Evolution of the subgenus Cyphelophorus (GenusHelophorus, Hydrophilidae,

Coleoptera): description of two new fossil species and discussion of

Helophorus tuberculatus Gyll.
JOHNV. MATTHEWS,
JR.
Terruin Sc4irnc.esDivision, Geologic,al Survey of Canada, 601 Booth Street, Ottawa, Canada KIA OE8
Received December 1, 1975

MATTHEWS. J. V., JR. 1976. Evolution of the subgenus Cyphelophorus (Genus Helophorus,
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Hydrophilidae, Coleoptera): description of two new fossil species and discussion of

Helophorus rubercrrlatus Gyll. Can. J . Zool. 54: 652-673.
Exposures of late Tertiary sediments in the North American Arctic contain well-preserved
fossils of Coleoptera (beetles). Two sites, Lava Camp in western Alaska and locality 2-73 on
Meighen Island (Canadian Arctic Archipelago), have yielded fragments referable to two fossil
species of the helophoran subgenus Cyphelophorus (Hydrophilidae). The only extant species in
the subgenus, Helophorus ruberculatus Gyll., is characterized by a distinctive array of elytral
tubercles, the intraspecific variation of which is documented here as an aid for distinguishingH .
tuberculatus from the two fossil species.
The two fossil species are obviously related to H . tuberculatus and probably represent
sequential species in the lineage leading to H . tubercularus. If so, they provide information on
evolutionary trends within the Cyphelophorus lineage, and when examined in the light of the
morphological features of species in other Helophorus subgenera, show that the tuberculate
elytron seen in H . tuberculatus is probably a derived character that developed by modification of
an elytron initially having raised alternate interstices.

MATTHEWS, J. V., JR. 1976. Evolution of the subgenus Cyphelophorus (Genus Helophorus,
Hydrophilidae, Coleoptera): description of two new fossil species and discussion of
Helophorus tuberculatus Gyll. Can. J . Zool. 54: 652-673.
For personal use only.

On a decouvert des fossiles bien conserves de coleopteres dans des sediments de la fin du
tertiaire, dans 1'Arctique nord-americain. Deux sites, le camp Lava dans I'ouest de 1'Alaska et la
localite 2-73 de I'ile de Meighen (Archipel arctique canadien) ont fourni des fragments apparten-
ant a deux esp&cesfossiles d'Helophorus, du sous-genre Cyphelophorus (Hydrophilidae). La
seule espece actuelle du sous-genre, H . tuberculatus Gyll., est caractkriste par I'arrangement
particulier des tubercules de I'elytre dont on examine ici la variation intraspecifique, afin de
faciliter la distinction entre H . tuberculatus et les deux especes fossiles.
Les deux espbces fossiles sont sans contredit apparentees 21 Helophorus tuberculatus et
representent sans doute une sequence dans la lignee conduisant 21H . tuberculatus. Si I'on accepte
cette hypothese, ces especes nous renseignent sur les tendances ivolutives de la lignCe de
Cyphelophorus et, si on tient compte des structures morphologiques des espbces appartenant a
d'autres sous-genres d'Helophorus, ellesdemontrent que I'ilytre tuberculi d'H. tuberculatus est
probablement un caractere CvoluO, apparu a la suite d'une modificationd'un elytre posstdant au
depart des interstices surCleves en alternance avec des interstices ordinaires.
[Traduit par le journal]

Introduction course, those preserved in amber are an excep-

The value of fossil beetles (Coleoptera) for
Quaternary paleoecological research is well
established (Coope 1970). One finding of such
studies in Europe (Coope 1970) and North
America (Matthews 1974a) is that evolution of
Coleoptera during the Pleistocene has been
negligible, implying that only by examination
of late Tertiary fossils will it be possible to
document, paleontologically, the latest episodes
of Coleoptera evolution. Unfortunately, be-
cause of poor preservation, most Tertiary beetle
fossils are inadequate for this purpose. Of
tion, but they contribute only minor information
because the occurrence of amber is so sporadic
and the beetle fossils contained in amber often
represent specialized groups of minute species,
many of which are not taxonomically well
understood in the extant fauna. Most reported
fossils of Tertiary beetles are impressions or
casts in fine-grained sediments, and although
in rare instances these yield detailed information
(e.g. Gersdorf 1969, 1971), more often they are
too poorly preserved to reveal critical structures
(e.g. Darlington 1967).
 MATTHEWS 653

A R C f l C
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A L A S K A
For personal use only.

C A N A D A

Fro. 1. Location map with sites mentioned in text. Locality 2-73 on Meighen Island is situated
approximately at the intersection of the 100th meridian and the line representing 80" N latitude.

My interest is in documenting the develop- distinctive hydrophilid beetle. It is ideally

