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AMPHIPACIFICA

JOURNAL OF AQ.UATIC SYSTEMATIC

BIOLOGY

Volume 3, Number 2
Nov. 15, 2001

AMPHIPACIFICA, Journal of Aquatic Systematic Biology (ISSN No. 1 189-9905) is published
quarterly by Amphipacifica Research Publications, 1710-1275 Richmond Rd., Ottawa, Ontario.,
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DEDICATION

The Journal AMPfflPACIFICA is dedicated to the promotion of systematic biology and
to the conservation of Earth’s natural resources.

This issue is dedicated to the memory of Ponald Evan McAllister, 1934 - 2001.

Cover design: Adapted from the title page of S. J. Holmes ( 1904). “Amphipod

Crustaceans of the Expedition.”

Harriman Alaska Expedition, pages 233-244.

AMPHIPACIFICA VOL. 3 NO. 2. Nov. 15, 2001

Donald Evan McAllister, 1934-2001:

A Tribute

During his career at the National Museum (now
Canadian Museum of Nature) as Curator of Fishes
( 1958 - 1986), Senior Research Scientist ( 1986 -1994),
Researcher Emeritus and Research Associate (1994 -
2001), Donald Evan McAllister made significant con-
tributions not only in his formal ichthyological and
curatorial studies, but also in his thoughtful advocacy
of responsible environmental management, and for
protecting global biodiversity. He founded and edited
two scientific journals, taught and inspired many indi-
viduals, and impressed all those he met as a warm,
friendly, humble, and patient man devoted to both his
family and his science. But above all else Don would
have been most pleased by a colleagues's characteriza-
tion of him as a true friend of the Earth.

Don was bom 23 August, 1934, in Victoria, B. C. He
took his B.A., M. A. (on freshwater sculpins of British
Columbia), and Ph.D. at the University of British Col-
umbia. His thesis for the latter, the Evolution of
Branchiostegals and Classification of Teleostome
Fishes, was published as a museum bulletin in 1968.
During his Ph.D. studies, he spent a year as an NSERC
Visiting Student at the Museum of Zoology, Univer-
sity of Michigan. He was the first ichthyologist on the
staff of the National Museum since its creation as part
of the Geological Survey in 1842, and was, unquestion-
ably, one of the most energetic, innovative, issue-ori-
ented and productive members of the staff. Although
Don lacked administrative ambition, or the patiencefor
it, he ran the ichthyology section with verve and infec-
tious energy. He attracted, trained and encouraged a
succession of assistant curators, technicians, contract-
ees, visiting scientists, post-doctoral students, assist-
ants and volunteers. His example of work ethic,
compassion, and concern for those around him left its
mark on everyone who passed though his section.

Don was, first and foremost, a collection builder. On
his arrival at the museum, there were fewer than 4,5(X)
specimens of fishes in the collection. Through every
means available to him, staff expeditions, donations,
purchases and trades, he built the collection into a
world-class resource for taxonomic and life-history
studies. By the year 2000, there were more than
585,000 specimens from all parts of the world, a
remarkable achievement on its own. But, at the same
time, Don was also a compulsive author who left a
comprehensive published record of his interests. His
bibliography exceeds 620 items* including scientific

1 ^

Donald Evan McAllister, 1934-2001

papers, books, popular articles, and book reviews,
ranging from his Ph.D. thesis and a Guide to the
Freshwater Sportfish of Canada, taxonomic revisions,
papers on computers in museums, methods of collect-
ing and preserving fishes, and the use of x-rays in
taxonomic studies. He was also the originator of two
environmental journals to which he contributed exten-
sive overviews and informed opinion.

From the time of his appointment as curator, Don
researched Arctic marine fishes with a view to a com-
panion book to the fishes of the other Canadian coasts.
He produced a preliminary checklist in 1960, followed
by keys and a series of papers. A distributional atlas,
more papers and a bibliography followed in the 1970s
and 1980s. He compiled an extensive database which
his successor, Brian Coad has continued. With the help
of a number of museum staff and the financial support
of the Department of Fisheries and Oceans, the long
envisioned book is close to completion.

He was a pioneer and advocate of the use of comput-
ers in museums which led to his devising a standard-
ized data sheet for recording detailed information in the
field. He also compiled an as yet unpublished compre-
hensive list of ichthyological terms and their defini-
tions. The computerized database of the fish collection

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 2

formed the basis for a list of the families and species of
fishes of the world which lists more than 20,000 valid
taxa.

Don held adjunct professorships at Carleton and
Ottawa Universities, taught ichthyology at the latter,
and participated on graduate student committees at
both. In 1974, he organized the American Society of
Ichthyologists and Herpetologists annual meeting in
Ottawa, only the third time the Society met in Canada
and the first meeting that was bilingual . Don was very
active on the Fish and Marine Mammals subcommittee
of the Committee on the Status of Endangered Wildlife
in Canada and wrote the first list of endangered fishes
in Canada in 1970.

In the late 1980s and throughout the 1990s, Don
increasingly applied his wealth of experience and his
social conscience to concerns for the planet. His
writing and activities on international committees were
directed at a wider audience than his ichthyological
colleagues, and he moved into the Museum’s newly-
created Biodiversity group. He had by this time
founded and edited the quarterly "Sea Wind", now in its
15th year of publication, and he mortgaged his house to
fund conservation and training programs organized by
its publisher, "Ocean Voice International". In 1991 he
originated and edited eight volumes of the quarterly
Canadian Biodiversity (later retitled "Global Biodi-
versity") published by CMN. In 2000, it was replaced
with "Biodiversity: A Journal of Life on Earth" and
published by the Tropical Conservancy. He was esp-
ecially active in efforts to conserve the world’s coral

reefs and finding non-destructive methods for the har-
vest of aquarium fishes by Philippine fishermen. He
participated in Canada's Biodiversity Convention Ad-
visory Group and advised on Canada’s strategy on the
Rio treaty signed in 1991.

During a period of museum downsizing in 1993, in
a characteristic selfless gesture, he volunteered early
retirement to free a position so that a younger staff
member could be retained. Following retirement he
did not "ease up" on editing the conservation-oriented
journals he had initiated, nor did he slacken his own
perceptive contributions to them.

Early in his career Don had made personal contribu-
tions to studies of invertebrate animals, largely through
his enthusiastic participation in two Museum expedi-
tions in the 1960s with Drs. Edward L. Bousfield
(coastal southeastern Alaska) and Arthur H. Clarke
(east coast of Hudson Bay), During the Alaska exped-
ition, Don’s keen eyes spotted several unusual amphi-
pods that were ectoparasitic on rockfish and greenling
cod that he had collected. Back at the museum lab,
most of the crustacean species proved new to science,
one of which, Paralafystius mcallisterU was named in
his honour. His passing has left a void that no single
individual is ever likely to fill.

Francis R. Cook, Charles G. Gruchy, and

Brian W. Coad

*A complete bibliography of Don McAllister's publications
has been compiled by Brian Coad and will appear in the
Canadian Field-Naturalist.

SEA WIND is published quarterly and may be obtained through membership or subscription from
OCEAN VOICE lOTERNATIONAL, P.O. Box 37026, 3332 McCarthy Road Ottawa, ON KIV OWO,
Canada, www.ovkca . PH; (613) 721-4541, Fax (613) 721-4562. Regular memberships: $25. per year.

AMPHIPACIFICA VOL. 3 NO. 2. Nov. 15, 2001 3

THE AMPHIPOD GENUS ALLORCHESTES IN THE NORTH PACIFIC REGION:
SYSTEMATICS AND DISTRIBUTIONAL ECOLOGY.

By E. A. Hendrycks^ and E. L. BonsHeld ^

Abstract

The talitroidean amphipod family Hyalellidae Bulycheva 1957, emend Bousfield, 1996, and herewith rede-
fined, encompasses the marine genera Allorchestes Dana, Parhyalella Kunkel , Exhyalella Stebbing, Marinohyale Ha
Lazo-Wasem & Gable, and the neotropical freshwater genus Hyalella Saussure. From the Pacific coast of North
America are newly described and figured Allorchestes rickeri n. sp. (S.E. Alaska to Central California) and A.
priceae n, sp. (S.E, Alaska to southern Vancouver Island). Character states of reproductive morphology are newly
utilized in redescribing and refiguring Allorchestes angusta, Dana (Alaska to California), A. bellabella Barnard
(Kamchatka and Bering sea to Oregon), A. carimta Iwasa (Northern Sea of Japan, Bering Sea and S.E Alaska), A.
malleola Stebbing (northern Sea of Japan and Okhotsk Sea), A. hirsuta Ishimaru (southern Sea of Japan), and un-
described species ofParhyalella and Hyalella. Numerical analysis of selected characters and character states, partly
developed recently by Lazo-Wasem & Gable, supports taxonomic recognition of family Hyalellidae and its
distinction from the closely related marine family Hyalidae, and indicates degrees of morphological similarity and
probable closeness of relationship between and among its five recognized genera. Species of Allorchestes dominate
littoral marine habitats of boreal and cool -temperate open and semi-protected sand bottoms having submerged algal
and phytal mats. Species of Hyalella occur in tidal freshwater portions ofestuaries and in coastal freshwater lakes
and ponds of western North America, from Alaska to Southern California.

INTRODUCTION

The talitroidean genus Allorchestes was first pro-
posed by Dana (1849), based on species shortly after-
wards described as A. compressa Dana 1853, and A.
novizealandiae Dana, 1853, from the Australian-New
Zealand region of the South Pacific. Member species
do share with member species of talitroidean family
Orchestiidae (now Talitridae) some apomorphic char-
acter states such as maxilla 1 palp minute or lacking,
and telson entire or apically cleft. However, the name
''Allorchestes^^ (“other orchestes”) is somewhat of a
misnomer since member species are mainly aquatic
and apparently non-saltatory, not terrestrial or semi-
terrestrial or capable of jumping actively in the air .

In the North Pacific region, Allorchestes angusta
was described from the coast of California by Dana
(1856), reaffirmed by Stimpson (1857), and subse-
quently found to be the common species of the entire
region (below). In 1854-55, Stimpson described
Allorchestes rubricorniSy A. penicillatuSy and A. jap-
onicus from coastal waters of Japan. Despite the in-
adequate descriptions and lack of figures, these species
continued to be listed in Bate (1862) and even in the
recent catalogue of Ishimaru ( 1994). Character states
of the eyes of A.japonicus suggested a species of Hyale
similar to H. pontica (see Della Valle 1893), but was
later transferred to the synonymy of A. compressa Dana
by Bamard&Karaman( 1991). Allorchestes seminuda

described by Stimpson (1857) from the Central Cali-
fornia coast, has not been recognized subsequently and
may be synonymous with A. angusta Dana. Stebbing
(1899,1906) described in detail A. malleolus from
seaweed in the Yellow Sea and Sea of Japan. During
the twentieth century, three further western Pacific
species were described: A. vladimiri Derzhavin, 1937
from the Sea of Okhotsk, A. carinata Iwasa, 1939 from
Okhotsk Sea and Sea of Japan, and A. hirsuta Ishi maru,
1995, from the Sea of Japan. Tzvetkova (1990) ex-
tended the ranges of A. carinata and A. bellabella to the
Kamchatka peninsular region. Previous records from
the Japanese region were summarized by Ishimaru
(1994).

Along the North American Pacific coast. Stout
(1913) described, but did not illustrate, Allorchestes
oculatus from the Laguna Beach region of S . Califomn-
ia, a species later synonymized with A. angusta Dana.
Significant contributions to the systematics of regional
species of Allorchestes were made by Barnard, includ-
ing redescriptions of A. angusta (1952, 1954, 1970,
1979), A. carinata (1979), and a formal new descrip-
tion ofA.Z>e//a^e//a( 1974, 1979, first figured in 1954).
North Pacific species, including those of Hawaii
(Barnard 1970), were compiled by Barnard &Karaman
(1991), and those of North America by Bousfield
( 2001 ) .

^ Research Division, Canadian Museum of Nature, Ottawa, Canada KIP 6P4.

^1710-1275 Richmond Rd., Ottawa, ON, Canada K2B 8E3.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 4

In a preliminary report on species of family Hyal-
idae in the North American Pacific region, Bousfield
(1981) included eight species of A//orc/ie5re5, These
and earlier records of A. angusta and A. bellabella Bar-
nard 1974 were summarized by Ricketts & Calvin
(1968), Barnard (1975), Austin (1985) and Staude
(1987).

Based mainly on the results of regional field expe-
ditions of the National Museum of Natural Sciences
(now the Canadian Museum of Nature), the present
study extends knowledge of the systematics and distri-
butional ecology of the genus Allorchestes and other
members of family Hyalellidae on the Pacific coast of
North America.

ACKNOWLEDGMENTS

For assistance with f ^eld work the authors are greatly
indebted to regional marine laboratories, notably the
Pacific Biological Station, Nanaimo, B. C., the Pacific
Environmental Institute, West Vancouver, B. C., the
Bamfield Marine Station, B. C., and the Friday Harbor
Laboratories, WA. Much of the detail has been ac-
knowledged in previous station lists (below). Dr.
Craig P. Staude, Dr. Colin B. Levings, the late Dr.
Josephine F. L. Hart, and the late Dr. Daniel B. Quayle
were especially helpful with the field collecting.

The authors are grateful to Drs. Peter Slattery and
Charles E. O'Clair for providing valuable study mate-
rial from the Bering Sea and Aleutian Islands regions.
Dr. Nina Tzvetkova, Zoological Museum, St. Peters-
burg, provided helpful commentary and loan of valu-
able western North Pacific material. We much value
the careful research of Dr. Shin-ishi Ishimaru on
Allorchestes hirsuta in Japanese coastal waters, and for
his kind permission to reproduce his published figures
of that species ( 1995) as deemed essential to the cohe-
sion of this study.

Taxonomic work on CMN amphipod collections
was performed ini tially by ELB as a staff member of the
National Museum of Natural Sciences at the Holly
Lane Laboratory, Ottawa, during the period 1979-
1984. Original line illustrations were prepared with the
capable assistance of artist Roy E. Zittin, Cupertino,
California. Marjorie Bousfield provided translations
of the Russian literature.

The work was carred out mainly in the research
laboratories of the Canadian Museum of Nature. The
authors are especially grateful to assistant collections
manager Judith C. Price for help in retrieving, cata-
loguing, and labeling of CMN amphipod material.

MATERIALS AND METHODS.

Station lists pertinent to field material utilized in
this study are provided in Bousfield (1958, 1963,
1968); Bousfield & McAllister ( 1962); and Bousfield
& Jarrett ( 1981),

The following abbreviations are used in the text and

figures:

Al-2

antenna 1, 2

BR

-

brood lamella

CLSP

-

clothespin spine

CX

-

coxal plate

DACT

-

dactyl

EP

-

abdominal side plate

GNl-2

-

gnathopod 1, 2

HD

-

head

LFT

-

left

LL

-

lower lip (labium)

MD

-

mandible

MX 1-2

-

maxilla 1. 2

MXPD

-

maxilliped

P3-7

-

peraeopods 3, 4, 5, 6, 7

PL

-

pleopod

PLP

-

palp

RET

-

retinacula

RT

-

right

SP

-

spine

T '

-

tel son

U

-

uropod

UL

-

upper lip (labrum)

UROS

-

urosome

X

-

enlarged

im

-

immature

juv

-

juvenile

ov

-

ovigerous

subad.

-

subadult

SYSTEMATICS

Family Hyalellidae Bulycheva

Hyalellidae Bulycheva, 1957: 18. Bousfield 1982:
270; 1996: 176.

Talitridae (part) Stebbing 1906: 523. Gurjanova 1951.
Hyalidae (part) (subfamily) Barnard 1970: 268. Bous-
field 1981: 176.

Talitroidea (part) Barnard & Barnard 1983: 161.
Type genus Hyalella Smith, 1874.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 5

Genera: Allorchestes Dana, 1849: 136; Exhyalella
Stebbing, 1917, revised Lazo-Wasem & Gable 2001:
51; Marinohyalella Lazo-Wasem & Gable, 2001: 67;
Parhyalella Kunkel, 1910: 74. Laso-Wasem & Gable
(2001) have recommended suppression of the enig-
matic genus Insula Kunkel, 1910, and we herewith
concur.

Diagnosis: Body smooth, occasionally carinate or pro-
cessiferous, mainly abdominally. Antennae relatively
short. Antenna 2 longer than antenna 1, peduncle
(male) occasionally stout and proximal flagellar seg-
ments conjoint (incrassate).

Mouthparts of generalized talitroidean form. Man-
dible, left lacinia 5-6 dentate; blades of spine row few
(2-4); molar seta tending to reduction or loss. Max-
illa 1, palp minute or lacking. Maxilla 2, inner plate
with 1-2 prominent inner marginal setae. Maxilliped,
outer plate short; palp moderate, dactyl unguiform.

Coxal plates 1-4 deep, subrectangular, posterior
marginal cusp (acclivity of Barnard 1979) vestigial or
indistinct. Gnathopods usually strongly sexually di-
morphic. Gnathopod 1 (male), often modified for pre-
amplexus; dactyl and propodal spines modified for
fitting into precopulatory notch of peraeon 2 of female.
Gnathopod 2 (male) powerfully subchelate; propodal
palm smoothly convex or weakly toothed; carpal lobe
well developed, interior margin lined with comb setae.

Peraeopods 3-4 slender, distal segments posteriorly
armed with singly inserted complex spines; dactyls
simple. Peraeopods 5-7, bases broad, hind margin
lacking ‘‘surge seta” and corresponding indentation;
distal segments typically slender and weakly spinose;
dactyls small to medium, lacking median seta or lock-
ing spine; basis of peraeopod7, posterodistal lobe lying
mediad of ischium.

Epimeral side plates ordinary, plate 2 deepest.
Pleopods normal, rami not sexually dimorphic; pedun-
cle with 2-3 small retinacula; inner rsimus proximally
with claviform "clothespin" spines. Uropods 1 - 2
short, peduncular distolateral and/or distomedial spines
ordinary; rami variously (or not) marginally spinose,
apical spines occasionally striate. Uropod 3 uniram-
ous ; ramus distinct, short, notfused to peduncle. Telson
plate-like; lobes fused basally, separated by di stal notch
only, or fused totally.

Coxal gills medium, sac-like, subequal. Sternal gills
(when present) paired, sublinear.

plates large, subrectangular or ovate and distally acute;
marginal hooked setae short to medium in length.

Distributional Ecology: Within the North Pacific
marine region, species of the genus Allorchestes are
largely endemic to cool-temperate waters of both east-
ern and western coasts. The tropical and warm-
temperate genus Parhyalella is represented in Taiwan
by P. kunkeli Lazo-Wasem & Gable and in the southern
Japan Sea by an undescribed species; in the Hawaiian
Islands by P. pietschmanni Schellenberg; and on the
Pacific coast of North America (Baja California) by P.
barnardi Lazo-Wasem & Gable. Species of Hyalella
occur in coastal fresh waters of North America north
approximately to the tree line in Alaska.

Remarks: Not all systematists (e.g., Barnard & Kara-
man 1991; Lazo-Wasem & Gable 2001) accept the
original concept of family Hyalellidae and generic
inclusions that were split off from family Hyalidae by
Bulycheva (1957). Barnard (1969) subsequently cre-
ated Chiltoniinae, within the Ceinidae, to encompass
the antipodean freshwater genera Chiltonia, Austro-
chiltonia, and Afrochiltonia. The aberrant hyalid genus
HyacheliaBdmdxdi 1967: 120),tentativelyplacedwithin
the talitroidean family Najnidae by Bousfield (1982),
was also placed wi thin Ceinidae by Barnard & Karaman
( 1991). However, its true family status remains enig-
matic.

Subfamily Chiltoniinae Barnard 1969: 467 (part), is
herewith removed from the Hyalellidae. Its genera,
along with marine genera Allorchestes and Parhyal-
ella, had been placed within family Hyalellidae by
Bousfield (1982, 1996). Chiltoniin genera lack: (1) a
carpal lobe in male gnathopod 2; (2) a precopulatory
notch in peraeon 2 (female); and (3) a corresponding
modification of the propod and dactyl of gnathopod 1
(male), all suggesting separate phyletic origin.

The Chiltoniinae continued to be recognized within
family Ceinidae by Barnard & Barnard (1983) and
Barnard & Karaman (1991). However, its principal
subfamily character states, involving sexually dimor-
phic and other characters, do not conform with those of
the type genus Ceina. Subfamily Chiltoniinae may
therefore require separate family recognition within
the Talitroidea, a proposal beyond the scope of the
present study. Subfamily Hyalellinae is therefore here
considered monotypic within family Hyalellidae.

Female: Peraeon segment 2 with anterodistal pream-
plexing (precopulatory) notch (Figs IB, 6). Brood

Classificatory status of famUy Hyalellidae
Barnard & Karaman (1991) have retained some

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 6

genera of family Hyalellidae within
family Hyalidae ( sens, lat.l . appar-
ently without formal numerical
analysis. Laso-Wasem & Gable
(2001) also include the genera
Parhyalella, Exhyalella, and
Marinohyalella within Hyalidae
( sens, lat.). However, all genera of
family Hyalellidae Bulycheva are
reliablydi stinguishedfromall mem-
bers of family Hyalidae ( sens, str.)
by the following character states:

1 . Maxilla 1, palp vestigial or lack-
ing (vs. distinct, 1-2 segmented);

2. Telson plate-like, apex entire or
narrowly cleft (vs. fully bilobate,
cleft to base).

Additionally withinfamily Hyal-
ellidae: coxae 1-4 are not poster-
iorly cuspate (vs. often distinctly
cuspate); gnathopod 2 (male), car-
pal lobe is always present and dis-
tinct (vs. lacking or vestigial); per-
aeopods 3-7, distal segments are
slender, weakly spinose (vs. stout,
strongly spinose); peraeopods 5-7,
baseslackposteriormarginal “surge
seta” and pit (vs. present) ;uropod 1,
distomedial and distolateral ped-
uncular spines ordinary (vs. often
strongly developed); and uropod 3
uniramous, or ramus fused to the
peduncle (vs. occ. unequally bi-
ramous, outer ramus not fused to
peduncle). Sternal gills are present
in one supergenus (Hyalella) (vs.
lacking in all Hyalidae including
freshwater members). Addition-
ally, the preamplexing notch (fe-
male) is present in ail genera and species of Hyalellidae
and is highly variable in form (Figs. 1, 6). The
preamplex-ing notch is yet known only within a few
subgroups of Hyalidae and, when present, is relatively
simple in form.

Although accepted by Tzvetkova ( 1990), Barnard
& Karaman (1991) did not recognize the new generic
and species names proposed by Bousfield (19^1) for
North American Pacific species within families
Hyalidae and Hyalellidae (e.g., within Allorchestes).
Bousfield (2001) tentatively recognized them in a list
of amphipod species of North America, but these

names are herewith regarded as nomena nuda and
suitably replaced in this study.

Characters and Character States.

In previous studies of hyalellid and hyalid amphi-
pods, characters and states most frequently selected for
species distinction have been those of the antennae,
mouthparts, gnathopods, uropods, and telson (e.g.,
Barnard 1979; Ishimaru 1995). In this study of North
Pacific hyalellids, those characters are also utilized, but
the taxonomic significance of reproductive morphol-

AMPMPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 7

KEY TO GENERA OF HY ALELLIDAE

1. Gnathopods 1-2 (female ) subsimiiar in size and form; uropod 1, marginal spines of outer ramus
few (1-2) or lacking; right and left mandibles, molar seta present; maxilla 1, j^p 1 -segmented . . 2.
Gnathopods 1-2 (female) unlike in form and size; uropd 1, marginal spines of outer ramus well
developed (3+); right and/or left mandible lacking molar seta; maxilla 1, palp essentially lacking
4.

2. Telson lobes narrowly separated distally (tel son cleft) Allorchestes (p. 7)

Telson lobes fused to apex (telson entire) 3.

3. Sternal gills present; uropod 1, outer ramus with marginal spines, freshwater Hyalella (p. 26)

Sternal gills lacking, uropod 1, outer ramus, marginal spines absent; marine Marinohyalella

4. Antenna 2 (male), peduncle very stout, basal flagellar segments conjoint; maxilla 1, palp lacking;

gnathopod 1 (male), unguis of dactyl often bidentate; one molar seta present . . Parhyalella (p. 28)
Antenna 2 (male), peduncle normally thickened, basal flagellar segments distinct; maxilla 1, palp
a single seta; gnathopod 1 (male), unguis of dactyl simple; both molar setae lacking; Exhyalella

ogy is emphasized. Thus, during preamplexus (precop-
ulatory "carrying" of the female), the male grasps the
female anteriorly, mainly utilizing gnathopod 1 , but the
"holding" action may be assisted by the stoutly en-
larged antenna 2 (Lazo-Wasem & Gable 2001).
Gnathopod 2 (male) may assist in positioning the
female and in agonistically fending off other males
(Borowsky 1984; Bousfield & Shih 1994). The dactyl
of gnathopod 1 (male) apparently fits into a pre-
amplexing "notch" located on the anteroventral margin
of peraeon segment 2 of the female (Figs. 1, 6). The
male dactyl and female "notch" are differently modi-
fied according to the species (Figs. 2, 6). Theantero-
distal palmar spines of the propod (male) may also be
specialized as "guiding" and "locking" spines, presum-
ably facilitating the grasping and holding action by the
male (Fig. lA). The form of the distal segments of
gnathopods 1 & 2 of the male varies by species (Figs.
2,3).Thecorrespondingportionsoffemalegnathopods,
used mainly infeeding and grooming, also vary accord-
ing to the species (Figs. 4, 5).

Allorchestes Dana

Allorchestes Dana, 1849: 136. Dana 1853: 883.
Stebbing 1906: 581. Bulycheva 1957: 1 1 1 (part). Bar-
nard, 1974: 90. Bousfield, 1981: 81; 1996: 178.
Bousfield 2001: 104.

non Allorchestes Chevreux & Fage (1925, part).
Gurjanova (1951 part). Krapp-Schickel (1974, part).

Type species: Allorchestes compressa Dana, 1853
(selected Chevreux & Fage 1925).

Species: Seven species occur in the North Pacific
region, including 2 species new to science, and 1
resurrected (see Barnard & Karaman 1991: 367).
Allorchestes angusta Dana, 1949 (= A. oculatus Stout,
1913?)(Alaska to California); A. bellabella Barn-
ard, 1974; Tzvetkova 1990 (Kamchatka, Aleutians to
Oregon) \A.carinata Iwasa, 1939 (Aleutians) ; A. /»>5M/a
Ishimaru, 1995 (Sea of Japan); A. malleola Stebbing,
1899 (= A. vladimiri Derzhavin, 1937?)(Westem Pa-
cific shores, N. Japan and Sea of Okhotsk); A. priceae
n. sp. (S.E. Alaska to southern British Columbia); and
A. rickeri n. sp. (S.E. Alaska to Central California).
Incerta sedis: Allorchestes ya/7<?rtica Stimpson, 1855(=
A. compressa Dana fide Barnard & Karaman 199 1 , but
listed separately by Ishimaru 1994.)

South Pticific: Allorchestes compressa Dana, 1853
(including A. compressa subsp. "W" Barnard 1974,
and A. compressa subsp. "V" Barnard 1974); A. novi-
zealandiae Dana, 1853 (see Hurley 1957).

Diagnosis: Body small to medium large, smooth or
weakly carinate posterodorsally. Eyes medium,
rounded. Antennae generally short, antenna 2 slightly
the longer, peduncle (male) stouter than in female;
gland cone small.

Mandible, left lacinia 5-6 dentate; both molar setae
present. Maxilla 1, palp minute 1 -segmented.
Maxilliped palp, dactyl unguiform, without whip seta.

Coxae 1-4, posterior marginal cusp or process (ac-
clivity of J. L. Barnard 1979) weak or lacking; coxa 4
large, broadly subquadrate. Gnathopods strongly
sexually dimorphic. Gnathopod 1, carpus and propod
usually hammer-shaped; dactyl often modified for

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

8

A. compressa subsp. V
(per Barnard 1979)

A. novizealandiae (Thomson, 1880)
(per Hurley 1957)

A. carinata twasa (per Barnard 1979)

A, hirsuta Ishimaru, 1995

A. bellabella Barnard, 1974

4. angusta Dana, 1856 A. maileola Stebbing, 1899

Hg. 2. Allorchestes spp. Male. Gnathopod 1. distal segments.

AMPHIPACinCA VOL. 3 NO. 2. Nov. 15. 2001 9

A. compressa subsp. V
(per Barnard 1979)

A. beUabeila Barnard, 1974

A, angusta Dana (per Barnard 1979)

4. novizealandiae (Thomson, 1880)
(per Hurley 1957)

4. priceae n. sp.

Fig. 3. Allorchestes spp. Male. Gnathopod2, distal segments.

AMPHIPACinCA VOL. 3 NO. 2. Nov. 15, 2001 10

KEY TO WORLD SPECIES OF ALLORCHESTES
(Character states of reproductive morphology illustrated in figures 1-6)

1. Gnathopod 1 (male), dactyl simple, tinmodified; mostly South Pacific species 2.

Gnathopod 1 (male) dactyl modified, inflated or bifid, not simple; North Pacific only 4.

2. Body dorsally smooth; P5-6 bases little broadened; South Pacific antiboreal only 3.

Pleon mid-dorsally carinated; P5-6, bases broadened, rounded behind; North Pacific only. . .
A. carinata Iwasa (p. 13)

3. Gnathopod 1 (male), dactyl exceeding toothed palm A. novizelandiae group

Gnathopod 1 (male), dactyl normal, not exceeding regular palm A. compressa group

4. Gnathopod 1 (male); dactyl inflated, as long as palm, unguis simple; propod palm processifer-

ous; peraeopods segments, comers with multiple blunt striated spines 5.

Gnathopod 1 (male); dactyl not infiated, shorter than palm; unguis bifid; palm smoothly convex;
peraeopods 5-7, hind comers of segments 4 & 5 with few simple spines only 6.

5. Gnathopod 1 (male) dactyl very inflated, little longer than wide; propod palmar process subacute;
coxa 4 squarish; urosome 1 not (or weakly) carinated; maxilliped palp dactyl short, nail long . .

A. bellabella Barnard (p. 14)

Gnathopod 1 (male) dactyl slightly inflated, distinctly longer than wide; propod palmar process
bluntly rounded; coxa 4 broadly rounded below; urosome 1 distinctly carinated; maxilliped
palp, dactyl slender, longer than segment A. priceae n. sp. (p. 16)

6. Uropod 1, outer ramus with marginal spines; antennae elongate; gnathopod 2 (male) palm sub-

marginally lined with long setose; telson squarish, deeply cleft A. hirsuta Ishimaru (p. 18)

Character states not so 7.

7. Uropod 3, ramus blunt, short (~l/2 length of peduncle); gnathopod 2 (male), propodal palm sub-

marginally with slender spines; western Pacific A. malleola (p. 20)

Uropod 3, ramus tapering, subacute, length about equal to peduncle; gnathopod 2 (male), palmar
margin lined with short spines; eastern Pacific 8.

8. Peraeopod 5, segment 4 normal, length longer than width; angle of preamplexing notch broadly

obtuse A. angusta (p. 24)

Peraeopod 5, segment 4 broadened, width = length; angle of preamplexing notch (female) nearly
right-angled A. rickeri n. sp. (p. 25)

preamplexus with female. Gnathopod 2 (male) large,
carpal lobe large, inner margin with comb setae.

Peraeopods 3-7 regular, slender, weakly spinose;
posterior margin of basis lacking “surge seta” and pit;
marginal spines simple or striate, locking spines ab-
sent; dactyls short, unarmed.

Epimeral plate 2 deepest; pleopods with 2-3
peduncular retinacula. Uropod 1, peduncule often
lacking distal outer marginal spines; outer ramus usu-
ally lacking marginal spines. Uropod 2, outer ramus
the shorter. Uropod 3 uniramous, ramus short, apex
with setae and spine(s).

Telson plate-like, apically notched, lobes separated
distally but not diverging.

Coxal gills sac-like, subequal. Sternal gills lacking.
Female: Precopulatory notch of peraeon segment 2
large, variable, often complex; locking slit also present
in more advanced species. Brood plates narrowly
spade-shaped to subrectangular, marginal setae short.

Distributional Ecology: Species of Allorchestes Dana
are restricted almost exclusively to the Pacific Ocean.
The animals occur mainly in mats of algae and sea
grasses along the open coast or in semi -protected

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

11

A. compressa subsp. V
(per Barnard 1979)

A. novizealandiae (Thomson, 1880)
(per Hurley 1957)

A. hirsuta Ishimaru, 1995

A. priceae n. sp.

A. angusta Dana (per Barnard 1979) A. malleola Stabbing, 1899

Fig. 4. Allorchestes and Hyalella species. Gnathopod l(female) , distal segments.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

12

A. compressa subsp. V
(per Barnard 1979)

4. carinata Iwasa (per Barnard 1979)

A. novizealandiae (Thomson, 1880)
(per Hurley 1957)

A hirsuta Ishimaru, 1995

A. angusta Dana (per Barnard 1979)

Hyalella azteca (Saussure, 1858)

Fig. 5. Allorchestes and Hyalella species. Gnathopod 2 (female) , distal segments.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

13

shallows, from the lower intertidal
zone to a depth of about 30 metres.

According to Barnard ( 1979:90),
known species are endemic to cool-
er waters of north and south Pacific
regions and the genus, as presently
constituted, therefore exhibits bi-
polardiscontiguity. Subsequentde-
scription of other new species, and
current studies on this genus and on
diverse generic groupings within
the Hyalidae tend to confirm this
conclusion (Hendrycks & Bous-
field, in prep.).

However, no species of All-
orchestes sens.str. occur in both the
north and south Pacific regions or
elsewhere. Thus, all previous
species allocations in the North At-
lantic region (e.g., in Chevreux &
Page 1925) have subsequently been
transferred to generic subgroupings
within Hyalidae (see also Barnard
1974; Krapp-Schickel 1974; 1993;
Bousfield 1981; Lazo-Wasem &
Gable 2001).

a. Hyale^azleca
female (4.9 mm)

b. Xoompressa
female (10.0 mm)

C. A-cuiturta
Female (19 mm)

d. fWtyaleHasiBeM
female (5. 5 mm)

e. A. beOabeSa
fem^{6.5mm)

f. A.f»icaaa
female (9.8 mm)

g.A.maHaola . h.Aanffusto lA.rlckari

female (5.6 mm) female (4.6 mm) female (4.0 mm)

Fig. 6. Preamplexing notch of peraeon segment 2 of mature females
of Allorchestes and other genera of family Hyalellidae.

Allorchestes carinata Iwasa
(Figs. 2, 3, 4, 5, 6c, 7)

Remarks: Several species of the
Australian-New Zealand region
have been synonymized with
Allorchestes compressa Dana (an
unclarified species, fide Barnard,

1974), but such action may have
doubtful validity. CMN material
collected by ELB and attributed to A. compressa, is
relatively large, smooth-bodied, and unlike species of
the North Pacific complex. Allorchestes novizealand-
iae (Thomson) also has several synonyms (Barnard &
Karaman 1991: 381). In both southern species com-
plexes, preamplexing morphological modification of
the male gnathopods are unlike those for North Pacific
counterparts, and in some populations, the degree of
distal fusion of the telson lobes appears variable within
males and females of the same species (authors' unpub-
lished data).

Except for its completely fused telson lobes, the
genus Marinohyalella, embracing the single species M
richardi from the Mediterranean region, appears virtu-
ally identical with genus Allorchestes.

Allorchestes malleoluscarinatuslwas3iA939:28S-2S9,
figs. 23, 24, pi. 21. Bulycheva 1857: 1 18-120, fig. 44.
Allorchestes carinata Barnard, 1979: 96-98, fig. 54.
Bousfield 1981: 81, figs. 12, 13. Tzvetkova, 1990: 45-
47, figs. 5-6. Barnard & Karaman 1991: 387.

Material Examined: Bering Sea & Aleutians, Alaska:

Stn. TSC-1, Square Bay, Cyril Cove, Amchitka, Aleutian
Islands, K. Chew coll., 1968 - 5 <S(S ( 1 slide mount?); 6 99
ov,NMNS Acc. 1976-30: Ibid., coll. J. Pahnisona, 1971 - 19
(slide mount) CMN ??; Dock at Constantine Harbor,
Amchitkal., C. E. O’ Clair coll,, 1968- 1 <S (damaged (slide

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 14

mount) - MNNS Acc. No. 1976-30; Ibid.. - Id* (23 mm)
(slide mount)(rig’d. specimen), 1 9 ov (22 mm) (slide
mount)(partly fig’d) + 3 cfcf . CMN Acc. 1976-30; Transect
Plot #4, Ml 4, St. Makarios Bay, Amchitka I. , G. J. T utmark,
C. E. O’ Clair coll., 1969 - 1 cf, 1 9ov. CMN Acc. No. 1976-
30; Ibid., 1 9 , CMN Acc. No. 1976-301.

Diagnosis: Male (23.0 mm). Body large, conspicu-
ously carinate mid-dorsally on posterior peraeon, pleon
and urosome 1. Head regular. Eyes medium, sub-
reniform, black. Antennae medium long. Antenna 1,
flagellum 16-20 segmented. Antenna 2, flagellum 13-
16-segmented, distal segments slender.

Mandible: left lacinia 5-dentate; spine rows with 3
blades. Maxilla 1, palp medium. Maxilla 2, inner plate
regular, marginal plumose seta strong. Maxilliped,
inner plate with oblique apex; outer plate medium; palp
segments 2 & 3 broad, massive; dactyl short, stout.

Coxal plates 1-3 broadly rectangular, coxa 3 with
slight posterior marginal cusp. Coxa 4as broad as deep,
lower margin slightly convex. Coxa 5 shallow,
aequilobate. Coxal gills relatively large, sac-like,
smallest on gnathopod 2.

Gnathopod 1, carpal lobe medium-deep, distal mar-
gin with ~30 long comb setae; propod medium deep,
broadening distally, palm smooth, nearly transverse;
guiding spine short, inset at an angle to palm; dactyl
stout, moderately thickened proximally, unguis rela-
tively large, simple, closing on locking spine at
posterodistal angle. Gnathopod 2, carpal lobe large,
lined distally with ^40 medium comb-spines; propod,
palm short, straight, lined with double row of short
spines.

Peraeopods 3 & 4, segment 6 lined with 3-5 singly
inserted spines. Peraeopods 5-7, bases wider than
deep, rounded behind; segments 4-5 slightly broad-
ened, postero-distal apices with clusters of short blunt
striated peg-spines; segment 6 lined anteriorly with 4-
5 pairs of short spines; dactyls stout.

Epimeral plates 2 deepest; plates 2 & 3, hind comers
squarish, not produced. Pleopods large, peduncles
with 2 retinacula. Marginal spines of peduncle of
uropod 1 lacking distally, of uropod 2 regularly spaced;
rami sublanceolate, with stout terminal spine; outer
ramus lacking marginal spines. Uropod 3, peduncle
stout, with 3 stout distal spines; ramus short, thick, apex
subtmncate, bearing single spine and several short
finely pectinate setae.

Telson squarish, lobes narrowly cleft 1/3 to base.

Female ov (19.3 mm): Peraeon segment 2 (Fig. 6)
with large squarish preamplexing notch; unguisial
groove relatively broad,deep; locking slit lacking;

postero-distal lobe large, deep. Gnathopod 1, carpal
lobe medium, distally with -20 longish comb-spines;
pro-pod medium deep, length subequal to carpus, palm
gently convex, nearly vertical. Cinathopod 2, carpal
lobe large, deep, with numerous (-50) medium comb-
spines; propod medium deep, palm slightly oblique.

Brood plates on gnathopod 2 large, broad, margins
with numerous short fine hooked setae.

Distributional Ecology; Subarctic and boreal pan-
Pacific, from the northern Sea of Japan, Okhotsk
Sea, and Kamchatka peninsula through Bering Sea and
the Aleutian island chain to SE Alaska and northern B.
C. Inhabits predominantly littoral and uppermost sub-
littoral zones (Barnard 1974). Specimens from the
shores of Olga Bay, eastern Kamchatka, occurred on
sandy, stony and rocky sediments, at depths of 0.5 - 4
m. (Tzvetkova 1990).

Remarks: Allorchestes carinata is a morphologically
primitive species, superficially distinguished by its
large size, mid-dorsal low posterior carinations, and
stout clavate spines on the peraeopods. In most
character states this large species appears intermediate
between members of the North and South Pacific gen-
eric subgroupings (Hg. 17).

