Overview of the Late Cretaceous fishes of the northern Western Interior Seaway

Abstract

The Cretaceous Western Interior Seaway covered North America to various extents from the Albian through Maastrichtian. During this time, the marine waters were home to a large diversity of fishes. Although the fish faunas of the southern parts of the seaway are fairly well known, the fishes of the northern part have not been as well documented. A number of new discoveries in recent years have greatly increased our knowledge of these
northern fishes. These faunas and fishes are reviewed here.

Full text

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Khosla, A. and Lucas, S.G., eds., 2016, Cretaceous Period: Biotic Diversity and Biogeography. New Mexico Museum of Natural History and Science Bulletin 71.
OVERVIEW OF THE LATE CRETACEOUS FISHES OF THE
NORTHERN WESTERN INTERIOR SEAWAY
ALISON M. MURRAY1 and TODD D. COOK2
1Department of Biological Sciences, University of Alberta, Edmonton, AB, T6G 2E9, Canada, ammurray@ualberta.ca; 2School of Science,
Pennsylvania State, Erie, The Behrend College, 4205 College Drive, Erie, PA, 16563-0203, USA, tdc15@psu.edu
Abstract—The Cretaceous Western Interior Seaway covered North America to various extents from the Albian
through Maastrichtian. During this time, the marine waters were home to a large diversity of shes. Although the
sh faunas of the southern parts of the seaway are fairly well known, the shes of the northern part have not been
as well documented. A number of new discoveries in recent years have greatly increased our knowledge of these
northern shes. These faunas and shes are reviewed here.
INTRODUCTION
The Cretaceous Western Interior Seaway (WIS) covered the west-
central part of North America to various extents from the Albian through
Maastrichtian (Smith et al., 1994). At times, the seaway stretched
all the way from the Gulf of Mexico in the south, to reach the Arctic
Ocean in the north. This great expanse of shallow, continental seaway
provided habitat for a diverse array of marine organisms, and formed
an important connection between the Western Tethys (modern Gulf
of Mexico area) and the Boreal Sea (Arctic Ocean). A northwestern
branch of the seaway, named the Hudson Arm, was probably present
during at least some of this time, forming a connection to the east with
the Eastern Tethys region (e.g., Blakey, 2014).
Shallow seaways proved to be optimal habitats for the evolution
of a number of sh taxa. The water depth, expanded coastal areas, and
paleoenvironments provided many niches for smaller shes to inhabit
and in which to diversify. The eastern Tethys Sea (modern Mediterranean
Sea) was the home of many small shes that are documented by the rich
fossiliferous deposits of Lebanon (i.e., Namoura, Sahel Alma, Hakel
and Hajula; Forey et al., 2003) and more recently discovered fossil
shes from Morocco (e.g., Cavin, 1999; Cavin et al., 2010; Murray et
al., 2013). Other eastern Tethyan shes inhabited the northern coastal
areas, such as the Dalmatian Coast or Komen (e.g., Radovčić, 1975;
Forey et al., 2003). The ichthyofaunas from the southernmost part
of the WIS have also become better known and well documented,
with recently described faunas from Mexico (e.g., Alvaredo-Ortega
and Than–Marchese, 2013; González-Rodríguez and Fielitz, 2008;
González-Rodríguez et al., 2013). Much work is continuing on the
Mexican WIS faunas.
Material from the central part of the WIS of the present day United
States has been known for many years, with elasmobranchs and bony
shes described from a wide variety of localities (see Russell, 1988).
In contrast, the northernmost faunas of the WIS have, until recently,
been less well documented. We here provide information on material
from Canada that documents the shes from the northernmost part of
the WIS from the Albian/Cenomanian through to the Maastrichtian. In
the southern parts of the Canadian WIS, most sh material is preserved
as isolated elements, with few previously described articulated sh,
other than notable exceptions such as sturgeon remains (Grande and
Hilton, 2009). Russell (1988) previously provided an overview of the
vertebrates of the Western Interior Seaway, including the sh faunas.
However, new research from the last decade, and still ongoing, is
providing much new information on shes from throughout the
Canadian WIS, from southern Alberta and Saskatchewan to the
Northwest Territories. We here provide an overview of the ichthyofaunas
from the northern Western Interior Seaway of Canada, including past
discoveries and covering new localities and shes that have recently
been documented. We also include several freshwater localities, which,
although clearly not part of the marine seaway, are important as they
document the retreat of the seaway in the later part of the Cretaceous,
and expand our understanding of the Cretaceous faunas by providing a
contemporaneous context for the marine shes.
