
nated vulnerable.

Acknowledgments

Our research on the Northern Bottlenose Whales
of the Gully, and the preparation of this report,
have been funded by World Wildlife Fund Canada,
the Whale and Dolphin Conservation Society, the
Natural Sciences and Engineering Research
Council of Canada and the Canadian Federation of
Humane Societies. Thanks to the Sea Education
Association for sighting records, and to David
Gaskin, Randall Reeves and Linda Weilgart for
useful comments on manuscripts.

Literature and Documents Cited

Amirault, D. L. 1995. Towards a conservation strategy
for the Gully, near Sable Island, Nova Scotia. Discus-
sion paper, Canadian Wildlife Service, Atlantic Region,
P.O. Box 1590, Sackville, New Brunswick E0OA 3CO.
8 pages.

Benjaminsen, T. 1972. On the biology of the bottlenose
whale, Hyperoodon ampullatus (Forster). Norwegian
Journal of Zoology 20: 233-241.

Benjaminsen, T., and I. Christensen. 1979. The natural
history of the bottlenose whale, Hyperoodon ampullatus
(Forster). Pages 143—164 in Behavior of marine animals.
Volume 3. Edited by H. E. Winn and B. L. Olla. Plenum,
New York.

Christensen, I. 1975. Preliminary report on the Nor-
wegian fishery for small whales: expansion of
Norwegian whaling to arctic and northwest Atlantic
waters, and Norwegian investigations of the biology of
small whales. Journal of the Fisheries Research Board of
Canada 32: 1083-1094.

Committee on low-frequency sound and marine mam-
mals. 1994. Low frequency sound and marine mam-
mals: current knowledge and research needs. National
Academy of Sciences, National Academy Press. 97
pages.

Cooke, J. G. 1991. Introduction and overview. Pages
4—18 in Dolphins, porpoises and whales of the world.
The IUCN Red Data Book. Edited by M. Klinowska.
IUCN, Gland, Switzerland.

pas

Dufault, S., and H. Whitehead. 1994. Floating marine
pollution in ‘the Gully’ on the continental slope, Nova
Scotia, Canada. Marine Pollution Bulletin 28: 489-493.

Faucher, A., and H. Whitehead. 1991. The bottlenose
whales of “the Gully.". Final report for 1988-1990 pro-
ject to WWF-Canada. World Wildlife Fund Canada,
Toronto.

Gowans, S., and H. Whitehead. 1995. Distribution and
habitat partitioning by small odontocetes in the Gully, a
submarine canyon on the Scotian Shelf. Canadian
Journal of Zoology 73: 1599-1608.

Klinowska, M. 1991. Dolphins, porpoises and whales of
the world. The IUCN Red Data Book. IUCN, Gland,
Switzerland.

Mead, J. G. 1989. Bottlenose whales Hyperoodon ampul-
latus (Forster, 1770) and Hyperoodon planifrons
(Flower, 1882). Pages 321-348 in Handbook of marine
mammals. Volume 4. River dolphins and the larger
toothed whales. Edited by S. H. Ridgway and R. Har-
rison. Academic Press, London.

Mitchell, E., and V. M. Kozicki. 1975. Autumn strand-
ing of a northern bottlenose whale (Hyperoodon ampul-
latus) in the Bay of Fundy, Nova Scotia. Journal of the
Fisheries Research Board of Canada 32: 1019-1040.

Parsons, J. 1995. Marine Mammal Monitoring Program--
Canadian Patrol Frigate Shock Trials, November 8-21,
1994. Final report to Department of National Defence,
Ottawa by John Parsons & Associates.

Reeves, R. R., E. Mitchell, and H. Whitehead. 1993.
Status of the northern bottlenose whale, Hyperoodon
ampullatus. Canadian Field-Naturalist 107: 490-508.

Rice, D. W. 1989. Sperm whale. Physeter macrocephalus

THE CANADIAN FIELD-NATURALIST

Vol. 111

Linnaeus, 1758. Pages 177—233 in Handbook of marine
animals. Volume 4. Edited by S. H. Ridgway and R.
Harrison. Academic Press, London.

Richardson, W. J., B. Wursig, and C. R. Greene. 1990.
Reactions of bowhead whales, Balaena mysticetus, to
drilling and dredging noise in the Canadian Beaufort
Sea. Marine Environmental Research 29: 135-160.

Richardson, W. J., C. R. Greene, C. I. Malme, and D.
H. Thomson. 1991. Effects of noise on marine mam-
mals. U.S. Department of the Interior, Minerals
Management Service, Atlantic OCS Region, Herndon,
Virginia.

Wade, J. A., G. R. Campbell, R. M. Procter, and G. C.
Taylor. 1989. Petroleum resources of the Scotian
Shelf. Geological Survey of Canada Paper 88-19.

Watkins, W. A., K. E. Moore, and P. Tyack. 1985.
Sperm whale acoustic behaviors in the southeast Carib-
bean. Cetology 49: 1-15.

Watkins, W. A., and W. E. Schevill. 1975. Sperm whales
(Physeter catodon) react to pingers. Deep-Sea Research
22: 123-129.

Whitehead, H., S. Gowans, A. Faucher, and S.
McCarrey. 1997. Population analysis of northern bot-
tlenose whales in the Gully, Nova Scotia. Marine
Mammal Science. 13: 173-185.

Winn, H. E., P. J. Perkins, and L. Winn. 1970. Sounds
and behavior of the northern bottlenose whale. 7th
Annual Conference on Biological Sonar and Diving
Mammals, Stanford Research Institute, Menlo Park,
California. Pages 53-59.

Accepted 8 July 1996

Status of the Sperm Whale, Physeter macrocephalus, in Canada*

RANDALL R. REEVES! and HAL WHITEHEAD?

'Okapi Research Associates, 27 Chandler Lane, Hudson, Quebec JOP 1HO
"Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1

Reeves, Randall R., and Hal Whitehead. 1997. Status of the Sperm Whale, Physeter macrocephalus, in Canada.
Canadian Field-Naturalist 1 11(2): 293-307.

The Sperm Whale (Physeter macrocephalus) is the largest of the toothed whales. Members of this species are extreme, or
distinctive, among cetaceans in many ways which include their large brain size, sexual dimorphism, diving behaviour,
social organization, communication system and ecological significance. They occur in all the deeper waters of the world’s
oceans that are not ice-covered, although females are generally restricted to areas where surface temperatures are warmer
than 15°C and depths are greater than 1000 m. Sperm Whales range widely with no clear divisions between populations.
Sperm Whales were depleted during two massive, world-wide hunts (peaking in approximately the 1840s to the 1860s and
the 1950s to the 1970s). The current world population likely numbers a few hundred thousand, but there are considerably
fewer Sperm Whales now than there were before whaling. Recovery of the population is slow because of a very low repro-
ductive rate, possibly compounded by the lingering effects of a socially disruptive hunt concentrated on large males. Sperm
Whales are found off both the east and west coasts of Canada with particular concentrations at the entrance to the Hudson
Strait, on the Scotian Shelf and west of Vancouver Island. Both sexes occur off western Canada, but females only occa-
sionally use the southern most waters off Atlantic Canada. Although no immediate threats to Sperm Whale populations are
known, these animals are vulnerable to particular kinds of human disturbance, and populations may be threatened, in the
long term, by increasing levels of pollutants in the oceans.

Le cachalot macrocéphale (Physeter macrocephalus) est le plus gros des odontocétes. Les membres de cette espéce sont
extrémes ou distinctif parmi les cétacés a bien des égards qui comprend leur large cerveau, leur dimorphisme sexuel, leur
comportement de plongé, leur organisation sociale, leur systeme de communication, et leur importance écologique. IIs
habitent toutes les eaux plus profondes des océans du monde ou il n’y a pas une couverture de glace, bien que les femelles
soient généralement limitées aux zones ou les températures a la surface sont plus chaudes que 15°C et la profondeur est
supérieure 4 1000 m. Les cachalots parcourent tout les océans et ils n’ont aucune division distincte entre les populations.
Le nombre des cachalots a été épuisé par deux chasses massives mondiales (apogée approximativement aux environs de
1840 a 1860 et encore de 1950 a 1970). Aujourd’hui, les cachalots sont considérablement moins nombreux qu’avant la
péche a la baleine bien que la population mondiale du jour est probablement quelques centaines de milles. Le récouvrement
de la population est lent 4 cause d’un bas taux de reproduction, et de l’effet possible d’une chasse socialement destructive
qui a concentrée son attention sur les gros males. Au Canada, on retrouve les cachalots au large des cOtes ouest et est avec
des centres de convergence dans |’entrée du Détroit de Hudson, sur la plate-forme néo-écossaise, et a l’ouest de Vile de
Vancouver. Au large de la Colombie-Britannique on peut retrouver les deux sexes, mais les femelles sont rarement trou-
vées dans les eaux les plus méridionales au large de |’ Atlantique canadien. Quoique on ne connaisse aucune menace immé-
diate envers la population des cachalots, mais ces baleines sont vulnérables a certains dérangements provoqués par les
humains et la population peut étre menacée, dans le long terme, par les niveaux de polluants croissant dans les océans.

Key Words: Sperm Whale, cachalot, cachalot macrocéphale, Physeter macrocephalus, Cetacea, Odontoceti, conservation
status, Canada.

The Sperm Whale (Physeter macrocephalus
Linnaeus 1758) is the world’s largest living odonto-
cete, or toothed whale (see front cover). Males grow
to a maximum length of 18 m and reach weights of
nearly 60 metric tons. Females are much smaller,
with a maximum length of about 11 m and a weight
of 24 metric tons (Rice 1989). The Sperm Whale is
extreme among mammals in a number of ways in
addition to its large size and sexual dimorphism. It
has the largest brain on Earth, and the largest geo-
graphical separation of adult males from adult
females. The aggregate world biomass is among the
highest of any wild mammal, and this distinction
probably would have been more marked in pre-

whaling times. Sperm Whales may dive deeper than
any other mammal, although they are rivalled by
Elephant Seals (Mirounga spp.) and Bottlenose
Whales (Hyperoodon spp.), and they seem to have
one of the most caring of marine mammal societies.
The world’s Sperm Whale populations were the tar-
get of two massive commercial hunts. The first,
using open rowed whale-boats and hand-thrown har-
poons, lasted from the early 18th Century to the
early 20th Century, peaking in the 1840s and 1850s.
Modern whalers aboard steam or diesel powered
catcher vessels with deck-mounted harpoon guns
took hundreds of thousands of Sperm Whales
between 1946 and 1978 (Best 1983).

*Reviewed and approved by COSEWIC 16 April 1996, status assigned: Not At Risk (NAR).

293

294

There is only one species of Physeter, although
the appropriate nomenclature and evolutionary histo-
ry of the Sperm Whale are matters of debate. Most
scientists and organizations (including the Inter-
national Commission for Zoological Nomenclature
and the International Whaling Commission) call the
Sperm Whale Physeter macrocephalus (Husson and
Holthuis 1974; Holthuis 1987; Rice 1989). Schevill
(1986, 1987), however, argued that Physeter catodon
was the proper Linnaean name for the species. The
Sperm Whale is generally classified in the suborder
Odontoceti, the toothed whales, but molecular evi-
dence has recently been interpreted as suggesting
that it is more closely related to the baleen whales
(suborder Mysticeti) and therefore that the odonto-
cetes are not monophyletic (Milinkovitch, Guillermo
and Meyer 1993).

This review discusses the conservation status of
the Sperm Whale, with particular attention to the
animals of the North Atlantic and North Pacific.

Distribution and Stock Identity

The Sperm Whale’s distribution spans virtually all
latitudes between the north and south polar ice caps.
It has the widest distribution of any marine mammal
except the Killer Whale (Orcinus orca) (Rice 1989).
The Sperm Whale, generally an animal of deep
waters, is particularly abundant off the outer edges
of the world’s continental and island shelves, in or
near upwelling areas, and in deeper basins of seas
and gulfs (Townsend 1935; Berzin 1971; Jaquet
1996). Females and young animals are generally
restricted to tropical and temperate waters whereas
mature males are more common at higher latitudes
(Rice 1989).

Both tagging experiments carried out with the
whaling industry and resightings of individually
identified animals indicate that female Sperm
Whales have ranges spanning very approximately
1000 km, although there is a great deal of variation
(Best 1979; Kasuya and Miyashita 1988; Dufault
and Whitehead 1995). Mature males have larger
ranges, migrating from high-latitude feeding grounds
to breed in tropical and subtropical waters (Best
1979). They also make considerable longitudinal
migrations (see below).

The delineation of Sperm Whale stocks was a
contentious issue at meetings of the Scientific
Committee of the International Whaling Com-
mission during the 1970s and early 1980s (e.g.,
International Whaling Commission 1980). Evidence
from different sources was often contradictory. Part
of the problem lay in the very different ranging pat-
terns of males and females, but the weight of evi-
dence suggests that there are no well-defined Sperm
Whale stocks — no areas in the ocean which Sperm
Whales rarely cross, or even areas which they rarely
cross at a particular time of year. For instance female

THE CANADIAN FIELD-NATURALIST

Vol. 111

Sperm Whales which visit the Galapagos Islands
also use waters off mainland Ecuador, 1000 km
away, where they mix with females which do not
visit the Galapagos (Dufault and Whitehead 1993).
In a study of variation in the maternally-inherited
mitochondrial DNA of female Sperm Whales, strong
group-level similarities among females were found,
indicating genetic relatedness within groups, but no
geographic structure at any scale: females in differ-
ent groups in the same area were as likely to share
mitochondrial haplotypes as those in different oceans
(Dillon 1996). Thus it seems that groups of female
Sperm Whales, and individual males, have ranges,
but that the population has no other clear geographic
structure.

North Atlantic

The overall North Atlantic distribution of adult
males extends north to at least 69°N in Denmark
Strait and to 71°N west of Jan Mayen and north of
Norway (Rice 1989). Sperm Whales are found in the
Caribbean Sea, the Gulf of St Lawrence and the
Mediterranean Sea (Viale 1977; Rice 1989; see
below). In the Atlantic, females and young males
rarely move north of the Subpolar Convergence (45
to 50°N, farther north in the east than the west).

Off the east coast of Canada, Sperm Whales have
not been reported north of Hudson Strait, although
they certainly move into higher latitudes (at least to
64°N) off West Greenland (Kapel 1979; Perkins et
al. 1982). The lack of records off the east coast of
Baffin Island may be due partly to the whales’ off-
shore distribution and the small amount of research
effort there. Large males are common at the eastern
entrance to Hudson Strait in late August and
September (Mitchell 1974; MacLaren Atlantic
Limited 1977; MacLaren Marex Inc. 1979), and
some enter Hudson Strait and Ungava Bay (Reeves
et al. 1986). Male Sperm Whales are present off, and
sometimes on, the continental shelf along the entire
east coast of Canada south of Hudson Strait (e.g.;
Mitchell 1974; 1975; Mitchell and Kozicki 1984;
Whitehead, Brennan, and Grover 1992; Parsons
1995; Figure 1). They are found with considerable
reliability in the northern part of the Gully (44°N,
59°W), a submarine canyon on the edge of the
Scotian Shelf (Whitehead, Brennan and Grover
1992). Sperm Whales are found in the Gulf of St.
Lawrence, usually only sporadically and in relatively
small numbers (Richard Sears, Mingan Island
Cetacean Study, 285 Green Street, St. Lambert,
Quebec; personal communication), although one to
four Sperm Whales, including one well-known indi-
vidual, “Tryphon”, were seen off Tadoussac in the
St. Lawrence Estuary quite consistently during the
summer months (May-October) between 1991 and
1995 (R. Michaud, GREMM, Tadoussac, Quebec;
personal communication).

| gs

120°

REEVES AND WHITEHEAD: STATUS OF THE SPERM WHALE

23

FiGuRE 1. Approximate distribution of Sperm Whales in waters near Canada. The approximate northern limit for females
is shown by a solid line in both the North Pacific and North Atlantic.

Sperm Whales have stranded in several parts of
eastern Canada, throughout the year (Table 1). Dead
Sperm Whales are reported in the Gulf of St.
Lawrence more often than might be expected judg-
ing by the infrequency of sightings. For example,
between 1982 and 1985 Sperm Whales represented
close to 4 percent of the total documented cetacean
mortalities in the Gulf (Béland et al. 1987).

The fishery for Sperm Whales off Nova Scotia
only took males (Mitchell 1975), and recent studies
of living Sperm Whales on and near the Scotian Shelf
have focused almost entirely on males (Mullins,
Whitehead, and Weilgart 1988; Whitehead, Brennan,
and Grover 1992). Recent strandings off eastern
Canada have also been principally of males (Table 1).
However, a female stranded in July 1994 in southern
Nova Scotia (Table 1), and groups of females have
twice been observed in the Gully: once on 23 June
1990 (Whitehead, Brennan and Grover 1992), and
once (tentative identification) on 15 July 1993 (H.
Whitehead, unpublished data). Thus, in the Atlantic,
the southern limit of the Canadian EEZ appears to
coincide approximately with the northern limit of the
range of female Sperm Whales (Figure 1).

A male tagged off Nova Scotia (42° 12’N, 65°
07'W;, 26 July 1966) was taken by whalers off north-
western Spain (44° 10’N, 11° 20’W; 7 August
1973), some 6000 km away (Mitchell 1975).

Another male taken in the northern part of Denmark
Strait (ca. 65°N, 29° 45’W) in early August 1981
apparently had been wounded the previous August
by whalers near the Azores about 3000 km to the
south (Martin 1982; Sigurj6nsson 1985). Several
Azorean-type hand harpoons (or harpoon fragments)
were found in Sperm Whales killed off northwestern
Spain (Aguilar 1985). Thus Sperm Whales move
across large parts of the North Atlantic, and the
International Whaling Commission has accordingly
managed whaling for Sperm Whales in the North
Atlantic on the assumption that there is only one
population.

However, the equator does not act as a barrier to
Sperm Whale movements, so that distinguishing
North Atlantic versus South Atlantic populations is
artificial. A young male tagged off Mauritania, West
Africa, in late November one year was killed west of
Cape Town, South Africa, in March somewhat more
than four years later (Ivashin 1967). This animal had
moved between an area near the Tropic of Cancer to
an area far south of the Tropic of Capricorn.

North Pacific

Sperm Whales are found throughout the North
Pacific, including the deeper waters of the Bering
Sea, the South China Sea, the East China Sea, the
Sea of Japan, the Sea of Okhotsk, the Gulf of Alaska

296

and the Gulf of California (Rice 1989). Females and
young males generally do not move north of the
Transitional Domain, just north of the Subarctic
Boundary (ca. 42°N in most of the North Pacific)
[Rice 1989]. Berzin and Rovnin (1966) gave the
northern boundary for the summer range of females
and young males in the eastern North Pacific as the
southern Gulf of Alaska, approximately in the lati-
tude of 50-51°N. The occasional occurrence of
females as far north as the Aleutian Islands may be
explained by exceptional incursions of warm water
into high latitudes (Pike and MacAskie 1969).

Sperm Whales are present in the deep waters off
the west coast of Canada (Figure 1), and strandings
have been reported at a low, but quite steady, rate
(Table 1). The whaling grounds for Sperm Whales
off Vancouver Island were in the open ocean up to
200 nautical miles (370 km) from shore. Large con-
centrations, including herds of 50 to 150 animals,
were encountered there from late spring through
early autumn (Pike and MacAskie 1969). Sperm
Whales are observed occasionally in Hecate Strait,
Dixon Entrance, Queen Charlotte Sound, and the
broader inside waterways north of Vancouver Island
(Pike and MacAskie 1969; H. Patterson, Marine
Mammal Research Group, Box 6244, Victoria, B.C.
V8P 5L5; personal communication).

Ninety percent of the Sperm Whales taken in
British Columbia were males averaging 11.6-14.0 m
long, and most of them were sexually mature (Pike
and MacAskie 1969). This strong bias in the catch
could have been at least partially due to legal restric-
tions, including requirements that only whales 35 ft
(10.7 m) or longer be hunted and that females
accompanied by calves not be taken. Females and
young are present in the deep waters off Vancouver
Island (Pike and MacAskie 1969) and, in some
years, the Queen Charlotte Islands (Reeves et al.
1985).

Tagged Sperm Whales have crossed much of the
North Pacific, as well as from the North Pacific to
the South Pacific (Kasuya and Miyashita 1988).
Despite this, there has been considerable inconclu-
sive debate as to whether the Sperm Whales in the
North Pacific constitute one, two, three or more
stocks (e.g., Rice 1974; Bannister and Mitchell 1980;
International Whaling Commission 1980; Kasuya
and Miyashita 1988; Donovan 1991).

Exploitation and Protection
North Atlantic

Sperm Whales were hunted by American and
European whalers on many grounds throughout the
North Atlantic, beginning in the early 1700s
(Starbuck 1878). The 18th and 19th Century hunting
grounds of the American whaling fleet included the
Labrador and Newfoundland coasts, the Gulf of St.
Lawrence, and the Grand Banks.

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Vol. 111

Modern whaling began in the 1860s. Although
rorquals were the principal targets of most modern
whaling operations in the North Atlantic, substantial
catches of Sperm Whales (100 or more per year)
were made in Iceland (mid-1950s to 1982), Madeira
(to 1981), the Azores (to 1987), and Spain (early
1950s to 1980), and moderate but regular catches
were made in Norway, the Faroe Islands, Portugal,
and Canada (Jonsgard 1977). Small numbers of
Sperm Whales continued to be taken in the Lesser
Antilles as recently as the 1980s (Price 1985; Reeves
1988).

Whalers in Newfoundland and Labrador caught
from 0 to 53 Sperm Whales per year (mean 6/yr)
from 1904 to 1970 (Mitchell 1974). Sperm Whales
did not become major targets of shore whaling in
Nova Scotia until 1970, when 25 were taken
(Mitchell 1974). From 1964, when operations began
at Blandford, through 1972, when commercial whal-
ing in Canada stopped, the total Nova Scotian catch
of Sperm Whales was 109 males and no females
(Mitchell 1975). Sperm Whales generally were taken
at Blandford only when Fin Whales (Balaenoptera
physalus) and Sei Whales (Balaenoptera borealis)
were unavailable, due either to their absence on the
grounds or to the fact that quotas had been filled
(Mitchell 1975).

There was no Sperm Whale catch limit in the
North Atlantic until 1977, when the International
Whaling Commission set a quota of 685 whales per
year (SigurjOnsson 1988). In 1980 the quota was
reduced to 273, divided by bilateral agreement
between Spain (173) and Iceland (100) and in 1981
it was further reduced to 130 (Iceland only; Spain
ceased whaling). The Sperm Whales in the North
Atlantic were classified as a protected stock begin-
ning in 1982. Catching at Madeira and the Azores
was unaffected by the regulations because Portugal
was not a member of the International Whaling
Commission.

Currently (1996) we are unaware of any directed
catch of Sperm Whales in the North Atlantic.

North Pacific

The American and European pelagic whaling
fleets began visiting Sperm Whale grounds in the
Pacific Ocean in the late 1700s (Starbuck 1878).
Thereafter, tens of thousands of Sperm Whales were
taken in the North Pacific by the American, French,
and British fleets (du Pasquier 1982; Best 1983;
Webb 1988). The open-boat fishery in the North
Pacific was especially intense between about 1825
and 1845 (Tillman and Breiwick 1983).

Modern whaling was introduced to the North
Pacific in the last decade of the 1800s, and floating
factories and shore stations proliferated during the
early decades of the 20th Century (Ohsumi 1980). A
list of reported Sperm Whale catches throughout the

1997

REEVES AND WHITEHEAD: STATUS OF THE SPERM WHALE

207

TABLE 1. Known strandings of Sperm Whales on Canadian coasts, including dead animals discovered at sea, during 1987
to 1995 (Q.C.I. = Queen Charlotte Islands; P.E.I. = Prince Edward Island) from R. Baird and H. Patterson (Marine
Mammal Research Group, Box 6244, Victoria, British Columbia), P.-Y. Daoust (University of Prince Edward Island
Veterinary College), M. Dillon (Department of Biology, Dalhousie University, Halifax, Nova Scotia), Z. Lucas (P.O. Box
3504, South Halifax, Nova Scotia), S. Todd (Whale Research Group, Memorial University of Newfoundland), R. Michaud
(GREMM, Tadoussac, Quebec), and D. McAlpine (New Brunswick Museum, Saint John, New Brunswick).

Date Position
Newfoundland/Labrador
11 June 1987 Red Harbour, Placentia Bay
8 July 1987 Penguin Island West, South Coast
1 March 1987 Conception Harbor, Conception Bay

30 September 1990
15 January 1991
6 March 1995

Nova Scotia
6 October 1990

Black Tickle, Labrador
Port aux Choix, West Coast
Lord’s Cove, Burin Peninsula

Sable Island

25 April 1991 Isle Petitgras, Cape Breton Island
— January 1992 Sable Island

— July 1994 Cherry Hill, S. Shore

— December 1994 River John, N. Coast

Prince Edward Island
13 December 1988
1 October 1989

Near Nail Pond, N.W.
Covehead Harbour, N.

5 October 1989 Malpeque Bay, N.

6 October 1989 N.W. of Malpeque

19 December 1991
20 December 1991
23 December 1991

New Brunswick

North Rustico, N.
Brackley Beach, N.
Brackley Beach

No. Length(m) Sex Comments
] 15 re) Dead
l 13 3 Dead
1 2 2
4 ? ? Entered harbour, driven out
1 Z ? Dead
1 15 3 Dead
2 ie ey Dead
13.8 3 Dead
2 14.5 3 Dead
HE6 3
? 3 Dead
1 9.8 2 Dead
| 2. ? Dead
1 12.8 re Dead, may have beached alive
Live stranding:
13.6 3 died on site, 5 pulled to open
water: | died and sank
1 11.4 3 Dead, probably from Cove-
head stranding, 45 km away
1 ? ? Dead, at sea, probably from
Covehead stranding
1 ? 3S
1 i 3 Dead
1 13.1 3 Dead

No strandings since 1987, altlhough some dead animals from Covehead, Prince Edward Island strandings washed ashore in

northern New Brunswick.

Quebec
13 June 1992
24 June 1992

British Columbia
26 June 1988
6 December 1988

Port Meunier, Anticosti Island
Sheldrake 50°15’N, 64°54’W

Goose Island 51°58’N, 128°26’W

Rose Pit, Queen Charlotte Islands

54°10’, 131°40’

17 November 1990
7 December 1990

22 May 1991

Tsusiat Falls 48°42'N, 124°56’W
Benson Pt, Nootka Island

49°47'N, 126°53’W

Graham Island 53°23'N, 133°11’W
White Point, Graham Island

Hecate Strait

2 November 1991
15 January 1993
1 February 1994

North Pacific from 1910 to 1976 totals more than a
quarter million, of which nearly 60 percent were
taken between 1963 and 1976 (Ohsumi 1980). Large
post-war catches were made from shore stations in
Japan and the Kuril Islands and by Japanese and
Soviet floating factories that operated all over the
North Pacific (Ohsumi 1980; Kasuya 1991).

98 nm W of Tofino 49°04'N, 128°27'W

1 je 5) 3 Dead
1 10.7 3 Dead
1 ee ? Dead
1 2 £4 Dead
1 ? 6? Dead
1 12:5 6? Dead
1 55 2 Dead
1 10 3S Dead
1 ? z Dead
1 v ? Dead

Shore-based whaling for Sperm Whales off west-
ern North America has a long history. Sperm Whales
were prominent in the catches of stations on
Vancouver Island and the Queen Charlotte Islands
(Pike and MacAskie 1969; minimum of 5937 taken
from 1919 through 1967), Kodiak and the Aleutian
Islands (Reeves et al. 1985), Washington State

298

(Scheffer and Slipp 1948), and California (Rice
1974). Shore whaling ended on the Canadian west
coast in 1967 and on the west coast of the United
States in 1971 (Ohsumi 1980).

A North Pacific Working Group was established
by the International Whaling Commission in 1951
and a Sperm Whale Subcommittee of the Scientific
Committee of the International Whaling
Commission in 1962. Based largely on the work of
these groups, the Commission began setting Sperm
Whale quotas for pelagic whaling in the North
Pacific in 1971. The quota was subdivided by sex
beginning in 1973. In 1975 North Pacific catch lim-
its were 6000 males and 4000 females (International
Whaling Commission 1977), and, in 1981, 890
males and 0 females (International Whaling
Commission 1982a). The Japanese officially
stopped catching Sperm Whales in the North Pacific
in 1988.

Worldwide

A worldwide ban on commercial whaling was
agreed by the International Whaling Commission in
1982 and took effect in 1985. By the end of the 1980s
the few remaining major shore-based whaling opera-
tions involving a catch of Sperm Whales (Iceland,
the Azores, Madeira and Japan) had ceased. The
Sperm Whale has been in Appendix I of the
Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES) since
1985. Trade in products, most importantly sperm oil,
teeth, and ambergris (a waxy substance found in the
large intestine of the Sperm Whale) is therefore ille-
gal in most countries (but not Japan and Norway,
both of which entered reservations against the
Appendix I listing). Since 1981, importation of
sperm oil and other Sperm Whale products (teeth
excepted) has been prohibited by the European
Union, and this has meant the closing of a formerly
important market (Klinowska 1991). The importa-
tion of marine mammal products has been banned in
the United States since 1972.

The only place in the world where a catch of
Sperm Whales is known to continue is at Lamalera,
Lembata (Lomblem on world maps), Indonesia,
where a few animals are taken per year by primitive
methods (Barnes 1991).

Population Sizes and Trends

All available estimates of Sperm Whale popula-
tion sizes are problematic. To obtain unbiased popu-
lation estimates, visual counts from ships or aircraft
must be corrected to account for their dives which
generally last at least 30 min (Papastavrou, Smith,
and Whitehead 1989; Watkins et al. 1993). Reliable
corrections for Sperm Whale dives are not available,
and estimates from visual counts are presented
uncorrected and so have a negative bias. Thus, sever-

THE CANADIAN FIELD-NATURALIST

Vol. 111

al well-conducted visual surveys provide useful min-
imum density estimates for particular ocean areas
(e.g.; Christensen, Haug, and Mien 1992; Wade and
Gerrodette 1993; Davis et al. 1995). Estimates from
visual surveys off North America include 756 (CV
0.49) from ship surveys off California (Barlow
1995), 892 (CV 0.99) from aerial surveys off
California (Forney, Barlow, and Carretta 1995), and
337 (CV 0.50) from aerial surveys of the continental
shelf and slope waters off the northeastern United
States (Blaylock et al. 1995).

Acoustic surveys of Sperm Whales, in which a
count is made of the number of vocalizations or
vocalizing animals heard through towed
hydrophones, have been attempted (e.g.; Leaper,
Chappell, and Gordon 1992; Davis et al. 1995), but
the methodology behind such censuses need further
development. In two cases mark-recapture estimates
of the number of animals visiting a restricted study
area have been made using individual photographic
identifications (Whitehead, Waters, and Lyrholm
1992; Childerhouse, Dawson, and Slooten 1995).
However, the wide ranging behaviour of this
species, and the lack of any clear stock divisions,
means that neither of these estimates refer to a dis-
crete population.

During the 1970s and early 1980s, considerable
effort was put into trying to determine the status of
Sperm Whale populations using data obtained during
whaling, principally catch per unit effort statistics
and changes in length and age distributions. There
are many problems with these estimates, including
the lack of clear stock divisions and the differences
in distribution and ranging behaviour of the different
segments of the population, with the result that none
of these estimates of the sizes of Sperm Whale popu-
lations are credible (Cooke 1986).

A fundamental problem when trying to assess the
status of Sperm Whale populations is determining
the effects of the open-boat hunt, which principally
took place in the 19th Century (Gambell 1983).
Using analyses of whalers’ logbooks, Whitehead
(1995) estimated that Pacific Sperm Whale popula-
tions may have been reduced to about 10 to 30% of
their pre-whaling size by 1860. Only slight recovery
was probably achieved by 1946, when modern whal-
ing for Sperm Whales became intense. As Sperm
Whaling started earlier in the Atlantic (see above),
the Atlantic populations may have been more seri-
ously affected than those in the Pacific (Whitehead
1995).

It is generally agreed that modern commercial
whaling made further substantial reductions in the
populations of adult male Sperm Whales in all
oceans (Gosho, Rice, and Breiwick 1984).
Populations of adult females were also reduced, but
less severely in most cases (Gosho, Rice, and
Breiwick 1984).

ee)

In the North Atlantic, catch-effort series from
Spain indicate that the hunted population of males
was reduced by approximately 80% between 1905
and 1981; that of females by about 50% (Gosho,
Rice, and Breiwick 1984). Data from Iceland and the
Azores, used in a length-specific model, indicate
similar but less extreme declines in those areas dur-
ing the same time-interval (Gosho, Rice, and
Breiwick 1984: their Table 5).

Models of North Pacific Sperm Whale populations
suggest pre-whaling numbers of a few hundred thou-
sand, with modern whaling depleting male numbers
by about 36% and females by 14% (Gosho, Rice,
and Breiwick 1984). However, as noted above, these
estimates have little credibility. There is an addition-
al problem in that the Soviets, who were important
exploiters of North Pacific Sperm Whales, seriously
misreported catches (Yablokov 1994). This was not
public knowledge when the status of North Pacific
Sperm Whale populations was examined in the late
1970s and early 1980s.

Estimates of Sperm Whale numbers from surveys
and photoidentifications suggest a few thousand
individuals inhabiting relatively small parts of the
world’s oceans (e.g.; Christensen, Haug, and Mien
1992; Whitehead, Waters, and Lyrholm 1992; Wade
and Gerrodette 1993) with a preliminary estimate of
some 140000 for the southern oceans, south of 30°S
(Butterworth and Geromont 1996). Thus, as Sperm
Whales continue to be sighted over deep waters
throughout the world, it seems likely that there are a
few hundred thousand alive today. The relationship
between their abundance now, and that in 1700,
before substantial whaling began, is unknown,
although males are probably depleted to a greater
extent than females.

Habitat

Although Sperm Whales can be found over any
deep water, they are most abundant near the edges
of continental shelves, large islands and offshore
banks and over submarine trenches and canyons
where vertical mixing of water masses creates zones
of high biological productivity (Berzin 1971;
Gulland 1974; Jaquet 1996). Females are rarely
found in waters less than 1000 m deep (H.
Whitehead, unpublished data). Males, although also
primarily deep water animals, are in some ocean
areas regularly found in waters of 200 to 1000 m
(Caldwell, Caldwell, and Rice 1966). A prime
example of relatively shallow water habitat of male
Sperm Whales is on the Scotian Shelf where they
were caught, and are seen, in water depths of 40 to
200 m (Mitchell 1975; Whitehead, Brennan, and
Grover 1992). In addition to their stricter habitat
requirements for depth, female Sperm Whales are
also generally restricted to waters with surface tem-
peratures warmer than about 15°C (Rice 1989).

