Coleoptera fossils: their potential value for dating and correlation of late

Cenozoic sediments
JOHN V . MATTHEWS
,JR.
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Terrain Sciences Division, Geological Survey of Canada, 601 Booth Street, Ottawa, Canada KIA OE8
Received November 23, 1976
Revision accepted for publication April 4,1977

Coleoptera (beetle) fossils play an important role in paleoecological research, but as yet have
contributed little information bearing on dating and correlation. Thereason for this is that most
Quaternary fossils represent extant species, precluding the evolutionary approach to dating,
while the rarity and poor preservation of Tertiary beetle fossils, many of which are from extinct
species, seriously limit their application to stratigraphic studies.
Tertiary beetle fossils recently discovered in Arctic Canada and Alaska are both well preserved
and abundant. Most of them represent extinct species that are closely related to living forms,
hence they have potential stratigraphic value. In one case treated herein comparison of fossils of
an Alaskan Tertiary species with those of a related species from the Beaufort Formation on
Meighen Island (Canadian Arctic Archipelago) implies that the latter sediments were deposited
less than 5.7 Maago. However, this conclusion requires testing because it is at odds with the date
on Meighen Island exposures reached by study of fossil plants. I submit that further study of the
insect fossils from the Beaufort Formation and other late Tertiary sites will help resolve such
problems of dating and correlation.
Quaternary beetle fossils have stratigraphic value even though fragments of that age represent
For personal use only.

for the most part only existing species. For example, it has been shown that late Pleistocene
fossils of stenothermal Coleoptera species can provide a sensitive record of climatic change, and
thus such fossils may be used for site to site correlation in areas where climatic history is well
documented. In exceptional cases beetle fossils appear to provide a more accurate basis for
correlation than even fossil pollen.

Les coleopteres fossiles jouent un r6le important en recherche paltocecologique mais jusqu'a
maintenant n'ont fourni que peu d'information utile B la datation ou la correlation. La raison de
cela est que la plupart des fossiles du Quaternaire representent des especes encore en existence,
ecartant ainsi I'approche evolutionniste a la datation, alors que la rarete et la mauvaise preserva-
tion des coleopteres fossiles du Tertiaire, dont plusieurs especes sont eteintes, limite serieuse-
ment leur application aux etudes stratigraphiques.
Les coleopteres fossiles du Tertiaire decouverts recemment dans 1'Arctique canadien et en
Alaska sont B lafois bien preserves et abondants. Laplupart d'entre eux representent des especes
eteintes qui s'apparentent de pres aux formes vivantes; ainsi ont-ils une valeur stratigraphique
potentielle. Dans un cas, traite dans cet article, la comparaison des fossiles d'une espece
provenant du Tertiaire d'Alaska avec ceux d'une espece parente dans la formation de Beaufort
sur I'ile de Meighen (archipet Arctique) indique que ces derniers sediments se sont deposes il y a
moins de 5.7 Ma. Toutefois, cette conclusion requiert verification puisqu'il y a conflit d'gge avec
les donnees de plantes fossiles provenant des affleurement de I'ile de Meighen. I1 m'apparalt que
des etudes supplementaires sur les insectes fossiles ide la formation de Beaufort et d'autres sites
de la fin du Tertiaire aideront B resoudre de tels problemes de datation et de correlation.
Les coleopteres fossiles du Quaternaire ont une valeur stratigraphique msme si les fragments
de cet 5ge ne representent pour la plupart que des especes vivantes. Par exemple, il a ete
dtmontre que parmi les fossiles de la fin du Pleistocene, des especes stenothermique de coleop-
teres peuvent fournir une indication sensible des changements climatiques et ainsi on peut utiliser
de tels fossiles pour des correlations d'un site a un autre dans les regions ou I'histoire climatique
est bien documentee. Dans des cas exceptionnels, les coltopteres fossiles semblent fournir une
base plus precise,de correlation que le pollen fossile.
[Traduit par le journal]
Can. J. Earth Sci., 14,2339-2347 (1977)