FISH-BEARING LOCALITIES
A number of localities in the Canadian part of the Western Interior
Seaway have provided sh material (Fig. 1). We here describe the
relevant localities in chronological age, starting with the oldest, and list
the sh taxa from each locality. The sh taxa and their relationships are
discussed after.
Albian
The oldest northern WIS locality to produce sh remains is the
Loon River Formation in the Northwest Territories. The Loon River
Formation has been dated as early to middle Albian but may be late
Albian (see Vernygora et al., 2016). From this locality, an enigmatic
sh was described as Erichalcis arcta by Forey (1975) as the presumed
most basal member of Clupeomorpha. Further analysis of this species
showed it to be a composite based on material of two separate
taxa. These have now been redescribed as a euteleost, keeping the
name Erichalcis arcta (Hermus et al., 2004), and the newly named
FIGURE 1. Map of western Canada with the general position of the
localities noted in the text. Numbered localities are: 1, Banks Island;
2, Anderson River; 3, Lac des Bois; 4, Loon River; 5, multiple sites in
the Fish Scales and Dunvegan formations; 6, Judith River Group; 7,
Bainbridge River, Carrot River and Etamami River localities.

256
Foreyclupea loonensis, an ellimmichthyiform clupeomorph (Vernygora
et al., 2016). A second species of Erichalcis, and E. conspicua, were
described from the middle Albian Christopher Formation, on Axel
Heiberg Island, Nunavut, Canada (Hermus et al., 2004), documenting
the roughly contemporaneous presence of this genus in the Cretaceous
Boreal Sea.
Cenomanian
The Cenomanian shes of the Canadian WIS come from three
formations, the Fish Scales and Dunvegan formations in Alberta,
and the Ashville Formation in Saskatchewan. The early Cenomanian
Fish Scales Sandstone (also called the Fish Scales Zone or the Fish
Scales Marker Bed) crops out in some areas of Alberta, from which
the named sh material has been collected; this layer has been dened
in the subsurface of east-central Alberta as the Fish Scales Formation
(Bloch et al., 1993). Most of the material from the marine shales of
this formation consists of isolated, disarticulated bones, scales, and
teeth; however, there is a single species, named for the zone (Xenyllion
zonensis), represented by disarticulated but associated bones of a small
paracanthopterygian (Wilson and Murray, 1996).
The Dunvegan Formation overlies the Fish Scale Formation in
at least some areas, and is dated at middle Cenomanian (Hay et al.,
2007). This formation crops out in northwestern Alberta, but has also
been sampled subsurface through drill cores. One particular drill core
provided the holotype, and only known specimen, of Tycheroichthys
dunveganensis, a small ellimmichthyiform sh missing only the tip of
the snout and tip of the caudal n rays (Hay et al., 2007). The Dunvegan
Formation is also the source of sturgeon material: a partial skull was
found in 2013 in the Peace River region of northwestern Alberta (Vavrek
et al., 2014). The Dunvegan Formation has produced a rather small
number of elasmobranchs. Cook et al. (2008) reported an assemblage
consisting of multiple lamniform taxa (Squalicorax, Archaeolamna,
Cretoxyrhina, Protolamna, Johnlongia) but only a single hybodont
(Meristodonoides rajkovichi) and a single batoid (Pseudohypolophus
mcnultyi) species.
The middle to late Cenomanian Belle Fourche Member of the
Ashville Formation in east-central Saskatchewan has provided a
wealth of disarticulated bonebed material. The material comes from
several localities, with an important locality being Bainbridge River
in the Pasquia Hills (Cumbaa and Tokaryk, 1999; Cumbaa et al., 2006,
2013). In contrast to the small number of elasmobranch species found
in the Dunvegan Formation, the Ashville Formation of the Carrot
River and Bainbridge River in the Pasquia Hills, which samples the
eastern margin of the WIS, has produced a rather large and diverse
contemporaneous assemblage with at least 20 chondrichthyans
(Cumbaa and Tokaryk, 1999; Cumbaa et al., 2006; Underwood and
Cumbaa, 2010). Bainbridge River also has a great taxonomic diversity
of actinopterygians, with 15 species, as well as other vertebrates
(Cumbaa et al., 2006). The reported elasmobranch taxa include a single
species of hybodont (Meristodonoides rajkovichi), several species
of Ptychodus, multiple lamniforms (Squalicorax, Palaeoanacorax,
Archaeolamna, Cretoxyrhina, Cretalamna, Carcharias, Roulletia),
two orectolobiforms (Orectoloboides, Cretorectolobus), and two
batoids (Rhinobatos incertusi, Cretomanta canadensis). Important
actinopterygian records include pycnodonts, a possible salmoniform
and acanthomorph remains (Cumbaa et al., 2013). The Carrot River
bone bed is found farther south in Saskatchewan; it is somewhat less
diverse in sh taxa than the Bainbridge River bone bed (Cumbaa and
Tokaryk, 1999).