ment and evolution of the Boreal and Arctic
flora and fauna, but to do this for Coleoptera,
fossils as well preserved as those originating
in Quaternary sediments must be available.
Several late Tertiary sites from Arctic North
America are currently yielding such fossils. One
i s the late Miocene Lava Camp locality in
western Alaska (Hopkins ei a/. 19311, and the
others are various exposures of the late Tertiary
Beaufort Formation on Banks Island and Meig-
hen Island in the Queen Elizabeth group of the
Canadian Arctic Archipelago (Matthews 19746;
Fig. I).
This paper, one of a projected series on late
Tertiary Arctic Coleoptera, deals with fossils
related to the species Helophorus (Cyphelo-
phorus) tuberculatus Gyll., a rare and very
suited for study because the characters by which
it may be recognized are those most likely to be
preserved on fossils. A detailed description is
given here of H. tuberculatus to facilitate com-
parison with, and description of, two un-
doubtedly extinct species of Helophorus from
the Lava Camp sediments in Alaska and a
Beaufort Formation locality on Meighen Island
(Fig. 1). When compared with H. tuberculatus
Gyll., the two newly described species seem to
form an evolutionary sequence within a single
lineage.
Eldredge and Gould (1972) point out that
paleontologists bring a "peculiar perspective to
the study of life," or at least one different from
that of neontologists. Nowhere is this more
obvious than when dealing with late Tertiary
 654 CAN. J. ZOOL. VOL. 54, 1976
insect fossils. for while it is clear to me that such
fossils are critical for construction and testing
of phylogenies, the meager fossil record of most
insect groups dictates that entomologists wishing
to decipher evolutionary history use methods
(e.g, phy logenetic systematics) whose implica-
tions appear to deny the usefulness of fossils.
Because late Tertiary beetle fossils from western
Alaska and Arctic Canada are so clearly related
(in some cases, conspecific) to extant species,
any attempt to reconstruct evolutionary history
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using the fossils must be made with the help
of the information revealed by extant species.
I have done this to assess the relationshi~sof
Cyphelophorus species, but the conclusion
reached, i.e. that all three species are members
of one lineage, is unacceptable under the pre-
cepts of phylogenetic systematics (Hennig 1966;
Brundin 1972; Cracraft 1974) or other cladistic
approaches to phylogeny (Eldredge and Gould
1972). However, it does seem the best explana-
tion of the facts at hand and, if valid, reveals
several evolutionary trends.
Methods
Specimens Examined
Heluplrorrrs t u b e r c t ~ l o ris~ a rarcly collected species,
For personal use only.
henm a large series or specimens is not available, For
this study I have examined 86 specimens of N. t d r r -
crrlurr#s, most of which are From North America but
a few are from northern and central Europe. In addition
I have examined a large series of Helophorus (Gephelo-
phorus) sibiricus Motsch., three specimens of the east
Asian species, H. (Gephelophorus) auriculatus Sharp, as
well as specimens of H. (Empleurus) nubilus F., H .
(Orphelophorus) arcticus Brown, H . (Orphelophorus)
obscurellus Popp., H. (Meghelophorus) grandis Illiger,
and Helophorus s.str.
Morphology
A number of special terms are used with reference to
morphological features of Helophorus. The traditional
ones and several new ones are illustrated in Fig. 2.
As with the pronotum of other Helophorus species,
that of H. tuberculatus possesses seven longitudinal
grooves separated by + raised intervals. The central or
median groove i s separated from the submedian grooves
by t h e internal intervals (Fig, 2). The submedian and sub-
marginal grooves delimit the middle intervals, while the
external pronotal interval is bordered by submarginal
and marginal grooves. The shape and width of the
suprapleural region located betwcen the notopleural
sutures and tbe edge of the prothorax (viewcd from below.
Fig. 2) are often important characters for distinguishing
some species and subgenera of Helophorus (Angus
1970~).
In diagnoses of Helophorus subgenera, the morpho-
logical characters most often cited are those of the
elytron. It possesses 11 coarsely punctate, complete
striae; the outer one is poorly developed and crowded
toward the true elytral margin. I n some species an inter-
mlary stria is present near the base of the elytron
betwen stria 1 (st. 1) and stria 2 (st. 2). The elytral region
bordered by st. 1 and the suture i s termed the sutural
interstice (int.);' the others are numbred consecutively
starting with int. 1, &tween st. I and st. 2. In H. tuber-
culatus, and at least one other species,ints. 2,4. 6, and 8
possess discontinuous ridga and (or) isolated tubercles.
The relative position of these features is more or less
constant and is designated here according to the inter-
stice in which they w u r , and their serial position from
the clytral base. Each elytral interstice has a median row
of small punctures often doubled or tripfed on the ridges
and lubercles. The 10th interstice is carinate in many
Helophorus species, including H. trrb.prcrr/otu~,and in
some specimens protrudes laterally more than the
epipleuron to form a pseudepipleuron (Fig. 2). A broad
impression trending obliquely rrom the suture toward
the elytral shoulder, and evident at least to int. 6, is
present in the basal one-third of the elytron of H.
rrrhercelurrrs. For illustrations of other morphological
terms see Fig. 2.
H. r~~her~rrlatus specimens tend to become covered in
life by an encrusted laycr which must be removed before
the morphological features mentioned above and shown
in Fig. 2 can be adequately observed. Cleaning is easily
accomplished in an ultrasonic bath or by careful use of a
brush and sudsy water.
Mensural Characters
Most taxonomic treatments of Coleoptera cite measure-
ments which are of little value for comparison with
fossils because they pertain either to the dimensions of
the entire, articulated insect or to parts such as tarsi and
eyes that are seldom preserved intact. Mensural char-
acters used here were chosen so as to be more readily
comparable with fossils, and therefore they apply to
those parts most often preserved (heads, pronota, and
elytra) and those dimensions that arc least likely to be
influcnced by the distortions of shape that often char-
acterize fossils. Figure 2 illustrnter the dimensions mea-
sured for this report. H. rrrb~rcirlarusspecimens have a
pronounced apicaI declivity on the elytron, causing the
mmsured length of the elytron to vary with its orienta-
tion. To alleviate this problem all elytral measurements
cited here were taken with theelytron similarly orientated,
j.e. with h t h endpoints of the elytron in the same plane
.
of focus at a magnification of 50 x The resultant figure
is somewhat larger than would be obtained if the basal
half of the elytron was oriented normal to the viewing
axis, as might s e m ro be proper for pinnetl museum
specimens, but it more closeIy approximates the length
of a fossil that has the elytron somewhat flattened,
like some of those discussed here.
Helophorus tuberculatus Gyll.
Fossils representing a number of genera and
families of Coleoptera are now known from late
'I follow Angus in using the term "interstice" rather
than "interval" to avoid confusion with the pronotal
"intervals."
 MATTHEWS