Introduction water inhabitants. This explains the rarity of

Unlike most invertebrate groups, insects are insect fossils because the geologic record is
now, and have been from their first appearance dominated by marine rather than terrestrial
in the Devonian, primarily terrestrial and fresh- sediments. It should, therefore, come as no sur-
 2340 CAN. J. EARTH SCI. VOL. 14, 1977

prise that the insects have contributed little or

nothing to stratigraphic studies-the traditional ARCllC
domain of invertebrate fossils. OCEAN
Well preserved fossils of Tertiary beetles, now
being recovered from arctic sites in North Amer-
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ica, may change this situation. Many of the fossils

represent extinct species closely related to mem-
bers of the existing fauna, and thus provide the
possibility for construction of reasonably detailed
phylogenies. When this is done, the fossils
themselves become potentially useful for pur-
poses of correlation and dating.
Tertiary insect fossils are rather rare, but insect
fossils, particularly those of beetles (Coleoptera)
are very abundant in Quaternary sediments. The
value of these Quaternary beetles for paleo- FIG. 1. Localities mentioned in text. Stippled zone
ecological studies has been well documented marks Beaufort Formation.
(Coope 1970); but because they represent extant
species, i.e. there has been little change at the their fossils will take on added stratigraphic
specific level during Quaternary time (Coope significance.
1970), they provide little evolutionary data appli- One example, already at hand, illustrates the
For personal use only.

cable to stratigraphicproblems. On the other hand way in which such fossils might be used for
Pleistocene fossils do have stratigraphic implica-correlation. The specimens involved come from
tions when whole assemblages of them reflect two widely separated sites: Lava Camp in western
known climatic perturbations. Thus they have Alaska and exposures of the Beaufort Formation
been used for correlation of late Pleistocene on Meighen Island in the Canadian Arctic
sediments, and in some exceptional instances the Archipelago (Fig. 1).
beetle fossils portray the changes in a known The Lava Camp site and its fossils were dis-
climatic sequence with greater accuracy and less cussed initially in Hopkins et al. (1971) and
time lag than fossil pollen, the best known means more recently in Matthews (1976b). One of the
of nonradiometric Pleistocene correlation. In the most significant aspects of the site is that a
following sections several examples show how radiometric date of approximately 5.7 Ma1 is
fossil beetles have been used and might be used associated with the insect fossils (Hopkins et al.
for dating and correlation of late Cenozoic 1971). The flora from the site is similar to those
sediments. representing the Clamgulchian Stage2 of southern
Alaska, while the K-Ar date and paleomagnetic
Late Tertiary Examples data show that deposition of the fossiliferous
Whereas most identifiable Pleistocene beetle sediments coincided with the short reversed
fossils can be assigned to existing species, many phase of Late Miocene Paleomagnetic Epoch 5
of the ones now coming from late Tertiary sites (Berggren and Van Couvering 1974; Matthews
in Arctic Canada and Alaska fall outside the 1976b).
range of variation of extant species (Matthews The Beaufort Formation crops out over large
1976a). This is true even of those fossils which areas of the western islands of the Canadian
refer to well documented, nondiverse, and mor- Arctic Archipelago (Tozer 1956; Thorsteinsson
phologically circumscribed genera; so it is -

unlikely that these Tertiary fossils represent lm.y. = Ma in SI units.

undiscovered or undescribed existing species. ZThe terms Seldovian, Homerian, and Clamgulchian
Instead most of them represent extinct species, refer to Alaskan time-stratigraphic units, defined on the
although many seem to be very closely related to basis of pollen and plant macrofossils (Wolfe et al. 1966).
The Seldovian Stage represents Middle Miocene; the
species in the present fauna (Matthews 1976~). Homerian, Middle and Late Miocene (before 8 Ma BP-
As the Tertiary evolutionary history of the more Triplehorn et al. (1977)), and the Clamgulchian, Late
commonly preserved beetle genera is established, Miocene (post 8 Ma) and Pliocene.
 MATTHEWS 2341
a. b.
and Tozer 1970) (Fig. 1) and as isolated outliers
on some of the eastern islands (Balkwill and
Bustin 1975; Wilson 1976). Beaufort Formation
sands with intercalated marine clays and peaty
I
Htm-"tuJ

\
Hm~omnsls

eeee0400
2.