Turonian
The eastern margin of the WIS is also preserved in the early
Turonian, with deposits of the Favel Formation, from which has
come a small diversity of sharks. Case et al. (1990) reported a single
species of Ptychodus (P. anonymus), several lamniforms (Squalicorax,
Cretoxyrhina, Dallasiella (reported as Cretodus), Carcharias,
Odontaspis) and two batoids (Rhinobatos, Cretomanta). Cumbaa
and Tokaryk (1999) identied articulated actinopterygian skeletons
of Xiphactinus, Apsopelix and Pachyrhizodus, as well as remains of
Enchodus, Gillicus, Ichthyodectes and Protosphyraena.
The early Turonian of the Canadian WIS is also represented at
the far northern end of the seaway by the Lac des Bois locality in
the Northwest Territories. Lac des Bois has provided a number of
osteichthyan taxa, including acanthomorphs, an amiiform, euteleost,
ichthyodectiforms, elopiforms and enchodontoid aulopiforms (Cumbaa
and Murray, 2008). Farther north, in the Boreal Sea rather than the WIS,
the Turonian Banks Island locality (Kanguk Formation) also contains
Avitosmerus, Ichthyodectes, and ?Osmeroides, as well as indeterminate
salmoniform, aulopiform and alepisauroid material (Wilson, 1978;
Cumbaa and Murray, 2008). The Banks Island specimens are all
disarticulated remains, rather than articulated shes as at Lac des Bois,
making identications and comparisons more difcult.
Farther south, the Watino localities in the early Turonian Kaskapau
Formation of northwestern Alberta have produced fragmentary
actinopterygian bones and scales of Belonostomus, Ichthyodectes,
Apateodus, Cimolichthys, and Enchodus and Dercetidae (Wilson and
Chalifa, 1989). Also recovered from the localities were numerous
elasmobranch taxa included two hybodontiforms (Meristodonoides,
Polyacrodus), a single species of Ptychodus (Ptychodus anonymus),
multiple lamniforms (Squalicorax, Archaeolamna, Cardabioodon,
Cretoxyrhina, Cretodus, Cretalamna, Scapanorhynchus, Johnlongia,
Dallasiella, Carcharias, Odontaspis), and a single batoid (Rhinobatos).
Just south of the Canadian–U.S. border, Siverson and Lindgren
(2005) reported several elasmobranchs from the mid-Turonian Fairport
Member of the Carlile Shale in east-central Montana. The taxa included
several species of Ptychodus, multiple lamniforms (Squalicorax,
Cardabiodon, Cretoxyrhina, Cretalamna, Johnlongia, Dallasiella,
Carcharias), and a single batoid (Cretomanta). Therefore, this
Turonian elasmobranch fauna was fairly widely distributed throughout
the seaway.
Coniacian–Santonian
Fishes from the Coniacian and Santonian are not well represented
in the Canadian WIS. There are some deposits of this age in the eastern
part of the seaway, in a locality on the Etomami River of Saskatchwan
(Cumbaa and Tokaryk, 1999). This locality has sediments from both
the Campanian Pierre Formation and Coniacian-Santonian Niobrara
Formation, with the only reported remains from the latter being bones
of Enchodus (Cumbaa and Tokaryk, 1999).
Santonian–Campanian
Material collected many years ago from the Puskwaskau
(Santonian-Campanian) Formation of northwestern Alberta and
curated in the Royal Tyrrell Museum and Canadian Museum of Nature
since 1973 and 1961, respectively, has recently been conrmed as
Xiphactinus audax Vavrek et al. (2016). This is the most northerly,
conrmed presence of Xiphactinus in the WIS.
The Campanian Pierre Shale has also provided remains of
Enchodus as well as a single Xiphactinus scale (Cumbaa and Tokaryk,
1999). Bardack (1965, 1968) reported material from the Boyne and
Pembina members of the Vermillion River Formation. The Boyne
Member is now included as part of the Carlile Formation, and is
considered older than Campanian (Nicholls and Russell, 1990;
Bamburak and Nicholas, 2009).The fauna from the Pierre Shale and
Carlile formations of Manitoba contain the elasmobranchs Ptychodus
and Scapanorhynchus, Squalicorax, and Cretalamna, as well as the
bony shes Gillicus, Xiphactinus, cf. Bananogmius, Apsopelix, and
Enchodus (Bardack, 1965, 1968)
Late Cretaceous deposits along the Anderson River in the Northwest
Territories have produced some poorly preserved sh; many of these
were not collected, but eld identications included Xiphactinus, cf.