Allorchestes bellabella Barnard
(Figs. 2,3,4,5,6e,8)

Allorchestes bellabella Barnard, 1974: 43. Barnard
1979: 94, figs. 52(part), 53. Bousfield 1991: 81, figs.
12, 13. Austin 1985: 594. Tzvetkova 1990: 40-45, figs.
3-4. Barnard & Karaman 1991: 367.

Allorchestes subcarinata Bousfield, 1981: 81, figs. 12,

13. Bousfield 2001: 104.

non A. angusta Barnard 1954: 21-23.

Material Examined: 41 males, 61 females, in 19 lots, from
Aleutians and SE ALASKA to Southern B.C. (see map,
fig. 18).

Alaska: Plot 1A2, #19, Amchitka I., Aleutians, C. E. O'
Clair, coll., 1971- 1 9?

SE Alaska, ELB Stns., June-July, 1961: A 16, South side
Port McArthur Bay, Kuiu I - 18 €?(?, 23 99 (mostly ov.);
A32,Wood Spit, Holkam Bay - 3cfcr (1 slide mount) (fig'd.
specimen); 2 99 , NMNS 61 - 123; A7 1 , Ankau Creek mouth
- 1 cf, 3 99 ov; A75, Kayak L, Wingham I, 1 9 ov (slide
mount); A 80, Nuchek Beach, Hinchenbrook I., 2 cfcf (1
slide mount), 1 9 (slide mount); A 136, Head of Thumb Bay,
Resurrection Bay - 1 cf, 1 9 ; A171, Puffin Bay, Baranof I.
(2 lots) - lcf,4 99 .

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

15

MXPD^

Fig. 7. Allorchestes carimta Iwasa, 1539. Male (23.0 mm); female 19.3 mm).
Low Intertidal, Amchitka, Alaska.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

16

British Columbia

Queen Charlotte Islands, ELB Stns., July, 1957.

H2, Kiusta Village, Parry Passage - 1 Cf , 14 99 -
North Central Coast, ELB Stns, July, 1964:

H4, Baker Pt, Aristazabel I., - 1 Cf; H23 Deadman’s Cove,

(21ots)-4C?C?,2 99 .

Southern Vancouver I. , ELB Stns, July, 1977.

B5a, Witty’ s Lagoon - 2 CfCT (slide mount, fig’d specimen),

5 99 (1 slide mount, partly fig’d); NMCC 1883-1520.

Diagnosis: Male (6.0 -10 mm). Body medium large,
very slightly carinate mid-dorsally on pleon segments
1-3. Head regular. Eye broadly subreniform, black.
Antennae medium long. Antenna 1, flagellum 9-14
segmented. Antenna 2, flagellum 11-15 segmented,
distal segments slender.

Mandible: left lacinia 5 1/2-dentate; left spine row
with 3 blades, right with 2 blades. Maxilla 1, palp
medium, distinct. Maxilla 2, inner plate regular, mar-
ginal plumose seta strong. Maxilliped, inner plate with
oblique apex; outer plate medium; palp segments 2 &
3 not enlarged; dactyl short.

Coxal plates 1-3 each with slight posterior marginal
cusp, coxae 2-3 deeply subquadrate, lower margins
gently rounded. Coxa 4 as broad as deep, lower margin
convex. Coxa 5 shallow, aequilobate. Coxal gills
medium, sac-like, subequal, slightly smallest on
gnathopod 2.

Gnathopod 1, carpus shallow, posterior lobe lined
distally with ~15 short comb-setae; propod medium
deep, narrowing slightly distally to a transverse palm
that is produced sharply forward at the posterodistal
angle; propodal guiding spine short, set nearly horizon-
tally above the posterior margin; propodal locking
spine short, set horizontally at base of palmar process;
dactyl strongly inflated, unguis small, simple, closing
below locking spine. Gnathopod 2, carpal lobe large,
with ~30 medium long marginal comb-setae; propod,
posterior margin relatively short, distally straight; palm
straight, submarginally lined with double row of me-
dium spines;dactyl lined posteriorly with -20spinules.

Peraeopods 3 & 4, posterior margin of segment 6
with 3-4 singly inserted complex spines. Peraeopods
5-6, bases not wider than deep, rounded behind; seg-
ments 4-5 broadened distally, posterodistal apices with
cluster of striated peg-spines; dactyls stout. Peraeopod
7, basis broadening posterodistally; posterodistal lobe
truncated; segment 6, anterior margin lined with 4-5
pairs of spines.

Epimeral plate 3, hind comer slightly acuminate.
Pleopod peduncles with 2 retinacula. Uropodsl & 2,

outer marginal spines of peduncle regularly spaced;
rami slightly tapering, outer ramus lacking marginal
spines. Uiopod 3, peduncle medium stout, with 2
distally striated spines; ramus medium short, thick,
tapering to blunt apex bearing 1 small spine and 8-10
short setae.

Telson squarish, lobes narrowly cleft 1/4 to base.
Female (5.0 mm): Peraeon segment 2 (Fig. 6) with
large regularly concave preamplexing notch; unguisial
groove lacking; locking slit short, vertical, dorsad of
mid-point of notch; postero-distal lobe large, deep.
Gnathopod 1, carpal lobe medium, distally with ~20
longish comb spines; propod medium deep, length
subequal to carpus, palm very slightly parachelate.
Gnathopod 2, carpal lobe large, deep, lined marginal-
ly with -40+ medium comb-spines; propod medium
deep, palm slightly convex.

Brood plates medium large, margins with numerous
short fine hooked setae.

Distributional Ecology: Tzvetkova (1990) encoun-
tered live material on rocky and sandy sediments in
Olga Bay, Kamchatka, at depths from 1.2 - 12 m.
Females (7- 10 mm) with eggs in the marsupium were
noted on 16 Aug., 1975 (Tzvetkova 1990). Known
from islands of Beringia, Aleutian Islands, S. E. Alaska
and B. C. south to Oregon, sometimes encountered in
the littoral plankton (Barnard 1979).

Remarks: The propod of gnathod 1 (male), with strong
posterio-distal tooth, inflated dactyl, and simple unguis
is distinctive of the species. It appears most closely
related to A. priceae (Figs. 9, 17).

Allorchestes priceae n. sp.

(Figs. 2,3,4,5,6f,9)

Allorchestes urocarinata Bousfield, 1981: 81, figs.

12, 13. (nomen nudum).— Bousfield 2001: 104.

Material Examined: SE Alaska, ELB Stns., 1961:

A171, Puffin Bay, Baranof I., July, 1980 - Holotype cf,
(slide mount), CMNC 2001-0025; Allotype 9 (slide mount)
CMNC 2001-0026, 1 , 1 QParatypes, CMNC 2001-0027.

Ibid . - -120 juvenile and immature specimens, Topotypes.
British Columbia:

Queen Charlotte Islands ELB Stns., 1957:

Stn. H2, Langara L, Queen Charlotte Islands - several imma-
ture specimens (possibly misidenlified as A. bellabellal)
Southern Vancouver I., ELB Stns., 1970:

Botanical Beach, Cape San Juan, low intertidal sand and
algae, July - 1 Cf , 1 9 -

AMmiPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 17

Fig. 8. Allorchestes bellabella Barnard, 1974. Male (6.0 mm); female (5.0 mm).

Witty’s lagoon, Vancouver L, B. C.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

18

Diagnosis: Male (13.0 mm). Body medium large,
conspicuously carinate mid-dorsally on urosome seg-
ment 1. Head regular. Eye medium large, broadly
subreniform, black. Antennae medium long. Antenna

1, flagellum 16-17 segmented. Antenna 2, flagellum
13-segmented, distal segments slender.

Mandible: left lacinia 5 dentate; right spine row
with3 blades. Maxilla Lpalpmedium. Maxilla 2, inner
plate regular, marginal plumose seta strong. Maxilliped,
inner plate with oblique apex; outer plate medium; palp
segments 2 & 3 broad, massive, dactyl slender.

Coxal plates 1-3 each with slight posterior marginal
cusp, lower margins gently rounded. Coxa 4 as broad
as deep, lower margin strongly convex. Coxa 5 shal-
low, aequilobate.

Gnathopod 1, carpus deep; propod medium deep,
broadening distally to transverse palm having rounded
tumescent posterodistal process; median amplexing
tooth striated, set at angle to palm; dactyl moderately
thickened medially, unguis small, simple. Gnathopod

2, carpal lobe large, with short marginal comb-setae;
propod, posterior margin relatively short, distally
straight; palm straight, lined with short spines.

Peraeopods 3 & 4, posterior margin of segment 6
with 3-4 groups of small complex spines. Peraeopods
5-7, segments 4-6 little broadened, margin and apices
with cluster of short blunt peg-spines.

Epimeral plates 2 & 3, hind comers slightly acumi-
nate. Uropods 1 & 2, outer marginal spines of peduncle
regularly spaced; outer ramus lacking marginal spines.
UropodS, ramus medium short, thick, apex blunt, with
short setae.

Telson squarish, lobes narrowly cleft 1/3.

Female ov (9.8 mm):J^eraeon segment 2 (Fig. 6)
withmediumlargesharplyconcavepreamplexingnotch;
unguisial groove large, broad, deep,narrowing sharply;
locking slit medium long,oblique,set dorsad of unguis-
ial groove; posterodistal lobe large, deep, strongly
convex anteriorly.

Gnathopod 1 , carpal lobe medium, distally with long
comb -spines; propod medium deep, length subequal to
carpus, palm gently convex, transverse. Gnathopod 2,
carpal lobe large, deep, with numerous medium comb-
spines; propod medium deep, palm slightly oblique.

Brood plates medium large, margins with numerous
short fine hooked setae.

Etymology: The species is named in honour of Judith
C. Price, CMN assistant collections manager of crusta-
ceans, whose ready help in recording, retrieving, and
cataloguing has greatly expedited the description of

many new taxa in the museum collections of Pacific
coastal marine amphipods.

Distributional Ecology: Recorded at outer coastal
stations from SE Alaska, the Queen Charlotte Islands,
and southern Vancouver I., in coarse sands of the lower
intertidal and inshore shallows. The distribution is
limited narrowly within SE Alaska to southern B. C.,
but might also be expected in suitable coastal habitats
of the northwestern United States.

Remarks; This medium-sized species is distinctive in
its relatively long, slender antennae, mid-dorsally
carinated urosome segment 1, and partly swollen sim-
ple-tipped unguis of gnathopod 1 (male). In balance of
character states it appears most closely allied with A.
hellabella Barnard (see also Hg. 17).

Allorchestes hirsuta Ishimaru
(Figs. 2, 3. 4, 5, 10)

Allorchestes hirsuta Ishimaru, 1995: 46, figs. 1-4.
Material Examined: None.

Diagnosis: Male (10.3 mm). Female ov (10.0 mm).
Ishimaru (1995) has provided full description and
figures of both sexes of this medium sized species from
the southern Sea of Japan. His figures, some of which
are reproduced here, also illustrate character states of
pleopods, brood plates, and other appendages that are
of generic as well as species diagnostic signficance,
and seldom figured and/or descibed by other authors.

Dr. Ishimaru did not illustrate the preamplexing
notch of the female of A. hirsuta. However, he did
examine this taxonomic feature and believed "the notch
of A. hirsuta (to be) relatively similar to that of A.
rickeri, (but) is subacute and smaller" (pers. com.).

Distributional Ecology: On floating Sargassum north
of Oki Island, southern Japan Sea. Not yet known from
the North American Pacific coast.

Remarks: The species belongs to the North Pacific
cluster of relatively small advanced species having
hammer- shaped male gnathopod 1 with split-tipped
unguis . It differs from A. angusta Dana in the elongate
antennae, strongly setose palmar margin of gnathopod
2 (male), carinate urosome, and marginally spinose
outer ramus of uropods 1 & 2, among other character
states.

uxxrx

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

19

Fig. 9. AUo^he^espri^ae n. sp. Male (13.0 mm); female ov (12.0 mm).
Stn A170, Puffin Bay, Baranof I., SE Alaska.

AMPHIPACIFICA VOL. 3 NO. 2. Nov. 15, 2001 20

Allorchestes malleola Stebbing

(Figs. 2, 3, 4, 5, 6g, 11)

Allorchestes malleolus Stebbing, 1899: 409, pi. 33A.
Iwasa 1939: 285, figs. 20-22. Tzvetkova 1967: 183.
Bousfield 1981: 81, figs 12, 13. Ishimaru 1994: 67.
Allorchestes vladimiri Derzhavin, 1937: 95-95, pi. 5
fig.2.Gurjanova 1951 : 822 ,fig. 575. Bulycheva 1957:
115, fig. 43.

Material examined: USSR: Stn. 1, Anama Bay, Shikotan,
S.Kurile I.,O.G.Kussakincoll.,July5, 1955-2cfd' (1 slide
mount), (fig’d specimen); 3 99 (1 slide mount), NMCC,
ACC 1979-267.

Japan: Usuziri, Hokkaido, A. Hirayama coll., June 26/78
- cS (9.0 mm) (slide mount), 9 ov (8.0 mm (slide mount),
CMN collns.

Diagnosis: Male (5-9 mm). Body medium, mid-
dorsally smooth. Head regular. Eye medium large,
broadly subreniform, black. Antennae short to me-
dium. Antenna 1, flagellum 9-12 -segmented. An-
tenna 2, flagellum 9-10-segmented.

Mandible: left lacinia 5 1/2 dentate; spine rows with
2-3 blades. Maxilla 1, palp medium. Maxilla 2, inner
plate regular, marginal plumose seta strong. Maxilliped,
inner plate with oblique apex; outer plate medium
broad; palp segments 2 & 3 broad, massive, dactyl
slender.

Coxal plates 2-3 subrectangular, each with slight
posterior marginal cusp. Coxa 4 squarish, not wider
than deep Coxa 5 shallow, aequilobate. Coxal gills
medium large, subequal, sac-like, slightly smallest on
gnathopod 2

Gnathopod 1, carpal lobe medium, lower margin
lined with -12+ medium-long split-tipped, finely pec-
tinate comb setae; propod medium deep, broadening
distally to nearly vertical palm that is proximo-medi-
ally convex; guiding tooth striated, elongate, set verti-
cally (parallel to, or at a slightly reverse oblique angle)
to palm, locking spine shorter, slightly sinuous, tip
reaching lower palmar margin; dactyl short, moder-
ately thickened proximally, ungui s small , tip unequally
bifid, closing above the locking spine. Gnathopod 2,
carpal lobe large, with -20 short marginal comb-setae;
propod, posterior margin relatively short, distally
straight; palm straight, submarginally lined with dou-
ble row of short spines; dactyl, inner margin lined with
fine spinules.

Peraeopods 3 & 4, posterior margin of segment 6
with 3-4 small spines. Peraeopods 5-7, bases broad,

(broaderthandeepinperaeopod7),hindmargin weakly

scalloped; segments 4-5 slightly broaded distally,
posteixxJistally produced, apices with clusters of dist-
ally striated spines; segment 6, anterior margin with 3-
4 pairs of spines; dactyls medium short.

Epimeral plates 2 & 3, hind comers sligthly acumi-
nate, not produced. Pleopod peduncles with 3 retinac-
ula. Uropodsl & 2, rami tapering, apex with 1 stout
striated spine and several small accessory spines; outer
ramus lacking marginal spines; peduncle of uropod 1
lacking marginal spines in distal half. Uropod 3,
peduncle short, deep, with 2-3 stout, striated, apico-
distal spines; ramus medium short, thick, apex blunt,
armed with 1 spine and 6 finely p»ectinate short setae.

Telson squarish, lobes narrowly separated, cleft 1/3-
1/2 to base.

Female ov (4.5-8 mm): Peraeon 2, pre-amplexing
notch shallow, angle very obtuse (140°+); ratio of
anterior: posterior lobes about 45:55); locking slit di-
rectly dorsad. Coxal plates large, deep; coxa 4 deeper
than broad.

Gnathopod 1, carpal lobe medium, distally with -14
longish finely split-tipped comb spines; propod me-
dium deep, length about twice depth and slightly longer
than carpus, palm very slightly parachelate . Gnathopod
2, carpal lobe large, deep, distally line with -24 me-
dium comb-spines; propod slightly deeper than in
gnathopod 1, palm transverse.

Brood plate on peraeopod 2 large, distally sharply
acute; margins with numerous medium length setae.

Distributional Ecology: Northern Sea of Japan,
Okhotsk Sea, to Kamchatka (Tzvetkova) low intertidal
to depth of 3-4 m (Derzhavin). Aleutian islands to SE
Alaska?

Remarks: This species has been synonymized with A.
angusta Dana, 1856, which it most closely resembles.
However, it is reliably distinguished by morphogical
characters of the key (p. 10), and by its non-overlapping
distribution in the central North Pacific region.

Bulycheva’s (1957) figures of the body, uropod 3
and telson identified by her as A malleolus Stebbing,
1 899, are apparently adapted directly from Derzhavin’s
figures of the 6-7 mm holotype male and 7-8 mm
allotype female of A. vladimiri. However, her indi-
vidual (enlarged) figures of the antennae, gnathopods,
peraeopods, and mouthparts may have been taken from
another specimen and possibly another species, since
significant differences may be discerned between her
figures and the corresponding individual figures of
Derzhavin (1937). Since this material has not been re-

AMFWIPAaHCA VOL. 3 NO. 2. Nov. 15, 2001

21

Fig. 10. Allorchesteshirsuta Ishimaru, 1995. Male ( 10.3 mm); female (10.0 mm),
oouthem Sea of Japan (after Ishimaru, 1995)

AMHBPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 22

Fig. 11. Allorchestes malleola Stebbing 1899. Male (5.0 mm); female ov (4.5 mm).
Shikotan, Kurile Islands, Okhotsk Sea

AMPHIPAaFICA VOL. 3 NO. 2. Nov. 15, 2001

23

Mg. 12 . AUorc^stes ^gusta Dma. Male (5.2 mm); female ( 4.3
atn. P704, Amphitrite Point, Vancouver I., B. C.

AMPMPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 24

examined, the synonymy is tentatively retained here.

Present study material of A. malleola Stebbing from
Hokkaido, Japan, kindly supplied by Dr. A. Hiwatari,
most closely resembles the individual parts figured by
Bulycheva (1957) and is here considered to be that
species. North American material from SE Alaska
west to the Aleutian island chain has been identified as
A. angusta, with character states trending to those of A.
malleola. Thus pleopod peduncules of northern speci-
mens tend to have 3 (rather than 2) retinacula.

However, material from the western Pacific identi-
fied and figured by earlier authors as A. malleola and A,
vladimiri differs from A. angusta in the more slender
form and positioning of amplexing spines of gnathopod
1 (male); the more rectangular form of coxae 2-4, the
broader basis of peraeopod 7, the shorter blunter ramus
of uropod 3, and the more obtuse angle of the pre-
amplexing notch of the mature female (FI g . 6) . Nagata
(1965) also records a 9 mm male specimen from the
Seto Inland Sea and an earlier record from Hokkaido
both of which may may prove to be A. malleola.

Allorchestes angusta Dana
(Figs 1, 2, 3, 4, 5, 6h, 12)

Allorchestes angustus Danei, 1856: 177. Barnard 1952:
20-23, pi 5, figs. 2-6.

Allorchestes angusta Barnard 1974: 42; 1979: 91 , figs.
50-52 (part). Bousfield 1981: 81, figs. 12, 13; 1996:
178, fig. 1.

non Allorchestes angustus Barnard 1954: 21-23. pi. 21
(= A. bellabella).

Allorchestes oculatus Stout, 1913: 651?

Material Examined: (About 600 specimens in 1 13
lots, SE Alaska to California. Numbers of specimens
in parentheses).

Alaska:

Aleutian Islands: Stn 1 A-2, Plot32, Amchitkal., Alaska, C.
E. O’ Clair coll., 1974 - 6 subadult specimens (large dark
reddish gray brown ring around eye).

Izenbak Lagoon, Unimak I., N. A. Powell coll., 1969 - 1 C?
(slide mount), NMNS 69-319.

SE Alaska: ELB Stns., June-July, 1961: A2(5); A7 (24); A8
94); A9 (5); A18 (6); A25 (1); A33, 2 lots (33 specimens -
1 slidemount); A57(1);A86(1); A88(4);A92(1);A99(14);
A114(5);A105(9); A115(1);A121 (4);A131 (1);A136
(29) A 140 (9) ; A 1 53 (3 1 ). Dixon Harbor, K. E. Conlan coll. ,
June 18/89 - 1 male (5.3 mm).

British Columbia:

Queen Charlotte Islands, ELB Stns. 1957: H5-2 lots (36);
H8a(65 cfcf&99);H9(l);W3a-21ots (4); W5 (1); W15
- 2 lots (2).

North Central coast, ELB Stns., 1964: HI, West Beach,
C:alvert I., July, 2 lots - 2 crd* , 299; H5 (2); H6 (2) H7 (2);
H10(3)-H11(25):H12(9); H18(2);H23(4); H26(1);H29
(1);H35(1);H40(6);H44(14);H48(20);H57(7). ELBStn.
P14, Porpoise Harbour, June, 1961 - 31 (S(S, 99-
South Central Coast, ELB Stns.. 1955: Ml -Savary I. (8);
M3 (6); M5 (27); Mil - White Rock (11).

Northern Vancouver Island, ELB Stns., July-Aug., 1959:
N13 (2); 01 (2); 02a (9); 06 (1 d'CJ', 2 99 - 1 slide mount);
V4b (1); VI 1 (1); V19, 2 lots (25); V27, 2 lots (4).

Southern Vancouver L, ELB Stns., July, 1955:

F4a (2); F6 (5); G2 (28); GIO, 2 lots (58); G13, Piper’s
Lagoon. 2 lots - 54 C?Cf.99; P2 (D; Glia, SE End
WickininnishBay- KJ (slidemount). ELB Stn.H44, Brady’s
Beachjuly 1964 - 1 <? (slide mt).

ELB Stns., July , 1970: P704, Amphitiite Point - 29 speci-
mens inch 1 (S (5.2 mm); 1 9 (4.3 mm), (fig’d specimens);
P705 (14); P708 (107 cfcJ, 99); P717 (1).

ELB Stns., May. 1975: Pic (4); P13 (40); P19a (5); P25 (1).
ELB Stns., May, 1976: B3 (1); B4(l).

ELB Stns., May, 1977: B5 (4); B6b, 2 lots (6); B16 (10).
Other B.C. material: Anthony I., Queen Charlotte Ids., G.
C. Carl coll., 1956 - 19; Stn. 13, Gulf Islands, D. Kittle coll.,
1972 - 50CfCf,99; Parry Bay, near Weirs Beach, KDH coll.,
1974 - 4 juveniles, BCPM 974-182-8; Stn. 6, Chemainus
estuary, D.V. Ellis coll., 1976 - 1 cf; Juene Landing, R.
Coates coll., 1976, 2 lots - 2 specimens (slide mounts),
RBCM Nos. 980-486; Salmon R. estuary, Vancouver I., R.
Coates coll., 1976, 2 lots- 17 cJcf, 99 (slide mount), RBCM
No. 977-471-8; Ibid - 33.

Wash.-Oregon, ELB Sms., July-Aug., 1966: W4 (9); W5

(10;W7(l);W8(30);W16(10);W28(l);W30,21ots(17);

W39 , 2 lots (7).

California: ELB Stn. C3, Baker's Beach, San Francisco,
July, 1959 - 1 juv.; Pt. Loma, San Diego, W. Klawe coll.,
1956 - 2 CfCf; Mission Bay, San Diego, W. D. Clarke coll.,
1956- 1 (S

Mexico: Turtle Bay, B. C., W. Klawe coll., 1956 - 1 (S.

Diagnosis: Male (5.2 mm; to 7 mm - Barnard 1979).

Body small to medium, mid-dorsally on urosome
smooth. Head regular. Eyes medium large, broadly
subreniform, black. Antennae medium. Antenna 1,
flagellum 1 1-12 segmented. Antenna 2, flagellum 1 1-
13-segmented, distal segments slender.

Mandible: left lacinia 5-dentate; left and right spine
rows with 2 blades. Maxilla 1, palp small. Maxilla 2,
inner plate regular, marginal plumose seta strong.
Maxilliped, inner plate with oblique apex; outer plate
medium; palp segments 2 & 3 broad, massive, dactyl
slender, unguis short.

Coxal plates 1-3 each with slight posterior marginal
cusp, lower margins sub-truncate Coxa 4 as broad as
deep, lower margin convex. Coxa 5 shallow,
aequilobate. Coxal gills medium large, sac-like, small-

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15. 2001 25

est on peraeopod 2.

Gnathopod 1, carpal lobe lined with 9-10 medium
finely pectinate comb setae; propod medium deep,
broadening distally to slightly oblique palm that is
convex medially; guiding spine short, thick, apically
striated, set vertically to palm; dactyl short, moderately
thickened proximally, unguis asymmetrically bifid,
closing on locking spine. Gnathopod 2, car^I lobe
large, with24closely set marginal comb-setae; propod,
posterior margin medium long, palm nearly straight,
lined with longish spines; inner margin of dactyl lined
with 16 spinules .

Peraeopods 3-4, posterior margin of segment 6 with
4-5 single small spines. Peraeopods 5-7, bases rela-
tively shallow, deeper than broad; segments 4-6 little
broadened, posterodistal margins and apices with pairs
or small clusters of simple spines.

Epimeral plates 2 & 3, hind comers sligthly pro-
duced, acute. Pleopod peduncles each with 2 retinacula.
Uropodsl & 2, marginal spines lacking on outer ramus
and distally on peduncle of uropod 1. Uropod 3,
peduncle short, with 1-2 apical spines; ramus slender,
tapering to subacute apex bearing a spine and 2-3 short
setae.

Telson squarish, lobes narrowly cleft 1/3 to base.

Female ov (4.5 mm): Coxal plates 2-4 large, deep;
coxa 4 deeper than broad. Peraeon segment 2,
preamplexing notch medium shallow, located nearly at
mid point of lower margin; notch angle obtuse (-125°);
locking slit positioned closely dorsad ; unguisial groove
lacking

Gnathopod 1, carpal lobe medium, distally with
longish comb spines; propod medium deep, length
subequal to carpus, palm gently convex, nearly trans-
verse. Gnathopod 2, carpal lobe large, deep, with
numerous medium comb-spines ; propod medium deep,
palm slightly oblique.

Brood plates medium large, margins with numerous
short fine hooked setae.

Distributional Ecology. Small specimens (males to 7
mm) occur from S. California north to British Colum-
bia. Slightly larger animals (8-9.3 mm) from SE
Alaska and the Aleutian chain trend toward the charac-
ter states of, but remain distinct from, those of A.
malleola of the western Pacific. According to Barn-
ard (1979), the species is phycophilous and mainly
intertidal, rarely subtidal.

Remarks: Barnard (1979) has synonymized the west-
ern Pacific material from Japan and USSR with the N.

American A. angusta, Dana’s type is from California
so that A. malleolus and A. vladimiri would be junior
synonyms. However, taxonomic evidence for the
distinctness of A. angusta includes the differentform of
the preamplexing notch of the female, as described .

Barnard (1979) and Barnard & Karaman (1991)
have included Allorchestes oculatus Stout, 1913, from
Laguna Beach, southern California, in the synonymy
of A. angusta Dana. Stout's unillustrated original des-
cription of the mature male of A. oculatus detailed its
colour in life, and the large size of the eyes that nearly
meet mid-dorsally. The brief outline otherwise pro-
vided little taxonomically useful information. Though
Dr. Barnard apparently examined fresh material from
south of Pt. Conception, such conspecificity would
seem unlikely. Thus, the range of well-illustrated ma-
terial of A. angusta extends from Central California,
dominantly through British Columbia, to Alaska, yet
very few inshore inverbrates occur both in the Bering
Sea ( Alaskan Aleutian Chain) and in southern Califor-
nia, south of Pt. Conception.

For the present study, CMN collections contain a
single 5.0 mm male specimen of Allorchestes sp. from
Mission Bay, San Diego, collected by C. D. Clarke,
June, 1956. Examination of the slide-mounted ap-
pendages revealed several differences from northern
specimens of A. angusta of slightly larger size (Fig. 1 2).
Espiecially noticeable in the San Diego specimen were:
(1) the relatively large eyes; (2) the relatively short
antennal flagella; (3) the relatively short palmar margin
of the propod of gnathopod 1 ; (4) the elongate ischium,
and small, weakly spinose propod of gnathopd 2; (5)
the squarish coxa 4; (6) the relatively large and deep
posterior lobes of coxae 5 & 6; and (7) the relatively
short propods (segment 6) of peraeopods 5-7. How-
ever, the limited nature of this material, lacking fe-
males and comparative series of specimens, renders the
nature of these differences inconclusive. The authors
therefore tentatively follow Barnard's synonymy of
Allorchestes oculatus within A. angusta, pending fur-
ther careful study of extensive regional material.

Allorchestes rickeri n. sp.

(Figs. 2, 3, 4, 5, 6i, 13 )

Allorchestes parva Bousfield, 1981 (nonien nudum):

81 ,fig5 12, 13.— Bousfield 2001: 104.

Material Examined: Approx. 135 <S(S, 155 99, 20 juv., in
41 lots, Northern B. C. to California. Number of specimens
in parentheses).

AMPHIPAanCA VOL. 3 NO. 2. Nov. 15, 2001 26

Alaska:

SJ:.Al«iska: ELB Stns., June-July, 1961: A22 (1), A80 (42
CJCJ, 99). A 140 (3).

British Columbia:

Queen Charlotte Islands, ELB Stns. , 1957 : W 1 (2) ; W8 (2) ;
W11(8);W12(2).

North Central Mainland, ELB Stns., July, 1964: H23 (18;
H41(18);H50(3).

North Vancouver I., ELB Stns., July, 1959: V27 (1).

South Vancouver I. ELB Stns., July-August, 1955: P6a,
South end Wickaninnish Bay - 81 cJc?, 99 (slide mount);
P6e (2). ELB Stns., July, 1970: P708, 2 lots (15); P714, CIo-
oose,21ots--350d’d‘. 99- ELB Stns., May, 1975:P14b(9).
ELB Stns., May, 1977: B6b (2 Cfd'); B 1 la (6), 2 lots (35);
B12b(2);B19b(10).

Other B, C, material: Pachena Bay, 20 ft. sand, P. Slattery
& C. Oliver coll., Apr. 17/83 - 42 cTc?, 5299, 20 juv. (slide
mounts); Port Louis, EAH coll., Jan 20/90 - CT, 9 (mating
pair).

Wash.-Oregon,ELB Stns., July-Aug., 1966: W8(l); W18,
Cf(3.0mm); 9ov(3.0mm);W22,21ots (3); W33(1);W34
(6); W35 (1); W36 (7); W40 (2); W46 (4); W57 (1); W61,
Neskowin Beach, Low intertidal -Id" (3.0 mm) (slide
mount), Holotype, CMNC 2001-0028; 9 ov (3.0 mm)(slide
mount), Allotype,CMNC 2001-0029; 2dd', 2 99 ,
Paratypes, CMNNos.2001-0030;lotNo.2--250cJ'd, 99
Topotypes. W63 (1).

California: Stn 89-1-3, Spanish Bay, Monterey peninsula,
K.E.Conlan coll., Jan. 2/89 -Id (5.8 mm), 1 9 -

Diagnosis: Male (3.0 mm): Body small, mid-dorsally
smooth. Head regular. Eye medium, broadly sub-
reniform, black. Antennae short. Antennal, flagellum
9-segmented. Antenna2,flagellum9-segmented, distal
segments slender.

Mandible; left lacinia 6-dentate; spine rows with 3-
4 blades. Maxilla 1, palp small. Maxilla 2, inner plate
regular, marginal plumose seta strong, Maxilliped,
inner plate with oblique apex; outer plate medium; p>alp
segments 2 & 3 medium broad, dactyl longish, slender.

Coxal plates 1-3 with very weak posterior marginal
cusp, lower margins very shallowly rounded. Coxa 4
large, nearly as broad as deep, lower margin evenly
convex. Coxa 5 shallow, aequilobate.

Gnathopod 1, carpus deep; propod medium deep,
broadening distally to a short convex palm, guiding
spine short, striated, set at angle to palm; dactyl thick-
ened basally, unguis unequally bifid, closing on short,
thick locking spine. Gnathopod 2, carpal lobe short,
with short marginal comb-setae; propod, posterior
margin relatively short, distally straight; palm nearly
straight, lined with short spines.

Peraeopods 3-4, posterior margin of segment 6 with
2-3 groups of small complex spines. Peraeopods 5-7,

bases broad, rounded; segments 4-6 broaded,
posterodistal margin and apices with cluster of short
simple spines.

EpimeraJ plates 2 & 3, hind comers nearly square.
Uropod 1, peduncle with proximal outer marginal
spines only, distally bare; outer ramus lacking marginal
spines. Uropod 3, peduncle thick, with 2 distal spines;
ramus shorter, slender, tapering to short-setose apex.
Telson subrectangular, lobes narrowly cleft 1/4.
Female ov (3.0 mm): Peraeon segment 2 (Fig. 6),
preamplexing notch medium, located anteriorly on
lower margin; notch angle steep, slightly obtuse
(-105®); locking slit positioned posteriorly dorsad;
unguisial groove lacking; posterior lobe relatively
shallow.

Gnathopod 1, carpal lobe medium distally with
longish comb spines; propod medium deep, length
subequal to carpus, palm gently convex, vertical.
Gnathopod 2, carpal lobe large, deep, with numerous
medium comb spines; propod medium shallow, palm
slightly parachelate.

Brood plates medium large, margins with numerous
short fine hooked setae.

Etymology: The patronym recognizes the outstanding
career contributions of the late W. E. Ricker to the
aquatic biological and fisheries sciences of North
America.

Remarks: In nearly all character states, Allorchestes
rickeri is clearly a member of the angusta group.
However, it is distinctive in the relatively large coxal
plates 1-4 , shallow propod of gnathopod 1 (male);
short, broadened segments 4& 5of peraeopods 5-7 and
the parachelate gnathopod 2, and relatively sharply
angled pre-amplexing notch in the female.

Hyalella Smith

Hyalella S. I. Smith, 1874: 645. Stebbing 1906: 574.
Bulycheva 1957: 181. Barnard & Barnard 19^: 708.
Bousfield 1996:183; 2001:104. Witt & Hebert 2000:
687.

Type species: Amphithoe azteca Saussure, 1858.

Species (North America): Hyalella (Hyalella) azteca
(Saussure, 1858);//.(//.) inermis Smith, 1874;//. (//.)
longicornis Bousfield, 1996; H. ( H.) muerta Baldinger,
Shepard, & Threloff, 2000; (H.) montezuma Cole &
Watkins, 1977; H. (H.) sandra Baldinger, Shepard, &

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15. 2001 27

Fig. 13. Allorchestes rickeri^ n. sp. Male (3.0 mm); female ov (3.0 mm).
Stn. W61, Neskowin Beach, Oregon. Low intertidal.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 28

Threloff, 2000; K (K) texana Stevenson & Peden,

1973 .

Diagnosis: Bodysniooth,mucronate,orprocessiferous.
Antennae short to medium. Antenna 2 (male),
peduncular segments regular, not stoutly expanded;
flagellar segments not incrassate (conjoint); gland cone
large, prominent.

Mouthparts regular. Mandible: molars with distal
plumose seta; left lacinia 5-dentate. Maxilla 1, palp
small, 1 -segmented. Maxilliped, palp unguiform,
dactyl without whip.

Coxal plates 1^ deep, subquadrate, lacking poster-
ior marginal cusps. Coxae 5 & 6 posterolobate. Gnath-
opods strongly sexually dimorphic; gnathopod 1 , dactyl
basally stout, unguis simple; propodal postero-distal
spines unmodified. Gnathopod 2 (male) propod large,
palm oblique. Peraeopods 3-7 slender, spines simple;
dactyls simple. Coxal gills large, sac-like, subequal.
Paired sternal gills ventrally on peraeon segments (2)
3-7.

Pleopod peduncles with 2 retinacula. Uropods 1 & 2,
rami marginally spinose. Uropod 3 uniramous; ramus
slender, apex with slender spines.

Telson entire; lobes fused apically.

Female: Peraeon segment 2, pre-amplexing notch
regular, simple. Gnathopod 1 regularly subchelate,
propod shorter than carpus. Gnathopod 2 subchelate to
parachelate; propod longer than carpus.

Brood plates relatively short and broad, marginal
setae short to medium.

Hyalella azteca (Saussure)

(Figs. 4, 5, 6a, 14)

Hyalella (H.) azteca Bousfield 1996: 183 (and
synonymies).

Material Examined:

SE Alaska: ELB Stns, June-July, 1961: A 166, Leo Anchor-
age, at Anclote stream, July 29 - 1 c? (4.5 mm); 1 9 ov (4.0
mm) (flg'd specimens), CMN collns.

British Columbia:

South Vancouver I; Horsefall, Departure Bay. Vancouver
I., Lot No 51, JFL Hart coll., July 6/33 *1 9- ov (4.0 mm)
Wash.-Oregon: ELB Stns., July-Aug., 1966:W23, Lake
Quisiault, at Falls Creek, July 22 - ~30 (S(S 9 specimens
(4-4.5 mm); W27, Lake Crescent, July 24 - 2 99 (4.7 mm),
4 juveniles, CMN colls.

Other U.S. material: Lake Leota, King Co., in Utricularia
bladders, R. Wallace coll. - 1 9 (2.5 mm). Warm Springs,
near Frenchglen, Hemey Co., (42^48'N., 118^54'N), in
spring 1280 m a.s.I., G. W. Courtney coll., July 7/87 - 6 d'd'.

6 99 (3.5 - 3.8 mm); CMN collns.

California: Malibu Lagoon State Park, D. Galli coll., Jan.
25/84 - 1 (S (5.0 mm); CMN collns. IZ 1984-39.

Distributional Ecology (Pacific drainages): Fresh
and slightly brackish waters of lakes rivers and upper
tidal portions of estuaries, Mexico and California north
approximately to the tree line of Alaska.

Remarks; The small mature male and female speci-
mens from springs at Frenchglen are quadrimucronate,
having postero-dorsal mucronations on peraeon seg-
ment 7 and all three pleon segments.

Parhyalella Kunkel
(see Figs. 6d, 15)

Parhyalella Kunkel, 1910: 74. Bulycheva 1957: 181.
Barnard & Karaman 1991: 372. Bousfield 1996: 182.
Lazo-Wasem & Gable 2001: 5.
im Parhyalella Krapp-Schickel 1993 (= Marino-
hyalella).

noaExhyalella Stebbing, 1917 (Type-F. natalensis)

Type species: Parhyalella batesoni Kunkel, 1910,
monotypy.

Species: Parhyalella barnardi Lazo-Wasem & Gable,
2001 (Baja California); P. congoensis Ruffo, 1953 (SE
Atlantic); P. kunkeli Lazo-Wasem & Gable, 2001 (Tai-
wan); P. nisbatae Lazo-Wasem & Gable, 2001 (Carib-
bean); P. pietschmarmi Schellenberg, 1938 (Hawaii);
P. rujfoU Lazo-Wasem & Gable, 2001 ; P. steelei Lazo-
Wasem & Gable, 2001; P. whelpleyi (Shoemaker,
1933) (Caribbean to Brazil); Parhyalella spp. Lazo-
Wasem & Gable, 2001 (Bahama Ids., Cuba, Rorida
Keys to US coast of Gulf of Mexico; also S. Korea).

Diagnosis: Antennae variable, mostly short. Anten-
na 2 (male), peduncle very stout; basal flagellar seg-
ments conjoint; gland cone small.

Mandibular left lacinia 5 l/-2-dentate; molar seta
lacking. Maxilla 1 lacking palp. Maxilliped palp,
dactyl unguiform, without whip.

Coxae 1-4 lacking posterior marginal process.
Gnathopods strongly sexually dimorphic. Gnathopod 1
(male), dactyl often bifid medially; gnathopod 2 (male)
propod large, palm setose or spinose, carpal lobe dis-
tinct. Peraeopods slender, homopodous, weakly spin-
ose; dactyls medium, without medial spine. Coxal gills
medium, sac-like, subsimilar. Sternal gills lacking.
Uropod 1 , peduncular spines ordinary; rami margin-

AMPHIPACIFICA VOL. 3 NO. 2. Nov. 15, 2001 29

Fig, 14. Hyalella azteca Saussure, 1858. Male (4.5 mm); female ov (4,0 mm).
ELB Stn. A 166, Leo Anchorage, at Anclote stream, SE Alaska.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 30

ally spinose. Uropod 3, peduncle relatively slender,

ramus short, apex with slender spines.

Telson plate-like, entire.

Female: Peraeon segment 2, preamplexing notch
ordinary, lacking locking slit and unguisial groove.
Brood plates slender, apex acute; marginal setae me-
dium short.