P
3
Hcmp*l

* /
5%, ,',I
:;,

zones are exposed over the entirety of Meighen

vo0 8" **-- -0- T(
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Island (Thorsteinsson 1961; Matthews 1974a, 4

*
C
.'-

1976~).The peats have yielded well preserved *.

plant macrofossils and beetle fragments (Hills *A
\
*'.

/ \
and Matthews 1974; Matthews 1974a, 1976~). C. d
Unfortunately, even though the Beaufort Form- 3. z 3.
*'
ation is reliably assigned to the late Tertiary, its , ;, *'
age is not as precisely known as is that of the ',
*, ,'
I'

*
8
'

Lava Camp locality. On the basis of plant macro- ,'h.

* *** ,#'#

fossils it has been referred to the Seldovian and f %

Homerian stages (Hills and Ogilvie 1970; Hills
et al. 1974: Hills and Bustin 1976J2
Some of the beetles from ~ i v aCamp are
congeneric with specimens from the Beaufort
Formation on Meighen Island, and recently
(Matthews 19763) several of these fossils have
been studied in detail. All of them belong to the
subgenus Cyphelophorus of the water beetle
For personal use only.
' /
*-. ** *+*
.* f
*. a==
FIG. 2. Alternate hypotheses on the phylogenetic
relationships of three beetle species. Helophorus coopei
Matth. and Helophorus meighenensis Matth. are extinct.
The former is from the Lava Camp site in Alaska; the
latter from Beaufort Formation on Meighen Island (see
Matthews (19766) for details). Dashed narrow lines
portray theoretical relationships based on elytral struc-

genus Helophorus, but the species from Lava
Camp (Helophorus coopei Matth.) appears to be
niore primitive than the one from Meighen
Island (H. meighenensis Matth.), and with respect
to elytral tubercles neither species is as structur-
ally advanced as Helophorus tuberculatus Gyll.,
the single extant member of the subgenus.
Although there are several possible interpreta-
tions of the evolutionary implications of the two
fossil and one extant species (Fig. 2), the one
preferred at the present time (Matthews 1976b)
is that the three form stages in the evolution of
one lineage (Fig. 2d). If so, then the fossils imply
that Beaufort Sediments on Meighen Island are
younger than 5.7 Ma (Clamgulchian) in age and
furthermore that Beaufort deposition was dia-
chronous since the more southern exposures on
Banks Island are undoubtedly older than 5.7 Ma
(Hills et al. 1974). Reconstructions of phylogenies
of other beetle taxa from Lava Camp and Beau-
fort Formation assemblages will be one means
of testing this conclusion. As with the case of
Cyphelophorous several of the fossils in those
assemblages represent taxa particularly suited
for phylogenetic analysis because of their low
diversity and certain preservable morphological
structures.
The known age of the Lava Camp beetle
fossils means that in the future they will continue
to be significant for long range correlation of
tures. Darkened portion of each species symbol in b, c, d,
indicates possible geological range of the species. Geologi-
cal ranges of species under alternative a are fixed by the
methodology that it reflects (see below). In b, c, d, species
are identified by number (see a for species name).
(a) Interpretation based on 'phylogenetic systematics'
principles (Hennig 1966; Griffiths 1972). Two hypo-
thetical ancestor species are required and each branching
results in two new species. (b) H. meighenensis arises as
a peripheral isolate of H. coopei; H. tuberculalus evolves
from H. meighenensis in the same manner. H . coopei
(most primitive species) could persist as a relict and be
a contemporary of H. meighenensis as for example at
time T I . (c) Same as b except H. tuberculatus envolves
directly from H. meighenensis (an example of phyletic
gradualism). H. coopei could persist after the first appear-
ance of H . meighenensis. (d) All three species are chrono-
species within one lineage. This option is favoured
(Matthews 19766) because it is the simplest explanation
of the facts at hand. Alternatives b or c agree better with
paleobotanical evidence for the age of the Beaufort
Formation (see text), but the paleobotanical evidence is
interpreted as in d.
late Tertiary sediments. But the fossils have an
equally valuable, though less spectacular, role
in local correlation. For example, beetle fossils
occur in organic zones within terrace alluvium
at two localities on the Kugruk River, approxi-
mately 25 km east of Lava Camp. Both exposures
were thought on stratigraphic and geomorphic
grounds to be of late Tertiary age (D. M. Hop-
kins, personal communication, 1973) but a
 2342 CAN. J. EARTH SCI. VOL. 14. 1977