HEAD FRONT VIEW

Y - s h a p e d groove

coronal rldge
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FIG.2. Morphological terms associated with Helophorus. Many of the terms are common to all
species of Helophorus, but the drawing represents H . tuberculatus Gyll. See text for remarks on
measuring length of the elytron.
 656 CAN. J. ZOOL , VOL. 54. 1976
Tertiary sediments in northern North America
(Hopkins et al. 1971; Matthews 1970b, 1974b),
and even though many of these seem to refer to
undescribed, possibly extinct, species, not all are
suited to the detailed evolutionary. type ~- of
analysis attempted here. For that purpose it is
desirable that the extant group to which the
fossils are related be (1) nondiverse taxonom-
ically to lessen the phylogenetic complexity
caused by cladogenesis; (2) easily distinguished
by external morphological characters likely to
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be preserved on fossils; (3) widely distributed;
and (4) not restricted to a specialized or rare
type of habitat. Helophorus (Cyphelophorus)
tuberculatus Gyll. meets these requirements
better than most beetle species. It ii very dis-
tinctive, at once recognized by features of the
elytron, and on this basis is placed in its own
subgenus. Though a rarely collected beetle, it is
Holarctic in distribution and inhabits the types
of local environment-poorly drained peaty bog
and riparian sites-that commonly occur in the
boreal regions of North America, Europe, and
Asia. Moreover, it is just such peaty sediment
that is most likely to be buried, preserved, and
later exhumed to be exposed in sedimentary
sequences.
For personal use only.
Because H. tuberculatus is so distinct mor-
phologically it has received less attention from
taxonomists than have other more complex
Helophorus groups (Angus 19706; McCorkle
1967). But the species is of great importance for
the discussion of the fossils described in this
paper; therefore those descriptions are prefaced
here by a detailed discussion-of H. tub&culatus,
focusing on the structure of the head, pronotum,
and elytron and problems associated with their
recognition as fossils.
Description
Head-Length, 0.38-0.50mm (mean = 0.44
mm); width, 0.60-0.80 mm (mean = 0.73 mm).
Black, shining, faint metallic reflections on some
specimens. Surface granulose (Fig. 6b), each
granule circular with a depressed setate puncture
at apex, setae curved; granulation slightly more
dense on epicranium, less dense to effaced near
apex of Y-shaped groove. The latter well
developed; lateral arms slightly sinuate at ends,
extended to margin of head; stem of Y-shaped
groove nearly linear, not greatly expanded ante-
riorly,+ extended posteriorly to cranial suture
(coronal ridge). Clypeus moderately domed at
centre, outer angles rounded with narrow lateral
bead of uniform width extending onto and +
across anterior clypeal margin. Labrum flat, with-
out pronounced shelf on upper surface, anterior
margin curved, setate. Mandibles falciform, uni-
dentate. Terminal articles of maxillary palps
symmetric. Antennae of nine articles.
Pronoturn-Length, 0.7-0.9 mm (mean = 0.80
mm); width, 1.06-1.36 mm (mean = 1.21 mm).
Black, shining, faint metallic reflections at
base on some specimens. Widest in anterior
half; sides straight to slightly sinuate, con-
verging posteriorly ; lateral margin slightly to
strongly denticulate with curved setae arising
between individual denticles; anterior angles
protruded; posterior margin in form of a broad
'V' with straight to slightly sinuate sides (Fig.
6d). Width between basal angles equal to or
less than distance between anterior pronotal
angles. Pronotal surface vaulted mediad of sub-
marginal grooves, reflexed laterad of submar-
ginal~.Grooves smooth, not deeply impressed,
,formed mainly by absence of granules; inner
three (median and submedians) better demar-
cated than outer four (submarginals and mar-
ginal~).Median groove usually widest at centre,
extended to base and there deeper or with
distinct pit; submedians distinctly sinuate, on
some specimens f confluent with median at
middle via zone of reduced granules on internal
intervals; submarginal grooves each with deep
pit at base, less sinuate than submedians, in
some specimens invaded by granules near
anterior margin; marginal grooves poorly de-
fined, in most specimens obscured by granules
basad of middle and f confluent with submar-
ginals via reduced granulation near middle of
external intervals. Suprapleural region, viewed
from below, strongly concave, shining, relatively
wide, tapered gradually to base.
Elytron-Length, 2.2-3.04 mm (mean = 2.66
mm); width, 0.48-0.64 mm (mean = 0.54 mm).
Black, shining, parallel sided before tapering
apex, apical declivity pronounced; shoulder
well developed, narrowly rounded; base abruptly
delimited, scutellar notch small (Fig. 6a). Eleven
coarsely punctate striae, little impressed between
punctures; 1lth not visible from above, reduced
to few punctures along true epipleural margin;
intercalary stria present, about 0.20 length of
elytron (range, 0.16-0.24), either joined or not
 MATTHEWS
to st. 1, f impressed. Interstices 11, each possess
median row of small punctures (doubled or
tripled at site of ridges and tubercles), with fine
recumbent hairs of length about equal to dis-
tance between punctures; alternate interstices of
some specimens slightly raised above others at
apex; sutural interstice usually more convex
and raised above int. 1 at middle; int. 10
strongly carinate except at base and apex,
forming a pseudepipleuron wider than true
epipleuron (viewed from below). Margin of
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pseudepipleuron not dentate. Interstices 2, 4, 6,
and 8 with interrupted ridges and (or) isolated
tubercles, variously developed but at relatively
constant positions (Fig. 2). Int. 8 slightly
carinate in apical half; int. 9 + carinate at
base. Int. 2 with basal ridge (2a) followed behind
oblique transverse elytral impression by tubercles
2b, 2c, 2d; tubercle 2b low and sometimes
nearly joined with 2c; 2c and 2d always prom-
inent. Interstice 4 with low ridge (4a) immediately
behind transverse elytral impression; in some
specimens 4a joined to 4b as long, low ridge,
on others in form of f discrete nodes. Tubercles
at 4b and 4c, the former well developed (Fig. 3).
Interstice 6 with ridge at base (6a), merged with
basal portions of ints. 7 and 8 to form swollen
For personal use only.
elytral shoulder. Tubercles at 6b and 6c, former
always well developed (Fig. 3). Interstice 8 with
low tubercle or ridge (8a) opposite 4b. True
epipleural margin markedly serrate at base and
apex, less so in between. Oblique transverse
elytral impression well developed, extended at
least to int. 4, often as far as ridge at 6a.
IntraspeciJic Variation
To understand the distinctive character of
the fossils discussed below, the elytral variation
of H. tuberculatus must be documented. For this
purpose the tubercles or ridges at 2b, 2c, 2d,
4a, 4b, 3c, 6b, 6c, and 8a (Fig. 2) have been
scored on a scale of 0-4 according to their
relative development. A score of 0 indicates that
at the expected tubercle site there is no sign of
tubercle development or even doubling of the
interstice puncture row that normally ac-
companies the tubercles. The score 1 indicates
doubling of the interstice puncture but no
visible bulging of the cuticle; 2 indicates slight
bulging; 3, definite bulging and doming, enough
to affect the linearity of the adjacent stria1
puncture rows; and 4, the most pronounced
657
state, with the sides of the tubercles nearly
vertical and abruptly delimited anteriorly and
posteriorly. The caption for Fig. 6a shows how
an elytron might be scored.
According to Fig. 3, tubercles 2c, 2d, 4b, and
6b are consistently among the best developed;
4a, 4c, 6c are more variable; and 8a and 2b
usually rather poorly developed. Figure 4
illustrates intraspecific variation of mensural
characters in H. tuberculatus compared with
analogous data from the fossils described below.
Habitat
The habitat requirements of H. tuberculatus
are poorly known. Though seemingly associated
with bog or riparian sites, it probably does not,
as the following examples indicate, live in the
water like many species of Helophorus s.str.
A. Smetana and J. M. Campbell (personal
communication, 1974) have collected H. tuber-
culatus in the Yukon Territory from litter
beneath alder bushes near a river, and also
at the side of a small alpine pool in association
with the beetle genera Pterostichus (Cryobius),
Acidota, Stenus, and Boreaphilus. Near Old
Crow (Yukon Territory) I collected a series of
24 specimens by vigorous trampling of the moss
and sedge-lined margins of ice-wedge polygon
pools within an area of sparse spruce cover.
The species may live in the mosses since nearby
pools surrounded entirely by sedges and grasses
failed to yield H. tuberculatus. In England and
the American Pacific Northwest, specimens have
also been collected in peat moss (Balfour-Brown
1958: McCorkle 1967) and burned areas
( ~ n & s1973), both contexts in which the tuber-
culate elytron has a camouflaging effect (R. B.
Angus, personal communication, 1975). Blatch-
ley (1910) records specimens from litter on the
south shore of Lake Michigan.
Distribution
It is difficult to plot the distribution of H.
tuberculatus because the beetles are rarely
collected. Figure 5 is an approximation based on
the localities of specimens I have seen, literature
citations, and personal communications. Note
that H. tuberculatus is an Holarctic species, with
a northern limit corresponding, in North
America at least, to about the position of
coniferous tree line (Angus 1974).
The species is not known from Alaska; how-
ever, this is probably due to inadequate col-
 CAN. J . ZOOL. VOL. 54, 1976
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T u b e r c l e s c o r e (see text)- o I 2 3 4
FIG.3. Elytral tubercle variation in a sample of H. tuberculatus Gyll. from Nearctic and Palaearctic
localities (n = 86). Curves represent smoothed histograms of the frequency of tubercle scores 0,1, 2,
3, and 4. See text and Fig. 2 for details on tubercle numbering and scoring.

lecting since it did live in central Alaska during
the Holocene (see Fossils) and occurs today at
sites like Dawson and Old Crow, near the
Alaska-Yukon border.
Fossils
In Europe numerous fossil beetle assemblages
have been documented and some of them (e.g.
Coope 1968) have yielded abundant Helophorus
fossils. In fact, the genus Helophorus is rep-
resented by at least a few specimens in almost
every British assemblage studied; yet to date
none of the fossils have been referred to H.
tuberculatus (G. R. Coope, personal com-
 MATTHEWS

Heod length
1

Heod width
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Pronotol length

Pronotol width

E l y t r o l width
For personal use only.