Distributional Ecology: Low intertidal and shallow
waters of protected coasts and estuaries, often associ-
ated with Enteromorpha in brackish pools and seeps.
Along tropical and warm-temperate shores of Pacific,
Atlantic and Indian Oceans, south to Central Chile, and
north to Ensenada, B. C., Mexico.

Remarks: The type locality of Parhyalella barnardi
is Ensenada, Baja California, Mexico, very close to the
southern limit of the present study region.

Parhyalella steelei Lazo-Wasem & Gable
(Fig. 6d)

Parhyalella steelei Lazo-Wasem & Gable, 2001: 37,
figs. 28-31.

Parhyalella pietschmanni Steele, 1973: 276, figs. 1-4.
Material Examined:

Ambatoloaka, Nosy B6, Mcidagascar, in drifting dead turtle
grass, D. Steele coll., July - December, 1963. - numerous cfcf
99, and imm. specimens, CMN collns.

Remarks: Steele (1973) detailed the life history of a
species of Parhyalella, recently described as P. steelei
Lazo-Wasem & Gable, that occurs commonly at Nosy
Be, Madagascar. He noted that the pre-amplexing
male utilized gnathopod 1 in grasping the female about
the level of peraeon segment 2.

The preamplexing notch of a female specimen from
the original material is illustrated in Fig. 6d. Its form
in other species of the genus, and in Exhyalella and
Marinohyalella remains to be fully described.

Parhyalella sp.

(Fig. 15)

Parhyalella whelpleyi Bousfield 19%: 181, fig. 2.
Parhyalella sp. Lazo-Wasem & Gable 2001: 43?

Material Examined: Isia Arenas, Cuba, sandy shore, LW.

- Cf (7.5 mm); 9 (5.5 mm) (slide mounts), CMN collns.

Remarks: Bousfield (19%) figured material from Isla

Arenas, Cuba, as Parhyalella whelpleyi Shoemaker,

1933. This figure, partly reproduced here (fig. 15),

differs in several respects from the more detailed fig-
ures and description of P. whelpleyi provided by Lazo-
Wasem and Gable (2001). The present species may
be similar to, or perhaps identical with, undescribed
species recorded by those authors from the Rorida
Keys, in CjCRL collections from along the Gulf coast
of Alabama and Mississippi (LeCroy, pers. comm.),
and in unidentified CMN collections of Parhyalella
from the Bahamas, Rorida, and the Gulf coast.

DISCUSSION
Phyletic Relationships

Family Hyalellidae has long been considered an
integral part erf, or closely related to, family Hyalidae
(Stebbing 1906; Bulycheva 1957; Barnard & Karaman
1991; Bousfield 19%). In a recent analysis of genus
Parhyalella, Laso-Wasem & Gable (2001) revived
genus Exhyalella Stebbing, 191 7, created the mono-
typic genus Marinohyalella for P, richardi Chevreux,
1902 of the Mediterraean region, but retained all three
genera with family Hyalidae ( sens. lat.V
Hg. 16 portrays character state similarities of world
genera and subgeneric groupings utilized here within
family Hyalellidae. The analytical treatment employs
a semi-phyletic modification of the UPGMA system of
Sneath and Sokal (1973), as in previous analyses of
other North Pacific amphipod taxa. Character states
are ordered plesio-apomorphically and the relative
phyletic placement of a given taxon is represented by a
numerical sum of plesiomorphic, intermediate, and
apomorphic character state values (0, 1, and 2, respec-
tively) in a Resio- Apomorphic (P.-A.) Index. The
matrix of 12 characters and corresponding character
states on which the resulting phenogram is based are
mainly those of Lazo-Wasem & CJable (2001 ), and may
be supplied on request.

The phenogram reveals two major generic sub-
groupings at and above the 50% similarity level; a
relatively primitive assemblage of genera and sub-
groups on the left (P.-A. index 5-10), and an advanced
generic group on the right (P. A. Index 1 1-13). The
former embraces the North and South Pacific genus
Allorchestes, the very closely related Mediterranean
genus Marinohyalella (-80% similarity), and the large
and diverse, but less closely similar, neotropical fresh-
water genus Hyalella. The advanced tropical and

AMPMPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 31

Fig. 15, Parhyalella sp. Male (7,5 mm); female (5.5 mm). Sandy shore, LW. Isla Arenas, Cuba.

warm-temperate marine genera Exhyalella, and the
Parhyalella complex, cluster at only the 65% level of
similarity, in our viewing justifying the recent decision
ofLascvWasem& Gable ( 2001 ) to separate the two
genera. The relatively close morphological similarity
of Marinohyalella to both Allorchestes and Hyalella,
and its former inclusion within genus Parhyalella,
tends to support the placement of all five genera withiri
family Hyalellidae as here defined.

Using a broader set of species level characters. Fig.
17 portrays character state similarities within known
species and subspecies of genus Allorchestes in rela-
tion to outgroup genera Parhyalella and Hyalella.
Characters and character states on which the resulting
phenogram is based are given in Table I. Cluster
analysis data on which the phenogram is based are
considered overly bulky and repetitive for publication
here, but may be supplied on request.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 32

Fig. 16. Phenogram of morphological similarities of genera
and species groups within family Hyalellidae

Two major subgroups of Allorchestes are discern-
ible above the 50% similarity level; a relative primitive
(P.-A. index 10-18) assemblage of species from the
southern hemisphere, and three large species from the
northernmost North Pacific region. These all share a
simple-tipped dactyl of gnathopod 1 (male), normally
spinose peduncle of uropod 1, and squarish, often
deeply cleft tel son. To the right is a cluster of 4 more
advanced species (P.-A. indices of 19-29) that share a
split-tipped unguis of the dactyl of gnathopd 1 (male)
as well as a disrupt! vely spinose peduncle of uropod 1,
and a broadly rectangular, shallowly clefttelson. With-
in the primitive subgroup, A. bella bella and A.priceae
are closely similar in the relatively elongate antennal

flagella, the swollen condition of the dactyl of gnatho-
pod 1 (male), and the ventrally rounded coxae 2-4.

Within the advanced angusta subgroup, A. hirsuta is
a relatively primitive western Pacific member. Close
similarity of the Asiatic A. malleola and the North Am-
erican A. angusta is here confirmed, but differences,
especially in their preamplexing morphologies, are
deemed sufficiently great to justify resurrections of
these twoforms as distinct species. Allorchestes rickeri
is a North American-endemic member of the angusta
complex, occuring mainly on shallow sandy
substrata,that is especially distinctive in the form of its
preamplexing morphology, in both males and females.

AMHflPACIHCA VOL. 3 NO. 2. Nov. 15, 2001

33

Fig. 17. Phenogram of morphological srailarities and possible phyletic relationshios
among species of Allorchestes-, in relation to Parhyalella and Hyalella outgroup genera.

Biogeography of North Pacific Hyalellidae
The distribution in the North Pacific Rim region of
7 species cS Allorchestes, northern records of the trop-
ical-warm temperate genus Parhyalella, and North
Amen^ records of the neotropical freshwater genus
Hyalella are summarized semi-diagramatically in Fig
18 With respect to the cold-temperate genus
Allorchestes, A. hirsuta and A. malleola occur in the
Western Pacific region only (Zone 1). Three species,
Allorchestes priceae, A. angusta. and A. rickeri occur
m the Eastern Pacific only (zones 3-9): Allorchestes
carimta, and A. hellabellaoccuTin both theeastemand

westemNorthPacificandoverlapthroughoutthe south-
ern Ben ng Sea and Aleutian Islands (zones 2 & 3). As
noted above, the more primitive species occur mainly
in the western Pacific region and the more advanced
species along North American Pacific shores.

The genus Parhyalella is represented on western

Pacific shores by P. iunfe/iand an undescribed species
from South Korea (Lazo-Wasem & Gable 2001). P.

pterscAmonni has beenfullydescribedfrom the Hawai-
ian islands by J. L. Barnard (1970). The reproductive
biology of a closely related Malagascan species, P.
steelei. has been described by Steele (1973). As noted
above, P. barmrdi was initially described from
Ensenada, Mexico, leading to speculation that the
species may occur also in southern California.

The freshwater genus //ya/e//a is represented by H.
azteca in streams and lakes throughout the North
American coastal region from the tree line of Alaska
southward to Mexico (Bousfield 1958; 1996). H.
azteca is typically dorsally bi-raucronate but a number
of genetic species currently being studied by Witt &
Hebert (2000), possibly embracing the 4-mucronate
van ety recorded here from Oregon, mayalsobe present.

AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 34

TABLE 1. CHARACTERS AND CHARACTER STATES: GNATHOPODS

CHARACTER

Plesiomorphic

0

STATE

Intermediate

1

Apomorphic

2

Head region

1. Antenna 1, flagellar length

long (>15 segmts)

13

short (9-11 segs.)

2. Antenna 2, flagellar length

Long

short

3. Maxilla 1, palp

Gnathopods (male)

large

small

lacking

4. GN 1 (male), median spine

horizontal

oblique

vertical

5. GNl, prop palm. p. d. angle

smooth

slightly, convex

stout process

6. GNl, prox. palm margin

straight

convex

7. dactyl unguis

normal

split-tipped

8. dactyl thickness

medially narrowing

medially swollen

9. GN2 propodal palm, ratio P/d

large (>2.6)

medium (2.5)

small (<2.4)

10. carpal lobe, length

short (<

long (>

Gnathopods (female)

1 1 . GN 1 , propod depth (L/W ratio) deep (< 1 .8)

medium (1.95)

slender (>2. 1 )

12. GN2, Propod depth (L/W ratio) deep (<1.8)

medium (1.9)

slender (>2.0)

13. GN2, palmar slope

oblique

vertical

parachelate

Peraeopods

14. P4 coxa, lower margin

square

rounded

15. P5, coxa 5 type

posterolobate

aequilobate

si. anterolobate

16. P 5, segment 4

normal width

broad (W =L)

17. P2, brood plate (female)

large num. setae

small, few setae

Uropods

18. Uropod 1, peduncular
outer marginal spines

uniform along

slight gap

distal gap

19. Uropod 3, Ramus length
(ram/ped ratio)

long (>0.8)

medium (0.7)

short (<0.6)

Telson

20. Depth of apical notch

deep (>l/3)

medium 1/3

shallow (< 1/3)

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AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 35

DISTRIBUTION
ZONES;
NORTH
PACIFIC
RIM

Hyalellidae:

AUorchestes sp

1

Western

Pacific

2

Bering
Sea and
Aleut-
ians

3

Prince

William

Sound

4

Cross Sd
to Dixon
Entrance
&Q.C.IJ

5

Nor.BX
to North
Vane. I.

/

6

Central
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S. Vanc.1

/

7

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ton state

8

Oregon
& North
Californ

1

9

Southern

Californ

A. malleola

X

•

A, hirsuta

X

A carinata

X

X

X

A, bellabella

X

X

X

X

X

•

X

A. priceae

•

X

X

X

X

•

A. angusta

X

X

X

X

X

X

X

7

•

A. rickeri

X

X

X

X

X

X

7

#

HyateUa spp.

X

X

X

X

X

X

X

Parhyalella spp

X

X

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AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 37

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Sneath, P. H. A., & R. R. Sokal 1973. Numerical
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AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 38

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AMPHIPACIHCA VOL. 3 NO. 2. NOV, 15, 2001 39

Systematics of the subterr^ean amphipod genus Stygobromm (Crangonyctidae) in Western
North America, with emphasis on species of the hubbsi group.

By Daqing Wang and John R. Holsinger

Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23507, USA

ABSTRACT

Twenty-eight new species of the Holarctic, freshwater amphipod genus Stygobromm are described from a variety
of subterranean groundwater habitats in western North America, including caves, springs, wells and the hyporheic
zone, and new data are given on species previously described from this region. A total of 53 species are covered
in this study, 44 of which are members of the hubbsi group. Twenty-three of the newly described species are assigned
to^s large species group, which is distinguished by the absence of sternal gills (processes). A phylogenetic analysis
affirms the monophyly of the hubbsi group and suggests that a relatively strong clade of 10 species exists in the far
western states of California, Oregon and Washington. A taxon-area cladogiam reveals a moderately strong
relationship between terminal taxa and physiographic provinces, which are designated as areas in the analysis. In
contrast to eastern North America, where many distinct species groups of Stygobromm are recognized, taxonomic
iversity is signifi^ntly reduced among the western species. A majority of species Stygobromm in western North
Amenca, especially those m the hubbsi group, are closely similar morphologically, leading to the conclusion that
^y of them are denved from a common ancestor foimerly widespread across much of the region

INTRODUCTION

Species of the large, widely distributed freshwater
amphipod genus Stygobromus are recorded from a
variety of subterranean groundwater habitats, includ-
ing especially caves, wells, springs and the hyporheic
zone beneath surface streams. All members of the
genus are stygobites (= stygobionts), inasmuch as they
are restricted to subterranean waters (or their resur-
gence) and are characterized morphologically by loss
of eyes and pigment and sometimes by attenuation of
the t^y. Stygobromus belongs to the Holarctic
amphipod family Crangonyctidae, which contains six
extant genera and 152 described species (plus many
undescribed). Although members of the family occur
in both North America and Eurasia, crangonyctids are
far more common and significantly more diverse in
North America than in Eurasia. Approximately 80
percent of the species in the family are stygobites, and
three of the six genera are exclusively subterranean
(Holsinger 1994a).

Prior to publication of the present paper,
Stygobromus included 101 described species, 98 of
which were recorded from subterranean habitats in
North America. Outside the continent, two species of
Stygobromus were reported from central Asia and one
from extreme eastern Europe near the Caspian Sea
(Holsinger 1987; Kulkina 1992). In this paper, we
describe 28 new species from the Cordilleran region of
western North America and provide brief synopses for
24 species previously described from this region. We
al so inci ude S. putealis (Hoi mes) from eastern Wi scon-

sin, which, while occurring far outside the western
Cordillera, is morphologically very similar to many
species in the west. A majority of species (44 of 53)
treated in this paper, including 5. putealis, are assigned
to the hubbsi group, which is a large complex of closely
similar species erected earlier by Holsinger (1974) to
encompass all species in the genus that, with one
exception, lack sternal gills (or processes) on the
pereonites. A revised diagnosis of the group is given
below. The description of 28 new species of
StygobromusfTom western North America, 23 of which
are assigned to the hubbsi group, brings the total
number ofdescribed species in the genus to 129. Soon
to be added to this total are descriptions of 22 more new
species from central and east-central North America
that are pending publication in another paper (Hol-
singer, ms.).

The first comprehensive taxonomic study of west-
ern North American species of Stygobromus was by
Holsinger (1974). In that paper, 17 new species were
described from Arizona, Califoinia, Orgeon, Montana,
Nevada, and Washington and the previously described
S. hubbsi Shoemaker, 1942 from Malheur Cave in
Oregon was partly redescribed. Between 1974 and the
present time, six more species were described from
western North America. These descriptions included
three species from hyporheic habitats and a spring in
Colorado by W ard (1977) ; one species from a cave and
one from a spring in the Canadian Rockies of Alberta,
respectively, by Holsinger (1980) and Bousfield and

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 40

Holsinger (1981); and one species from caves on Van-
cxiuver Island, British Columbia by Holsinger and
Shaw (1986, 1987). In addition, the monotypic genus
Stygonyx, a morphologically closely similar sister ge-
nus of Stygobromus, was described from a phreatic
water habitat in northern Oregon by Bousfield and
Hoi singer ( 1989) . Twoundescribed {non-hubbsi group)
species have also been discovered in Alaska in recent
years (see Holsinger et al. 1997) but will be described
in another paper.

The principal objectives of the present study are: (a)
to describe 28 new species of Stygobromus that have
been discovered in groundwater habitats of western
North America in recent years, primarily during the last
two or three decades; (b) update taxonomic and distri-
butional data on previously known species from this
region; (c) further clarify the taxonomic and geo
graphic limits of the hubbsi group species vis-a-vis
other (non-member) species from western North Am-
erica; (d) conduct a cladistic analysis with the dual pur-
pose of sorting out phylogenetic relationships among

westcmspeciesofStygobromuSyespecldllyinihQhubbsi

group, and examining the relationship between
phylogeny and geographic distribution; and e) formu-
late a plausible hypothesis that explains the geographic
distribution of numerous, morphologically closely
similar stygobitic species over a large part of the highly
varied and rugged terrain of the western Cordillera of
North America.

We have included distribution maps for all western
species of Stygobromus, and a table showing the fre-
quency of occurrence of species in different kinds of
subterranean groundwater habitats. A phylogenetic
anal ysi s was conducted, and the alignment of species in
this paper follows the sequence established on the
cladogram in which characters are partially weighted
and ordered. This cladogram is also converted to a
taxon-area cladogram in an attempt to examine the
relationship between clades and their geographic dis-
tribution. Species assigned to the hubbsi group are
listed first in the Systematics section, followed at the
end of the section by nine non-hubbsi group species.

ACKNOWLEDGMENTS

We are grateful to the many biologists who assisted
with the fieldwork and/or donated specimens for study .
We also thank those collectors who furnished us with
pertinent ecological data from their field notes. Al-
though we have credited all those who assisted with
collections of material in the "materials examined"

sections under the various species, we are especially
grateful to the following biologists who provided us
with substantial numbers of specimens and detailed
information on the habitats they sampled; Cheryl B.
Barr, Andrew Boulton, Steven P. Canton, Jim Chester,
Greg W. Courtney, Scott J. Harden, David B. Herbst,

IbrahimMohammad,MarilynMyers,StewardB.Peck,

Margaret E. Ver Hey, D. Craig Rudolph, and James V .
Ward.

In addition to the collectors, we thank the following
museums for lending us additional pertinent materials:
Bernice P. Bishop Museum (MBB), Canadian Mu-
seum of Nature (CMN), Essig Museum of Entomology
(EME) of the University of California at Berkeley,
Montana Entomology Collections (MTEC), Museum
of Southwestern Biology (MSB), and Thomas Burke
Memorial Washington State Museum (TBMWSM).
Drs. Kent E. Carpenter and James D. Thomas read the
manuscript and made a number of helpful comments on
the phylogenetic analysis and interpretation of the
results. Dr. Thomas also commented on the taxonomy
and made several useful suggestions. We are espe-
cially grateful to Dr. E. L. Bousfield for encouraging us
to publi sh our manuscript in Amphipacifica, and for his
valuable editorial assistance and many useful sugges-
tions regarding arrangement of the figures. His com-
ments and suggestions, along with those of Drs. Car-
penter and Thomas resulted in significant improve-
ments of the paper. We also owe a special debt of
gratitude to Donald Emminger of the Graphics Office
at Old Dominion University for his professional assist-
ance with preparation of the distribution maps.

Finally, we thank the National Science Foundation
for its generous support of our systematics research
through a PEET (Partnerships for Enhancing Expertise
in Taxonomy) grant to JRH (DEB-9521752). This
kind of support is essential for the survival of taxonomy
in the 21st Century.

METHODS AND MATERIALS

Conventional methods for collecting subterranean
amphipods have been discussed in previous papers
(Holsinger 1967, 1972). Additional comments on
collecting methodology as it applies to certain condi-
tions in western North America are found in Ward
(1977) and Ward and Holsinger (1981). In the labora-
tory all measurements were made in millimeters to the
nearest tenth with the aid of a calibrated micrometer
disc. Total length refers to the length of the body
excluding appendages, i.e., length from base of anten-

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 41

na 1 to base of telson. The figures were prepared with
the aid of a Leitz drawing tube from appendages
mounted in Faure's mounting medium (a modification
of the conventional Hoyer's medium) on glass slides.
Drawings were sometimes made from appendages
mounted in undiluted glycerin, in which case these
structures were later moved to the more permanent
Faure’s medium for further clearing and storage. Ap-
pendages were not drawn to any particular scale unless
noted to the contrary. A legend for figure symbols is
provided on page 147.

With the exception of one species, holotype speci-
mens of species described in this paper are deposited in
the National Museum of Natural History (Smithsonian
Institution) under the catalog numbers of the former
United States National Museum (USNM). The excep-
tional holotype is deposited in the Canadian Museum
of Nature collections (CMNC). Collections deposited
in other museums are indicated by the museum abbre-
viation given above under ’Acknowledgments". All
other specimens examined in this study pertaining to
the new species are designated as paratypes and are
deposited in the research collection of the second
author (JRH) at Old Dominion University. This mate-
rial will eventually be transferred to the Smithsonian
Institution.

In species descriptions, nomenclature for setal pat-
terns on segment 3 of the mandibular palp follows the
now widely used standard introduced by Stock (1974)
(see Koenemann & Hoi singer 2001 , fig. 2g, for a recent
application). According to this convention, the five
different sets of setae that may occur on this segment
and are designated by upper case letters A, B, C, D, and
E. For the species treated in this paper, as well as most
other species of Stygobromus, D (row of short setae on
inner margin) and E (cluster of longish setae on apex)
are the most common and are always present, B and C
are less common and often absent, whereas A is almost
always absent.

Because vandalism and pollution have become ma-
jor threats to many cave habitats, we have elected to
protect these sensitive environments from further de-
struction by providing only the name, and a very
general location, of caves listed in the "material exam-
ined" sections for each species. This protocol applies
only to caves, inasmuch as springs, hyporheic habitats,
wells, etc. are generally less vulnerable to vandalism or
frequent visitation. Thus, if there is legitimate need for

a precise cave location, such information can be ob-
tained either from a published state speleological sur-
vey (available for some states) or the JRH database
upon request. State cave surveys have been published
for California (Halliday 1962), Colorado (Parris 1973),
andWashington(Halliday 1963). Additional informa-
tion is available on caves in Oregon (Greeley 1971),
and supplementary information on Washington caves
was published in the "Guidebook of the 1972 Conven-
tion of the National Speleological Society."

SYSTEMATICS

Stygobromus Cope

Stygobromus Cope, 1872:422; Aprocrangonyx
Stebbing, 1899:422; Stygonectes W, P. Hay, 19(B:
430; Synpleonia Creaser, 1934:1.

Diagnosis. A recent, detailed diagnosis of Stygobromus
is that of Holsinger (1978). However, the principal
diagnostic character of the genus is the uniramous third
uropod (U3), in which the ramus is 1-segmented,
shorter than the peduncle, sometimes vestigial or ab-
sent, and when present armed with 1 to several short
spines. The taxonomic and geographic relationship of
Stygobromus to other genera in the family Crangonyct-
tidae is discussed by Holsinger (1986a, 1986b).

The hubbsi group

Diagnosis. Distinguished from all other members of
the genus Stygobromus (with the exception of one aber-
rant species in North Carolina) by the absence of sternal
gills (processes) on the pereonites and the following
combination of characters: mature females larger than
mature males; propod of gnathopod 2 usually larger
than, but sometimes subequal in size to, propod of
gnathopod 1 ; posterior margin of propod of gnathopod
1 typically shorter than palm, usually without setae but
sometimes withfew short, submarginal setaejust proxi-
mal to the defining angle; shape of bases of pereopods
5, 6 and 7 variable but often relatively narrow and not
much expanded posteriorly, and often lacking distinct
distoposterior lobes; telson usually as long as broad or
little longer than broad, apical margin typically with
shallow notch.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 42

Stygobromus saltuarisy new species

(Fig. 1)

Material examined. OREGON. Lane Co.: hyporheic/seep
area, Trail Creek, Willamette National Forest, HOLOTYPE
9(on slide mounts in part)(CMNC 2001-0020), 1 9paratype
(CMNC 2001-0021), G. W. Courtney, 25 June 1987.

Diagnosis. A medium-sized hyporheic species, related
to S, oregonensis in structure of pleonal plates but
distinguished by the following: gnathopods 1 and 2
with proportionately longer posterior margin; palm of
gnathopod 2 concave; uropod 1 and 2 with fewer long
and stout spines on rami and peduncle; uropod 3
peduncle broad; and telson lacking notch and with
fewer spines. Largest 9» 5.0 mm; cf unknown.
Female. Antenna 1: 66 percent length of body, 80
percent longer than antenna 2; primary flagellum with
14 segments. Antenna 2: flagellum with 6 segments.

Mandibles subequal: spine row with 4 plumose
spines; palp segment 2 with row of 2 rather long setae
on inner margin; palp segment 3 bearing 1 B seta, 4-5
C setae, few D setae, and 3 E setae, lacking A setae.
Inner lobes of lower lip absent. Maxilla I: inner plate
with 7 apical, plumose setae; palp with 4 stiff setae or
slender spines apically. Maxilla 2: inner plate with
oblique row of 8 plumose setae on inner margin.
Maxilliped: inner plate with 1 bladelike spine, 3 plum-
ose spines, and 3 naked setae apically, and 1 stiff setae
on inner margin; outer plate with short setae on inner
margin and 1 small bladelike spine on or near apex.

Gnathopod 1 : propod shorter than that of gnathopod
2; palm straight or slightly concave about 2 times
longer than posterior margin, armed with 13-14 spine
teeth in double row ; defining angle with 3 spine teeth on
outside, 3 shorter ones on inside; posterior margin
without setae; 4 superior medial setae; 2 or 3 inferior
medial setae; dactyl nail rather long; coxa about as
broad as deep, margin with 2 setae. Gnathopod 2:
propod subrectangular, 2 times longer than broad; palm
slightly concave medially and armed with 13 spine
teeth in double row ; defining angle with 2 spine teeth of
unequal length on outside, 2 shorter spine teeth on in-
side ; posterior margin approximately 70 percent length
of palm, with 3-4 sets of doubly inserted setae; 6
superior medial setae, singly inserted; 3 singly inserted
inferior medial setae; coxa little broader than deep,
margin with 4 setae.

Pereopods 3 A: coxal plates about as deep as broad,
margins with 3 setae. Pereopod 6 little longer than
pereopod 7, about 55 percent length of body, and 22
percent longer than pereopod 5. Pereopods 5-7: bases

of about as broad proximally as distally; posterior
margins convex; distopostcrior lobes well developed;
anterior and posterior margins with variable number of
spines and setae; segments 4, 5 and 6 of pereopods 5-
7 with longish, slender spines; dactyls of pereopods 5-
7 relatively elongate, that of pereopod 6 about 25
percent length of corresponding propod. Coxal gills
present on pereopods 2-6, absent from 7. Brood plates
slightly expanded distally.

Pleonal plates: posterior margin of plates 1 and 2
slightly concave or nearly straight, with 1 setule near
distopostcrior comer, that of 3 convex, each with 1
setule near distopostcrior comer; distopostcrior cor-
ners rounded and indistinct; ventral margin of plate 2
with 2 spines, that of plate 3 with 4 spines. Uronites
free. Uropod 1: inner ramus little longer than outer
ramus, about 80 percent length of peduncle, with 7
spines; outer ramus with 6 spines; peduncle with 6
spines. Uropod 2: inner ramus longer than outer ramus,
subequal in length to peduncle, with 7 spines; outer
ramus with 4 spines; peduncle with 4 spines. Uropod 3 :
peduncle usually not bearing small setae; ramus ap-
proximately 50 percent length of peduncle, with 3
apical spines.

Telson approximately 30 percent longer than broad,
narrowing distally; apical margin with tiny median
notch between spine clusters, bearing 10 relatively
long spines.

Distribution and ecology. This species is only known
from its type-locality, the elevation of which is about
640 m above sea level (G. W. Courtney, pers. comm.).
The larger female (5.0 mm) in the series of two has
setose brood plates.

Etymology. The epithet saltuaris is from the Latin,
meaning “forest,” in reference to the location of the
type-locality in the Williamette National Forest.

Stygobromus rallus, new species
(Figs. 2,3)

Material examined. WASHINGTON. Whitman Co.: Rock
Lake Spring, about 28 km S. of Cheney, HOLOTYPE 9
(USNM 1000069), 3 and 3 99 paratypes, 2 juvs., I.
Mohammad, 2 July 1992; Spokane Co.: Millers Spring No.
2, 2 cTcT, 299, 1- Mohammad, 5 Dec. 1992.

Diagnosis. A relatively large groundwater species,
closely related to S. duplus in gnathopods and uropods
1-2, but distinguished from that species as follows:

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 43

Fig. 1. Stygobromus saltuaris^ n. sp. Female (4.7 mm). Lane County, Oregon.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 44

Fig. 2. Stygobromm nUlus, n. sp. Female (7.6 mm). Whitman County, Washington.

AMFMPAaHCA VOL. 3 NO. 2. NOV. 15, 2001 45

Fig. 3. Stygobromm raUus, n. sp. Female (7.7 mm); male (4.5 mm). Whitman Co., Washington.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 46

propod of gnathopod 1 with short posterior margin and
more teeth on palmar margin; gnathopod 2 with more
teeth on palmar margin; bases of pereopods 4-7 more
narrow; uropod 3 with more spines on rami; telson
lacking notch on apical margin and with longer and
stouter spines. Largest 9» 1 1.0mm;largestcJ’, lO.Omm.

Female. Antenna 1: 70 percent length of body, 62
percent longer than antenna 2; primary flagellum with
18 segments. Antenna 2: flagellum with 7 segments.

Mandibles subequal: spine row with 6 plumose
Spines; palp segment 2 with row of 13 rather long setae
on inner margin; palp segment 3 bearing 3 B setae, 16
D setae, and 4 E setae, lacking both A and C setae. Inner
lobes of lower lip vestigial. Maxilla I : inner plate with
8 apical, plumose setae; palp with 6 stiff setae or
slender spines apically. Maxilla 2: inner plate with
oblique row of 9 plumose setae on inner margin.
Maxilliped: inner plate with 1 bladelike spine, 11
plumose setae, 5 naked setae apically, and 4 coarse
setae on inner margin; outer plate with setae on inner
margin and apex, some slightly plumose, and 1 small
spine.

Gnathopod 1: propod subequal in size to that of
gnathopod 2; palm sli ghtly convex and about five times
longer than posterior margin in length, armed with 27
spine teeth in double row; defining angle with 4 spine
teeth on outside, 4 shorter ones on inside; posterior
margin with row of 4 setae; approximately 8 superior
medial setae, mostly triply inserted; 5-6 singly inserted
inferior medial setae; dactyl nail rather short; coxa
rather narrow and broader than deep, margin with 2
setae. Gnathopod 2; propod palm straight or weakly
convex and armed with 29-30 spine teeth in double
row ; defining angle with 4 spine teeth of unequal length
on outside, 2 shorter spine teeth on inside; posterior
margin about 50 percent length of palm, with 3 sets of
mostly triply inserted setae; 10 superior medial setae,
mostly triply inserted; 5 singly inserted inferior medial
setae; coxa little broader than deep, margin with 3
setae.

Pereopod 3: coxal plate broader than deep, margin
with 4 setae. Pereopod 4: coxal plate broader than deep,
reaching about 30 percent length of basis, margin with
4 setae. Pereopod 6 little longer than pereopod 7, about
78 percent length of body, 43 percent longer than
pereopod 5. Pereopods 5-7: bases narrowing in larger
specimens, broader proximally than distally; posterior
margins convex; distoposterior lobes distinct (but less
so on pereopod 7); anterior and posterior margins with
variable number of spines and setae; segments 4, 5 and

6 of pereopods 5-7 with longish, slender spines ; dactyls
of pereopods 5-7 relatively slender and elongate, that of
pereopod 6 about 25 percent length of corresponding
propod. Coxal gills present on pereopods 2-6, absent
from segment 7. Brood plates somewhat narrowing
distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 setule near distoposterior comer, that of 2 and 3
less convex, each with 1 setule near distoposterior
comer; distoposterior comers tiny but distinct; ventral
margin of plate 2 without spines, that of plates 3 with 3
to 4 spines. Uronites free. Uropod 1: inner ramus little
longer than outer ramus, about 70 percent length of
peduncle, with 9 spines; outer ramus with 9 spines;
peduncle with 14 spines. Uropod 2: inner ramus longer
than outer ramus, subequal in length to peduncle, with
8 spines; outer ramus with 8 spines; peduncle with 6
spines. Uropod 3: ramus approximately 45 percent
length of peduncle, with 4 apical spines.

Telson little longer than broad, apical margin typi-
cally with tiny median notch between spine clusters,
bearing 14 - 18 relatively long spines.

Male. Differing from female as follows: Gnathopod
propods shorter and narrower, palm of propod 2 longer
in relation to posterior margin and with fewer teeth on
palmer margin. Uropod 1: peduncular process approxi-
mately 25 percent length of outer ramus, narrowing
distally, upper margin minutely serrate distally; inner
ramus with 7 spines; peduncle with 8 spines. Uropod 2:
inner ramus with 9 spines, peduncle with 6 spines.
Telson with 14 long apical spines.

Distribution and ecology . In addition toMil lers Spring
no. 2 and Rock Lake Spring (the type-locality),
Mohammad ( 1995) reported this species from Millers
Springs no. 3 and 4, Spokane Co., WA and Dragon
Spring, Adams Co., Washington. This species co-
occurs with S. lotus in Rock Lake Spring and Millers
Spring No. 2

Etymology. The epithet rallus means “thin” or “nar-
row,” in reference to the bases of pereopods 5-7,
which are relatively narrow in comparison to those of
the sympatric 5. lotus.

Remarks: This is the same species that Ibrahim
Mohammad named Stygobromus chrissi in hi s unpub-
lished 1995 master’ s degree thesis at Eastern Washing-
ton University.

AMPHIPAOHCA VOL. 3 NO. 2. NOV. 15, 2001 47

Fig. 4. Stygobromus Umbus, n. sp. Female (10.0 mm). Culberson County, Texas.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

48

Stygobromus limbus, new species
(Fig. 4)

Material examined TEXAS. Culberson Co.: Border Cave,
HOLOTYPE 9 (on slide mounts in part) (USNM 1000067),
W. C. Welboum, 1 Nov. 1976; 12 99. paratypes, 16juvs.,S.
J. Harden and C. T. Lindblom, 4 July 1985; 8 99, 1 cf, S. J.
Harden and C. T. Lindblom, 15 Aug. 1986.

Diagnosis. A relatively large cavemicolous species
possibly related to S. azisonensis in structure of the
gnathopods but distinguished from that species as
follows: maxillae 1 and 2 with more setae on inner
plate; maxilliped with more long setae on inner plate;
distoposterior lobe of pereopod 6 better developed;
coxal plates 3-4 with more setae on margin; pleonal
plates 2-3 with more spines on ventral margin; uropods
1-2 with more spines on rami and peduncle; and tel son
with shorter apical spines and slight notch. Largest 9,
12.5 mm; cT unknown.

Female. Antenna 1: 50 percent length of body, 45
percent longer than antenna 2; primary flagellum with
19-20 segments. Antenna 2: flagellum with 7 seg-
ments.

Mandibles subequal: spine row with 6-7 plumose
spines ; palp segment 2 with row of 8 long setae on inner
margin; palp segment 3 bearing 1 B seta, 11 D setae,
and 5 setae E, lacking both A and C setae. Inner lobes
of lower lip absent. Maxilla 1 : inner plate with 1 1 api-
cal, plumose setae; palp with 8 stiff setae subapically.
Maxilla 2: inner plate with oblique row of 13 plumose
setae on inner margin. Maxillip^: inner plate with 3-
4 bladelike spines, 5 plumose spines, 1 naked seta
apically, and 5 plumose setae on inner margin; outer
plate with setae on inner margin and apex, and single
small spine near apex.

Gnathopod 1 : propod smaller than that of gnathopod
2; palm straight and approximately 2 times longer than
length of posterior margin, armed with 20 spine teeth in
double row; defining angle with 2 spine teeth on
outside, 4 shorter ones on inside; posterior margin
lacking setae; 2-3 superior medial setae; 2-3 inferior
medial setae; dactyl nail rather long; coxa broader than
deep, margin with 3 setae. Gnathopod 2; piopod
subrectangular, deeper than broad; palm straight or
slightly concave and armed with 20 spine teeth in
double row; defining angle with 1 spine tooth on
outside, 1 shorter spine tooth on inside; posterior mar-
gin 50 percent as long as palm, with 2 sets of doubly

inserted setae; 10 superior medial setae, triple inserted;

5inferior medial setac;coxadeeperthan broad, margin

with 3 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 6 setae. Ptereopod 4: coxal plate relatively broad,
reaching about 35 percent length of basis, margin with
6 setae. Pereopod 6 little longer than pereopod 7, about
55 percent length of body, 30 percent longer than
pereopod 5. Pereopods 5-7: bases little broader proxi-
maJly than the distall y; posterior margins weakly con-
vex; distoposterior lobes poorly developed; anterior
margin with 3-5 spines, posterior margins with numer-
ous spines; segments 4, 5 and 6 of pereopods 5-7 with
longish, slender spines; dactyls of pereopods 5-7 rela-
tively slender, that of pereopod 6 aj^>roximately 20
percent length of corresponding propixl. Coxal gills
present on pereopods 2-6, absent from 7. Brood plates
little expanded distally.

Pleonal plates: posterior margin of plate 3 convex,
with 2 setules near distoposterior comer, that of 1 and
2 1 ess conv ex, each al so with 2 setule near di stoposterior
comer; distoposterior comers rounded and indistinct;
ventral superior margin of plate 2 with 2 spines, that of
plate 3 with 3 spines. Uronites free. Uropod 1: inner
ramus little longer than outer ramus, about 75 percent
length of peduncle, with 15 spines; outer ramus with 1 1
spines; peduncle with lOspines. Uropod 2: inner ramus
longer than outer ramus, little longer than peduncle,
with 16 spines; outer ramus with 6 spines; peduncle
with 5 spines. Uropod 3: peduncle usually without
small setae; ramus approximately 50 percent length of
peduncle, with 2 apical spines.

Telson little longer than broad, tapered distally;
apical margin with small notch between spine clusters,
bearing 8-9 relatively long spines.

Distribution and ecology. This species is known only
from its type-locality, a cave developed in gypsum
bedrock, where it has been collected on several occa-
sions from phreatic lakes on the cave’s lower level.
Samples were collected from rocks in deep lakes with
the stygobiont isopod Lirceolus nidulus Lewis (see
Lewis 2001). All of the females collected by Harden
and Lindblom in the summer of 1985/86 were ovig-
erous.

Etymology. This species is named for the type-local-
ity, Border Cave. The epithet limbus is from the Latin,
meaning “border”.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 49

Stygobromusfontinalis, new species
(Figs. 5, 6)

Stygobromus sp. B; - Ward and Holsinger, 1981 : 64-
67.

Material examined. COLORADO. Rio Blanco Co.: spring,
ca. 32 km northwest of Rio Blanco, HOLOTYPE 9 (USNM
1000056), 2 cfcf and 1 9paratypes, J. V. Ward, 19May 1977;
5 CfcJ*, 5 99, V, Ward, 20 May 1977; spring at mouth of
Stewart Gulch, ca. 32 km north-northwest of Rio Blanco, 1
9, L V. Ward, 2 Nov. 1979.

Diagnosis. A medium-sized groundwater species,
closely related to S. holsingeri in gnathopods and
uropods 1-2, but distinguished from that species as
follows: maxilliped with 3 bladelike apical spines on
inner plate; gnathopods with more teeth on palmar
margin of propods; palmar margin of gnathopod 1
straight or slightly convex; maxillae 1 and 2 with more
setae on inner plate; telson proportionately longer.
Largest 9, 7.5 mm; largest cf, 6.5 mm.

Female. Antenna 1: 60 percent length of body, 57
percent longer than antenna 2; primary flagellum with
13 segments. Antenna 2; flagellum with 6 segments.

Mandibles subequal: spine row with 6 plumose
spines; palp segment 2 with row of 8 rather long setae
on inner margin; palp segment 3 bearing 1 B seta, 13 D
setae, 3 E setae, lacking both A and C setae. Inner lobes
of lower lip vestigial. Maxilla 1: inner plate with 9
apical, plumose setae; palp with 8 stiff setae or slender
spines apically. Maxilla 2; innerplate with oblique row
of 10 plumose setae on inner margin. Maxilliped: inner
plate with3 bladelike spines, 5 plumose spines, 2 naked
setae apically, and 4 plumose setae on inner margin;
outer plate with setae on inner margin and face, and on
or near apex.

Gnathopod 1 : propod shorter than that of gnathopod
2; palm slightly convex and about 30 percent longer
than ^sterior margin in length, armed with 27-28 spine
teeth in double row ; defining angle with 3 spine teeth on
outside, 2 shorter ones on inside; posterior margin
lacking setae; 2 superior medial setae; about 6 singly
inserted inferior medial setae; dactyl nail rather short;
coxa much broader than deep, margin with 3 setae.
Gnadiopod 2: propod longer than broad; palm straight
or slightly concave and armed with 21 spine teeth in
double row; defining angle with 1 long spine tooth on
outside, 3 shorter spine teeth on inside; posterior mar-
gin about 50 percent length of palm, with 1 cluster of
setae; 9 superior medial setae, most doubly inserted;

about 4 singly inserted inferior medial setae; coxa
broader than deep, margin with 2 setae.