TABLE1. Preliminary list of Coleoptera fossils from TABLE

1 (Concluded)
Kugruk River sites
Fossil
Fossil localities"
localities"
Taxa 1224 1226
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Taxa 1224 1226

Scolytidae
Carabidae Trypodendroncf. T. lineatum ( O l i ~ . ) ~ x
Carabus truncaticollis Eschzb x Scolytus cf. S. piceae S W . ~ x
Pelophila cf. P. borealis Payk. x Pityophthorus ~ p . ~ x
Notiophilus cf. N. aeneus Herbst.b-c,d x Orthotomicus cf. 0 .caelatus ( E i ~ h o f f ) ~ x
Notio~hilusSD. x
~ l a ~ h ; sp.
us' 'Location: 122+Kugruk River, north bank, 1.6 km downstream
Blethisa catenaria Brown from mouth of Ch~cagoCreek (6S054'20" N ; 162"28'12" W), Alaska.
Diacheilapolita Fald. 1226Approximately 1.4 km upstream from mouth of Relndeer
Creek (65"50'40" N ; 162'25'58'' W), Alaska. Both samples collected
Asaphidion cf. A. alaskanurn Wick. by R. Rose Union Oil Co. o f Calrforn~a,Anchorage, Alaska.
Asaphidion sp. bNamed ipecies or a closely related one represented in the Lava
Camp assemblage (see text).
Bembidion cf. B. pseudocautum Lth. CN. aeneus is currently restricted to eastern North America (Fig. 3).
Bembidion spp. 'Not known from Alaska and in general having the northern limit
across southern and central Canada.
Agonum sp. eFossils from Lava Camo * mav - remesent
- a soecies ancestral to this
Pterostichus (Cryobius) spp. one.
fIdentified by D . Bright, Biosystematics Research Institute, Canada
Pterostichus cf. P . corvinus Dej.b~d Department of Agriculture, Ottawa, Canada.
P. haematopus Dej.
P. sublaevis Sahlb. preliminary examination of the beetle fossils
Amara alpina Payk.
Amara sp. (Table 1) suggests a different conclusion. One
For personal use only.