E l y t r o l length

FIG. 4. Mensural data for H. tuberculatus Gyll. Horizontal line, 3 SD (a) about mean (vertical
line); zone delimited by arrows, observed range of measurements; stippled bar, l a either side of
mean; black bar, confidence interval (95%) of mean. All dimensions in millimetres. Note scale change
for 'elytral length.' Triangle signifies value of comparable feature on holotype and referred pronotum
and head of H. mekhenensis a s p . Circle signifies value of comparable feature on holotype of H.
coopei n.sp.

rnunjcation, 1975). In North America, A.
tubercutatus fossils are known only from four
localities : (I) a mid-Wisconsin interstadial peat
from the Titusville, Pennsylvania, section (G. R.
Coope, personal communication, 1975; Totten
1971; White and Totten I965); (2) the 12 100-
year-old peat from MalIoy Pit located near
Lockport, New York, and the sauth shore of
Lake Ontario (Matthews and Miller, in prepara-
tion; Miller 1973); (3) an organic zone within
the late Wisconsin Champlain Sea sediments
exposed near Mt. St. Hillaire, Quebec (V. K.
Prest, personal communication, 1975); and (4)
a 7280-year-old peat horizon at McGee Pit
near Tofty in west central Alaska (Matthews,
unpublished; Matthews 1970~).I have seen only
the fossils (all elytra) from the last three sites,
and of these only the ones from the McGee
section and Mt. St. Hillaire are well enough
preserved for comparison with elytra of extant
specimens (Fig. 8). The Lockport assemblage
included several elytral fragments, which, though
 660 CAN. J. ZOOL. VOL. 54, 1976
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FIG.5. Inferred distribution of Helophorus tuberculatus
Gyll. (hatched area) based on specimens seen by the
author and on literature citations. 0, literature citations;
m, localities of specimens examined for this report,
as follows (with number of specimens). CANADA, Yukon
For personal use only.
Territory: Dawson City (1); 8 mi NW Keno Hill (1);
Old Crow area (24). British Columbia: Summit Lake,
mi 392, Alaska highway (6); Summerland (1). Alberta:
McMurray (lo); Edmonton (I); Waterton (I). Manitoba:
Winnipeg (4); Seddons Corner (1); "Aweme" (near
Brandon) (1). Ontario: Ottawa (12); Prince Edward
County (1); Brimley (7). Quebec: Montreal (2); Mistas-
sini Lake (3). Nova Scotia: Lockeport (1). UNITED
STATES-Horn Mountain Club, Michigan (2). FINLAND-
(2). FRANCE-Pontarlier (1). GREAT BRITAIN-Scotland:
Coatbridge (3); Glasgow (1).
clearly recognizable by their tubercles as be-
longing to H . tuberculatus, are too poorly pre-
served to be scored as in Fig. 3.
Because specimens of H . tuberculatus are
structurally distinctive, complete or partially
articulated fossils of the species are unlikely to
be confused with any other helophorine, even H.
auriculatus Sharp, an Asian species that, like H.
tuberculatus, possesses elytral tubercles (Fig. 6e).
Isolated fragments, particularly of heads, might
not be so readily identified, especially using
only the character of the surface sculpture. H.
nubilus is similar to H. tuberculatus in the form
and type of surface granules and even the more
distinctive irregular polygonal domed granules
found in H . crinitus, H. obscurellus, and H.
arcticus (Angus 1970a, 1970b) may resemble
those in H. tuberculatus (compare Fig. 6b
and 6c). H. crinitus, however, is much larger
than H. tuberculatus (R. B. Angus, personal
communication, 1975) and in the other three
species the sides of the clypeus diverge more
toward the eyes (e.g., Fig. 6c). H. nubilus also
has the stem of the Y-shaped groove expanded
anteriorly. Fossil heads of H. obscurellus and H.
arcticus that retain mandibles can be dis-
tinguished from H. tuberculatus by the bidentate
apex of their mandibles,
Despite similarities of surface sculpture, the
pronota of the above-mentioned species can be
easily distinguished from those of H. tuber-
c u l a t ! ~by~ larger size (H.crinifus),nontapering
suprap!euraI area (H. crinitus, H. nubilus),
better development of the pronotal grooves (H,
obscus~llus, H. nubihs, H. crinitus), and total
absence of pronotal grooves ( H . arcticus).
Helophorus (Gephelophorus) auriculatus Sharp,
a southeast Asian species, is apparently the only
helophorine besides H. tuberculatus and the
fossils described below to have a tuberculate
elytron (Angus 1970a). Nevertheless fossil elytra
of H. tuberculatus and H. auriculatus are un-
likely to be confused because the latter is some-
what larger (3.3-4.3 mm in the three specimens
I have examined), the tubercles are sometimes
nearly absent (R. B. Angus, personal com-
munication, 1975) or in any case less developed
(Fig. 6e), and the prominent pseudepipleuron
is slightly dentate or serrate, whereas the true
epipleuron is only sparsely serrate, much less
so than on H . tuberculatus. Although colour
distinctions are not always reliable for identifica-
tion of fossils, the elytron of H. auriculatus
is dark brown, not black like H. tuberculatus.
Angus ( 1 9 7 0 ~states
) that the intercalary striae of
H . auriculatus are 0.25 the length of the elytra,
but under the mensural scheme outlined above
the three H. auriculatus specimens I have seen
yielded values (0.19, 0.21, and 0.18) not greatly
different from those recorded for H. tuber-
culatus.
Assessment of the relationships of Helophorus
subgenera is beyond the scope of this paper;
however, mention must be made of several
previous opinions specifically relating to
Cyphelophorus. McCorkle (1967), in his treat-
ment of the Nearctic fauna, considered Cyphelo-
phorus to be allied with the subgenera Gephelo-
phorus and Meghelophorus because all three
 MATTHEWS 661

have nine-segmented antennae, intercalary striae,
and short stiff recurved setae on the elytron,
pronotum, and head. Furthermore, he con-
sidered H. (Gephelophorus) sibiricus Motsch.
(= H. fennicus Gyll.) to be closest to H. tuber-
culatus because it possesses "vestiges" of
swellings on the elytra. According to this line
of reasoning, the sister species to H. sibiricus, H.ment of a right elytron. Strial punctures as in
auriculatus with its tuberculate elytron, would
be considered even more closely related to H.
tuberculatus. But H. auriculatus and H. sibiricus
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4a; well-developed tubercles at 2c, 4b, and 6b;
no doubled puncture row or swelling at posi-
tions 4c and 8a. Interstice 8 not at all carinate;
int. 9 evenly convex along entire length. Basal
third of elytron with transverse oblique im-
pression extended from st. 1 to int. 4.
PARATYPES : GSC-433 13 (Fig. 7b). Basal frag-
holotype. Ridges 2a and 6a slightly more
developed than holotype. Base of int. 4 with
only a few doubled punctures. Carinate anterior

differ markedly from H. tuberculatus in other
characters (including some on the elytra); hence
there seems little reason to consider the two
subgenera, Gephelophorus and Cyphelophorus,
to be closely related. Angus (1973), in a discus-
sion of some Palearctic Helophorus species,
expresses another opinion on Cyphelophorus, i.e.
that it is intermediate in structure between the
two subgenera Empleurus and Trichelophorus,
both of which have the alternate elytral in-
terstices keeled.
Helophorus (Cyphelophorus) coopei n.sp.
Description
HOLOTYPE: GSC-43312, right elytron; slightly
flattened, missing portions near shoulder and (see Fossil Localities section).
For personal use only.
portion of int. 10 evident, smooth (not serrate),
disappearing before shoulder. True elytral mar-
gin serrate at base, less so posteriorly. Scutellar
notch small as in H. tuberculatus.
DISPOSITION OF TYPE MATERIAL: Holotype and
paratype in the Geological Survey of Canada
(GSC) type collection: GSC, 601 Booth Street,
Ottawa, Ontario KIA OE8.
DERIVATION OF SPECIFIC EPITHET: After G.
Russell Coope, pioneer investigator of Quater-
nary insect fossils.
DISTRIBUTION: Known only from the type
locality, Lava Camp Mine (65O53.04' N, 163'7.9'
W, Fig. I), Seward Peninsula, western Alaska.
AGE: About 5.7 million years, late Miocene

scutellar region (Fig. 7a).