REEVES AND WHITEHEAD: STATUS OF THE SPERM WHALE

299

General Biology
Reproduction

Females begin to ovulate (and usually conceive
for the first time) at seven to 13 years of age when
they are 8.3 to 9.2 m long (Rice 1989). Intervals
between births are usually about four to six years,
although older females have much reduced reproduc-
tive rates (Best, Canham, and Macleod 1984).
Following a gestation period of about 15 to 16
months, a single 4 m calf is born (Best, Canham, and
Macleod 1984). In most areas there is a peak calving
season of about three months during the spring
(Best, Canham, and Macleod 1984). Lactation seems
to last about two years although the duration varies,
and estimates are complicated by both allosuckling —
suckling by females other than the mother — and
occasional suckling of older (up to 13 yr) juveniles
(Best, Canham, and Macleod 1984; Gordon 1987b).

Sexual maturation in males is gradual, beginning
at about seven to 11 yr of age and a length of 8.7 to
10.3 m (Rice 1989). The relative size of the testes,
and density of spermatozoa, increase during the
male’s late 20s (Rice 1989). These physiological
changes are consistent with what is known about
breeding strategies in male Sperm Whales. Males in
their late teens and early twenties are largely segre-
gated from females by latitude. On the prime breed-
ing grounds they would probably generally be out-
competed by older, and much larger, males, so it
pays them to remain at high latitudes, investing in
growth rather than seeking mating opportunities
(Whitehead 1994). However, there is some overlap
in distribution, and young sexually mature males will
attend females whom they encounter, and may
obtain some mating opportunities (Best, Canham,
and Macleod 1984; Whitehead, Brennan, and Grover
1992). Large males, in their late 20s and older,
migrate back to low latitudes to breed (Best 1979).
The duration, frequency and timing of such migra-
tions are unknown.

While on the breeding grounds the large males
rove between groups of females spending a few
hours or so with each (Whitehead 1993). The large
males do fight each other, but rarely (Best 1979;
Kato 1984; Clarke, R. and Paliza 1988; Whitehead
1993). Watkins et al. (1993) suggested that males on
a breeding ground may establish dominance hierar-
chies. This, together with synchronized oestrus
among grouped females (Best and Butterworth
1980), may account for the indications of paternal
relatedness among grouped females shown by genet-
ic analysis (Richard et al. 1996). Mating itself has
not been convincingly described.

Population Structure and Social Behaviour

The fundamental element of the social structure of
the Sperm Whale is the matrilineal family unit: about
12 genetically related females and their offspring

300

who are permanent companions (Best 1979;
Whitehead, Waters and Lyrholm 1991; Richard et al.
1996). Most, if not all, females seem to spend their
lives with their female relatives. The number of indi-
viduals in a family unit is highly variable
(Whitehead, Waters, and Lyrholm 1991). The units
may travel with other units for periods of days as
coordinated groups of about 20 to 25 animals
(Whitehead, Waters, and Lyrholm 1991).

Within groups, it is usually difficult to distinguish
particular family units, so that most descriptions of
social behaviour are at the level of the group. Calves
accompany different adult members of their group
during their mothers’ dives, and groups containing
calves have more staggered dives than those without,
suggesting active babysitting (Gordon 1987b;
Whitehead 1996a). This form of alloparental care
would be in addition to allosuckling (see above), and
might be the most important function of the family
units and groups (Best 1979; Whitehead 1996a).
Sperm Whale calves are vulnerable to predators,
such as Killer Whales, but can be successfully
defended by coordinated actions of members of their
group (Arnbom et al. 1987; Jefferson, Stacey and
Baird 1991).

The cohesive nature of the female society is also
shown by the structured ranks, about 500 m long and
arranged perpendicular to the direction of move-
ment, that groups may form while foraging
(Whitehead 1989), and social gatherings at the sur-
face. For periods of several hours, members of a
group may remain at or close to the surface (Figure
2), lying close to one another, often almost motion-
less (Whitehead and Weilgart 1991). Particular
behaviour, such as rolling and touching, breaches
(see front cover) and vocalizations, such as codas
(see below), are characteristic of such social periods
(Whitehead and Weilgart 1991).

The age at which males leave the family units
seems to be highly variable but may average about
six years (Best 1979; Richard et al. 1996). After dis-
persal the males are found in bachelor schools with
other males of about their own age (Best 1979). As
the males age, they generally live in smaller schools
and at higher latitudes (Best 1979).

The structure of schools of young bachelors has
been little studied. However, by the time males reach
the age of about 20 yr, bachelor schools seem to con-
sist of little more than loose aggregations of similar-
sized whales, probably in the same area because of
food concentrations (Whitehead, Brennan and
Grover 1992: Childerhouse, Dawson and Slooten
1995). Such males rarely cluster close together.
While an underlying social structure may exist, and
is perhaps indicated by multiple strandings of males
of about this age (e.g. Table 1), there is little sign of
it over the spatial and temporal scales used in recent
boat-based studies of living animals (Whitehead,
Brennan, and Grover 1992; Childerhouse, Dawson,

THE CANADIAN FIELD-NATURALIST

Vol. 111

and Slooten 1995). The largest males appear essen-
tially solitary, and some evidence suggests that they
avoid one another on the breeding grounds
(Whitehead 1993).

Aggregations of 50 to as many as several hundred
Sperm Whales are sometimes encountered
(Caldwell, Caldwell, and Rice 1966; Whitehead and
Weilgart 1990). These are temporary gatherings,
probably formed in response to clumped prey.

Diving

A typical Sperm Whale dive cycle consists of a
dive to roughly 400 m below the surface for about
40 min followed by breathing at the surface for
about 8 min, with the whale maintaining a speed of
approximately 4 km/hr throughout (Gordon 1987a;
Papastavrou, Smith and Whitehead 1989;
Whitehead, Brennan, and Grover 1992; Watkins et
al. 1993). However, there is considerable variation.
The longest timed dive was well over two hours
(Watkins, Moore, and Tyack 1985), and the discov-
ery of a fresh bottom-dwelling shark in the stomach
of a Sperm Whale captured in over 3195 m of
water, as well as other data, suggests that the
species is capable of dives of several kilometres
(Clarke 1976).

Feeding

Sperm Whales prey mainly on mesopelagic and
bathypelagic cephalopods of more than 55 species
from 36 genera (Kawakami 1980). Most of these
squid are quite small, weighing a few hundred grams
(Clarke, Martins, and Pascoe 1993). However, they
do take much larger animals at times including the
Giant Squid (Architeuthis sp.) which can approach
the size of the Sperm Whales themselves (Fiscus and
Rice 1974). Large males prey on larger species of
cephalopod and on larger individuals of a given
species than do females and younger males (Best
1979; Clarke 1980; Best, Canham, and Macleod
1984). A variety of large demersal and mesopelagic
fishes is also taken, especially by adult males in high
latitudes, for example in Denmark Strait (Kawakami
1980; Martin and Clarke 1986). Items occasionally
found in Sperm Whale stomachs include large crus-
taceans, parts of seals, stones, and man-made flotsam
and jetsam (Rice 1989). However, there is no evi-
dence that Sperm Whales prey on mammals regular-
ly. Most stomachs examined from Sperm Whales
taken off Vancouver Island during late spring, sum-
mer, and fall contained food, mainly squid
(Moroteuthis robusta and Gonatus sp.), Ragfish
(Acrotus willoughbyi) and Rockfish (Sebastodes sp.)
[Pike and MacAskie 1969].

Clarke, Martins, and Pascoe (1993) suggested that
the cephalopod prey of Sperm Whales fall into two
main classes. Smaller, neutrally buoyant, luminous
animals without much musculature are easily detect-
ed visually and captured; whereas larger, faster, non-

ie

REEVES AND WHITEHEAD: STATUS OF THE SPERM WHALE

301

FIGURE 2. Grouped Sperm Whales at surface, Galapagos Islands, 1985.

luminescent animals must be actively chased using
sonar (see below).

Sperm Whales apparently feed at all times of day,
year-round (Rice 1989). Lockyer (1981) developed
an energy budget for Sperm Whales. She concluded,
judging mainly from an analysis of stomach capaci-
ties and assuming a diet of squid providing 800
kcal/kg, that Sperm Whales weighing less than 15
metric tons consume about 3 percent of their body
mass daily. This rate probably increases to about 3.5
percent for the 30— to 50-ton males.

Acoustics

The principal vocalization of the Sperm Whale is
a loud (source level 171dB re 1uPa), broad-band (ca.
0.1 to 20kHz) click (Worthington and Schevill 1957;
Backus and Schevill 1966; Watkins 1980; Weilgart
1990; Goold and Jones 1995). These clicks are used
in a number of ways.

During deep foraging dives, the clicks are heard in
long trains with inter-click intervals of about 0.8 s
for males and 0.5 s for females (Mullins, Whitehead,
and Weilgart 1988; Whitehead and Weilgart 1990;
Goold and Jones 1995). Most authors (e.g., Norris
and Harvey 1972; Gordon 1987a; Weilgart 1990;
Goold and Jones 1995) believe that these clicks are a
form of echolocation, allowing the whales to detect
prey at ranges of several hundred metres.

During social periods Sperm Whales produce
codas, consisting of from two to about 20 clicks
“arranged into a patterned series (Watkins and Schevill
1977). Codas are often heard as exchanges, and they
seem to function as communication (Watkins and
Schevill 1977; Weilgart and Whitehead 1993).

Other click-based vocalizations include creaks
(click trains with high repetition rates) heard during

both foraging and social periods (Gordon 1987a;
Whitehead and Weilgart 1991), and slow clicks:
loud, reverberant clicks with an inter-click interval
of about 6 s heard principally from mature males on
the breeding grounds (Weilgart and Whitehead
1988). A few, much quieter, non-click vocalizations
are occasionally recorded from Sperm Whales
(Gordon 1987a).

Limiting Factors
Mortality

Estimates of total natural mortality rates for
Sperm Whales vary between 0.05 and 0.09 per year
(Rice 1989). Young Sperm Whales are preyed upon
by Killer Whales and possibly by large sharks (Best,
Canham, and Macleod 1984; Arnbom et al. 1987),
but adult Sperm Whales probably experience almost
no natural predation (Jefferson, Stacey, and Baird
1991). Lambertsen (in International Whaling
Commission 1987) suggested that nematodes can
inflict penetrating ulcers in the stomachs of Sperm
Whales, in rare instances possibly causing death.

Sperm Whales strand, singly or multiply (up to 72
animals), alive or dead throughout the world from
time to time (Rice 1989; Table 1). The causes of the
strandings are generally unclear.

During 1988 and 1989 at least 32 dead male
Sperm Whales were discovered in northern Europe,
about as many as the total reported from the area
during the previous 100 years (Christensen 1990). In
late 1994, another 20 Sperm Whales were found
dead in and around the North Sea. It is not clear what
caused these deaths, although pollutants might have
contributed to the rash of deaths in what is one of the
more polluted parts of the world’s oceans (Joiris
1995). In general, Sperm Whales have pollutant

302

loads somewhere between those of the often highly
contaminated inshore odontocetes, and the much less
affected baleen whales (Aguilar 1983).

Some mortality is caused by entanglement in gill-
nets (Notarbartolo di Sciara 1990; Barlow et al.
1994; IWC 1994) and other fishing gear (Haase and
Félix 1994). Ingestion of marine debris such as plas-
tic buckets and netting could pose serious health
risks to individuals (IWC 1987). Sperm Whales may
be killed or injured by collisions with ships. A
severely injured, but living, Sperm Whale observed
south of Nova Scotia on 10 May 1994, in position
42°12’'N, 65°21’W (J. Conway, Department of
Fisheries and Oceans, Maritime Region, Halifax,
Nova Scotia; personal communication), was likely
hit by a ship.

Disturbance

Sperm Whales seem to be particularly sensitive to
acoustic disturbances within the frequency range of
their own clicks. They consistently interrupted their
sound production in the presence of underwater puls-
es made by pingers or submarine sonar (Watkins and
Schevill 1975; Watkins, Moore, and Tyack 1985),
and deserted an area of abundance after the onset of
seismic testing with air guns (Davis et al. 1995: 244-
249). Sperm Whales sometimes react to close
approaches (<200 m) of a survey vessel by abrupt
dives (Davis et al. 1995: 232-233). High-speed
whalewatching vessels affected the behaviour of
male Sperm Whales off Kaikoura, New Zealand,
especially when the boats were not handled cautious-
ly (Gordon et al. 1992).

Climatic/Oceanographic Factors

Off the Galapagos Islands, Sperm Whales have
greatly reduced feeding success when sea tempera-
tures are warmer than average, as in El Nifio years
(Whitehead, Papastavrou, and Smith 1989). In such
conditions, their residence periods near the Islands
are shorter, and their movements more directed
(Whitehead 1996b). If these effects are general, then
global warming could have a serious impact on
Sperm Whale populations.

Residual Effects of Whaling

We do not know how populations of Sperm
Whales have been affected by past whaling.
However there is concern that the effects may persist
longer than would normally be expected for an
exploited population.

The social organization and segregation of the
sexes of this species have led to a consideration of
the possibility that a reduction in the ratio of socially
mature males to females, caused by selective whaling
for bulls, could result in a decrease in the rate of con-
ception of females (e.g., Mitchell 1977; IWC 1980;
May and Beddington 1980; Whitehead 1987). There
is evidence that this happened in the southeast Pacific
(Clarke, Aguayo, and Paliza 1980). A particularly

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Vol. 111

low calving rate off the Galapagos Islands during the
past ten years may be related to the small number of
mature males in the area (Whitehead 1990). The
Galapagos are close to the Peruvian whaling grounds
where males were especially heavily hunted in the
1970s and early 1980s (Ramirez 1989).

Special Significance of the Species

A crude guess at the total amount of cephalopods
consumed annually by the world Sperm Whale pop-
ulation is 110 to 320 million metric tons (Clarke
1977), roughly comparable with the total world-wide
catch of all marine species. A high proportion of the
Sperm Whale’s diet consists of ammoniacal squid
that are unacceptable as human food. Moreover, the
direct harvesting by humans of most of the Sperm
Whale’s prey species is probably impractical. As pri-
marily a mesopelagic macroteuthophage, the Sperm
Whale probably has little direct competition for food
resources from other marine mammals or humans.
Its closest cetacean competitors are ziphiid whales
that feed in the same depth zones but generally prey
on smaller squids (Rice 1989). Non-cetacean poten-
tial competitors for food are Elephant Seals
(Mirounga spp.) and some of the larger squids (e.g.,
Dosidicus gigas) that are themselves preyed upon by
Sperm Whales.

Although the meat of Sperm Whales is not consid-
ered palatable in much of the world (Berzin 1971), it
is eaten in parts of Japan (Freeman 1988), eastern
Indonesia (Barnes 1991), and the Lesser Antilles
(Price 1985), despite warnings that it may contain
unsafe amounts of organic mercury (Beary 1979).
Other valuable products obtained from Sperm
Whales include: spermaceti, a waxy substance from
the whale’s head that has been widely used to make
candles and cosmetics (e.g., cold creams); sperm oil,
used as a temperature- and pressure-resistant lubri-
cant and industrial wax; ambergris, a concretion
formed in the large intestine of the Sperm Whale,
used as a fixative in perfumes; and ivory teeth, used
for carving (Berzin 1971). Sperm oil was generally
more valuable than that obtained from mysticete
whales because of its alcohol content; the alcohols
were used for manufacturing many kinds of deter-
gent (Bailey, Carter, and Swain 1952). Sperm Whale
liver is rich in vitamin A (Bailey, Carter, and Swain
1952). Synthetic or natural substitutes are now avail-
able for most Sperm Whale products.

Since 1988, watching living Sperm Whales has
become an important activity in a few parts of the
world. The most successful operation is in Kaikoura,
New Zealand, but whale-watch tours primarily
directed at Sperm Whales are also run regularly from
Dominica in the West Indies, Andenes in Norway,
the Azores, and Japan (Hoyt 1995). In other parts of
the world, including the Galapagos and Antarctica,
Sperm Whales are among the attractions of more

1997

general nature-tourism operations. A number of
other sites with deep water close to shore, including
Sri Lanka and some Caribbean islands, have consid-
erable potential for the development of whale-watch-
ing based on Sperm Whales (Hoyt 1995). Canada,
with its broad continental shelves keeping the Sperm
Whales generally away from the coasts, appears to
have little potential for whale-watching based princi-
pally on Sperm Whales. However, in the last few
years the Sperm Whales appearing off Tadoussac,
Quebec (see above), have been an attractive but spo-
radic target of the local whale-watching tour opera-
tors, who focus mainly on rorquals (Michaud, per-
sonal communication).

The Sperm Whale is a creature of extremes. It has
the largest brain on Earth and (probably) the most
powerful natural sonar system, as well as the greatest
geographical separation of the sexes. In their sexual
dimorphism, dispersion, biomass, diving ability, and
longevity Sperm Whales are at, or close to, the maxi-
ma of all mammals. No animal on Earth is much like
the Sperm Whale anatomically or ecologically, and
its unusual social system is only paralleled closely
among the elephants (Weilgart, Whitehead, and
Payne 1996). Combinations of these and other
attributes have given the Sperm Whale particular
significance in a range of areas of human activity,
including aboriginal cultures [such as the New
Zealand Maoris (Anonymous 1996)], commerce
(Starbuck 1878) and literature (Melville 1851).

Evaluation

Sperm Whale populations were likely well below
pre-whaling levels in 1946 when modern whaling for
them became especially intense, were reduced sub-
stantially between 1950 and 1980 (see above), and
can only recover very slowly (see below). Thus the
current world population is certainly much smaller
than it was before the start of whaling for Sperm
Whales in about 1710. However, recent abundance
estimates and sighting rates in different ocean areas
suggest a world population of more than 100 000
(see above).

Whatever the current level, it will not rise fast.
The most recent estimates of Sperm Whale popula-
tion parameters by the Scientific Committee of the
International Whaling Commission (IWC 1982b)
suggest a maximal rate of increase of about 0.9% per
year. Residual effects of whaling (see above), and
other factors, may slow even this low rate, leading to
very long recovery times.

The lack of any clear stock structure means that
the effects of human disturbance, whether from
whaling or pollution or other sources, will not be
contained within a clearly defined subset of the
aggregate population. However, this attribute also
promotes resilience at smaller spatial scales: if
Sperm Whales are removed from an area, but the

REEVES AND WHITEHEAD: STATUS OF THE SPERM WHALE

303

world population is healthy, then it is likely that
recolonization will take place before too long.

Probably the greatest current threat to Sperm
Whale populations is chemical pollution. If toxic
loads continue to accumulate in their prey, so that
levels in individual animals rise, and this affects
reproduction, mortality or social behaviour, then a
putative population increase of less than 1% per year
could easily become a population decrease. Acoustic
pollution couid be an additional detrimental factor.
The bycatch of Sperm Whales in fishing gear is sub-
stantial and regular in some areas. This means that
the species is being “exploited”, indirectly, in many
more areas and at a greater scale than suggested by
catch statistics alone. As whaling for Sperm Whales
has declined, the use of fishing gear that is hazardous
for cetaceans, especially gillnets, has expanded
(IWC 1994). A third area of threat to Sperm Whale
populations is the possibility of breakdown in the
current international framework for the management
and protection of whale populations (through the
International Whaling Commission) and resultant
widespread, unregulated whaling.

Considering the enormous range of the species, its
aggregate numerical abundance, and its virtually
complete protection from commercial exploitation at
present, the Sperm Whale is not endangered.
Although vulnerable (principally because of its very
slow potential rate of increase which could easily be
reversed) and threatened by pollution and other fac-
tors, the threats are probably not sufficiently imme-
diate or clear that a COSEWIC designation would be
appropriate or useful at this time. If large-scale com-
mercial whaling should resume, or trends of
increased pollutant levels or mortality become appar-
ent, then the status of the Sperm Whale should be re-
evaluated by COSEWIC.

Canada would improve the prospects of the
Sperm Whale, as well as other marine life, by any
actions which reduce marine pollution or support
international agreements regulating the exploitation
of marine resources, such as the International
Whaling Commission.

Acknowledgments

This review was funded by the Department of
Fisheries and Oceans and World Wildlife Fund
Canada through a grant to Okapi Wildlife
Associates, Hudson, Quebec. We thank Robin Baird
and Heather Patterson (British Columbia), Pierre-
Yves Daoust (Prince Edward Island), Mary Dillon
and Zoe Lucas (Nova Scotia), Sean Todd
(Newfoundland/Labrador), Robert Michaud
(Quebec) and Donald McAlpine (New Brunswick)
for information on strandings, and Jerry Conway
(Department of Fisheries and Oceans, Scotia-Fundy
Region), Robert Michaud and Richard Sears for
sightings.

304

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Weilgart, L., and H. Whitehead. 1993. Coda vocaliza-
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744-752.

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Whitehead, H. 1987. Social organization of sperm whales
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Whitehead, H. 1989. Formations of foraging sperm
whales, Physeter macrocephalus, off the Galapagos
Islands. Canadian Journal of Zoology 67: 2131-2139.

Whitehead, H. 1990. Assessing sperm whale populations
using natural markings: recent progress. Reports of the
International Whaling Commission (special issue) 12:
377-382.

Whitehead, H. 1993. The behaviour of mature male
sperm whales on the Galapagos breeding grounds.
Canadian Journal of Zoology 71: 689-699.

Whitehead, H. 1994. Delayed competitive breeding in
roving males. Journal of Theoretical Biology 166:
127-133.

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Whitehead, H., S. Brennan, and D. Grover. 1992.
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Accepted 8 July 1996

Notes

First Breeding Record of Red-breasted Merganser, Mergus serrator,
on Axel Heiberg Island, Northwest Territories

THEO HOFMANN!, JOHN W. CHARDINE2, and HANS BLOKPOEL?

!Department of Biochemistry, University of Toronto, Toronto, Ontario MSS 1A8
2Canadian Wildlife Service, P.O. Box 21276, St. John’s, Newfoundland AlA 5B2
3Canadian Wildlife Service, 49 Camelot Drive, Nepean, Ontario K1A OH3

Hofmann, Theo, John Chardine, and Hans Blokpoel. 1997. First breeding record of Red-breasted Merganser, Mergus ser-
rator, on Axel Heiberg Island, Northwest Territories. Canadian Field-Naturalist 111(2): 308-309.

A female Red-breasted Merganser with six young was observed on the river flowing from Buchanan Lake into Mokka
Fjord on Axel Heiberg Island, Northwest Territories, on 9 August 1993. This is some 900 km north of the northern edge of

the known breeding range in eastern Canada for this species.

Key Words: Red-breasted Merganser, Mergus serrator, breeding record, Axel Heiberg Island, Northwest Territories.

The Red-breasted Merganser (Mergus serrator)
breeds extensively throughout the boreal regions of
Canada. The northern limit of its general breeding
range extends from about 67°N on Baffin Island
through 65°N in western Hudson Bay to 70°N in the
western Arctic (Godfrey 1986). In addition, breed-
ing has been recorded on Utuk Lake (72°30'N)
(Godfrey 1986) and on two occasions near Pond
Inlet, 72°25’N (Renaud et al. 1981) and sightings of
single birds have been made in the northern part of
Baffin Island at Oliver Sound, 72°N, 78°W, and
Arctic Bay 73°N, 85°W (Godfrey 1986). In Alaska
the northernmost breeding sites are near
Wainwright, 70°N, 160°W (Bellrose 1976). On the
west coast of Greenland the known breeding range
extends to about 74°N (Bellrose 1976). Also in
Greenland, a female Red-breasted Merganser was
seen on 2 July 1979 at Constable Bugt (84°N,
32°W), near the most northerly point of land in the
world (Hjort 1982). In Europe the breeding range
extends to Nord Kapp (71°N, 25°E) on Magergya,
the northernmost island of Norway (Harrison 1982;
Flint et al. 1984) and in European Russia to the
northwest corner of the South Island of Novaya
Zemlya at 74°N, 54°E. In Asian Russia the north-
ernmost breeding area is in the Taymyr area around
75°N, 105°W (Flint et al. 1984).

On 9 August 1993, we saw a female Red-breasted
Merganser with six young, estimated age 15 and 25
days, at Mokka Fjord (79°30.0'N, 87°23.9’W) on
Axel Heiberg Island, Northwest Territories, near the
outlet of the river flowing from Buchanan Lake into
the fjord. The habitat at Mokka Fjord consisted
mainly of shingle with scattered tundra vegetation.

The riverbed, flood plain and edges of the river were
largely devoid of vegetation. This agrees well with
the general habitat for Red-breasted Merganser in
the Arctic described by Snyder (1957). This breeding
record of Red-breasted Merganser seems to be the
most northerly record reported. Furthermore, as a
sighting it is the second most northerly one in the
world after the sighting of a single female at the
north end of Greenland (Hjort 1982).

What are the chances that the ducklings were able
to leave the area before freeze-up? Based on the
estimated age of the brood and a fledging period of
at least 59 days (Gooders and Boyer 1986), we pre-
dict a fledging date around mid-September. At that
time freeze-up of freshwater lakes and rivers would
normally have begun because average maximum
and minimum daily temperatures in the area are
well below 0°C (Anonymous 1984). Thus, to be
successful, the brood would have had to move out
of the lake and river system into the marine waters
of Mokka Fjord well before this time. This is plausi-
ble because Red-breasted Mergansers are the most
marine of the merganser ducks and are known to
move their broods from freshwater into marine areas
during chick development (A. J. Erskine, personal
communication). According to the weekly ice com-
posite charts published by Environment Canada
(Anonymous 1993), ice coverage of marine waters
in the area of Mokka Fjord was only 30% on 26
August 1993, 50% on 9 September 1993, stayed
around 90% until 30 September 1993, and reached
100% only by 7 October 1993. This would have
given the brood sufficient time to fledge and fly
south before complete freeze-up.

308

1O97

Acknowledgments

H. B. and J. C. thank S. Blyth, Blyth and Co.
Travel Inc., and D. Wells of Marine Expeditions for
having them aboard the Kapitan Khlebnikov as
resource persons while giving them the opportunity
to make observations for the Canadian Wildlife
Service on the pelagic distribution of sea-birds. We
also thank T. Lang, Department of Zoology,
University of Toronto, for information on age
determination of young ducks and the staff of the
Royal Ontario Museum library for their help with
the literature search. A. J. Erskine, A. J. Gaston, W.
E. Godfrey and A. Reed commented on an earlier
version.

Literature Cited

Anonymous. 1984. Principal Station Data: A Summary
of Hourly Weather Observations, Climate Normals
and Extremes for Canadian Principal Climate Stations.
Eureka. Report PSD/DSP 79, Atmospheric Environment
Service, 48 pages. Supply and Services Canada,
Ottawa.

Anonymous. 1993. Weekly Ice Composite Charts, East-
ern Canada. Northern Arctic Ice Centre, Environment
Canada, Ottawa.

NOTES

309

Bellrose, F. C. 1976. Ducks, Geese and Swans of North
America. Stackpole Books, Harrisburg, Pennsylvania.
Flint, V. E., R. L. Boehme, Y. V. Kostin, and A. A.
Kusnetzov. 1984. A Field Guide to Birds of the USSR,
including Eastern Europe and Central Asia. (English
Translation) Princeton University Press, Princeton, New

Jersey.

Godfrey, W. E. 1986. The Birds of Canada, Revised
Edition. National Museum of Natural Sciences, Ottawa,
Ontario.

Gooders, J., and T. Boyer. 1986. Ducks of North Amer-
ica and the Northern Hemisphere, Facts on File Inc.,
New York.

Harrison, C. 1982. An Atlas of the Birds of the Western
Palearctic. William Collins Sons & Co. Ltd., London,
United Kingdom.

Hjort, C. 1982. Red-breasted Merganser, Golden Plover
and Fieldfare in Eastern North Greenland 1979. Dansk
Ornitologisk Forening Tidsskrift 76: 71 [in Danish].

Renaud, W. E., W. G. Johnston, and K. J. Finley. 1981.
The avifauna of the Pond Inlet region, N.W.T. American
Birds 35: 119-124.

Snyder, L. L. 1957. Arctic Birds of Canada. University of
Toronto Press, Toronto, Ontario.

Received 5 October 1995
Accepted 11 August 1996

Sagina (Caryophyllaceae) Range Extensions in Canada:
S. japonica New to Newfoundland; S. procumbens New to the

Northwest Territories

RICHARD K. RABELER! and JOHN W. THIERET

‘Herbarium, North University Building, University of Michigan, Ann Arbor, Michigan 48104
"Herbarium, Department of Biological Sciences, Northern Kentucky University, Highland Heights, Kentucky 41099

Rabeler, Richard K., and John W. Thieret. 1997. Sagina (Caryophyllaceae) range extensions in Canada: S. japonica new to
Newfoundland; S. procumbens new to the Northwest Territories. Canadian Field-Naturalist 111(2): 309-310.

Sagina japonica is reported as new to Newfoundland from St John’s in 1992; S. procumbens is reported as new to the

Northwest Territories from Yellowknife in 1994.

Key Words: Sagina japonica, Sagina procumbens, Caryophyllaceae, Newfoundland, Northwest Territories

Except for Sagina nodosa, with its relatively
showy flowers (petals 3-4.5 mm long, about twice
as long as sepals), the Canadian representatives of
Sagina (Caryophyllaceae) are easily overlooked,
weedy plants with inconspicuous flowers (petals to
2.5 mm long, shorter than but slightly surpassing
the sepals). We report here Canadian range exten-
sions of two species of this genus: Sagina japonica
and S. procumbens.

Sagina japonica is an addition to the flora of
Newfoundland. Collection data are: NEWFOUND-
LAND: weed in downtown St. John’s, Thieret 58108,
19 August 1992 (KNK, DAO). It was collected from

a crack between a sidewalk and the base of a build-
ing along Water Street. Many of the ca. 25 plants in
the group were enmeshed in a mass of black threads.
Nearby grew several plants of Erucastrum gallicum
(Brassicaceae). The source of the seeds that estab-
lished these urban weeds is unknown. The few
Canadian occurrences of S. japonica were summa-
rized by Crow (1978) as Nanaimo and Prince
Rupert, British Columbia; and Ottawa, Ontario. No
additional Canadian collections are known to us.
Scoggan (1978) did not include the species. Crow
(1978) reported the species only from Oregon in the
United States; Mitchell and Tucker (1991) cited

310

collections of this “overlooked adventive” also
from Connecticut, Massachusetts, New York, and
Pennsylvania. The species is almost certainly more
widespread in North America than present herbari-
um records indicate.

The Newfoundland vascular plant atlas (Rouleau
and Lamoureux 1992) reported two species of
Sagina (S. nodosa and S. procumbens) from that
province. Two additional species — S. caespitosa
and S. saginoides — were added to the flora by Hay
et al. (1994). With the discovery of S. japonica, the
genus is now represented in Newfoundland by five
species.

Sagina japonica is an annual characterized by its
5-merous flowers, its dark brown tuberculate or peb-
bled seeds lacking a dorsal groove, and its glandular
pubescent stems, pedicels, and calyces. The St.
John’s plants were noticeably viscous.

Sagina procumbens is an addition to the flora of
the Northwest Territories. Collection data are:
NORTHWEST TERRITORIES: garden weed, Yellow-
knife, 19 July 1994, Hils & Thieret 1504 (KNK,
DAO). It was collected from the garden area planted
with native species in front of the Tourist
Information Bureau in Yellowknife. As with S.
Japonica in St. John’s, there were about 25 individu-
als; their seed source, too, is unknown. In eastern
Canada the species occurs in Quebec and all the
Maritime Provinces; in the west, in British Columbia

THE CANADIAN FIELD-NATURALIST

Vol. 111

(Crow 1978; Scoggan 1978). Crow (1978) recorded
an Alaskan Panhandle collection that approaches the
Yukon.

Sagina procumbens is a perennial spreading by
offshoots and is characterized by its procumbent to
ascending stems and its primarily 4-merous flowers
with white-margined sepals.

Acknowledgment
We thank William J. Cody for checking Canadian
records of Sagina japonica and S. procumbens for us.

Literature Cited

Crow, G. E. 1978. A taxonomic revision of Sagina
(Caryophyllaceae) in North America. Rhodora 80: 1-91.

Hay, S. G., A. Bouchard, and L. Brouillet. 1994.
Additions to the flora of Newfoundland. III. Rhodora 96:
195-203.

Mitchell, R. S., and G. C. Tucker. 1991. Sagina japonica
(Sw.) Ohwi (Caryophyllaceae), an overlooked adventive
in the northeastern United States. Rhodora 93: 192-194.

Rouleau, E., and G. Lamoureux. 1992. Atlas of the vascu-
lar plants of the island of Newfoundland and of the
islands of Saint-Pierre-et-Miquelon. Fleurbec, Saint-
Henri-de-Lévis, Quebéc.

Scoggan, H. J. 1978. The Flora of Canada. Part 3 —
Dicotyledoneae (Saururaceae to Violaceae). National
Museum of Natural Sciences Publications in Botany,
Number 7: 545-1115.

Received 3 May 1996
Accepted 21 October 1996

Use of Active Beaver, Castor canadensis, Lodges by Muskrats,

Ondatra zibethicus, in Wyoming

Mark C. McKinstry, Rory R. KARHU, and STANLEY H. ANDERSON

Wyoming Cooperative Fish and Wildlife Research Unit, Box 3166, Laramie, Wyoming 82071.

McKinstry, Mark C., Rory R. Karhu, and Stanley H. Anderson. 1997. Use of active Beaver, Castor canadensis, lodges by
Muskrats, Ondatra zibethicus, in Wyoming. Canadian Field-Naturalist 111(2): 310-311.

Three Muskrats (Ondatra zibethicus) were observed using active Beaver (Castor canadensis) lodges in Wyoming.
Muskrats were observed carrying cut vegetation into active lodge entrances and one Muskrat was trapped in a run as it was
leaving the lodge. Muskrats may use Beaver lodges in an attempt to decrease predation and alleviate extreme environmen-

tal conditions.

Key Words: Beaver, Castor canadensis, denning, Muskrats, Ondatra zibethicus, predation.

Lodge use by Beaver is common in northern cli-
mates and is thought to limit predation and alleviate
extreme environmental temperatures (Buech et al.
1989). Rutherford (1964) hypothesized that preda-
tion pressure directly influences both the size and
quality of dams and lodges. The average lodge has
more than one plunge hole with a chamber located in
the center near the water’s surface for feeding, rest-
ing, or sleeping (Dieter and McCabe 1989).