Cimolichthys, cf. Pachyrhizodus, and cf. Enchodus from the Santonian-
Campanian Smoking Hills Formation, and cf. Acipenser, cf. Apsopelix,
cf. Enchodus and cf. Stratodus from the Campanian-Maastrichtian
Mason River Formation (Russell 1967, 1988; Yorath and Cook, 1984).
Campanian
Campanian marine WIS deposits are present in the southern part
of Alberta, left by the Bearpaw Sea. However, the Campanian deposits
that have produced most of the shes in southern Alberta are brackish
or freshwater deposits, rather than fully marine (Brinkman, 1990;
Brinkman et al., 2004). From the marine to brackish shore localities
of the Foremost Formation (Belly River Group), Brinkman et al.
(2004) reported a number of elasmobranch taxa, including a hybodont,
Synechodus, an orectolobiform, two lamniforms (Archaeolamna,
Synodontaspis) and two batoids (Ischyrhiza, Rhinobatus). The only
bony shes that Brinkman et al. (2004) reported from these deposits
are Belonostomus, Paralbula, Actractosteus and an acipenserid. More
recently, sh teeth and other isolated remains have been reported from
another microfossil site in marine deposits of the Foremost Formation;
the additional genera identied from there are the elasmobranchs
Odontaspididae indet. and Pseudomyledaphus, the chimaeriform

257
Elasmodus, and the actinopterygians Lepisosteus and Horseshoeichthys
(Cullen et al., 2016).
The freshwater Campanian deposits, some in the Foremost
Formation and others of the Oldman and Dinosaur Park formations,
have produced the elopiforms Paralbula and Coriops (although this
latter may be an osteoglossomorph; see Neuman and Brinkman, 2005),
esocoids, amiids, Paratarpon, and indeterminate teleosts (Brinkman,
1990; Brinkman et al., 2004; Neuman and Brinkman, 2005). Campanian
freshwater deposits of the Dinosaur Park Formation have also produced
the northernmost record, and only Canadian characiform, indicating a
much warmer environment during this time (Newbrey et al., 2009). The
Lethbridge Coal Zone of the Dinosaur Park Formation has produced
several marine chondrichthyans and actinopterygians (Beavan and
Russell, 1999; Brinkman et al., 2005). The reported elasmobranch taxa
include a single species of hybodont (Meristodonoides montanensis,
as Hybodus montanensis), an orectolobiform (Cretorectolobus), three
lamniforms (Archaeolamna, Carcharias, Odontaspis) and four batoids
(Protoplatyrhina, Ischyrhiza, Myledaphus, Ptychotrygon). Beavan
and Russell (1999) also reported holocephalan material (Elasmodus).
Additional elasmobranch taxa recorded by Brinkman et al. (2005) are
Cretalamna, Squatina, Eucrossorhinus, Archaeotriakus and Ischyrhiza.
The actinopterygians from the area include Acipenser, Enchodus, a
polyodontid, Atractosteus, Paralbula, and a number of unidentied taxa
(Beavan and Russell, 1999; Brinkman et al., 2005). Farther south, in
contemporaneous deposits in Montana, Case (1978, 1979) reported two
holocephalans (Elasmodus and Ischyodus), two species of hybodont,
two species of Synechodus, two orectolobiforms (Cretorectolobus,
Chiloscyllium), several lamniforms (Squalicorax, Archaeolamna,
Carcharias, Odontaspis), a carcharhiniform (Archaeotriakis)
and several batoids (Protoplatyrhina, Ischyrhiza, Ptychotrygon,
Myledaphus). Konishi et al. (2014) identied lamniforms (Squalicorax
and unidentied taxa) from the overlying upper Campanian Bearpaw
Formation of Alberta. Ursichthys longiparietalis, a partially articulated
aulopiform preserved in the gut of a mosasaur, is from the lower part
of the Bearpaw Formation of southern Alberta south of Lethbridge
(Newbrey and Konishi, 2015).
Towards the end of the Cretaceous, the Western Interior Seaway
had narrowed signicantly, and it eventually lost its southern
connection to the western Tethys (Gulf of Mexico) at the end of the
Campanian. However, a number of freshwater deposits are known
in the area from this time. Articulated freshwater shes include the
Campanian osteoglossomorph Cretophareodus (Li, 1996) from the
Oldman Formation of southern Alberta, and the sturgeon Priscosturion
longipinnis from the Judith River Formation of southern Alberta and
Montana (Grande and Hilton, 2006, 2009).