Length, 3.04 mm; width, 0.54 mm. Black, Comparison and Relationships
shining, lacking metallic reflections (wet or dry). Helophorus coopei is distinguished from all
Striae: 11 complete plus intercalary stria; other species of Helophorus except H. auri-
punctate, impressed between punctures; I lth less culatus, H. tuberculatus, and the next described
impressed, confined to margin of epipleuron. fossil species by its possession of a tuberculate
Intercalary atria abaut 0.20 total elytral length, elytron. It is distinguished from H. auriculatus
not merged with st. 2 at base, isolated from st. 1 by the smaller size of the elytron, more poorly
posteriorly, impressed between punctures. Inter- developed carina on int. 10, and the restriction
stices : 11, moderately convex with single medial of denticles or serration to the true elytral
puncture row except at sites of ridges and tuber- margin. It differs from H. tuberculatus and the
cles and at base of int. 4. Some punctures retain next described fossil species by its more finely
small recumbent setae. Sutural interstice not developed striae (striae impressed between
markedly convex or raised above int. 1. Inter- punctures and individual punctures smaller), and
stice 10 carinate except at: apex (base obscured the less developed array of elytral tubercles,
by break), i n form of pronounced pseudepi- notably the absence of tubercles at positions 4c
pleuron that is wider than a true epipleuron and 8a and the incipient character of tubercle
(viewed from below). True elytrnl margin 2d (Figs. 7a, 8). Total tubercle score is well
strongly serrate at base and apex, less so between. outside range of variation of H. tuberculatus
Interstices 2, 4, and 6 with low tubercles or (Fig. 8).
ridges. Ridges present at 2a, and 6a and zone of H. coopei is referred to the subgenus Cyphelo-
doubled interstice punctures at base of int. 4. phorus because of its possession of a system of
Interstices 7 and 6 joined at base in form of elytral tubercles, intercalary stria, and similar-
raised elytral shoulder. Tubercles: 2b, with ity to H. tuberculatus in size and the develop-
doubled puncture row; 2d and 6c with doubled ment of the pseudepipleuron.
puncture row and slight swelling; low ridge at In Fig. 7a the H. coopei elytron appears to
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652
CAN. J. ZOOL. VOL. 54. 1976
 MATTHEWS 663
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For personal use only.

FIG. 6. (a) Helophorus (Cyphelophorus) tuberculatus Gyll. (Ottawa, ,Ontario. W.H.H., 192-, Can.
Natl. Coll.) Total elytral tubercle score (see text and Fig. 2): 2b,2; 2c, 3; 2d, 4; 4a, 3; 4b, 4; 4c, 3; 66, 4;
6c, 3; 8a, 3; total = 29. (b) H. tuberculatus Gyll. (Merivale, Ontario, IV-28-39, T. N. Freeman,
Can. Natl. Coll.). Head lacking labrum, mouthparts, and portion posterior to coronal ridge. (c)
Helophorus (Orphelophorus) arcticus Brown (Churchill, Manitoba, VI-16-1937, W. J. Brown, Can.
Natl. Coll.) Head lacking labrum, mouthparts, and portion posterior to coronal ridge. Compare shape
of clypeus with that of H. tuberculatus in Fig. 66. (d) H. tuberculatus Gyll. (Old Crow area, Yukon
Territory, VI-15-1975, J. V. Matthews). Left half of pronotum. Note development of granules on
the intervals, extension of median groove to pronotal base, and tendency for marginal and sub-
marginal grooves to be confluent. (e) Helophorus (Gephelophorus) auriculatus Sharp. (Tygosan
Island, Chusan Archipelago, People's Republic of China, Walker Coll. 93-18, BM-9616.) Left elytron.
Note weakly developed tubercles on interstices 2, 4, and 6. Color pattern not visible. (f) H. (Gephelo-
phorus) sibiricus Motsch. (Tangle Lakes, Alaska, VI-14-21-1969, J. V. Matthews.) Left elytron.
Note raised alternate interstices. Color pattern not visible. Scale bar = 0.5 mm.
FIG. 7. (a) Helophorus (Cyphelophorus) coopei n.sp. Holotype (GSC-43312). Lava Camp Mine,
Alaska. Late Miocene. Right elytron. Elytral tubercle score (see text and Fig. 2): 26, 1; 2c, 3; 2d, 1;
4a, 2; 46, 3; 4c, 0; 66, 3; 6c, 1: 8a, 0; total = 14. Compare tubercle development with Fig. 6a. (b)
H. (Cyphelophorus) coopei n.sp. Paratype (GSC-43313). Lava Camp Mine, Alaska. Late Miocene.
Basal portion of right elytron. Note basal part of intercalary stria between st. 1 and st. 2, carinate
portion of interstice 10, serrate margin, and tendency for puncture doubling at base of int. 4 (directly
above 'b' in photo). Scale bar = 0.5 mm.
 CAN. J. ZOOL. VOL. 54, 1976

Helophorus
T u b e r c l e Score
tubercu/otus G y l l . I
I
Left elytron I
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Helophorus
meiphenensis n. sp.

Heloplr o r u s
coopei n. sp.
For personal use only.

Total Tubercle Score ( n =86)

FIG. 8. Range of tubercle scores (stippled zone) for H. tuberculatus compared with tubercle de-
velopment on fossils of H. tuberculatus and holotypes of H. coopei n.sp. and H. meighenensis n.sp.
Horizontal bar, 3 SD (o) either side of mean (vertical bar); zone delimited by arrows, observed range
of scores (n = 86); stippled bar, 1 o either side of mean; black bar, confidence interval (95%) of mean.
A, scores of Quaternary fossils from McGee cut, Alaska; B, scores of Quaternary fossils from Mt. St.
Hillaire, Quebec (see text). Note that H. coopei and H. meighenensis differ most from H. tuberculatus
with respect to posterior tubercles; both fossil species are outside the 3 o range of H. tuberculatus.
FIG. 9. (a) Helophorus (Cyphelophorus) meighenensis n.sp. Holoty pe (GSC-43314) Locality 2-73,
Meighen Island, N.W.T. Late Miocene? Right elytron. Elytral tubercle score (see text and Fig. 2):
2b, 3; 2c, 3; 2d, 3; 4a, 1;4b, 3; 4c, 1;66.4; 6c, 2; 8a, 1 ;total = 21. (b) H. (Cyphelophorus) meighenensis
n.sp. GSC-43319. Locality 2-73, Meighen Island, N.W.T. Late Miocene? Pronotum, right half, some-
what contorted. Compare granule development on inner intervals with Fig. 6d and note that (1)
median groove terminates before base; (2) because of contortion of specimen, part of pleural region
is visible outside margin. (c) H. (Cyphelophorus) meighenensis n.sp. GSC-43318. Locality 2-73, Meighen
Island, N.W.T. Miocene? Head, clypeus, and posterior portion greatly contorted. (d) Clypeal margin
(enlarged) of GSC-43318. Note angularity of the corner; and disposition of the lateral bead with
respect to corner and anterior margin (upper margin in photograph). This feature is not due to dis-
tortion of the fossil. (e) Clypeal margin (enlarged) of H. tuberculatus. Compare with Fig. 9d. Scale
bar = 0.5 mm; Fig. 9e and d are same magnification.
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19
For personal use only.
 a66 CAN. J. ZOOL. VOL. 54. 1976