Buech et al. (1989) examined lodge temperatures
in Beaver of northern Minnesota and found that tem-
peratures were within the thermoneutral zone for
Beavers year-round. Microclimatic air temperatures
in various Muskrat structures (lodges, burrows, and
push-ups) were close to the thermoneutral zone for
single, adult Muskrats (10° to 25° C) (MacArthur and
Aleksiuk 1979). Other non-lodge-building mammals
may use these lodges to reduce exposure to both

1997

environmental factors and predation (Lovallo et al.
1993). Use of inactive Beaver (Castor canadensis)
lodges has been recorded in Bobcats (Felis rufus)
(Lovallo et al. 1993) and is thought to occur in Black
Bears (Ursus americanus) (Ronald Grogan, 1996,
Wyoming Cooperative Fish and Wildlife Research
Unit, personal communication), although it has never
been documented. Abandoned Beaver lodges or bank
dens would certainly be attractive to semi-aquatic
mammals and would be accessible to terrestrial ani-
mals if the entrance was not flooded.

During a reintroduction project in spring and fall
of 1994 and 1995, we live-trapped Beavers through-
out Wyoming, concentrating on small creeks and
rivers. While Muskrats were frequently observed in
Beaver ponds, and were incidentally trapped in
snares set for Beaver on dams, on three different
occasions we observed Muskrats apparently using
lodges occupied by Beaver. Two Muskrats were
observed swimming in and out of separate occupied
Beaver lodges (NE1/4, SW1/4, Sec 20. T. 19 N. R.
78 W., Carbon County, Wyoming) and we trapped a
third Muskrat at an underwater lodge entrance
(NE1/4, SE1/4, Sec 20. T. 18 N. R80 W., Carbon
County, Wyoming) in a snare set for Beaver. Both
Muskrats observed using lodge entrances had cut
vegetation in their mouths and one of them entered
the lodge via a 1-m-wide canal created by Beavers.
The one Muskrat trapped at the lodge entrance was
dead but it appeared that it had been trapped as it
was leaving the lodge. On all three occasions
Beavers were live-trapped at the same entrances
where Muskrats were observed or trapped, indicating
that the lodges were actually used by both species.

While Muskrats typically construct huts or houses
out of available vegetation (Proulx and Gilbert
1984), river dwelling Muskrats almost exclusively
construct bank dens (Fuller 1951; Boutin and
Birkenholz 1987). We could not determine what part
of the Beaver lodge Muskrats were using. However,
because Beaver are aggressive (Hill 1982) and do
not tolerate invading Beaver from adjoining
colonies, we suspect that Muskrats would excavate
their own living quarters.

In creeks and rivers, Beaver lodges, which are
more stable and long-lasting than Muskrat houses
(Chamberlain 1951), could be used by Muskrats for
protection from the weather and predators. Perry
(1982) suggested that Muskrats would use aban-
doned shelters of other animals but we found no ref-
erence to this behavior in our literature review.

Acknowledgments
We thank the Wyoming Game and Fish
Department, U.S. Fish and Wildlife Service, Jack’s

NOTES

oid

Plastic Welding, Ducks Unlimited, National Rifle
Association, Rocky Mountain Elk Foundation,
National Fish and Wildlife Foundation, and the
George E. Menkens Memorial Scholarship for fund-
ing this research. Numerous landowners provided
unlimited access for Beaver trapping efforts. F.
Lindzey, C. Martinez del Rio, and three anonymous
reviewers provided helpful comments on previous
drafts of the manuscript.

Literature Cited

Boutin, S., and D. E. Birkenholz. 1987. Muskrat and
round-tailed muskrat. Pages 314-325 in Wild furbearer
management and conservation in North America. Edited
by M. Novak, J. A. Baker, M. E. Obbard, and B.
Maloch. Ontario Trappers Association, North Bay,
Ontario.

Buech, R. R., D. J. Rugg, and N. L. Miller. 1989.
Temperature in beaver lodges and bank dens in a near-
boreal environment. Canadian Journal of Zoology 67:
1061-1066.

Chamberlain, J. L. 1951. The life history and manage-
ment of the muskrat on Great Meadows Refuge. M.S.
thesis, University of Massachusetts, Amherst. 68
pages.

Dieter, C. D., and T. R. McCabe. 1989. Factors influenc-
ing beaver lodge-site selection on a prairie river.
American Midland Naturalist 122: 408-411.

Fuller, W. A. 1951. Natural history and economic impor-
tance of the muskrat in the Athabasca-Peace Delta,
Wood Buffalo Park. Canadian Wildlife Service. Wildlife
Management Bulletin Serial 1, Number 2. 98 pages.

Hill, E. P. 1982. Beaver. Pages 256-281 in Wild mam-
mals of North America: biology, management, and eco-
nomics.. Edited by J. A. Chapman, and G.. A.
Feldhamer. Johns Hopkins University Press, Baltimore,
Maryland.

Lovallo, M. J., J. H. Gilbert, and T. M. Gehring. 1993.
Bobcat, Felis rufus, dens in an abandoned beaver,
Castor canadensis, lodge. Canadian Field-Naturalist
107(1): 108-109.

MacArthur, R. A., and M. Aleksiuk. 1979. Seasonal
microenvironments of the muskrat (Ondatra zibethicus)
in a northern marsh. Journal of Mammalogy 60:
146-154.

Perry, R. H., Jr. 1982. Muskrats. Pages 282-325 in Wild
mammals of North America: biology, management, and
economics. Edited by J. A. Chapman, and G. A.
Feldhamer. Johns Hopkins University Press, Baltimore,
Maryland.

Proulx, G., and F. F. Gilbert. 1984. Determining muskrat
population trends by house counts. Journal of Wildlife
Management 48: 917-922.

Rutherford, W. H. 1964. The beaver in Colorado: Its
biology, ecology, management and economics. Colorado
Department of Game and Fish, Game Research Division
Technical Bulletin 17, Denver. 49 pages.

Received 8 May 1996
Accepted 10 October 1996

312 THE CANADIAN FIELD-NATURALIST Vol 74

Fall - Early Winter Home Ranges, Movements, and Den Use
of Male Mink, Mustela vison in Eastern Tennessee

RICHARD T. STEVENS!*, Tom L. ASHwoop?, and JONATHAN M. SLEEMAN?

'Department of Biology, Tennessee Technological University, Cookeville, Tennessee 38505

"Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee 37831

’Department of Comparative Medicine, The University of Tennessee College of Veterinary Medicine, Knoxville
Tennessee 37901

Present address: Department of Biology, University of Memphis, Memphis, Tennessee 38152

Stevens, Richard T., Tom L. Ashwood, and Jonathan M. Sleeman. 1997. Fall - early winter home ranges, movements, and
den use of male Mink, Mustela vison, in eastern Tennessee. Canadian Field-Naturalist 111(2): 312-314.

Three male stream-dwelling Mink (Mustela vison), one juvenile and two adults, were radiotracked on 97 occasions during
fall and early winter in eastern Tennessee. Home range estimates ranged between 5663 m and 11083 m and average home
range was the largest reported for the species. Mink made overnight movements of up to 4300 m within their home ranges,
and used between 8 and 24 dens within their home ranges. Bank burrows, such as abandoned Woodchuck (Marmota
monax) and Muskrat (Ondatra zibethicus) burrows and cavities within exposed tree roots, were the most often used den
type (57% of dens). Male Mink home ranges and movements are potentially influenced by feeding requirements, access to

females, and territoriality, but data on the extent of influence by each of these factors is limited at present.

Key Words: American Mink, Mustela vison, home ranges, movements, radiotelemetry, Tennessee.

Although Mink (Mustela vison) are found
throughout most of North America north of Mexico
(Linscombe et al. 1982), and are economically
important furbearers (Spencer 1990), they have been
the subject of relatively few ecological investiga-
tions. Radiotelemetry has been used to study Mink in
Europe, (Gerell,1970;. Linn and Birks, 1931;
Dunstone and Birks 1983) and the western United
States (Whitman 1981; Arnold and Fritzell 1987),
but there have been no published studies of Mink
home ranges and movements in eastern North
America. The objective of this study was to deter-
mine the population status, home ranges, move-
ments, and den use of Mink in eastern Tennessee.

Methods

This study was conducted along the three largest
streams (East Fork Poplar Creek, Bear Creek, and
White Oak Creek) originating on the U. S.
Department of Energy Oak Ridge Reservation
(35°56’ N 84°18’ W), in Oak Ridge, Tennessee.
Land surrounding these streams was primarily
mature Oak (Quercus spp.) and Hickory (Carya
spp.) forest (Kitchings and Mann 1976).

Mink were captured with two door 15 X 15 X 61
cm or 23 X 23 X 81 cm Tomahawk live traps
(Tomahawk Live Trap Co., Tomahawk, Wisconsin)
using barrier tunnels (Eagle and Sargeant 1985).
Captured Mink were transported to The University
of Tennessee College of Veterinary Medicine in
Knoxville, Tennessee. Mink were anesthetized and
intraperitoneal radiotransmitters (Model #IMP/200/L,
Telonics, Inc., Mesa, Arizona) were implanted under
aseptic conditions. Venous blood, ectoparasite, and
fecal samples were collected, sex — age class were
determined (Whitman 1981), and standard mammal

body measurements were taken. All methods used
in this study were approved by the Oak Ridge
National Laboratory and The University of
Tennessee Animal Care and Use Committees.

One daily diurnal radiolocation was obtained for
each Mink using the homing technique (Mech 1983).
Fixed-wing aircraft equipped with directional anten-
nae were occasionally used when Mink could not be
located from the ground. Home ranges were calculat-
ed as meters of waterway used by Mink. Because
Mink typically restricted movements to narrow ripar-
ian zones (Gerell 1970; Whitman 1981), reported
home ranges were linear. Typical methods of calcu-
lating home ranges as polygons would overestimate
home range sizes (Whitman 1981) and would also be
less ecologically meaningful. Movements were cal-
culated as Minimum Nightly Movements (MNMs),
the meters of waterway between two successive
diurnal locations (Arnold and Fritzell 1987).
Tributaries that were not known to be utilized during
the 24h period were not included in MNMs. It was
not always possible to calculate MNMs, as occasion-
ally Mink could not be located on consecutive days.

Results

Four male Mink, one juvenile and three adults,
were captured in 6073 trap nights. Three of these
male Mink, all captured in October or November,
were successfully radiotracked.

Home ranges averaged 7519 m and ranged between
5663 m and 11038 m (Table 1). Mink often used
tributaries of the main stream, and were found in trib-
utaries in 54% of radiolocations. Tributaries contribut-
ed between 13% and 43% of total home ranges.

Overnight movements of Mink (MNMs) were
variable and ranged from 0 to 4300 m. The longest

1997

NOTES

313

TABLE 1. Home ranges (HR) of three male Mink in eastern Tennessee.

Period Mass at
Mink Radiotracked Capture (g)
M1 4 November 1993-26 January 1994 660
M3 8 October 1994—28 October 1994 900

M4 2 November 1994-2 February 1995 = 925

Mean
Standard Error

MNM of each Mink was 1446 m, 4000 m, and 4300
m. Mink were found in the same den as the previous
day in 22% to 50% of radiolocations. Mink were not
monitored at night, and it was likely they left dens to
forage and simply returned to the same den.

Mink used many dens within their home ranges
(range = 8—24 dens). Mink used certain dens on > |
occasion and all used certain dens for periods > 3
consecutive days. The longest consecutive period of
use for a single den was 8 days. In 51-53% of radi-
olocations Mink were found in a previously
unrecorded den. Mink most often (57% of dens)
denned in bank tunnels, primarily either abandoned
Muskrat (Ondatra zibethicus) or Woodchuck
(Marmota monax) dens or holes between exposed
tree roots. Other frequently used dens included slash
piles and dense shrub entanglements (13% of dens)
and culverts (8% of dens). All identified dens were
within approximately 25 m of either the main stream
channel or a tributary of the main stream. Most
(78%) identified dens were on the stream or tributary
bank, within 2 m of the water.

Discussion

Although comparisons with previously reported
home ranges must be made with caution due to sam-
ple size, male Mink in this study had the largest aver-
age home range reported. Introduced male Mink were
found to have an average home range of 2630 m in
Sweden (Gerell 1970), and 2530 m (Dunstone and
Birks 1983) and 4480 m (Linn and Birks 1981) in
separate studies in Great Britain. The only
radiotelemetry study of stream-dwelling Mink in
North America was conducted by Whitman (1981),
who reported an average home range size of 5250 m.
This was smaller than our home range estimate; how-
ever, he did not include tributary movements in his
estimate, and thus it may have underestimated home
range size. Mink in this study had an average of 4948
m of main stream channel in their home ranges,
which is similar to the results of Whitman (1981).

Home range estimates reported here were pre-
breeding season home ranges. Mink begin breeding
in late February or early March (Enders 1952), and
the latest date a Mink was radiotracked in this study
was 17 February (Table 1). Mink in Sweden (Gerell

Number of Main Stream Tributary Total
Locations Channelin HR (m) in HR (m) HR (m)
35 5109 748 5857
15 3205 2458 5663
47 6531 4507 11038
4948 25/1 TS19
964 1087 1786

1970), North Dakota (Eagle and Sargeant 1985), and
Manitoba (Arnold and Fritzell 1987) extended their
home ranges significantly during the breeding season
because of movements in search of females.
Therefore, if radiotracking had continued into the
breeding season, home ranges may have been signif-
icantly larger.

Mink have relatively large home ranges for carni-
vores (Arnold and Fritzell 1987). Spacing in solitary
males is thought to be an evolutionary compromise
between competing demands for food and females
(Sandell 1989). Other male mustelids, such as
Martens (Martes americana), were thought to main-
tain relatively large home ranges to insure access to
one or more females (Katnik et al. 1994). However,
because Mink leave their pre-breeding season home
ranges to search for females during the breeding sea-
son, home ranges during the non-breeding season
should be driven primarily by feeding requirements.
Gerell (1970) reported that male Mink in Sweden
were territorial and that territoriality was a major
determinant of home range size. Whitman (1981),
however, found no evidence of territoriality in Mink
in Idaho, and even found Mink of the same sex den-
ning within meters of each other. As no two Mink
inhabiting the same stream were simultaneously
radiotracked in this study, no data on territoriality
was collected.

Presence of suitable den sites can be an impor-
tant factor in the distribution and habitat utilization
of Mink (Gerell 1970; Linscombe et al. 1982).
Bank tunnels, such as abandoned muskrat burrows
and natural cavities among exposed tree roots were
the most often used den sites in this study, as well
as in previous studies by Marshall (1936), Gerell
(1970), and Shladweiler and Storm (1969).
Whitman (1981) reported logjams to be the most
common den type in Idaho (33% of dens), but bank
tunnels were also used often (30% of dens). As in
this study, these previous studies found that Mink,
especially males, use many different dens within
their home ranges. Marshall (1936) and Melquist et
al. (1981) reported that Mink choose daily den sites
based on proximity to current foraging areas.
Because home ranges were large, it was necessary
to maintain many different dens.

314

There is a need for more ecological investigations
of Mink in North America to elucidate factors influ-
encing movements, habitat use, and home range sizes.
Even basic aspects of Mink ecology such as territorial
behavior are poorly understood, and existing data
(Gerell 1970; Whitman 1981) are contradictory.

Acknowledgments

We thank E. Ramsay and the staff of the
Department of Comparative Medicine at The
University of Tennessee College of Veterinary
Medicine for their help, and M. J. Harvey, A. B.
Sargeant, J. Spencer, and S. Ratnayeke for helpful
advice. T. W. Arnold, J. S. Whitman, and two
anonymous reviewers provided helpful comments on
the manuscript. This research was supported in part
by an appointment to the U. S. Department of
Energy Graduate Student Research Program at the
Oak Ridge National Laboratory administered by Oak
Ridge Institute for Science and Education.

Literature Cited

Arnold, T. W., and E. K. Fritzell. 1987. Activity pat-
terns, movements, and home ranges of prairie mink.
Prairie Naturalist 19: 25-32.

Dunstone, N., and J. D. S. Birks. 1983. Activity budget
and habitat usage by coastal mink (Mustela vison
Shreber). Acta Zoologica Fennica 174: 189-191.

Eagle, T. C., and A. B. Sargeant. 1985. Use of den exca-
vations, decoys, and barrier tunnels to capture mink.
Journal of Wildlife Management 49: 40-42.

Enders, R. K. 1952. Reproduction in the mink.
Proceedings of the American Philosophical Society 96:
691-755.

Gerell, R. 1970. Home ranges and movements of the
mink Mustela vison Shreber in southern Sweden. Oikos
21: 160-173.

Katnik, D. D., D. J. Harrison, and T. P. Hodgman. 1994.
Spatial relations in a harvested population of marten in
Maine. Journal of Wildlife Management 58: 600-607.

THE CANADIAN FIELD-NATURALIST

Vol. 111

Kitchings, J. T., and L. K. Mann. 1976. A description of
the terrestrial ecology of the Oak Ridge Environmental
Research Park. Oak Ridge National Laboratory, Oak
Ridge, Tennessee. ORNL/TM-5073. 58 pages.

Linn, I. J., and J. D. S. Birks. 1981. Observations on the
home ranges of feral American mink (Mustela vison) in
Devon, England, as revealed by radiotracking. Pages
1088-1102 in Proceedings of the Worldwide Furbearer
Conference. Edited by J. Chapman and D. Pursley.
Worldwide Furbearer Conference, Inc., Frostburg,
Maryland. 2056 pages.

Linscombe, G., N. Kinler, and R. J. Aulerich. 1982.
Mink. Pages 629-643 in Wild Mammals of North
America: Biology, Management, and Economics. Edited
by J. Chapman and G. Feldhamer. The Johns Hopkins
University Press, Baltimore, Maryland. 1147 pages.

Marshall, W. H. 1936. A study of the winter activities of
the mink. Journal of Mammalogy 17: 382-392.

Mech, L. D. 1983. Handbook of animal radio-tracking.
University of Minnesota Press, Minneapolis,
Minnesota. 108 pages.

Melquist, W. E., J. S. Whitman, and M. G. Hornocker.
1981. Niche differentiation and coexistence of mink and
river otter. Pages 187-220 in Proceedings of the
Worldwide Furbearer Conference. Edited by J. Chapman
and D. Pursley. Worldwide Furbearer Conference, Inc.,
Frostburg, Maryland. 2056 pages.

Sandell, M. 1989. The mating tactics and spacing patterns
of solitary carnivores. Pages 164-182 in Carnivore
behavior, ecology, and evolution. Edited by J. L.
Gittleman. Cornell University Press, Ithaca, New York.
620 pages.

Schladweiler, G. L., and G. L. Storm. 1969. Den-use by
mink. Journal of Wildlife Management 33: 1025-1026.
Spencer, J. 1990. The mink manual. Outdoor World

Press, Inc. Humansville, Missouri. 128 pages.

Whitman, J. S. 1981. Ecology of the mink (Mustela
vison) in west-central Idaho. M.S. thesis. University of
Idaho, Moscow, Idaho. 101 pages.

Received 9 May 1996
Accepted 4 October 1996

1997

NOTES

215

Observations of a Possible Cleaning Symbiosis Between Painted
Turtles, Chrysemys picta, and Snapping Turtles, Chelydra serpentina,

in Central Ontario

MARGARET A. KRAWCHUK, NICOLA KOPER AND RONALD J. BROOKS

Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1

Krawchuk, Margaret A., Nicola Koper, and Ronald J. Brooks. 1997. Observations of a possible cleaning symbiosis
between Painted Turtles, Chrysemys picta, and Snapping Turtles, Chelydra serpentina, in central Ontario. Canadian

Field-Naturalist 111(2): 315-317.

Painted Turtles (Chrysemys picta) were observed removing chlorophytic algae and leeches (Placobdella spp.) from
Snapping Turtles (Chelydra serpentina) in a pond in central Ontario. We propose that a facultative mutualistic association
exists between the two species in that the Painted Turtles derive a nutritional benefit from the leeches and algae and the
Snapping Turtles benefit by having parasites removed. This may be the first reported instance of a cleaning symbiosis

between species of reptiles.

Key Words: Painted Turtle, Chrysemys picta, Snapping Turtle, Chelydra serpentina, mutualism, cleaning symbiosis, leech,

Placobdella parasitica.

The interactions between an organism and its
biotic environment fall into five main categories
based on how the activity affects the organism:
competition, predation, parasitism, mutualism and
commensalism (Begon et al. 1990). Described
interactions between turtles and other vertebrate
members of their ecological community tend to fall
into the first three of these categories. Mutualistic
and commensalistic interactions involving turtles
are rarely reported and, where such interactions
have been observed, they are usually categorized as
the latter (Grier 1984; Begon et al. 1990), in which
one species benefits and the other does not. For
example, Painted Turtles (Chrysemys picta) have
been reported to approach and climb on the cara-
pace of Snapping Turtles (Chelydra serpentina)
apparently to bask, but it appeared that there was
no benefit to the Snapping Turtles (Legler 1956). In
a similar example, Painted Turtles basked on the
head of a Water Buffalo (Syncerus caffer)
(MacGeorge 1928 in Boyer 1965). In another they
basked on the nesting platform of a Common Loon
(Gavia immer) and even on the nesting loon itself
(Gelatt and Kelley 1996), but because the turtles
primarily used the platform and not the loon to
bask, this relationship probably does not qualify as
being commensal.

In rare instances, turtles have been observed in
apparently mutualistic interactions, i.e. a symbiotic
interaction where both species receive a benefit,
usually food (Grier 1984). For example, on two
occasions a Common Grackle (Quiscalus quiscula)
was observed removing leeches (Placobdella para-
sitica) from map turtles (Graptemys ouachitensis)
(Vogt 1979). Vogt speculated that this interaction
reduced leech infestations in some populations of
Map Turtles and referred to the interaction as
cleaning symbiosis, defined as a cleaner organism

feeding on ectoparasites, scales, mucus or other
materials from the surface of a cooperating host
(Gorlick et al. 1978; Snelson et al. 1990). Other
examples in chelonians involved removal of exter-
nal parasites from tortoises (Vogt 1979). In these
few cases, the turtle or tortoise was the passive
partner in the interaction and observations suggest-
ed that the cleaning behaviour might have been
unique to one or two individuals. Here we describe
observations of an apparent cleaning symbiosis
between two species of turtles and discuss whether
this behaviour qualifies as the first reported
instance of mutualistic relationship between species
of turtles.

We observed the turtles at Wolf Howl Pond in
Algonquin Provincial Park, Ontario (45°34'N,
78°41'W). The pond is a 1.7-ha area of open water
and floating bog mats (Schwarzkopf and Brooks
1985; Lefevre and Brooks 1995) and sustains popu-
lations of approximately 200 Painted Turtles and 5-
10 Snapping Turtles. Most of these turtles are indi-
vidually marked with aluminum tags and carapacial
notches (Lefevre and Brooks 1995). Throughout the
nesting seasons of 1995 and 1996 we observed
(between 1630 and 2200h) the nesting activity of
both species of turtle from an embankment border-
ing the east side of the pond.

During the nesting season, females of both
species slowly swam up and down alongside the
pond bank waiting for an opportunity to nest.
Snapping Turtles nested in both early morning and
evening, and the Painted Turtles nested in the late
afternoon and evening. On several occasions in
1995 and 1996, male and female Painted Turtles
were observed pursuing female Snapping Turtles as
the latter floated or moved slowly in the water. In
this relaxed position, the majority of the skin sur-
face of the Snapping Turtle was exposed due to the

316

species’ small plastron, and the full extension of the
legs below the body and of the head forward. After
approaching a Snapping Turtle, Painted Turtles
investigated both the carapace, which often sustains
chlorophytic algal growth, and the skin, which is
commonly infested by both adults and broods of
leeches (Placobdella parasitica and P. ornata)
(Brooks et al. 1990). The Painted Turtles periodical-
ly explored beneath the Snapping Turtle’s body and
around the legs, neck and tail and were observed to
remove and eat both the algae and leeches. The
Snapping Turtles did not change position or appear
to react during these encounters. For example, at
1934h on 14 June 1996, several Painted Turtles
were observed feeding on two Snapping Turtles,
and at 1910h on 18 June 1996, male Painted Turtle
J6 was observed feeding on leeches on a passive
Snapping Turtle. Similarly, at 2000h on 4 July
1996, Painted Turtle J46 approached female
Snapping Turtle T6 and fed on leeches or algae on
T6. J46 continued to follow T6 as the Snapping
Turtle slowly swam beside the bank.

Painted Turtles have also been observed showing
aggression towards conspecifics who approached.
At 1945h on 13 June 1995, female Painted Turtle
C16 attempted to climb onto the carapace of an
unidentified female Snapping Turtle. When a sec-
ond, unidentified Painted Turtle approached the
Snapping Turtle, C16 lunged towards the other
Painted Turtle, which immediately swam away. C16
subsequently swam around and under the Snapping
Turtle for several minutes, and lunged twice
towards a third female Painted Turtle which
approached the same Snapping Turtle. This uniden-
tified female Painted Turtle immediately swam
away. C16 did not appear to bite the other Painted
Turtles. Similarly, at 1910h, 21 June 1996, female
Painted Turtles Y1 and A19 were observed swim-
ming around and feeding from a passive Snapping
Turtle. Yl was then seen to lunge violently, with
her mouth open, towards A19, and to snap at her.
A19 immediately swam away, but returned to feed
from the Snapping Turtle after Y1 had left. A19
removed leeches from the base of the tail and from
the hind leg pockets of the Snapping Turtle.

These associations were observed periodically
throughout the nesting seasons of 1995 and 1996.
We observed Painted Turtles feeding on leeches
and algae on Snapping Turtles on 10 separate occa-
sions. Although our observations occurred during
the nesting season this does not preclude occur-
rence of this behaviour at other times when we
were not observing them as intensively.

Lake Sasajewun (45°35'N, 78°30'W) is located
20 km east of Wolf Howl Pond within Algonquin
Park and is inhabited by Snapping Turtles, but not
by Painted Turtles (RJB, unpublished data). As in
Wolf Howl Pond, the turtles of Lake Sasajewun

THE CANADIAN FIELD-NATURALIST

Vol. 111

have been studied intensively over the past 15
years. Captures of Snapping Turtles were made pre-
dominantly in May and June but also continued
throughout July and August in both Lake Sasajewun
and Wolf Howl Pond, and numbers and locations of
leeches were recorded for all individuals. Using
two-tailed t-tests we compared leech abundance
within and between sites. Sample sizes were rela-
tively small, however significantly more leeches
were observed on Snapping Turtles at Lake
Sasajewun in July and August than on Snapping
Turtles at Wolf Howl Pond in the same interval (t =
4.44, df = 2, P = 0.0471). The mean number of
leeches found on turtles at Sasajewun was 48, and at
Wolf Howl Pond the mean was 3. We also found an
increase in leech numbers between May/June and
July/August at Lake Sasajewun (t = -3.925, df = 3,
P = 0.0294), whereas a decrease occurred at Wolf
Howl Pond (¢ = 2.941, df = 27, P = 0.00664). In
contrast, for the Painted Turtles at Wolf Howl Pond,
leeches were observed on only 13 of 113 individu-
als captured in May, which was not significantly
different from 2 of 25 individuals captured in June
(x? = 1.23, df = I, P > 0.25). This indicates that a
decrease in the leech population in the pond was not
the reason for the decrease in leech abundance on
the Snapping Turtles. Though many factors could
be contributing to the different levels of parasitism
by leeches, we speculate that the disappearance of
leeches from C. serpentina may be due in part to
their removal by Painted Turtles. This seems plausi-
ble given that Painted Turtles outnumber Snapping
Turtles by 20:1 in Wolf Howl Pond. The higher
density of leeches on Snapping Turtles in May may
occur because the Painted Turtles do not feed until
water temperatures rise above 15°C in late May
whereas the leeches are already active and seeking
hosts (Brooks et al., 1990). In June and July, when
water temperatures rise above 15°C, turtles begin
feeding extensively (Sexton 1989; this study). We
did not observe Painted Turtles remove leeches
from each other, perhaps because leeches were
much less abundant on Painted Turtles and even if
this feeding did occur it would be brief.

This interaction between Painted and Snapping
turtles and leeches may be akin to the mutual asso-
ciation observed between Oxpeckers (Buphagus
erythrorhynchus and B. africanus) in Africa and
their mammalian hosts which are usually large
ungulates (Benson 1973). The birds feed on the
ticks, blood-sucking flies and tissues of their hosts.
Through this association, the mammals have pests
removed while the birds receive food. As well, in a
marine environment, the Neon Cleaner Goby
(Gobiosoim oceanops) of the Caribbean, and the
Cleaner Fish (Labroides dimidiatus) of the Indo-
Pacific rid larger fish of their ectoparasites (Zander
1974). Similarly, the Painted Turtle receives food

oe 0

and the Snapping Turtle has leeches and algae
removed. In many cases, the “cleaners” are strongly
adapted physiologically and ecologically to per-
form ectoparasite removal and then to derive their
food primarily from this source (Snelson et al.
1990). In contrast, the behaviour exhibited by
Painted Turtles is likely to be sporadic and faculta-
tive, not obligate. Nevertheless, we suggest that this
behaviour may affect the numbers of leeches on
Chelydra and may provide the Painted Turtles with
a significant food source.

Although the biological significance of the
observed interactions between Painted Turtles and
Snapping Turtles at Wolf Howl Pond remains
uncertain, it is clear that this behaviour has occurred
repeatedly and involved several different turtles
over at least two years. Therefore, we suggest these
interactions could represent the first reported exam-
ple of a cleaning symbiosis between turtle species
and the first case of a turtle being an active partner
in a mutualistic interaction.

Literature Cited

Begon, M., J. L. Harper, and C. R. Townsend. 1990.
Ecology: individuals, populations and communities.
Blackwell Scientific Publications, Boston, Massa-
chussetts.

Benson, C. W. 1973. Starlings, Old World orioles and
Drongo. Page 375 in Grzimek's Animal Life
Encyclopedia, Volume 9, Birds III. Edited by H. C.
Bernhard Grzimek Van Nostrand Reinhold Company,
New York.

Boyer, D. R. 1965. Ecology of the basking habit in turtles.
Ecology 46: 99-118.

Brooks, R. J., D. A. Galbraith, and J. A. Layfield. 1990.
Occurrence of Placobdella parasitica (Hirudinea) on
snapping turtles, Chelydra serpentina, in south eastern
Ontario. Journal of Parasitology 76: 190-195.

Gelatt, T. S., and J. D. Kelley. 1995. Western painted tur-

NOTES

S17

tles, Chrysemys picta bellii, basking on a nesting com-
mon loon, Gavia immer. Canadian Field-Naturalist 109:
456-458.

Gorlick, D. L., P. D. Atkins, and G. S. Losey, Jr. 1978.
Cleaning stations as water holes, garbage dumps and
sites for evolution of reciprocal altruism. American
Naturalist 112: 341-353.

Grier, J. W. 1984. Biology of Animal Behaviour. Times
Mirror/Mosby, Toronto, Canada.

Isenhart, F. R., and D. F. Desante. 1985. Observations of
scrub jays cleaning ectoparasites from black-tailed deer.
Condor 87: 145-147.

Lefevre, K., and R. J. Brooks. 1995. Effects of sex and
body size on basking behavior in a northern population
of the painted turtle, Chrysemys picta. Herpetologica 51:
217-224.

Legler, J. M. 1956. A social relation between snapping and
painted turtles. Transactions of the Kansas Academy of
Science 59: 461-462.

Nagle, R. D., R. C. van Loben Sels, and J. D. Congdon.
1995. Observations of agonistic and mutualistic interac-
tions between birds and non-avian vertebrates on the E.
S. George Reserve, Michigan. Canadian Field-Naturalist
109: 464-466.

Schwarzkopf, L., and R. J. Brooks. 1985. Application of
operative environmental temperatures to analysis of
basking behavior in Chrysemys picta. Herpetologica 41:
206-212.

Snelson, F. F., Jr., S. H. Grober, F. L. Murru, and T. H.
Schmid. 1990. Southern stingray, Dasyatis americana:
host for a symbiotic cleaner wrasse. Copeia 1990:
961-965.

Vogt, R. C. 1979. Cleaning/feeding symbiosis between
grackles (Quiscalus: Icteridae) and map turtles
(Graptemys: Emydidae). Auk 96: 608-609.

Zander, L. D. 1974. Blenmies and gobies. Pages 181-184
in Grzimek's Animal Life Encyclopedia. Volume 5,
Fishes II Amphibians. Edited by H. C. Bernhard
Grzimek. Van Nostrand Reinhold Company, New York.

Received 24 June 1996
Accepted 22 October 1996

318

THE CANADIAN FIELD-NATURALIST

Vol. 111

Weissia brachycarpa (Nees & Hornsch.) Jur. at Niagara Falls,

A Moss New to Ontario

P. M. ECKEL

Clinton Herbarium, Buffalo Museum of Science, Buffalo, New York 14211

Eckel, P. M. 1997. Weissia brachycarpa (Nees & Hornsch.) Jur. at Niagara Falls, a moss new to Ontario. Canadian Field-

Naturalist 111(2): 318-319.

A collection of the rare moss Weissia brachycarpa at Niagara Falls, is the first record for Ontario, the third Canadian
province in which the species is known to occur. This plant is a noteworthy addition to the unusually rich biodiversity of

the Niagara River Gorge.

Key Words: Moss, Weissia brachycarpa, Ontario, Niagara Falls.

During a study of the mosses of the Niagara River
Gorge the following moss was discovered:

Weissia brachycarpa (Nees & Hornsch.) Jur. Canada:
Ontario, Welland, Regional Municipality of Niagara,
43°30'N, 79°10'W, City of Niagara Falls; base of wooded
embankment overlooking the Horseshoe (Canadian) Falls;
in spray on calcareous clay in lawn; 28 May 1991, Eckel
9302250 (Buffalo Museum of Science).

According to Ireland et al. (1987) Weissia
brachycarpa in Canada has been found in two
provinces: British Columbia at one end of the coun-
try and Nova Scotia at the other. Stations reported
for the United States by Crum and Anderson (1981)
are located south to North Carolina and west to
Texas, but otherwise the New World distribution is
largely eastern North American. The Old World dis-
tribution includes temperate Europe east to the
western states of the former Soviet Union. It is con-
sidered rare throughout its range (Crum and
Anderson 1981).

Weissia brachycarpa has been recorded recently
in New York State in nearby Niagara County
(Eckel 1987) and just across the Niagara River from
the Ontario station on Goat Island in the city of
Niagara Falls, New York (Eckel and Eckel 1988).
All collections were found at the margins of grassy
areas by roadsides or in lawns, a habitat typical for
the species. These are the only stations recorded in
New York.

As with other species in the genus, now enlarged
by the incorporation of Astomum (Stoneburner
1985), the plants cannot be identified if sterile.
Capsules are necessary for their determination. The
capsules of Weissia brachycarpa are distinctive in
the smallness of the capsule opening, perhaps indi-
cating a tendency of the species toward cleistocarpy,
a feature of the subgenus Astomum. The capsules
and leaves tend to be broader than a more common
species, Weissia controversa Hedw., with which it
could most easily be confused.