Maastrichtian
Elasmobranchs reported from the late Maastrichtian Hell Creek
Formation in Montana include a hybodont (Lonchidion), several
orectolobiforms (Protoginglymostoma, Chiloscyllium, Restesia) and
two batoids (Ischyrhiza, Myledaphus) (Estes et al., 1969; Bryant,
1989; Cook et. al., 2014). Hoganson and Murphy (2002) reported the
multiple lamniforms (Cretalamna, Serratolamna, Paranomotodon,
Carcharias, Odontaspis), a carcharhiniform (Galeorhinus), several
batoids (Pseudohypolophus, Ischyrhiza, Peyeria, Ptychotrygon,
Myledaphus), and holocephalans (Ischyodus) from the same formation
in North Dakota. Future work may well allow us to document these
taxa in southern Alberta as well. Currently, Maastrichtian marine shes
are poorly represented in Canada, with only brackish water deposits
of the upper beds of the Horseshoe Canyon Formation from which
was reported the marine shark Cretoxyrhina (Larson et al., 2010).
However, this species is strictly marine, and this occurrence would be
the youngest known, so we suspect it may have been reworked or that
there is another explanation for its presence in this formation.
The Canadian Maastrichtian sh-bearing deposits are found in
southern Alberta, and essentially all represent freshwater environments.
Horseshoeichthys armiserratus is an ellimmichthyiform clupeomorph
from the Horseshoe Canyon Formation in Dry Island Buffalo Jump
Provincial Park in south-central Alberta (Newbrey et al., 2010a). The
Scollard Formation also crops out in this park, and a relatively new
locality, dubbed Pisces Point, is producing signicant new articulated
specimens (e.g., Newbrey et al., 2013a; Murray et al., 2016). Scollard
Formation shes are also represented by many disarticulated remains
in the same area (Murray et al., 2016). Elasmobranchs reported from
the Scollard Formation include an orectolobiform (Chiloscyllium) and
a batoid (Myledaphus) (Cook et al., 2014).
By the end of the Maastrichtian, the WIS was reduced to a narrow
body of water in the far north, with a few scattered and isolated water
bodies in central Canada, and disappeared completely in the Paleocene
(e.g., Blakey, 2014). From this time on, the freshwater ichthyofaunas
took over the area, with a large number of diverse taxa represented
in the Paleogene Paskapoo Formation, including paracanthopterygians
(Massamorichthys, Lateopisciculus), the osmerid Speirsaenigma, an
osteoglossiform (Joffrichthys), and the pike, Esox tiemani (Wilson
and Williams, 1991; Li and Wilson, 1996; Murray, 1996; Murray and
Wilson, 1996).
DIVERSITY AND ENVIRONMENTS OF FISHES IN THE
NORTHERN WESTERN INTERIOR SEAWAY
The shes recovered from the northern part of the Western Interior
Seaway represent a wide variety of taxa. More primitive forms, such as
sturgeons and amiiforms, are represented, along with groups that are
now completely extinct, such as the pycnodonts and ichthyodectids.
Some of the earliest records of acanthomorph shes are found in the
northernmost part of the seaway. Some taxa are spread throughout the
seaway, and others have been found only in very restricted regions. In
this section we comment on the taxa that are currently known from the
northern WIS, in taxonomic order.
Elasmobranchs
Underwood and Cumbaa (2010) noted that the mid-Cenomanian
Bainbridge assemblage from Saskatchewan was dominated by pelagic
lamniforms consistent with other WIS Cretaceous assemblages. The
Dunvegan assemblage of Alberta supports this observation (Cook et
al., 2008). Unlike assemblages reported from the southern seaway (e.g.,
Welton and Farish, 1993; Cappetta and Case, 1999) both Canadian
assemblages have rather limited benthic (and/or nektobenthic)
durophagus shark species and batoid diversity, likely indicating an
adverse environment in this area, possibly due to the persistent anoxic
bottom waters (Cook et al., 2008; Cumbaa et al., 2010; Underwood and
Cumbaa, 2010).
The paucity of benthic selachian species within the northern region
of the WIS appears to have continued into the Turonian (Siverson and
Lindgren, 2005; Cook et. al, 2013). By the Campanian, however, there
is an increase in both batoid and small benthic orectolobiform species
(Case, 1978, 1979; Brinkman, 1990; Beavan and Russell, 1999). The
diversity of batoids appears to continue into the late Maastrichtian
(Hoganson and Murphy, 2002).