Pereopod 3 : coxal plate deep, about as broad as deep,
margin with 2 setae. Pereopod 4: coxal plate relatively
broad and deep, reaching about 30 percent length of
basis, margin with 6 setae. Pereopod 6 little longer than
pereopod 7, about 67 percent length of body, 50 percent
longer than pereopod 5. Pfcreopods 5-7: bases of about
as broad proximally as distally; posterior margins con-
vex; distoposterior lobes developed but not prominent;
anterior and posterior margins with variable number of
spines and setae; segments 4, 5 and 6 of pereopods 5-
7 with longish, slender spines; dactyls of pereopods 5-
7 relatively slender and elongate, that of pereopod 6
about25percentlengthofcorrespondingpropod. Coxal

gills present on pereopods 2-6, absent from 7. Brood
plates slightly narrowing distally.

Pleonal plates: posterior margin of plate 1 with 1
setule near distoposterior comer, those of 2 and 3
weakly convex, each with 1 setule near distoposterior
comer; distoposterior comers rounded and indistinct;
ventral margin of plate 2 with 2 spines, that of plate 3
with 4 spines. Uronites free. Uropod 1: inner ramus
little longer than outer ramus, about 67 percent length
of peduncle, with 8 spines; outer ramus with 8 spines;
peduncle with 1 1 spines. Uropod 2: inner ramus longer
than outer ramus, subequal in length to peduncle,
heavily spined, with 19 spines, many toward distal end;
outer ramus with 6 spines; peduncle with 4 spines. Uro-
pod 3: peduncle without setae; ramus approximately 33
percent length of peduncle, with 3 apical spines.

Telson little longer than broad, gently tapered dist-
ally; apex with tiny median notch between spine clus-
ters, bearing 11-12 relatively long spines.

Male. Differing from female as follows: Gnathopod
propods broader, palm of propod 2 with more teeth on
margin. Uropod 1 : peduncular process 20percent length
of outer ramus, sharply pointed distally, upper margin
minutely serrate; inner ramus with 8 spines; peduncle
with 6 spines. Uropod 2: inner ramus with 24 spines,
peduncle with 4 spines. Telson with 12 apical spines.

Distribution and ecology. The species is recorded
from its type-locality and a spring at the mouth of
Stewart Gulch, both in Rio Blanco County, Colorado.
The salinity of the type-locality springs was about
1,000 mg/L at the time of collection, 19 and 20 May,
1977 (Ward and Holsinger 1981).

Etymology. The fontinalis is from the Latin,

AMHflPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 50

Fig. 5. Stygobromusfontiruilis, n. sp. Female (7.5 mm). Rio Blanco County, Colorado.

AMPHIPACinCA VOL. 3 NO. 2. NOV. 15, 2001

51

I'xCA

MX2 y mxpd

fis.<k RioB,„coCo,C„l<,^„,

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 52

Fig. 7. Stygobromus jemezemisy n. sp. Female (6.5 mm). Sandoval County, New Mexico

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 S

Fig. 8. Stygobromusjmezensis, n. sp. Female (6.5 mm); male (4.2 mm).

Sandoval Co., New Mexico

AMPHIPACinCA VOL. 3 NO. 2. NOV. 15, 2001 54

meaning “a spring/’ with refererence to the spring
habitat of the species.

Stygobromus jemezensis, new species
(Figs. 7,8)

Material examined. NEW MEXICO. Sandoval Co.:
unnamedwellatBland,HOLOTYPE9(USNM 1000064), 1
9 and 1 <? paratypes (MSB), B. Wilson, 29 Aug. 1994.

Diagnosis. A medium-sized groundwater species,
closely related to S. arizonensis in structure of
gnathopods but distinguished from that species as
follows: propod of gnathopod 1 with more teeth on
palmar margin; pereopod 6 with distinct distoposterior
lobe; posterior margins of pleonal plates with 1 setule;
uropod 1-2 with more spines on rami (similar to 5.
holsingeri); uropod 3 with proportionately broad pe-
duncle; telson with short spines and slight apical notch.
Largest 9, 7.0 mm; largest d* 4.5 mm.

Female. Antenna 1: 52 percent length of body, 57
percent longer than antenna 2; primary flagellum with
14 segments. Antenna 2: flagellum with 6 segments.

Mandibles subequal: spine row with 7 plumose
spines; palp segment 2 with row of 8 rather long setae
on inner margin; palp segment 3 with 10 D and 5 E
setae, lacking A, B and C setae. Inner lobes of lower lip
absent. Maxilla 1; inner plate with 7 apical, plumose
setae; palp with 10 stiff setae or slender spines apically.
Maxilla 2: inner plate with oblique row of 7 plumose
setae on inner margin. Maxilliped: inner plate with 1
bladelike spine, 3 plumose spines, 2 naked setae on or
near apical margin, 3 plumose setae on inner margin;
outer plate with numerous short setae on inner margin
and 1 bladelike spine and several setae on or near apex.

Gnathopod 1 : propod slightly shorter than propod of
gnathopod 2; palm straight or slightly concave, about
half length of posterior margin, armed with 22 spine
teeth in double row; defining angle with 3 spine teeth on
outside, 3 shorter ones on inside; posterior margin
without setae; superior medial setae 4, doubly inserted;
2-5 inferior medial setae; dactyl nail rather long; coxa
deeper than broad, margin with 3 setae. Gnathopod 2:
propod subrectangular, longer than broad; palm armed
with 19-20 spine teeth in double row; defining angle
with 1 very long spine tooth on outside, 1 shorter spine
tooth on inside; posterior margin about 33 percent
length of palm, with 2 setae; 9 superior medial setae,
doubly or triply inserted; 5 or 6 singly inserted infer-

ior medial setae; coxa deeper than broad, margin with
5 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 5 long setae. Pereopod 4: coxal plate relatively
broad and deep, reaching about 45 percent length erf
basis, margin with 8 setae. Pereopod 6 little longer than
pereopod 7, about 66 percent length of body, 30 percent
longer than pereopod 5. Pereopods 5-7: bases little
broader proximally than distally; posterior margins
convex; distoposterior lobes distinct but not promi-
nent; anterior and posterior margins with variable
number of spines and setae; segments 4, 5 and 6 of
pereopods 5-7 with longish, slender spines; dactyls of
pereopods 5-7 relatively slender and elongate, that of
pereopod 6 typically 33 percent length of correspond-
ing propod. Coxal gills present on pereopods 2-6,
absent from 7. Brood plates narrowing distally.

Pleonal plates: posterior margins of plate 1 and 3
convex, each with 1 setule near distoposterior comer,
that of 2 less convex, with 1 setule near distoposterior
comer; distoposterior comers rounded and distinct;
ventral margin of plate 2 with 2 spines, that of plate 3
with 3 spines. Uronites free. Uropod 1: inner ramus
little longer than outer ramus, about 90 percent length
of peduncle, with 10 spines; outer ramus with 7 spines;
peduncle with 7 spines. Uropod 2: inner ramus longer
than outer ramus, subequal in length to peduncle, with
17 relatively long spines; outer ramus with 7 long
spines; peduncle with 2 spines. Uropod 3: peduncle
without setae or spines; ramus approximately 30 per-
cent length of peduncle, with 2 apical spines.

Telson little longer than broad; apical margin with
tiny median notch between spine clusters, bearing 12
relatively short spines.

Male. Differing from female as follows: gnathopod
propods smaller, palm of propod 2 longer in relation to
posterior margin and with more teeth on palmar mar-
gin. Uropod 1: peduncular process present and reach-
ing about 25 percent length of outer ramus, bluntly
rounded distally, upper margin slightly serrate; inner
ramus with 8 spines; peduncle with 6 spines. Uropod 2:
inner ramus with 16 spines, peduncle with 3-4 spines.
Telson with 15 short apical spines.

Distribution and ecology. The species is known from
its type-locality, where it was collected from a deep
water well in the Jemez Mountains of New Mexico.

Etymology. The species is named for the location of
the type-locality in the Jemez Mountains.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 55

Stygobromus herbsUy new species
(Figs. 9, 10)

Material examined. NEVADA. Hko Co.: Ruby Lake Na-
tional Wildlife Refuge, spring No. 121, HOLOTYPE 9
(USNM 1000059), 9 cfcf and 6 99 paratypes (2 partly on
slide mounts), D. B. Herbst, 10 June 1997; spring No. 229,
2 99 , D. B. Herbst, 10 June 1997; spring No. 129, 13 d'd', 5
99 , 1 juv., D. B. Herbst, 9 June 1997, spring No. 103, 5 99 ,

1 Cf , D. B. Herbst, summer 1997; White Rne Co. : Ruby Lake
National Wildlife Refuge, spring No. 208, 1 cf, 18 juvs, D.
B. Herbst, 14 Sept. 1997; spring No. 191, 4 cfcf, 17 99 , 16
juvs., D. B. Herbst, 13 Sept. 1997, spring No. 180,4cfcf, 15
99 , 12 juvs, D. B. Herbst, 12 Sept. 1997; spring No. 223, 5
cfcf, 2 99 , 9 juvs, D. B. Herbst, 14 Sept. 1997; spring No.
217, 22 juvs, D. B. Herbst, 14 Sept. 1997; spring No. 219, 3
cfcf, 1 9 , 26 juvs, D. B. Herbst, 14 Sept. 1997.

Diagnosis. A medium-sized species, closely similar to
S, grahami and 5. gradyU but distinguished from those

2 species as follows: maxilla 1 and maxilliped with
fewer setae on inner plate; palms of gnathopod 1 and 2
with more spines; coxal plate of gnathopod 2 broader
than deep; lacking di stoposterior lobes of pereopod 6 or
poorly developed; pleonal plate 2 with fewer ventral
spines; uropod 3 with relatively longer peduncle; and
telson apical margin with very tiny notch. Largest 9,
8.5 mm; largest cf 6.0 mm.

Female. Antenna 1; 50 percent length of body, 30
percent longer than antenna 2; primary flagellum with
17 segments. Antenna 2: flagellum with 7 segments.

Mandibles subequal: spine row with 6-7 plumose
spines; palp segment 2 with 10 rather long setae on
inner margin; palp segment 3 with 12 D setae, 6 E setae,
lacking A, B and C setae. Inner lobes of lower lip small
to vestigial .Maxilla 1 : inner plate with 7 apical, plumose
setae; palp with 10 stiff setae or slender spines apically.
Maxilla 2: inner plate with oblique row of 12 plumose
setae on inner margin. Maxilliped: inner plate with 2
bladelike spines, 3 plumose spines, 3 naked setae
apically and 2 plumose spines on inner margin; outer
plate with setae on inner margin and apex and 5 or 6
small serrate spines on or near apex.

Gnathopod 1 : propod smaller than that of gnathopod
2; pal m straight or slightly convex and about 60 percent
longer than posterior margin, armed with 24 spine teeth
in double row; defining angle with 3 apine teeth on
outsicfe, 2 shorter ones on inside; posterior margin with
2 setae; 3 superior medial setae; 10 inferior medial
setae; dactyl nail long; coxa about 33 percent deeper
than broad, margin with 4 setae. Gnathopod 2: propod
little longer than broad, palm straight or slightly con-

cave medially and armed with 26 spine teeth in double
row; defining angle with 1 long spine tooth on outside,
3 shorter spine teeth on inside; posterior margin ap-
proximately 50 percent length of palm, with row of
setae; 14mostly double-inserted superior medial setae;
about 12 singly inserted inferior medial setae; coxa
broader than deep, margin with 10 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 8 setae. Pereopcx! 4: coxal plate relatively broad
and deep, about as deep as broad, reaching about 33
percent length of basis, margin with 1 1 setae. Pereopod
6 little longer than pereopod 7, about 53 percent length
of body, 13 percent longer than pereopod 5. Pereopods
5-7: bases rather narrow, broader proximally than
distally; posterior margins slightly convex, disto-pos-
terior lobes indistinct; anterior and posterior margins
with variable number of spines and setae; segments 4,
5 and 6 of pereopods 5-7 with longish, slender spines;
dactyls of pereopods 5-7 relatively slender and elon-
gate, that of pereopod 6 typically 30 percent length of
corresponding propcxl. Coxal gills present on pereopods
2-6. Brood plates little expanded distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 setule near distoposterior comer, that of 2 and 3
less convex, each with 1 setule near distoposterior
comer; distoposterior comers rounded; ventral margin
of plates 2 and 3 with 4 spines. Uronites free. Uropod
1 : inner ramus little longer than outer ramus, about 65
percent length of peduncle, with 12 spines; outer ramus
with 7 spines; peduncle with 13 spines. Uropod 2: inner
ramus longer than outer ramus, subequal in length to
peduncle, with 1 1 spines; outer ramus with 5 spines;
peduncle with 5 spines. Uropod 3: peduncle usually
bearing 2 small setae; ramus approximately 30 percent
length of peduncle, with 4 apical spines.

Telson little longer than broad, apical margin entire
or with tiny median notch between spine clusters,
usually bearing 12 spines, some relatively long.

Male. Differing from female as follows: Gnathopod
propods narrow; palm of propod 2 shorter in compari-
son with posterior margin. Uropod 1: peduncular proc-
ess approximately 33 percent length of outer ramus,
bluntly rounded distally, lower margin minutely ser-
rate; inner ramus with 8 -9 spines; peduncle with 7-8
spines. Uropod 2: inner ramus with 12-14 spines,
peduncle with 10-13 spines. Telson with 10-12 apical
spines.

Distribution and ecolc^. This species is recorded
from a series of springs in the Ruby Lake National
Wildlife Refuge, Elko and White Pine Cos., Nevada.

AMPMPAaHCA VOL. 3 NO. 2. NOV. 15, 2001 56

Fig. 9. Stygobromus herbsti, n. sp. Female (8.5 mm). Elko County, Nevada.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2CX)1 57

Fig.lO. Stygobromus herbsti, n. sp. Female (8.5 mm); male (5.5 mm). Elko County, Nevada.

58

AMPFDPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

Etymology. The species is named in honor of its
collector, Dr. David B. Herbst.

Stygobromus trinus^ new species
(Figs. 11, 12)

Material examined. CAUFORNIA. Trinity Co.: Hall City
Cave, about 54.4 km W. of Redding. HOLOTYPE Q
(USNM 1000072), 9 paratypes, D. C. Rudolph, B. Martin
and S. Winterath, 11 Apr. 1979.

Diagnosis. A medium-sized cavemicolous species
closely similar to 5. grahami but distinguished as’
follows: maxilla 2 with fewer setae on inner plate-
palms of gnathopods 1 and 2 straight; coxal plate of
pereopod 3 broader than deep; pleonal plates 1 and 2
with more ventral spines; uropodS with tiny ramus; and
apical margin of telson with very shallow notch. Size rf
largest 9 7.0 mm, (? unknown.

Female. Antenna 1: 60 percent length of body, 80
percent longer than antenna 2; primary flagellum with
18 segments. Antenna 2: flagellum with 7 segments.

Mandibles; spine row with 6 plumose spines; palp
segment 2 with 8 long setae on the inner side; palp
segment 3 with 10-1 1 C setae, few D setae, 3 E setae,
lacking A and B setae. Iimer lobes of lower lip vestigial
or absent. M^lla 1: inner plate with 9 apical, plumose
setae; palp with 6 stiff setae or slender spines apically.
Maxilla 2; inner plate with oblique row of 10 plumose
setae on inner margin, Maxilliped: inner plate with 2

bladelikespines,3plumosespine,2nakedsetaeapically,

and 3 plumose setae on inner margin and apex; oute^
plate with 1 bladelike spine and 5 or 6 small serrate
pines on or near apex.

Gnathopod 1: propod smaller than that of gnathopod

2; palm straight and about 50 percent longer than
posterior margin, armed with 22 spine teeth in double
row; defining angle with 3 spine teeth on outside, 3
shorter ones on inside; posterior margin lacking setae*

4 supenor medial setae; 4 -5 inferior medial setae, all
singly inserted; dactyl nail short; coxa approximately 2
times deeper than broad, margin with 4 setae. Gnatho
fxxl 2: propod subrectangular, longer than broad; palm
sinuate (concave medially) and armed with 23 spine
teeth in double row; defining angle with 1 long spine
tooth on outside, 2-3 shorter spine teeth on inside-
posterior margin about 33 percent length of palm, with
2 sets of triply inserted setae; 8 doubly inserted superior
medial setae; 4 inferior medial setae, singly inserted;
coxa deeper than broad, margin with 4 setae.

Pereopod 3: coxal plate about as broad as deep.

margin with 5-6 setae. Pereopod 4: coxal plate broader
than deep, reaching about 40 percent length of basis
margin with 5 setae. Pereopod 6, slightly longer than’
pereopod 7, about 60 percent length of body, 33 percent
longer than pereopod 5. Pereopods 5-7: bases as broad
or slightly broader proximally; posterior margins nearly
straight; distoposterior lobes distinct but not well de-
veloped; anterior and posterior margins with variable
number of spines and setae; segments 4, 5 and 6 of

pereopods5-7withvariablespines;dactylsofpereopods

5-7 not elongate, that of pereopod 6 about 30 percent
length of corresponding propod. Coxal gills present on
pereopods 2-6, absent from pereopod 7. Brood plates
narrowing distally.

Pleonal plates: posterior margin of plates 2 and 3
slightly convex, with 1 seta each near distoposterior
comer, that of 1 less convex, with 1 setule near
distoposterior comers; distoposterior comer rounded
and indistinct, ventral margin of plate 1 with 4 spines;
ventral margin of plate 2 with 3 spines, that of plate 3
also with 4 spines. Uronites free. Uropod 1: inner
ramus little longer than outer ramus, about 80 percent
length of peduncle, with 9 spines; outer ramus with 8
spines; peduncle with 8 spines. Uropod 2: inner ramus
longer than outer ramus, longer than peduncle, with 12
spines; outer ramus with 6 spines; peduncle with 2
spines. Uropod 3: peduncle without setae; ramus ap-
proximately 25 percent length of peduncle, with 3-4
apical spines.

Telson longer than broad, subrectangular; apical
mar^n with tiny median notch between spine clusters,
bearing 10-12 relatively long spines.

Distribution and ecology. This species is known only
from Its type-locality. Hall City Cave, where it was
collected from wood debris near the edge of a deep pool
(D. C. Rudolph, pers. comm.).

Etymology. The epithet trinus is from the Latin, which
IS based on the geographic place name. Trinity County.

Stygobromus tritus Holsinger, 1974

Stygobromus tritus Holsinger, 1974: 8-11, figs. 2-3

[type-locality; well at Victor Crossing, Ravalli Co.
Montana], ’’

Material examined. MONTANA. Missoula Co.: shallow
water wells (2.6-6.6 m depth) in Missoula, 14 99, 4 <f<S 1
juv., M. E. Ver Hey, May 1986.

Diagnosis. A rather small subterranean species distin-

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 5S

guished by the diagnosis and description of Holsinger
(1974). Largest 9, 5.0 mm; largest cJ, 4.5 mm.

Distribution and ecology. This species is recorded
from a well at Victor Crossing, Ravalli Co. Montana
(see Holsinger 1974) and wells in Missoula County
(see above). Females ranging in length from 4.5 to 5.0
mm have setose brood plates and are sexually mature.

Taxonomic remarks: Re-examination of male speci-
mens revealed an apically serrate peduncular proMss
on uropod 1, which was not described in the original
description by Holsinger (1974).

Stygobronws boultoni, new species
(Fig. 13)

Material examined. ARIZONA. Maricopa Co.; PVC well
no 1 at Sycamore Creek, about 32 km NE Hioemx,
HOLarYPE$(USNM 1000051),4$9paratypes,3juvs.,

A Boulton, 12 Mar, 1990; PVC well no. 1, 2 juvs., A.
Boulton, 2 Mar, 1990; PVC well no. 3 on Sycamore Creek,

1 9, A. Boulton, 2 Mar, 1990.

Diagnosis. A small-sized goundwater species, very
similar to 5. arizonensis but distinguished as follows:
palms of gnathopod 1 and 2 oblique and with fewer
teeth; maxilla 2 with fewer setae; posterior margin of
gnathopods 1 and 2 longer; coxal plates 3 deeper; bases
of pereopods 6 and 7 more narrow and with fewer
spines on posterior margin; telson with longer apical
spines; uropods 1-2 with more long and thick spines on
rami; pleonal plates 2 and 3 with more spines on
posterior margin. Largest 9 4.0 mm, <S unkown.

Female. Antenna 1: 54 percent length of body, 43
percent longer than antenna 2; primary flagellum with
14 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 4-5 plumose
spines; palp segment 2 with row of 3 rather long setae
on inner margin; palp segment 3 with 4 C setae, 5 E
setae, lacking A, B and D setae. Inner lobes of lower lip
vestigial.Maxilla 1: inner plate with 4 apical, plumose
setae; palp with 5 stiff setae or slender spines apically.
Maxilla 2: inner plate with oblique row of 4 plumose
setae on inner margin. Maxillip^: inner plate with 1
bladelike spine, 2 plumose spines, and 3 naked setoe
apically, and without plumose spines on inner margin;
outer plate with 3 setae on inner margin and 4 setae on
or near apex.

Gnathopod 1 : propod smaller than that of gnathopod
2; palm straight or slightly concave and 2 times longer

than posterior margin, armed with 11 spine tee* in
double row; defining angle with 2 spine teeth on
outside, 2 shorter ones on inside; postenor marpn
lacking setae; fewer superior medial setae; laclang
inferior medial, dactyl, nail long; coxa broader ^
deep; margin with 2 setae. Gnathopod 2:
subrectangular, deeper than broad; palm armed with 9
spine teeth in double row; defining angle with 1 long
spine tooth on outside, 2 shorter spine teeth on inside;

posterior margin 50 percentof palm in length, wi* 1 set

of setae; 3 superior medial setae, singly inserted; few
inferior medial setae; coxa broader than deep, margin
with 2 or 3 setae.

Pereopod 3; coxal plate about as broad as deep,
margin with 3 setae. Pereopod 4: coxal plate about as
broad as deep, reaching about 30 percent length o

basi s, margin with 2 setae. Pereopod 6 little longer than

pereopod 7, about 70 percent length of body, 30 pei^nt

longer than pereopod 5. Pereopods 5-7; bases little

broader proximally than distally; posterior margiiw
nearly straight; distoposterior lobes not develope^
anterior and posterior margins with variable number d
spines and setae; segments 4, 5 and 6 of pereop^s 5-
7 with longish spnes; dactyls of pereopods 5-7 rela-
tively slender, that of pereopod 6 typically 25 percent
length of corresponding propod. Coxal gills present on
pereopods 2-6, absent from 7. Brood plates little nar-
rowing distally. , , . , J T

Pleonal plates: posterior margin of plate 1 ana 4
convex, each with 3 spines near distoposterior comer,
that of 3 less convex, with 5 setae near distoposterior
comer; distoposterior comers of plate 2 and 3 rounded;
ventral mar^n of plate 2 with 1 spine, that of plate 3
with 2 spines. Utonites free. Uropod 1; inner ramus
little longer than outer ramus, about 80 percent length
of peduncle, with 8 spines; outer ramus with 6 spines;
peduncle with 5 spines. Uropod 2; inner ramus longer
than outer ramus, subequal in length to peduncle, with
7 spines; outer ramus with 5 spnes; peduncle with 2
spines. Uropod 3; peduncle not bearing setae; ramus
approximately 30 percent length of peduncle, with 2
apical spines.

Telson longer than broad, gently tapered distally;
apical margin with small median notch between spine
clusters, bearing 9 relatively long sjanes.

Distribution and ecology. This species is known only
from wells of PVC ppe drilled into substi^ of Syca-
more Creek, 32 km northeast of Phoenix, Arizona.
According to Dr. Andrew Boulton (pers. comm.), the
material was collected during a study of interstitial

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15. 2001

60

...p. Female (7.0™,. 7™^. Coan,.. Cif™,.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 61

¥lgA2.Stygobromustrinus,n,sp. Female (7.0 mm). Trinity County, California,

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

62

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 63

fauna utilizing phreatic pumping through PVC wells
from a depth of approximately 180 cm.

Etymology. This species is named in honor of its
collector. Dr. Andrew Boulton.

Siygobromus duplus, new species
(Figs. 14, 15)

Material examined. WASHINGTON. Spokane Co.: well
No. 7 in Spokane Valley, near Millwood, HOLOTYPE9 (on
slide mounts) (USNM 1000055). L. A. Fuste, 21 Apr. 1981;
well No. 5 in Spokane Valley, 1 <S paratype , L. A. Fust6, 25
Feb., 1981.

Diagnosis. A medium-sized groundwater species, re-
lated to S. mysticus and S, latus in structure of the
gnathopods, but distinguished from those species as
follows: gnathopod 2 with shorter posterior margin;
bases of pereopods 5-7 much narrower and proportion-
ately longer than other segments; distoposterior lobes
of pereopod 5-7 poorly developed or absent; uropods 1
and 2 with more spines on rami and peduncles; telson
proportionately broader with long spines. Largest (and
only known) 9 5.4 mm; largest (and only known) <S ,
5.0 mm.

Female. Antenna 1: 70 percent length of body, 90
percent longer than antenna 2; primary flagellum with
16 segments. Antenna 2: flagellum with 6 segments.

Mandibles subequal: spine row with 6 plumose
spines; palp segment 2 with row of 7 rather long setae
on inner margin; palp segment 3 bearing 2 B setae, 7 D
setae, and 3 E setae, lacking A and C setae. Inner lobes
of lower lip absent.Maxilla 1 : inner plate with 8 apical ,
plumose setae; palp with 5 stiff setae or slender spines
apically. Maxilla 2: inner plate with oblique row of 10
plumose setae on inner margin. Maxilliped: inner plate
with 2 bladelike spines, 3 plumose spines, and 3 naked
setae apically, and 3 coarse setae on inner margin ; outer
plate with numerous setae along inner margin and 5
setae on or near apex.

Gnathopod 1: propod shorter than propod of
gnathopod 2; palm slightly convex and about 35 per-
cent longer than posterior margin, armed with 12 spine
teeth in double row ; defining angle with 2 spine teeth on
outside, but no shorter ones on inside; posterior margin
with 2 setae; 6 superior medial setae, in unevem row;
few inferior medial setae; dactyl nail rather long and
slender; coxa deeper than broad, margin with 2 setae.
Gnathopod 2: propod subtriangular, almost 2 times
longer than broad; palm armed with 15-16 spine teeth
in double row; defining angle with 1 long and 2 shorter

teeth on outside, no spine teeth on inside; posterior
margin rather short, only about 50 percent length of
palm, with 3 sets of mostly doubly inserted setae; 4
superior medial setae; 6 inferior medial setae; coxa
about as deep as broad, margin with 4 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 5 setae. Pereopod 4: coxal plate deeper than broad,
reaching about 35 percent length of basis, margin with
5 setae. Pereopod 6 little longer than pereopod 7, about
70 percent length of body, 25 percent longer than
pereopod 5. Pereopods 5-7: bases little broader proxi-
mally than distally; posterior margins convex proxi-
mally, but not greatly expanded; distojx>sterior lobes
poorly developed or vestigial; anterior and posterior
margins with variable number of spines and setae;
segments 4, 5 and 6 of pereopods 5-7 with longish,
slender spines; dactyls of pereopods 5-7 not greatly
elongate, that of pereopod 6 approximately 33 percent
length of corresponding propod.Coxal gills present on
pereopods 2-6, absent from 7. Brood plates somewhat
narrowing distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 setule near distoposterior comer and 4 spines on
ventral margin, that of 2 and 3 also convex, each with
2 setae near distoposterior comer; distoposterior cor-
ners rounded and indistinct; ventral margin of plate 2
with 2 spines, that of plate 3 with 2 spines. Uronites
free. Uropod 1: inner ramus little longer than outer
ramus, about 75 percent length of peduncle, with 1 1
spines; outer ramus with 8 spines; peduncle with 1 1
spines. Uropod2: inner ramus longer than outer ramus,
subequal in length to peduncle, with 7 spines; outer
ramus with 5 spines; peduncle with 5 spines. Uropod
3: ramus short and only approximately 25 percent
length of peduncle, with 2 apical spines.

Telson little longer than broad, gently tapered dist-
ally; apical margin with tiny median notch between
spine clusters, bearing 10-1 1 mostly longish spines.

Distribution and ecology. To date this species is
known only from two specimens from two test wells in
the Spokane Valley near Millwood, Spokane County,
Washington.

Etymology. The epithet duplus is from the Latin,
meaning “double or 2 times,” which refers to the 2 sets
of setae on the posterior margin of the propod of
gnathopod 2.

Remarks: The single male specimen was lost in
handling after initial examination.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 64

Fig. 14. Stygobromus duplus, n. sp. Female (5.4 mm). Spokane County, Washington.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

65

Fig.l5. Stygobromusduplus, n.sp. Female (5.4 mm). Spokane County, Washington.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

Slygobrnmus utahensis, new species
(Fig. 16)

Material examined. UTAH. Duchesne Co.: Pole Creek

^ of

~2128ni,HOLOTYPE9(USNM 1000074), 7099 ISCfd’

paratypes, 1 juv., S. J. Peck, 5 Aug. 1979,

Diagnosis. A medium-sized cavemicolous species ap-
parently very similar to S. arizonensis in structure of

gnathopods anduropodI-2, but distinguishedfrom this

species as follows; coxal plates 3^ broader than deep-
teses of pereopods 6-7 with fewer spines on
distopostenornwgin; pleonal plates 1-3 lacking spines
on ventral margin and fewer setules near disloposterior
comer; peduncular process of uropod 1 of male sharply
rwinted aptcally; and telson with only slight notch.
Largest 9, 5.5 mm, largest cf, 4.0 mm.

Female. Antenna 1; 55 percent length of body, 50
^rcent longer than antenna 2; primary flagellum with
14 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 8-9 plumose
spines ; palp segment 2 wi th row of 6 long setae on inner

margin;palpsegment3bearinglBseta,9Dsetae and

3 setae E, lacking both A and C setae. Inner lobes of
lowerlip vestigial.Maxilla 1: innerplate with 8 apical
plumose setae; palp with 4 stiff setae subapically’
Maxilla 2: inner plate with oblique row of 7 plumose
setae on inner margin. Maxilliped: inner plate with 2
bladehkespines, 1 plumosespine, 1 nakedsetaapically,
Md 3 setae on inner margin; outer plate with setae oii
inner margin and 4-5 on or near apex.

Gnathopod Lpropod smaller than that of gnathopod

2; palm straight or slightly concave, 2 times longer than
postenw margin, armed with 17 spine teeth in double
row; defining angle with 2 spine teeth on outside, 2
shorter ones on inside; posterior margin lacking setae;

4 doubly inserted superior medial setae; 3 singly in-
serted infenor medial setae; dactyl nail rather long and
sharply pointed; coxa about 2 times broader than deep
mar^n with 2 setae. Gnathopod 2; propod longer than’
broad; palm straight or slightly convex, strongly ob-
lique, armed with 13 spine teethindouble row; defining

angle with 1 spine tooth on outside, 2shorterspine teeth

on inside; posterior margin about 30 percent length of
Nm, with 2 sets of doubly inserted setae; 7 suLor
m^al setae, most doubly inserted; 5 inferior medial
setae, most singly inserted; coxa about as broad as
deep, margin with 3 setae.

Pereopod 3: coxal plate slightly broader than deep
margin with 3 setae. Pereopod 4: coxal plate broader

66

than *ep, reaching about 30 percent length of basis
margin with 2 setae. Pereopod 6 little longer than’

pereopod7,about58peroentlength(rfbody,20peicent

longer than pereopod 5. Pereopods 5-7: bases little

broaderproximallythan the distally; posterior margins

convex; distoposterior lobes poorly developed and
^most indistinct; anterior and posterior margins with
2-4 spnes and setae; segments 4, 5 and 6 erf pereopods
^7 with Icmgish, slender spines; dactyls of pereopods
5-7 relatively slender, that of peree^ 6 about 35
percent length of corresponding propod. Coxal gills
jwesent on pereopods 2-6, absent from 7. Brxxxl plates
little expanded distally.

Pleonal plates; posterior margins of plate 1-3
ranvex, each with 1 setule near distoposterior comer
disto^stenw comers rounded and indistinct; ventral
^gin of pJates 1, 2 and 3 lacking spines. Uronites
tw. Urc^xxl 1: inner ramus subequal to outer ramus
about 60 percent length erf peduncle, with 8 spines’
outer i^us with 8 spines: peduncle with 6 spines
Uropod 2: inner ramus Itmger than outer ramus, little
longer than peduncle, with 8 spines; outer ramus with
spines; peduncle with 3 spines. Uropod 3; peduncle
not beanng small setae; ramus approximately 30 per-
cent length erf peduncle, with 2 apical spines.

®jsonlittlelongerthanbroad,gentlytapercddistallv-

apical margin with small median notch between spine
clusters, bearing 8 relatively long spines.

Male. Differing from female as follows: Gnathopod
prop^s smaller and shorter, palm of propod 2 longer in
relation to posterior margin and with fewer teeth on
palm^ margin. Uropod 1: peduncular process lOper-
cent length of outer ramus, sharply pointed distally
upper margin minutely serrate; inner ramus with 5
spines; peduncle with 6 spines. Uropod 2: inner ramus

with 5 spines, peduncle with 2 spines. Telson with 8
apical spines.

Distribution and ecology. This species is known only
irom Its type-locality.

Etymology. The species is namedfor the state of Utah.

Stygobromus intersUtiaUs, new species
(Figs. 17, 18)

Materiale^ined.cOLORADO.EIPasoCo.:hyporheicin

Monument Creek below Palmer
Lake, HOLOTYPE 9 (USNM 1000063), 2 and 2 M

paratypes, J. V. Ward, 5 Oct 1980; Monument Creek, ca. 30
km N of Colorado Springs, 1 O’, J, v. Ward, 27 July 1979;

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 67

Fig.l6. Stygobromus utahensis, n. sp. Female (5.5 mm); male (4.0 mm). Duchesne County, Utah.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 68

Fig. 17. Stygobromus interstitialis, n. sp. Female (3.6 mm). H Paso County, Colorado

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 69

MXPD

Fig.l8. Stygobromus interstitialis, n. sp. Female (3.6 mm); male (3.0 mm) El Paso Co,, Colorado.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 70

Monument Creek in Colorado Springs, 1 9, S. R Canton, 4
Nov. 1987.

Diagnosis. A small groundwater species, very similar
to S. coloradensis but distinguished as follows: max-
illae 1 and 2 with more setae on inner plates; posterior
margin of palm of propod 2 shorter; pleonal plate 3
lacking ventral spines; uropods 1-2 with more long,
thick spines on rami; uropod 3 with very short ramus;
telson with longer apical spines and slight notch be-
tween spine clusters. Largest 9, 3.6 mm; largest cf , 3.0
mm.

Female. Antenna 1: 50 percent length of body, 55
percent longer than antenna 2; primary flagellum with
12 segments. Antenna 2: flagellum with 5 segments.

Mandibles: spine row with 4-5 plumose spines; palp
segment 2 with row of 3 setae on inner margin; palp
segment 3 bearing 1 B seta, 4 C setae, and 4 E setae,
lacking both A and D setae. Inner lobes of lower lip
vestigial/absent.Maxilla 1: inner plate with 6 apical,
plumose setae; palp with 5 stiff setae or slender spines
apically. Maxilla 2: inner plate with oblique row of 6
plumose setae on inner margin. Maxilliped: inner plate
with 1 bladelike spine, 2 plumose spines, 1 naked seta
apically, and 3 plumose spines on inner margin; outer
plate with shortsetaeoninner margin, 3 lightly plumose
setae and 1 small bladelike spine on or near apex.

Gnathopod 1: propod smaller than that of gnathopod
2; palm straight or slightly concave medially, approxi-
mately 2 times longer than posterior margin, armed
with 10 spine teeth in double row; defining angle with
2 spine teeth on outside, 2 shorter ones on inside;
posterior margin without setae; 4-5 singly inserted
medial setae, dactyl nail rather long; coxa about 25
percent broader than deep, margin with 1 seta.
Gnathopod 2: propod longer than broad; palm straight
and armed with 9 spine teeth in double row; defining
angle with 1 long spine tooth on outside, 2 shorter spine
teeth on inside; posterior margin about 50 percent
length of palm, with 1 set of triply inserted setae; 2
superior medial setae, 2 singly inserted inferior medial
setae; dactyl nail relatively short; coxa little broader
than deep, margin with 1 seta.

Pereopod 3: coxal plate about as broad as deep,
margin with 1 seta. Pereopod 4: coxal plate about as
broad as deep, reaching about 25 percent length of
basis, margin with 1 seta. Pereopod 6 little longer than
pereopod 7, about 50 percent length of body, 36 percent
longer than pereopod 5. Pereopods 5-7: bases about as
broad proximally as distally; posterior margins con-
vex; distoposterior lobes poorly developed; anterior

and posterior margins with variable but few spines and

setae; segments 4, 5 and 6 of pereopods 5-7 with

longish, slender spines; dactyls of pereopods 5-7 rela-
tively slender, that of pereopod 6 approximately 30
percent length of corresponding propcxl. Coxal gills

presentonpereopods2-6,absentfrompeieopod7.Brood

plates slightly narrowing distally, but not setose (fully
mature) in material examined.

Pleonal plates: posterior margins of plate 1 - 2
weakly convex, each with 1 setule near distoposterior
comer; distoposterior comers rounded and indistinct;
ventral margin of plate 1 with 1 spine, that of plate 2
with 3 spines. Uronites free. Uropod 1: inner ramus
slightly longer than outer ramus, about 75 percent
length of peduncle, with 8 spines; outer ramus with 7
spines; peduncle with 5 spines. Uro-pod 2: inner ramus
longer than outer ramus, subequal in length to pedun-
cle, with 9 spines; outer ramus with 4 spines; peduncle
with 3 spines. Uropod 3: peduncle not bearing setae;
ramus tiny, approximately 20 percent length of pedun-
cle, with 2 apical spines.

Telson longer than broad, apical margin with tiny
median notch between spine clusters, bearing approxi-
mately 8 relatively long spines.

Male. Differing from female as follows: gnathopod
propods smaller and narrower, pal m of propod 2 longer
in comparison to posterior margin, with more teeth on
palmar margin. Uropod 1: peduncular process 30 per-
cent length of outer ramus, sharply pointed distally,
inner ramus with 7 spines; outer ramus with 5 distal
spines; peduncle with 5-6 spines. Uropod 2: inner
ramus with 9-10 spines, peduncle with 3-4 spines.
Telson with 9 apical spines.

Distribution and ecolc^iy. This species is recorded
from hyporheic habitats in the deep gravel alluvium of
Mounument Creek at three different sites (see “mate-
rial examined”) in El Paso Co., Colorado.

Etymology. The epithet interstitialis is from the Latin,
meaning “interstitial,” in reference to the habitat of this
species.

Stygobromus puteanus Holsinger, 1974

StygobromusputeanuslioX^mgitr, 1974: ll-13,figs.4-
5 [type-locality: an unnamed well near Three Forks,
Gallatin Co., Montana].

Diagnosis. A medium-sized subterranean species simi -

71

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

lar to S. tritus but distinguished from that species by
the straight to slightly convex palms of gnaAopodal
propods, broader bases of pereopods 5-7; and having
mor^pical spines on uropod 3 and telson. Ingest 9,

6.5 mm; (? unknown (see Holsinger 1974).

Distribution and ecology. This species is known only
from its type locality.

Stygobromus permaki Ward, 1977

Stygobromus permaki Ward, 1977: 458-461, figs. 3-5
[type-locality; hyporheic in North Fork of the South
Platte River, Jefferson Co., Colorado].

Material examined. COLORADO. Arapahoe Co.: inttisti-
tial waters of “shore zone" with Bou-Rouch pump in Sou*

Platte River,Uttleton(ca. 2.5km belowChatfieldreservoir),

IQ i(? S P Canton, 9 Feb. 1988; Jefferson Co.: hyporheic

habitats’ in North R.rk of South S R »

as follows: 0.48km W of S. Hatte, 2 99. 26 M^. 1985, 2g,
5Apr. 1985and599,9Apr. 1985 ; 0.8 km \^S. Hatte, 3^,
ICJ 5Apr. 1985; behind S. Platte Hotel,599,5Apr. 1985,

7 QQ 9 Apr. 1985. and 14 99, 5 Nov. 1985; 100 m
downstream from jet. with N. Fork, 3 99, 5 Apr. 1985, near
gaging station in S. Platte, 2 99, 9 Apr. 1985.