Other mosses found associated at the Ontario sta-
tion were Amblystegium varium (Hedw.) Lindb.,
Barbula convoluta Hedw., B. unguiculata Hedw.,

Fissidens taxifolius Hedw., Phascum cuspidatum
Hedw., Physcomitrium pyriforme (Hedw.) Hampe,
Pleuridium subulatum (Hedw.) Rabenh., and Pottia
truncata (Hedw.) Fuernr. All are typical of dis-
turbed soil. Species found here but not recorded for
Welland County (now part of the Regional
Municipality of Niagara) according to Ireland and
Cain (1975) are Barbula convoluta, B. unguiculata,
Phascum cuspidatum and Pleuridium subulatum.
The last species is known from only seven other sta-
tions in Ontario (Ireland and Ley 1992). Specimens
of these mosses are deposited at BUF.

The Niagara River Gorge and its environs have
supported a rich tradition of bryophyte interest
beginning with such notables in the history of bryol-
ogy as Thomas Drummond, collecting on the
Canadian side in the early nineteenth century, con-
tinuing with John Macoun, Leo Lesquereux and T.
P. James, among others, in the latter part of the
nineteenth century. It was also then when Judge
George W. Clinton, member of the Regents of the
University of the State of New York, made system-
atic collections of interesting mosses on both sides
of the Niagara River for distribution to various
herbaria. Nearly all of these individuals included
specimens collected at Niagara in exsiccatae distrib-
uted in the past century. That the moss flora still has
populations of interest is supported by recent dis-
coveries, such as Eucladium verticillatum (Brid.)
Bruch & Schimp. in BSG in Ontario (Eckel 1990)
and Didymodon australasiae var. umbrosus
(C.Muell.) Zand. (Eckel 1986) and Bryum rubens
Mitt. (Eckel and Shaw 1990) on the American side
of the River.

The gorge of the Niagara River is the focus of the
distributive power of converging transient human
populations. It exists in the midst of several dense
transportation networks, not to mention its function
as a biological conduit for the Great Lakes water-
shed, the waters of which are constricted into a
strait, bisecting and thereby creating the Niagara
Peninsula. Such a concentration of vectors for dias-
pore movement are coupled with the gorge of the

1997

River: an unusual haven of relatively undisturbed
complexes of microhabitats for the capture and
retention of such diaspores. These conditions pre-
date European migrations into the region, with the
consequence that the gorge environment and the
microhabitats along its rim contain nearly 75 per-
cent of the biodiversity (of vascular plants) known
for western New York and adjacent Ontario (Eckel,
manuscript in preparation). The gorge appears to
represent a biological sink for the chance seed or
spore that happens into it. The extraordinary biodi-
versity of the Niagara River and gorge environment
was noted earlier by David F. Day (1887).

Despite a return to the locality in every year sub-
sequent to the original collection, I have been
unable to rediscover the populations of Weissia
brachycarpa. Whether this is due to their naturally
ephemeral nature, to chemical lawn management, or
to any of a number of other pressures characteristic
of the natural and artificial environment of the area,
is unknown.

Acknowledgments

Richard Zander determined the specimen collect-
ed at Niagara and the manuscript has benefited from
his suggestions and comments. Permission to study
the floras of the Niagara Gorge has been granted to
the author by the Niagara Parks Commission,
Ontario, and the New York State Department of
Parks, Recreation and Historic Preservation. This
study has been supported in part by a grant from
the Niagara Frontier Chapter of the Adirondack
Mountain Club.

NOTES

319

Literature Cited

Crum, H. A. and L. E. Anderson. 1981. Mosses of Eastern
North America. Columbia University Press, New York.

Day, D. F. 1887. Catalogue of the Niagara Flora. Annual
Report of the Commission for the State Reservation at
Niagara for the Year 1887: 67-133. Also reprinted as
pamphlet, Troy, New York, 1888.

Eckel, P. M. 1987. Mosses new and rare for New York
State. Rhodora 89: 375-379.

Eckel, P. M. 1986. Didymodon australasiae var. umbrosus
(Musci: Pottiaceae) new to eastern North America.
Bryologist 89: 70-72.

Eckel, P. M. 1990. Eucladium verticillatum (Musci) sec-
ond Ontario station. Evansia 7: 15.

Eckel, P. M., and M. P. Eckel. 1988. Weissia hedwigii on
Goat Island, second locality in New York State. 1988.
Clintonia 3(3): 8—9

Eckel, P. M., and J. Shaw. 1990. Bryum rubens from
Niagara Falls, new to New York State. Bryologist
94(1): 80-81.

Ireland, R. R., G. R. Brassard, W. B. Schofield, and D.
H. Vitt. 1987. Checklist of the mosses of Canada II.
Lindbergia 13: 1-62.

Ireland, R. R., and R. F. Cain. 1975. Checklist of the
Mosses of Ontario. National Museums of Canada
Publications in Botany 5, Ottawa.

Ireland, R. R., and L. M. Ley. 1992. Atlas of Ontario
Mosses. Canadian Museum of Nature, Syllogeus
(Canadian Museum of Nature, Ottawa, Ontario) 70:
1-138.

Stoneburner, A. 1985 [1986]. Variation and taxonomy of
Weissia in the southwestern United States. II.
Taxonomic treatment. Bryologist 88: 293-314.

Received 19 July 1996
Accepted 20 October 1996

320

THE CANADIAN FIELD-NATURALIST

Vol. 111

The Moss Tortella alpicola Dix. New to Alberta and the
Yukon Territory with a Discussion of its Range and Comments

on Related Species

P. M. ECKEL

Clinton Herbarium, Buffalo Museum of Science, Buffalo, New York 14211-1293.

Eckel, P. 1997. The moss Tortella alpicola Dix. new to Alberta and the Yukon Territory with a discussion of its range and
comments on related species. Canadian Field-Naturalist 111(2): 320-322.

Tortella alpicola Dix., once considered rare as only a few isolated stations had been recorded in North America, is shown
to have a more widespread distribution. Characteristics distinguishing this species from the common and widespread

Tortella fragilis and T. tortuosa are presented in a key.

Key Words: Tortella alpicola, Tortella fragilis, Tortella tortuosa, Tortella nitida, Alberta, Canada, Yukon Territory, rare

bryophytes, moss.

Among the collections of undetermined bryophyte
material of the Geography Department of the
University of Calgary (those not sent to the
Provincial Museum of Alberta in Edmonton in 1995
and examined subsequent to that year), specimens of
Tortella alpicola Dix. came to light. The North
American distribution of this species, due to taxo-
nomic difficulties, has not yet been adequately docu-
mented. The University of Calgary specimens, and
three others seen from Canada at other herbaria, rep-
resent the present known distribution of the species
in that country (herbarium code throughout follows
that designated by Holmgren et al. 1990):

Tortella alpicola:

Alberta: Red Deer River Watershed, provincial campsite at
junction of James River and Willson Creek on
Cochrane-Nordegg Forestry Trunk Road, approx.
51°48’N, 115°03’W, elev. 4400 feet, June 23, 1961, C.
D. Bird 6249; with Encalypta vulgaris (Geography
Department, University of Calgary); Bow River
Watershed, 3 miles WSW of Exshaw, SW17-24-9-W5,
51°03’'N, 115°13’W, elev. 4300 ft., Upper Foothills
Forest, north-facing slope with steep limestone expo-
sures, on May 19, 1976, Glennis Lewis no. 36, with
Encalypta rhaptocarpa and Gymnostomum aeruginosum
(BUF); Jasper National Park, Dryas drummondii flat 2-3
miles below Athabaska Glacier, July 23, 1959, W. A.
Weber B-2278 (COLO).

Northwest Territories: Ellesmere Island, lake east of
Taconite Inlet, 82°52’N, 78°05’W, cliff of metamorphic
rock, 50 m, May 21, 1969, G. Brassard 4136 (CANM).

Yukon Territory: Selwyn Mountains, Itsi Range, North end
of unnamed lake, south of Fuller Lake, 62—57'N,
130-09’W. 1600 m, in mesic Betula pumila-Vaccinium
uliginosum dominated tundra on east and north facing
slopes, siliceous rock outcrops of granite and slate pre-
sent, with small amounts of calcareous rock cropping
out, August 10, 1978, D. H. Vitt s.n. (MO).

This species, as Sarconeurum tortelloides Greene,
was first described from Antarctica (Greene et al.
1970). Robinson (1972) placed the species into the
genus Tortella. Zander and Hoe (1979) included a

specimen from Hawaii in the distribution under the
name Tortella fragilis var. tortelloides (Greene)
Zand. & Hoe, as Zander and Hoe considered it only
marginally distinct from Tortella fragilis (Hook. &
Wils in Drumm.) Limpr. After study of material of
the genus Tortella from throughout Arctic North
America, Zander (manuscript in preparation) con-
cluded that the variety was a good species, as origi-
nally determined by Robinson.

Following Zander’s decision, Eckel (1991) pub-
lished the discovery of the species in the contermi-
nous United States from the state of Wyoming,
including reference to a station in the Northwest
Territories of Canada. The taxon was discussed
again by Hyvoenen (1991), who indicated, based on
his examination of specimens of Tortella fragilis
from the southern hemisphere, that Tortella tortel-
loides should be placed under Tortella fragilis as a
variety again. My examination of specimens at H
from Tierra del Fuego collected by H. Roivainen,
some of which were annotated by Hyvoenen, has
shown all of these specimens to be Tortella fragilis
var. fragilis. None of them were what is indicated
here as Tortella alpicola. Other specimens at CHR,
H, NY and ALTA of both Tortella fragilis and T.
tortuosa collected in the southern hemisphere were
all either correctly determined or were species other
than T. alpicola. So far, the only known southern
hemisphere stations for the latter species are in
Antarctica. The species has been found in Colombia
(Churchill and Linares 1995), and may prove to be a
member of the Andean flora. Zander (1993) reported
an earlier name for Tortella tortelloides based on the
type of T. alpicola Dixon (1930), collected from a
station in the Himalayas.

As a result of my examination of numerous speci-
mens of the genus Tortella from North American
herbaria, as well as material from the southern
hemisphere, Tortella alpicola, is reconfirmed taxo-
nomically as a distinct species. Some account of its
distribution has begun to emerge, at least in North

1997

America. It is apparently a species of more widespread
occurrence than previously thought.

With clarification of its distinctive characters,
Tortella alpicola is readily differentiated from the
two other species in the genus with which it is most
often confused: T. tortuosa (Hedw.) Limpr. and T.
fragilis, which are usually robust plants: T. tortuosa

NOTES

a2!

can reach 6 cm, 7. fragilis 5 cm in height, and both
are typically two or more cm tall. Since T. alpicola is
usually only one cm in height (rarely 1.5 cm), the
former two species can most often be differentiated
by height alone. It is when specimens of any one of
the three species are only one cm in height or less
that the real confusion arises between them:

Key to North American specimens of Tortella tortuosa, T. fragilis and T. alpicola when stems are

only one cm high:

. Leaf apices not deciduous (youngest leaves at the stem apex not propaguloid) with nearly subulate apices composed of
ummnimemurc eels leat -colls/S—Iils unm Wide ciiiessy 222.0 1.se2.0...s00c0sscdes<scalvinns (osvadcosbucdesselecencesdedecdduceacsschen Tortella tortuosa.
. Leaf apices deciduous (youngest leaves at the stem apex with deciduous, propaguloid modifications) the apices
mucronate with fewer cells, occasionally to five cells, leaf cells 9—-13(-14) pm Wide ..0........ceeececeeeecessseceesseeceseeeeseeeesees a
2. Propaguloid apices of youngest leaves in one straight segment without articulations, with smooth, straight margins
bordered by elongate, smooth cells; stems usually red-tomentose with fine, conspicuous tomentum ....... Tortella fragilis.
2. Propaguloid apices of youngest leaves articulated into several “barrel-shaped” segments, with crenulated-papillose

margins of quadrate cells; stems without tomentum, or rhizoids sparse and COaPSE€ .............::cceeesseeeeeeeees Tortella alpicola.

There are other less definite distinctions, such as
the snowy white hyaline basal cells in Tortella alpi-
cola, while these regions in the other two taxa have
a yellowish tint. Both Tortella tortuosa and T. frag-
ilis have conspicuous red tomentum, but T. alpicola
does not. The largest specimens of 7. alpicola seen
tend to be more rhizoidiferous than is typical, while
small variants of T. fragilis tend to have less
tomentum, but at least some stems can be found in
a collection that are red-tomentose. The clear,
smooth elongate cells on the upper margin of espe-
cially young leaves of Tortella fragilis is a critical
character in distinguishing this species from both T.
alpicola and forms of T. tortuosa with which it may
be confused (Zander, manuscript in preparation).

Among the specimens seen from the southern
hemisphere, a tiny example of T. fragilis from
Campbell Island in Antarctica proved difficult to
differentiate from T. alpicola, mainly because of
the apparent absence of any propaguliferous leaf
apex modifications of any distinction. Although the
apices of the leaves were very narrow and decidu-
ous (fragile), most of them possessed the clear bor-
der of thicker-walled elongate cells and, with dili-
gence, one or two apices could be found with thick-
ened modifications. The larger average leaf cells of
some of these reduced specimens also duplicate the
typical size of cells of T. alpicola. The specimen
was, however, coarse, with blackened tomentum at
the very base. The marginal border has proved criti-
cal in naming similar specimens of T. fragilis.

Actually, the species that is most similar to T.
alpicola, even to the barrel-shaped segmentation in
the apices of the young leaves, is small, narrow-
leaved forms of Trichostomum tenuirostre (Hook.
& Tayl.) Lindb. Specimens of the latter species,
however, are readily separated by the thicker-
walled cells on the leaf margins at the base, whose
cells do not form the characteristic vee-shape of the

genus Tortella. There are also occasional low and
distant teeth on the upper leaf margin.

Specimens of Tortella alpicola seen to date are
associated with shaded or exposed, wet or dry
rocks, crevices and ledges of granite, quartzite,
sandstone and calcareous outcrops on cliffs and in
canyons, but also in wet, mesic tundra in the Yukon
at elevations between 900 and 3300 m. In addition
to its Canadian range, its general distribution so far
noted is North America in Alaska, Arizona,
Colorado, Idaho, Montana, Nebraska, Utah,
Wyoming; South America in Colombia, Asia in
India, Pacific Islands in Hawaii, and Antarctica
(Alexandra Island).

That a peculiar species of Tortella occurs in our
flora has shown itself in the literature in various
ways. Tortella nitida (Lindb.) Broth. was ascribed
to the North American flora by Haring (1938) and
Flowers (1973). This has been suggested to be a
variety of Tortella tortuosa by Crum and Anderson
(1981), who also state that material cited as T. niti-
da by Haring was in fact either T. fragilis or T. tor-
tuosa. Material from Utah, described by Flowers,
does not appear to have been seen by them.

Tortella nitida, a European species, however,
has basal cells gradually, not abruptly, differentiat-
ed from the laminal cells, a shining costa on the
dorsal leaf surface with no distinctive subulate
propaguloid leaf apex, and leaf cells to 10 um wide.
Tortella nitida, by reason of its basal cells and
other characters, has been thought to possibly be a
fragile-leaved form of Trichostomum crispulum (R.
Zander, personal communication). North American
material examined for the present study that was
named Tortella nitida is variously T. alpicola, T.
fragilis, T. tortuosa and Trichostomum tenuirostre,
with Tortella alpicola specimens predominating in
the western United States of Utah and Colorado.
Tortella alpicola differs from T. nitida immediately

322

in the brightly distinct, clear basal cells set off
from the green laminal cells as well as the other
characters mentioned. One specimen cited as T.
nitida from Alaska proved to be Trichostomum
tenuirostre, distinguished by the basal cells not in a
vee-shape, bulging-mammillose cells, and tiny
papillae often arranged in two parallel rows over
each lumen.

Specimens of T. alpicola from North America
have also been called T. rigens N. Alberts. (Weber
1973) due to the wide leaf cells (ca. 14 um), but
plants of that species in Sweden are longer, 1.5—3
cm high, and lack apical propagulae (see also
Nyholm 1989). Examination of material of this
species from Sweden showed T. rigens to have a
strong morphological similarity to T. fragilis and to
be different in many characters from T. alpicola,
especially in leaf shape and color.

Acknowledgments

I thank Stewart Scott, Department of Geography,
University of Calgary, for providing me with a col-
lection of undetermined specimens to examine,
collected by C. D. Bird and his associates.

I thank the curators of bryophytes at ALTA,
CANM, CHR, COLO, H, MO, and NY for allow-
ing the examination of specimens from their
herbaria. I am grateful to Richard Zander, who per-
mitted me access to his unpublished manuscript
discussing the genus Tortella in Arctic North
America, and who reviewed the present paper.
Outside reviewers contributed substantially to
improvements in the manuscript.

Literature Cited

Churchill, S. P., and E. L. Linares C. 1995. Prodromus
Bryologiae Novo- Granatensis: Introduccion a la Flora
de Musgos de Colombia. Partes 1 y 2. Biblioteca “Jose
Jeronimo Triana” 12. Santafe de Bogota.

THE CANADIAN FIELD-NATURALIST

Vol. 111

Crum, H. A., and L. E. Anderson. 1981. Mosses of
Eastern North America. 2 Volumes. Columbia
University Press, New York.

Dixon, H. N. 1930. Additions to the moss flora of the
North-Western Himalayas. Annales Bryologici 111:
51-70.

Eckel, P. M. 1991. Tortella tortelloides (Musci:
Pottiaceae) new to North America. Bryologist 94: 84-87.

Flowers, S. 1973. Mosses: Utah and the West. Brigham
Young University Press, Provo, Utah.

Greene, S. W., D. M. Greene, P. D. Brown, and J. M.
Pacey. 1970. Antarctic moss flora. 1. The genera
Andreaea, Pohlia, Polytrichum, Psilopilum, and
Sarconeurum. British Antarctic Survey Science Report
64: 1-118.

Haring, I. M. 1938. Tortella. In Moss Flora of North
America. Edited by A. J. Grout, 1(3): 165-170.
Newfane, Vermont.

Holmgren, P. K., N. H. Holmgren, and L. C. Barnett.
1990. Index Herbariorum. Part 1. Edition 8. New York
Botanical Garden, Bronx, New York.

Hyvoenen, J. 1991. Tortella fragilis (Pottiaceae) report-
ed for Southern South America. Bryologist 94:
416-418.

Nyholm, E. 1989. Illustrated Flora of Nordic Mosses.
Fasc. 2. Pottiaceae-Splachnaceae-Schistostegaceae.
Nordic Bryological Society. Copenhagen, Denmark.

Robinson, H. E. 1972. Observations on the origin and
taxonomy of the Antarctic moss flora. Pages 163-177 in
Antarctic Terrestrial Biology. Edited by G. A. Llano.
Antarctic Research Series 20. American Physical Union,
Washington, D.C.

Weber, W. A. 1973. Guide to the mosses of Colorado.
Institute of Arctic and Alpine Research, University of
Colorado, Occasional Paper Number 6.

Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of
harsh environments. Bulletin of the Buffalo Society of
Natural Sciences 32: i-vi + 1-378.

Zander, R. H., and W. J. Hoe. 1979. Geographic dis-
junction and heterophylly in Tortella fragilis var.
tortelloides (=Sarconeurum tortelloides). Bryologist
82: 84-87.

Received 12 August 1996
Accepted 29 October 1996

News and Comment

Editor’s Report for The Canadian Field-Naturalist Volume 110 (1996)

Research, observation, synthesis, or tribute
manuscripts submitted to The Canadian Field-
Naturalist totalled 125 in 1996, an increase of 30
over 1995. Reviews were completed for all but 21 by
the end of the year. Of the 104 returned to authors,
72 were revised and accepted within the year. In
addition, 37 revisions of manuscripts submitted in
1995 and 1 submitted in 1993 were received and
accepted. Of the 110 accepted, 55 were published in
1996 and 55 are in galley, or submitted to the printer
to be set, for 1997. These will fill issues 111(1), (2),
and (3).

Totals for circulation to members and subscribers
to The Canadian Field-Naturalist in 1996 together
with those of 1995 are given in Table 1. Total copies
distributed were only down by 22 despite a drop in
memberships of 30.

Issue mailing dates for volume 110 were: (1) 30
April, (2) 4 July, (3) 22 November 1996, and (4) 2
April 1997. The volume totalled 762 pages; the
largest single issue (1) was 254 pages. This, the
largest in the journal's history, was a special issue:
“A Life with Birds: Percy A. Taverner, Canadian
Ornithologist, 1875” a biography by John L.
Cranmer-Byng (see Editor’s Report for Volume 109
The Canadian Field-Naturalist 110(2): 360-363 for
acknowledgements to the numerous people who
assisted with this issue).

Special thanks are due to Joan Hebden who pre-
pared the tribute to the late honourary member,
Claude Eugene Garton (1907-1996). This appeared
in 110(3): 554-557, and the many people in the
Thunder Bay area who assisted her, the editor, or
both, with this article are acknowledged in the
tribute individually.

The number of articles and notes in volume 110 is
summarized in Table 2 by topic. The totals for Book
Reviews and New Titles are given in Table 3, and
the distribution of published pages among issues in
Table 4.

Prepared this year for Volume 111, number 1, was
another Special Issue: “Orchids in the Ottawa
District: Floristics, Phytogeography, Populations
Studies and Historical Review” by Joyce Reddoch
and Allan Reddoch. This study synthesized over 30
years of detailed study, as well as historical observa-
tions back to the 1850s, in the area 50-km radius
from the Parliament Buildings by the authors and
members of the Ottawa Field-Naturalists’ Club, and
provides original data for this area for 44 species
included. Prepared for 111(2) were six COSEWIC
status reports for Fish and Marine Mammals
approved in 1995.

M.O.M. Printers, 300 Parkdale Avenue, Ottawa,
set and printed the journal and special thanks are due
Emile Holst and Eddie Finnigan and their staff.
Wanda J. Cook proof-read the galleys for the vol-
ume. Bill Cody continued as Business Manager, a
role he has shouldered well since he was appointed
“Treasurer’s Assistant” 21 May 1947 (soon after ini-
tially joining the Club 7 December 1946). The title
of the expanding post was changed to “Business
Manager” 15 December 1948. [Special thanks to the
informal club historian, Dan Brunton, for this infor-
mation from his notes on the Minutes of OFNC
Council]. Bill was assisted by Lois Cody who for-
mally relinquished her long-time role of assistant to
the Business Manager during the year. Bill also
oversaw the compilation, and proof-read and edited,
the Index for volume 110 which was diligently pre-

TABLE 1. The 1996 circulation of The Canadian Field-Naturalist (with 1995 figures in parenthesis). Membership totals
from Membership Committee analysis, Annual Report of the Ottawa Field-Naturalists' Club (page 334); subscription totals
compiled by W. J. Cody. Forty percent of membership dues and 100% of subscription payments go to production of The
Canadian Field-Naturalist. Members can vote on Club affairs, subscribers can not.

Canada
Memberships
Family & individual 987 (1017)
Subscriptions
Individual 186 (170)
Libraries 198 (206)
TOTALS ESTE. £1395)

USA Foreign Totals
Sek) Sab) 1024 (1054)
62. (61) 6° .@) 254 (239)

267 (266) 46 (46) ait) G18)
361 (358) 57 (60) 1789 (1811)

323

324

TABLE 2. Number of articles and notes published in The
Canadian Field-Naturalist Volume 110 (1996) by major
field of study.

Subject Articles Notes Total
Mammals i? 14 31
Birds 11 9 20
Amphibians

and reptiles Z 1 3
Fish | 2. 9
Invertebrates 2 0 pf
Plants 8 2 10
Other* 1* 0 1*
Totals 48 28 76

*COSEWIC Fish and Marine Mammal Subcommittee
report 110(3). In addition, there was one tribute article
(Claude Eugene Garton) in News and Comment section
110(3).

pared by Leslie Durocher. Wilson Eedy continued as
Book-Review Editor, including compilation of New
Titles, despite his ever-increasing world travel on
other duties (his detailed annual report will appear
separately). George La Roi continued to serve as
Coordinator of the Biological Flora of Canada (a
summary of this series will appear separately).

All article and note manuscripts submitted to the
Canadian Field-Naturalist are normally reviewed by
at least one associate editor and usually by one or
more additional reviewers as well. This combination
of associate editors who review the majority of man-
uscripts submitted in a field through the year plus
outside specialists (chosen by the editor or an associ-
ate editor, or selected from alternates suggested by
authors) who may see only one (but sometimes
more) CFN manuscript(s) a year maintains a balance
of both consistency and freshness of opinion. The
editor makes the final decisions on manuscripts and
corresponds with the authors, largely based on the
advice of all reviewers. I am indebted to all who
returned reviews in 1996 (if more than one review,
the number is in parenthesis):

Associate Editors (listed also on inside front cover
of each issue): Mammalogy: William O. Pruitt, Jr.,

TABLE 3. Number of reviews and new titles published in
Book Review section of The Canadian Field-Naturalist
Volume 110, by topic.

Reviews New Titles
Zoology 18 87
Botany 3 = fi
Environment 15 74
Miscellaneous 3 12
Young Naturalists 0 +4
Totals 49 268

THE CANADIAN FIELD-NATURALIST

Vol. 111

Department of Zoology, University of Manitoba,
Winnipeg (34; plus 3 reviewed by others); Warren
B. Ballard, Wildlife Research Unit, Faculty of
Forestry, University of New Brunswick, Fredericton,
and [28 August] Arizona Game and Fish
Department, Phoenix (17; and additional comments
on 16 reviewed by others); Ornithology: Anthony J.
Erskine, Sackville, New Brunswick (33); W. Earl
Godfrey, Canadian Museum of Nature, Ottawa,
Ontario (9); Fish and Marine Mammals: Robert R.
Campbell, Woodlawn, Ontario (3; plus 15
COSEWIC status reports); Ichthyology: Brian W.
Coad, Canadian Museum of Nature (2);
Entomology: R. Anderson, Canadian Museum of
Nature (5); Botany: Charles D. Bird, Erskine,
Alberta (17). Joining the Associate Editors in 1997
will be Paul Catling, Agriculture Canada, Ottawa. In
addition Claude Renaud, Canadian Museum of
Nature, gave special assistance with editing the
french abstracts of the COSEWIC fish and marine
mammal status reports.

Additional reviewers: Brad Andres, U.S. Fish and
Wildlife Service, Anchorage, Alaska; Todd Arnold,
Department of Wildlife Humbolt State University,
Arcata, California; Darlene Balkwill, Canadian
Museum of Nature; J. F. Barr, Guelph, Ontario;
Spencer C. H. Barrett, Department of Botany,
University of Toronto; Rene J. Belland, Devonian
Botanical Garden, University of Alberta, Edmonton;
J. Roger Bider, Department of Renewable Resources
MacDonald College of McGill University and the
Ecomuseum, Ste. Anne de Bellevue, Quebec (3);
D. A. Boag, Brentwood Bay, British Columbia (3);
Stan Boutin, Department of Biological Sciences,
University of Alberta, Edmonton, Alberta; Jeff
Bowman, New Brunswick Cooperative Fish and
Wildlife Research Unit, University of New
Brunswick, Fredericton (3); Ronald J. Brooks,
Department of Zoology, University of Guelph; Luc
Brouillet, Herberier Marie-Victorin, Institute
botanique, Université de Montreal; Daniel F.
Brunton, Ottawa (2); Ludwig Carbyn, Canadian
Wildlife Service, Edmonton (2); Paul M. Catling,
Agriculture Canada, Ottawa (6); Nigel Caulkett,
Department of Veterinary Anesthesiology,
Radiology, and Surgery, Western College of
Veterinary Medicine, Saskatoon, Saskatchewan;
Jacques Cayouette, Agriculture Canada, Ottawa;
Robert G. Clark, Canadian Wildlife Service,
Saskatoon, Saskatchewan; W. J. Cody, Agriculture
Canada, Ottawa (2); William J. Crins, Ontario
Ministry of Natural Resources, Huntsville; Vince
Crichton, Manitoba Department of Natural
Resources, Winnipeg (3); Peter Ross Croskery,
Grimsby, Ontario; Stephen Darbyshire, Agriculture
Canada; A. W. Diamond, Department of Biology,
University of New Brunswick, Fredericton; George
W. Douglas, British Columbia Conservation Data

1997

NEWS AND COMMENT

325

TABLE 4. Number of pages published in The Canadian Field-Naturalist Volume 110 (1996) by section (number of manu-

scripts in parenthesis).
see

76 (13)
23: (12)
17 (20)

Eas
243 (1)

Issue number

Articles

Noies

News and Comment
Book Reviews - - £G. Cr?)
Index 9 (1) - -
Notices - = - =

2 (2)

Advice to Contributors ie awe

Total pages: 254 132

-3- -4- Total

146 (17) 127,402) 592 (48)
EZ... (6) 18 (10) 52 (28)
Fisk23) 1 (2) 34 (44)
14 (18) 14 (13) 43 (49)
ay eee 29 (14) 38 (2)
Ey ies Zz (2 2 2)
- - 1 4) wth)
184 192 762

*Total pages for book review section include both reviews and new titles listings but parenthesis figures include only the

number of reviews.

Centre, Victoria; David Duncan, Saskatchewan
Wetland Conservation Corporation, Regina; Graham
Forbes, Wildlife Research Unit, Faculty of Forestry,
University of New Brunswick, Fredericton; Bill
Freedman, Department of Biology, Dalhousie
University, Halifax, Nova Scotia (2); Chuck Geale,
Tree Canada Foundation, Ottawa; Lynn Gillespie,
Canadian Museum of Nature; Mark Fuller, Raptor
Research and Technical Assistance Centre,
National Biological Service, Boise, Idaho; Frank S.
Gilliam, Department of Biological Sciences,
Marshall University, Huntington, West Virginia;
Karan Graham, Department of Zoology, University
of Guelph, Ontario; Patrick T. Gregory, Depart-
ment of Biology, University of Victoria, British
Columbia (2); Graham Griffiths, Department of
Biological Sciences, University of Alberta,
Edmonton; Thomas Herman, Biology Department,
Acadia University, Wolfville, Nova Scotia; Stuart
Houston, Saskatoon, Saskatchewan; Richard L.
Hutto, Division of Biological Sciences, University
of Montana, Missoula; Robert Ireland, Nepean,
Ontario (5); Ross D. James, Sunderland, Ontario
(3); Stephen Jenkins, Department of Biology,
University of Nevada, Reno (3); William Jensen,
North Dakota Game and Fish Department,
Bismarck; Tony Keith, Canadian Wildlife Service,
Hull, Quebec; Brina Kessel, University of Alaska
Museum, Fairbanks, Alaska; Edward Kott,
Department of Biology, Sir Wilfred Laurier
University, Waterloo; Paul A. Krausman, College
of Agriculture, School of Renewable Natural
Resources, University of Arizona, Tucson,
Arizona; Karl W. Larsen, Alberta Pacific Forest
Industries Inc., Boyle; Yves de Lafontaine, Saint-
Laurent Vision 2000, Environment Canada,
Montreal, Quebec; Raymond LeClair, Department
de Cimie-Biologie, Université du Quebec a Trois
Riviéres; Ross Lein, Department of Biology,
University of Calgary; Diane L. Leudorf,
Department of Biology, York University, North
York, Ontario; Nora Lewis, Department of Animal

Science (Agriculture), University of Manitoba,
Winnipeg; Lawrence E. Licht, Department of
Biology, York University, North York, Ontario; W.
Bruce McGillivray, Provincial Museum of Alberta,
Edmonton; Mike Malhiot, Ontario Ministry of
Natural Resources, Wingham; John Maunder,
Newfoundland Museum, St. John’s; Norton Miller,
Biological Survey, New York State Museum,
Albany; L. David Mech, U.S. Fish & Wildlife
Service, N. Central Forest Experiment Station, St.
Paul, Minnesota; David Nagorsen, Vertebrate Unit,
Royal British Columbia Museum, Victoria (5); R.
W. Nero, Manitoba Natural Resources, Winnipeg;
Thomas W. Nudds, Department of Biology,
University of Guelph, Ontario; Martyn E. Obbard,
Ontario Ministry of Natural Resources, Maple;
Henri Ouellet, Canadian Museum of Nature; Gerry
Parker, Canadian Wildlife Service, Sackville, New
Brunswick (2); Lawrence Powell, Department of
Biology, University of Calgary, Alberta (2);
William F. Porter State University of New York,
College of Environmental Science and Forestry,
Syracuse; William B. Preston, Manitoba Museum
of Man and Nature, Winnipeg (2); Gilbert Proulx,
Alpha Wildlife and Management Limited,
Sherwood Park, Alberta (6); Scott Redhead,
Agriculture Canada, Ottawa (2); William D. Reese,
Department of Biology, University of Southwestern
Louisiana, Lafayette; Randall Reeves, Okapi
Wildlife Associates, Hudson, Quebec (2); T. E.
Reimchen, University of Victoria, British
Columbia; Claude Renaud, Canadian Museum of
Nature; A. A. Reznicek, University of Michigan
Herbarium, Ann Arbor; Fred Scott, Wildlife
Museum, Acadia University, Wolfville, Nova
Scotia; Spencer G. Sealy, Department of Zoology,
University of Manitoba, Winnipeg; Dianne Secoy,
Biology Department, University of Saskatchewan,
Regina; Bradley Sinclair, Department of
Environmental Biology, Ontario Agricultural
College, University of Guelph; J. M. Soper,
Ottawa; K. W. Stewart, Zoology Department,

326

University of Manitoba, Winnipeg (2); Tom
Sullivan, Faculty of Forestry, University of British
Columbia, Vancouver; Edmund S. Telfer, Canadian
Wildlife Service, Edmonton; Gary J. Umphrey,
Department of Zoology, University of Western
Ontario, London; Leslie A. Viereck, Institute of
Northern Forestry, Fairbanks, Alaska (2); Marc-
Andre Villard, Department of Biology, University
of Moncton, New Brunswick; Dennis Voigt,
Ontario Ministry of Natural Resources, Maple;
Robin Walsh, Fairview, Alberta; Patrick J.
Weatherhead, Department of Biology, Carleton
University, Ottawa; Michelle Wheatley, Kanata,
Ontario (2); Heather Whitlaw, Wildlife Research
Unit, Faculty of Forestry, University of New
Brunswick, Fredericton and Phoenix, Arizona (5)

THE CANADIAN FIELD-NATURALIST

Vol. 111

[with apologies for a typo in her name in the last
Editor’s Report in the apology for one in a previous
report]; P. M. Youngman, Ottawa.

I am also indebted to David Moore, President
of the Ottawa Field-Naturalists’ Club, the Club
Council, Chairman Ron Bedford and the
Publications Committee of the OFNC for their sup-
port, to the adminstration and staff of The Canadian
Museum of Nature for allowing me to continued
use of its library and the facilities at the Macoun
Nature Centre at Aylmer, and to Joyce for continu-
ing encouragement at home throughout another
year.