be less parallel sided than that of H. tuber- at base and slightly converging with st. 1
culatus, but this is the result of the orientation posteriorly; on other two, intercalary stria
of the fossil and the fact that it has suffered deeply impressed but effaced before base and
some postdepositional flattening. The last fact not at all tending to converge with either st. 1
also explains why the true elytral flank of the or st. 2. Other striae as on holotype. Sutural
holotype (Fig. 7a) is so much more visible than interstice of two specimens raised more above
the elytral flank of H. tuberculatus in Fig. 6a. int. 1 at middle than on holotype. Interstices 2,
4, and 6 with tubercles, ridges or zones of
Helophorus (Cyphelophorus) meighenensis n.sp. doubled punctures at normal positions. Tubercle
Description 2a less developed than holotype on two speci-
HOLOTYPE : GSC-43314, complete right elytron mens; 2b of one obscure, consisting at most of
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(Fig. 9a). doubled puncture zone; remaining specimens

Length, 3.00 mm; width, 0.6 mm. Black, with 2b as on holotype. GSC-43315 with int. 4
shining, slight metallic reflection (wet). Striae: at base slightly convex and possessing doubled
11 complete plus intercalary stria; punctate; interstice puncture row. Tubercles 46, 4c, and
first deeply impressed, outer ones less so and 6a as on holotype. Only GSC-43317 with region
consist of coarse puncture series. Stria 11 with of 8a intact, but shows no tubercle development
punctures less numerous, confined to elytral or interstice puncture doubling. Two specimens
margin. Intercalary stria, about 0.17 total with int. 9 slightly more convex basally than on
elytral length, merging basally with st. 2, holotype. Int. 10 distinctly carinate on all
isolated and not converging posteriorly with st. paratypes, but on GSC-43317 less so than on
1, little impressed between punctures. Inter- holotype-a possible preservation effect. Trans-
stices: 11, convex to flat with single medial verse elytral impression evident on all three
puncture row, except at sites of ridges and paratypes.
tubercles and at base of int. 4. Sutural inter- DISPOSITION OF TYPE MATERIAL: Holotype and
stice narrower than int. 1, more convex than paratypes in the Geological Survey of Canada
and raised above int. 1 at middle. Interstice 10 type collection: GSC, 601 Booth Street, Ottawa,
For personal use only.

carinate except at base and apex; true elytral Ontario KIA OE8.
margin markedly serrate along entire length. DERIVATION OF SPECIFIC EPITHET : After Meig-
Interstices 2, 4, and 6 with low tubercles or hen Island, Queen Elizabeth Group, Canadian
ridges. Ridges at 2a and 6a, the latter combined Arctic Archipelago, Northwest Territories.
with ints. 7 and 8 at base to form elytral shoulder. DISTRIBUTION: Known only from the type
Tubercles well developed at 26, 2c, 2d, 4b, and locality, Meighen Island, N.W.T. (Fig. I),
6b; 4a and 4c represented by doubled puncture locality 2-73 (79"50' N, 99'15' W), Beaufort
row with little evidence of swelling; 4a tending Formation sample JVM 5b.
to merge with 4b; 6c low but definitely bulging. AGE: Latest Miocene to Pliocene (see Fossil
Interstice 8 not carinate, but slightly more Localities section).
convex in apical half; doubled interstice row
but no swelling at position 8a; int. 9 slightly Referred Fossils
carinate in basal fourth. Transverse oblique im- These include one head GSC-43318 (Fig. 9c)
pression on basal third poorly developed. and a partial pronotum GSC-43319 (Fig. 9b)
PARATYPES: Three elytral fragments: GSC- from the same assemblage as the holotype and
43315, right elytron missing portion posteriorad paratypes. Their similarity to the heads and
of a break across tubercle 2d to behind 4c pronota of H. tuberculatus suggests they rep-
and 66; GSC-43316, right elytron lacking por- resent H. meighenensis; however, inasmuch as
tions posteriorad of 4b and laterad of interstice they were not directly associated with the holo-
6; GSC-43317, left elytron, lacking portion type or any of the paratypes of H. meighenensis
posteriorad of a break crossing anterior to 2c, and represent parts other than the elytra, they
longitudinally through 4b, and obliquely laterad are not treated here as part of the type series.
behind position of tubercle 8a to elytral margin. Head-Fig. 9c. Somewhat smaller than H.
One specimen, GSC-43317, with intercalary tuberculatus; length, 0.36 mm; width, 0.62 mm.
stria less impressed than in holotype, formed of Colour: black, shining. Differing from H.
four or five punctures only, merged with st. 2 tuberculatus by the more polygonal shape of
 HEWS 667

surface granules (compare Figs. 9e and 9d) and tion. The fossil species is referred to the sub-
the form of the clypeus, which on the fossil is genus Cyphelophorus because of its similarity to
more angulate at the corner with the lateral bead H. tuberculatus.
not extended significantly onto anterior margin.
Mandibles and labrum present; the former Fossil Localities
falciform unidentate; the latter evenly rounded, Lava Camp Mine
similar to that of H. tuberculatus. This site, previously described by Hopkins
Pronotum-Fig. 9b. Slightly wider than H. et al. (1971), is an exposure of frozen, woody,
tuberculatus; length, 0.78 mm; width (by extra- peaty alluvium and pond sediments capped by a
polation) 1.42 mm. Colour : black, shining. reversed polarity basalt dated at 5.7 f 0.2
Similar to H. tubercuIatus except for the less million years before present. Because the peat
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19

serrate lateral margin, deeper and narrower and wood is charred and structures at the basal
pronotal grooves, and reduced surface granula- contact of the basalt suggest flow of lava over a
tion on intervals (caused by raised intergranular wet substrate (Hopkins et al. 1971), the organic
area). Marginal pronotal groove narrower than sediments from which the fossils come may be
in H. tuberculatus, obliterated posteriorly and assumed to be nearly contemporaneous with
anteriorly; better delimited medially with no the extrusion of the basalt. The basalt and
tendency for confluence with submarginal groove underlying sediments have been referred to
across the external interval. Median, submedian, the Pliocene (Hopkins et al. 1971); but,
and submarginal grooves more incised than on under the present concept of Tertiary chronol-
H. tuberculatus. Submedians markedly sinuate ogy, derived from research on deep-sea cores,
and parallel sided, forming a deep trench at the 5.7-million-year date represents the late
base; median terminated abruptly before base, Miocene (Berggren and Van Couvering 1974).
somewhat obscure at anterior margin. Surface The study of deep-sea sediments has also
granulation best developed on external intervals; provided welcome confirmation of the validity
on middle and internal intervals less defined, of the absolute age determination for the Lava
nearly reduced to a flat punctate surface, Camp site, since Paleomagnetic Epoch 5,
For personal use only.