Chondrostei, Acipenseriformes
There are two sturgeons known from the Canadian portion of the
WIS. Grande and Hilton (2006) described Priscosturion (originally
as Psammorhynchus, a preoccupied name) from Montana. They later
described an articulated partial specimen of this species that was
recovered from deposits of the Campanian Judith River Group of
southern Alberta (Grande and Hilton, 2009). The second sturgeon is
an acipenserid from the middle Cenomanian Dunvegan Formation of
north-central Alberta (Vavrek et al., 2014). This specimen is a partial
articulated skull, being the anterior half of the parasphenoid and most
of the left and right parietals, from a large sh that had an estimated
length of 5 m (Vavrek et al., 2014). Although sturgeons are rare in the
WIS fossil record, this may be a factor of poor preservation of these
predominantly cartilaginous shes.
Pycnodontiformes
Pycnodonts have only been reported from the Cenomanian
Bainbridge River bone beds representing the eastern margin of the
northern WIS. The pycnodonts in the Bainbridge River deposits are not
as common as some taxa, but not rare, with between 50-100 specimens
reported (Cumbaa and Tokaryk, 1999: table. 1). Pycnodonts are often
reconstructed as shallow reef shes (e.g., Nursall, 1996). They became
broadly distributed in the shallow Cretaceous seas of the Tethys,
particularly during the Cenomanian and Turonian (Nursall, 1996).
Without further fossil nds, we are unable to suggest whether this
localized occurrence of pycnodonts in the WIS is the result of a unique
paleohabitat or paleoenvironment, or if perhaps pycnodonts only
managed to expand into the eastern areas and were blocked by deeper
waters from crossing the WIS. The lack of pycnodonts elsewhere in the
WIS may also be related to preservational bias, with the Bainbridge
bonebeds capturing an extremely rich diversity of isolated elements,
compared to other deposits preserving a smaller diversity of articulated
remains.

258
Amiiformes
Amiiform shes are represented in the earlier Canadian WIS by
marine caturids, but are later represented by the freshwater amiids in
younger deposits. Caturids are represented in the Turonian Lac des Bois
locality and in the Cenomanian Bainbridge River bonebed (Cumbaa et
al., 2013). The Lac des Bois record is based on a single specimen which
is a partial articulated tail, whereas the Bainbridge River specimens
are isolated bones, including toothed elements that are considered
uncommon in the locality (Cumbaa et al., 2013). Campanian amiids
are present in southern Alberta. Maastrichtian freshwater amiids are
represented in the Scollard Formation (see below).
Teleostei
Crossognathiformes, Pachyrhizodontidae
The only crossognathiform shes known from the northern WIS
are both members of the Pachyrhizontidae (Pachyrhizodontoidei).
Aquilopiscis wilsoni from the Turonian Lac des Bois locality (Cumbaa
and Murray, 2008) is named on the basis of multiple articulated
skeletons. Bone bed material from the Cenomanian Bainbridge
River, Saskatchewan, has produced remains of Elopopsis sp. and
Pachyrhizodus minimus (Cumbaa et al., 2013).
Ichthyodectiformes, Ichthyodectidae
Ichthyodectiforms are generally fairly large shes and are
represented in the WIS by three ichthyodectid genera, Xiphactinus,
Ichthyodectes and Gillicus, as well as remains that cannot be determined
to genus. Xiphactinus has been recovered from the Cenomanian
Ashville Formation at Carrot River, Saskatchewan, Santonian-
Campanian Puskwaskau Formation in north-central Alberta, Turonian
Favel Formation of southern Saskatchewan, and the Campanian Pierre
Formation at Etomami River, also in Saskatchewan (Cumbaa and
Tokaryk, 1999; Vavrek et al., 2016). Therefore, Xiphactinus ranges
in the northern WIS from south to fairly far north, and from east to
west. Many of these specimens represent quite large sh with some
articulated and fairly complete (pers. obs.).
Ichthyodectes has been recovered from the same Turonian Favel
and Cenomanian Ashville formations in Saskatchewan. This genus is
also present in the Turonian localities of Lac des Bois in the northern
part of the seaway and Banks Island in the Boreal Sea (Wilson, 1978;
Fielitz, 2002; Cumbaa and Murray, 2008). These records are based on
isolated scales (pers. obs.).
Gillicus is the least common of the ichthyodectiforms in the
northern WIS. It has only been reported from the Turonian Favel
Formation in southern Saskatchewan, representing the eastern side of
the seaway (Cumbaa and Tokaryk, 1999).