FRANCIS R. COOK
Editor

Global Biodiversity: The Canadian Museum of Nature

Volume 6, Numbers 3 and 4, winter 1996 and
spring 1967, respectively, are both 48 pages, edited
by Don E. McAllister and published by Canadian
Museum of Nature. For subscription or more infor-
mation contact Global Biodiversity Business
Manager Dawn Arnold, or Subscription Manager
Susan Swan, Canadian Museum of Nature, P. O.
Box 3433, Station D, Ottawa, Ontario, Canada KOG
1RO; Phone (613) 993-5908; Fax (613) 990-0318;
e-mail darnold @mus-nature.ca.

Volume 6, Number 3, Winter 1996: The impact
of seaweed introductions on biodiversity by Robert
E. De Wreede — Schoolyard greening: The biodiver-
sity connection by Ann Coffey — Can we predict the
fluctuations of ecosystem species richness? by Marc
Proulx — Do medicinal plants have a place in today's
medicine? by N. N. Malo and R. Roy — Eco-
Products: Ecofoam, duplex printers, and energy
efficient computers — Views: Evolution will not
evolve us by Anurag A. Agrawal — A Portrait of
Biodiversity: Bombus occidentalis robbing
Ipomopsis aggregala — Biodiversity Reports:
International Coral Reef Symposium; The Second
World Congress on the Preservation and
Conservation of Natural History Collections; Global
Biodiversity Forum; IUCN-WCC; Convention of
Biological Diversity: COP-3; Biodiversity News —
Cyberdiversity — Biodiversity meetings — Book and
periodical niche — The Last Word: West Africa gets
going by Jean Lauriault.

Volume 6, Number 4, Spring 1997: Biodiversity,
ecodiversity, sociodiversity: three aspects of diver-
sity, Part 1 by Pierre Dansereau [with emphasis on
the first of the terms, subsequent issues will cover
the other two] — Lighting a sacred fire by Naigzy

Gebremedbin [from Eritrea, based on a speech to
the Ministerial section of the Conference of the
Parties of the Biodiversity Convention (COP-2) in
Jakarta in 1995] — Biodiversity and the Bible by
Calvin B. DeWitt — Impressions of the Amazon at
age 12 by Don Chao — A Portrait of Biodiversity:
Large marble butterfly, Euchloe ausonides ssp. —
Limitations of endangered species lists by Lee
Harding — Initiatives and Successes: International
Summer School on Biodiversity & Systematics; A
Charter for Earth: A code of conduct for humans in
regard to nature; A new industry: conch farming] —
Biodiversity News: British Columbian and Nova
Scotia get top marks in protecting wilderness;
Celebrating the Portuguese centennial of oceanogra-
phy; Global hazardous chemicals agreement on its
way; Population, environment, and Canadian
Policy; Canadian environmental philanthropy —
donating your lands for conservation purposes;
Canadian study on Biodiversity Law and Policy
now available; New Fish Protection Act for British
Columbia; Storm-water pollution survey has good
news and bad news; LEAD Canada now accepting
applications; Discontinuation of the ATBI [= All
Taxa Biodiversity Inventory; Costa Rica];
Petawawa National Forestry Institute closes;
Environment Canada and State of Sao Paulo under-
take watershed project; New citizen's handbook on
wetlands available now; We've moved! —
Cyberdiversity: Biodiversity and the Internet —
Biodiversity Meetings — Book and Periodical Niche
— Index Volume 6 — The Last Word: Pierre
Dansereau, the barefoot ecologist by Jacques
Prescott.

FRANCIS R. COOK

£997

Rana-Sauria

The newsletter of the Quebec Amphibian monitor-
ing program and the Atlas of the Amphibians of
Quebec, Volume 3, Number 2, December 1996. This
is the first issue prepared by David Rodrigue, who has
replaced Sylvie Mattee, the first editor of the news-
letter and co-author with Roger Bider of the first three
Quebec Atlas editions. This issue begins with “A
Word from the new coordinator’, and is followed by
“The amphibian population follow-up: four years
later; 1996 objectives; Top species for 1996”; “The
coordinator’s year end comments: The survey roads,

NEWS AND COMMENT

327

The reproductive sites”; “Urban sprawl”; “It only hap-
pens to others ...”; “Participants in the 1996 amphib-
ian population monitoring program” and “The Atlas
of Amphibians and Reptiles of Quebec”. More infor-
mation on the atlas, the monitoring programs, and the
newsletter are available from David Rodrigue, St.
Lawrence Valley Natural History Society, 21125 ch.
Ste-Marie, Ste-Anne-de-Bellevue, Quebec H9X 3L2.
Telephone: (514) 457-9449; Fax: (514) 457-0769;
e-mail ecomus @total.net.

FRANCIS R. COOK

CITES Reports: 1992 Annual Report for Canada

This CITES (Convention on International Trade
in Endangered Species of Wild Fauna and Flora)
report (Number 20) issued for Canada by the
Canadian Wildlife Service, Environment Canada,
received at The Canadian Field-Naturalist 15
January 1997, is dated “November 1996” and cov-
ers imports and exports of whole animals, plants and
their parts or derivatives for Canada in 1992. It was
prepared by Jean R. Robillard and Francois Pilon.
After a 4-page of introduction, 84 pages list transac-
tions in species designated on Appendix I and II in
six tables: Living specimens (1) exported or re-
exported; (2) imported; Dead specimens, parts and

derivatives (3) exported or re-exported; (4) import-
ed; and transactions made under (5) temporary
import/export certificates and (6) scientific
import/export certificates. In 1992, the federal gov-
ernment issued 114 Import Permits (102 in 1991),
3986 (2714) Export Permits, 95 (111) Temporary
Export/Import Certificates, and 38 (34) Scientific
Certificates. Provincial/Territorial governments
issued 10393 (3174). Much of the latter increase is
attributed to the placing of the Black Bear on
Appendix II, 11 June 1992.

FRANCIS R. COOK

Froglog: Newsletter of the Declining Amphibians Populations Task Force of the World
Conservation Union’s Species Survival Commission

Number 20, January 1997: More seed grants avail-
able — The DAPTF in Prague — Amphibian re-intro-
duction symposium report [15 August 1996 at the
Royal Botanical Gardens, Hamilton] — The status of
amphibians in Slovenia — Nyctibrachus major in
Malnad, India — Stability for Swedish treefrogs? —
Toads in the United Arab Emirates — The effect of
aquatic acidification on Rana pipiens — USA nation-
al coordinator’s Report — Froglog shorts —
Publications of interest.

Number, 21, March 1997: Anonymous donation
boosts seed grant funds — Hungarian Working Group
report and the Year of the Salamander —
Conservation meeting in Hungary — Amphibins in

Portugal — Herbicides kill frogs — 1996 IUCN Red
List — Georgia Herp Atlas — U.S. amphibian teaching
packs — Taiwan amphibian status — Report from the
Urals — Toad Mortality — Froglog shorts —
Publications of interest.

Froglog is available by writing John W.
Wilkinson, Department of Biology, The Open
University, Walton Hall, Milton Keynes, MK7 6AA,
United Kingdom; e-mail DAPTF@open.ac.uk; or on
the World Wide Web at the following URL:
http://acs-info.open.ac.uk/info/newsletters/FROGLOG.html

FRANCIS R. COOK

328

THE CANADIAN FIELD-NATURALIST

Vol. 111

Ontario Natural Heritage Information Centre Newsletter

Volume 3, Number 1, Winter 1996/1997 contains
12 pages of news items including a report on the
Ontario Ministry of Natural Resources Ecologists
Meeting 9-10 October 1996. Also featured is a
Community Ecology section: Rare Communities, an
article on Glaciere Talus at the spectacular gorges at
Ouimet Canyon and Cavern Lake Canyon northeast
of Thunder Bay. There is an announcement that a
Vegetation Community Ranks List is now available;
progress in the Mapping of Rare Vegetation
Communities from Original Land Survey Notes
Project; and an update on the Vegetation Community
Catalogue. An atlas of Ontario Dragonflies and
Damselflies database is reported as reaching 4000
records from the published literature, with compila-
tion of records from naturalists and museum collec-
tions proceeding. An Ontario perspective of

Renew Report #6: 1995-96

This bulletin, 32 pages, has been issued by the
(Canadian) Committee on the Recovery of Nationally
Endangered Wildlife. The latter involves non-govern-
mental organizations and federal, provincial and terri-
torial agencies in coordinating and promoting wildlife
conservation. It is chaired by David Brackett, Director
General of the Canadian Wildlife Service. This report
has an Executive Summary review of the year | April
1995 to 31 March 1996 of successes and failures for
species at risk, followed with individual mention
(updates of progress to date, or admissions of insuffi-
cient time to organize a committee yet) for 42 species
of those designated Endangered, Threatened or
Vulnerable by the Committee on the Status of
Endangered Wildlife in Canada (COSEWIC). There is
a separate “1995 list of Species at Risk Designated by
COSEWIC”. Also included are a section on funding
sources for RENEW, profiles of featured Team

Sea Wind

Volume 10, Number 4, October-December 1996,
contains features on Sea Reform — Counterpart to
Land; News from OVI [Ocean Voice International];
A charter for Earth: A code of conduct for humans
in regard to nature; Seahorse Conservation in the
Central Philippines: A community-based approach;
The economic benefits of tourism in the marine
reserve of Apo Island, Philippines; The impacts of
fishing gear on seafloor habitats; Coral reef conser-
vation in Indiana. A section titled Sea News
includes the startling item that the [UCN has deter-

RENEW (Recovery of Nationally Endangered
Wildlife) is given, as are 1996 Botanical highlights,
notice of the NHIC Amphibian and Reptile and Rare
Vascular Plant lists, and progress on the Trent
University Herbaria record computerization. There is
an account of the Site District Boundary Refinement
and Revision in Site Regions 6 and 7. News and
notes cover the IUCN World Conservation Congress
in Montreal in October 1996; International Visitors
at the NHIC; the 1996 A. D. Latornell Conservation
Symposium; and a New MNR Vulnerable,
Threatened and Endangered Species List. As well,
there is a focus item on the indefatigable Mike
Oldham, the botanist/herpetologist of the National
Heritage Information Section since its inception.

FRANCIS R. COOK

Leaders Diane Amirault and Doug Janz, a listing of
all Recovery Team members, a summary of the Status
of Recovery Plans, Province and Territory addresses
for more information and the 16 organizations that are
RENEW members.

The work being done is impressive and the support
credible but, arguably, a sour note is the occasional
glib plan and action outlined for “re-introduction” of
some apparently extirpated Canadian populations.
Once lost, a population may not be accurately restored
with stock obtained elsewhere, and, although this may
not be noticeable to the public, what is the point?
Such attempts may fail more often than they succeed,
and irregardless of “success” are a squandering of pre-
cious funds that are sorely needed to preserve habitats
of populations of species which have actually sur-
vived so far in Canada.

FRANCIS R. COOK

mined, based on analysis of 35 years of data from
500 scientists worldwide, that (world-wide) 5205
vertebrates are endangered: including nearly 25%
of mammals, 25% of amphibians, 20% of reptiles,
34% of fish. Concluding sections, The Booke
Nooke contains six book and journal reviews; On
the Net details on seven web sites or e-imail
addresses; and Conferences lists four upcoming
notable meetings, with two more included on the
inside back cover.

FRANCIS R. COOK

1997

NEWS AND COMMENT

329

Richards Natural History Education Award by Federation of Ontario Naturalists

The Richards Memorial Award for excellence in
Natural History Education was presented to the
Ottawa Field-Naturalists’ Club at the 65th
Anniversary Dinner of the Federation of Ontario
Naturalists in Toronto, 7 November 1996.
Accepting the big silver trophy on behalf of the
OFNC was Bill Cody. The Federation of Ontario
Naturalists was founded in 1931 to represent natu-
ralists on provincial issues. At that time it consisted
of seven nature clubs and 28 individual members.
Today it represents 83 clubs (including The Ottawa
Field-Naturalists’ Club) and associated groups and
15 000 individual naturalists. The Richards Award
is made in recognition of a special contribution to
natural history education. Mr. Norman W. J.
Richards, after whom the award is named, was a
forester who took a keen interest in improving nat-
ural history education.

nea

i: Hpk:

fii Ss

e:

The Richards Natural History Education Award
was presented to the OFNC in recognition of the
contribution made by our scientific journal, The
Canadian Field-Naturalist; the first time in the his-
tory of the award that a team rather than an individ-
ual has been recognized by it. Much of the success
of this journal over the last half of this century is
due to Bill Cody, the Business Manager for nearly
50 years, first appointed “Treasurer’s Assistant” 21
April 1947; retitled the following year) and to
Francis Cook, present and longest serving editor
(over 20 years: 1962-1966; 1981-present) who has
continued many traditions initiated by predeces-
sors, most recently (since 1942) Harold Senn, Bob
Hamilton, Ted Mosquin and Lorraine Smith. These
efforts combine with those of the Associate Editors,
outside reviewers, Editor’s and Business Manager’s
assistants, copy editor, index compilors, the sup-

The cup symbolizing the Norman W. J. Richards Memorial Award with (left and right standing) Dave Moore,
President, and Ron Bedford, Chair, Publications Committee, The Ottawa Field-Naturalists’ Club; and (seated)
William J. Cody, Business Manager, and Francis R. Cook, Editor, The Canadian Field-Naturalist.

330

RICHARDS
Bo al | ey ti ten
AWARD

Presented to:

The Ottawa Field-Naturalist Club

hac Ly {yg

In recogninon of an excepional contribution in Nazural History Educasion

Z November 7, 1996
Gin, GU

Presiden Mu Dae
/

/

FEDERATION OF
Ontario Naturalists

The Richards Education Award certificate
7 November 1996.

port of The Ottawa Field-Naturalists’s Club
Publication Committee, Council Club, member-
ship, and the authors who have submitted to the
journal. We are pleased to see these efforts offi-
cially recognized for their provincial, national and
international significance.

Two recent special issues are especially note-
worthy. They are: Volume 109, Number 3; The
History of the Exploration of the Vascular Flora of
Canada, Saint-Pierre et Miquelon and Greenland
and; Volume 110 Number 1; A Life with Birds:
Percy A. Taverner. Funding special issues requires
a departure from normal practise since much of the
revenue from page, table, figure and reprint
charges assessed authors and their institutions is
unavailable. Normally, such charges cover 40 to
50% of the cost (contributions from membership
dues usually amount to just over 10%). Upon a
recommendation by the Publications Committee,
the Council approved the withdrawal of funds
from Club financial reserves to compensate for this
loss of revenue. The Vascular Flora issue was co-
sponsored by the Missouri Botanical Garden and

THE CANADIAN FIELD-NATURALIST

Vol. 111

Recipients of the Norman W. J. Richards Memorial Award
for excellence in Natural History Education presented
annually by the Federation of Ontario Naturalists

1979 William Stirling

1980 Audrey Wilson

1981 Kenneth Strasser

1982 Anne Robertson

1983 Ron Strickland, Ron Tozer
1984 Audrey Wilson

1985 Clarke Birchard

1986 Douglas Sadler

1987 Alan Watson

1988 Robert Dorney

1989 Horace Milson

1990 —

199] Bill Wilson

1992 Sandy Symmes

1993 Jim Rule

1994 Anne Range

1995 Kittie Fells

1996 The Ottawa Field-Naturalists’ Club

the Taverner issue costs were partly recovered
through additional sales of specially hard-bound
copies.

The Federation of Ontario Naturalists instigated
the Richards Education award in 1979. This is the
17th time that it has been presented. Some previous
winners that may be recognized by Club members
are Doug Sadler of Peterborough for his books on
nature interpretation, Dan Strickland and Ron Tozer
for the interpretation programs in Algonquin Park,
Anne Robertson of Kingston for her work with
junior naturalists and Sandy Symmes for natural his-
tory interpretation with the Credit Valley
Conservation Authority. We work regularly with
Sandy who is the co-ordinator of regional councils
for the Federation of Ontario Naturalists.

FRANK POPE
Past President, The Ottawa Field-Naturalists’ Club

Minutes of the 118th Annual Business Meeting of
The Ottawa Field-Naturalists’ Club, 14 January 1997

Place and Time: Canadian Museum of Nature,

Metcalfe and McLeod Streets, Ottawa, 7:30 p.m.

Chairperson: Dave Moore, President

Attendance:

Dave Moore opened the meeting by asking mem-
bers to spend the first half hour reviewing copies of
the minutes of the previous meeting, the Treasurer’s
Report, and the reports of the Committees.

1. Minutes of the Previous Meeting
No errors or omissions were identified. It was
moved by Frank Pope (2nd Tom Reeve) that the
minutes be accepted.
(Motion Carried)

2. Business Arising from the Minutes
There was no business arising from the minutes.

3. Treasurer’s Report

Gillian Marston opened her remarks by noting the
favourable report from the Club’s Auditor. In
reviewing details of the financial statements, she
noted, among other things, a decrease in Member’s
Equity from $233,483 to $212,315 and an increase in
the de Kiriline-Lawrence Fund from $12,749 to
$15,432. In response to a question from the floor,
Gillian explained that the Fund had been established
in honour of Louise de Kiriline Lawrence to support
the conservation of natural areas, and gave several
examples of how this could be done. She noted an
increase in the cost of publishing The Canadian
Field-Naturalist, in what appears to be a trend. In
response to a question from the floor, it was agreed
that the last line of the Statement of Operations-
CEN, should read “Expenses over Income”.

It was moved by Gillian Marston (2nd Ken
Young) that the Treasurer’s report be accepted.

(Motion Carried)

4. Committee Reports

Dave Moore introduced each of the Committee
reports and asked for comments and questions. There
were none. He noted that the Executive Committee
did not meet in 1996.

It was moved by Tom Reeve (2nd Colin Gaskell)
that the Committee reports be accepted.

(Motion Carried)

Forty-two persons attended the meeting.

5. Nomination of the Auditor
It was moved by Ken Young (2nd Frank Pope)
that Janet Gehr continue as Auditor for another year.
(Motion Carried)

6. Report of the Nominating Committee

As Chairperson of the Nominating Committee,
Frank Pope presented the report of the Committee.
He explained that calls for nominations had been
placed in both The Canadian Field-Naturalist and
Trail & Landscape and that a special insert had been
included with membership renewal forms requesting
nominations for a Treasurer. No nominations were
forthcoming from these initiatives. The Committee
was unable to identify a candidate for the position of
Treasurer but will continue its effort to find one. The
Committee presented the following list of candidates
for the 1997 Council (new members are indicated
with an asterisk):

Dave Moore
Michael Murphy

President
Vice-President

Vice-President (vacant)
Recording Secretary Dave Smythe
Corresponding Secretary Lee Cairnie
Treasurer (vacant)
Other Council Members Ron Bedford
Stephen Bridgett*
Fenja Brodo
Bill Cody

Francis Cook
Ellaine Dickson
Barbara Gaertner*
Alan German

Jeff Harrison
Philip Martin*
Patricia Narraway
Isabel Nicol*
Frank Pope

Tom Reeve

Stan Rosenbaum*
Chris Traynor
Ken Young
Eleanor Zurbrigg*

Jou

332

Seven members of the 1996 Council chose not to
stand for re-election: Eileen Evans, Colin Gaskill,
Christine Hanrahan, Cendrine Huemer, Anne
MacKenzie, Gillian Marston, and Jane Topping.
Frank thanked them for their service on Council and
said that he was certain they would continue to con-
tribute in other ways.
It was moved by Frank Pope (2nd Mike Murphy)
that the proposed slate be accepted.
(Motion Carried)

7. New Business

Dave Moore informed the meeting that the Club
had received recognition for its work in the form of
two awards in 1996. In June, Jeff Harrison received
the City of Ottawa Environmental Award,
Community Group, on behalf of the Fletcher
Wildlife Garden and the Club. A framed Certificate
recording this event is now installed in the
Interpretive Centre. In October, the Club learned that
the Federation of Ontario Naturalists had awarded it
the 1996 Richards Natural History Education Award
for the Club’s work in publishing the Canadian
Field-Naturalist. Bill Cody attended the Award
Ceremony in Toronto and returned with a handsome
trophy which was displayed at the meeting and will
be in the Club’s possession for the next year.

THE CANADIAN FIELD-NATURALIST

Vol. 111

8. Presentation

Mike Murphy gave a presentation on the history
of the Birds Committee. The Club has always had
some part of itself associated with the study of birds;
an Ornithology Branch in the early years, and a Bird
Census Committee established in the late 1910s,
which subsequently evolved into the Bird Records
Committee and the Bird Feeder Committee in 1971.
In 1980, the Council approved the formation of the
present Birds Committee as one of twelve Standing
Committees of the Club. Its mandate includes the
maintenance of bird records, feeders, the “birding
Hotline”, and other birding activities. New activities
include helping the Fletcher Wildlife Garden to
organize the Taverner Cup competition. Mike con-
cluded his presentation by inviting members to con-
tact the Committee’s Chairperson, Chris Traynor, for
further information.

9. Adjournment
At 8:50 p.m., it was moved by Tom Reeve (2nd
Ellaine Dickson) that the meeting be adjourned.
(Motion Carried)

DAVE SMYTHE
Recording Secretary

Committee Reports for 1996 to The Ottawa Field-Naturalists’ Club

Awards Committee

The following awards were presented at the
Annual Soiree held on April 26, 1996:
1995 — MEMBER OF THE YEAR AWARD: Bob Bracken
for the sharing of his knowledge of the flora and
fauna of the National Capital Region on numerous
field trips.

1995 — ANNE HANES NATURAL HISTORY AWARD:
Marilyn Light for her eleven-year study of the popu-
lation dynamics of orchids in Gatineau Park.

1995 — CONSERVATION AWARD FOR MEMBER: Jeff
Harrison for his work as Chairperson of the Fletcher
Wildlife Garden Committee and his co-authorship of
the “Urban Naturalist” in the Ottawa Citizen.

1995 — CONSERVATION AWARD FOR NON—MEMBER:
Kit Chubb and the Avian Care and Research Centre
at Verona Ontario, for care and rehabilitation of
injured birds.

In addition to the above awards, the President’s
Prize was given to Dave Smythe for his work in
keeping Club records since 1991.

BILL ARTHURS

Birds Committee
The Committee continued its regular activities
including the Fall Bird Count, the Ottawa-Hull

Christmas Bird Count, the Dunrobin Christmas Bird
Count (a new responsibility), tending the Bird Status
Line and the Rare Bird Alert. It also provided leaders
for Club field trips and articles for Trail &
Landscape.

The Committee maintained bird feeders at five
locations and organized the annual Seedathon where
almost $1000 was raised to provide seed for the
feeders.

The Bird Records Subcommittee was reinstated and
met four times to review the list of rare bird reports.

In resonse to Council's request for new projects,
planning was started for a birding contest for school-
age children, to create and promote interest in bird-
ing and to foster field skills.

Certificates of appreciation were sent to members
of the community who have given generously of their
time, energy, and hospitality to Ottawa’s birders.

Members of the Committee participated in the
trial run of the Fletcher Wildlife Garden’s Taverner
Cup competition.

CHRIS TRAYNOR

Computer Management Committee

The focus of 1996 activities was on upgrading the
Club’s computer hardware. A high performance per-
sonal computer was purchased for use as a desktop

1997

publishing system for Trail & Landscape. An exist-
ing machine will be reconfigured for the Editor of
The Canadian Field-Naturalist, to provide access to
electronic mail for communication with authors
across Canada.

The Committee took advantage of the conve-
nience and efficiency of electronic mail to conduct
much of its business.

The Committee assisted in planning the eventual
integration of the accounting and membership record
systems, by providing technical support concerning
computer software. The previous manual accounting
system was moved to a computer-based package,
using the existing membership records to establish
the initial “customer” base for the new accounts.

ALAN GERMAN

Conservation Committee
The Conservation Committee represented the
OFNC in a variety of conservation and policy issues
during 1996. Members of the Committee participat-
ed in the Marlborough Forest Advisory Committee.
They continued to support enhancement of the envi-
ronmental policies of the Regional Municipality of
Ottawa Carleton (RMOC) Official Plan through par-
ticipation in focus groups such as the Wetlands
Working Group and the Rural Sounding Board.
Various submissions were made by OFNC mem-
bers to the Minister of Heritage, Parks Canada, the
National Capital Commission, committees of the
Ontario Legislature, and the Ontario Municipal
Board as well as local and regional municipalities, to
advocate the protection and enhancement of natural
areas in the Ottawa-Hull area.
MICHAEL MURPHY

Education & Publicity Committee
The following major events were attended by mem-
bers of the Club:

— Ottawa Boat & Sportsman show (first time)

— Ottawa Home Show (for Fletcher Wildlife

Garden)

— Ottawa Regional Science Fair (judging)

— Environment Week (at Statistics Canada)

The first two shows generated more publicity for
OFNC than any other activities to date due to the
size and geographical diversity of the audience.

Of special note, were the many events held at the
Fletcher Wildlife Garden Interpretive Centre, where
the Committee maintains its display as part of the
permanent infrastructure. The Committee also held
all of its meetings there since the Spring of 1996.

TOM REEVE

Excursions & Lectures Committee

The Committee met four times during 1996, to
plan 62 activities for the enjoyment and education of
Club members. The schedule included eight monthly

MINUTES OF 118TH ANNUAL BUSINESS MEETING

a>

meetings featuring a diverse array of guest speakers
and topics.

Single day bus excursions were organized to
Presqu'ile Provincial Park, Derby Hill Bird
Observatory, the Montreal Botanical Gardens,
Chaffey's Locks, Cooper Marsh (annual picnic), and
Gatineau Park. A weekend outing to observe Spruce
Grouse in Algonquin Park was again well received
by the participants.

Two indoor workshops were conducted, one on
bird identification, by sight and sound, and the other
on fern identification. The balance of the program
consisted of 43 full or half-day outings occurring
predominantly within the Ottawa district. Only one
trip had to be cancelled due to the leader's commit-
ment to an out-of-town event.

COLIN GASKELL

Executive Committee
The Executive Committee did not meet in 1996.
DAVE MOORE

Finance Committee
The Finance Committee met three times in 1996.
The main items considered were the annual accounts
and budget of the Club, the treatment of the Fletcher
Wildlife Garden in the Club’s accounting system and
a recommendation to Council concerning a donation
in kind.
KEN YOUNG

Fletcher Wildlife Garden Committee

Significant improvements to the physical environ-
ment of the Garden were made this year. The model
garden was completed. The sedge meadow has
become an amphibian pond with an island and deep
pool where eggs of four species of frogs were intro-
duced. Two major plantings were made in the new
woods. In the old woodlot, more woodland plants
were brought in and some buckthorn removed. New
signs were installed in various locations.

The Interpretive Centre was improved with the
addition of furniture, exterior security lighing, the
start of a reference library, and new exhibits. New
printed material was made available including the
first checklist of Butterflies of the Ottawa District.
Many Council and Committee meetings were held at
the Centre.

Twelve program events were conducted attracting
about 1000 people. Major events included Earth
Day, an Environment Fair, Migratory Birds Day, and
a Butterfly Gardening workshop.

On 19 May, the Bill Holland trail was dedicated,
with about 50 people including Bill's family in atten-
dance. The first Taverner Cup competition was held
on 25 May with five teams competing. It is hoped
that in future years this competition will help to fund
the Garden’s operation.

334

A grant of $2400 was received from the Friends of
the Environment Foundation, and $1000 from
Consumers Gas. For its effort in developing the
Garden, the Club, through the Garden, was awarded
the City of Ottawa's first Environmental Achievment
Award for a community based project.

JEFF HARRISON

Macoun Field Club Committee
The Committee met five times during the year to
plan the weekly schedule for the children and young
people who are members of the Club. Lectures and
workshops conducted by local naturalists and scien-
tists were successfully arranged, as were field and
camping trips, led mainly by Committee members.
The Committee is concerned by a drop in recruitment
at the youngest age levels, and notes that even if this
is corrected, a difficult gap will ripple through the
Club's age-segregated groups in the years ahead.
ROBERT LEE

Membership Committee

The total membership paid up for 1996, was 1024,
a decrease of 30 from 1995. There were 123 new
members, including 56 Family, 66 Individual and 1
Life. A detailed breakdown is given below with the
1995 figures in brackets for comparison.

Four complimentary one-year memberships were
presented to outstanding young participants in the
Annual Science Fair.

Unfortunately, in 1996 we lost two of our
Honorary members, Claude Garton, a member since
1945 and an Honorary member since 1985, and Dr.
W. G. “Bill” Dore, an active member since 1932 and
an Honorary member since the early 1970s.

This year we added four members to our “Golden
Anniversary” membership list: Dr. N. Polunin, a Life
member now living in Switzerland, and three well-
known local residents, Clarrie Frankton, Jack Gillett

THE CANADIAN FIELD-NATURALIST

Vol. 111

and Vic Solman. Again we are so fortunate to know
and learn from these delightful friends.
PATRICIA NARRAWAY

Publications Committee

The Publications Committee oversees and advises
on all aspects or the Club’s Publications. It met twice
in 1996.

Five issues of The Canadian Field-Naturalist
(CFN) were published in 1996: Volume 109, Issues
3,4 and Volume 110, issues 1,2,3. They contained
822 pages, 32 articles, 35 notes, 10 COSEWIC arti-
cles, 82 book reviews, 278 new titles, 1 commemo-
rative tribute, and 33 pages of News and Comments.

Volume 109, issue 3, was a special issue published
in cooperation with the Missouri Botanical Garden,
containing three articles: reviews of the history of the
exploration of the vascular flora of Canada, Saint-
Pierre and Miquelon, and Greenland, by James
Pringle. Volume 110, issue 1, contains a biography of
P. A. Taverner by John Cranmer-Byng entitled “A
Life with Birds: Percy A. Taverner, Canadian
Ornithologist, 1875-1947”. Preparatory work was
completed for another special issue in 1997.

All but one of the Associate Editors of the CFN,
accepted reappointment in 1996. [The retired associ-
ate Diana Laubitz, invertebrates, has retired from her
Emeritus position at the Canadian Museum of
Nature].

Volume 30 of Trail & Landscape was published in
4 issues that contained 180 pages. Popular subjects
were about birds and the Fletcher Wildlife Garden.
Two of the issues were accompanied by a single
page issue of the Green Line.

During 1996, the Club approved a further grant of
$1000 towards the publication of “The Butterflies of
Canada” by the University of Toronto Press, authors
P. Hall, D. Lafontaine, and R. Layberry. This volume
is expected to appear in the early summer of 1997.

RONALD BEDFORD

Membership of The Ottawa Field-Naturalists Club, 1996.

(subscriber totals not included, see page 323).

CANADIAN

359% (355)
13 (14)
403 (432)
camels @ lea
Bo. . 9)
199325)

188 (192)

[oo7

MINUTES OF 118TH ANNUAL BUSINESS MEETING

a35

The Ottawa Field-Naturalists’ Club Financial Statement:

for the Year end 30 September, 1996
Auditor’s Report

To: The Members of THE OTTAWA FIELD-NATURALISTS’ CLUB:

I have audited the balance sheet of The Ottawa
Field-Naturalists’ Club as at September 30, 1996,
and the statements of operations and members’ equi-
ty. These financial statements are the responsibility
of the organization’s management. My responsibility
is to express an opinion on these statements based on
my audit.

Except as explained in the following paragraph, I
conducted my audit in accordance with generally
accepted auditing standards. Those standards require
that I plan and perform an audit to obtain reasonable
assurance whether the financial statements are free of
material misstatement. An audit includes examining
evidence supporting the amounts and disclosures in
the financial statements. An audit also includes
assessing the accounting principles used and signifi-
cant estimates made by management, as well as eval-
uating the overall financial statement presentation.

In common with many non-profit organizations,
The Ottawa Field-Naturalists’ Club derives some of

its revenue from memberships, donations, and fund
raising activities. These revenues are not readily sus-
ceptible to complete audit verification, and accord-
ingly, my verification was limited to accounting for
the amounts reflected in the records of the organiza-
tion.

In my opinion, except for the effect of the adjust-
ments, if any, which I might have determined to be
necessary had I been able to satisfy myself concern-
ing the completeness of the revenues referred to in
the preceding paragraph, these financial statements
present fairly, in all material respects, the financial
position of the OFNC as of September 30, 1996, and
the results of its operations for the year then ended in
accordance with generally accepted accounting
principles.

JANET M. GEHR
Chartered Accountant
North Gower, Ontario
January 11, 1997

336 THE CANADIAN FIELD-NATURALIST

The Ottawa Field-Naturalists’ Club
Balance Sheet
September 30, 1996

1996 1995
ASSETS
CURRENT ASSETS
Gash 2 hee 240,662 270,404
Accounts Receivable ...... 14,904 9,808
Interest Receivable ......... 3,023 4,299
Prepaid Expenses... 2:5: 1,608 1,608

Fixep (Note'3)!.2.22...2..5 0 0
LAND - Alfred Bog ........ 3,348 3,348

263,545 289,467

LIABILITIES, FUNDS AND MEMBERS’ EQUITY

CURRENT LIABILITIES

Accounts Payable............. 12,418 22,800
Deferred Income .............. 13,836 12,068

26,254 34,868
FUNDS (Note 4).........:...52.. 17,976 14,616
LIFE MEMBERSHIPG............ 7,000 6,500
CLUB RESERVES). .0.ic22.2.2.00 100,000 100,000
GENERAL EQUITY ............-- M2315 133,483

263,545 289,467

The Ottawa Field-Naturalists’ Club
Statement of Members’ Equity
September 30, 1996

1996 1995
EXCESS INCOME (EXPENDITURES)
The Ottawa Field-
Naturalists’ Club.......... (3,070) 1,476
Canadian Field-Naturalist (18,015) 15,126

(21,085) 16,602

OTHER INCOME (ALLOCATIONS)
Donations - Misc. upon

membership renewal .... 3,551 3.503
Allocation to
Kiriline-Lawrence Fund (3,634) (2,719)
(83) 784
TOTAL INCOME 2. 2 (21,168) 17,386

MEMBERS’ EQulirty,
Beginning of Year........ 233,483 216,097

MEMBERS’ Egulirty,
Pune Of VAM... i500: ZZ 315 233,483

The Ottawa Field-Naturalists’ Club
Statement of Operations - OFNC
Year Ended September 30, 1996

1996
INCOME
Membersnips......205..:-s0.<.-. 14,879
T&L Subscriptions and
Back Issues. oo 186
Interest 2S. eee 2,626
Other SalésAi. ce2) ee 3,268
Special Publications......... 366
Total Income: .0csdc<seucs% 21325
EXPENSES
OPERATIONS EXPENSES
Affiliation (GCS -...<.--5-<c-2--- 2,193
Computer tcc ee 2,299
Memberships. 22 L677
Office’assistant).. 2.05 1,012
Operavions::.4.342. Soe 3,584
OFNC GST Rebate :.....2.4 778

Total Operations Expenses — 11,539

CLuB ACTIVITY EXPENSES (Net)

PRWATES cc oeece sees se eee 187
SOMeG Sue ere rer (191)
Birds 2 wes dae gee
Conservation: cy... 280
Education and Publicity ... 1,337
Excursions and Lectures .. (22)
Fletcher Wildlife Garden . 1,237
Macoun/Glubie x. ccccccccss 0c 968
Trail & Landscape............ 8,733

Total Club Activity Expenses 12,856

24,395

EXPENSES OVER INCOME... (3,070)

Vol. 111

1997 MINUTES OF 118TH ANNUAL BUSINESS MEETING 337

The Ottawa Field-Naturalists’ Club
Statement of Operations - CFN
Year Ended September 30, 1996

1996
INCOME
Baemberships...............:-... 9,878
DUOSCLIPHONS.................-+. 28,821
SG 38,699
ANNE) sedis wdaceese ensue 9,002
Publication charges.......... 17,270
BE MUMNETS.............000..- 316
Interest and exchange....... 12,429
Total Income................ TING
EXPENSES
PMSA ooo a. 2. sees 67,534
2S 5,306
APG MION ..3..c<ci.205........26- 5,646
aes sp icnssendsvsee> 2,411
Office assistant................. 5,428
Cpice SUNPIICS ............:.... 2,509
2G 84
oor 4,500
rm Gs. Rebate .:........... 2543
95,731
EXPENSES OVER INCOME...... (18,015)

1995

9,699
ZT

35,470

6,542
31,026
316
16,485

89,839

49,557
6,495
6,743
2,475
4,997
2,344

145
4,500
(2,543)

74,713

15,126

The Ottawa Field-Naturalists’ Club
Notes To The Financial Statements
September 30, 1996

1. Authority and Activities

The Ottawa Field-Naturalists’ Club is a non-profit
organization incorporated under the laws of Ontario
(1884). The Ottawa Field-Naturalists’ Club promotes
the appreciation, preservation and conservation of
Canada’s natural heritage; encourages investigation
and publishes the results of research in all fields of
natural history and diffuses information on these
fields as widely as possible. It also supports and
cooperates with organizations engaged in preserving,
maintaining or restoring environments of high quali-
ty for living things.