Diagnosis. A small hyporheic species, apparently re-
lated to S. puteanus but distinguished by longer
gnathopod propod 1 with more oblique palm; more
teeth on palmar margin; fewer marginal setae on coxal
plates 3 and 4; more spines on posterior margin ot
pleonal plates, fewer spines on ventral margins; and
uropod 3 with proportionately larger ramus. Largest 9,
4.3 mm; largest cJ, 4. 1 mm.

Distribution and ecology. This species is knownfrom
hyporheic habitats in the North Fork of South Platte
River, Jefferson County, and interstitial waters of the
shore zone, Arapahoe County, Colorado (see also
Pennak and Ward 1986). As noted later, this species
was often collected with 5. coloradensis in hyporheic
habitats in the South Platte River.

Stygobromus urospinatus, new species
(Figs. 19, 20)

Stygobromus sp. A. - Waid & Holsinger, 1981 : 64-67.

Material examined. COLORADO. Larimer Co^Bellvue
Spring, about 1 .6km SEof Bellevue, HOLOTYPE9 (USNM
1000073) 13 99 paratypes, J. V. Ward, 15 Sept 1977; 1 Cf,
16 99. J. V. Ward, 2 Apr. 1978; 3 (S(S, 3 99, J- V. Ward, 29

Jan. 1978; Redstone spring no. l.westofHorsetoothR^r-

voir 1 <? J. V. Ward, 30 Oct 1976; spnng at base of S[mng

SLli>amofHorsetoothReservoir.999,J.V.Wm^^

Apr. 1975.

Diagnosis.Amedium-sizedgroundwaterspeciessinu-

lar to S. holsingeri but differing as follows: palm^
gnathopods 1 and 2 straight; coxal plates of perwpods
3 and4d^r than broad; pleonal plates 1-3 withf ewer
ventral spnes but more setules on postenOT margin,
uropods 1-3 more heavily spinose; and telson with

numerous and longer spines on apical margin. Largest

9, 7.0 mm; largest cf, 6.0 mm.

Female. Antenna 1; 46 percent len^h «f »

percent longer than antenna 2; primary flagellum
\2 segments. Antenna 2: flagellum with 8 segments.

Mandibles subequal; spine row with 8 plumose
spines; palp segment 2 with about 12 long set« on
inner margin, segment 3 beanng 1 tong se ,

setae, approximately 14Dsetae.and5Eseme, lacking

A setae. Inner lobes of lower hp vestigial or absent.
Maxilla 1: inner plate with 10 apical, plumose setae;
palp with 13 stiff setae or slender spines apically.
Maxilla 2; inner plate with oblique row of 11 plumose
setae on inner margin. Maxilliped: inner plate with 2

bladelikespines,3plumosespines,lnakedsetaapicaly

and 2 plumose setae on inner margin; outer plate with
setae on inner mar^n and 5 setae on or near apex.
Gnathopod 1 : propod subequal in size to gnathojxxl

2 ;palmstraightandnearly 2 timeslongerthan^stenor

margin, armed with 26 spine teeth in double row,
defining angle with 3 spine teeth on outside, 4 shorter
ones on inside; posterior marpn without setae; beanng

single superiormedial seta and lacking infenor medial

setae; dactyl nail short; coxa approximately 2 times
broader than deep, margin with 3 setae. Gn^opi^ 2:
propod longer than broad; palm long and straight,
armed with 29 s{»ne teeth in double row ; defimng angle
with 1 long spine tooth on outside, 3 shorter spine teeth
on inside; posterior margin relatively short, only ap-
proximately 33 percent length of palm, with 3 tets
setae, 2 mostly triply inserted; beanng 4 doubly in-
serted superior setae and 4 singly inserted inferiOT

m«halsetae;coxalittle broader thandeep, margin with

5 setae.

Pereopod 3: coxal plate broader than deep, margin
with4setae.Pereopod4: coxal plate broader than deep,
reaching approximately 25 percent length of basis,
margin with 6 setae. Pereopod 6 little longer than

pereopod7,about70percentlengthofbody,33 percent

longer than pereopod 5. Pereopods 5-7; bases nanow.

AMPHIPAOHCA VOL. 3 NO. 2. NOV. 15, 2001

72

Fig. 19. Stygobrommurospinatu,,n.sf. Female (7.0 mm). Larimer County, Colorado.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001

73

Fig. 20. Stygobromus urospimtus^ n. sp. Female (7.0 mm); male (6.0 mm)

Larimer County, Colorado.

AMPmPAQHCA VOL 3 NO. 2. NOV. 15, 2001 74

about as broad ptoximally as distally; posterior mar-
^ns nearly straight; distoposterior lobes almost indis-
tinct, not broa^y rounded; anterior and posterior mar-
gins with variaUe number of spnes and setae; seg-
ments 4, 5 and 6 of pereopods 5-7 with slender spines;
dactyls of pereopods 5-7 relatively slender and elon-
gate, that of pereopod 6 typically about 25 percent
length of corresponding propod. Coxal gills present on
pereopods 2-6, absent from pereopod 7.

Pleonal plates: posterior margins weakly convex,
with 7 setules on posterior margin of plate 3 and 5
setules on margins of plates 1 and 2; distoposterior
comer indistinct; ventral margin of plates 1 and 2
bearing 1 spine, that of plate 3 hearings spines. Uron-
ites free. Uropod 1: heavily spinose, especially inner
ramus; inner ramus subequal in length to outer ramus,
about 70 percent length peduncle, with 18 long
spines; outer i^us with 15 spines; peduncle with 19
long, strout spines. Uropod 2; heavily spinose overall,
inner ramus longer than outer ramus and peduncle, with
30 long, strout spines; outer ramus with 14 mostly long
spines; peduncle with 10 spines. Uropod 3: peduncle
without setae; ramus appoximately 50 percent length
of peduncle, with 5 apical spines (3 relatively long).

Telson longer than broad, little broader distally,
apical margin with tiny median notch between spine
clusters, bearing 16 relatively long, strout spines.

Male. Differing from female as follows: gnathopod
propods slightly broader; palm of propod 2 slightly
longer in comparison with posterior margin and bear-
ing more sets of setae. Uropod 1 : peduncular process 25

percentlengthrfouterramus, tapered toapoint distally,

inner ramus with 15-17 long spines; peduncle with 20
spines. Uropod 2: inner ramus with 28-35 long spines

peduncle with 10-12 spines. Telson with 16-19aiMcai
spines.

1-2 [type-locality: hyporheic in North Fork of the

South Platte River, Jefferson Co., Colwado].

Materlalexamined.COLORADO.JeffeisonCo.:hyporheic

in North Fork of South Platte River, by S. P. Canton as
follows: upstream of confluence with mainstream 1 Q 4
A^ 1984; 0.48 km W of S. Platte, 1799,26 Mar. l^Salid
39$.5Apr. 1985;0.8kmWofS.Platte,1799,5Apr 1985-

just below jet. with N. Fork, S. Platte. I 9. 5 Apr. 1985; 100
m downstream from jet. with N. Fork, 2 99 5 Apr 1985-
upstream from Dam Rock, 7 99. 9 Apr. 1^; behind S.’

S’ ^ ® 1“®* upstream from jet.

PI* ft ^ S.

Platte, 25 femlae, 9 Apr. 1985; S. Platte. 2 99, 9 Apr. 1985.

Diagnosis. A very small hyporheic species, distin-
guished by possession of 4 apical plumose setae on
inner plate of maxilla 1 ; oblique row of 4plumose setae
on margin of inner plate of maxilla 2; gnathopodal
propods small and robust, palmar margins slightly
oblique and distinctly serrated, few teeth on palmar
margin; uropod 3 with 1 short and 1 longer spine.
Largest 9, 3.3 mm; largest cf, 3.0 mm.

Distribution and eulogy. This species is known from
hyporheic habitats in the North Fork of South Platte
River, Jefferson Co., Colorado (see Pennak and Ward
1986). In most collections made in March, April and

November 1985, this species was found together with

S. pennaki.

Stygobromus blinni, new species
(Hg. 21)

Distribution and ecology. This species is recorded
from Beilvue Spring, the type-locality, and two other
spnngs near Horsetooth Reservoir, west of Ft. Collins.
Itwascollectedtogetherwith 1 specimenafS.holsineeri
on30Oct. 1976fromRedstcme Spring no. 1 . All sites
are in Larimer County.

E^molr^. The epithet urospimtus isfiom the Latin,
meaning “spines on uropods,” in reference to the heav-
ily spinose uropods of this species.

Stygobromus coloradensis Ward, 1977
Stygobromus coloradensis Ward, 1977: 452-456, figs

Material examined. ARIZONA. Coconino Co.: Roaring
Spnngs Cave, on the north rim of the Grand Canyon near
nght Angel Trail and Bright Angel Creek, HOLOTYPE Q
(USNM 1000050), 2 99 paratypes, D. Blinn, 28 Sept. 1994.

Diagnosis. A medium-sized cavemicolous species,
very similar to S. arizonensis in gnathopods and S,
holsingeri in uropod 2, but distinguished from these 2
species by the following: maxilla 2 with fewer setae on
inner plate; maxilliped with bladelike spines on outer
plate (unlike S. holsingeri)\ coxal plate 4 broader than
deep; bases of pereopods 5-7 narrow and with more
seme on the margin; uropod 1 with more long and stout
spines on rami and peduncle; uropod 2 with more long
and stout spines on rami; urop^ 3 ramus smaller;

telson notched slightly. Largest 9,6.0 mm, d* un-
known.

AMPMPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 75

MXPD

Fig. 21. Stygobromus blinni, n. sp. Female (5.8 mm). Coconino County, Arizona.

76

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001

Female. Antenna 1: 67 percent length of body, 25
percent longer than antenna 2; primary flagellum with
16 segments. Antenna 2: flagellum with 7 segments.

Mandibles subequal: spine row with 7-8 plumose
spines; palp segment 2 with row of 6 rather long setae
on inner margin; palp segment 3 bearing 1 long B seta,
5 C setae, few D setae, 4 E setae, lacking A setae. Inner
lobes of lower lip vestigial or absent.Maxilla 1: inner
plate with 5 apical, plumose setae; palp with 7 stiff
setae or slender spines apically. Maxilla 2: inner plate
with oblique row of 6 plumose setae on inner margin.
Maxilliped: inner plate with 2 bladelike spines, 3
plumose spines, and 2 naked setae apically, and 2
plumose spines on inner margin; outer plate with setae
on inner margin and apex, and 1 lightly plumose spine
on or near apex.

Gnathop^ 1: propod smaller than that of gnathopod

2, palm straight and little longer than posterior margin,
armed with 17 spine teeth in double row; defining angle
with 2 spine teeth on outside, 2 shorter ones on inside;
posterior margin lacki ng setae; superior medial setae 3 ;

2 inferior medial setae; dactyl nail rather long; coxa
rather narrow, deeper than broad, margin with 3 setae.
Gnathopod 2: propod oblique, deeper than broad; palm
straight and armed with 17 spine teeth in double row;
defining angle with 1 long spine tooth on outside, 2
shorter spine teeth on inside; posterior margin about 50
percent length of palm, with 1 set of setae near defining
angle; 8 - 9 superior medial setae, doubly inserted; 4
singly-inserted inferior medial setae; coxa deeper than
broad, margin with 4 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 6 setae. Pereopod 4: coxal plate relatively broad
and deep, reaching about 33 percent length of basis,
margin with 5 setae. Pereopod 6 little longer than

pereopod7,about80percentlengthofbody, ISpercent

longer than pereopod 5. Pereopods 5-1: bases as broad
proximally as distally; posterior margins slightly con-
vex; distoposterior lobes weakly developed, that of 7
indistinct; anterior and posterior margins with variable
number of spines and setae; segments 4, 5 and 6 of
pereopods S7 with longish, slender spines; dactyls of
pereopods 5-7 relatively slender and elongate, that of
pereopod 6 about 45 percent length of corresponding
propod. Coxal gills present on pereopods 2-6, absent
from 7. Brood plates somewhat narrowing distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 senile near distoposterior comer, that of 2 and 3
less convex, each with 2 setae near distoposterior
comer; distoposterior comers rounded; ventral margin
of plate 2 with 1 spine, that of plate 3 with 2 spines.

Uronites free. Uropod 1 : inner ramus little longer than
outer ramus, about 75 percent length of peduncle, with
8-9 spines; outer ramus with 8-9 spines; peduncle with
12 spines. Uropod 2: inner ramus longer than outer
ramus, subequal in length to peduncle, with 10 spines;
outer ramus with 6 spines; peduncle with 4 spines.
Uropod 3: peduncle usually lacking small setae; ramus
approximately 30 percent length of peduncle, with 2
apical spines.

Telson little longer than broad, apical margin with
small median notch between spine clusters, bearing 10-
1 1 relatively long spines.

Distribution and ecology. This species is only known
from its type-locality. It was collected from a slow-
moving stream with a fine sediraent/gravel substrate,

aboutl.6kminside the cave (DeanBlinn.pers. comm.).

Etymology. It is pleasure to name this species for its
collector. Dr. Dean Blinn.

Stygobromus holsingeri Ward, 1977

StygobromusholsingeriWard, 1977; 461-465, figs. 6-
8 [type-locality: small unnamed spring/seep, 3.2 km
W. of Ft. Collins, Larimer Co., Colorado]. Ward and
Holsinger 1981: 64-65.

Material examined. COLORADO. Larimer Co.: several
habitat types associated with interrupted stream drainage
near Horsetooth Reservoir, W of Ft. Collins, approximately
240 specimens. J. V. Ward, May 1976 - Jan. 1978 (most in
1977) as follows (number of collections in parentheses):
Devils Gulch (1); Redstone site 1-5 (12); Redstone LS (2)-
Redstone NB (5); Redstone SB (3); Soldier Canyon (7).

Diagnosis. Amedium-sizedgroundwaterspecies, simi-
lar to S. anzonensis but distinguished by inner plate of
maxilliped with plumose setae extending partway down
innermargin,gnathopodal propod 1 with more oblique
palm, 1 or2fewerteethonpalmarmargin;distoposterior

lobes of pereopod 6 and 7 better developed; telson
shorter with a shallower notch and more spines on
apical lobes (see Ward 1977; Ward and Holsinger 1981
for details). Largest 9 5.7 mm; largest (3 5.4 mm.

Distribution and ecology. This species is recorded
from seeps, springs and interrupted stream drainage in
the vicinity of Horsetooth Reservoir in Larimer Co.,
Colorado (see Ward and Holsinger 1981 for further
details).

77

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

Stygobromus arizonensis Holsinger, 1974

Stygobromusarizonensis^dmigtx, 1974: 47-49,figs^
28-29 [type-locality: small cave on Hying “H” Ranch
near Fort Huachuca, Cochise Co., Arizona].

Material examined. ARIZONA. Cochise Co.: Mine Cave,
south of Siena Vista, 2 99, R- B. Pape, 17 Sept. 1994; S^m
Cruz Co.: bog springs, Madera Canyon, 2 (S<S, 1 juv., W. D.
Sheppard, 3 June 1993.

Diagnosis. A relatively small to medium-sized
cavemicolous species, distinguished by the diagnosis

and description ofHolsinger(1974).Largest97.0ram;

largest cJ* 5.0 mm.

Distribution and ecology. This species is recorded
from 4 localities (2 caves, 1 mine and 1 spring) m
southeastern Arizona. Mine Cave is a natural cave that
was broken into by miners and is located on the oppo-
site side of Huachuca Mountains from the type-locality
cave on Hying “H” Ranch (R. B. Pape, pers. comm.).
The specimens from this cave were collected from a
pcx>l approximately 1.8 m deep and 61 m from the
entrance. Bog spring is a spring/seep located approxi-
mately 48 km NNW of Fort Huachuca.

Stygobromus quatsinensis Holsinger and Shaw, 1987

Stygobromus n. sp. Holsinger and Shaw, 1986. 79.
Stygobromus quatsinensis Holsinger and Shaw, 1987:
222-229, figs. 1-3 [type-locality: Thanksgiving Cave,
Vancouver Island, British Columbia, Canada], Hol-
singer etal. 1997:347-348.

Diagnosis. A medium-sized groundwater species, dis-
tinguished from most other members of the hubbsi
group by presence of 2 or 3 submarginal setae on distal
part of posterior margin of propod of gnathopod 1,
inserted below defining angle; and absence of distal
peduncular process on uropod 1 of male. Largest 9 9.0
mm; largest d* 7.0 mm.

Distribution and ecology. This species was originally
described from two cave populations on Vancouver
Island, British Columbia, Canada (Holsinger and Shaw
1986, 1987). Subsequently, the species has beenfound
in caves elsewhere on Vancouver Island and in karst
groundwater habitats (caves and resurgences) on sev-
eral islands in the Alexander Archipelago of southeast-
ern Alaska (Holsinger etal. 1997). Ovigerous females

(6.0 -7.0 mm) occurred in two July samples from
Alaska.

Stygobromus wahkeenensis, new species
(Hgs. 22, 23)

Stygobromus sp.- Bousfield and Holsinger, 1989:
968.

Material examined. OREGON. Multnomah Co.; railroad
bank spring, Wakkeena Creek in Columbia River gorge
(elev. 183 m), HOLOTYPE 9 (on slide mounts m part)
(USNM 1000075), R. W. Wisseman, 17 Mar. 1989; seep or
hyporheic habitat associated with Wahkeena Creek,
km SSW of Bonneville, 3 99 paiatypes, 2 juys^ C^C
2001-0023), G. W. Couitney, 19 Apr. 1985, 1 9, 1<1^ (CMNC
2001-0024), 6 Mar. 1986.

Diagnosis. A small groundwater species distinguished
as follows: palm of gnathopod 2 straight or slightly

concavemediallyandwithfewteethonpalmarmargin;

maxilliped with bladelike spine cat outer plate; bases of
peieopods 5-7 rather narrow; pleonal plates 2-3 with 3
ventral spines each; uropod 1-2 moderately spinose;
and telson longer than broad. Largest 9, 4.0 mm,
largest cT, 3.0 mm.

Female. Antenna 1: 50 percent length of body, 46
percent longer than antenna 2; primary flagellum with
14 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 4 plumose
spines ; palp segment 2 with row of 4 long setae on inner
margin; palp segment 3 with 6 D setae and 4 E setae,
lacking A, B and C setae. Inner lobes of lower lip
absent-Maxilla 1: inner plate with 7 apical, plumose
setae; palp with 5 stiff setae or slender spines apically.
Maxilla 2: inner plate with oblique row of 6 plumose
setae on inner margin. Maxilliped: inner plate with 1
bladelike spine, 4 plumose spines, and 3 naked setae
apically, and 1 plumose spine on inner margin; outer
plate with setae on inner margin and apex, and small

bladelike spine on or near apex.

Gnathopod 1: propod 1 subequal in size to that of
gnathopod 2 but of different shape; palm straight or
slightly concave and approximately 65 percent longer
than posterior margin, armed with 16 spine teeth in
double tow; defining angle with 2 spine teeth on
outside, 2 shorter ones on inside; posterior margin
without setae; 4 superior medial setae; 4-5 inferior
medial setae; dactyl nail short; coxa about 2 times
broader than deep, margin with 1 seta. Gnatho^ 2;
propod longer than broad; palm straight or slightly

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

78

Fig. 22 . Stygobromuswahkeenemis,n.sp. Female (3.8 mm). Multnomah County,

Oregon.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 79

AMPMPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 80

Fig. 24. StygobrommUinemis,n.^j>. Female (9.0 mm). Lane County.

Oregon.

amphipacihca

VOL. 3 NO. 2. NOV. 15, 2001 81

Fig. 25. Stygobromus lanensis^ n. sp.

I. Female (9.0 mm). Lane County, Oregon.

AMPMPACIHCA VOU3 NO. 2.

NOV. 15. 2001 82

OTiiMve medially and armed with 14 spine teeth in
doubk row; defining angle with 1 long spine tooth on
outside, 2 shorter spine teeth on inside; posterior mar-
&n 30 percent length c£ palm, with 2 sets of doubly
malted setae; 6-7 doubly inserted superior medii
setae, 2 iirfenw medial setae; coxa broader than deep
margin with 1 seta.

Pereopod 3: coxal plate about as broad as deep

ZIT r*/ 4: coxal plate little’

broader than deep, reaching about 30 percent length of

^reop(rf7,about60percentlengthQfbody,40percent

longer than pereopod 5. Pereopods 5-7: bases about as
bioadproximally as distally;posteriormargins weakly
conyex; distopostenor lobes well developed; anterior
and posterior margins with variable number of spines
and setae; se^ents 4. 5 and 6 of pereopods 5-7 with
lon^sh slender spines; dactyls of pereopods 5-7 rela-

hydyslenderandelongate,thatQfpereopod6approxi-

^tely 35 percent length of corresponding propod
Co^ gills present on pereopods 2-6, absent fro^
Br<^ plates sublinear or slightly narrowing distally
Fl^nal plates: posterior margins of plate 1 - 3
str^ght or slightly convex, with 1 setule each near

comers rounded

Md toat of plate 3 indistinct; ventral margin of plates 2
Md 3 each with 3 spines. Uronites free. Uropod 1-
inner ramus little longer than outer ramus, about 45
^rcent length of peduncle, with 7 spines; outer ramus
with6spines;peduncle with 1 1 spines. Uropod 2- inner

^us ongerthanouter ramus, littlelongerthanpedun-

cle, with 7 spines; outer ramus with3 spines; peduncle

with spine. Uropod3:peduncleustJlyb4ringlt

2 sm^l setae; ramus approximately 45 percent length
of peduncle, with 3 apical spines. ®

Telsonsubrectangular,longerthanbroad,apexwith

ttny mediOT notch between spine clusters, bean^g 9-10
relatively long spines. iw

contained 18 and 9 specimens of
5fyg^yx co^tneyi Bousfield and Holsinger (see
Bousfield and Holsinger 1989).

Ecology. The epithet wahkeenensis is in reference
to the associaUon of this species with Wahkeena Creek.

Stygobromus lanensis, new species
(Rgs. 24, 25)

10000651 S W (USNM

V "• ^ hyporheic/seep area

Trail Creek, Willamette National Forest, 1 9 and 1 cT para-

1 JUV. (CMNC 2001-0022), G. W. Courtney, 22^y

Wagi^is.Amedium-sizedgroundwaterspecies,simi-

ar to5. arizonensis but distinguished as follows- pos-
tenor margin of gnathopod 1 shorter; distoposterior
lotes of pereopod 6 distinct; pleonal plate 3 with 1
setule Ml postenor margin and 2 spines on ventral
m»gm; uropods I and 2 with more long spines on rami
onl “ore apical spines and

oriyslight apical notch. I^gest9.9.0mm; largest cf

V ’ ) mm.

follows: gnathopod

^I«dsbroader,palmofpropod 2 shorterinrelatiS

^toor margn, with fewer teeth on palmar margin.
Uro^ 1. peduncular process 30 percent length of

s“™“'’"“^°'^'"8‘^**«''y’“PPcrmargin™
senate, inner ramus with 6 spines; peduncle with 3

with 2 spines. Telson with 10-12 apcal spines.

^ibution and ecology. This species is recoided
Ae type-locality spring and a nearby seep and/or
hyporfieic zone associated with Wahkeena Creek in

Female. Antenna 1: 56 percent length of body, 80
^rcent longer than antenna 2; primary flagellum with

A 6 segments

Mandibles: spine row with6-7plumose spines palp
segment 2 with row of 10 setae on inner m^gin; Sp

^ckingAandCsetae.Innerlobesoflowerlipvestigial
^UalMiiner plate with6apical,plumosesetae;^lp

with 5 snff seme apically. Maxilla 2: inner plate ^th
obl^e row of 6 plumose setae on inner margin. Max-
dhped. inner plate with I bladelike spine, 3 plumose
spines, and 2 naked setae apically. iLking

few setae on inner margin and 4 setae and 1 bladelike
spine on or near apex.

Gnathopod 1. propod slightly shorter than that of
3 Umes lOTpr than posterior margin, armed with 17

Zh r 'I* 1 ‘'Pine

^th on outside, 1 shorter one on inside; posterior
M^n lactang setae; 1-2 superior medial setae and 3
infenor medial selae; dactyl nail short, all singly in-

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 83

serted; coxa about 2 times broader than deep, margin

with 4 setae. Gnathopod 2; ’

palm convex mediaily and amed with 21-22 spine
^th in double row; defining angle with 1 long spine
tooth on outside, 3 shorter spine teeth on inside; poste-
rior margin approximately 50 percent length of palm
with 3 sets of setae; 5 superior medial setae, doubly
inserted; 4 inferior medial setae; coxa deeper than

broad, margin with 4 setae.

Pereopod 3; coxal {date about as deep as br^,
margin with 4 setae. Pereopod 4: coxal plate little
deeper than broad, reaching about 35 percent length of
basis, margin with4setae. Pereopod 6 little longer than

pereopod7,about 60 percentlength of body,50 percent

longer than pereopod 5. Pereopods 5-7: bases broader
pioximally than distally; posterior margins nearly
stimght; distoposterior lobes poorly developed; ante-
rior and posterior margins with vanable number
spines and setae; segments 4, 5 and 6 of pereopods S
7 with longish, slender spines; dactyls of
7 not particularly elongate, that of pereopod 6 ab^t 30
percent length of corresponding propod. Coxal ^lls
present on pereopods 2-6, absent from pereopod .
Brood plates slightly narrowing distally.

Pleonal plates: posterior margins of plates 1-3 con-
vex, each with 1 setule near distoposterior corner,
distoposterior comers rounded and incUstinct; ven^
marginsof plates l,2and3 each with2 spines. Uronites
free Uropod 1: inner ramus slightly longer than outer

ramus, about 80 percentlength of peduncle, with 1^11

spines; outer ramus with 10 spines; peduncle with 1
spines . Uropod 2: inner ramus longer than outer ramus,
longer than peduncle, with 7 spines; outer ramus wiA
6 s(Hnes; peduncle with 4 spines. Uropod 3; peduncle
usually bearing 1 small setae; ramus approximately 45
percent length of peduncle, with 4 apical spines.

Telson subrectangular, longer than broad; apical
mar^n with tiny median notch between spine clusters,
bearing 12-13 relatively long spines.

Distribution and ecology. This species is only known
from its type-locality, where it was discovered ^
pumping from a cobble/gravel fill at a depth of 50- 100

cm.

Etymology. This species is named for Lane County,
Oregon.

Stygobromus sierrensis Holsinger, 1974
Stygobromus sierrensis Holsinger, 1974: figs.

20-21 [type-locality: an unnamed spring in Trosi Can-
yon, Sierra Co., California].

Diagnosis. A medium-sized groundwater
distinguished by shallow coxal plates cf ^thopod 2

and pereopods3 and4;2 ventral spines each Mpleon^

plates; proportionately short ramus of ^pod 3; ^d
near absence of a{Mcal notch of telson. Largest 9 . 7 .
mm; largest cJ , 5.2 mm.

Distribution and ecology. This species is known only
from its type-locality.

Stygobromus putealis Holmes, 1909

Stygobromusputealis Holmes, 1909:77-78,figs. [type-
locality: well at Waupun, Dodge Co., Wisconsin.

Diagnosis.Amedium-sizedgroundwatersp^ies,mor-

phologically closely similar to many members of the
hubbsi group much farther west. This species lacks
sternal processes and setae on the posterior margin of
the propod of gnathopod 1. Largest 9, 6.0 mm; largest

cf , 4.0 mm.

Distribution and ecology. To date, all collections of
this species have been from wells in

Wisconsin, approximately 56 km east of the Drrftless

Area.” It is the only species of the hubbsi group foun
in central North America east of the western Cordillera.

Remarks: A detailed redescription of this species is
being prepared by Holsinger (ms.).

Stygobromus lacicolus Holsinger, 1974

Stygobromus lacicolus Holsinger, 1974: 4447 figs.
26-27 [type-locality : Lake Tahoe at Cave Rock, Doug-
las Co., Nevada].

Diagnosis. A medium-sized, deep-lake dwelling
stygomorphic species, that occurs sympatrically wi
S, tahoensis but is distinguished from that specify
longer palms and more spines on gnathopodal propods;

deeper coxal plates of pereoix>ds3and4,broader^^^^^

of pereopods 5-7;fewer spines on uropods 1 and2;ancl

longer ramus of uropod 3. Largest 9 , 6.0 mm; largest
cf , 5.5 mm.

Distribution and ecology. This species is only known

from samples taken frran Lake Tahoe. It is recorded

AMPMPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

84

Fig. 26. Stygobrommmyersae, Female (5.0 nun). Inyo County. California.

AMPHlPAaFlCA VOL. 3 NO. 2. NOV. 15. 2001 85

MXPD

Fig. 27. Stygobromus myersae, n. sp. Female (5.0 mm); male (3.8 mm). Inyo County, California.

AMPHIPAOHCA VOL. 3 NO. 2. NOV. 15, 2001
v^ous sites in Uke Tahoe in H Dorado and

SO™)

Stygobromus sheldoniVlcAsmset, 1974

l^yg^romus sheldoni Holsinger, 1974: 37-40, figs.

[type-locality: bog spring (elevation 1,920 m)
tnbutary to Sagehen Creek. Nevada Co., California].

86

Diagn^ A medium-sized groundwater species dis-
hngmshed as foilows: propod of gnathopS 1 with
relatively long postenor margin; propod <rf gnathopod
2 propoilKmately smaller; posterior margin of j^eo^

CesTle o’" '“ckingapical notch,

i-argest 9, 6.0 mm; largest cf, 4.5 mm.

nn7 f T 7 Station

Diagnosk A small to medium-sized groundwater spe-
cies probably closely related to both S. mackensiei Md
5. iierre/isw. It differs from the latter by smaller size at
sexual matunty, fewer spine teeth on palms of propod

^ P™Po«ionately smalls

gnatho^ propod 2 of male, and proximally broader
tese^f pereopods 5-7. Largest 9. 5.5 mm; largest cf.

Distribution ecology. This species is recorded

1 ^ «l®''a‘'ons between

1915 and piO m (above sea level) that feed Segehen

Creek in Nevada Co., California (see Holsinger 1974 ).

7 by Barr and

Shepard both the male and female are at least 1.5 mm

onger ttan any of the specimens described by Hol-

reZ H ^ ‘be prevLsly

insteadtT i®" '^Ser female has I

instead of 2, apical spines on the ramus of uropod 3; and

12, instead of 10, apical spines on the telson. The larger

‘be gnathoiL

propods and 2^ more apical spines on the telson.

Stygobromus myersae, new species
(Figs. 26, 27)

Material examined. CAUFORNIA. Inyo Co.: Black Can-

Myers, 17

» CTcf, 6 99, 3 juvs., M. Myers, 4 Aue 1996'
unnamed hillside spring, ca. 7km S of Black Canyo/sp^’

Sjuvs., M. Myem, 29 July 1998; 7 juvs., unnamed hLile
spnng m Marble Canyon ca 7 km 8
Sp.mg,3Nov. 1998;NlvADA.;:^,^^rcXsS''

Myem,^ ^ 9?. M.

Female. Antenna I: 46 percent length of body 45

2= primary flagellum with
10 ^gmen s. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 6-7 plumose
spines; palp segment 2 with row of 4 rather long setae
on inner margin; palp segment 3 with 8 D setae 4 E
setae, lacking A, B and C setae. Innerlobes of lower lip
vestigial or absent.Maxilla 1 : inner plate with7apical
plumose setae; palp with 7 stiff setae or slender spines’
apically. Maxilla 2: inner plate with oblique row rf 7

wr?7!7r Ma»'bped: inner plate
ladelike spine, 3 plumose spines, 3-4 naked
setee apcally, and 2 plumose spines on inner margin-
outer plate with setae on inner margin and setae and 1
or z small spines on apex.

bttle smaller than propod rf

fu7i°7 f ’ 7" ^b^btly conver^d

subequal in length to posterior margin, armed with 16
spine teeth in double row; defining angle with 3 spine
eeth on outside, 3 shorter ones on inside; posterior
margn lactang setae; 3 superior medial setae, 4 singly
msei^d irfenor medial setae; dactyl nail rather short

roxab^derthandeep,marginwith3selae.Gnathopod

2. propod relauvely small, longer than broad; p^
stiteght and ^ed with 16 spine teeth in double Sv-
jMmng angle with 1 very long spine tooth of unequal
^gth on outside. 1 shorter spine tooth on insL;

of ni^tly doubly inserted (occasionally triply) setae; 6

do^lyinse^d superior medial setae,4sin^yinserted

itfenor m^al setae; coxa about as broad as deep,
margin with 3 setae. ”

‘ban broad,

mjgin with 4 setae. Pfereopod 4: coxal plate about as
^p as broad, re^hing about 40 percent length cf

tt;J!?rS!!i7 J b little longer

an peiwpod 7, about 62 percent length of body 33

I^ent lOTger than pereopod 5. Pereopods 5-7: ^es

atom as broad proximally as distally; posterior mar-

'°b®® developed,
roady rounded; anterior and posterior margins ^th
viable "Mberof spines and setae; segmente4, 5and

6ofpereoptos5-7withlongish. slender spines;dactyls
of pereopods 5-7relativelyslenderandelongate.thatrf

87

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15. 2001

sponding propcxi. Coxal gills present on
6^bsentfrom 7. Brood plates little expimded distally.
Pleonal plates: posterior margin of plate 1

with 1 setuleneardiste^lerior owner, those of2and

3 less convex, each with 1 setule near distopwtenor
comer; distoposterior comers rounded and iiAstmct
ventral margin of plate 2 without spines, that cf plate 3
with 1 spine. Uionitesfree. Uropod 1; inner ramus htUe
longer than outer ramus, about 90 percent length o
peduncle, with 7 spines; outer ramus with 7 spines;
peduncle with 7 spines. Uropod 2: inner ramus longer
to outer ramus, subequal in length to p^uitele, with
6 spines; outer ramus with 5 spines; peduncle w,* 3

spines.UropodS; peduncleusually without small setae,

ramus approxinmtely 55 percent length of peduncle,
with 2 apical spines.

Telson subrectangular, longer than br^d; aja
margin typically entire, occasionally with tiny medi^
notch or “break” between spine clusters, bearing 12
relatively long spines.

Male. Differing from female as follows: gnatho^
propods proportionately shorter, palm of pro^ 2
slightly longer in relation to posterior margin and with
fewer teeth on palmar margin. Uropod 1: pedunculm
process 33 percent length of outer ramus, sharply
pointed distally, upper margin minutely serrate; inner
ramus with 5 spines; peduncle with4spmes. Urop^2:
inner ramus with 7 spines, peduncle with 4 spines.
Telson with 12- 14 apical spines.

Distribution and ecology. This species is recorded
from two springs, approximately 7 km apart, in tayo
Co., California, and one spring in Esmeralda Co^
Nevada, about 64 km to the east. Samples from Black
Canyon Spring were obtained through a PVC pi^
pushed into the substrate and with an aquanum net (M.
Myers pers. comm.). The water temperature in the
spring was about ICW:. Elsewhere, in the unnamed
spring in Marble Canyon, the water temperature was
about 12.8”C. Females ranging in size from 5.0 - 6.0
mm have setose brood plates and are sexually mature.

Etymology. This species is named in honor of its
collector, Marilyn Myers, who collected several go^
samples of this species and furnished useful data on its

ecology.

Stygobromus cherylae, new species
(Figs. 28, 29)

Materlalexamined. CALIFORNIA. SonomaCo.: unnamed
spring box, just east of Maacama Creek Bndge, beside

Highway 128, approximately 19.5 km east of GeyseiviUe
HOLCy^PE9 (USNM 1000052). and 60
and JRH), C. B. Barr and W. B. She^. 21 ftb. 2^.
spring box and feeder pipe in the same locauon 1 <?. 1 ju^
C B Bair, 5-7 Oct 1995; feeder pipe at spnng Iw 193 km
it of G^yserville. 1 9, 2 juvs.. 2^28 Apr. 1996; p^ at

unnamed spring on hillside, 19.5km east of Geyserville. 99,

33 juvs., 26-28 Apr. 1996.

ably closely related to me granunu r-

fardiertotheeast.butcharacterizedbysin^lerit^esat

sexual maturity. Further distinguished from S. gra-
hami by having fewer setae on inner plates rf maxillae

andmaxilliped;neariystraighttoslightlycoiivexp^ms

of propods of gnathopods 1 and 2; wxal plates 1 and 2
proportionately smaller; pleonal plates 2 an
ftw« ventral spines; apical margin of telson with tiny
notch. Largest 9 , 7.5 mm; largest <f, 4.5 mm.

Female. Antenna 1: 60 percent lengA of ^y, ^
percent longer than antenna 2; pnmary flagellum wi
“8 segments. Antenna 2: flagellum ^ segmen^
ftodibles subequal: spine row with 5-6 plumose
spines; palp segment 2 with 6 long setae on itmer
margin, 2 on outer margin; palp segment 3 wi* 5 C
setae, about 10 D setae and 5 E setae, 'aching
and B setae. Inner lobes of lower lip absent.\tolla L
inner date with 8 apical plumose setae; palp with 5 stiff
setae or slender spnes apically. Maxilla 2: inner plate
with oblique row of 9 plumose setae on inner margin.
Maxilliped: inner plate with 2 bladelike spnes, 2
plumose spine, and 2naked setae apically, and 2^se
plumose setae on inner margin; outer plate with setae
on inner margin and apex, and 1 bladelike spine on or

”%iShopod l:propodsmallerthanthatofgnathop<^

2- palm slightly convex or straight and approxiinately
2 times longer than posterior margin, armed with 22
spine teeth in double row; defining angle with 3 spine
teeth on outside, 2 shorter ones on inside; postenor
margin without setae; 4 doubly inserted superior me-
dial setae;4or5inferiormedial setae;dactyl nail short,
coxa approximately 2 times broader than deep, margin
with 3 setae. Gnathopod 2: propod deeper than broad;
palm ccmvex medially and armed with 21-22 spine
teeth in double row; defining angle with 1 long spine
tooth on outside, 1 shorter spne tooth on inside; inte-
rior mar^n about 25 percent of length palm, with 2 sets

of doubly or triply inserted setae; 9 - 10 doubly inserted
superior medial setae; 4 or 5 singly inserted infenot
medial setae; dactyl nail shorter; coxa broader than
deep, margin with 4 setae.

AMPHIPACIHCA VOL. 3 NO. 2. NOV.

15, 2001 88

n*.2*. P™i.,7.0™,. Sono™ County,

AMHUPAaHCA VOL.3 N0.2. NOV. 15. 2001 89

Fig. 29. Stygobromus cherylae, n. sp. Female (7.0 mm); male (4.5 mm). Sonoma Co., California.

AMPKPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

Pereopod 3: coxal plate about as broad as deep
margn with 3 setae. Pereopod 4 : coxal plate about as
broad as deep, reaching about 33 percent length of
basis, margin with 4 setae. Pereopod 6, slightly longer
than pereopod 7, about 57 percent length of body 33
percent longer than pereopod 5. Pereopods 5-7: b^s
as broadproximally as distally; posteriormargins vari-
ably convex; distoposterior lobes moderatedly well
developed, anterior and posterior margins with vari-
able number of spines and setae; segments 4 , Sand 6of
pereopods 5-7 with longish, slender spines; dactyls of
pereopods 5-7 relatively slender, that of pereopod 6
approximately 30 percent length of coiresponding
propod. Coxal gills present on pereopods 2-6, absent

d^staH^"^”*^ ^ slightly narrowing

Pleonal plates: posterior margin of plate 3 convex,
with 1 setule near distoposterior comer, thatrfplates 1
and2iessconvex,eachwith 1 setule neardistoposterior
comer; distoposterior comers rounded and indistinct-
ventral margin of plates 2 and 3 each bearing 3 spines!
Urorates free. Uropod 1 : inner ramus little longer than
outer ramus, about 75 percent length of peduncle, with
10spines;outerramus with 1 1 sjanes; peduncle with 12
spines. Uropod 2: inner ramus longer than outer ramus,
onger than peduncle, with 10 spines; outer ramus with
spines; peduncle with 3 sjanes. Uropod 3: peduncle
not l^ng setae; ramus approximately 30 percent
length of peduncle, with 3-4 apical spines.

Telson longer than broad, apical margin with tiny
median notch between spine clusters, bearing 14-16
relatively long spines.

Mrie. Differing from female in a number of characters
as follows; sexually mature specimens approximately
3.0 mm shorter; antenna 1 with longer esthetascs, and
presence of peduncular process on uropod 1, which is
approximately 20^25 percent length of outer ramus.
Gnathopod propods smaller, palm of propod 2 longer
than posterior margin, with fewer teeth on palmar
margin. Uropod 2: inner ramus with 9-1 1 spines, pe-
duncle with3-5 spines. Telson with 13- 15apical spines.