especially behind the posterior end of median straddling the Miocene-Pliocene boundary, con-
groove. tains an interval of reversed magnetic polarity
dated at 5.7-5.58 million years (Berggren and
Comparison and Relationships
Van Couvering 1974; Ryan 1972), remarkably
Helophorus meighenensis differs from H.
close to the age of the reversed polarity Lava
auriculatus by the same elytral characters that
Camp basalt.
separate the latter from H. tuberculatus. Though
During the late Miocene, Seward Peninsula
very similar to H. tuberculatus, the elytron of H.
and particularly the Lava Camp region probably
meighenensis can be distinguished by its posses-
possessed lowland deciduous-coniferous forests
sion of only incipient tubercles at 8a and 4c
and an insect fauna similar in taxonomic com-
plus the relatively poor tubercle development
position to that of the present-day North
at positions 2c, 2d, 4b, and 6c. The score for American Pacific Northwest (Hopkins et al.
each individual tubercle is within the range of
1971). Helophorus (Cyphelophorus) was originally
H. tuberculatus, but the total tubercle score is listed as present in the Lava Camp fossil as-
clearly not (Fig. 8, bottom).
semblage (Hopkins et al. 1971, p. 223), but it was
Providing the pronotum and head mentioned only after more of the sediments were processed
above refer to H. meighenensis, that species is
and searched for fossils that a relatively com-
further distinguished from H. tuberculatus by plete elytron, here described as H. coopei,
the slightly more angulate clypeal margin with
confirmed the original determination and pro-
its lateral bead not evenly prolonged onto the
vided the necessary morphological criteria for
front, the deeper longitudinal grooves on the distinguishing the fossil from Helophorus tuber-
pronotum, failure of the median groove to
culatus.
reach the base,2 and less pronounced granula-
'R. B. Angus (personal communication 1975) has seen Meighen Island
3ne Siberian specimen of H. tuberculatus that approaches Helophorus meighenensis and other Meighen
.his condition. Island insect fossils previously discussed (Mat-
 668 CAN. J . ZOOL. VOL. 54, 1976
thews 19743) come from the Beaufort Formation,
a unit of unconsolidated sands, gravels, and
organic sediments exposed over the entirety
of Meighen Island (Thorsteinsson 1961), the
western parts of other islands in the Queen
Elizabeth group (Tozer and Thorsteinsson 1964)
and as isolated outliers on Axel Heiberg Island
(Balkwill and Bustin 1975) and possibly Elles-
mere Island (D. Hodgson, personal communica-
tion, 1974).
Meighen Island locality 2-73, like other
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19
exposures studied to date, consists primarily
of several tens of metres of alluvial sand with
sporadic interbedded lenses of peaty sediments
representing old channel fillings. These peats
usually contain plant and insect fossils, the best
preserved of the latter coming from a locality
2-73 peat horizon which can be recognized at
several other exposures (= "Lower Peat Hori-
zon" of Kuc 1973, 1974a). The peat typically
occurs near the top of the exposures, about 9 m
below a felted moss peat (= "Upper Peat
Horizon" of Kuc 1973) and 19-20 m below the
top of the exposure and a thin cap of till or
glacial lag deposits.
The Beaufort Formation is not precisely
dated, although Hills et al. (1974) have shown
For personal use only.
that exposures on southern Banks Island (Fig. 1)
are correlative with the early to mid-Miocene
Seldovian Stage of Alaska (Wolfe et a[. 1966).
The exceptional preservation of Beaufort Forma-
tion fossils on Meighen Island could be taken
as an indication that the northern Beaufort
sediments are of Pleistocene interglacial age
(Kuc 1973; M. Kuc. personal communication,
1973). However, preliminary study of insect and
plant macrofossils from Meighen Island (Mat-
thews 19746; Hills and Matthews 1974) reveals
assemblages with closest affinities to the present-
day northern coniferous and taiga zone more
than 800 km south of Meighen Island (Fig. 1).
In contrast, Pleistocene interglacial sediments
from islands such as Banks and Bathurst (Fig. I),
located well south of Meighen Island, record a
much smaller northern movement of the boreal
and tundra biota (Blake 1974; Kuc 19743).
Meighen Island fossils thus portray a climate
that is much warmer than that of an Arctic
interglacial; consequently, the age of the Meig-
hen fossils and the sediments from which they
come is assumed to be older, at least of late
Tertiary age.
A more precise estimate of the age of the
Beaufort Formation on Meighen Island will
be possible once some of the well-preserved
plant fossils are studied in detail. Microfossils
now under study from marine clays underlying
the locality 2-73 exposure will also undoubtedly
provide data for correlation of Meighen Island
Beaufort sediments with marine Tertiary se-
quences in other parts of the Arctic Basin.
Evolution and Phylogeny
As a large group of numerically abundant and
taxonomically diverse animals, the insects are
notable for their meager fossil record. Con-
sequently, entomologists seeking knowledge of
the evolutionary history of insect taxa must
often rely only on the information embodied
by the extant species. This approach, neces-
sarily cladistic in principle, usually involves
search for derived characters and establishment
of sister groups using the methodology known
as "phylogenetic systematics" (Hennig 1966).
Phylogenetic systematics has been the subject
of heated debate (Darlington 1970; Brundin
1972) and much discussion by both neontologists
and paleontologists alike (Cracraft 1974; Schaef-
fer et al. 1972; Brundin 1972; Kavanaugh 1972).
Many paleontologists would, I believe, take
issue with some of the implications arising from
rigid interpretation of the tenets of phylogenetic
systematics (e.g. see Darlington 1970 for a good
discussion of these problems); nevertheless, its
use has resulted in phylogenetic hypotheses that
are both consistent with alternate evidence and
suitably formulated for testing in the light of
possible future evidence (Whitehead 1972).
Fossils provide one of the best means of
testing phylogenies; therefore it is imperative
that neontologists not disregard fossil evidence
when seeking evolutionary history via the
phylogenetic systematics approach. It is equally
important that the paleontologist not neglect
the wealth of information offered by extant
taxa when formulating phylogenetic hypotheses,
especially in cases such as this one in which the
fossils are so clearly related to an extant species.
The following discussion relies heavily on the
characteristics of extant Helophorus species, but
involves implicit assumptions that deviate some-
what from those of both the phylogenetic
systematists and phyletic gradualists (Cracraft
1974; Eldredge and Gould 1972; Schaeffer et al.
1972). These assumptions are (1) fossils should
be treated under the same nomenclatural
 MATTHEWS 669
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FIG. 10. Phenocline illustrating elytral tubercle development in several species of Helophorus.
Drawings are of the midportion of a left elytron including sutural (s), second (2), and fourth (4)
interstices. (a) H. (Cephelophorus)sibiricus condition, characterized by raised alternate interstices with
tendency to form doubled puncture row on raised interstices. (b) H. (Gephelophorus) auriculatus
condition, characterized by possession of low tubercles and interrupted ridges on alternate inter-
stices. (c) H. (Cyphelophorus) coopei n.sp. See Fig. 7a. ( d )H. (Cyphelophorus) meighenensis n.sp. See
Fig. 9a. (e) H. (Cyphelophorus) tuberculatus condition, with pronounced tubercles and interrupted
ridges on alternate interstices. See text for further comment.