2. Significant Accounting Policies

Membership, subscriptions and donations are
recorded as received. All other revenues and expen-
ditures except for inventory are accounted for on an
accrual basis. Memberships are allocated to The
Canadian Field-Naturalist publication on a predeter-
mined percentage.

Supplies, records, tapes and other items held for
resale are expensed when purchased.

Fixed assets acquired after 1989 are expensed.
Fixed assets acquired prior to 1990 were recorded at
cost and depreciated on a straight line basis.

3. Fixed Assets

1996 1995
Ostet hse ee ee oe 16,746 16,746
Accumulated Depreciation..... 16,746 16,746
Net Book Valve. -2:0000.208 0 0

4. Funds

1996 1995
Seedatho tte K Sie eee 964 585
Anne Hanes Memorial Fund... 870 870
de Kiriline-Lawrence Fund.... 15,432 12,749
PRU BOS 28s ccicscadensseccasecs 451 412

5. Statement of Operations - CFN

The CEN operating deficit for the current fiscal
year was a result in part of the publication, without
charge, of a special issue.

The Problem of Invading Alien Trees and Shrubs: Some Observations
in Ontario and a Canadian Checklist

PAUL M. CATLING

8 Scrivens Drive, R.R. #3, Metcalfe, Ontario KOA 2P0

Catling, Paul M. 1997. The problem of invading alien trees and shrubs: some observations in Ontario and a Canadian
checklist. Canadian Field-Naturalist 111: 338-342.

A list is presented of 180 taxa (mostly species) of woody plants which have been introduced into Canada from another
country, or introduced from one region of Canada to another, and have been reported as spreading in their newly colonized
Canadian range. Some of these woody aliens have proven very aggressive, forming monospecific stands, displacing native
plants and animals, and changing native communities, with the ultimate result being the loss of native biodiversity. The
majority of the invading aliens are from Europe, with only 25 (14%) from within North America. They were introduced
and distributed for varying reasons including use as ornamentals, crops, wildlife enhancement plants, and sources of fibre
and drugs. Effective management requires both taxonomic, phytogeographic and ecological data. Strict restrictions on
entry and commerce and biological controls are all important considerations. A more widespread appreciation of the prob-

lem is a fundamental need.

Key Words: Biodiversity, aliens, invasions, weeds, trees, shrubs, Canada.

Some people have speculated that introduced
species are the greatest threat to global biodiversity.
“Even if the destruction of natural habitats such as
rain forests and wetlands ceased tomorrow, the insid-
ious effects of introduced species would continue to
erode the world’s biodiversity” (Clout 1995). In
Canada, alien invaders of natural habitats, such as
Purple Loosestrife (Lythrum salicaria; see Mal et al.
1992), have recently emerged as a new and very con-
spicuous class of weeds that form dense monospecif-
ic stands displacing native plants and animals and
changing native communities, with the ultimate result
being the loss of native biodiversity (ecotypes,
species and communities). Of course, they have been
present for a long time, but their harmful effects are
now becoming more obvious. Unlike the traditional
weeds of row crops, pastures and lawns, which are
mostly herbaceous plants, this newly defined group
includes a significant proportion of trees and shrubs.
A recent survey (White et al. 1993), prepared under
contract to the Canadian Wildlife Service, identified
55 important invasive aliens of natural habitats, of
which 18 (32%) were woody plants. The buckthorns
were listed among the four principal invasive aliens
of uplands, and Glossy Buckthorn (Rhamnus frangu-
la, see also Catling and Porebski 1995) was was rated
as one of the six principal invasive aliens of native
wetlands. How many of these invading woody aliens
are there in Canada and what are they? What kinds of
problems are associated with them? Is our state of
knowledge adequate to cope with the problems? The
following observations and checklist respond to these
and related questions.

Problems
Aerial photography and mapping of woodlands in
some urban (and some not so urban) areas of south-

ern Ontario may suggest some impressive forest
remnants, and a claim that a certain portion of the
landscape is “natural”. A closer look, however, may
reveal that the forests are not dominated by Sugar
Maple, but the European Norway Maple, or that
underneath a canopy of native trees is a thicket of
alien shrubs beneath which nothing else can grow. A
lack of understanding of the impact of alien trees and
shrubs can result in inappropriate assumptions about
forest quality. Such a lack of understanding has even
resulted in field surveys that have mistaken a forest
of aliens for a native forest!

The extent of woody alien domination in many
urban situations in southern Ontario has led some
biologists and planners to consider abandoning bio-
diversity protection within cities. However, special
features of urban areas, like nutrient inputs, relative-
ly high disturbance levels, pollution, and/or relative-
ly heavy “seed rain” of cultivated species, do not
appear to limit alien invasion. In fact, some inva-
sions appear to begin in urban centres and then move
out into rural areas (Catling and Porebski 1995).

Despite a great deal of attention that has been ~
focussed on old growth woodlands and wetlands,
some of the most serious problems in protectng
native biodiversity in eastern North America centre
on the decline of dry to mesic open habitats due to
both control of fires and natural succession to woody
growth. Aliens have sometimes played a major role
here in invading openings more quickly, and aggres-
sively dominating them more completely, than
would any native woody plant. European Buckthorn
(Rhamnus cathartica) is an example of a woody
alien that has become very common over large por-
tions of the southern Ontario landscape and should
be a major concern with regard to protection of
native biodiversity.

338

1287,

The contributions of conservationists and wildlife
managers to to the problems with invading aliens is a
source of embarassment. To appreciate the problem,
one need only consider the planting of Scots Pine
(Pinus sylvestris) in native open habitats that have
become thoroughly choked with this aggressive
weed tree. Plantings of Multiflora Rose (Rosa multi-
flora), Honeysuckles (Lonicera spp.), and Autumn-
Olive (Elaeagnus umbellata) in the name of
“wildlife enhancement” have contributed a great deal
to invading alien problems. In many cases, the dam-
age may have been done without much improvement
in either conservation or wildlife enhancement.
There is little evidence that exotics are generally
superior to native species (e.g., Harty 1986).

Invading woody aliens are not only a threat to the
biodiversity of natural ecosystems; some influence
Canadian economy in more direct ways. With regard
to agriculture, some of the most serious invaders of
natural ecosystems in eastern Canada, the buck-
thorns, are directly relevant as major alternate hosts
of pests, such as the aphids on potato crops, and
pathogens such as rusts on cereal crops. These buck-
thorns are covered by provincial weed acts in three
provinces. Other invading woody plants are utilized
as crops, as food and cover for wildlife, were devel-
oped for horticultural purposes, are planted for soil
improvement, erosion control, or as windbreaks.
Their naturalization is the cost of their use for these
various purposes, and represents an impact of
wildlife management, agriculture or horticulture, etc.,
on Canadian biodiversity. Biodiversity threatened by
woody invasive aliens includes native germplasm
resources and potential biocontrol agents.

Inadequacy of Information

With recent literature and herbarium collection
data, the list of taxa of alien trees and shrubs spread-
ing in Canada was increased by 96 species out of a
total of 168. Over half of these spreading aliens are
rare or local. However, the two texts providing the
basic list are relatively recent (1966-1997,
1978-1979) suggesting that alien trees and shrubs
have been neglected by botanists. It is unlikely that
the number of native plants, crop relatives, or weeds
directly impacting upon row crops, in these works
could be increased by more than 50%. How can one
requiring accurate data be confident of the informa-
tion and identification when the text used is known
to be lacking more than 50% of the possibilities,
even if the likeliest possibilities are included?

A few recent manuals (e.g., Hinds 1986) treat
woody invasive aliens comprehensively, but the spe-
cialized weed and woody plant literature provides a
remarkable contrast. The only woody plant in most
weed books was poison ivy. Various books featuring
trees and/or shrubs have at least some deficiencies in
terms of inadequate coverage. The most recent manu-

CATLING: INVADING ALIEN TREES AND SHRUBS

339

al of Canadian trees (Farrar 1995) for example, helps
much more than its predecessors because it includes
introduced and naturalized (spreading) species, but
often with relatively little information compared to
that provided for the native species covered. Despite
numerous references to the value of various trees, and
often minor values such as ornaments and carving, the
problems associated with invading alien tree species
in Canada are not well presented.

It appears that the environment, wildlife, systemat-
ics and ecology related sectors have the most com-
plete understanding of the problems associated with
invading alien trees and shrubs. Those traditionally
(and most) concerned with weeds (i.e., the agricultur-
al sector), and those most concerned with woody
plants (i.e., the forestry and horticultural sectors),
appear to have a less complete understanding.
Certainly, the available literature provides some of
the explanation for this. Although the systematics and
ecology sectors may be the most logical ones to deal
with the problem, the agriculture and forestry sectors,
in particular, have potentially important roles, espe-
cially in relation to management and control.

Solutions

Analysis of the list (Table 1) provides some useful
perspectives on management. Since the majority of
species on the list are cultivated plants, there are pos-
sibilities for some control through consideration of
alternative utilization of native species or of use of
less aggressive cultivars. Some outstanding exam-
ples of the substitution of native species for exotics
in wildlife enhancement are now available (e.g.,
Harty 1986). The majority of species on the list, and
the most serious problem species, are Eurasian, and
are more or less without close relatives in Canada.
This suggests that it is likely that the problems of
dominance could be a consequence of lack of natural
enemies in Canada, and this in turn leads to the opti-
mistic conclusion that biocontrol agents may be
found and introduced to effectively deal with the
problem. Consideration of aliens as “guilty until
proven innocent” (i.e., keeping all nonindigenous
species out of the country until they are demonstrat-
ed to be safe, Ruesink et al. 1995) appears to be a
very sensible approach.

Management of invading alien woody plants
requires both accurate and efficient means of identi-
fication and biological information. Trees and shrubs
are not as well known as many people think. If they
were, it would not have been possible to add four
new species to the native trees of Ontario over the
past 10 years. Even basic taxonomic work is inade-
quate. For example, the classification of some
groups such as the taxa of Rubus requires more
study. The hybrids of the alien Betula pendula
invading open habitats in the lower Ottawa valley,
including a wetland recently designated as signifi-

340

cant on a worldwide scale, also require further study
to enable accurate identification. European
Buckthorn has two close Asian relatives (R. davurica
and R. utilis) and is itself rather variable. Although
the potential for effective biocontrol has been identi-
fied, a sound basis for it is lacking until a field sur-
vey and taxonomic research can determine authorita-
tively the origin and identity of the plants that have
become a major component of the vegetation of
much of the eastern Canadian landscape.
Information on effects of alien woody plants on the
natural ecosystem is largely limited to anecdotal
observations and assumptions, instead of data. The
assembly and review of biological information
required for effective management is at most prelim-
inary. Although a small proportion of the spreading
alien woody plants have serious impacts on native
biodiversity, their impacts are rapidly increasing and,
it may be appropriate to deal with small problems
before they turn into much larger ones and the dam-
age is done. Probably the greatest obstacle to effec-
tive solutions however, is the lack of a more wide-
spread appreciation of the problem, and access to
satisfactory information.

The Checklist (Appendix 1)

A preliminary list of alien trees and shrubs grow-
ing in Canada without cultivation was prepared
using Boivin (1966-1967) and Scoggan (1978-1979),
and this was expanded through a survey of other
more recent literature (e.g., Anderson 1988; Douglas
et al. 1989-1994; Hinds 1986) and herbarium collec-
tion data. “Alien” plants are those species estab-
lished in parts of Canada beyond their native ranges.
“Invading” refers to the fact that these plants are
increasing (to a greater or lesser extent) in both num-
ber of individuals and area covered through natural
(unaided by humans) reproduction and dispersal. A
species was included if there was any report or evi-
dence of natural reproduction or spread, even if the
evidence was weak (as for Maclura pomifera, for
which, however, there are a number of reports of
escape in Michigan). Slightly shrubby (suffrutescent)
species such as the perennial, decumbent, Tree
Lupine (Lupinus arboreus Sims), naturalized on
southern Vancouver Island, are not included.

The current list includes 180 taxa (mostly species).
Some not included in identification keys in Canadian
literature are included in texts covering the United
States (Gleason and Cronquist 1991), and many are
included in Bailey’s (1954) manual of cultivated
plants. The majority of the species on the list are in
southern Ontario and\or southern British Columbia.
Most are introduced from Europe, with only 25
(14%) from within North America (either from the
United States, such as Robinia pseudoacacia or from
one part of Canada to another, such as Rubus alleghe-
niensis and Vaccinium corymbosum introduced to
British Columbia from eastern Canada). Although the

THE CANADIAN FIELD-NATURALIST

Vol. 111

native North American species are fewer, some have
been identified as problems (Acer negundo, Robinia
hispida). The Pea family contains more invading
North American natives than any other family. Some
of the invasive North American natives are remark-
able when one considers their distant and restricted
native ranges (e.g., Catalpa bignonioides, Maclura
pomifera, Robinia viscosa).

Literature Cited

Anderson, A. B. 1988. Shrubby Evening Primrose,
Calylophus serrulatus, adventive in Wellington County,
Ontario. Canadian Field-Naturalist 102(4): 737-738.

Bailey, L. H. 1954. Manual of cultivated plants.
MacMillan Co., New York. 1116 pages.

Boivin, B. 1966-1967. Enumération des plantes du
Canada. Le Naturaliste canadien 93: 253-274, 371-437,
583-646, 989-1063, 94: 131-157, 471-528, 625-655.

Catling, P. M. and Z. S. Porebski. 1994. The history of
invasion and current status of glossy buckthorn,
Rhamnus frangula, in southern Ontario. Canadian Field-
Naturalist 108(3): 305-310.

Clout, M. 1995. Introduced species: the greatest threat to
global biodiversity. Species 24: 34-36.

Farrar, J. L. 1995. Trees in Canada. Canadian Forest
Service and Fitzhenry and Whiteside Ltd., Markham,
Ontario. 502 pages.

Gleason, H. A. and A. Cronquist. 1991. Manual of vascu-
lar plants of northeastern United States and adjacent
Canada. New York Botanical Garden, Bronx, New
York. 910 pages.

Harty, F. M. 1986. Exotics and their ecological ramifica-
tions. Natural Areas Journal 6(4): 20-26.

Douglas, G. W., G. B. Straley and D. Meidinger.
1989-1994. The vascular plants of British Columbia. 4
parts. British Columbia Ministry of Forests, Victoria.

Hinds, H. R. 1986. Flora of New Brunswick. Biology
Department, University of New Brunswick, Fredericton.
885 pages.

Mal, T. K., J. Lovett-Doust, L. Lovett-Doust and G. A.
Mulligan. 1992. The biology of Canadian weeds. 100.
Lythrum salicaria. Canadian Journal of Plant Science
72: 1305-1330.

Meikle, R. D. 1985. Nomenclatural notes on some willow
hybrids. Watsonia 15(3): 273-274.

Ruesink, J. L., I. M. Parker, M. J. Groom and P. M.
Kareiva. 1995. Reducing the risks of nonindigenous
species introductions. Bioscience 45(7): 465-477.

Scoggan, H. J. 1978-1979. The flora of Canada. 4 parts.
National Museum of Canada, Ottawa. 1711 pages.

White, D. J., E. Haber and C. Keddy. 1993. Invasive
plants of natural habitats in Canada. Canadian Wildlife
Service. 121 pages.

Voss, E. G. 1985. Michigan flora, part II, Dicots
(Saururaceae- Cornaceae). Cranbrook Institute of
Science Bulletin 59. 724 pages.

Received 14 February 1996
Accepted 28 October 1996

Appendix 1.

List of invading trees and shrubs in Canada, including
both native North American species (preceded by “+’)
introduced and spreading beyond their native range in

1997 CATLING: INVADING ALIEN TREES AND SHRUBS 341

Canada, and alien species (from outside North America),
based on Boivin (1966-1967) and Scoggan (1978-1979) as
well as consideration of more recent literature and herbari-
um specimens. The list is intended to include all species for
which there is some evidence of introduction and natural
spread. Asterisks (*) indicate invasive aliens of natural
habitats identified by White et al. (1993).

PINACEAE
Larix decidua Mill., EUROPEAN LARCH
Picea abies (L.) Karst., NORWAY SPRUCE
*Pinus sylvestris L., SCOTS PINE
Pinus mugo Turra, MUGHO PINE
Pinus nigra Arnold, AUSTRIAN PINE
Pinus virginiana Miller, SCRUB PINE

SALICACEAE

Populus alba L., WHITE POPLAR

Populus x berolinensis (Regel) Dippel (P. nigra var.
italica x laurifolia)

Populus x canadensis (P. deltoides x nigra), CAROLINA
POPLAR

Populus x canescens (Aiton) Smith (P. alba x tremula)

+Populus deltoides Bartr. ex Marsh (including the western
ssp. monilifera (Ait.) Eckenw. and the eastern ssp. del-
toides and intermediates), COTTONWOOD

Populus x heimburgeri B. Boivin (P. alba x tremuloides)

Populus nigra L., BLACK POPLAR (including the
Lombardy Poplar, var. italica Du Roi)

Salix alba L., WHITE WILLOw (including var. alba with
silky-hairy branchlets, var. calva Mey. with brown
glabrate branchlets, and var. vitellina Stokes with yellow
glabrate branchlets)

Salix capraea L., GOAT WILLOW

Salix cinerea L., GREY WILLOW

Salix daphnoides Villars, WILLOW

Salix elaeagnos Scop., WILLOW

Salix fragilis L., CRACK-WILLOW

Salix x jesupii Fern. (S. alba x lucida), JEsup’s WILLOW

Salix myrsinifolia Sal. (S. nigricans), WILLOW

Salix x pendulina Wenderoth (S. babylonica x fragilis),
WILLOW

Salix pentandra L., BAY-LEAVED WILLOW

Salix x rubens Schrank (S. alba x fragilis), WILLOW

Salix x sepulcralis Simonkai (S. alba var. vitellina x
babylonica), WEEPING WILLOW

Salix purpurea L., BASKET-WILLOW

Salix viminalis L., OSIER WILLOW

MYRICACEAE
+Myrica californica C. and S., WAX MYRTLE

BETULACEAE
*Alnus glutinosa (L.) Gaertn., BLACK ALDER

Betula alba L. (B. pubescens), EUROPEAN WHITE BIRCH
*Betula pendula Roth., EUROPEAN BIRCH

FAGACEAE
Quercus robur L., ENGLISH OAK

ULMACEAE
Ulmus glabra Huds., WYCH ELM
Ulmus procera Salisb., ENGLISH ELM
Ulmus pumila L., SIBERIAN ELM

MORACEAE
+Maclura pomifera (Raf.) Schneid., OSAGE ORANGE
* Morus alba L., WHITE MULBERRY

CANNABINACEAE
Humulus lupulus L. var. lupulus, Hop

BERBERIDACEAE
Berberis thunbergii DC., JAPANESE BARBERRY
Berberis vulgaris L., COMMON BARBERRY
Berberis x ottawensis Schneid., HYBRIDE x BARBERRY
+Mahonia repens (Lindley) G. Don, MAHONIA

GROSSULARIACEAE
Philadelphus coronarius L., MOCK-ORANGE
Philadelphus inodorus L., MOCK-ORANGE
Philadelphus pubescens Loisel, MOCK-ORANGE
Ribes diacanthum Pallas, SIBERIAN CURRANT
Ribes grossularia L., GOOSEBERRY

+Ribes missouriense Nutt., MISSOURI GOOSEBERRY
Ribes nigrum L., BLACK CURRANT

+Ribes odoratum Wendland f., BUFFALO CURRANT
Ribes rubrum L. (R. sylvestre, R. sativum), EUROPEAN RED

CURRANT

Ribes uva-crispa L., GARDEN GOOSEBERRY

ROSACEAE
+Aruncus dioicus (Walter) Fern., GOAT’S BEARD
Cotoneaster acutifolia Turcz., COTONEASTER
Cotoneaster bullatus Boiss., BULLATE-LEAVED
COTONEASTER
Cotoneaster horizontalis Decne., CREEPING COTONEASTER
Cotoneaster melanocarpa Lodd., COTONEASTER
Cotoneaster simonsii Bak., SIMON’S COTONEASTER
Crataegus laevigata (Poiret) DC. (C. oxyacantha),
ENGLISH HAWTHORN
Crataegus monogyna Jacq., ENGLISH HAWTHORN
Cydonia oblonga Miller (Pyrus cydonia), QUINCE
Malus baccata L., SIBERIAN CRAB
Malus pumila Miller (M. sylvestris, Pyrus malus), APPLE
Malus prunifolia Willd., CHINESE APPLE
Prunus avium L., SWEET CHERRY
Prunus cerasifera Ehrh., CHERRY PLUM
Prunus cerasus L., SOUR CHERRY
Prunus domestica L., DAMSON PLUM (including var.
domestica with slightly pubescent or glabrous twigs and
var. insititia (L.) Poir. with densely pubescent twigs)
Prunus laurocerasus L., CHERRY-LAUREL
Prunus mahaleb L., MAHALEB CHERRY
Prunus padus L., EUROPEAN BIRD-CHERRY
Prunus persica (L.) Batsch, PEACH
Prunus spinosa L., SLOE
Prunus tomentosa Thunb., MANCHU CHERRY
Pyrus communis L., PEAR
Rosa canina L. var. canina, DOG-ROSE
Rosa centifolia L., CABBAGE-ROSE
Rosa cinnamomea L., CINNAMON ROSE
Rosa eglanteria L., SWEET-BRIER
Rosa hugonis Hemsley, HUGO ROSE
Rosa micrantha J. E. Smith, ROSE
*Rosa multiflora Thunb., MULTIFLORA-ROSE
Rosa odorata Sweet, TEA-ROSE
Rosa pumpinellifolia L., ROSE
Rosa rubiginosa L., ROSE (possibly better combined with
R. eglanteria)
Rosa rugosa Thunb., JAPANESE ROSE
Rosa spinosissima L., BURNET-ROSE
Rosa tomentosa Sm., DOWNY ROSE
+Rubus allegheniensis T. C. Porter, ALLEGHENY
BLACKBERRY
Rubus caesius L., EUROPEAN DEWBERRY

342

*Rubus discolor Weihe and Nees (R. procerus),
HIMALAYAN BLACKBERRY
Rubus idaeus L. var. idaeus, RASPBERRY
Rubus illecebrosus Focke, STRAWBERRY-RASPBERRY
Rubus laciniatus Willd., EVERGREEN BLACKBERRY
Sorbaria sorbifolia (L.) A. Br., FALSE SPIRAEA
Sorbus aucuparia L., EUROPEAN MOUNTAIN-ASH
Spiraea chamaedryfolia L., SPIRAEA
Spiraea corymbosa Raf., SPIRAEA
Spiraea japonica L. f., JAPANESE SPIRAEA
Spiraea salicifolia L., SPIRAEA
Spiraea thunbergii Siebold ex Blume, THUNBERG’S SPIRAEA

FABACEAE
+Amorpha fruticosa L. var. fruticosa, FALSE INDIGO
*Caragana arborescens Lam., CARAGANA

Caragana frutex (L.) Koch, RUSSIAN PEA-SHRUB
+Cladrastis kentuckea (Dum.-Cors.) Rudd. (C. lutea),

YELLOW-WoOoD

Colutea arborescens L., BLADDER SENNA
*Cytisus monspessulanus L., BROOM

Cytisus scoparius (L.) Link, SCOTCH BROOM
+Gleditsia triacanthos L., HONEY-LOCUST

Genista tinctoria L., DYER’S GREENWEED

Laburnum anagyroides Medic., GOLDEN CHAIN TREE
+*Robinia hispida L., BRISTLY LOCUST
+Robinia luxurians (Dieck) Schneid., LOCUST
+Robinia pseudo-acacia L., BLACK LOCUST
+Robinia viscosa Vent., CLAMMY LOCUST

Ulex europaeus L., GORSE

RUTACEAE
Ruta graveolens L., RUE

SIMAROUBACEAE
Ailanthus altissima (Mill.) Swingle, TREE-OF-HEAVEN

AQUIFOLIACEAE
Ilex aquifolium L., ENGLISH HOLLY

CELASTRACEAE
Celastrus orbiculatus Thunb., ORIENTAL BITTERSWEET
Euonymus alata (Thunb. Siebold, SPINDLE-TREE
Euonymus europaea L., SPINDLE-TREE
Euonymus fortunei (Turcz.) Hand.-Mazz., SPINDLE-TREE

ACERACEAE
Acer ginnala Maxim. (A. tatarica L.), AMUR MAPLE
+*Acer negundo L., MANITOBA MAPLE (including var.
interius (Britt.) Sarg. with velvety-pubescent twigs and
var. negundo with glabrous twigs)
*Acer platanoides L., NORWAY MAPLE
Acer pseudo-platanus L., SYCAMORE MAPLE

HIPPOCASTANACEAE
Aesculus hippocastanum L., HORSE-CHESTNUT
+Aesculus glabra Willd., OHIO BUCKEYE

RHAMNACEAE

*Rhamnus cathartica L., COMMON BUCKTHORN
Rhamnus davurica Pall., BUCKTHORN

*Rhamnus frangula L., GLOSSY BUCKTHORN

TILIACEAE
Tilia cordata Miller, SMALL-LEAVED LINDEN
+Tilia heterophylla Vent., WHITE BASSwooD
Tilia x vulgaris Hayne (T. cordata x platyphyllos),
LINDEN

THE CANADIAN FIELD-NATURALIST

Vol. 111

MALVACEAE
Lavatera thuringiaca L., TREE-MALLOW
Lavatera trimestris L., TREE-MALLOW

THYMELAEACEAE
Daphne mezereum L., DAPHNE

ELAEAGNACEAE
Elaeagnus angustifolia L., RUSSIAN OLIVE
Elaeagnus umbellata Thunb., AUTUMN-OLIVE
Hippophae rhamnoides L., SEA-BUCKTHORN

ONAGRACEAE
+Calylophus serrulatus (Nutt.) Raven, SHRUBBY EVENING
PRIMROSE

ARALIACEAE
Aralia spinosa L. (or more likely A. elata (Mig.) See.) ,
DEVIL’ S-WALKING-STICK
* Hedera helix L., ENGLISH Ivy

ERICACEAE
Calluna vulgaris (L.) Hull, HEATHER
+Vaccinium corymbosum L., HIGHBUSH BLUEBERRY

OLEACEAE
Fraxinus excelsior L., EUROPEAN ASH
Ligustrum vulgare L., PRIVET
*Syringa vulgaris L., LILAC

APOCYNACEAE
Vinca major L., LARGE PERIWINKLE
*Vinca minor L., COMMON PERIWINKLE

SOLANACEAE
Lycium barbarum L. (L. halimifolium), MATRIMONY-VINE
Lycium chinense Miller
Solanum dulcamara L., NIGHTSHADE

BIGNONIACEAE
+Catalpa bignonioides Walt., CATALPA

CAPRIFOLIACEAE

Lonicera x bella Zabel, (L. morrowii x tatarica), PRETTY

HONEYSUCKLE

Lonicera caprifolium L., WOODBINE

Lonicera chrysantha Turcz., HONEYSUCKLE

Lonicera etrusca Santi, ETRUSCAN HONEYSUCKLE

Lonicera japonica Thunb., JAPANESE HONEYSUCKLE

Lonicera maackii (Rupr.) Maxim., MAACK’S

HONEYSUCKLE

Lonicera morrowii Gray, MORROW’S HONEYSUCKLE

Lonicera xnotha Zabel., NOTHA HONEYSUCKLE

Lonicera periclymenum L., WOODBINE HONEYSUCKLE
+Lonicera prolifera (Kirchn.) Rehd., GRAPE- HONEYSUCKLE
+Lonicera sempervirens L., TRUMPET-HONEYSUCKLE
*Lonicera tatarica L., TATARIAN HONEYSUCKLE

Lonicera xylosteum L., HONEYSUCKLE

Sambucus ebulus L., DWARF ELDER

Sambucus nigra L., EUROPEAN ELDER
+Symphoricarpos orbiculatus Moench, INDIAN-CURRANT

Viburnum lantana L., WAYFARING TREE

Viburnum opulus L. var. opulus, GUELDER-ROSE

ASTERACEAE
+Iva frutescens L. var. oraria (Bartlett) Fern and Griscom,
MARSH-ELDER

Excluded

Salix babylonica L., WEEPING WILLOW (most reports of this
are likely referable to S. x sepulcralis Simonk. (S. alba
var. vitellina x babylonica, see Meikle 1985)

Book Reviews

ZOOLOGY

The Minds of Birds

By Alexander F. Skutch, with line drawings by Dana
Gardner. 1996. Texas A&M University Press, College
Station. 200 pp., illus. U.S. $29.95.

A book by Alexander Skutch is always a delight
to read. With years of meticulous and intimate obser-
vation of an avifauna that most of us can only expe-
rience in brief, tantalising glimpses, his perspective
is a highly individual one. He is at his best when
describing those observations, but in this book he
tackles a larger and very challenging topic, the work-
ings of birds’ minds. Like his other books, this vol-
ume is full of anecdotal accounts of bird behaviour,
many drawn from the literature and some from his
own observations. These have been selected to pro-
vide an overview of the enormous range of activities
that birds are capable of, and by inference, to pro-
vide insight into their mental abilities.

His thesis is that birds are far less stupid than most
people think. He suggests that they are “feeling crea-
tures whose own capacity for enjoyment increases
the total value of life on Earth”, and argues in favour
of a certain amount of anthropomorhism in interpret-
ing animal behaviour. There are 17 chapters, dealing
with various aspects of birds’ mental abilities: these
include Emotions, Play, Aesthetic Sense, Mental
Conflicts, Intelligent Birds, and Apparently Stupid
Behaviour. And indeed, many of the anecdotes do
suggest “intelligence” and consciously purposeful
behaviour, as with the Blue Jay that tore a strip of
paper from the floor of its cage and used it to poke
between the bars to bring food in the adjoining cage
within reach. Skutch favours such anecdotal evi-
dence, as he feels that structured experiments often
remove the element of spontaneity that may reveal
the capacity for original thought in a bird.

The difficulty with all this is that the real objec-
tions to anthropomorphism in the first place were that
it sometimes led to conclusions that more careful
analysis revealed to be wrong. There is certainly a
case to be made for interpretations being too mecha-

nistic (although perhaps more formerly than today),
but even on Skutch’s own terms I had difficulty with
some of his arguments. To give one example, if
through sexual selection birds have evolved an aes-
thetic sense akin to our own, as he postulates, how
does one account for such aesthetically unprepossess-
ing species as crows? To say that crows are aestheti-
cally pleasing to other crows would seem to destroy
the rest of the argument, and to argue that crows lack
aesthetic sense just as some persons do, sidesteps the
issue, even if it is legitimate to compare differences
between individuals of a species with differences
between species. Surely birds are a world of their
own, in the last analysis unknowable in the human
terms that Skutch espouses. He mounts an eloquent
case for birds as intelligent, purposeful beings, but I
suspect his views on such matters will be unlikely to
convince anyone who is not already convinced.

This, then, is not a book for the ornithologist seek-
ing an analysis of the latest in research on birds’ men-
tal capacities, and those who view their birds in these
rigorous terms could well find it irritating. For the
rest of us, Skutch has provided a concise summary of
birds’ abilities and of the incredible variety of their
behaviours, and in this he has succeeded admirably.
Anyone looking for a readable and thought-provok-
ing overview of birds’ abilities should find it fascinat-
ing. Indeed, even the skeptic might well be impressed
by this concise picture of the entire rich tapestry of
bird behaviour. You may not always agree with the
author’s views but they are always stimulating, and
provide a balance to the rather mechanistic views of
bird behaviour that prevail elsewhere. Skutch has
unique insights into these fascinating creatures that
delight and intrigue so many of us, and his ideas
deserve thoughtful consideration.

CLIVE E. GOODWIN

1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8

Martens, Sables, and Fishers: Biology and Conservation

Edited by S. W. Buskirk, A. S. Harestad, M. G. Raphael,
and R. A. Powell. 1994. Comstock Publishing
Associates, Cornell University Press, Ithaca, New York.
484 pp., illus. U.S. $65.00

Never before has such a synthesis of available
information and current research been compiled for

the genus Martes. As stated in the book’s preface, the
objectives of the editors were to provide a compre-
hensive review of our current knowledge of martens,
sables, and fishers; identify areas where information
is lacking; and suggest areas for additional research.
The authors have unequivocally met these objectives.

343

344

This book is an extension of the “Symposium on
the biology and management of martens and fishers”
held in Laramie, Wyoming, in 1991. All but three of
the chapters were based on presentations made at
this symposium. Although only one third of the pre-
sentations appear as chapters, those that do represent
a comprehensive cross-section of Martes biology
and conservation.