Elopomorpha
Two records of Osmeroides represent the only marine elopiform
shes reported from the northern WIS. These are from the northern
end of the seaway, recovered from Lac des Bois, and the Boreal
Sea represented by Banks Island (Fig. 1). This distribution suggests
Osmeroides preferred cooler or more open water than was available in
the more southern area of the seaway.
Euteleostei incertae sedis
A small euteleostean sh, Avitosmerus canadensis, was described
from the Turonian Lac des Bois locality (Fielitz, 1996, 2002).
Additional, better preserved, material collected later allowed further
details of this sh to be determined (Murray and Cumbaa, 2015).
However, this sh remains incertae sedis, mainly because of a lack of a
phylogenetic framework for early euteleost shes. Distinctive isolated
bones, such as the preopercle, document the presence of Avitosmerus on
Banks Island in the Turonian Boreal Sea as well.
Cumbaa et al. (2013) noted the presence of a possible salmoniform
in the Bainbridge locality. Future study of these small, early teleost
shes may eventually provide us a better understanding of their
relationships.
Clupeomorpha, Ellimmichthyformes
The extinct order Ellimmichthyiformes forms the sistergroup to
the Clupeiformes, which contain all living clupeomorph shes (as well
as some fossil forms). The extinct order is almost global in distribution,
and the earliest ellimmichthyiform is known from the early Early
Cretaceous. Members of the order survived in marine, estuarine, and
later fresh waters through until the Eocene.
Two ellimmichthyiforms are known from the Canadian WIS. The
older Canadian species is Tycheroichthys dunveganeneis, from middle
Cenomanian deposits (Dunvegan Formation) in north-central Alberta
(Hay et al., 2007). Tycheroichthys was recovered from a subsurface
drill core taken south of Grande Prairie. The core fortuitously
captured the whole sh (hence the generic name meaning lucky sh)
from a depth of greater than 1300 meters. The second northern WIS
ellimmichthyiform is the younger (Maastrichtian) Horseshoeichthys
armigserratus. This species is represented by a single, partial articulated
sh, from freshwater deposits of the Horseshoe Canyon Formation of
central Alberta (Newbrey et al., 2010a). According to the analysis of
Murray and Wilson (2013), the older Tycheroichthys is a member of
the family Paraclupeidae, whereas Horseshoeichthys is a more basal
ellimmichthyiform related to Armigatus. The biogeography of these
shes is still not well understood.
Aulopiformes, Enchodontoidei
Enchodontoids are known from throughout the WIS. In the
Turonian, Enchodus, Cimolichthys and Ornatipholis are found in the
northern Lac des Bois locality (Fielitz, 1996; Cumbaa and Murray,
2008). Three species of Enchodus are found in the Cenomanian
Belle Fourche Member of the Ashville Formation (Bainbridge River
bonebed). These have been identied as Enchodus cf. E. gladiolus and
cf. E. shumardi, and the third is just referred to Enchodus sp. (Cumbaa
et al., 2013). Enchodus has also been found in the Campanian Pierre
Shale and Carlile Formation on the Etomami River of Saskatchewan
(Cumbaa and Tokaryk, 1999), as well as the Campanian Lethbridge
Coal Zone (Brinkman et al., 2005). Enchodontoids were also widely
distributed in the Tethys Sea, and are found in deposits from the
Cenomanian through Maastrichtian of North America, Europe,
Morocco and the Levant (Fielitz, 2004).
Acanthomorpha, Paracanthopterygii and Polymixiiformes
Acanthomorph shes rst appear in the fossil record in the Albian–
Cenomanian (González-Rodríguez and Fielitz, 2008; González-
Rodríguez et al., 2013) and Cenomanian (Alvarado–Ortega and Than–
Marchese, 2013) of the southern WIS, in Mexico. Farther north, two
acanthomorphs are know from earliest Cenomanian deposits, one from
Utah and the other from the Canadian WIS; respectively, these are
Xenyllion stewarti from the Mowry Formation (Newbrey et al., 2013b)
and X. zonensis from the Fish Scales Formation, formerly part of the
Shaftsbury Formation of Alberta (Wilson and Murray, 1996). Xenyllion
is a paracanthopterygian most closely related to Sphenocephalus from
the late Campanian of Europe (Wilson and Murray, 1996).
Two polymixiiform shes have recently been discovered in the Lac
des Bois (Turonian) locality. Boreiohydrias dayi and Cumbaaichthys
oxyrhynchus are both represented by single specimens and are rare
in the deposits (Murray and Cumbaa, 2013; Murray, 2016). A third,
poorly preserved, acanthomorph specimen from Lac des Bois remains
unnamed and of uncertain afnities (Murray and Cumbaa, 2013).