Trichocelluscf. T. mannerheimi assemblage (1226, Table I), like that at Lava

R.F. Sahlb. Camp (Hopkins et al. 1971; Matthews, un-
Dytiscidae published results), contains fossils of extinct
Colymbetes sp. species, many of which are most closely related
Gyrinidae to extant species with northern distributional
Gyrinus sp. limits far south of the Seward Peninsula. The
Hydrophilidae other assemblage (1224, Table l), is much more
Helophorus cf. H. splendidus J. Sahlb. diverse, but most important the majority of
Hydrobius sp. fossils refer to extant species, many of which are
Georyssidae living today in forest-tundra regions near Lava
Georyssus sp.b*d Camp. Thus the beetle fossils show that the two
Staphylinidae Kugruk River sites are of different age. One
Omalinae certainly contains late Tertiary sediments, though
Oxytelinae
Bledius sp.
possibly somewhat younger than those at Lava
Camp, while the other probably represents a
Aleocharinae
Gymnusa sp.
Pleistocene intergla~ial.~ In the western part of
the Arctic Archipelago it is sometimes difficult
Scarabaeidae to distinguish Beaufort Formation from Quater-
Aegialia sp.
nary sediments. There, as in the Alaskan example,
Byrrhidae
Morychus? sp.
Coleoptera fossils are of value, for the overall
Simplocuria sp. composition of Beaufort assemblages is quite
Elateridae
different from those of Quaternary age.
In cases such as the two mentioned above,
Lathridiidae part of the reasoning concerning correlation or
Stephostethus sp.
lack of correlation relies on the fact that certain
Chrysomelidae genera have become locally extinct during the
Donacia sp.
course of Cenozoic climatic change. Extinctions
Curculionidae
Lepidophorus lineaticollis Kirbye x -
3An interglacial age for sample 1224 (Table 1) is also
Vitawitus thulius Kissingerb X
suggestedby plant macrofossils such as conifer seeds and
Grypidius cf. G . equiseti Fab. X
one fruit of Alisma (cf. A. plantagoaquatica L.), a genus
Notarius sp. X
which presently does not occur within 1500 km of the
Lepyrus sp. X
fossil locality (Hulten 1968; Matthews 19746, p. 833).
 MATTHEWS
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FIG. 3. Distribution of some species of beetles and
For personal use only.
other insects (Cicadellidae = leafhoppers) which are
either represented in the fossil assemblages from the two
Tertiary sites or are closely related to extinct species in
those assemblages. In all cases the indicated taxa are
presently extinct in northern North America, hence the
fossils at Lava Camp and Meighen Island mark a
particular stage in the evolution of the North American
fauna.
of a more regional extent may also have strati-
graphic significance. For example, among the
Lava Camp fossils are several heads very similar
to those of the Asian ground-beetle species,
Dromius rufficollis Motsch., as well as fragments
of a species closely related to or conspecific
with Notiophilus aeneus Herbst, a ground beetle
presently confined to eastern North America
(Fig. 3). Both species are extinct over the entirety
of northwestern North America. It may already
be confidently assumed that any Alaskan assem-
blage containing fossils of either is at least as
old as early Pleistocene, and the value of fossils
of these species for establishing the minimum
age of sediments can only increase as the time of
their regional demise becomes better known.
The same rationale applies to the Meighen
Island Beaufort Formation assemblages, which
likewise contain a few fossils of Palearctic or
geographically isolated North American insects
(Fig. 3).
Quarternary Examples
One of the triumphs of the initial work of
G. R. Coope in England, in fact the very basis
2343
for use of fossil beetles in paleoecological studies,
was the discovery that the rate of evolution of
Coleoptera has been very slow. Even early
Pleistocene fossils are referable to extant species
(Matthews 1974~)and some existing species
extend well back into the late Tertiary (Matthews
1970, 1976~).This means that the evolutionary
approach to dating and correlation cannot be
used with Pleistocene beetle fossils. A possible
exception involves the rove beetle species
Tachinus apterus Makl. Figure 4 shows the
differences in flight wing development and
elytral proportions of T. apterus elytra from
two assemblages, one early Pleistocene in age
and the other late Pleistocene, at Cape Deceit
in western Alaska (Matthews 1974~).The differ-
ence in elytral size seems to represent an intra-
specific evolutionary trend, and if so T. apterus
fossil elytra, which occur at a number of Alaskan
and northwest Canadian sites, could have chron-
ological ~ignificance.~ Intraspecific evolution has
undoubtedly occurred in other Coleoptera line-
ages during the Quaternary, but it is unlikely
that many such cases will be documented since
it is seldom that numerous well preserved fossils,
like those of T. apterus, are found in independ-
ently dated early and late Pleistocene contexts
at a single site. Thus, in practice, the evolution-
ary approach using fossil beetles will probably
not be an important tool for dating Pleistocene
sediments.
Recent studies in England (Ashworth 1972;
Coope and Brophy 1972; Osborne 1974) show,
however, that stratigraphic and chronologic
4Complicating this statement is the fact that some
authors (e.g. Ullrich and Campbell 1974) consider
T. apterus to be a complex of two very similar species
(T. apterus Makl. and T. brevipennis J. Sahlb.), neither of
which can be distinguished by elytral characters alone.
I believe that T. brevipennis is more likely a subspecies of
T. apterus; however, if it is a distinct species then it must
be noted that other diagnostic Tachinus fragments from
the Cape Deceit site show that both T. brevipennis and
T . apterus are probably represented by the elytra in the
late Wisconsin sample (Fig. 4). Only fossils of the T.
apterus type have been found in the early Pleistocene
assemblage. These facts mean that the diagram in Fig. 4
may not portray temporal change in elytral size of a
single species as suggested. However, unless all fossil
elytra of T. apterus (sensu Ullrich and Campbell 1974)
in the late Wisconsin sample fall within the overlap zone
of late Wisconsin and early Pleistocene elytral dimension
fields (Fig. 4), the trend mentioned above still applies,
and it is highly probable if, as I believe, T. brevipennis is
only a mainland subspecies of T. apterus (see distribution
of T. apterus and T . brevipennis in Ullrich and Campbell
(1974)).
 CAN. J . EARTH SCI. VOL. 14, 1977