Martens, Sables, and Fishers consists of an intro-
ductory chapter followed by an additional 29 chap-
ters divided into seven major parts. These parts
include evolution and biogeography, population
ecology, management of populations, status and con-
servation of Holartic Martes, habitat ecology, man-
aged habitats, and physiology and reproduction. sev-
eral of the chapters emphasize original research
although the majority are comprehensive reviews of
the chosen topic. Use of tables and figures is not

Chickadees, Tits, Nuthatches & Treecreepers

By Simon Harrap and David Quinn. 1995. Princeton
University Press, Princeton, New Jersey. 464 pp., illus.
$49.50.

Each new volume that treats a worldwide group of
birds following the “Harrison” style (Harrison 1983)
seems to rise a notch in quality over its predecessors.
This latest contribution is no exception. The text is
scholarly and the art work first rate. It covers all the
species and subspecies of the world’s chickadees
(or tits), true nuthatches, treecreepers, and the
Wallcreeper.

A couple of weeks ago I suggested to a friend that
the Black-capped Chickadee was truly a Canadian
bird. It is always cheerful, gets along well with oth-
ers, withstands heat or cold, is always friendly, and
is generally ignored - truly a Canadian! This impish
little bird and its cousins are ignored no more; they
have a book of their own.

My first impression flipping through the book was
how many of the species are similar. The Black-
capped Chickadee has close relatives in the Marsh,
Sombre, Caspian, Willow, and Songer tits and
Carolina Chickadee. These are all small, perky
black, grey and white birds with a strong superficial
resemblance. True, they differ in fine points of
plumage, song, behaviour, and sometimes in distrib-
ution. Often the extremes of differences in a single
species (i.e., the subspecies) are more substantial
than between separate species. To keep this problem
in proper perspective, it is essential for the user to
read the author’s notes on splitting, lumping, sub-
species, and superspecies before consulting the
species accounts. Hybrids also occur and are dealt
with by the author, including an illustration if appro-
priate.

THE CANADIAN FIELD-NATURALIST

Vol. 111

excessive and augments the text well. The book con-
tains an incredibly comprehensive bibliography and
a functional index.

This book is not for the casual reader; rather, it is
a scientific treatise of our current knowledge of
Martes. Contributors to this holistic treatment repre-
sent a group of dedicated wildlife biologists, many
who have devoted much of their career to study this
group of carnivores. Martens, Sables, and Fishers
will surely become a classic reference because of its
broad appeal to a variety of individuals with interests
in wildlife biology and forest ecology.

JERROLD L. BELANT

U.S. Department of Agriculture, National Wildlife Re-
search Center, 6100 Columbus Avenue, Sandusky, Ohio
44870

The book covers 110 species in a series of very
thorough species accounts. Each account follows the
now-standard format of identification, sex/age,
voice, distribution, habitat population habits, biolo-
gy, description, moult, geographical variation, rela-
tionships, and references used. This generally results
in an exhaustive and detailed narrative for the wide-
spread and well known species. The text is less
detailed for the few species that are range- or politi-
cally-isolated. In general, this group of birds is wide-
spread and relatively common. Only a few species
(or are they subspecies?) are rare and these are in
remote and less accessible areas. For example, the
White-browed Nuthatch is endemic to Mount
Victoria, Myamar (Burma). Is this a geographical
variation of the common White-tailed Nuthatch of
the adjacent Himalayas? Is it more widespread than
previously thought? These are difficult questions to
research in a politically repressive region. Each
species has a range map, with additional detailed
maps added to the text, if warranted. A small, but in
my Opinion an important item, included by the
author is an explanation of the terms used to describe
colour. I long ago learnt that others use very differ-
ent words from me to describe a specific shade of
colour. In this instance the most different use of
words is for “flesh” - described by the author as the
colour of raw salmon.

The text is supported with illustrations by David
Quinn, one of the artists who illustrated the recent
“Warblers” volume (Curson et al. 1994). It is difficult
for me to believe that the over 500 full representa-
tions and numerous head, tail, and flight details can
be bettered. He has captured the ‘jizz’ correctly, pro-
vided anatomical precision and maintained a lively

1997

artistic feeling. While I have been impressed by the
art work in many recent books I think Quinn has that
extra-something that makes him the new Archibald
Thorburn (Thorburn was the outstanding illustrator of
wildlife in Europe in the 1900s). My only surprise
was Quinn’s positioning of Coal and Spot-winged
Tits on the plate. I would have thought it advisable to
have at least one or two heads turned to show the dis-
tinctive head (or nuchal) spot. This feature is present
but is difficult to discern without close inspection.
This book is a fine addition to the enthusiastic
birder’s library and should solve many gift problems.
Backyard feeder watchers should also consider buy-
ing it as a reference to those species that are the most
reliable of our small bird companions. Whether a
fanatic or casual observer the purchaser will derive
much insight and, certainly, tremendous pleasure in

BooK REVIEWS

345

owning this fine new book. For me it has raised
some new challenges. In future, I will be trying to
sex nuthatches and, in the appropriate locations, I
will be looking for hybrid chickadees. It has also
reminded me of my failure to date to find a Wall-
creeper in Europe, something I must rectify soon.

References

Harrison, Peter. 1983. Seabirds - an identification guide.
Houghton Mifflin Company, Boston. 448 pages.

Curson, J., D. Quinn, and D. Beadle. 1994. Warblers of
the Americas. Houghton Mifflin Company, Boston. 252

pages.

Roy JOHN

#1 613 McPherson Avenue, Saskatoon, Saskatchewan S7N 0X7

Larvae of the North American Caddisfly Genera (Trichoptera)

By Wiggins, Glenn B. 1996. 2nd edition. University of
Toronto Press, Toronto. 457 pp., illus. $110.00.

The text of the second edition has been virtually
rewritten with very few paragraphs remaining
unchanged from the 1977 edition, although the same
general style is retained, as well as the original illus-
trations.

The classification (pp. 8—13) is based on phyloge-
netic evidence, whereas the families and genera are
arranged alphabetically, consequently the page refer-
ences given appear to be “jumbled”. The Trichoptera
are currently divided into three suborders and six
superfamilies. Eight families and 13 genera are
added. The arrangement of families and genera may
be objectional to some readers, however, as pointed
out in the Introduction, this book is primarily a refer-
ence work.

In general, for each genus, there is a page of text
giving distribution and species (general distribution

and number of species worldwide and for North
America), morphology, case or retreat, biology, and
remarks. On the opposite page are excellent draw-
ings of the mature larva, dorsum of head and thorax,
legs, anal leg, case if applicable, etc. It is gratifying
to read that of the 149 genera recognized in North
America, the larvae of at least one species are known
for all but four genera.

The number of references has nearly doubled,
which is indicative of activity among caddisfly
workers. This, plus the fact that eight families and 13
genera are added, validates the need for this revision.
The book is highly recommended and is worth the
price.

EDWARD C. BECKER

Crop Protection Program, Eastern Cereal and Oilseed
Research Centre, Agriculture and Agri-food Canada, Ottawa,
Ontario K1A 0C6

The Scientific Basis for Conserving Forest Carnivores: American Marten,
Fisher, Lynx, and Wolverine in the Western United States

Edited by Leonard F. Ruggeriero, Keith B. Aubrey, Steven
W. Buskirk, L. Jack Lyon, and William J. Zielinski.
1994. General Technical Report RM-254. Fort Collins,
Colorado, U.S. Department of Agriculture, Forest
Service, Rocky Mountain Forest and Range Experiment
Station, Fort Collins, Colorado. 184 pp. + map.

The reason for this compilation of information is to
assess the scientific basis for conservation of forest
carnivores. Extensive literature reviews for each of the
four species are presented with a discussion of man-

agement considerations and information needs. The lit-
erature reviews for each species are sufficiently read-
able to provide even lay readers with exciting general
information and yet sufficiently thorough to provide
scientists with useful detail and outlines of the avail-
able data. Although the geographical focus of this
book is the northwest states, much of the information
is reasonably transferable to other parts of their ranges.

One of the most important features of this volume
are the list of literature cited in each of the chapters.

346

The writers of the species’ accounts have impressive
credentials and have done standout service in gather-
ing extensive detailed information, much of it from
obscure and unpublished sources.

The scientists involved in the evaluation of the
scientific information found, not surprisingly, that
there are “substantial gaps” in the knowledge neces-
sary for assessing conservation status. For example,
they recognize that the much of what we know about
the Wolverine or Lynx is largely based on studies
conducted in Canada or Alaska. The authors caution
readers that other weaknesses in the current informa-
tion include high variability in the data, unreliability,
and the difficulty and expense of research. As a
result, the useful knowledge base for developing the
conservation strategies for forest carnivores is
extremely limited.

Following the species accounts, the authors have
included a chapter on the scientific basis for con-
serving forest carnivores and another on information
needs and a research strategy, all designed to prevent
the decline of their populations. The appendices
include distribution maps for each species in the
northwest from 1961 to 1982 and from 1983 to
1993, and a brief review of a system to gather data
on forest carnivores in National Forest system lands.

The authors recognize the paramount importance of
habitat quality in regulating the populations of forest
carnivores. They conclude that mesic late-succession-
al forest stands are vital in forest carnivore conserva-
tion. They further emphasize some of the more press-
ing research needs, including dispersal and migration
and the effect on them of habitat quality and land-
scape. One of the most important characteristics of
these forest carnivores is the large geographic area

The Last Panda

By George B. Schaller. 1993. University of Chicago
Press, Chicago. 291 pp. U.S. $13.95.

For some people, learning about high-profile con-
servation and rescue operations can produce feelings
of comfort and relief: Someone is Saving a Species
from Extinction. These projects can soothe the con-
science of those not directly involved, and many of
them contribute financially to worthwhile conserva-
tion projects.

The glamour projects which pull in substantial
contributions tend to be those involving the attrac-
tive, probably cuddly, and photogenic species, a
description the panda fulfills to a T.

“...but the panda is one of those endangered

species which can be a symbol to change our
concept of ourselves, a new design in the strategy
of human survival” (page xvi).

THE CANADIAN FIELD-NATURALIST

Vol. 111

over which they range, although little is known about
it. The authors would like to see more research on
how habitat is saturated with home ranges, how these
vary within the sexes of the animal, and how home
ranges and habitat attributes are related.

There are also political and legal complications to
this scientific situation. A story in the 18 February
1996 Los Angeles Times noted that a number of
environmental and animal-protection groups have
sued the U.S. Fish and Wildlife Service and
Department of the Interior for refusing to add
the Lynx to the Endangered Species List.
Environmentalists contend that senior Fish and
Wildlife Service and Department of the Interior offi-
cials are refusing to list it as a means of avoiding the
hassle this might trigger from conservative lawmak-
ers from the west. Senior officials and the
Washington headquarters claimed that “there was a
lack of conclusive evidence of biological vulnerabil-
ity or real threats to the species in the contiguous 48
states.” The Wolverine and Fisher have also been
recommended for listing under the Endangered
Species Act.

In any case, the authors have done a commend-
able job to make clear the absolute necessity of con-
tinuing studies of rare forest carnivores. Effective
conservation strategies for them will be needed to
insure their persistence. We need to hope that mea-
sures such as these will be successful.

MICHAEL M. J. Morris

Mono Cliffs Outdoor Education Centre, R. R. #1, Orange-
ville, Ontario LOW 2Y8

China would seem to have devoted considerable
resources to protect its appealing pandas, and vast
sums of money and resources have been poured into
the projects by China and by wildlife organizations,
primarily World Wildlife Fund (WWE), and the New
York Zoological Society (NY25). George Schaller,
Director of the NYZS Conservation Division,was
appointed by WWF to head the panda project in 1980.
Schaller has already written The Great Pandas of
Wolong describing the initial scientific investigation.
This present account is largely about the behind-the-
scenes political and financial ramifications which
dominated the project and which left the panda’s wel-
fare lagging far down the priority list. He is a good
writer, and by the words he chooses conveys far more
than what is actually written. He does describe some
of his own field surveys which contributed to the
WWE Plan for Panda Conservation published in

1997

1989, but this is not the main theme. His recommen-
dations to sustain and perhaps increase panda popula-
tions, the WWF-Chinese government negotiations
and the eventual use of the considerable financial
investment and equipment, are interesting insights
into Panda Politics, and by extension some other
wildlife projects. Very often the state-of-the art equip-
ment was under used or not used at all. The greatest
threat to pandas is People: Pandas require large tracts
of different species of bamboo in order to flourish and
breed. Interference in the ecosystem by deforestation
and new settlements in the panda ranges is one prob-
lem; add to that capture by hunters (for pelts, meat,
and body parts, for which there is a large market) and
by researchers; and the periodic death of the bamboo.
These factors have had appalling effects on popula-
tions - which were none too large to begin with.
Loans of pandas to western zoos for display, and
which fill zoo coffers, bring in needed foreign curren-
cy, but deprive wild populations of breeding-age ani-
mals. The policy of capturing pandas, ostensibly for
breeding under controlled conditions, has been mostly
unsuccessful.

BooK REVIEWS

347

Schaller’s forecast for the future is sanguine. Wild
panda populations are barely surviving and the sci-
entists’ recommendations have not been acted on.
One such recommendation is to link the separated
reserves in Sichuan into one large one by creating
corridors between the panda ranges. This could be
the last hope for that area, but there is a lack of
cooperation between the various agencies which are
responsible for the ranges: the Chinese Ministry of
Forest & Development, The Environmental
Protection Office, and The China Association of
Zoological Gardens. These agencies, and others have
their own bureaucracy and agenda. China is not
unique.

This is a very interesting inside look at the diffi-
culties of co-operative conservation between two
different cultures. The lessons learned, though, can
be applied universally. Highly recommended if you
really care about all endangered species.

JANE E. ATKINSON

255 Malcolm Circle, Dorval, Quebec H9S 1T6

The Birds of Great Slave Lake, Northwest Territories, Canada
By Jacques Sirois. 1994. Ecology North, Yellowknife, N.W.T. 36 pp. $3.00

The Birds of the Northwest Territories

By Jacques Sirois and Doug McRae. 1994. Canadian Wildlife Service, Yellowknife, N.W.T. 28 pp. Free.

These two pamphlets share several features — a
common author, a physical description and map
detailing the areas they cover, a description of sites in
which to birdwatch, precautions to take while birding,
a short but adequate introduction of the literature per-
taining to their respective regions, a Canadian
Wildlife Service address for further information or to
report sightings, an indication of abundance and status
of species in their areas of coverage, and a checklist of
confirmed and unconfirmed bird species.

Because it covers Great Slave Lake and a 50-km
area around the lake the first booklet contains more
details than the general work on the Northwest
Territories. The Great Slave Lake book gives brief
information on access to birdwatching areas and sug-
gestions for timing of visits. The book lists 15
“Highlights” and 20 “Curiosities”. It covers abun-
dance and status of selected species. The 7-page
checklist gives only common names for 237 con-
firmed and 10 unconfirmed species. Its shirt-pocket
size allows it to be carried easily. A quote from John
Richardson entitled “Thoughts From The Past” does
not add anything and probably should have been
deleted to allow a more complete discussion of status
and abundance of species.

The booklet covering the entire Northwest
Territories is physically larger (14 X 21.5 cm) but
has fewer details on the logistics a birdwatcher
faces. Surprisingly there are only 77 more con-
firmed species for the whole Northwest Territories
(i.e., 314) than for the Great Slave Lake area. There
are 16 unconfirmed species for the Northwest
Territories. Common and Latin names are given in
a 10-page species list along with explicit indication
of the status of each species in four broad geo-
graphical regions: Western Cordillera (west of the
Mackenzie and Liard Rivers), Mainland (area east
of these rivers), Arctic Island (marine water and
islands north of 60°N), and Bay Islands (islands and
water of Hudson Bay south of 60°N and James
Bay). These areas are perhaps too broad, for exam-
ple the Mainland region contains boreal, subarctic,
low arctic, and mid-arctic zones, although criteria
for distinction of these zones is not discussed.
Included in the reference section is a listing of five
general birdwatching guides. A criticism is that
several settlements are not shown on the map.
People unfamiliar with the north may not realize
there are settlements making access to these areas
somewhat easier.

348

While both of these card-covered, stapled refer-
ences will need to be used in conjunction with a
field guide, they make an excellent starting point
for beginning birders within and for birders from
outside the localities they cover. The book on Great
Slave Lake is available from Ecology North - a
non-profit organization dedicated to public educa-
tion and environmental conservation. The work

Bird Song

By C. K. Catchpole and P. J. B. Slater. 1995. Cambridge
University Press, New York. viii + 248 pp., illus. U.S.
$32.00.

Bird fans and professionals alike will be grateful
for this highly readable synthesis of the vast litera-
ture of this active topic of investigation. Both
authors are well known for their own contributions,
and have collaborated successfully by attending to
Tinbergen’s four famous programmatic aspects of
behaviour, namely causation, development, function,
and evolution.

The book opens with a general consideration of
signals and communication, sound as a modality, the
terminology for components of bird song, and tech-
niques of recording and analysis, with appropriate
emphasis on the usefulness of interactive digital
sound emitters and the problem of pseudoreplication
in experimental design. More emphasis on the validi-
ty and reliability of recognizing components would
have been desirable because so much of subsequent
issues such as repertoire size depends on this.

Production and reception of sound are examined
in terms of anatomy and physiology, including neur-
al changes and localization of sounds. The develop-
ments of song includes key phenomena such as tem-
plates, memory, social factors, accuracy, mimicry,
and phylogenetic distribution. Especially valulable
here are the presentations of recordings, and the dis-
cussion on the functionality of learning.

BOTANY
A History of the Orchid

By Merle A. Reinikka. 1995. Timber Press, Portland,

Oregon. 324 pp., illus. U.S. $29.95.

Orchids, an exotic topic for a Canadian field natu-
ralist. Journeys into dense greenery, life everywhere,
all around, breathing. When I agreed to review A
History of the Orchid by Merle A. Reinikka, I hoped
to read at least a few of those exotic stories of adven-
tures into the unknown of early collectors and natu-

THE CANADIAN FIELD-NATURALIST

Vols iPr

covering the Northwest Territories is free from
the Government of the Northwest Territories
Department of Economic Development and
Tourism.

DON ALBRIGHT

Box 3, Goodfare, Alberta TOH 1T0O

There are chapters on the transmission of song
(including attenuation and degradation, the role of
habitat, and interpretation of critical features) and the
timing of song in terms of breeding, hormones, the
dawn chorus (whose possible causes are carefully
considered), and competitive aspects. Under diverse
types of social recognition ranging from species to
individuals in territorial defence, the diversity of
experimental techniques employed are well
described. The role of song in sexual selection is out-
lined in depth, particularly stimulation for extra-pair
copulations and aspects of repertoire size. A superb
chapter on themes and variations (which also pro-
vides the subtitle for the book) fully investigates
such major subjects as the organization of repertoire,
duets and choruses, and the evolution and function
of versatility. Finally, variation in space and time is
considered in such terms as dialects and cultural
change. The resulting book is an excellent exposition
written in a clear and flowing style.

The layout is generous and well illustrated, with
references and index. Particularly valuable are the
descriptions of case studies and the presentation of
alternative interpretations for controversial issues.
The wide audience for whom this book is intended
will welcome it.

PATRICK W. COLGAN

Canadian Museum of Nature, P.O. Box 3443, Station D,
Ottawa, Ontario K1P 6P4

ralists. On reading the book, it is immediately evi-
dent that there is an astounding amount of historical
information of interest to someone interested in the
history of botany or taxonomy or orchid culture. I
noted that the book itself is a revised and fuly updat-
ed version of a 1972 edition by the same name, an
established and respected book. There are many
great stories too. A potential buyer should know that

1997

this author is not a story-teller like Peter
Matthiessen. Mr. Reinikka tells his stories fairly bare
boned and so you supply much of the flesh.

I will give a few tidbits I gained from this infor-
mative book. The name Orchid comes from the
Greeks referring to the testiculated bulbs or tubers of
their local plants. The interest was herbal and even
up to the 17th century orchids were prescribed (fol-
lowing Dioscorides’s Doctrine of Signatures) as
remedies for male sexual problems. Field naturalists
observing the plants noticed the flowers had no seeds
but a dust and concluded that orchids were products
of animal semen. This theory was developed to fair
complexity backed by “observant field naturalists”.

The orchid as a flower made its impact in the
West when in 1818 a Cattleya labiata arrived acci-
dentally in England. It was either the packing materi-
al or used to tie some other tropical specimens. A
curious naturalist named William Cattley chose to
try to grow it for some reason. He was successful
and what a success.

The original location was discovered by a Dr.
Gardener in the then semi-remoteness near Rio De
Janerio where the plant thrived. But by the time
word got back to England, this area was totally
destroyed to make room for coffee plantations. An
intense and well-funded search was undertaken but
this plant was not found for another 50 years.
Meanwhile other species were found and the orchid
craze was in full gear.

Plants were bought and sold for small fortunes.
Rain forests that these plants depended on were often
razed to obtain them. One group of collectors in an
effort to obtain the showy and desirable Odonto-
glossium crispum, cut down 4 000 trees to collect
10 000 plants. Ruthless is the word that comes to
mind. Some parts of Central and South America
remain to this day without natural orchids. I did also
note that the collectors, who as a group inflicted such
damage, themselves encountered great dangers and
death in this new land so different from England.

The Ontario Naturalized Garden

By Lorraine Johnson. 1995. Whitecap Books, North
Vancouver. 188 pp., $18.95.

For the average gardener or would-be gardener,
with limited knowledge of natural history or ecolo-
gy, really good information on naturalizing has been
far too scattered in arcane publications to be of
much use. In addition, the rapid increase in publica-
tions on aspects of naturalizing only adds to the
confusion — which authority to consult? Many of
the current books cover a wide geographical area,

Book REVIEWS

349

The last two-thirds of the book is dedicated to bio-
logical notes of 51 people who played prominent
parts in the history of the orchid. To give you an idea
of this I have given a few points that I learnt from
reading one of these biographies.

Charles Darwin states in a letter in 1861, two years
after his ground breaking On the Origin of Species by
Means of Natural Selection, “I never was more inter-
ested in any subject in my life than this of Orchids,”
[In letter to Sir Joseph Hooker quoted on page 181].
He spent considerable time and energy helped by
sons and friends doing field studies of orchid pollina-
tion. He was very impressed by the complex symbiot-
ic relationships that existed between various orchid
and certain insects. He made his famous prediction of
the moth with a foot long proboscis (Xanthopan mor-
gani praedicta) upon viewing the unusually long nec-
tar tube of the Madagascar orchid species
Angraecum sesquipedale. This prediction came out of
a background of intense field study and observation,
not idle speculation. By the way, this and many other
“historic” orchids do make wonderful house plants.
Darwin published a book on the relationship of
insects and orchids in 1862 which he felt was the
needed evidence to uphold his theories, but which
also laid the foundation for any further work on that
branch of orchidology.

As you may have concluded by now, A History of
the Orchid is packed full of facts, details, and infor-
mation. It lacks an overall coherence because it cov-
ers many diverse aspects like taxonomy, cultivation,
hybridization, collecting, and scientific application.
It does do exactly what the author intended, namely,
give a concise history of all aspects of orchids. It
doesn’t speculate, deviate, or stray much. It is thick
reading, if read cover to cover. On the other hand it
is clear and thorough and would serve as an excel-
lent reference book on orchid history for a library or
for someone especially interested in orchids.

STROM EEDY
R.R. 1, McDonalds Corners, Ontario KOG 1MO

have limited subject matter, or have little applica-
tion to Ontario. This is a what makes this book so
worthwhile and timely. At long last Ontario garden-
ers, and naturalists who want to naturalize, have a
book that can advise them on what they can do in
their yards, why they should do it, and what plants
they should use.

The book is divided into three parts and has an
extensive bibliography. Part One gives the basic
arguments for naturalizing, introducing such impor-
tant concepts as the ecological value of indigenous

350

species, an introduction to Ontario’s ecological
zones, some philosophical underpinnings as to the
community benefits of naturalizing, and an overview
of how settlement has contributed to the increase of
aliens at the expense of native species.

Part Two is the practical “how-to” of naturalizing,
covering the types of native plant communities and
describing their habitats, plus design of a native
plant garden, plant propagation, acquisition of native
plants, and species lists for various habitats.

Part Three discusses such important issues as
weeds, lawns, and wildlife, and where they fit into
the naturalization process. In summary, the book
touches all the bases while concentrating on the most
practical ways to encourage people to naturalize.

It is hard to underestimate the importance of the
naturalization movement and how a timely, personal
and popular book with good ideas could make it
grow. As Johnson recognizes in her book, native
plant gardening will grow because its key ideas
make a lot of common sense as well as environmen-
tal sense. Who can argue with ideas that encourage
species diversity and reduced use of fertilizers, toxic
sprays, and water? It is not surprising to learn that
Lorraine Johnson is a Director of the Canadian
Wildflower Society. The book encapsulates much of
the Society’s philosophy and native plant knowledge
as set down in its magazine, Wildflower. No wonder
that its editor, James Hodgins, refers to this book as
“a pioneer work that will be a touchstone for years to
come”’. I heartily concur.

As a naturalist and an environmentalist, I have
long marvelled at the quality and intellectual rigour
of Wildflower. I believe the widespread adoption of
its philosophy and ideas have more potential to
change the average person’s attitudes and relation-
ship with nature than any other movement. It’s based

ENVIRONMENT

THE CANADIAN FIELD-NATURALIST

Vol. 111

on the idea that anyone with a little knowledge and
interest in the natural world can, through working
with native plants, begin to understand their relation-
ship with nature. This practical hands-on approach is
intended to bring about a personal ecological awak-
ening and attitudinal shift from dominance of nature
to partnership with nature.

As we have become increasingly urban, and our
contact with wild areas increasingly remote, we have
created an environment where our planning, engi-
neering and gardening practices are mostly hostile to
the natural order. At long last we have a tool that can
help to redress the balance.

In addition to practical advice, this book includes
an extensive section entitled Further Reading and
Resources. Here are listed important books and
resource materials on ecology and the environment,
gardening, native plants and natural history, maga-
zines, native plant sources, and organizations.

In its purest form, naturalization has the potential
to introduce people to plant communities, and
through attempts at restoration, it offers a way to
start learning about how they function. Wildlife gar-
dening, planting not just to create plant communities,
but to attract insects, amphibians, reptiles, mammals,
and birds, is a logical extension. The whole move-
ment, through the diversity of ways it draws in
people’s interests, is the key to widespread adoption.
As someone working in the naturalizing movement,
it’s wonderful to have a thoughtful and practical
book that I can heartily recommend to expert and
would-be naturalizers.

JEFF HARRISON

33 Glen Avenue, Ottawa, Ontario K1S 2Z6 [Chairman,
Fletcher Wildlife Garden, Ottawa Field-Naturalists Club].

Canada’s Biodiversity: The Variety of Life, its Status, Economic Benefits, Conservation

Costs, and Unmet Needs

By Ted Mosquin, Peter G. Whiting, and Don E. McAllister.
1995. Canadian Museum of Nature, Ottawa. 293 pp.,
illus. $45.00.

A conservation axiom has developed that to main-
tain biodiversity is to ensure a healthy environment.
Any activities that do otherwise will lead to a loss of
ecological integrity. As a buzzword of the 1990s,
biodiversity is a double-edged sword; the public and
decision-makers understand and support the princi-
ple of diversity, but they do not comprehend the
complexity of the issue, the extent of ignorance on
the part of the proponents, or the enormity of the
task of managing what is in effect, the entire planet

from alleles to the Arctic. A new book Canadian
Biodiversity: The Variety of Life, its Status,
Economic Benefits, Conservation Costs, and Unmet
Needs is an important step in rectifying the igno-
rance and complexity of biodiversity in Canada and
a timely arrival as governments and managers, com-
mitted to the Rio Convention on Biological
Diversity, begin to address the formidable task of
maintaining biodiversity.

There are many people, including scientists and
managers, who believe species diversity is biodiver-
sity. In fact, there are three levels of biodiversity;
genetic, species, and community/ecosystem. All lev-

1997

els are discussed throughout the book, often, quite
correctly as interrelated parts of the same system.
But, more important than the addition of genes and
communities, is the recognition of the processes that
create diversity at these levels. To their credit, the
authors of Canada’s Biodiversity use a separate
chapter to underline the importance of such ecologi-
cal functions as soil creation, nutrient cycling, dis-
persal, stability, and plant succession to understand-
ing and maintaining biodiversity.

The remaining chapters cover a wide range of
topics on the status, economics, and management of
biodiversity in Canada. The issue of exactly what
constitutes biodiversity is well established, and I
found the inclusion of domestic species with native
species a good means of incorporating agricultural
interests with those of biodiversity. However, con-
cerns over the conservation of natural biodiversity
outweigh concerns over agricultural and domestic
species and I was surprised that a stronger priority
to the role of native diversity was not evident
throughout.

Much of the value of this book relates to the
wealth of data in the 100 pages of appendices.
Fourteen appendices present data on the number, dis-

The Greening of Ethics

By Richard Sylvan and David Bennett. 1995. The White
Horse Press, Cambridge, U.K. and the University of
Arizona Press, Tucson. 269 pp. Cloth U.S. $45; paper
es 2295.

Sylvan and Bennett teach philosophy in
Australian universities, and are both very interested
in environmental philosophy in particular. The
Greening of Ethics is based on the results of a
UNESCO project to investigate environmental
ethics in Australia. It was written in order to estab-
lish what environmental ethics is and what it is not,
in which directions environmental ethics are going,
and to recommend methods and actions for promot-
ing them. The resulting discussion swings wildly
from basic environmental good sense (Reduce,
Reuse, Recycle) to some very convoluted and foggy
propositions. New words have been coined to
describe the proposals, and commonplace words are
often used in new and inappropriate ways. Some of
the theories are so difficult to follow one wonders
whether the authors themselves have a clear concept
of them. When an inappropriate word is used, it is
often followed by the dictionary definition, but sure-

Book REVIEWS

351

tribution, and rarity of species, the status of protected
areas, and the economic value and costs of biodiver-
sity. Thus, in one reference, a reader can determine
that Canada has two recognized species of
Thysanura (silverfish), 27 of 29 national marine
regions are unrepresented by protected areas, and
that wild fungi are worth up to 30 million dollars
annually. Many topics are covered in detail but sev-
eral jurisdictions lack the data on many topics and
future editions will be required to complete some
appendices, particularly the measures and costs of
conserving biodiversity.

The only real limitation of the book is that the
scope of the coverage, as evidenced in the title, does
not permit the level of detail necessary to make this
book a stand-alone reference on any particular topic.
However, as an overview of biodiversity in general,
and for Canada in particular, the book should be con-
sidered integral to every Canadian interested in
nature conservation.

GRAHAM J. FORBES

Faculty of Forestry and Environmental Management,
University of New Brunswick, Fredericton, New
Brunswick E3B 6C7

ly giving a dictionary definition of a word used out
of context does not make it legitimate!

Is it possible that the environmental movement in
Australia has lagged behind North America? If this
is the case it may explain why some of the early, ele-
mentary environmental principles of the 1970s are
given prominence in the book.

The text also focuses on the “Deep Green” and
“Deep Ecology” theories, and on the application of
philsophical systems to them. There are mathemati-
cal formulae and figures such as “Annular picture of
moral rings in object space (and the position of
Humans)” (page 141). This is an indication that this
is far from an easy book to read, but it may appeal to
intellectuals, trained in philosophy, who are on the
leading edge of environmental ethics and who, by
advocating environmental theories and philosophy,
play their part in changing attitudes towards the
environment.

JANE ATKINSON

255 Malcolm Circle, Dorval, Quebec H9S 1T6

332

THE CANADIAN FIELD-NATURALIST

Vol. 111

Wildlife and Recreationists: Coexistence Through Management and Research

Edited by Richard L. Knight, and Kevin J. Gutzwiller.
1995. Island Press, Washington, D.C. 372 pp., illus.
Cloth U.S. $49.95; paper U.S. $27.50.

The objective of this book was to provide “practi-
cal value in helping minimize or ameliorate the neg-
ative impacts of recreation on wildlife”. I believe the
authors have achieved this objective and more.

The book consists of 21 chapters which are com-
bined into four major topics broadly defined as
General Issues, Specific Issues, Case Studies, and
Ethics and Answers. General Issues provide con-
ceptual overviews of broad topics relevant to the
theme of this book including historical and antici-
pated trends of outdoor recreation, human dimen-
sions of wildlife management, and the origin of
wildlife responses to recreationists. Specific Issues
provides reviews of more defined aspects of recre-
ationist-wildlife interactions such as indirect effects
of recreationists on wildlife and responses of
wildlife to noise. Case Studies brings the reader a
series of well-researched papers on highly-publi-
cized issues including the effects of hunting on
waterfowl, beach recreation and nesting birds, and
waterborne recreation and the Florida manatee.
Most of the chapters include several tables and fig-
ures, many of which draw upon the authors own

Conservation Biology in Theory and Practice

By G. Caughley and A. Gunn. 1996. Blackwell Science,
Cambridge. 459 pp., illus. U.S. $44.95.

In a world of declining biological resources an
understanding of conservation would seem to be
knowledge of interest to individuals involved in
slowing or halting the decline. The purpose of
Conservation Biology in Theory and Practice is to
provide this knowledge and/or means of obtaining it
for this target group. The authors’ secondary aim is
to demonstrate the need to understand the ecology of
species in trouble and the necessity of applying sci-
entific method to determine why species are driven
into decline and how this can be reversed.

The transfer of information is accomplished with a
great deal of reliance on case studies of animal
species. Caughley and Gunn provide the reader with
examples of why it is best to look at the ecology of
the organism(s) in question. They also introduce the
human element of the problem. The human element
often being peoples’ economic aspirations which can
be seen as part of the problem but also part of the
answer.

The information is supplied in a concise and high-
ly readable manner. The use of case studies helps in
bringing a sense of reality to the problem. They help
demonstrate past failures and successes. They also

research or similar scientific studies which augment
the text well. Although written by a group of well-
known researchers, this book is written in a manner
easily understood by scientists and laypeople alike.

Beyond the fact that the authors bring together a
myriad of fundamentally similar topics relative to
effects of human activity on wildlife, I believe the
most beneficial aspect of this book is its practical
approach. Rather than simply state there is a problem
and something should be done about it, many of the
chapters provide realistic solutions to alleviate the
problem and suggest areas of future work to provide
a greater understanding of the problem to improve
management efforts.

Wildlife and Recreationists provides considerable
insight and awareness of the problems and needs for
management of the complex and imperative issue of
human-wildlife coexistence. It will appeal to a broad
cross-section of society and is recommended to any-
one with an interest in our relationship with, and
stewardship for, the environment.

JERROLD L. BELANT

U.S. Department of Agriculture, National Wildlife
Research Center, 6100 Columbus Avenue, Sandusky, Ohio
44870

point out the great difficulty in determining sources
of the problem and possible solutions. The major
problem is often lack of data.

The authors emphasize over and over the need to
follow standard scientific methodologies. They
repeatedly demonstrate the lack of actual facts when
dealing with organisms threatened with extinction.
One quote: “Scientific ignorance should be recog-
nized for what it is and not treated as anything else
forces one to confront this lack of information”
emphasizes what the authors see as one major
problem.