90

Distributionandecology. This species is only known
irom Its type-locality.According to C. B. Barr (pers.
<^mm.), who collected most of the specimens, some of
the waterfrom the spring is diverted at the spring head
mtoafeederpipe that carries water downhill toa spring
box. Specimens were obtained from both the feeder
pi^ (by netting) and spring box. The large 21 February
roUection was made by jacking through roots and
aebns in the upper spring box.

It is a pleasure to name this species for
Uhei^ B. Barr in recognition of her collections of
^phipodsfrom springs in western North America and

forsuj^lyingoursmdy with useful data cm habitats and

collecting techniques

Remwks. A second, smaller (undescribed) species of
Stygobromus, occurs with 5. cherylcu in the type-
I^ity, but was not detected until the collection of 21
Febr^ 2000 was examined. The second species,
which IS distingmshed by significantly smaller size at
sexud maturity, proportionately shorter appendages

broaderbasesQfpereopods6and7,andfewerspineson

the uropods and telson, will be described in a subse-
quent paper.

Stygobromus cowani, new species
(Fig. 30)

Material examined. CAUFORNIA. Napa Co.: roadside
pipe from unnamed spring. Soda Canyon Road, approxi-
mately 6.4 km north of junction with Silverado Trail and
about 6.4 km east of Y ountville, HOLOTYPE 9 (on 2 slide
mounts in part) (USNM 1000053), 1 c? (in poor shape) 1
JUV., C. B. Bair, 23 Feb. 1991.

Diagnosis. A small groundwater species closely re-
lated to S. gradyi but distinguished as follows: maxilla

lwith8-9setaeoniimerplate;maxilla2andmaxilliped

wiA fewer setae on inner plate; lower lip without inner
lobes; palm rfpropod of gnathopod 2 slightly concave-

coxal plate of pereopods 3 and 4 deeper than broad ; and

ventral margin of pleonal plate 2 with fewer spines
Largest 9, 4.6 mm; <f ca. 3.5 mm.

Female. Antenna 1; 52 percent length of body, 76
^rcent longer than antenna 2; primaiy flagellum with
4 segments. Antenna 2: flagellum with 6 segments.
Mandibles subequal.- spine row with 6-8 plumose
spines; palp segment 2 with 5 long setae on inner
rnwgin; palp segment 3 wi th S6 C setae, 5 D setae and
5 E setae, lacking both A and B setae. Inner lobes of
lowerhp vestigial orabsent. Maxilla 1: inner plate with
9 apical plumose setae; palp with 5 stiff setae or slender

spines apically.Maxilla2;innerplate with oblique row

ct lOplumose setae on innermargin. Maxilliped: inner
plate with 3 blade-like spines, 3 plumose spines and 1
naked seta apically on inner margin; outer plate with
setae oninner margin and 10-12 lightly plumose setae

on or near apex.

Girathopod Upropod shorter than that of gnathr y^

2; palm straight or slightly concave medially, little

AMPHIPACIFICA

VOL. 3 NO. 2. NOV. 15, 2001 91

MXPD

F\g.dti.Stygobromuscoyvani,n.sp. Female (4.6 mm). Napa County, California.

AMMPAaHCA VOL. 3 NO. 2 . NOV. 15, 2001

longer than posterior margin, armed with 1 6 spine teeth
m double row; defining angle with 3 spine teeth on
^tside, 4 shorter ones on inside; posterior margin not
^ng setae; 3 singly inserted superior medial setae;
smglyinserted inferior medial setae;dactyl nail short;
OTxa about as broad as deep, margin with 3 setae
Onathopod 2 : propod longer than broad; palm straight
or very weakly caicave, and armed with 16 spine teeth

mdoublerow;defininganglewithllong spine toothon

outside, 2 shorter spine teeth on inside; posterior mar-

gmap^owmatelySOpercentlength of palm, with 1 set

of tnply inserted setae; 5 -6 singly inserted superior
medial setae; 4 singly inserted inferior medial setae;

COM little deeper than broad, margin with 3 setae.

Pereo^ 3: coxal plate about as deep as broad,
i^gn with3 setae. Pereopod4: coxal plate relatively
broad, broader than deep, reaching about 40 percent
length of basis, margin with 5 setae. Pereopod 6 little
teger than pereopod 7, about 44 percent length of
y. 1 1 percent longer than pereopod 5 in length.
Perec^s 5-7: bases broader proximaJly than distally;
postenor margins convex but more strongly convex in
pereopod 7; distoposterior lobes relatively well devel-
oj^, broadly rounded; anterior and posterior margins
with viable number of spines and setae; segments 4 ,

5 and 6 of pereopods 5-7 with longish, slender spines-
dactyls of pereopods 5-7 relatively slender, that of
pereopod 6 typically 33 percent length of coirespond-
mg propod. Coxal gills present on pereopods 2-6

absentfr^pereopod7.Broodplatessublinear,slightly

broader distally. ® ^

Pleonal plates: posterior margin of plate 1 weakly
Mnvex, with 1 setule near distoposterior comer, that of
2 and 3 less convex, each with 1 setule near distopost-
enor comer; distoposterior comers rounded, indistinct-
venti^ margin of plate 2 with 2 spines, that of plate 3
wiA 3 spines. Uronites free. Uropod I: inner ramus
little longer than outer ramus, with 7 spines, about 75
percent length of peduncle; outer ramus with 7 spines-
peduncle with 7 spines. Uro[x>d 2 : inner ramus longed
than outer ramus, subequal in length to peduncle, with
6 spines; outer ramus with 4 spnes; peduncle with 2
spines. Uropod 3; peduncle narrow, lacking small
setae; ramus also narrow, approximately 30 percent
length of peduncle, with 2 apical spines.

Telson little longer than broad; apcal margin with

ny median notch between spine clusters, with 1 1-12
relatively long spines.

92

sha^. The male specimen is covered by fungi and can
not be measured accurately or dissected.

E^oI^.This species is namedfor David Cowan, in
recopition of his able assistance with the cdlection of
amphipods from subterranean waters in California.

Stygobromus tahoensis Hdsinger, 1974

Stygobromustahoensis msinger, 1974: 4(M4 lies
^25 [type-locality: Lake Tahoe, at about 61 m depth,

tetween Skunk Harbor and Tahoe City, Placer Co
California]. *’

htoterial examined. CAUFORNIA.: Lake Tahoe in Chara
^ at 61n, depth, 15 99, 3 <?(? paiatypes, E R. Byron, fall

Diagnosis. A medium-sized, deep-lake dwelling

st)jc»norphicspecies,apparentlyrelatedto5.sierren«i

Md S. sheldoni, but distinguished from these 2 species
by the diagnosis and description of Holsinger (1974)
Largest 9 , 6.5 mm; largest <f, 5.0 mm.

Distributionand ecology. This species is known only
rom Lake Tahoe, where a majority of specimens were
collected at depths of 60A95 m (see Holsinger 1974).

Stygobromus mackenziei HdlBinggr, 1974
Stygobromus mackenzieiHolsingeT, 1974:32-35 fip,

18-19 [type-locality: Empire Cave, Santo Cruz’ Co ’
California].

Material ex^lned. CALIFORNIA. Santa Cruz Co.: Em-
pire Cave, 5 9, D. C. Rudolph, D. Cowan and B. van Ingen
22 Apr. 1979; 2 99. T. S. Briggs, 4 Dec. 1983.

Diagnosis. A small cavemicolous species, distantly
related to species of the Mother Lode region of CaJifor-

maand distinguished by havlng4apicalspinesoninner
plate(rfmaxilli^:Iightlyspinedpalmsofgnathopodal

prop^s, relatively large, deep coxal plates rf
gnathopods Md pereopods 3 and 4; proportionately

r 5-7, lightly spined peduncles

of uropc^s 1 and 2 , and 8 apical spines on telson.
Largest 9 , 6.0 mm; largest cf, 4.0 mm.

amphipacihca

VOL. 3 NO. 2. NOV. 15, 2001 93

April 1979 sample measured 6.0 mm and had setose
brood dates. One specimen was ovigerous with 7
embryos in the brood pouch. This collectitm dso con-
tained 1 specimen of Stygobromus impenalis n. sp.

(describedbelow). The December 1983 collection was

taken from a flooded room with 2 specimens o
Calasellus califomicus (see Lewis 2001) and 2
stygobiont ist^s d the genus Caecidotea.

Stygobromus oregonensis Holsinger, 1974

StygobromusoregonensisHMn^r, 1974: 13-16,figs.

6-7 [type-locality: small unnamed cave near Roseburg,
Douglas Co., Oregon].

Material examined. OREGON. Douglas Co.: a small cave
near Roseburg (type-locality), 2 99 topotypes, Jim Riggs, 1
Jan. 1983.

guished by having fewer plumose setae on inner plates
onmaidllae 1 and 2; fewer apical spines on inner plate

of maxilliped;andfewer apical spines onapical margin

of telson. Largest 9, 6.5 mm; largest cf, 5.5 mm.

Distribution and ecology. This species is known only
from a lake in the type locality, Malheur Cave, itoey
Co Oregon. Accorthng to John Markham (pers.
cot^.), who is involved with a recent research pro-
ject in the cave, amphipods are attracted to baited traps
left in the cave lake. A recent map and descnption ot

Stygobromus mysticus Holsinger, 1974
Stygobromus mysticus Holsinger, 1974: 49-52, figs.

30-31 [type-locality: subterranean habitat, Greenview,

Siskiyou Co., California].

Diagnosis. A moderately cavemicolous species easily

distinguishedfromotherspeciesofthe/i«fr^«groupby

numerous long, stiff setae and slender spines on
peduncular segments 4 and 5 of antenna 2; proportion-
ately large propod gnathopod 1 with very long, heavily
spined palm; convex palm of gnathopodal propod 2
with double row of 16 spine teeth; convex posterior
margins of bases of pereopods 5-7; midlateral position
of posterior subacute distoposterior comers of pleonal
plates 2 and 3 ; and larger ramus of uropod 3. Largest
9, 13.0 mm; cf, unknown.

Diagnosis. A medium-sized subterranean species, ^

tinguished by possession of relatively smallgnathopod

propods; deep coxal plates; broadly expanded toses rf
pereopods 5-7; narrow outer rami of uropod 1 and 2,
and long, distally tapering telson. Largest 9, 8.0 mm;
cf, unknown.

Distribution and ecology. This species is known only
from subterranean habitat, possibly from a well, in the
Klamath Mountains, Siskiyou Co., California
(Holsinger 1974).

Distribution and ecology. This species is known only
from type-locality. Two samples, containing only fe-
males, have been collected from a pool in the bottom of
the cave. According to Jim Riggs (pers. comm.), the
cave is a single passage developed in the Eocene-aged
Umpqua formation, which was formed by faulting an<^
or fi ssuring in sandstone or siltstone conglomerate. It is
not developed in basalt as stated earlier by Holsinger
(1974).

Stygobromus hubbsi Shoemaker, 1942

Stygobromus hubbsi Shoemaker, 1942: 1-6, figs. 1-2
[type-locality: Malheur Cave, Harney Co., Oregon].
See Holsinger (1974) for earlier synonymy.

Diagnosis. Corresponding to the redescription by
Holsinger (1974) . A medium-sized cavemicolous spe-
cies, closely allied morphologically with species of
Stygobromus from east-central California (especially
S. grahami, S. harai, and 5. wengerorum) but distin-

Stygobromus hyporheicus, new species
(Figs. 31, 32)

iterial examined. CAUFORNIA. Marin Co.: gravel-bed

rilJlittentstieam.CronanCreek,HOLOTYPE9(U^^

D T Till V 1997.

Diagnosis. A medium-sized interstitial species, similar
to S. grahami and S. wengerorum but distinguished
from those as follows; maxillae 1 and 2 withfewer setae

on inner plates; palms of gnathopod 1 and 2 convex or
straight, and with more spine rows; coxal plates 3 and
4 broader than deep; pleonal plates 2 and 3 with fewer
ventral margin spines, but with more posterior margin
spines; uropods 1-3 with more spines on peduncl^ ^d
rami; telson 2 times deeper than broad. Largest 9» 7.0
mm; Cf, unkown.

Female. Antenna 1: 70 percent length of body, 78
percent longer than antenna 2; primary flagellum with

AMPMPACIHCA VOL. 3 NO. 2. NOV.

Fig. 31 . Stygobromus hyporheicm, n.

sp. Female (7.0 mm). Marin County, California.

AMPMPAanCA VOL. 3 NO. 2. NOV. 15, 2001 95

¥i%.i2.Stygobrommhyporheicm,n.s^. Female (7.0 mm). Marin County, California.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15. 2001 96

21 segments. Antenna 2: flagellum with 7 segments.

Mandibles; spine row with 7-8 plumose spines;palp
segment 2 with row of 11 setae on inner margin,
segment 3 bearing 3-4 C setae, approximately 8 D
setae, 3 E setae, lacking A and B setae. Inner lobes of
lowerlip vestigial or absent. Maxilla 1: inner plate with
8 apical plumose setae ; palp with 7 stiff setae or slender
spines apically or subapically. Maxilla 2: inner plate
with oblique row of 9 plumose setae on inner margin.
Maxilliped: inner plate with 2 bladelike spines, 9
plumose spines, 2 naked setae apically and 1 plumose
seta on inner margin; outer plate with few setae, 1
bladelike spine, and 8-9 lightly plumose setae on or
near apex.

Gnathopod 1 : propod smallerthanthatof gnathopod
2; palm straight or slightly convex and about 67 percent
longer than posterior margin, armed with 35 spine teeth
in double row; defining angle with 3 spine teeth on
outside, 3 shorter ones on inside; posterior margin
lacking setae; 3 superior medial setae; 3 -4 inferior
medial setae ; dactyl nail short; coxa broader than deep,
margin with 3 setae. Gnathopod 2: propod deeper than
broad; palm straight or slightly convex and armed with
34 spine teeth in double row; defining angle with 2
spine teeth of unequal length on outside, 2 shorter spine
teeth on inside; posterior margin approximately 30
percent length of palm, with 2-3 sets of doubly inserted
setae; 6 doubled inserted superior medial setae; 3
inferior medial setae, 2 medial setae; coxa slightly
broader than deep, margin with 4 setae.

Pereopod 3: coxal plate broader than deep, margin
with 8-9 setae. Pereopod 4: coxal plate broader than
deep, reaching approximately 50 percent length of
basi s, margin with 8-9 setae. Pereopod 6 slightly longer
than pereopod 7, about 70 percent length of body, 30
percent longer than pereopod 5. Pereopods 5-7: bases
broader proximally than distally; posterior margins
convex proximately; distoposterior lobes developed;
anterior and posterior margins with variable number of
spines and setae; segments 4, 5 and 6 of pereopods 5-
7 with longish spines; dactyls of pereopods 5-7 rela-
tively elongate, that of pereopod 6 approximately 33
percent length of corresponding propod. Coxal gills
present on pereopods 2-6, absent from pereopod 7.
Brood plates slightly narrowing distally.

Pleonal plates: posterior margin of plate 2 convex,
with 4 setae, that of 1 and 3 less convex, plate 1 with 1
setule near distoposterior comer, plate 3 with 4 setules;
distoposterior comers rounded; ventral margin of plates
1 and 2 without spines, that of plate 3 with 4 spines.
Uronites free. Uropod 1: inner ramus little longer than
outer ramus, about 80 percent length of peduncle, with

9-10 spines; outer ramus with 9 spines; peduncle with
10 spines, Uropod 2: inner ramus longer than outer
ramus, little longer than peduncle, with 8 spines; outer
ramus with 7 sjnnes; peduncle with 3 spines. Uropod 3:
peduncle without setae; ramus approximately 33 per-
cent length of peduncle, with 4 or 5 apical spines.

Tel son much longer than broad, apical margin with
small median notch between spine clusters, with 9-10
relatively long spines.

Distribution and ecology. This species is known from
the hyporheic zone of a gravel-bedded intermittent
stream, Cronan Creek, in Golden Gate National Rec-
reational Area, which is adjacent to Point Reyes Na-
tional Seashore. The specimens were collected from a
depth of 40 cm. The water temperature was 16° C.

Etymology. The epithet hyporheicus is from the Latin,
meaning “hyporheic,” in reference to the habitat of this
species.

Stygobromus elliotti Holsinger, 1974

Stygobromus elliotti Holsinger, 1974: 16-20, figs. 8-9
[type-locality: Deadhorse Cave, Skamania Co., Wash-
ington].

Material examined. WASHINGTON. Skamania Co.:
Deadhorse Cave, 1 9 (TBMWSM), R. L. Crawford, 8-9 Oct.
1977; Little Red River Cave (lava tube), 1 9, 1 cf and 1 (J
juv., R. L. Crawford and C. M. Serger, Oct. 1983.

Diagnosis. A medium-sized cavemicolous species ap-
parently distinguished from other species of the hubbsi
group by the long and heavily setose palpal segment 2
of maxilliped; broadly rounded outer lobes and short
lateral processe of lower lip; and proportionately un-
even posterior margins of the bases of pereopods 5-7.
Largest 9, 9.5 mm; largest cf, 7.0 mm.

Distribution and ecology. This species is recorded
from small streams in three lava caves in Skamania Co.,
Washington, twoof which (Deadhorse Cave and Upper
Falls Creek Cave system) are also inhabited by the
stygobiont isopod Salmasellus howarthi (see Hols-
inger 1974; Lewis 2001).

Stygobromus gradyi Holsinger, 1974
(Fig. 33)

Stygobromus gradyi HoXunger, 1974: 23-26, figs. 12-

AMFWPACIFICA VOL.3 N0.2. NOV. 15, 2001 97

Fig. 33. Stygobromus gradyi Holsinger. Female (5.0 mm). Amador County, California.

AMPHIPAOHCA VOL. 3 NO. 2. NOV. 15, 2001 98

Fig. 34. Stygobromuslatus^n,s^> Female (5.6 mm). Spokane County, Washington.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 99

MXPD

Fig. 35. Stygobromus latus^ n. sp. Female (5.6 nun); male (6.0 mm). Spokane County, Washington.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 100

13 [type-locality: Crystal Palace Cave, Tuolumne Co„

California].

Material examined. CALIFORNIA. Amador Co.: Fern
Frond Cave,799,6cfcf, D. C. Rudolph, S. Winterath, E. van
Ingen and D. Cowan, 15 Apr. 1979; unnamed spring. Ma-
sonic Cave, 1 9. Lulubell Cave, 1 9, F- G. Howaith, 17 Oct.
1987; Tuolumne Co.: small unnamed spring, ca. 14.8 km
southeast of Angels Camp, 1 9. D- Cowan, 19 Nov. 1978.

Diagnosis. A medium-sized cavemicolous species
closely related to S. grahami but differing from that
species by smaller size at sexual maturity; lacking inner
lobes on lower lip; proportionately smaller gnathopodal
propod 2 of 9, with shorter and slightly convex palm;

4 spines on ventral margins of pleonal plates 2 and 3;
distally serrate peduncular process of male uropod 1.
Largest 9, 7.0 mm; largest (5*, 4.5 mm.

Distribution and ecology. This species is recorded
from a small, unnamed springs and four caves in
Amador and Tuolumne Cos., California.

Remarks. This species is very relatively similar
morphologically to S. grahami and S. harai, and is
therefore illustrated as reference point for these species
(see Holsinger 1974).

Stygobromus lotus, new species
(Figs. 34, 35)

Material examined. WASHINGTON. Spokane Co.; Millers
Spring No. 1, about 2.6 km S. of Cheney, HOLOTYPE 9
(USNM 1000066), 5 99 and 2 <S<S paratypes, I. Moham-
mad, 5 Dec. 1992; Millers Spring No. 2, 3 99, 1- Mohammad,

5 Dec. 1992; Whitman Co.: Rock Lake Spring, 1 (f, 3 99, 1-
Mohammad, 2 July 1992.

Diagnosis. A medium-sized stygobiont species, simi-
lar to S. mysticus in uropod 3 and broad bases of per-
eopods 5-7, but distinguished as follows: gnathopods 1
and 2 with more teeth on palmar margin; bases of per-
eopods 4-7 much broader and proportionately longer
than restof segments; uropod 2 with more spines on
inner ramus; telson lacking notch, proportionately deep
and with more apical spnnes. Largest 9, 7.5 mm; largest
cf, 6.0 mm.

Female. Antenna 1: 70 percent length of body, 50
percent longer than antenna 2; primary flagellum with
13 segments. Antenna 2: flagellum with 6 segments.

Mandibles subequal: spine row with 5-9 plumose
spines; palp segment 2 with row of 4 long setae on inner
margin; palp segment 3 bearing 2 A setae, 1 B seta, 2-
3 C setae, row of few D setae, and 4 E setae. Inner lobes
of lower lip absent-Maxilla 1: inner plate with 10
apical, plumose setae; palp with 3 stiff setae or slender
spines apically or subapically. Maxilla 2: inner plate
with oblique row of 1 1 plumose setae on inner margin.
Maxilliped: inner plate with 3 bladelike spines, 3
plumose spines, 1 naked setaapically/ subapically and 1
plumose setae on inner margin; outer plate with fewer
setae on inner margin and 3 lightly plumose setae on or
near apex.

Gnathopod 1 propod smaller than that of gnathopod
2; palm subequal in length to posterior margin, armed
with 15 spine teeth in double row; defining angle with
3 spine teeth on outside, 2 shorter ones on inside;
posterior margin lacking setae; few inferior medial
setae; dactyl nail rather short; coxa broader than deep,
margin with 2 setae, one of them longer. Gnathopod 2
propod subrectangular, longer than broad; palm straight
or little convex and armed with 13-14 spine teeth in
double row; defining angle with 2 spine teeth of un-
equal length on outside, 2 shorter spine teeth on inside;
posterior margin subequal to palm in length, with 2 sets
of triply inserted setae; few medial setae; coxa broader
than deep, margin with 5 setae.

Pereopod 3: coxal plate much deeper than broad,
margin with 5 long setae. Pereopod 4: coxal plate large,
broad and deep, reaching about 60-65 percent length of
basi s, margin with 7 setae. Pereopod 6 little longer than
pereopod 7, about 46 percent length of body, 1 5 percent
longer than pereopod 5. Pereopods 5-7: bases very
broad; posterior margins convex and greatly expanded,
expecially that of 7; distoposterior lobes well devel-
oped, broadly rounded; anterior and posterior margins
with variable number of spines and setae; segments 4,
5 and 6 of pereopods 5-7 with longish, slender spines;
dactyls of pereopods 5-7 relatively short, that of
pereopod 6 about 20 percent length of corresponding
propod. Coxal gills present on pereopods 2-6, absent
from 7. Brood plates narrow but apparently submature
on material examined.

Pleonal plates: posterior margin of plate 1 nearly
straight, with 1 seta near distoposterior comer, that of
2 and 3 less convex, each with 1 setule near distopost-
erior comer; distoposterior comers rounded; ventral
margin of plate 2 with 3 spines, that of plate 3 with 5
spines. Uronites free. Uropod 1: inner ramus little
longer than outer ramus, about 66 percent length of

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 101

peduncle, with 6 spines; outer ramus with 6 spines;
peduncle with 7 spines. Uropod 2: inner ramus longer
than outer ramus, longer than peduncle, with 1 1 spines ;
outer ramus with 3 spines; peduncle with 2 spines.
Uropod 3: peduncle usually bearing 1 or 2 small setae;
ramus approximately 45-50 percent length of pedun-
cle, with 4 apical spines.

Telson little longer than broad, gently tapered
distally ; apical margin with tiny median notch between
spine clusters, bearing 12-13 relatively long spines.

Male. Differing from female as follows: gnathopod
propods slightly shorter, palm of propod 2 shorter in
relation to posterior margin and with more teeth on
palmar margin. Peduncular process of uropod 1 about
25 percent length of outer ramus, pointed distally,
upper margin minutely serrate; inner ramus with 6
spines; peduncle with 5 spines. Uropod 2: inner ramus
with 9 spines, peduncle with 3 spines. Telson with 8
apical spines.

Distribution and ecology. This species is recorded
from six springs located to the south and southwest of
Cheney, WA (Mohammad 1995). They include
Giffords, TNWR, Millers Springs 1 and 2, Spokane
Co.; Rock Lake Spring, Whitman Co.; and Dragon
Spring, Adams Co. This species co-occurs with Stygo-
bromus rallus (describled above) in Millers Spring No.

1 and Rock Lake Spring.

Etymology. The epithet lotus is from the Latin, mean-
ing “broad,” in reference to the broad bases of the
pereopods, which easily distinguishes this species from
most others in the genus Stygobromus.

Remarks. This species was not described but was
mentioned by Ibrahim Mohammad in his 1995 unpub-
lished Master’s thesis in the Department of Biology at
Eastern Washington University, Cheney. He referred
to it as '‘^Stygobromus species (stocky body).”

Stygobromus gaUawayaCy new species
(Figs. 36, 37)

Material examined. CALIFORNIA. Butte Co.: an un-
named spring on Rock Creek, about 20.8 km N of Chico,
HOLOTYPE9 (USNM 1000057), 2 d*Cf paratypes, 6juvs.,

J. Gallaway, 28 July 1997; 1 9, 1 fragment. J. Callaway 15
Oct 1996.

Diagnosis. A medium size stygobiont species closely
related to S. grahami but distinguished from that spe-

cies as follows: gnathopod 2 with straight or slightly
concave palmar margin; pleonal plates 2 and 3 with
more spines on ventral margin; uropod 1 with propor-
tionately longer peduncle and with fewer spines; pe-
duncle of uropod 2 with 2 spines; peduncle of uropod
3 with 1-3 spines and proportionately long and broad;
telson only slightly notched. Largest 9, 6.0 mm;
largest cf,5.0mm.

Female. Antenna 1: 63 percent length of body, about
70 percent longer than antenna 2; primary flagellum
with 15 segments. Antenna 2: flagellum with 6 seg-
ments.

Mandibles subequal: spine row with 9 plumose
spines ; palp segment 2 with row of 6 long setae on inner
margin; palp segment 3 with 8 D setae, 7 E setae,
lacking A, B and C setae. Inner lobes of lower lip small .
Maxilla 1: inner plate with 11 apical, plumose setae;
palp with 8 stiff setae or slender spines apically. Max-
illa 2: inner plate with oblique row of 12 plumose setae
on inner margin. Maxilliped: inner plate with 3 blade-
like spines, 4 plumose spines, 2 naked setae apically,
and 5 plumose spines on inner margin; outer plate with
setae on inner margin and 3 bladelike spines on or near
apex.

Gnathopod 1 : propod smaller than that of gnathopod
2; palm about 2 times longer than posterior margin,
armed with 20 spine teeth in double row ; defining angle
with 2 spine teeth on outside, 3 shorter ones on inside;
posterior margin without setae; 2 superior medial setae;
4-6 inferior medial setae; dactyl nail short; coxa ap-
proximately 60 percent broader than deep, margin with
4 setae. Gnathopod 2: propod subrectangular, palm
slightly concave medially and armed with 16 spine
teeth in double row; defining angle with 1 long spine
tooth on outside, 2 shorter spine teeth on inside; poste-
rior margin about 33 percent length of palm, with 2 sets
of triply inserted setae; 8 superior medial setae, doubly
inserted; 4-5 inferior medial setae, 3 long medial setae;
coxa broad as deep, margin with 5 setae.

Pereopod 3: coxal plate broader than deep, margin
with 4 setae. Pereopod 4: coxal plate relatively broad
and deep, reaching about 50 percent length of basis,
margin with 8 setae. Pereopod 6 little longer than
pereopod 7, about 67 percent length of body, about 33
percent longer than pereopod 5. Pereopods 5-7: bases
almost as broader proximally as distally; posterior
margins slightly convex; distoposterior lobes well de-
veloped; anterior and posterior margins with variable
number of spines and setae; segments 4, 5 and 6 of
pereopods 5-7 with short spines; dactyls of pereopods

AMKflPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 102

Fig. 36. Stygobromus gaUawayae, n. sp. Female (6.0 mm). Butte County, California.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 103

Fig. 37. Stygobromus gaOawayae, n. sp. Female (6.0 mm); male (4.8 mm). Butte Co., California.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 104

Fig. 38. Stygobromus harai Holsinger. Female (6.0 mm), Tuolumne County, California.

y-T

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 105

MXPD

Fig. 39. Stygobromus harai Holsinger. Female (6.0 mm). Tuolumne County, California.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 106

5-7 relatively slender and elongate, that of pereopod 6
about 25percentlengthofcorrespondingpropod. Coxal
gills present on pereopods 2-6, absent from 7. Brood
plates slightly narrowing distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 setule near distoposterior comer, that of 2 and 3
less convex, each with 1 setule near distoposterior
comer; distoposterior comers rounded; ventral margin
of plate 2 with 6 spines, that of plate 3 with 5 spines.
Uronites free. Uropod 1 : inner ramus little longer than
outer ramus, about 75 percent length of peduncle, with
9 spines; outer ramus with 8 spines; peduncle with 10
spines. Uropod 2: inner ramus longer than outer ramus,
subequal in length to peduncle, with 8 spines; outer
ramus with 7 spines; peduncle with 2 spines. Uropod 3:
peduncle not bearing setae; ramus approximately 30
percent length of peduncle, with 4 apical spines.

Telson longer than broad; apical margin typically
with tiny median notch between spine clusters, bearing
12 relatively long spines.

Male. Differing from female as follows: gnathopod
propods smaller, palm of propod 2 longer in relation to
posterior margin and with fewer teeth on margin.
Uropod 1: peduncular process 33 percent length of
outer ramus, narrowing sharply distally, upper margin
minutely serrate ; inner ramus with 5-6 sfnnes ; peduncle
with 5 spines. Uropod 2: inner ramus with 6-7 spines,
peduncle with 3 spines. Telson with 13- 14 short apical
spines.

Distribution and ecology. This species is known only
from its type-locality, a spring next to Rock Creek in
Butte Co., California. The temperature of the spring
was about 2QPC (J. Gallaway, pers. comm.).

Etymology. This species is named in honor of its
collector, Ms. Jody Gallaway, who is thanked for
collecting this material and allowing us to study it.

Stygobromus harai Hoi singer, 1974
(Figs. 38,39)

Stygobromus haraiWAsiXigtr^ 1974: 26-29, figs. 14-15
[type-locality: Pinnacle Point Cave, Tuolumne Co.,
California].

Material examined. CALIFORNIA. Tuolumne Co.:
Windeler Cave, near Vellecito, 1 9, B. Ehr, 21 Feb. 1976; 1
9, 2cfcf,W.R. Elliott, 14Jan. 1978; unnamed spring, 15 km

southeast of Sonora, 1599. D. C. Rudolph, S. Winterathand
D. Cowan, 19 Feb. 1979.

Diagnosis. A medium-sized cavemicolous species,
closely similar to S. gradyi and S. grahami but differing
from these 2 species as follows: fewer apical spines on
inner plate of maxilliped; inner margin of posterior
angle of gnathopodal propod 1 with mostly unforked
spine teeth; gnathopodal propod 2 of male proportion-
ately a little larger than propod 2 of female; coxal plates
of gnathopod 2 and pereopods 3 and 4 not as deep;
ramus of uropod 3 with 3 apical spines; and proportion-
ately longer telson about one-third deeper than broad.
Largest 9, 7.5 nun; largest d* , 7.0 nun.

Distribution and ecology. This species is recorded
form two caves, one mine tunnel, and a spring, all in
Tuolumne Co., California . It was collected from a pool
in Windeler Cave and from beneath rocks and organic
debris in the unnamed spring. The spring sample con-
tained 1 ovigerous female (6.0 mm) with 8 embryos in
the brood pouch.

Remarks. This species is closely similar morphologi-
cally to S. grahami and less so to S. gradyi, and is ill-
ustrated here as a reference point for these species.

Stygobromus wengerorum Hoi singer, 1974

Stygobromus wengerorum Holsinger, 1974: 29-32,
figs. 16-17 [type-locality: Bower Cave, Mariposa Co.,
California].

Material examined. CALIFORNIA. Mariposa Co.: Centi-
pede Cave, 1 cT, 6 99» S. J. Shimek, 30 July 1975.

Diagnosis. A relatively large cavemicolous species,
closely similar to butdifferingfrom other cavemicolous
species of the Mother Lode region by proportionately
larger gnathopod propods with more spine teeth on
palms and more spines on ventral margins of pleonal
plates. Largest 9 » 10.0 mm; largest (and only known)
cf, 5.5 mm.

Distribution and ecology. This species is known only
from two caves in Mariposa Co., California.

Taxonomic Remarks: The peduncular process present
in the male is small and triangular in shape. Male
otherwise like female.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 107

Fig. 40. Stygobromus grahami Holsinger. Female (9.0 mm). Calaveras County, California.

Stygobromus grahami Holsinger, 1974
(Figs. 40,41,42)

Stygobromus grahami Holsinger, 1974: 20-22, figs.
10-11 [type-locality: Cave ofthe Catacombs, Calaveras
Co., California].

Material examined. CALIFORNIA. Amador Co.: Fern
Frond Cave, 1^,6 99» D. C. Rudolph, S. Winterath, E. van
Ingen and D. Cowan, 15 Apr. 1979; Masonic Cave, 3 99
(BBM), F. G. Howarth, 17 Oct. 1988; Calaveras Co.: Cave
City Cave (= California Caverns), M. Demarest, 5 <S<S , 22
Mar. 1996 and 31 specimens (CT, 9 and juvs.), 2 Apr. 1996;
Shaws Cave, 5 99, D. C Rudolph, S. Winterath and E. van
Ingen, 17 Apr. 1979.

Diagnosis. A medium-sized cavemicolous species dif-
fering from other species of the hubbsi group by 15 or
16plumose setaeonapexof inner plate of maxilla 2; 12
or 13 apical spines on inner plate of maxilliped; row of
forked spine teeth on the inside margin of the posterior
angle of gnathopodal propod 1 ; deep coxal plates of
gnathopod 2 and pereopod 3 and 4; more spines on
ventral margins of pleonal plates 2 and 3 (except S.
hubbsi). Largest 9, 9.0 mm, largest Cf, 5.0 mm.

Distribution and ecology. This species is recorded
from eight caves in Amador and Calaveras Cos., CA. It

hasbeencollectedtogetherwithS.gro^fynnFemFrond

(small stream) and Masonic caves in Amador County.

Remarks. This species is a rather typical member of
the hubbsi group and is illustrated here as a reference
point for comparison with other species (see also Meth-
ods and Materials).

Stygobromus rudolphi, new species
(Figs. 43, 44)

Material examined. CALIFORNIA. Santa Barbara Co.:
Montgomery Spring in Los Padres National Forest,
HOLOTYPE 9 (USNM 1000070), 6 99. 1 c5’ paratypes, D.
C. Rudolph, 10 May 1974.

Diagnosis. A medium-sized species closely similar to
S. grahami and S. gradyi but distinguished from these
2 species as follows: palm of gnathopod 1 straight;
coxal plates 1 -2 with fewer setae; coxal plates 3 and 4
broader than deep and with fewer setae ; pleonal plate 2
with fewer ventral spines; peduncle of uropod 3 with a
long spine; telson with more spines on apical margin.
Largest 9, 6.0 mm, largest d", 3.0 mm (probably
immature).

Female. Antenna 1: 52 percent length of body, 57
percent longer than antenna 2; primary flagellum with
13 segments. Antenna 2: flagellum with 7 segments.

Mandibles subequal: spine row with 7-8 plumose
spines; palp segment 2 with row of 6 long setae on inner
margin; palp segment 3 with 3-4 C setae, 9 or more D

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 108

Fig. 41. Stygobromus grahami Holsinger. Female (6.7 mm). Amador County, California.

AMPMPAaFlCA VOL. 3 NO. 2. NOV. 15, 2001 109

OPL

Fig.42. Stygobromus grahami Holsinger. Female (6.7 mm); male (5.5 mm) Amador Co., California

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 110

Fig. 43. Stygobromus rudolphi, n. sp. Female (7.0 mm). Santa Barbara County, California,

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

111

Fig- 44. Stygobromus rudolphi, n. sp. Female (7.0 mm); male (3.0 mm)
Santa Barbara County, California.

AMPHIPACIHCA VOL. 3 NO, 2. NOV. 15, 2001 11

setae, 4 E setae, lacking A and B setae. Inner lobes of
lower lip small to vestigial-Maxilla 1: inner plate with
12 apical, plumose setae; palp with 6 stiff setae or
slender spines on apical margin. Maxilla 2: inner plate
with oblique row of 12 plumose setae on inner margin.
Maxilliped: inner plate with 2 bladelike spines, 4
plumose spines, 3 naked setae apically, plumose spines
and/or coarse setae absent from inner margin; outer
plate with numerous naked setae on inner margin and
lightly plumose setae on or near apex.

Gnathopod 1 : propod smaller than that of gnathopod
2, palm straight, 67 percent longer than posterior mar-
gin, armed with 21 spine teeth in double row; defining
angle with 3 spine teeth on outside, 3 shorter ones on
inside; posterior margin not bearing setae; 3-4 superior
medial setae, singly inserted; 6-7 inferior medial setae;
dactyl nail small; coxa broader than deep, margin with
4 setae. Gnathopod 2: propod deeper than broad; p^m
sinuate (concave medially) and armed with 20 spine
teeth in double row; defining angle with 1 long spine
tooth on outside, 2 shorter spine teeth on inside; poste-
rior margin less half length of palm, with 2 sets of
doubly inserted setae; 5 medial setae; coxa broader
than deep, margin with 5 setae.

Pereopod 3: coxal plate broader than deep, margin
with 6 setae. Pereopod 4: coxal plate broader and
deeper than that of pereopod 3, reaching about 40
percent length of basis, margin with 6 setae. Pereopod
6 little longer than pereopod 7, about 60 percent length
of body, about 50 percent longer than pereopod 5.
Pereopods 5-7: bases about as broad proximally as
distally ; posterior margins convex; distoposterior lol^s
well developed; anterior and posterior margins with
variable number of spines and setae; segments 4, 5 and
6 of pereopods 5-7 with longish spines; dactyls of
pereopods 5-7 relatively slender and elongate, that of
pereopod 6 about 33 percent length of corresponding
propod. Coxal gills present on pereopods 2-6, absent
from pereopod 7. Brood plates slightly expanded
distally.

Pleonal plates: posterior margins of plate 1 and 3
convex, each with 1 setule near distoposterior comer,
that of 2 less convex, with 1 setule near distoposterior
comer; distoposterior comers rounded and distinct;
ventral margin of plate 2 with 1 spine, that of plate 3
with 4 spines. Uronites free. Uropod 1: inner ramus
subequal to outer ramus, about 80 p)ercent length of
peduncle, with 10 spines; outer ramus with 9 spines;
peduncle with 8 spines. Uropod 2: inner ramus longer
thjui outer ramus, little longer than peduncle and with
8 spines; outer ramus with 7 spines; peduncle with 3

spines. UropodS: peduncle bearing one spine apically;
ramus approximately 33 percent length of peduncle,
bearing 5 apical spines.

Telson longer than broad, apex with tiny median
notch between spine clusters, bearing 14 relatively
long spines.

Male. Differing from female as follows: palm of
gnathopod 2 shorter in relation to posterior margin;
palmar margin convex and with fewer teeth; Uropod 1 :
peduncular process 25 percent length of outer ramus,
inner ramus with 6 spines; peduncle with 6 spines.
Uropod 2: inner ramus with 7-8 spines, peduncle with
3-4 spines. Telson with 14-17 long apical spines.

Distribution and ecology. This species is known only
from its type-locality, Montgomery Spring,which is
apparently developed in sandstone. Specimens were
collected from a water tank fed by a small seep-like
spring in the San Rafael Mountains (D. C. Rudolph,
pers. comm.).

Etymology. This species is named in honor of its
collector, D. Craig Rudolph, who discovered it during
a survey of the subterranean invertebrates of Califor-
nia.

Stygobromus imperialis, new species
(Rgs. 45, 46)

Material examined. CALIFORNIA. Santa Cruz Co.. Em-
pire Cave, HOLOTYPE 9 (on slide mounts in part) (USNM
1000062), D. C. Rudolph, D. Cowan and B. van Ingen, 22

Apr. 1979.

Diagnosis. A relatively large cavemicolous species
closely similar to S. grahami but distinguished as
follows: palp segment 2 of mandible with 6 rather long
setae on inner margin; maxillae 1 and 2 with more setae
on inner plates; posterior margin of propod of gnatho-
pod 2 with 1 -3 setae; coxal plate of pereopods 3 and 4
deeper and with more setae; more spines on ventral
margins of pleonal plates 2 and 3; uropod 3 with fewer
apical spines. Largest 9, 9.2 mm, cJ* unknown.