Flight Wings
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T apferus
For personal use only.

l i0 112 1.4
E l y t r a l l e n g t h(mm)
FIG.4. Elytral and flight wing reduction within the Tachinus apterus Makl. lineage during the
Pleistocene at Cape Deceit in western Alaska. Stippled and hatched zones represent the fields for
elytral measurements from the two samples. Dark lines and bars on vertical and horizontal axes mark
mean and first standard deviation for each field. Dashed lines = regression of length on width for
each assemblage. Note smaller size of late Pleistocene fossils, a trend which seems to be correlated
with flight wing reduction during the Pleistocene.

information may result from study of the tax-
onomic composition of Pleistocene assemblages.
Coope and Brophy, for example, were able to
establish faunal units based on the presence of
certain stenothermal species, and the succession
of these units provides a detailed picture of late-
glacial climatic change. Of added significance is
that the beetles apparently responded more
quickly to late-glacial climatic change than
plants, hence the climatic succession based on
evidence of beetle fossils is more accurate than
that based on pollen. In other words the beetle
fossils provide a better basis for site to site
correlation. The use of Coleoptera fragments in
this manner is dependent on one important
prerequisite-a sound knowledge of the tax-
 MATTHEWS
onomy and distribution of species in the con-
temporary fauna. Workers in western Europe
possess such information; we in North America
are less fortunate because large gaps remain in
our understanding of the present Nearctic
Coleoptera fauna. For this reason the use of
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Pleistocene beetle fossils for correlation and
dating of North American sediments is, at
present, a blunt instrument and only very crude
estimates of age are possible. For example,
beetle fossils identified by the author from peats
on Bathurst Island (Arctic Archipelago) (Fig. l),
where beetles do not live today, suggest only
that the sediments from which they come are of
interglacial age (Blake 1974). Nevertheless, this
is an important conclusion because previous 14C
dates on these peats allowed the possibility that
they could be of either interstadial or inter-
glacial age. The beetle fossils imply a climate
warmer than that of the present; hence, an
interglacial. Similarly, fossils of the ground-
beetle Oodes americanus Dej. from peat formerly
For personal use only.
presumed to be of interstadial age on one of the
Magdalen Islands (Gulf of St. Lawrence) show
instead that the peat is more likely of Sangamon
interglacial age (Prest et al. 1976).
Discussion
Remarks made above and elsewhere (Coope
1970, 1975) indicate that fossil beetles, unlike
fossil mammals, have only limited value for
dating and correlation of Pleistocene sediments.
The most promising cases are those in which
intraspecific evolution can be demonstrated or
where changes in the composition of fossil
beetle assemblages have accurately and quickly
tracked chronologically documented climatic
fluctuations (Coope 1975).
The composition of fossil beetle assemblages
of Tertiary age may also provide information
of value for correlation, and the fact that certain
extant species not present in North America
today are known to have lived or had close
relatives living there during the Tertiary, augurs
well for use of fossils of such beetles as indices
of minimum age. Some recently discovered
Tertiary fossils probably represent totally extinct
species. As their evolutionary relationships and
temporal ranges are defined, fossils of such
species will take on increased stratigraphic value.