The reader is provided with chapter summaries,
references, and index to aid in using the book for
later reference. I did note a few of the references in
the text failed to correspond with their listing at the
back of the book. On the whole though one is pro-
vided with a good overview of the subject of conser-
vation with a clear and concise introduction to the
methodologies required. The book could be used as
text at the undergraduate level, or a handy reference
for those interested in staving off extinction.

M. P. SCHELLENBERG

434 4th Avenue SE, Swift Current, Saskatchewan
S9H 3M1

1997

Macroecology

By James H. Brown. 1995. University of Chicago Press,
Chicago. xiii + 269 pp., illus. Cloth U.S. $42.50; paper
UWS.$15.75.

A large proportion of today’s ecological studies
provides information dealing with a specific taxon,
microsite, or defined population. Macroecology dis-
cusses approaches and techniques to study the ecolo-
gy of the biosphere or the “bigger picture”. James
Brown writes this monograph, in his own words, as
“a blatant advocacy” looking at the positive features
of macroecological research. The macroecology
approach provides a possible opportunity to see the
big picture despite the loss of detail found with the
more standard reductive and inductive procedures.

The author arranges the material for his mono-
graph in 13 chapters. Each chapter ends with con-
cluding remarks. Brown describes macroecology as
“way of studying relationships between organisms
and their environment that involves characterizing
and explaining statistical patterns of abundance, dis-

Fundamentals of Conservation Biology

By M. L. Hunter. 1995. Blackwell Science, Cambridge,
Massachusetts. xiv + 482 pp., illus. U.S. $42.95.

In recent years the field of conservation biology
has captured a large amount of publicity. There are
several new courses offered by universities to cover
the subject area but, as with any new subject area,
text books were missing. When Malcom Hunter
started writing this text, there were very few possible
texts. Recently this has changed with a number of
books being published to fill the gap Fundamentals
of Conservation Biology is intended to be a text for
an introductory course in conservation biology and
an introduction for readers with a general interest in
the field.

The book has 17 chapters divided into four main
sections: Biodiversity and its importance, Threats to
Biodiversity, Maintaining Biodiversity, and The
Human Factors. Each chapter is provided with sub-
sections, listed in the table of contents for easy refer-
ence, ending with a summary. The chapters also con-
tains at least one case study to bring the material
some sense of reality. For the intrigued, further read-
ings and discussion questions are provided in each
chapter. The discussion questions should provide

BOOK REVIEWS

353

tribution, and diversity” by drawing on a variety of
disciplines such as biogeography, paleobiology, sys-
tematics, and conservation biology. He does not
attempt to provide a complete literature review but
does provide a healthy well-organized overview of
the literature. Nor does he make an attempt to pro-
vide a final statement on the subject but hopes the
book will contribute to its own obsolescence.

James Brown has authored a book which reveals
some of his excitement for the subject. A book with
its ultimate value, he wishes, to be determined by the
impact upon the reader. The impact on this particular
reader has been positive. As a result Macroecology
will be placed on the shelf for easy access and
strongly recommended to those with an interest in
the larger ecological picture.

M. P. SCHELLENBERG

434 4th Avenue SE, Swift Current, Saskatchewan
S9H 3M1

some lively discussion and self exploration. All 17
chapters are arranged to provide a coherent progres-
sion through the material. For ease of reference to
specific information within the text, an index and list
of references are provided at the end of the text.

Dr. Hunter has provided an interesting introduc-
tion to conservation biology. He has attempted to
suggest a change from anthropocentric to biocentric
perspective. The intrinsic value of organisms is dis-
cussed. He raised a number of interesting questions
including “if beaver and reef building corals can
shape landscapes in positive ways, why can’t
people? Can people improve natural ecosystems?”
His references tended to be mostly from the field of
conservation even for areas where other sources are
available in greater quantity. His discussion of cer-
tain topics tended to be one sided occasionally. But
overall Dr. Hunter has provided a good general
introduction and a good read for those with a general
interest in the field.

M. P. SCHELLENBERG

434 4th Avenue SE, Swift Current, Saskatchewan
S9H 3M1

354

THE CANADIAN FIELD-NATURALIST

Vol. 111

Drinking Water: Refreshing Answers to All Your Questions

By James M. Symons. 1995. Texas A&M University
Press, College Station, Texas. Canadian distributor
U.B.C. Press, Vancouver. 160 pp., illus. Cloth $36.95;
paper $16.95.

Although this book is one of a “Natural
Environment Series’’, it is aimed at householders, not
at naturalists. It is written in question-and-answer
format, with 128 questions. Here is No. 120: “Does
drinking water contain calories, fat, sugar, caffeine,

or cholesterol?” The answer is “No”, accompanied
by a sketch of a glass of water overprinted with a list
of the above-named substances, each followed by
“OQ mg”, and a list of the ingredients, to wit, hydro-
gen and oxygen. To be fair, some of the questions
are mildly interesting.

E. C. PIELOU
S-13, C-17, RR 1 Denman Island, British Columbia VOR 1T0

Eastern Deciduous Forest: Ecology and Wildlife Conservation

By Richard H. Yahner. 1995. University of Minnesota
Press, Minneapolis. xvii + 220 pp., illus. U.S. $18.95.

It is difficult to know how to classify this book.
Heavily burdened with references and with a dry
writing style this thin volume is clearly not aimed at
a general audience. And yet, with its superficial
treatment of the topics it can only make for a very
junior level textbook.

Yahner is ambitious in his coverage. In 10 chap-
ters he covers everything from the history of the
eastern deciduous forest since the last ice age to the
current ecological concerns over forest fragmenta-
tion, and the future of the forest. In between, he
tackles succession, plant-animal interactions, biodi-
versity, and atmospheric threats such as acid rain.
In trying to be all encompassing too often chapter
sections resemble little more than lists of topics.
More detailed examination of some of the studies

MISCELLANEOUS

River out of Eden

By Richard Dawkins. 1995. BasicBooks, New York. 172

+ xili pp. U.S. $28.00.

Richard Dawkins is (in)famous for his perceptive
and witty books on Darwinism, and so a reader
approaches this one with anticipation. Those who
have not previously read him will be delighted, but
those already familiar with his previous writings will
be disappointed. The title refers to the river of DNA
coursing over the temporal landscape of biology.
Dawkins expands on the central role of natural selec-
tion in this river, the nature of speciation, and the
view of genetic material as information underlying
complexity, embryology, and adaptation. He then
takes a historical view in order to address issues of
inbreeding, sexual reproduction, phylogenies, molec-

mentioned might have provided greater insight into
the complexity of the issues. There are also a few
careless mistakes in the text: typos and in one
instance, the misuse of the word fauna when flora
was intended.

The book also suffers from an obsessiveness with
references. For example, the self-evident statement
“Forests are an economic and a recreational treasure
for all Americans” is supported by three references,
occupying a full line of text!

Despite these problems, the book does provide a
broad survey of forest ecology and perhaps would
make a useful text for an introductory course in
ecology.

DAVID SEBURN

Seburn Ecological Services, 930 River Road South, R.R. 4,
Kemptville, Ontario KOG 1J0

ular clocks, and an African Eve. Dawkins expounds
his thorough-going adaptationism well by analyzing
natural design, its gradual appearance with plenty of
time and variation, and pertinent examples, especial-
ly from animal behaviour. Confounding those who
search for purpose, he demonstrates the blind effects
of natural selection maximizing genetic survival
through instances of sex ratio, competition, and the
evolution of senescence, but is adamant in drawing
no moral. Finally, Dawkins focusses on “the replica-
tion bomb”, the consequences of molecular replica-
tion exploding with information, like the memes of a
chain letter, with life crossing a series of thresholds
from genotypes through phenotypes and conscious-
ness to space travel. .

1997

The treatment of these topics is generally light,
zesty, and certainly less detailed than in his previous
books. As ever he is messianic in his Darwinism,
strident (against the arguments of religionists, and,
within science, of supporters of punctuated equilib-
ria), and provocative (“there is more poetry in
Mitochondrial Eve than in her mythological name-
sake’). Some claims jar, such as the suggestion that
Ronald Fisher could be regarded as Darwin’s great-
est twentieth-century successor, and reference to
“purebreds” by one so versed in modern genetics.
And the announcement that 1953, the year of the
double helix, “will come to be seen ... as the end of
mystical and obscurantist views of life’ may apply to
some people but certainly not to any society at large.

Overall this volume is neither a general intro-
duction to Darwinism nor an advanced work
exploring some central theme such as the evolving
intellectual framework of Darwinism (cf. Depew
and Weber’s Darwinism Evolving reviewed in The
Canadian Field-Naturalist 109: 515) or evolution-

Book REVIEWS

S35

ary ethics (cf. Ruse’s Evolutionary Naturalism
reviewed in The Canadian Field-Naturalist 110(3):
568). An author who has worked hard to conclude
that “Nature is neither kind nor unkind” might
have been expected to say at least something about
ethics.

Subtitled A Darwinian View of Life, this is a vol-
ume in a Science Masters Series of “short, easy-to-
read, attractive books” conveying cutting-edge ideas
to a broad audience. It is on a par with the one other
volume of this series that I have seen, but is shallow
compared with Dawkins’ excellent other books.
Hence the conclusion to date, although the list of
future contributors is stellar, must be that these vol-
umes are light introductions, fine as such, but pale
reflections of substantial works.

PATRICK W. COLGAN

Canadian Museum of Nature, P.O. Box 3443, Station D,
Ottawa, Ontario K1P 6P4

Diary of William W. Judd: Sifton Botanical Bog (Byron Bog) London, Ontario, 1981-1993

By W. W. Judd. 1995. Phelps Publishing, London. iti + 25
pp. $5.00.

This booklet is the author’s third diary on the
Byron Bog of London, Ontario. The previous two
covered the years 1961-1966 and 1967-1980,
although I have not seen them. Anyone expecting
rich personal observations by an eminent scientist on
the changing natural landscape of the Byron Bog
will be sorely disappointed. Judd’s entries are terse,
sometimes almost in point form: date (occasionally
even this is missing), weather, time of visit, plants in
bloom, and animals seen or heard. Each year’s

NEw TITLES

Zoology

*American marten, fisher, lynx, and wolverine: survey
methods for their detection. 1995. Edited by W. J.
Zielinksi and T. E. Kucera. General Technical Report
PSW-GTR-157, Pacific Southwest Research Station, U.S.
Department of Agriculture, Albany, California. 163 pp.

*American Museum of Natural History: 125 years of
expedition and discovery. 1995. By L. Rexer and R.
Klein. Harry N. Adams, New York. 256 pp., illus.

*Amphibian biology, volume 1: the Integument. 1994.
Edited by H. Heatwole and G. T. Barthalmus. Surrey
Beatty, Chipping Norton, Australia. 418 pp., illus. U.S.
$92.

entries average only a page in length, with the
remaining 12 of 25 pages occupied by appendices.
The appendices thoroughly cross-reference the diary
by people referred to in the diary and plants and ani-
mals mentioned in the text. Judd evidently knows a
great deal about the bog; it is a shame he has not
shared it more fully with the rest of us.

DAVID SEBURN

Seburn Ecological Services, 930 River Road South, R.R.
#4, Kemptville, Ontario KOG 1JO

*Amphibian biology, volume 2: social behavior. 1995.
Edited by H. Heatwole and B. Sullivan, Surrey Beatty,
Chipping Norton, Australia. pp.419-710, illus.

*The amphibians and reptiles of the Yucatan Peninsula.
1996. By J. C. Lee. Cornell University Press, Ithaca. 500
pp., illus. U.S. $175.

*The birds of British Columbia: Volume 3: passerines
(flycatchers through vireos). 1997. By R. W. Campbell,
N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W.
Kaiser, M.C. E. McNall, and G.E.J. Smith. University of
British Columbia Press, Vancouver. c635pp., illus. $95.

+The breeding birds of Québec: atlas of the breeding
birds of southern Québec. Les oiseaux richeurs du

356

Québec. 1996. Edited by J. Gauthier and Y. Aubry.
Association québecoise des groupes d’ornithologues,
Province of Québec Society for the Protection of Birds
and Canadian Wildlife Service, Montreal. Editions
Multimondes, Saint-Foy. (English and French, review
copy is English) 1302 pp., illus. $149.95.

Coming of age with elephants, a memoir. 1996. By J.
Poole. Hyperion, N.Y. xi+288 pp., illus. U.S. $24.95.

*Cranes: their biology, husbandry, and conservation.
1996. Edited by D. H. Ellis. G. F. Gee, and C. M.
Mirande. Hancock House, Surrey, B. C. xii+307 pp., illus.
$69.95.

+Demography of the northern spotted owl. 1996. Edited
by E. D. Forsman, S. De Stefano, M. G. Raphael, and R. J.
Gutierrez. Studies in Avian Biology No. 17. Proceedings
of a Workshop, Fort Collins, Colorado, December, 1993.
Cooper Ornithological Society, Camarillo, California. 122
pp., illus. U.S. $20.

Eagle’s plume: preserving the life and habitat of
America’s bald eagle. 1996. By B.E. Beans. Scribner,
New York. 318 pp., illus. U.S. $25.

The earth dwellers: adventures in the land of ants.
1996. By E. Hoyt. Simon and Schuster, New York. 319
pp., illus. U.S. $24.

The ecology of migrant birds: a neotropical perspec-
tive. 1996. By J. H. Rappel. Smithsonian Institute. P.,
Washington. xvi + 269 pp., illus. U.S. $35.

*Encyclopaedia of Canadian Fishes. 1996. By B. Coad.
Canadian Museum of Nature, Ottawa. 928 pp., illus. $100.

*Field guide to the birds of the Middle East. 1996. By R.
F. Porter, S. Christensen, and P. Schiermacker-Hansen. T
& A.D. Poyser, London, England. Xv + 460 pp., illus. +
plates. £29.95.

*The historical atlas of breeding birds in Britain and
Ireland 1875-1900. 1996. Compiled by S. Holloway. T &
A Poyser, London, England. 476 pp., illus. £25.

Hyperiid amphipods (Amphipoda, Hyperiida) of the
world ocean. 1996. By M. E. Vinogradov, A. F. Volkov,
and T. N. Semenova. Science Publishers, Lebanon,
Michigan. 650 pp. U.S. $178.

Larvae of marine bivalves and echinoderms. 1997. By
V. E. Kasyanov, G. A. Kryuchova, V. A. Kulikova, and L.
A. Medvedva. Science Publishers, Lebanon, New
Hampshire c350pp.

+ Life of the flycatcher. 1997. By A. F. Skutch. University
Oklahoma Press, Norman. 192., illus. U.S. $40.

Mammals of the Soviet Union, volume 1, part 3: pin-
nipeds and toothed whales (Pinnipedia and Odonto-
celi). 1996. By V. G. Heptner. Science Publishers,
Lebanon, New Hampshire. cl000pp. U.S. $225.

Mammals of the Soviet Union, volume 2, part la: sea
cows, wolves, and bears. 1997. Edited by V. G. Heptner.
Science Publishers, Lebanon, New Hampshire. c750pp.
U.S. $195.

*+Metapopulations and wildlife conservation. 1996.
Edited by D. R. McCullough. Island Press, Washington. x
+ 429 pp., illus. Cloth U.S. $55: paper U.S. $28.

THE CANADIAN FIELD-NATURALIST

Vol. 111

*Monitoring bird populations by point counts. 1995.
Edited by R. C. John, J. R. Sauer, and S. Droege. General
Technical Report PSW-GTR-149, Pacific Southwest
Research Station, U.S. Department of Agriculture,
Albany, California. 187 pp.

*Pleistocene amphibians and reptiles in North America.
1995. By J. A. Holman. Oxford University Press, New
York. 243 pp. U.S. $65.

*Snakes in question: the Smithsonian answer book.
1996. By C. H. Ernst and G. R. Zug. Smithsonian
Institution Press, Washington. xii + 203 pp., illus. +
plates. Cloth U.S. $49; paper U.S. $24.95.

The venomous and poisonous marine invertebrates of
the Indian Ocean. 1996. By F. E. Russell and R. Nagab-
hushanam. Science Publishers, Lebanon, New Hampshire
c270 pp. U.S. $ 89.

*The wood duck and the mandarin: the northern wood
ducks. 1996. By. L. L. Shurtleff and C. Savage.
University of California Press, Berkley. 232 pp., illus.
US. $34.95.

Botany

+The alpine flora of the Rocky Mountains, Volume 1:
the middle Rockies. 1997. By. R. W. Scott. University
Utah Press, Salt Lake City. 768 pp., illus. U.S. $110.

*Balancing act: environmental issues in forestry. 1996.
By H. Kimmins. 2nd edition. University British Columbia
Press, Vancouver. 305 pp., illus. $29.95

Conifers: the illustrated encyclopaedia. 1996. By. D. M.
van Gelderen and J. R. P. van Hoey Smith. 2 volumes.
Timber Press, Portland, Oregon. 706 pp., illus. U.S. $125.

Contemporary plant systematics. 1997. By R. R.
Kowal, K. J. Sytsma, M. Clayton, M. J. Ward, and D.
Woodland. Andrews University Press, Berrien Springs,
Michigan. 680 pp., illus. + CDROM. U.S. $59.95.

Fungi on rhododendron: a world reference. 1996. By
D. F. Farr, H. B. Esteban, and M. E. Palm. Parkway
Publishers, Boone, North Carolina. 192 pp. U. S. $45.

Meeting the tree of life: a teacher’s path. 1997. By J.
Tallmadge. University Utah Press, Salt Lake City. 220 pp.
Cloth U.S. $44.95: paper U.S. $18.95.

*Michigan flora, part III: dicots (Pyrolaceae-
Compositae). 1996. By E. G. Voss. Cranbrook Institute
of Science, Broomfield Hills, Michigan. xix + 622 pp.,
illus. U.S. $15.

Mycology in sustainable development: expanding con-
cepts and vanishing borders. 1997. Edited by I. H.
Chapela and M. E. Palm. Proceedings of an international
Workshop, San Diego, 9 August 1995. Parkway
Publishers, Boone, North Carolina. c150 pp., U.S. $40.

Environment

+The abstract wild. 1996. By J. Turner. University
Arizona Press, Tucson. xvii + 136 pp. Cloth U. S. $32.50;
paper U.S. $15.95.

*British Columbia: a natural history. 1996. By. R. and S.
Cannings. Greystone Books, Douglas and McIntyre,
Vancouver. 310 pp., illus. $45.

1997

*Conservation of Great Plains ecosystem: current sci-
ence, future options. 1995. Edited by S. R. Johnson and
A. Bouzaher. Kluwer Academic Publishers, Hingham,
Massachusetts. xvii + 434 pp., illus. U.S. $192.

+Deeper ecology: essays on ecological spirituality. 1996.
By D. A. Doyle. Wild Side Publishing, Eureka, California.
47 pp., U.S. $6.95 + U.S. $3 shipping + U.S. $10 foreign.

+Ecological census techniques: a handout. 1996. Edited
by W. J. Sutherland. Cambridge University Press, New
York. xv + 336 pp., illus. Cloth U.S. $74.95; paper U.S.
$27.95.

Ecology of a polar oasis: Alexandra Fiord, Ellesmere
Island, Canada. 1996. Editd by j. Svoboda and B.
Freedman. Captus Press, North York, Onatrio. 286 pp.,
illus. $38.50.

Environmental geology of urban areas. 1997. Edited by
N. Eyles. Geological Association of Canada, Memorial
University, St. John’s Newfoundland. 608 pp. $95; U.S.
$81.

Environmental harm: civil actions and Compensation.
1996. By. M. D. Faiela, H. B. Kolm, R. Kligman, and J.
Swaigen. Butterworths, Markham. 688 pp. $176.

Environmental law: a guide to concepts. 1996. By. B.
Pardy. Butterworths, Markham. $53.

+Evolution and environment in tropical America. 1996.
Edited by J.B.C. Jackson, A. F. Budd, and A. G. Coates.
University Chicago Press, Chicago. ix + 425 pp., illus.
Cloth U.S. $27.50.

The forgiving air: understanding environmental
change. 1996. By R. C. J. Somerville. University of
California Press, Berkley. xiv + 195 pp., illus. U.S.
$21.95.

*Invasions of the land. 1995. By M. S. Gordon and E. C.
Olson. Columbia University Press, N. Y. xv + 312 pp.,
illus.

Islands in space and time. 1996. By D. Campbell.
Houghton Mifflin, New York. pp., illus. U.S. $35.

Krakatau: the destruction and reassembly of an island
ecosystem. 1996. By I. Thornton. Harvard University
Press, Cambridge, Massachusetts. xi + 346 pp., illus. U.S.
$39.95.

+Measuring and monitoring biological diversity: stan-
dard methods for mammals. 1996. Edited by D. E.
Wilson, F. R. Cole, J. D. Nichols, R. Rudran, and M. S.
Foster. Smithsonian I. P., Washington. xxvii + 409 pp.,
illus. Cloth U.S. $49; paper U.S. $22.50.

+Misreading the Africa landscape: society and ecology
in the forest-savanna mosaic. 1996. B. J. Fairhead and
M. Leach. Cambridge University Press, New York. xvii +
354., illus. Cloth U.S. $55; paper $19.95.

+The natural classroom: a directory of field courses,
programs, and expeditions in the natural sciences.
1996. By J. R. Edelman. North American Press, Golden,
Colorado. 279 pp. U.S. $32.50.

*The natural history of pollination. 1996. By M. Proctor,
P. Yeo, and A. Lack. Harper Colins, London. 479 pp.,
illus. £16.99.

BoOoK REVIEWS

a59,

*Nature guide to Ontario. 1997. Edited by W. Wake.
University Toronto Press., Toronto. c500 pp., illus. Cloth
$65; paper $19.95.

+Null models in ecology. 1996. By N. J. Gotelli and G. R.

Graves. Smithsonian Institute Press, Washington. xvi +
368 pp., illus. U.S. $30.

Okoboji wetlands: a lesson in natural history. 1996. By
M. J. Lannoo. University Iowa Press, Iowa City. xiv + 156
pp., illus. Cloth U.S. $29.95; paper U.S. $14.95.

*Restoring nature’s place: a guide to naturalizing
Ontario parks and greenspace. Ontario Parks
Association, Toronto and Ecological Outlook Consulting,
Schomberg. v + 227 pp., illus. $49.

Riparian ecosystem recovery in arid lands: strategies
and references. 1996. By M. K. Briggs. University
Arizona Press, Tuscon. xv + 159 pp., illus. Cloth U.S.
$45; paper U.S. $19.95.

*In search of nature. 1996. By E. O. Wilson. Island Press,
Washington. x + 214 pp., illus. U.S. $19.95.

The shoresaver’s handbook: a citizen’s guide. 1996.
By. T. Coomble. Lyons & Burford, New York. 138 pp.,
illus. U.S. $12.95.

Sixty years of biology: essays on evolution and devel-
opment. 1996. By J. T. Bonner. Princeton University
Press, Princeton. x + 143 pp., illus. U. S. $24.95.

*The song of the dodo: island biologeography in an age
of extinctions. 1996. By D. Quammen. Scribner, New
York. 702 pp., illus. U.S. $32.50.

+Wild China. 1996. By J. MacKinnon. MIT Press,
Cambridge, Massachusetts. 208 pp., illus. U.S. $40.

World resources 1996-1997: a guide to the global envi-
ronment. 1996. By the World Resources Institute. Oxford
University Press, New York. xiv + 365 pp., illus. U.S.
$24.95.

Miscellaneous

*+Darwin’s dream pond: drama in the Lake Victoria.
1996. By T. Goldschmidt. Translated by S. Marx-
Macdonald. MIT Press, Cambridge, Massachussetts. 274
pp., illus.

The late Devonian Mass extinction: the Frasnian/
Famennian crisis. 1996. By G. R. McGhee, Jr. Columbia
University Press, New York. xi + 303 pp., illus. U.S. $20.

+Minutes of meetings 1936-1939 of the MclIlwraith
Ornithological Club, London, Ontario, Canada. 1997.
By W. W. Judd. Phelps Publishing, London, Ontario.
(order from author, 50 Hunt Club Drive, London, Ontario
N6H 3Y3). 108 pp. $10.

Plants, people, and culture: the science of ethnobotany.
1996. By M. J. Balick and P. A. Cox. Freeman, New
York. vii + 228 pp., illus. U.S.$ 32.95.

Summer stargazing. 1996. By T. Dickinson. Firefly
Books, New York. 64 pp., illus. U.S. $18.95.

The time before history: five million years of human
impact. 1996. By C. Tudge. Scribner, New York. 366 pp.,
illus. U.S. $27.50.

BOOKS FOR YOUNG NATURALISTS

Anytime weather everywhere. 1996. By M. H. Mogh
and B. G. Levine. How the Weather Works, Rockville,
Maryland. 106 pp., illus. U.S. $15.95.

Backyard birds of summer. 1996. By C. Lerner. Morrow
Junior, New York. 48 pp., illus. U.S. $16.

Bizzare insects. 1996. By M. J. Anderson. Enslow,
Springfield, New Jersey. 48 pp., illus. U.S. $15.95.

Blood-feeding bugs and beasts. 1996. By L. P. Kite.
Millbrook Press, Brookfield, Connecticut. 48 pp., illus.
U.S. $15.90.

Bloodsuckers. 1996. By. R. Knapp. Enslow, Springfield,
New Jersey. 48 pp., illus. U.S. $15.95.

A brief green history of planet earth. 1996. A. F. Alper.
PPI Publishing, Dayton. 141 pp., illus. U.S. $6.95.

Call of the wolves. 1996. By M. Berger. Newbridge, New
York. 16 pp., illus. + teaching guide. U.S. $14.95.

City kids and city critters: activities for urban explor-
ers. 1996. By J. W. Roberts and C. Huelbig. Learning
Triange Press, New York. 153 pp., illus. U.S. $19.95.

Dirty, rotten, dead: a worm’s-eye view of death, de-
composition, and life. 1996. By J. Emory, Harcourt,
Brace, San Diego. 48 pp., illus. U.S. $15.

How do animals sleep? 1996. By M. Berger. Newbridge,
New York. 16 pp., illus. + teaching guide. U.S. $14.95.

Invertebrates. 1996. By A., V., and R. Silverstein.
Twenty-first Century Books, New York. 64 pp., illus. U.S.
$16.98.

Janice vanCleave’s rocks and minerals: mind-boggling
experiments you can turn into science fair projects.
1996. By J. vanCleave. Wiley, New York. 90 pp., illus.
WS. $995.

Life. 1996. By R. Snedden. Chelsea House, New York. 48
pp., illus. U.S. $15.95.

THE CANADIAN FIELD-NATURALIST

Vol. 111

Mammals. 1996. By Time-Life Books, Alexandria,
Virginia. 64 pp., illus. U.S. $15.

The microscope book. 1996. By S. Levine and L.
Johnstone. Sterling, New York. 80 pp., illus. U.S. $19.95.

Monerans and protists. 1996. By A., V., and R.
Silverstein. Twenty-first Century Books, New York. 64
pp., illus. U.S. $16.95.

Natural treasures fieldguide for kids. 1996. By E.
Biesiot. Roberts Rinehart, Boulder. 64pp., illus. U.S.
$12.95.

The night sky. 1996. By R. Kerrod. Benchmark Books,
Tarrytown, New York. 64 pp., illus. U.S. $16.95.

Plants of the dessert; Plants of the forest; Plants under
the sea; and The vegetation of rivers, lakes, and
swamps. 1996. By A. Llamas. Chelsea House, New York.
each 32 pp., illus. U.S. $14.95.

Reptiles. 1996. By Time-Life Books, Alexandria,
Virginia. 64 pp., illus. U.S. $15.

Seismosaurus: the longest dinosaur and Utahraptor:
the deadliest dinosaur. 1996. By D. Lessem. Carolrhoda,
Minneapolis. each 40 pp., illus. U.S. $14.95.

Super science fair sourcebook. 1996. By H. M. Iritz.
McGraw-Hill, New York. viii + 248 pp., illus. U.S.
$17.95.

Vertebrates. 1996. By A., V., and R. Silverstein. Twenty-
first Century Books, New York. 64 pp., illus. U.S. $16.95.

World’s weirdest birds. 1996. By M. I. Roberts. Troll,
Mahwah, New Jersey. 32 pp., illus. U.S. $12.50.

You animal: putting humans in their place. 1996. By J.
Booth. Harcourt, Brace, SanDiego. 48 pp., illus. U.S. $15.

Young explorer’s guide to undersea life. 1996. By P.
Armstrong. Monterey Bay Aquarium, Monterey,
California. 64 pp., illus. U.S. $16.95.

yavailable for review
*assigned for review

l

\

- Notes

First breeding record of Red-breasted Merganser, Mergus serrator,
on Axel Heiberg Island, Northwest Territories
THEO HOFMANN, JOHN W. CHARDINE, and HANS BLOKPOEL

Sagina (Caryophyllaceae) range extensions in Canada: S. japonica new
to Newfoundland; S. procumbens new to the Northwest Territories
RICHARD K. RABELER and JOHN W. THIERET

Use of active Beaver, Castor canadensis, lodges by Muskrats, Ondatra zibethicus, in Wyoming
MaRK C. MCKINSTRY, RorY R. KARHU, and STANLEY H. ANDERSON

Fall — early winter home ranges, movements, and den use of male Mink,
Mustela vision, in eastern Tennessee
RICHARD T. STEVENS, TOM L. ASHWOOD, and JONATHAN M. SLEEMAN

Observations of a possible cleaning symbiosis between Painted Turtles,
Chrysemys picta, and Snapping Turtles, Cheldra serpentina, in central Ontario
MARGARET A. KRAWCHUK, NICOLA KOPER, and RONALD J. BROOKS

Weissia brachycarpa (Nees & Hornsch.) Jur. at Niagara Falls,
a moss new to Ontario P. M. ECKEL

The moss Tortella alpicola Dix. new to Alberta and the Yukon Territory
with a discussion of its range and comments on related species P. M. ECKEL

News and Comment

Editor’s Report for The Canadian Field-Naturalist Volume 110 (1996) — Global Biodiversity: The
Canadian Museum of Nature — Rana-Sauria — CITES Reports: 1992 Annual Report for Canada —
Froglog: Newsletter of the Declining Amphibians Populations Task Force of the World
Conservation Union’s Species Survival Commission — Ontario Natural Heritage Information Centre
Newsletter — Renew Report # 6, 1995-96 — Sea Wind — Richards Natural History Education
Award by the Federation of Ontario Naturalists

Minutes of the 118th Annual Business Meeting of The Ottawa Field-Naturalists’ Club, 14 January 1997

The problem of invading alien trees and shrubs: Some observations in Ontario and a Canadian checklist
PAUL M. CATLING

Book Reviews

Zoology: The Minds of Birds — Martens, Sables, and Fishers: Biology and Conservation —
Chickadees, Tits, Nuthatches & Treecreepers — Larvae of the North American Caddisfly Genera
(Trichoptera) — The Scientific Basis for Conserving Forest Carnivores: American Marten, Fisher,
Lynx, and Wolverine in the Western United States — The Last Panda — The Birds of Great Slave
Lake, Northwest Territories, Canada — The Birds of the Northwest Territories — Bird Song

Botany: A History of the Orchid — The Ontario Naturalized Garden

Environment: Canada’s Biodiversity: The Variety of Life, its Status, Economic Benefits, Conservation
Costs, and Unmet Needs — The Greening of Ethics — Wildlife and Recreationists: Coexistence
Through Management and Research — Conservation Biology in Theory and Practice —
Macroecology — Fundamentals of Conservation Biology — Drinking Water: Refreshing Answers to
All Your Questions — Eastern Deciduous Forest: Ecology and Wildlife Conservation

Miscellaneous: River Out of Eden — Diary of William W. Judd: Sifton Botanical Bog (Byron Bog),
London, Ontario, 1981-1993

New Titles

Mailing date of the previous issue 111(1): 2 May 1997

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309

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320

325
331

338

343
348

350

354
333

THE CANADIAN FIELD-NATURALIST Volume 111, Number 2 1997

Articles

The decline and current status of the dune race of the Dwarf Cherry,
Prunus pumila var. pumila, on the Canadian shores of the lower Great Lakes :
PAUL M. CATLING and BRENDON M. H. LARSON 187

A taxonomic study of the grass genus Glyceria (mannagrass) in British Columbia
ROSARIO RUIZ DE ESPARZA and JACK MAZE 194.

Coyote, Canis latrans, visitations to scent stations in southeastern Alberta
SYBILLE WOELFL and MANFRED WOELFL 200:

Beaver, Castor canadensis, home range size and patterns of use in the
taiga of southeastern Manitoba: I. Seasonal variation MICHELLE WHEATLEY 204:

Beaver, Castor canadensis, home range size and patterns of use in the
taiga of southeastern Manitoba: II. Sex, age, and family status MICHELLE WHEATLEY 211

Beaver, Castor canadensis, home range size and patterns of use in the |
taiga of southeastern Manitoba: II. Habitat variation MICHELLE WHEATLEY 217}

Food preference and ad libitum intake of wild-captured Sitka Mice,
Peromyscus keeni sitkensis
ED O. REESE, JEFFREY C. BARNARD, and THOMAS A. HANLEY 223

Summer food habits and population density of Coyotes, Canis latrans,
in boreal forests of southeasten Québec CLAUDE SAMSON and MICHEL CRETE 228

Egg composition and post-DDT eggshell thickness of the American White Pelican,

Pelecanus erythrorhyncos SHAWN C. BUGDEN and ROGER M. EVANS 234

Population growth of Moose, Alces alces, in Labrador |
TONY E. CHUBBS and JAMES A. SCHAEFER 238
Resurgence of breeding Merlins, Falco columbarius richardsonii,

in Saskatchewan grasslands C. STUART HOUSTON and KEITH A. HODSON 243
Rare and endangered fishes and marine mammals of Canada:

COSEWIC Fish and Marine Mammal Subcommittee Status Reports XI. R. R. CAMPBELL 249
Status of the Nootsack Dace, Rhinichthys sp., in Canada J. D. MCPHAIL 258
Status of the Northern Fur Seal, Callorhinus urinsus, in Canada

ROBIN W. BAIRD and M. BRADLEY HANSON 263
Status of the Lacs des Loups Marins Harbour Seal,

Phoca vitulina mellonae, in Canada R. J. SMITH 270!

Updated Status of the Sea Otter, Enhydra lutris, in Canada
JANE C. WATSON, GRAEME M. ELLIS, THOMAS G. SMITH, and JOHN K. B. FORD 277

Status of the Northern Bottlenose Whale, Hyperoodon ampullatus,
in the Gully, Nova Scotia
HAL WHITEHEAD, ANNICK FAUCHER, SHANNON GOWANS, and STEPHEN MCCARREY 287 |

Status of the Sperm Whale, Physeter macrocephalus, in Canada
RANDALL R. REEVES and HAL WHITEHEAD 293:

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