Female. Antenna 1: about 58 percent length of body,
70 percent longer than antenna 2; primary flagellum
with 18 segments. Antenna 2: flagellum with 6 seg-
ments.

Mandibles subequal, spine row with 8-9 plumose
spines; palp segment 2 with 6 rather long setae on inner

113

AMFWPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

mar^n; palp segments with 4-5 C, 13 D and 4 E setae,
lacking both A and B setae. Inner lobes of lower lip
small to vestiglal.Maxilla 1; inner plate with 13 apical
plumose setae; palp with 9 stiff setae apcally or
subapically. Maxilla2: inner plate with oblique row of
14 plumose setae on inner margin. Maxilliped: inner
plate with 3 bladelike spines, 2 plumose spines, and 3
naked setae apically/subapically, 3 i^umose spines on
inner margin; outer plate with setae on inner margin; 7
setae and 1 bladelike spine on or near apex.

Gnathoixxi: propod smaller than that at gnathopod
2; palm 2 times longer than posterior margin, armed
with 30 spine teeth in double row; defining angle with

4 spine teeth on outside, 6 shorter ones on inside;

^terior margin withoutsetae;8superior medial setae,

5 infenor medial setae; dactyl nail rather shorter; coxa
about 33 percent deeper than broad, margin with 3
setae. Gnathopod 2: propod subrectangular, longer
than broad; palm sinuate, armed with 25 - 26 spine teeth
in double row; defining angle with 2 spne teeth of
unequal length on outside, 3 long spine teeth on inside-
posterior margin about 1/3 length of palm, with few
setae; 8 superior medial setae, 9 inferior medial setae;

d^tyl nail shorter; coxa broader than deep, margin
with 5 setae.

Pereopod 3: coxal plate deeper than broad, margin

with 9 setae. Pereopod 4: coxal plate relatively broad
anddeep,subquadrate,reachingabout45percentIength

of basis, margin with 9 setae. Pereopod 6 little longer
than pereopod 7, about 65 percent length of body, 20
^rcent longer than pereopod 5. Pereopods 5-7: bases
broader proximally than distally; posterior margins
convex (but not greatly expanded); distoposterior lobes
m^erately well developed, broadly rounded; anterior
and postenor margins with variable number of spines
^d setae; segments 4, 5 and 6 of pereopods 5-7 with
longish, slender spines; dactyls of pereopods 5-7 rela-
hvely shorter, that of pereopod 6 typically 25 percent
lengthtrf-COTrespondingpropod. Coxal gills present on
pereopods 2-6, absent from pereopod 7. Brood plates
somewhat narrowing distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 seta near distoposterior comer, that of 2 and 3
lessconvex, each with 1 seta near distoposterior comer
distoposterior comers rounded; ventral margin of plate^

2 with 3 spines, that of plate 3 with 5 spines. Uronites
Iree. Uropod 1: inner ramus little longer than outer
ramus, about 60 percent length of peduncle, with 7
spines; outer ramus with 9 spines; peduncle with 10
spines. Uropod 2, inner ramus longer than outer ramus
subequal m length to peduncle, with 9 spines; outer

ramus with 5 spines; peduncle with 4 spines. Uropod 3,
peduncle not bearing setae; ramus approximately 30
percent length of peduncle, with 2 apical spines.

Telson little longer than broad, gently tapered
distally, apical margin with tiny median notch between
spine clusters, bearing about 10 relatively long spines.

Distribution and ecology. The species is known only
from a single female specimen.This species was col-
lected from a cave pool up to 1 m in depth (D. C.
Ruldolph, pers. comm.). It was taken together with 5
specimens of Stygobromus tnackenziei.

Etymology. The epithet imperialis is from the Latin,
meaning “imperial,” in reference to the type-locality!
Empire Cave.

Other Species

The following nine species from western North Amer-
ica have not been assigned to the hubbsi group and are
treated separately in this section. They are distin-
guished from species of the hubbsi group by possess-
ing sternal gills (or processes).

Stygobromus canadensis Holsinger, 1980

Stygobromus canadensis Holsinger, 1980: 290-297,
figs. 1 [type-locality: Castleguard Cave, Alberta,
Canada]; Holsinger, 1981: 93 - 95; Holsinger et al
1983: 546-547.

Diagnosis. A small to medium-sized subterranean spe-

cies,distinguishedfromallotherspeciesof5(yg(?^r<?/MW5

by presence of paired sternal gills on pereonites 2-5,
also distinguished by having few segments of pleopod
rami, and having simple, paired lateral sternal gills on

pereopods 6 and 7 . Largest 9, 5.0 mm; largest d* ,55

mm.

Distribution and ecology. To date this species is
known only from a stream in its type-locality,
Castleguard Cave. This is a large alpine-type cave
situated partly beneath a glacier and located about 90
km south of Cadomin in Banff National Park, Alberta,
Canada (see Holsinger 1980; Holsinger et al. 1983).

Stygobromus idahoensis, new species
(Figs. 47, 48)

Materialexamined.IDAHO.UmhiCo.;Shallowhyporhei^

habitat at mouth of Wilson Creek, a tributary to the middle

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 114

Fig. 45. Stygobromus imperious, n.sp. Female (9.2 mm). Santa Cruz County, California.

AMKBPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 115

Fig. 46. Stygobromus imperious, n.s^. Female (9.2 mm). Santa Craz County, California.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 116

Fig. 47. Stygobromus idahoensis, n. sp. Female (3.5 mm). Lemhi County, Idaho.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 117

Fig. 48. Stygobromm idahoensis, n. sp. Female (3.5 mm); male (3.0 mm). Lemhi County, Idaho.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 1 18

fork of Salmon River [45° 02 '57'’ N, 1 14° 43 3 1 ' ’ W.), coll.
-HOLOTYPE 9 (USNM 1000061), 1 CJ’and399pamtypesi
P. Koetsier, probably late 1986 (no exact date on the collec-
tion label).

Diagnosis: A small groundwaterspecies, distinguished
by small size, rather broad bases of pereopods 5 - 7,
especially 7; 2 long spine teeth at defining angle of
propod of gnathopod 2; and apically heavily spinose
uropods 1 and 2. Largest 9,3.5mm, largest d*, 3.0 mm.

Female. Antenna 1: 47 percent length of body, 40
percent longer than antenna 2; primary flagellum with
13 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 5-7 plumose
spines; palp segment 2 with row of 3 rather long setae
on inner margin; palp segment 3 bearing 1 B seta, 6 D
setae, and 4 E setae, lacking both A and C setae. Inner
lobes of lower lip small to vestigial. Maxilla 1, inner
plate with 6 apical, plumose setae; palp with 5 stiff
setae or slender spines apically. Maxilla 2: inner plate
with oblique row of 5 plumose setae on inner margin.
Maxilliped: innerplate with 1 bladelikespine,4plumose
spines, and 2-3 naked setae apically, and 2 plumose
setae on inner margin; outer plate with setae on inner
margin and 6-7 lightly plumose setae on or near apex.

Gnathopod 1: propod little smaller than piopod of
gnathopxxi 2; palm straight, little longer than posterior
margin, armed with 11 spine teeth in double row;
defining angle with 2 spine teeth on outside, 2 shorter
ones on inside; posterior margin lacking setae; 3 supe-
rior medial setae; 2 -3 inferior medial setae; dactyl nail
moderately short; coxa approximately about as broad
as deep; margin with 4 setae. Gnathopod 2, propod
longer than broad; palm straight or weakly convex and
armed with 1 1 spine teeth in double row ; defining angle
with 2 long spine teeth on outside, 2 shorter spine teeth
on inside; posterior margin shorter than palm, with 1 or
2 setae; few superior medial setae doubly and singly
inserted; 2 singly inserted inferior medial setae; dactyl
nail moderately long; coxa deeper than broad; margin
with 3 setae.

Pereopod 3, coxal plate deeper than broad, margin
with 4 setae. Pereopod 4, coxal plate relatively broad
and deep, reaching about 55 percent length of basis,
margin with 9 setae. Pereopod 6 little longer than
pereopod 7, about 54 percent length of body, 20 percent
longerthanpereopod5. Pereopods 5-7, bases broadest
proximally; posterior margins of 5 and 7 strongly
convex and expanded, that of 6 less so; distoposterior
lobes distinct and broadly rounded; anterior and poste-

rior margins with variable number of spines and setae;

sepients 4, 5 and 6 of pereopods 5-7 with longish
spines; dactyls of pereopods 5-7 relatively slender and
elongate, that of pereopod 6 about 35 -40 percent
length (rf corresponding propod. Coxal gills present on
pereopods 2-6, absent from 7. Simple, lateral sternal
gills present on pereonites 6 and 7. Brood plates
slightly narrowing distally.

Pleonal plates: posterior margins of plates 1 - 3
weakly convex, with 1 setule near distojX)sterior cor-
ner; distoposterior comers rounded and indistinct; ven-
tral margin of plate 1 with 1 spine, that of plates 2 and
3 with 2 spines each. Uronites free. Uropod 1: inner
ramus little longer than outer ramus, 10 percent longer
than length ofpeduncle, with 8 spines, 1 elongate; outer
ramus with 7 spines, 3 elongate; peduncle with 4
spines. Uropod 2: rami heavily spinose apically; inner
ramus longer than outer ramus, subequal in length to
peduncle, with 1 1 spines; outer ramus with 5-6 robust
spines; peduncle with 2 spines. Uropod 3: peduncle
without setae, ramus only 30 percent length of pedun-
cle, with 2 apical spines.

Telson subquadrate, about as long as broad, apical
margin with tiny median notch between spine clusters,
beanng 10 relatively long spines, some as long as
telson.

Male. Differing from female as follows: gnathopod
propods narrower, palm of propod 2 shorter in relation
to posterior margin and with fewer teeth on palmar
margin. Uropod 1 : peduncular process about 50 per-
cent length of outer ramus, tapering to blunt point
distally, upper margin minutely serrate toward apex;
inner ramus with 7-8 spines; peduncle with 5 spines.
Uropod 2: inner ramus with 9-12 spines, peduncle with
3 spines. Telson, with 13-14 long apical spines.

Distribution and ecology. This species is known only
fromits type-locality, whereaccordingto Peter Koetsier
(pers. comm.), the material was collected with a Surber
bottom sampler.

Etymology. This species is named for the state of
Idaho.

Stygobromus glaciaUs, new species
(Fig. 49)

Material examined. MONTANA. Glacier Co.: Zoo Cave,
about 16 km W of Babb Glacier Nat. Park, HOLOTYPE 9
(USNM 1000058), 1 9 paratype, J. Chester, 27 Aug. 1977;
Flathead Co.: Algal Cave. 2 99. J. Chester, 25 Aug. 1977;

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 119

Fig. 49. Stygobromus glacialisj n. sp. Female (6.5 mm). Glacier County, Montana.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 120

4 99, P- Hendricks, 27 Sept. 1999; Logan Creek springhead,
Logan Pass, 100(+) specimens, (9, CT juv.) (MTEC), J.
Giersch, 20 Aug. 2000; stream in an unn 2 imed cave on Trail
Creek in Flathead National Forest, ca. 12.8 km S of Canadian
border, 1 9, J- A. Stanford, 25 Nov. 1980.

Diagnosis. A relatively small to medium-sized
cavemicolous species, distinguished from all other
species of Stygobromus by the possession of 3 pairs of
lateral sternal gills on pereonites 5, 6 and 7; and further
distinguished by slender bases of pereopods 5-7, and
several setules each on posterior margins of pleonal
plates. Largest 9, 7.0 mm; largest cf , 4.5 mm.

Female. Antenna 1: 40 percent length of body, 40
percent longer than antenna 2; primaiy flagellum with
13 segments. Antenna 2: flagellum with 6 segments.

Mandibles subequal: spine row with 7-8 plumose
spines; palp segment 2 with row of 9 long setae on inner
margin; palp segment 3 bearing 2 B setae, 9 D setae, and
4 setae E, lacking both A and C setae. Inner lobes of
lower lip vestigial or absent. Maxilla 1: inner plate with
7 apical, plumose setae; palp with 8 stiff setae apic-
ally. Maxilla 2: inner plate with oblique row of 7
plumose setae on inner margin. Maxilliped: inner plate
with 1 bladelike spine, 5 plumose spines, 2 naked setae
apically, and 3 plumose spines on inner margin; outer
plate with setae on inner margin and 3 lightly plumose
setae on or near apex.

Gnathopod 1: propod smaller than that of gnathopod
2; palm straight and almost 2 times longer than poste-
rior margin, armed with 18 spine teeth in double row;
defining angle with 2 spine teeth on outside, 1 shorter
one on inside; posterior margin lacking setae; 4 doubly
inserted superior medial setae; 5 inferior medial setae;
dactyl nail long and slender; coxa approximately 2
times broader than deep, margin with 2 setae. Gnatho-
pod 2: propod longer and broader than first pereopod,
longer than broad; palm straight or little convex medi-
ally, armed with 20 spine teeth in double row; defining
angle with 1 spine tooth on outside, 3 shorter spine teeth
on inside; posterior margin 33 percent length of palm,
with 2 sets of doubly inserted setae; 7-8 superior
medial setae doubly and triply inserted, about 5 singly
inserted inferior medial setae; coxa broader than deep,
margin with 3 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 1 setae. Pereopod 4; coxal plate relatively broad
and about as broad as deep, reaching about 45 percent
length of basis, margin with 7 setae. Pereopod 6 little
longer than pereopod 7, about 50 percent length of
body, 25 percent longer than pereopod 5. Pereopods 5-

7: bases little broader proximally than distally; poste-

riormarginsslightlyconvex;distoposteriorlobespoorly

developed and indistinct; anterior and posterior mar-
gins with 2-4 spines and setae; segments 4, 5 and 6 of
pereopods5-7 with slender sjxnes ; dactyls of pereopods
5-7 relatively slender and elongate, that of pereopod 6
approximately 30 percent length of corresponding
propod. Coxal gills on pereopods 2 - 6, absent from 7.
Three pairs of long, slender simple lateral sternal gills
present on pereonites 5-7, longest on 6 and 7. Brood
plates rather narrow, not setose on specimen illustrated.

Pleonal plates: posterior margins of plates 1-3
convex, each with 2 cx*3 setules; distoposterior comers
rounded and indistinct; ventral margins of plates 1-3
lacking spines. Uronites free. Uropod 1: inner ramus
little longer than outer ramus, about 80 percent length
of peduncle, with 7 spines; outer ramus with 7 spines;
peduncle with 6 spines. Uropod 2; inner ramus longer
than outer ramus, little longer than peduncle, with 5
spines; outer ramus with 7 spines; peduncle with 3
spines. Uropod 3: peduncle usually bearing 1-2 small
setae; ramus approximately 20 percent length of pe-
duncle, with 3 apical spines.

Telson longer than width, apical margin with small
median notch, bearing 12 relatively long spines.

Distribution and ecology. The species is recorded
from Zoo Cave in Glacier County and Algal Cave (=
West Tunnel Cave), an unnamed cave on Trail Creek
and Logan Creek springhead in adjoinning Flathead
County. Algal and Zoo caves and the spring are in
Glacier National Park, whereas the unnamed cave is
just outside the park’s western boundary. According to
Paul Hendricks (pers. comm.), who made the most
recent collection of this species from Algal Cave in
1999, the specimens were taken from a series of three
silt-bottom pools with a slight flow between them. He
also reported seeing 30-40 amphlpods that were not
collected. Stygobiont isopods, Salmasellus stegona-
thrix Bowman also inhabit the pools (Lewis 2001 ). The
2 ovigerous females measured 5.8 mm long and each
had 2 embryos in the brood pouch, although 4 embryos
were lost in the vial. The other females were 5.0 mm
long and had setose brood plates but were not ovigerous.
The temperature of the Algal Cave stream pools was
6.5°C.

A very recent (August 2000) collection of this spe-
cies was made by Joe Giersch, who obtained a large
sample of more than 100 individuals form the
Springhead of Logan Creek in Logan Pass at an elevat-
ion of 2159 m. According to J. Giersch (pers. comm.)

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 121

the species is abundant in gravels in the substrate of the
spring flow.

Etymology. The epithet glacialis is from the Latin,
meaning “ice” or ‘Trozen” in reference to the occur-
rence of this species in Glacier National Park.

Stygobromus secundus Bousfield & Holsinger, 1981

Stygobromus secundus Bousfield and Holsinger, 1981:
1827-1830, figs. 1 [type-locality: a spring 24 km south-
west of Rocky Mountain House, Alberta, Canada].

Diagnosis. A small to medium-sized groundwater spe-
cies,distinguishedfromallotherspeciesofStygobromus
bypresenceofsimple,medianstemalgillsonpereonites
2 and 3 and simple, paired, lateral sternal gills on
pereonites 4, 5, 6 and 7; and also distinguished by
comparatively large ramus of uropod 3 (but less than
half length of peduncle); and absence of a peduncular
process on uropod 1 of the male. Largest 9, 5.2 mm;
largest d* , 4.0 mm.

Distribution and ecology. This species is known only
from its type-locality (see Bousfield and Holsinger
1981).

Stygobromus obscurus Holsinger, 1974

Stygobromus obscurus Holsinger, 1974: 55-58, figs.
34-35 [type-locality: well at Victor Crossing, Ravalli
Co., Montana].

Diagnosis. A small to medium-sized subterranean spe-
cies, apparently unrelated to other known species of
Stygobromus in western North America and distin-
guished by: relatively slender propods of gnathopods;
shallow coxal plates of gnathopod 2 and pereopcxls 3
and 4; narrow bases of pereopods 5-7 which lack
distoposterior lobes; 2 pairs of simple, lateral sternal
gills on pereonites 6 and 7; absence of ventral spines on
pleonal plates; heavily spinose uropods and telson; and
relatively long spines on uropods 3 and telson. Largest
9, 7.0 mm; d unknown.

Distribution and ecology. This species is known only
from its type-locality, collected along with two other
species of Stygobromus (see Holsinger 1974).

Stygobromus montanensis Holsinger, 1974

Stygobromus montanensis Holsinger, 1974: 52-55,

figs. 32-33 [type-locality: well at Victor Crossing,
Ravalli Co., Montana].

Diagnosis. A small to medium-sized subterranean spe-
cies, distinguished by possession of few setae on pos-
terior margin of gnathopod propod 1 and presence of 2
pairs of tiny, simple lateral sternal gills on pereonites 6
and 7; 4 relatively long spines on ramus uropod 3
(Holsinger 1974). Largest 9. 5.5 mm; d unknown.

Distribution and ecology. This species is known only
from its type-locality, where it has been collected with
twoother species of 5tygc?^romw5(see Holsinger 1974).

Stygobromus curroae, new species
(Figs. 50,51)

Material examined. NEW MEXICO. Taos Co.: ground-
water discharge into Gallina Creek, Sangro de Cristo Moun-
tains. near Taos, HOLOTYPE 9 (USNM 1000054), 2 99
paratypes, J. Curro, 23 Nov. 1994 and 1 d, 5 July 1994.

Diagnosis. A relatively small goundwater species,
with some resemblance to S. holsingeri and S. ariz-
onensis but easily distinguished as follows: maxillae 1
and 2 with fewer setae on inner plates; palm of propod
of gnathopod 1 with more spine teeth; coxal plates 3
and 4 with more setae; relative long distally expanded
telson, with 18 - 20 apical spines and rather deep notch;
uropods 1-2 with more long and thick spines on rami;
and tiny, simple, lateral sternal gills on pereonites 6 and
7. Largest 9, 4.8 mm; largest d, 4.5 mm.

Female. Antenna 1: 55 percent length of body, 40
percent longer than antenna 2; primary flagellum with
12 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 6 plumose
spines; palp segment 2 with row of 6 setae on inner
margin; palp segment 2 with 6 long setae in inner
margin; palp segment 3 bearing 1 B seta, 5 C setae, 9 D
setae, and 3 E setae, lacking A setae. Inner lobes of
lower lip absent. Maxilla 1: inner plate with 6 apical,
plumose setae; palp with 6 stiff setae or slender spines
apically. Maxilla 2: inner plate with oblique row of 6
plumose setae on inner margin. Maxilliped: inner plate
with 3 bladelike spines, 2 plumose spines, 1 naked seta
apically, and 1 plumose spine on inner margin; outer
plate with setae on inner margin, and 1 lightly plumose
seta and 1 bladelike spine on or near apex.

Gnathopod 1: propod little smaller than that of
gnathopod 2; palm straight or slightly convex, approxi-
mately 2 times longer than posterior margin, armed

AMPMPAaHCA VOL. 3 NO. 2. NOV. 15, 2001 122

Fig. 50. Stygobromus curroae, n. sp. Female (4.4 mm). Taos County, New Mexico.

AMPHIPAaHCA VOL. 3 NO. 2. NOV. 15, 2001 123

Fig. 51. Stygobromus curroae, n. sp. Female (4.4 mm); male (4.5 mm). Taos Co., New Mexico.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 124

with 16 spine teeth in double row; defining angle with
1 long spine tooth on outside, 2 shorter ones on inside;
posterior margin without setae; 1 superior medial setae;
2-3 inferior medial setae; dactyl nail relatively short;
coxa little broader than deep, margin with 5 setae.
Gnathopod 2: propod deeper than broad; palm straight
or slightly convex and armed with 15-16 spine teeth in
double row; defining angle with 1 long spine tooth on
outside, 2 shorter spine teeth on inside; posterior mar-
gin about 50 percent length of palm, with 1 or 2 setae;
4-5 doubly inserted superior medial setae; 4 inferior
medial setae ; coxa li tde broader than deep, margin with
5 setae.

Pereopod 3: coxal plate deeper than broad, margin
with 7 setae. Pereopod 4: coxal plate relatively broad
and deep, reaching about 50 percent length of basis,
broadly convex margin with 10 setae. Pereopod 6 little
longer than pereopod 7, about 77 percent length of
body, 42 percent longer than pereopod 5. Pereopods 5-
7: bases broader proximally than distally; posterior
margins convex; distoposterior lobes well developed;
anterior and posterior margins with variable number of
spines and setae; segments 4, 5 and 6 of pereopods 5-
7 with longish, slender spines; dactyls of pereopods 5-
7 relatively slender and short, that of pereopod 6
approximately 25 percent length of corresponding
propod. Coxal gills present on pereopods 2-6, absent
from pereopod 7. Two pairs of tiny, simple lateral
sternal gills on pereonites 6 and 7.Brood plates little
expanded distally, but not setose in material examined.

Pleonal plates: posterior margin of plates 1-3
convex, each with 1 setule near distoposterior comer;
distoposterior comers rounded, indistinct; ventral mar-
gin of plate 2 with 3 spines, that of plate 3 with 1 spine.
Uronites free. Uropxxl 1: inner ramus spinose, little
longer than outer ramus, about 95 percent length of
peduncle, with 15 mostly robust spines; outer ramus
with 10 spines; peduncle with 9 spines. Uropod 2: inner
ramus longer than outer ramus, heavily spinose, longer
than peduncle, with approximately 20 spines; outer
ramus with 9 spines, 1 very large; peduncle with 4
spines. Uropod 3: peduncle usually bearing 1 or 2 small
setae; ramus approximately 55 percent length of pe-
duncle, with 2 apical spines.

Telson 25-30 percent longer than broad, gently
expanded distally; apical margin typically with rela-
tively deep V-shaped median notch between spine
clusters, bearing 19-20 relatively long spines.

Male. Differing from female as follows: gnathopod
propods slightly broader, palm of propod 2 shorter in
comparison with posterior margin, with more teeth on

palmar margin. Uropod 1: peduncular process sharply
pointed distally, with lightly serrate upper margin,
approximately 20 percent length of outer ramus; inner
ramus with 14-15 spines; peduncle with 7 spines.
Uropod2: innerramus withapproximately22-25spines,
peduncle with 4-5 spines. Telson with 18-21 apical
spines.

Distribution and ecology. This species is known only
from its type locality, where it was sampled from the
hyporheic zone with a PVC pipe and pumpfrom adepth
of 30 cm (J. Curro, pers. comm.).

Etymology. This species is named in honor of its
collector, Julia Curro.

Stygobromus simplex^ new species
(Fig. 52)

Material examined. COLORADO. Rio Blanco Co.: Spring
Cave, southeast of Meeker, HOLOTYPE 9 (on slide mounts
inpart)(USNM 1 00007 1 ), M. Tate and K. Tate, 4 Oct., 1995.

Diagnosis. A medium-sized cavemi colons species that
resembles 5. kolsingeri in structure of gnathopods and
uropods 2-3, but is distinguished from that species as
follows: maxilla 1-2 with fewer setae on inner plate;
maxilliped with bladelike spines on inner plate;
gnathopods 1-2 with more teeth on palmar margin;
bases of pereopods 3-7 with fewer spines on posterior
margins; pleonal plates 2-3 with more ventral margin
spines; and presence of relatively long, lateral sternal
gills. Largest 9, 5.0 mm, cf, unknown.

Female. Antenna 1: 44 percent length of body, 34
percent longer than antenna 2; primary flagellum with
10 segments. Antenna 2: flagellum with 5 segments.

Mandibles subequal: spine row with 4-6 plumose
spines; palp segment 2 with row of 6 rather long setae
on inner margin; palp segment 3 with 8 D setae, 4 E
setae, without A, B and C setae. Inner lobes of lower lip
vestigial or absent. Maxilla 1 : inner plate with 6 apical,
plumose setae; palp with 6 stiff setae or slender spines
apically. Maxilla 2: inner plate with oblique row of 6
plumose setae on inner margin. Maxilliped: inner plate
with 2 bladelike spines, 3 plumose spines, 2 naked setae
apically, and 2 plumose setae on inner margin; outer
plate with setae on inner margin and apex.

Gnathopod 1: propod little shorter than that of
gnathopod 2; palm weakly convex, about 2 times
longer than posterior margin, armed with 21-22 spine
teeth in double row ; defining angle with 2 spine teeth on

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001

125

Fig. 52. Stygobromus simplex^

n.sp. Female (5.0 nm). Rio Blanco County, Colorado.

AMHflPAaFICA VOL. 3 NO. 2. NOV. 15, 2001 126

Fig. S3. Stygobromuswardi,n.sp. Female (3.7 mm). Eagle County, Colorado.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001

127

AMPMPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 128

outside, 2 shorter ones on inside; posterior margin
without setae; 1 or 2 superior medial setae;8* lOsingly
inserted inferior medial setae; dactyl nail rather long
and sharply pointed; coxa deeper than broad, margin
with 1 seta. Gnathopod 2: propod about 2 times longer
than broad; palm slightly convex and armed with 19-20
spine teeth in double row; defining angle with 1 long
spine tooth on outside, 2 shorter spine teeth on inside;
posterior margin about 50 percent length of palm, with
1 set of triply-inserted setae; 3 or 4 superior medial
setae; 3 singly-inserted inferior medial setae; coxa
about as broad as deep, margin with 4 setae.

Pereopod 3; coxal plate deeper than broad, margin
with 2 setae. Pereopod 4: coxal plate relatively broad
and deep, reaching about 40 percent length of basis,
margin with 4 setae. Pereopod 6 little longer than
pereopod 7, about 76 percent length of body, 20 percent
longer than pereopod 5. Pereopods 5-7; bases rela-
tively narrow, but little broader proximally than dist-
ally; posterior margins weakly convex, not expanded;
distoposterior lobes poorly developed, almost indis-
tinct; anterior and posterior margins with variable
number of spines and setae; segments 4, 5 and 6 of
pereopods 5-7 with longish spines; dactyls of pereo-
pods ^7 relatively slender and elongate, that of pereo-
pod 6 approximately 35-40 percent length of corre-
sponding propod. Coxal gills present on pereopods 2-
6, absent from 7. Relatively long, simple, lateral ster-
nal gills present on pereonites 6 and 7. Brood plates
little expanded distally.

Pleonal plates: posterior margins of plate 1-3
weakly convex, with 1 setule near distoposterior cor-
ner, distoposterior comers rounded and indistinct; ven-
tral margins of plates 2 and 3 with 2 spines each.
Uronitesfree. Uropod 1: inner ramus little longer than
outer ramus, about 60 percent length of peduncle, with
9 spines; outer ramus with 6 spines; peduncle with 7
spines. Uropod 2; inner ramus longer than outer ramus,
longer than peduncle, with 9-10 spines; outer ramus
with 5-6 spines; peduncle with 3 spines. Uropod 3:
peduncle usually bearing 2 small setae; ramus approxi-
mately 35 percent length of peduncle, with 2-3 apical
spines.

Telson little longer than broad, gently tapered
distally; apical margin with tiny median notch between
spine clusters, bearing 10-12 spines.

Distribution and ecology. This species is known only
from a single female. The type locality is one of the
longest caves in Colorado (Parris 1973) and contains a
stream and lake.

Etymology. The epithet simplex is from the Latin,
meaning “simple” or “uncomplicated,” in reference to
the simplicity of a species based on a single specimen.

Stygobromus wardi, new species
(Figs. 53,54)

Material examined. COLORADO. Eagle Co.: Fulford
Cave, S. of Eagle, HOLOTYPE 9 (USNM 1000076), 3 Cfd’
and 4 99 paratypes, 1 juv., J. V. Ward and J. R. Holsinger,

6 July 1980; Blue Butt Cave, 6 99, J- V. Ward and J. R.
Holsinger, 6-7 July 1980.

Diagnosis. A small cavemicolous species, morpho-
logically similar to S. holsingeri and 5. coloradensis in
structure of gnathopods and uropod 1-2 but distin-
guished from these species as follows: antenna 1 about
2 times as long as antenna 2; maxillae 1-2 with more
setae on inner plate; coxal plates 3-4 broader than deep;
bases of pereopods 6-7 with fewer spines on posterior
margin; pleonal plates 2-3 with fewer spines on ventral
margin; presence of lateral sternal gills (processes);
telson broader than deep and with longer apical spines.
Largest 9, 5.0 mm; largest cJ* , 3.5 mm.

Female. Antenna 1: 46 percent length of body, ap-
proximately 2 times longer than antenna 2; primary
flagellum with 18 segments. Antenna2: flagellum with
5 segments.

Mandibles subequal: spine row with 5-6 plumose
spines; palp segment 2 with 3 long setae on inner
margin; palp segment 3 bearing 1 B seta, 6 D setae, 3
E setae, lacking A and C setae Inner lobes of lower lip
small to vestigial. Maxilla 1: inner plate with 7 apical,
plumose setae; palp with 4 stiff setae subapically.
Maxilla 2: inner plate with oblique row of 6 plumose
setae on inner margin. Maxilliped: inner plate with 1
bladelike spine, 3 plumose spines, 3-4 naked setae
apically, and 1 plumose spine on inner margin; outer
plate with 7 or 8 setae on inner margin, 4 setae on or near
apex.

Gnathopod 1 ; propod subequal in size to gnathopod
2; palm straight, approximately same length as poste-
rior margin, armed with 7 spine teeth in double row;
defining angle with 2 spine teeth on outside, 2 shorter
ones on inside; posterior margin lacking setae; without
superior medial setae; 1 -3 inferior medial setae;dactyl
nail rather long; coxa broader than deep, margin with 1
seta. Gnathopod 2: propod longer than broad; palm
straight or slightly concave and armed with 8 spine
teeth in double row ; defining angle with 2 spine teeth of

AMPMPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 129

unequal length on outside, 2 shorter spine teeth on
inside; posterior margin little shorter than palm, with 1
set of doubly or triply inserted setae; 3 superior medial
setae, one rather long; 2 inferior medial setae; coxa
deeper than broad, margin with 1 seta. Pereopod 3:
coxal plate deeper than broad, margin with 2-3 setae.
Pereopod 4: coxal plate subquadrate, about as broad as
deep, reaching about30 percent length of basis, margin
with 3 setae. Pereopod 6 little longer than pereopod 7,
about 65 percent length of body, 33 percent longer than
pereopod 5. Pereopods 5-7: bases little broader proxi-
mallythanthedistally;posteriormarginsnearlystraight
or slightly convex; distoposterior lobes indistinct; an-
terior and posterior margins with fewer (2-4) spines
and setae; segments 4, 5 and 6 of pereopods 5-7 with
slender spines; dactyls of pereopods S7 relatively
slender and elongate, that of pereopod 6 typically up to
40-45 percent length of corresponding propod. Coxal
gills present on pereopods 2-6, absent from 7; Two
pairs of simple lateral sternal gills/processes on
pereonites 6 and 7. Brood plates slightly expanded
distally.

Pleonal plates: posterior margin of plate 1 convex,
with 1 setule near distoposterior comer, that of 2 and 3
less convex, each with 1 setule near distoposterior
comer; distoposterior comers rounded; ventral mar-
gins of plates 1 and 2 without spines, that of plate 3 with
2 spines. Uronites free. Uropod 1: inner ramus little
longer than outer ramus, about 80 percent length of
peduncle, with 5 spines; outer ramus with 5 spines;
peduncle with 4 spines. Uropod 2: inner ramus longer
than outer ramus, little longer than peduncle, with 7
spines; outer ramus with 5 spines; peduncle with 2
spines. Uropod 3: peduncle without setae; ramus ap-
proximately 30 percent length of peduncle, with 3
apical spines.

Telson little broader than long, lateral margins
slightly convex; apical margin with shallow median
notch between spine clusters, bearing 8 mostly long
spines.

Male^ Differing from female as follows: gnathopod
propods proportionately shorter, palm of propod 2
longer in relation to posterior margin and with fewer
teeth. Uropod 1: peduncular process present, reaching
about 25 percent length of outer ramus, pointed di stally,
upper margin minutely serrate; inner ramus with 5
spines; peduncle with 5 spines. Uropod 2: inner ramus
with 8 spines, peduncle with 3-4 spines. Telson with 6-
9 apical spines.

Distnbution and ecology. This species is known only
from small streams in two caves in Eagle Co., Colo-

rado, at elevations of approximately 2902 m and 3,048
m (above sea level). It was collected from gravels and
assorted rubble in a rapid, high gradient stream in
Fulford Cave, one of the larger and better known caves
in the state (Parris 1973), where the water temperature
was 4.5°C. In Blue Butt Cave, 6 specimens were col-
lected from the silty substrate of the cave stream by a
combination of Surber bottom sampler and overnight
drift net. A single female (4.0 mm) in the July sample
from Blue Butt Cave had 2 large embryos in the brood
pouch. The water temperature near the entrance to this
cave was 8 °C.

Etymology. This speciesisnamedinhonorof Prof. Dr.
James V. Ward, who helped with the discovery and
collection of this species and has made many other
significant contributions to the study of subterranean
amphipods in Colorado.

Stygobromus species

The following populations were recognized as two
distinct species in this study but are not described
because of the lack of mature specimens or enough
material. These specimens and detailed habitat data
were donated by Richard Casey and Gary Byrtus from
the Alberta Environmental Center in Vegreville, AB,
Canada.

Species No. 1. A single immature male (ca. 4.0
mm), collected by G. Byrtus from a groundwater well
on the grounds of the Glencoe Golf and Country Club,
just west of Calgary, Alberta, Canada. This species has
small, simple lateral sternal gills; telson with small
apical notch; uropod 3 with 2 apical spines; and
gnathopod 2 larger than gnathopod 1 . Its overall mor-
phology is rather typical for species of the hubbsi group
with exception of the sternal gills. This specimen was
collected along with a second undescribed species,
which is listed immediately below (see species no. 2).

Species No. 2. Eleven tiny specimens from: a)
groundwater well on grounds of Glencoe Golf and
Country Club, just west of Calgary (3 specimens col-
lected by G. Byrtus, 19 Oct. 1993); b) groundwater
wells on grounds of Kananaskis Country Golf Course,
60 km west of Calgary (7 apparently immature speci-
mens in 5 samples collected by G. Byrtus from 3 June
1992 to 1 June 1993); c) groundwater well on grounds
of Canmore Golf and Curling Club, about 70 km west
of Calgary (1 tiny specimen collected by G. Byrtus, 4
May 1993).

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 130

PHYLOGENY

A phylogenetic analysis was performed on the 32
morphological characters in Tables 1 and 2, utilizing
PAUP3.1 and MacClade3.01 (MaddisonandMaddi-
son 1993) on a Macintosh computer (OS-7). The
choice of characters used in the analysis is based on
descriptions from the literature and illustrations. Most
characters are binary, but three are multi state. Each
series of runs was started with the Heuristic Search
option. For the first run, only minimum trees were kept
by Random Stepwise Addition, TBR Branch Swap-
ping, and MULPARS options activated. For subse-
quent runs, all trees in memory from the preceding runs
were used, activating MULPARS option and keeping
all trees that were as short as, or one step shorter than,
those loaded into memory. These steps were repeated
until no shorter trees could be found. From the result-
ing trees, strict and majority rule consensus trees were
calculated.

In the first-run, characters were treated as unord-
ered and unweighted (randomly reversible states and
equally weighted), assuming that, for example, any
number of spines on a certain appendage could evolve
directly. In the second run, characters were ordered and
partly weighted. Multistate characters were assumed to
have evolved in reversible linear transformation series.
Some characters were considered more important and
were weighted accordingly (see Table 1). In the fol-
lowing discussion, the alternative trees resulting from
the analyses are referred to as "unordered-unweighted"
and “ordercd-weighted.” Although the two trees are
not identical and have different lengths, there are many
similarities between their topologies.

Polymorphic characters, for example, varying
numbers of long setae on the inner plate of maxilla 1,
were coded according to the scaled method, but not
strictly (see Wiens 1995) because the presence of four
or five long setae on the inner plate of maxilla 1 could
not be treated as two states. In most cases, it would have
been more satisfactory to code polymorphic characters
in relation to their frequencies. If the majority of the
specimens had four long setae, thenfour setae would be
treated as a coded state. However, such information is
not always available because some species descrip-
tions are based on an inadequate number of specimens.
In this case relevant information of related species is
used as a reference.

In some of the species, the occurrence of a charac-
ter showed a variable or transitional state, for example,
the number of apical and subapical spines on the inner
ramus of uropod 1 and uropod 2 (e.g., character 25).

These variations were treated as separate morphologi-
cal conditions. Another problem encountered during
the analysis was the pronounced degree of interspecific
variation. For example, the shape of propod palm of
gnathopods 1 and 2 is straight in species A, concave in
species B and convex in species C. Two different
approaches were used to deal with it. For the first,
variable characters were coded as question marks if
their morphological/evolutionary significance was not
clear. Alternatively, they were coded according to the
majority of their frequency occurrence in all species
(Wiens 1995).

In our analysis, we chose representative species of
three non-stygobromid genera in the family
Crangonyctidae as outgroups, including Synurella
ambulans (central Europe), Crangonyx forbesi (east-
central North America), and Bactrurus brachycaudus
(east-central North America). Among the three, only
B. brachycaudus is a stygobite; the other two species
are stygophiles, inasmuch as they are recorded from
both surface and subterranean groundwater habitats
and are not completely troglomorphic. All three of the
outgroup genera share important morphological char-
acters with Stygobromus, and their probable
phylogenetic relationship to this genus and with other
crangonyctids is discussed and illustrated elsewhere
(Holsinger 1994a, 1994b).

Assumptions about morphological adaptations of
amphipods to subterranean groundwater environments
have been discussed in the literature (Stock 1981;
Notenboom 1991; Holsinger 1994; Koenemann et al.
1998). Troglomorphisms, such as reduction and/or
loss of eyes and pigment and attenuation of certain
appendages, which are often treated as apomorphies,
are attributed to hypogean adaptation. Alternatively,
each corresponding plesiomorphic character state
(coded as state 0 in our matrix), may be the condition
found in putative surface (epigean) ancestors or rela-
tives of the ancestors. The large number of homoplasies
found in amphipod crustaceans tends to severely limit
the number of usable or reliable characters available for
a cladistic analysis, as well as to impede the choice of
effective outgroups. In light of this, our choice of
outgroups is probably reasonable, inasmuch as we
selected representatives of three of the six extant gen-
era in the family Crangonyctidae, and only one of
which is fully adapted morphologically to subterran-
ean waters.

Because the analysis resulted in a relatively large
number of cladograms, many of which were very
similar, we present two representative 50% majority-

AMPMPACinCA VOL. 3 NO. 2. NOV. 15, 2001 131

cE

5. ambulans

C. forbcsi

B. bnichycauduf^

impcrlalls

rudolphi

grahaml

wengerorum

harai

gallawayae

Utus

gradyl

hyporheicus

elllotti

hubbil

interstitialls

utBhensIs

obicurtu

Mituaris

tdahocnaU

canadensis

coloradensis

rallus

blinni

trinus

herbsti

jemezensis

bovltoni

duplus

tritus

puteanus

pennaki

cowani

urospinatus

fontinalis

limbus

curroae

mystic us

montanensis

glacialis

secundtts

lanensls

quatsinensis

simplex

arizonensis

holsingerj

wardi

myersae

sheldoni

lacicotus

wahkeenensis

puteaiis

cherylac

sicrrensis

oregonensis

tahoensis

mackenzici

Fig. 55. Maj ority-rule consensus tree of the first-run using three non-Stygobromus species as outgroups;
characters not weighted and not ordered (Cl: 0.16; RI: 0.50; tree length: 232).