This report mentions as one example some
fossils of the subgenus Cyphelophorus and their
2345
potential value for dating northern exposures
of the Beaufort Formation. The conclusion
reached-that Beaufort sediments on Meighen
Island are younger than those from the Alaskan
Lava Camp site, hence younger than the Alaskan
Homerian Stage-is at odds with that implied
by the recent discovery of cones of the extinct
spruce, Picea banksii Hills and Ogilvie, on
Meighen Island (Hills and Ogilvie 1970; Hills
and Bustin 1976). Picea banksii is thought to be
the direct lineal ancestor of white spruce (Picea
glauca), and since fossils of the latter species are
present at the Lava Camp site, Meighen Island
Beaufort exposures are assumed (Hills and
Ogilvie 1970; Hills and Bustin 1976) to be older,
not younger, than Lava Camp. This discrepancy
could result from the fact that fossil cones of
P. banksii from Meighen Island occur at a level
stratigraphically lower than the unit yielding the
critical Helophorus fragments; however, I doubt
that the age difference of Meighen Island expo-
sures is that great. Thus either my interpretation
of Helophorus (Cyphelophorus) phylogeny is in-
correct or Picea banksii is not the immediate
ancestor of P. glauca. It should be noted that
both arguments are founded on the acceptance
of 'phyletic gradualism' (evolution within one
lineage) as a possible mode of speciation and
evolution. Several recent papers (Schaffer et al.
1972; Eldredge and Gould 1972) have questioned
the reality of gradual phyletic change. Gingerich
(1976), on the other hand, provides a convincing
paleontological case for the importance of both
phyletic gradualism and branching (cladism) in
evolution. Recognizing these various arguments,
I have attempted, while favouring one particular
explanation of Cyphelophorus phylogeny, to
provide several possible alternative explanations.
Obviously, the same alternatives should be con-
sidered in attempts to decipher the relationship
of Picea banksii and other spruce species.
These remarks are not intended as a critique
of the paleobotanical approach to dating of the
Beaufort Formation, but only to show that
there now exists an opportunity to check such
correlations using insect fossils. Such fossils,
specifically those of beetles, will probably come
to play an ever more important role in dating
of the Beaufort Formation and other northern
Tertiary sequences. At such sites Coleoptera
fossils are less abundant than palynomorphs, but
unlike palynomorphs many of them can be
 2346 CAN. J. EARTH S(21. VOL. 14. 1977
identified to the specific level, and they are not
as likely to have been rebedded from older units.
Beetle fossils representing more than 66 genera
and 87 species have been recovered at Beaufort
Formation exposures on Meighen Island, show-
ing that Coleoptera assemblages from that
Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 04/24/20
region are also more diverse taxonomically than
assemblages of pollen or plant macrofossils.
Moreover, many of the fossil beetles from
Meighen Island, Lava Camp, and other Tertiary
exposures represent widely distributed taxa,
furthering the possibility of their use in long
range correlation.
Acknowledgments
I have profited from discussions with L. V.
Hills (University of Calgary), G. R. Coope
(University of Birmingham), and G. E. Ball
(University of Alberta). Thanks are extended to
D. M. Hopkins for permission to mention un-
published samples and to the staff of the Coleop-
tera Unit, Biosystematics Research Institute, Ot-
For personal use only.
tawa, for continuing help in identifying Coleop-
tera fossils and for allowing access to collections.
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