Despite the presence of acanthomorphs in the northern and southern
WIS in the Campanian, representatives of this group have only been
reported from the Santonian in the middle (U.S.) part of the seaway
(Bardack, 1976; Stewart, 1996).
FRESHWATER FISHES
Numerous micro-fossil sites from Late Cretaceous deposits
preserve an abundance of freshwater sh taxa in southern Alberta (e.g.,
Brinkman and Neuman, 2002; Neuman and Brinkman, 2005). These
are the subject of ongoing research to identify the taxa and interpret
their diversity and distribution.
Two important freshwater localities are known from Maastrichtian
deposits, which document the retreat of the WIS just prior to the end
of the Cretaceous. These are the One-Four locality in southern Alberta
and the Pisces Point locality in central Alberta. Pisces Point, in the
Scollard Formation, preserves articulated material that is only just
starting to be described (e.g., Newbrey et al., 2010b, 2013a; Murray
et al., 2016). Remains of pike have been found (pers. obs.), although
these specimens have not yet been fully prepared or studied. These
Maastrichtian fossils may prove to be related to Esox tiemani, currently
the oldest known member of the genus, from freshwater deposits of
the Paleocene Paskapoo and Ravenscrag formations of Alberta and
Saskatchewan (Wilson, 1980).
Osteoglossomorphs have also been recovered from Pisces Point.
There are at least two species represented, at least one of which is
a basal osteoglossiform. Osteoglossiformes are also known from

259
older, Campanian, deposits; Cretophareodus was the rst articulated
osteoglossomorph described from Cretaceous deposits of Alberta, from
freshwater deposits of the Oldman Formation, near Steveville, in the
southern part of the province. This sh is often excluded from analyses
of relationships because of its poor preservation, but it is considered a
member of Osteoglossidae (Wilson and Murray, 2008).
The Scollard Formation has previously produced articulated
skulls and isolated elements of an amiid, Cyclurus (=Kindleia) fragosa
(Grande and Bemis, 1998). Remains of this sh are found nearby but
just slightly lower in section than Pisces Point (pers. obs.). Additionally,
many isolated bones and teeth that were previously documented from
the Scollard Formation are now being compared with the articulated
material from Pisces Point to better determine their taxonomic identity
(Murray et al., 2016). The disarticulated material includes isolated teeth
of the batoid Myledaphus (Cook et al., 2014). The late Maastrichtian
Hell Creek Formation in Montana has produced freshwater and brackish
hybodonts, orectolobiform and batoid selachians (Cook et al., 2014);
therefore a greater diversity of elasmobranch taxa may eventually be
found on the Canadian side of the border.
The second important freshwater Maastrichtian locality is One-
Four in southern Alberta. Newbrey et al. (2009) reported the presence
of characiform shes from this site. Characiformes are tropical shes,
intolerant of marine waters, so these sh are believed to have moved
through freshwater connections from South America to reach northern
North America during a time of globally high temperatures (Newbrey
et al., 2009). These remains are an important indicator of past climates
and freshwater, and thus terrestrial, connections that existed at the end
of the Cretaceous.
CONCLUSIONS
One of the difculties for palaeontologists in documenting the
faunas of northern Canada is the inaccessibility of some localities.
Those in the far north, such as Lac des Bois, Northwest Territories,
or sites in the high Arctic, require expensive air transport to reach
the sites. Other northern fossils have been found and recovered using
boats, by researchers oating down rivers in northern Alberta to reach
areas inaccessible by road. And yet other new sites are quite easily
accessible in areas that have long been known for dinosaur remains,
but where delicate sh fossils were long overlooked. In the past 25
years, since Russell (1988) published his review of the vertebrate
faunas, there has been a signicant amount of collecting and discovery
of new specimens and localities sampling the Late Cretaceous marine
deposits of the Western Interior Seaway in Canada. This has greatly
increased our understanding of the diversity of shes from the northern
part of the seaway. Many of these shes show clear relationships with
contemporaneous faunas from the western and eastern Tethys, but
others appear to have been restricted to the cooler northern waters.
Active collecting is still on-going and will continue to improve our
understanding of the shes and their environments in the northern
waters of the Western Interior Seaway.
ACKNOWLEDGMENTS
Our thanks to very helpful reviews by Don Brinkman and Rob
Holmes. We would also like to acknowledge the eld work of many
researchers, in particular Don Brinkman, Stephen Cumbaa, Mike
Newbrey, and Matthew Vavrek, which has led to the discoveries of
many new shes as summarized here.
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