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15. 2001 132

S, ambuliB*

C.forbftit

B. brachycaudtu

wmrdi

linplex

corroa*

nonUBcasIs

obscuruf

sccuBdus

Klaclalis

ctBade&sia

Idabocnsb

ImptrUlii

rudolphl

Srahami

wtBscrorum

baral

gallawayat

lalui

gradyi

clliom

hyporbeieui

mystleua

bubbil

orc^oaensls

maekanzlcl

Cabocatli

eowiBl

chary laa

myarsaa

sbaldoni

Uckolus

puUalls

ficrrcnsls

lanculs

wabktaacnsis

quatalBCBSii

arizOACosts

boUlogcri

bUBBl

coloradcBiif

vrosplaatui

pcaaikl

puCaanui

InCcrstUUlis

utakcnsii

duplua

bottltoni

trltua

triaua

htrbaU

Jamezcnats

foBtlnalis

Umbua

Fig. 56. Majority-rule consensus tree of the second-run using three non-Stygobromus species as
outgroups; characters partially weighted and ordered (Cl: 0.16; RI: 0.51; tree length: 229).

AMWflPACIHCA VOL. 3 NO. 2. NOV. 15, 2001

133

Table 1: Ust of characters and character states used in the phylonegetic analysis for species of

otherwise indicated, characters were onlered and

weighted by 1 m the second ran.

1. Ratio of length of antenna 1 to that of body:
state 0: 50% or less; state 1; more than 50%

broad; state 1; broader than deep.

Maxillae

2. Plumose setae on inner plate of maxilla 1:

state 0: more than 10; state 1: equal to or less than 10

3. Plumose setae in oblique row on inner plate of maxilla 2:

state 0; more than 1 1 ; state 1: equal to or less than 1 1

Mandibles

4. -B" setae on segment 3 of mandibular palp:

state 0: present; state 1: absent

5. Spines in spine row of mandibles:

state 0: more than 8; state 1: equal to or less than 8

Gnathopods 1 and 2

6. Shape of propod palm of gnathopod 1:

state 0: straight or sinuate; state 1 : concave; state 2: convex
The palm shape of the propods of gnathopods 1 and 2
vanes in species of Stygobromus. The terms straight, con-
cave and convex refer to the general shape of the piopod
palm. However, the palm of gnathopod 2 may be variably
straight or sinuate. Because the distinction between the two
is unclear, we treat "straight or sinuate" as a single state,

7. Spine teeth on propod palm of gnathopod 1:

state 0: equal to or less than 28; state 1: more than 28
In second-run: ordered and weighted 2

8. Shape of propod palm of gnathopod 2:

state 0: straight or sinuate; state 1: concave; state 2'
convex.

In second-run: ordered and weighted 2.

9: Ratioof length of posterior margin of propod of gnathopod
2 to length of propod palm:
state 0: more than 30%; state 1: 30% or less,

10. Sets of setae, posterior margin of propod, gnathopod 2-
state 0: present; state 1: absent

14. Setae on coxal margin of pereopods 3-4:

state 0: equal to or less than 6; state 1: more than 6.

15. Coxa of pereopods 4:

state 0: deeper than or approximately as deep as broad;
state 1: broader than deep.

16. Pereopod 6 longer than 7:
state 0: yes; state 1: no.

17. Shape of bases of pereopods 6 and 7:
state 0: normal; state 1: broad

In second-run: ordered and weighted 2.

The term “normal ” refers to the ratio of breadth to depth
(or length) of the bases seen in most species of Stygobromus.
However,somespecies,e.g.,5.i/M/i/Mj, S. rallusandS. latus,
have broad bases in comparison with the normal shape .

18. Distoposterior lobe of pereopod 6 and 7:
state 0: present; state 1: absent or indistinct

Sternal Gills/Processes

19. Sternal gill(process):

state 0: present; state 1: absent
In second-runs ordered and weighted 8.

This character plays an important role in separating the
species of Stygobromus into hubbsi group (without sternal
gills on pereonites) and Xioxi-hubbsi species group (with
sternal gills on pereonites, typically 6 and 7).

Pleonal plates

20. Setules on posterior margins of pleonal plates 2 and 3*

state 0: 1; state 1: 2 or more.

21. Spines on ventral margin of pleonal plate 3:
state 0: more than 3; state 1:1-3; state 2: absent
In second-run: ordered and weighted 4.

11. Coxa of gnathopod 2:

state 0: broader than or approximately as broad as deep
state 1: deeper than broad.

12. Superior medial setae on propod of gnathopod 2*
state 0: present; state 1: absent

Pereopods

13. Coxa of pereopod 3:

state 0: deeper than or approximately as deep as

Uropods

22. Some apical spines on inner ramus of uropod 1 nearly as
long as that of peduncle:

state 0: no; state 1: yes

23. Spines on peduncle of uropod 1:

state 0: equal to or less than 12; state 1: more than 12.

24. Length of inner ramus of uropod 1 compared to that of
peduncle:

state 0: equal to or shorter; state 1: longer.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 134

TABLE 1 (CONT'D).

25. Spines on inner ramus of uropod 2:

state 0: equal to or less than 12. state 1; more than 12.

26. Length of inner ramus of uropod 2 compared with that of
peduncle:

state 0; equal to or shorter; state 1: longer.

Telson

29. Apical notch of telson:

state 0: present; state 1: absent or indistinct
In second-run: ordered and weighted 2.

30. Spines on apical margin of telson:
state 0: 8 or less; state 1: more than 8.

27. Ratio of length of ramus of uropod 3 to length of
peduncle:

state 0: 25 % or less; state 1: more than 25 %

In second-run: ordered and weighted 3.

28. Spines on apical margin of uropod 3:
state 0: 1-2; state 1: 3 or more.

rule consensus trees: unordered-unweighted and or-
dered-weighted (Figs. 55, 56). It is important to note
that the consensus trees are very similar in their compo-
sition of some clades. For example, both trees support
an almost identical monophyletic Califomia-Oregon-
Washington clade, consisting of S. imperialis, S. rud-
olphi, S. grahami, S. gallawayae, S. lotus, S. wen-
gerorum, S. harai, S. gradyi, S. elliotti, and S. hubbsi
(Figs. 55, 56). The only difference between the two is
the inclusion of S. mysticus in the second-run tree (Fig.
56). In addition, the two trees also share three identical
doublets, consisting of S. myersae-S. sheldoni, S. inter-

stitialis-S. utahensis, and S. herbsti-S, jemez^nsis.
However, the ordered-weighted consensus tree ap-
pears preferable overall to the alternative unordered-
unweighted consensus tree, inasmuch as it supports the
monophyly of the hubbsi group and shows a stronger
agreement between geographic distribution and termi-
nal taxa (see discussion under Biogeography, below).
In establishing the monophly of the hubbsi group, the
subset of nine western species with sternal gills (proc-
esses) becomes an outgroup to the hubbsi group spe-
cies, which completely lack these structures. Un-
doubtedly, a significant amount of the difference be-
tween the trees can be attributed to the added weight
given to the absence of sternal gills (see Table 1).

HABITAT DIVERSITY

Species of Stygobromus have exploited a wide
variety of subterranean groundwater habitats through-
out the generic range (Holsinger 1974, 1978;Wardand
Holsinger 1981). In western North America, species
have been collected from at least four different (gener-
alized) groundwater habitats and deep lakes (Table 3).
The groundwater habitats include a) caves, b) springs,
c) wells, and d) hypohreic zone. The number of species
collected from the five generalized habitat types in-

31. Apical spines of telson as long as telson:

state 0: no; state 1: yes.

32. Ratio of telson length to width:

state 0: 1.5 times or less; state 1: more than 1.5 times.

eludes; 19 from caves based on 47 records; 29 from
springs based on 53 records; 9 from wells based on 16
records; 9 from hyporheic zones based on 39 records;
and 2 from a surface deep lake (Lake Tahoe), based on
15 records.

The generalized aquatic cave habitat actually en-
compasses three distinctly different, within-cave
macrohabitats, including drip/seep pools, streams, and
phreatic lakes, but these are not differentiatedper se in
Table 3. Springs generally represent the resurgence of
groundwater at the surface, although some may be
seep-like, with intermittent flow contingent on sea-
sonal precipitation and elevation of the groundwater
table. Some seeps, however, are recognized as a
distinct groundwater habitat called the hypo-
telminorheic. Wells are more of a means of access to
groundwater aquifers than a habitat per se. They
usually consist of conventional domestic water wells
but sometimes include PVC pipes driven into the
substrate of a surface stream or into a nearby phreatic
aquifer. The hyporheic habitat is defined as the
underflow of groundwater beneath surface streams and
is accessed either by pumping water though PV C pipes
or sampling directly into the gravel substrate of the
stream. Thus, the difference in habitat for amphipods
collected from a well of the PVC type or directly from
the hyporheic zone is not always clear. A surface deep
lake is not a typical groundwater habitat, although two
species of Stygobromus are recorded from Lake Tahoe
in western North America (Holsinger 1974) and one
from a deep lake in Siberia (Holsinger 1987). It is
assumed that these species probably gain access to lake
waters via groundwater seeps developed in the walls or
floor of the lake basin.

With respect to the number of cave-dwelling spe-
cies, there is a significant contrast between the eastern
and western North American stygobromid fauna. In a

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 135

Table 2. Character data matrix (question marics refer to unknoiMi character states).
Taxon/Character 12 3 4 5 6 7 8 9 10 1112 1314 1516171819 20 2122 232425 26272829 303132

1 B, brackfcauBrns

1 0 0

0 0

2 0 2

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

1 C.forbtii

0 1 0

0

1

2 1 2

0

0

0

0

0

0

0

0

0

0

0

0

0

I

I

0

0

0

0

0

0

1

0

0

3 S. ambukats

1 0 0

0

1

1 1 1

0

0

1

1

0

0

0

0

1

0

0

0

0

0

1

0

1

0

0

0

0

0

0

0

4 imperious

1 1 1

0 0

0 0 0

0

1

1

I

1

1

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

1

0

0

5 cowan/

1 0 0

0

1

0 0 0

0

1

1

1

0

0

1

0

0

0

I

1

1

0

0

0

0

1

0

0

0

1

0

0

6 /rima

1 0 1

0

1

0 0 0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

0

0

1

1

1

1

0

0

7 hyporheicus

1 0 0

1

0

2 1 0

0

0

0

0

1

1

0

0

0

0

1

1

0

0

0

0

0

I

0

1

0

1

0

1

8 mdotphi

1 ! 1

0

1

0 C 1

1

0

0

1

1

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

0

0

9 gaUawayae

1 1 I

0 0

0 0 0

1

0

0

0

0

1

1

0

0

0

1

0

0

0

0

0

0

0

0

1

1

1

0

0

10 cherylae

1 0 0

0

1

2 0 0

1

1

0

0

I

0

1

0

0

0

1

0

1

0

0

0

0

1

0

1

1

1

0

0

1! nyersae

0 0 0

0

1

0 0 0

0

0

1

1

0

1

0

0

0

0

I

0

1

0

0

0

0

1

0

0

1

1

0

1

12 kerbsii

1 0 1

0

1

0 0 0

0

0

0

0

0

I

0

0

0

1

1

0

0

0

1

0

1

0

1

1

0

1

0

0

13 uroapinatus

I 1 1

1

1

0 1 0

0

0

0

0

0

0

0

0

0

1

I

1

1

0

1

1

1

1

0

1

0

1

1

0

14 fimtiHolis

1 0 1

1

1

2 1 0

0

0

0

0

0

0

0

0

0

0

1

?

?

0

0

0

1

1

0

1

0

1

0

0

15 inierstUiaUs

1 0 0

1

I

0 0 0

1

1

0

0

0

0

0

0

0

i

1

0

2

1

0

0

0

0

1

0

0

1

1

0

16 wardi

0 0 0

]

1

0 0 0

0

1

0

1

0

0

0

0

0

1

1

7

?

0

0

0

0

1

0

1

0

1

1

0

17 blitmi

1 0 0

I

1

0 0 0

I

1

0

0

1

0

0

0

0

0

1

?

7

0

0

0

0

0

0

0

0

I

0

0

IS /emexentis

1 0 0

0

1

0 0 0

0

0

0

0

1

1

0

0

0

0

1

0

0

0

0

0

1

1

1

0

0

1

0

0

19 kmensis

1 0 0

I

1

0 0 0

1

0

0

0

0

0

1

0

0

0

1

0

1

0

1

0

0

1

0

I

0

1

0

0

20 wahkeenensis

1 0 0

0

1

0 0 0

0

1

0

0

1

0

1

0

0

0

1

0

I

0

0

0

0

1

0

1

0

1

0

1

21 roBta

1 0 0

1

1

2 0 0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

1

0

0

0

0

1

0

1

1

0

22 IcOus

1 1 1

1

0

0 0 0

0

0

0

1

0

1

1

0

1

0

1

0

0

0

0

0

0

1

0

1

1

1

0

1

23 utahensis

1 0 0

1

0

0 0 0

1

0

0

0

0

0

0

0

0

1

1

0

2

0

0

0

0

0

0

0

0

1

0

0

24 limbus

1 1 1

1

1

0 0 0

0

0

0

0

1

0

0

0

0

0

I

1

0

0

0

0

I

0

0

0

0

1

0

1

25 bo$titom

1 0 0

0

1

1 0 1

1

1

0

1

1

0

0

0

0

1

1

1

1

0

0

0

0

0

1

0

0

I

0

0

26 hubbsi

0 0 0

0

1

0 1 0

1

0

0

0

I

1

0

0

0

0

1

0

0

0

I

0

0

0

0

1

0

1

0

0

27 tritus

1 0 0

0

I

0 0 1

0

0

0

0

1

0

0

0

0

I

I

1

1

0

0

0

0

0

1

1

0

1

0

0

28 puteanui

1 0 0

0

1

0 0 0

0

0

0

0

1

1

0

0

0

0

1

1

1

1

I

0

0

0

1

1

1

1

1

0

29 oregonensis

0 0 0

0

1

0 1 0

0

0

1

0

1

0

1

1

0

0

1

0

0

0

0

0

0

1

0

1

1

1

0

0

30 elliotti

1 1 0

0 0

0 1 1

1

0

0

0

1

1

0

0

0

0

1

0

1

0

0

0

1

0

0

1

1

1

0

1

31 grahami

1 1 !

0

1

0 0 0

I

1

1

0

0

1

0

0

0

0

1

0

0

0

0

0

0

1

0

1

0

1

0

0

32 gratfyi

1 1 1

0 0 0 0 0

1

0

1

I

I

1

1

0

0

0

1

0

1

0

0

0

0

0

0

I

0

1

0

0

33 harai

0 1 1

0 0

0 0 1

1

0

0

1

1

0

0

0

0

0

1

0

0

0

0

0

0

I

0

1

0

I

0

0

34 wengerortm

1 1 1

0

1

2 0 1

1

0

0

I

0

I

0

0

0

0

1

0

0

0

0

0

0

1

0

1

0

1

0

1

35 macketaiei

0 0 0

0

1

0 0 0

0

0

0

1

1

0

1

0

0

0

1

0

0

0

0

0

0

0

0

I

0

0

0

0

36 sierrensis

0 0 0

0 0 0 0 1

1

1

0

0

1

0

0

0

0

0

1

0

1

0

0

0

0

1

I

1

1

1

0

0

37 jAe/(tfonf

1 0 0

0 0

0 0 0

1

0

0

1

0

0

0

0

0

0

1

0

1

0

0

0

0

1

0

0

1

1

0

1

38 loAoensu

0 0 0

0

1

0 0 0

0

0

0

1

1

0

0

0

0

0

1

0

0

0

0

0

0

1

0

1

1

0

0

0

39 lacictybis

0 0 0

0

1

0 0 1

1

1

0

1

0

0

1

0

0

0

1

0

1

0

0

0

0

1

0

1

0

1

0

1

40 arkoiteHsu

1 0 0

0

1

0 0 0

0

1

0

0

0

0

1

0

0

I

1

1

1

0

0

0

0

1

0

0

0

1

0

0

41 pennaki

1 0 0

1

1

0 0 0

0

0

0

1

1

0

0

0

0

0

1

I

1

1

1

0

1

0

0

1

0

1

1

0

42 cotoradensis

0 0 0

1

1

0 0 0

0

0

0

0

1

0

0

0

0

0

I

0

1

0

0

0

0

0

0

0

0

1

1

0

43 holah^ri

1 0 0

1

1

0 0 0

0

I

0

0

0

0

0

0

0

0

1

1

1

0

0

0

0

1

0

0

0

1

1

0

44 putecdis

0 0 0

1

1

0 0 0

1

1

0

0

1

0

1

0

0

0

1

0

1

0

0

0

0

1

0

1

1

1

0

1

45 dt^bts

I 0 0

1

1

2 0 0

1

0

0

0

0

0

0

0

0

1

I

1

1

0

0

0

0

0

1

0

0

I

0

0

46 scdtw^

1 0 0

1

I

0 0 1

1

0

0

0

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

I

0

47 quatsinensis

1 0 0

1

1

0 0 0

0

I

0

1

0

0

1

0

0

0

1

0

1

0

0

0

0

1

0

1

0

1

0

0

48 mysticuM

0 1 0

0

1

0 0 0

0

0

1

0

0

I

0

0

i

0

1

0

1

0

0

0

0

0

0

1

0

1

0

1

49 canadensis

0 0 0

1

1

0 0 0

0

0

0

1

0

0

1

0

0

0

0

0

1

0

0

t

0

0

0

0

1

0

0

0

SO curroae

1 0 0

1

1

0 0 0

0

1

1

0

I

1

1

0

0

0

0

?

?

0

0

0

1

1

0

0

0

1

0

1

51 gkfdaiis

0 0 0

1

1

0 0 0

0

0

0

0

1

1

0

0

0

0

0

0

0

0

0

0

0

1

1

1

0

1

0

0

52 idahoensis

0 0 0

I

1

0 0 0

0

1

0

0

1

0

0

0

0

0

0

0

2

0

0

1

0

0

0

0

1

1

1

0

S3 nundanensts

1 0 0

1

1

0 0 0

0

0

1

0

1

I

0

0

0

0

0

0

1

0

0

0

0

1

0

]

0

I

1

0

54 o6icfirttf

1 0 0

1

1

0 0 0

1

0

1

1

0

0

0

0

0

1

0

0

2

0

1

0

0

0

0

1

0

1

1

0

55 secundus

t 0 0

1

1

0 0 0

0

0

0

0

0

1

0

0

0

0

0

1

1

0

0

0

0

1

0

I

0

1

0

0

56 tinqalex

0 0 0

0

1

2 0 2

0

I

0

0

0

0

0

0

0

1

0

0

1

0

0

0

0

1

0

0

0

1

0

0

Note: Boarunu brachyaatdus, CrmgoiQfxforbtsi and SynwreUa ambuUtns are outgroup species

AMPHIPACIRCA VOL. 3 NO. 2. NOV. 15, 2001 136

study of 48 species of Stygobromusirom eastern North
America, primarily from the Appalachians and Coastal
Plain (Holsinger 1978), 41 species were recorded from
caves, whereas in the present study of 53 species from
western North America (including one from wells in
Wisconsin), only 21 species are reported from caves.
Moreover, in the east approximately 63 percent of the
records of Stygobromus are from caves and approxi-
mately 21 species are known only from caves. In
comparison, in the west approximately 28 percent of
the records are from caves and approximately 16 spe-
cies are known only from caves. This significant
regional difference may well be explained by the avail-
ability of cave habitats. On a broad scale, the number,
as well as density, of caves is many magnitudes greater
ineastemNorthAmeiicathaninwestemNorthAmerica.
Conversely, there are 39 (+) records for nine species of
Stygobromus from the hyporheic habitat in western
North America, whereas there are very few records for
species of Stygobromus from hyporheic habitats in
eastern North America. The gravel substrates of many
river systems in the west appear to be rich in exploit-
able interstitial spaces, which are in turn essential to
hyporheic habitat structure.

BIOGEOGRAPHY

As shown on the maps in figures 57-61, the geo-
graphic distribution of 53 species of Stygobromus in
western North America (including 5. putealis from
Wisconsin) is widespread and scattered, although dis-
tinct clusters of species occur in north-central Califor-
nia, western Oregon-southern Washington, northwest-
ern Montana-southern Alberta, and north-central Colo-
rado. Many of these species display highly insular
ranges, and 24 are currently known only from single
localities. Moreover, with the notable exception of S.
quatsinensis (see Holsinger et al. 1997), those species
recorded from more than one locality are restricted to
small ranges.

Taxonomic diversity, based on number of species
groups, and species richness within the genus
StygobromusdiXt significantly greaterin North America
east of the Great Plains than in the in North America
westofthe Great Rains (Holsinger 1986b). That these
two regions are generally comparable in size and are
occupied by species of the same genus tends to justify
this comparison. In the east, approximately 94 species
are either described (ca. 72) or are in the process of
being described (descriptions of 22 in ms.), and most of
them can be assigned to one of approximately 16
species groups. However, in the west there are 52

described species, 44 of which are assigned to the
hubbsi group. The remaining nine species, excepting
S. canadensis which appears to represent a distinct
group ( Holsinger 1980), are not assigned to a group.
Moreover, except for the presence of sternal gills, sev-
eral of the nine non-hubbsi group species are similar
morphologically to members of ^e hubbsi group.

A two-fold hypothesis was proposed earlier
(Holsinger 1986b) to explain the reduced taxonomic
diversity in thisregion vis-a-vis the significantly greater
diversity in a comparable-sized part of eastern North
America, and to account for the geographic distribu-
tion of numerous, morphologically closely similar
stygobitic species of Stygobromus over a large part of
the geologically variable terrane of the western Cordill-
era of North America.

First, many physiographic regions in eastern and
central United States with large numbers of species,
such as the Appalachians, Interior Low Rateaus, Oz-
arks, and Edwards Rateau, are characterized locally by
extensive karst terranes that contain numerous cave
and subterranean groundwater habitats. In contrast,
karst terranes in western North America are greatly
limited in extent, and cave density is much lower.
Thus, the availability of karst groundwater habitats is
significantly less in western North America and has
undoubtedly played a major role in limiting the diver-
sity of Stygobromus species. The difference in the
geological history of the two regions has also been
important, inasmuch as potential habitats in the west
probably developed more recently than those in the east
because of the pronounced tectonic activity in the
western Cordillera since the middle Cenozoic. In
contrast, geological activity in the east during the same
time frame has been relatively stable, allowing the
existence of comparatively older and less perturbed
habitats. The more humid climate of eastern North
America would also enhance the development and
recharge of groundwater aquifers, whereas in the west,
the overall drier climate would not be as favorable to
recharge.

Second, the comparative lack of taxonomic diver-
sity in the west may also be a reflection of the pro-
nounced geological activity in the Cenozoic which, in
turn, is largely responsible for the relative youth of
groundwater habitats in this regiion. The widespread
distribution of so many closely similar species of
Stygobromus suggests that they may be derived from a
relatively recent, widely distributed ancestor. The
recent, rapid, and often violent development of present
landforms in the west during the Cenozoic would have

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 137

Table 3. Records of species of Stygobromus from five kinds of generalized habitats

in western North America.

Species

caves^

springs

weils^

hyporheic^

deep lake^

arizonensis

3

1

-

-

-

blirmi

1

-

-

'

-

boultoni

-

-

2

-

-

canadensis

1

-

-

-

-

cherylae

-

1

-

-

-

coloradensis

-

-

-

10(+)

cowani

-

1

-

-

*

curroae

-

-

-

1

-

duplus

-

-

2

-

-

elUotti

3

-

-

-

-

fontinalis

-

2

-

-

-

gallawayae

-

1

-

-

-

glacialis

3

-

-

-

grahami

8

-

-

-

-

gradyi

4

-

-

-

-

harai

3

1

-

-

-

herbsti

-

10

-

-

-

holsingeri

-

1

-

10(+)

-

hubbsi

1

-

-

-

-

hyporheicus

-

-

-

1

-

idahoensis

-

-

-

1

-

imperialis

1

-

-

-

-

interstitialis

-

-

-

3

-

jemezensis

-

-

1

-

lacicolus

-

-

-

-

5

lanensis

-

1

-

1

-

lotus

-

6

-

-

limbus

1

-

-

-

-

mackenziei

1

-

-

-

-

montanensis

-

-

1

-

-

myersae

-

3

-

-

-

mysticus

-

-

1

-

-

obscurus

-

-

1

-

-

oregonensis

1

-

-

-

-

pennaJd

-

-

-

m+)

putealis*

-

-

5

-

puteanus

-

-

1

-

-

quatsinensis

9

8

-

-

-

rudolphi

-

1

-

-

-

rallus

*

5

-

-

-

saltuaris

-

-

-

1

-

sierrensis

-

1

-

-

-

sheldoni

-

5

-

-

-

secundus

-

1

-

-

-

simplex

1

-

-

-

-

tahoensis

-

-

-

-

10

trimis

1

-

-

-

-

tritus

-

-

2

-

-

urospinatus

-

3

-

-

-

utahensis

1

-

-

-

-

wardi

2

-

-

-

-

wengerorum

2

-

-

-

-

wahkeenensis

-

1

-

-

1- Includes 1 mine record each for 5. arizonensis and S.harai and caves in limestone or basalt
for all others; 2- Includes PVC wells driven into stream beds; 3- Includes interrupted streams
for S. holsingerv, 4- Includes only 2 species from Lake Tahoe (precise habitat is unknown).
♦This species is recorded from Wisconsin.

AMMPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 138

fragmented the range of a widespread, ancestral
groundwater species into numerous isolated
populations, ultimately resulting in the evolution of the
many scattered but closely similar species (Holsinger
1974). In comparison, species of Stygobromus in
eastern North America living in groundwater habitats
developed in geologically older and more stable terr-
anes probably have a longer evolutionary history,
which would, in turn, account for their greater taxo-
nomic diversity.

The occurrence of S.putealis in southeastern Wis-
consin just east of the Driftless area (Rg. 57) may give
us a better idea of the age and former potential distribu-
tion of the hubbsi group or its immediate ancestor. The
occurrence of this species approximately 1400 km east
of the western Cordillera is strong evidence of a former
continuous distribution of the hubbsi group or its im-
mediate past ancestor across what is now the Great
Plains (Holsinger 1986b). The present disjunction
between S. putealis and related species in the west may
well have developed through progressive aridity in the
Miocene or destruction of groundwater habitats by
glaciation in southern Canada or the northern United
States during the Pleistocene. Whichever is the correct
explanation for the disjunction, it is probable that a
continuous distribution of the hubbsi group across the
Great Plains is at least as old as the early Reistocene or
even as old as the Miocene.

The pluvial lake system of the Pleistoecene might
have potentially affected the former distribution of
some species of the hubbsi group. It is well established
that an extensive pluvial lake system existed in parts of
the western Cordillea during glacial maxima of the
Pleistocene, which would have effected elevation of
groundwater tables and expansion of groundwater aq-
uifers. However, most of these lakes were developed
in parts of the Basin and Range and Columbia Plateau
and would potentially have i mpacted the di stribution of
relatively few species of Stygobromus, The majority of
species occur the mountainous areas to the east and
west of the provinces with most of the pluvial lakes.

To obtain a better picture of the relationship be-
tween the taxa (species) and their geographic distribu-
tions, we converted the taxon cladogram in Rgure 56 to
the taxon-area cladogram in Rgure 62 by substituting
areas for terminal taxa (species). The areas are major
physiographic provinces of western North America,
and the rationale for using them as areas i s because each
is defined by its unique geomorphogy, climate, and
historical geology. Moreover, each area has endemic
species that are so far unrecorded in any other area.

although one exception may be S. pennakU which has
been recorded from one locality just outside the South-
ern Rocky Mountain area on the extreme western
margin of the Great Plains (Rg. 59).

Within the hubbsi group, the strongest agreement
between taxa and areas is the large clade of 15 species
(extending from tahoensis to imperialis on the
cladogram), which occurs in the western part of the
region. These species are recorded from the Coastal
Range, Cascades, Sierra Nevada, and Columbia Ra-
teau. Moreover, a strong nested subset within this clade
is composed of S.grahamU S. harai, ^dS.wengerorum,
which live in caves in the Mother Lode region of
eastern California (see also Holsinger 1974), and S.
rudolphi from a spring to the southwest in the Coastal
Range. Ten of these 15 species make up the
monophyletic Califomia-Oregon-Washington clade
discussed under Phylogeny, above. A second reason-
ably strong clade is also found in the western part of the
region and consists of eight species from the Coastal
Range, Sierra Nevadaand Cascades (species extending
from S. quatsinensis to S, cherylae on the cladogram).
However, this clade is not fully congruent geographi-
cally, inasmuch as it also contains S. myersae from just
east of the the Sierra Nevada in the Basin and Range,
and more importantly, S.putealisiiom Wisconsin. The
fact that S. putealis is imbedded in the middle of the
cladogram is a further indication that this species,
despite its huge geographic separation from the west-
ern Cordillera, is a bona fide member of the hubbsi
group.

Almost all remaining species on the cladogram are
recorded from the central and eastern parts of the
western Cordillera and occur in different sections of
the Rocky Mountains, Colorado Plateaus, Basin and
Range, and one in the Great Plains (just east of the
Basin and Range in far western Texas) . The position of
S. trims, S. r alius, and S. saltuaris on the cladogram
(Rg. 62) is unclear and incongruent with geographic
distribution. However, it i s clear that the hubbsi group
species appear to be split roughly into eastern and
western subsets on opposite sides of the Basin and
Range-Columbia Rateau (see Rgs. 59-61). Twenty-
three species are recorded from the Cascades-Sierras
Nevada-Coastal Range on the western side of the
Cordillera, and 20 species are recorded from the Rocky
Mountains and Colorado Plateau on the eastern side.
However, only seven species are recorded from the
Basin and Range-Columbia Plateau and four of them
occur near the edge of this combination of provinces.
The extreme aridity that has developed throughout

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 139

Fig. 57. Geographic distribution of 44 species of Stygobromus in the hubbsi group in North America. A closed circle indicates
one to many closely proximate locality records for a species. 1, S. arizonensis; 2, S. blinnv, 3, S. boultoni\ 4, S. cherylae; 5,
S. coloradensis; 6, S. cowani\ 7, S. duplus", 8, 5. elliotti\9, S. fontinalis; 10, S, gallowayaei 11,5. gradyi\ 12, S. grahami\ 13,
S. harai\ 14, S. herbsti\ 15, 5. holsingeri; 16, 5, hubbsi; 17, S. hyporheicus; 18, S. imperialis; 19, S. interstitialis; 20, S.
jemezensis; 21,5. lacicolous; 22, 5. lanensis; 23, 5. latus; 24, 5. limbus; 25; 5. mackenziei; 26, 5. myersae; 27, 5. mysticus;

28. 5. oregonensis; 29, 5. pennaki; 30, 5. putealis; 31,5. puteanus; 32, 5. quatsinensis; 33, 5. rallus; 34, 5, rudolphi; 35, 5.
saltauris; 36, 5. sheldoni; 37, 5. sierrensis; 38, 5. tahoensis; 39, 5. trinus; 40, 5. tritus; 41, 5. urospinatus; 42; 5. utakensis;

43 . 5. wahkeenensis; 44, 5. wengerorum. See figs. 59-6 1 for greater detail. The heavy dashed line represents the approximate
southern extent of Pleistocene glaciation. DA = the Wisconsin Driftless Area.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 140

Fig. 58. Geographic distribution of non-hubbsi group species of Stygobromus in the western Cordillera of North America:
1, S. canadensis; 2, S. curroae; 3, S. glacialis; 4, S. idahoensis; 5, S. montanensis; 6, S. obscurus; 7, S. secundus; 8, S.
simplex; 9, S. wardi', 10, specimens of 1 or 2 new species of Stygobromus from 3 localities west of Calgary that cannot be
described without additional material . See figs. 59 and 60 for greater detail. The heavy dashed line represents the approximate
southern extent of Pleistocene glaciation.

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 141

Fig. 59. Geographic distribution of Stygobromus in Colorado, New Mexico and far western Texas. A closed circle indicates
one to many closely proximate localities for a species (number of recorded localities in parentheses). Hubbsi group: S. limbus
(1 loc.); 2, S,jemezensis (1 loc.); 3, S. inter stitialis (3 Iocs.); 4, S. coloradensis (8 or 9 Iocs.); 5, S.pennaki (7 or 8 Iocs.); 6,
S. holsingeri (approx. 25 Iocs.); 7, S. urospinatus (3 Iocs.); 8,; 9, S.fontinalis (2 Iocs.). Other species: 9, S. wardi (2 Iocs.);
10, S. simplex ( 1 loc.) ; 11, S. curroae ( 1 loc.). Insert: closed circles, S. holsingeri', open circles, S. urospinatus', x, both species
from same locality.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 142

Fig. 60 . Geographic distribution of in Idaho, Montana, Oregon, and Washington, USA; and Albertaand British

Columbia, Canada. A closed circle indicates one to several closely proximate localities for a species (number of recorded
localities in parentheses), hubbsi group: 1, S. hubbsi ( 1 loc.); 2, 5. oregonensis (I loc.); 3, S. lanensis{l loc.); 4, S. saltauris
(1 loc.); 5, S. wahkeenensis (2 Iocs.); 6, S, elliotti (3 Iocs.); 7, S. latus (3 Iocs.); 8. 5. rallus (2 Iocs.); 9, S. duplus (2 Iocs.);
10 S. tritus (2 Iocs.) ; 1 1 , 5. puteanus ( 1 loc.) ; 12, S. quatsinenesis (4 loctdities on Vancouver Island; 26 in southeastern Alaska
not shown). Other species: 13, S. idahoensis (1 loc.); 14, S. montanensis (1 loc.); 15, S. obscurus (1 loc.); 16, S. glacialis
(3 Iocs.); 17, S. canadensis (1 loc.); 18, S. secundus (1 loc.); 19 & 20, S. spp.(2).

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 143

Fig. 61. Geographic distribution of Stygobromus in Arizona, California, Nevada and Utah. A closed circle indicates one to
several closely proximate localities for a species (number of recorded localities in parentheses), lubbsi group: 1 , S. mysticus
( 1 loc.) ; 2, S. trinus ( 1 loc.) ; 3, 5, gallowayae ( 1 loc) ; 4, S. cherylae ( 1 loc.) ; 5, S. cowani ( 1 loc.) ; 6, S. hyporheicus ( 1 loc.) ;
7, S. imperialis ( 1 loc.) ; 8, S. mackenziei ( 1 loc.) ; 9, S. rudolphi ( 1 loc.) ; 10, 5. myersae (3 Iocs.) ; 1 1 , 5. wengerorum (2 Iocs.) ;
12, S. harai (4 Iocs.); 13, S, grahami (8 Iocs.); 14, S, gradyi (5 Iocs.); 15, S. sierrensis ( 1 loc.); 16, S. sheldoni (4 or 5 Iocs.);
17, S. tahoensis (9 or 10 sites in Lake Tahoe); 18, S. lacicolous (5 sites in Lake Tahoe); 19, S, herbsti (10 Iocs.); 20, S.
arizonensis (4 Iocs.), 21, S. boultoni (2 Iocs.); 22, S. blinni (1 loc.); 23, S. utahensis ( 1 loc.). Insert: open circles, S. grahami;
closed circles, S. gradyi; x, localities with both S. grahami and S. gradyi; closed triangles, S. harai; closed boxes, S.
wengerorum.

AMPHIPACIFICA VOL. 3 NO. 2. NOV. 15, 2001 144

S. ambulini

C. forbul

B. brachycaudiu

wardl

SR

simplex

COP

curroae

SR

montsBensis

NR

obscurus

NR

fceuBdus

GP

glaciaiis

NR

esnadensis

NR

idaboensis

NR

impcrialas

CR

rudolpfal

CR

grahaml

SN

wcBcarorum

SN

liaral

SN

gallswayac

SN

latus

CLP

gradyl

SN

cllioUi

CM

hyporbcicus

CR

mystictts

CR

hubbsl

CLP

oragancniis

CR

macktnzlal

CR

tahoeniU

SN

cowaoi

CR

chcrylae

CR

myarsaa

BR

abeldoBi

SN

tacicolus

SN

pUtMlJf

Wisconsin

ticrrcjuis

SN

laDcnsb

CM

wahkacnensb

CM

qumtetiieiisif

CR

artzoociisis

BR

bolsingerl

SR

blianl

COP

coloradcnsit

SR

itroiplnatus

SR

pc&aaki

SR

putcaBus

NR

lAterslUIalis

SR

utabcnsls

MR

duplus

NR

bQuUoni

BR

trltus

NR

(rtaus

CR

berbsti

BR

jcmexcBsls

SR

foatlnallt

COP

limbus

GP

rallus

CLP

saltuarb

CM

Fig. 62. Taxon-area cladogram converted from the majority-rule consensus tree in Fig. 56. Areas (see text) (with
their number of species in parentheses) are as follows: Basin and Range (BR) (4 spp.); Colorado Plateaus COP
(3 spp.); Northern Rocky Mountains NR (8 spp.); Middle Rocky Mountains (MR) (1 sp.); Southern Rocky
Mountains (SR) (8 spp.); Columbia (Lava) Plateau (CLP) (3 spp.); Cascade Mountains (CM) (4 spp.); Sierra
Nevada Mountains (SN) (9 spp.); Coastal Range (CR) ( 10 spp.); Great Plains (GP) (2 spp.); and Wisconsin ( 1 sp.).

AMPHIPACIHCA VOL. 3 NO. 2. NOV. 15, 2001 145

much of the Basin and Range and Columbia Plateau
since the Pleistocene (see Mincldey et al. 1986) has
probably led to the development of dispersal barriers
for groundwater amphipods and contributed to physi-
cal isolation of species of the hubbsi group to opposite
sides.

It is also important to note that all of the nine non-
hubbsi group species occur on the eastern side of the
western Cordillera and that six of them are recorded
from areas that were glaciated or were in close proxim-
ity to glaciation during the Pleistocene. Although
some of these species appear to occupy a position close
to the ancestor(s) of the hubbsi group, others, such as
5. canadensis^ may not.

ADDENDUM

Subsequent to this study and during the last phase of
manuscript preparation, a significant number of am-
phipod samples from groundwater habitats or
groundwater-associated habitats in western North
America were sent to JRH for determination. Although
it was too late to include a detailed study of this mater-
ial in the present paper, a very brief summary will be
made. These samples were collected from approxi-
mately 22 different localities in five different states and
appear to represent approximately 1 1 undescribed spe-
cies of Stygobromus, some of which can be assigned to
the/iw^/75igroup,andonepossiblytoanewcrangonyctid
genus. The collections were made in the following
localities by the persons indicated in parentheses: 3
sites (stream sediments) in Santa Clara County (Steve
Fend) and a spring in Somona County (C. B. Barr and
W. D. Shepard), California; 1 site (hyporheic ?) in
Boise County, Idaho (Gary Lester ?); hypohreic and
deep lake sites in Jackson, Josephine and Klamath
counties, Oregon (R. W. Wisseman); 2 springs and 1
cave in Cache County, Utah (Candace Brindza);
hyporheic habitats in Queets River, Olympic National
Park, Jefferson County (Holly Coe) and East Lakes,
Cascades National Park, Whatcom County (Amy Hill),
Washington. In addition, undetermined species of
Stygobromus, one of which is apparently S. glacialis,
are reported by Stanford et al. (1994) and J. V. Ward
(pers. comm.) from groundwater habitats in the Flat-
head River catchment in northwestern Montana.

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Legend for Figures

Al,2

antennae 1, 2

OP

outer plate

ABD -

abdomen

P3-7

peraeopods 3, 4, 5, 6, 7

BRPL -

brood lamella

PLP

pleopod

CX

coxal plate

RT

right

EP 1,23 -

abdominal side plates 1, 2, 3

T

tel son

GN1,2 -

gnathopods 1, 2

Ul,23 -

uropods 1, 2, 3

HD

head

UL

upper lip (incl. epistome)

IP

inner plate

X

part enlarged

LFT

left

cf

male

LL

lower lip (labium)

9

female

MD

mandible

MXl

maxilla 1

Note: All appendages are those of the female specimen.

MX2 -

maxilla 2

, unless preceded by the male (d*) symbol. Individual

MXPD -

maxilliped

appendages

are enlarged to visually suitable scales.

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