system as extant species; (2) ancestors of extant
species are potentially knowable; (3) in inter-
preting phylogeny one should avoid giving a
priori significance to the age or stratigraphic
For personal use only.
closer relationship to other Helophorus sub-
genera than to Cyphelophorus. This is evidence
that tuberculate elytra have evolved indepen-
dently in the two Helophorus subgenera, just

position of fossils (Schaeffer et al. 1972; Eldredge
1971); and (4) even though cladogenesis is
probably the most important style of evolution
and agrees with the concept of allopatric specia-
tion (Eldredge 1971; Eldredge and Gould
1972), phyletic gradualism, or evolution within
one lineage, is an equally likely process in
certain cases.
Figure 10 is a phenocline expressing different
degrees of elytral tubercle development in
Helophorus, from the state of having raised
alternate interstices but no tubercles in H.
sibiricus, to the well-developed tubercles of H.
tuberculatus. Justification for the position of the
fossil species in the phenocline is found in the
taxonomic descriptions.
Because the possession of elytral tubercles
is rare among Helophorus species, the tuber-
culate elytron is considered to be a derived
character. This means that the polarity of the
phenocline in Fig. 10 is from H. sibiricus (most
primitive) to H. tuberculatus (most derived);
however, relationships should not be read in-
discriminately from the phenocline, for, as in-
dicated earlier, H. sibiricus and H. auriculatus
belong to a subgenus (Gephelophorus) that shows
as tuberculate elytra have arisen indepen-
dently in other Hydrophilidae (Hydrochus
nitidicollis Mulsant) and several other Coleoptera
families (Curculionidae, Grypus equiseti F.;
Derodontidae, Peltastica tuberculata Mann.;
Lathridiidae, Aridius nodger Westw.). One might
conclude that each of the fossil species described
here also acquired its elytral tubercles in-
dependently, but I consider this to be unlikely
both because of the rarity of elytral tubercles
among Helophorus species and the other
similarities (size, color, pseudepipleural de-
velopment) that the fossils share with H. tuber-
culatus. Therefore, within the subgenus Cyphelo-
phorus elytral tubercles are considered to be
homologous and the character transformation
illustrated in Fig. 10 may be used to formulate a
phylogeny (Fig. 11). On the basis of elytral
tubercle development H. tuberculatus and H.
meighenensis are sister species and the two com-
bined are the sister group of H. coopei. Ac-
cording to the principles of phylogenetic system-
atics each pair of sister groups must share a
common hypothetical ancestor, but as Ross
(1974, p. 203) has indicated, if a fossil species in
a series possesses all the characters expected of
 CAN. J . ZOOL. VOL. 54, 1976

H. t u b r r c u l o t u s G y l l . H.m r i g h s n r n r i r n.a p t H. c o o p s i n.sp.t

Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19

FIG.11. Theoretical cladistic relationships (solid narrow line) and proposed phylogeny (bold line
and symbols) of the subgenus Cyphelophorus. Two interpretations of phylogeny are expressed in the
diagram: (1) that H. coopei and H. meighenensis are sequential or chronospecies in one lineage
terminating with the extant H. tuberculatus; and (2) that branching (cladogenesis) had occurred.
For personal use only.

the hypothetical ancestor, then it must be that
ancestor. Both of the fossil species discussed here
do embody the elytral characters expected of the
common ancestor, and I consider them to be
sequential or chronospecies within one lineage
(Fig. 11, alternative 1). The present monobasic
character of the subgenus Cyphelophorus seems
to support this conclusion, unless, of course,
it is argued that the subgenus was more diverse,
with all species except H. tuberculatus becoming
extinct during the later Tertiary or Pleistocene.
I find this contingency difficult to support in
view of the apparent persistence of northern
Coleoptera species in the face of the drastic
climatic changes and range disruptions that oc-
curred during the Quaternary (Coope 1973;
Ullrich and Coope 1974).
Alternative 2 in Fig. 11 is the cladistic view
of Cyphelophorus history, agreeing with the
concept of evolution in stages of "punctuated
equilibria," as outlined by Eldredge and Gould
(1972). However, it is a more complicated
hypothesis than alternative 1 in that the splits
which it portrays imply several periods of
geographical isolation. It would be incorrect
to assume that the islandic location of the type
locality of H. meighenensis suggests one of the
required geographical isolations, because at the
time that H. meighenensis lived, the Arctic Archi-
pelago was probably a single land mass con-
nected to the rest of the continent. If the ancestral
Cyphelophorus species had attained a Holarctic
distribution by the Miocene, then the speciation
suggested in Fig. 11 could have occurred during
the separation of North America and Asia
by a seaway in Bering Strait region about 10
million years ago (Hopkins 1967; Nelson et al.
1974). The duration of this separation was
probably several million years, but what we
know about the habitat requirements of H.
tuberculatus and the distribution of the forest-
tundra border during the Pleistocene glacials and
interglacials (Hopkins 1972) makes it likely that
the present disjunction in the distribution of H.
tuberculatus (Fig. 5 ) is at least as old. Since it
has not resulted in any apparent divergence of
populations on either continent, one wonders if
a late Tertiary disruption of similar duration
could have been any more effective as the cause
of speciation. Of course it is not major dis-
 MATTHEWS
ruptions such as this that Eldredge and Gould
(1972) claim as the cause for episodic speciation.
Instead they see such changes taking place
rather quickly in small isolated peripheral
populations. But the Pleistocene record is replete
with cases of disruption or isolation of Coleop-
tera populations in small refugia without
speciation having occurred (Coope 1970). Thus
despite the persuasive argument that phyletic
gradualism is a poor way to explain the evolution
of fossil species (Eldredge and Gould 1972), it
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19
may be the best means of explaining the evolu-
tion of Cyphelophorus.
Several evolutionary implications follow from
the assumption that H. coopei, H. meighenensis,
and H. tuberculatus are sequential species. The
most obvious of these relates to the steps in-
volved in the evolution of tuberculate elytra.
Though not closely related to Cyphelophorus,
the two species H. sibiricus and H. auriculatus
of the subgenus Gephelophorus show what types
of elytral modifications might have occurred
before the evolution of H. coopei. The ancestor
of the two Gephelophorus species probably had
an elytron like that of H. sibiricus (alternate
interstices raised, Fig. lo), and therefore evolu-
tion of tubercles on the elytron of H. auriculatus,
For personal use only.
and by analogy on that of H. coopei, involved
apical reduction of the alternate interstices
except at certain positions where remnants of
the raised interstice were preserved and en-
hanced. Continued enhancement of the elytral
tubercle complement is displayed by the three
Cyphelophorus species (compare apical tubercle
scores of H. coopei and H. tuberculatus, Fig. 8),
but they also show that some of the elytral
tubercles (e.g. 8a) formed after the portion of the
interstice in which they occur had been reduced.
Alternate interstices of all three Cyphelo-
phorus species are ridgelike near the base and
there seems to be little distinction in the relative
development of these ridges on the three species.
In contrast, nearly all of the fossils, compared
with only one of the 86 H. tuberculatus speci-
mens, possess a zone of double interstice punc-
tures near the base in int. 4. Perhaps this dif-
ference, though subtle, is evidence that some
reduction of the interstice ridges has occurred
since the evolution of H. coopei. The range of
variation exhibited by H. tuberculatus elytra
seems adequate for continued formation of
isolated tubercles, since a few of the specimens
67 1
examined had the ridge at 4a weakly divided
into two low tubercles; this condition pre-
sumably represents the derived, high-fitness
phenotype which would be favored by selection.
Selection pressures responsible for the elytral
changes revealed by the fossil species may be
associated indirectly with late Tertiary climatic
cooling, one result of which was probably early
formation of discontinuous permafrost in the
polar areas of the world. This would have led to
a prevalence of cold, organic soils, low in
available nutrients, and with poor drainage, the
type of conditions presently responsible for
the many small moss-ringed ponds found in
today's subarctic areas. H. tuberculatus lives
in the mosses at the margin of such ponds. Its
tubercles seem to help camouflage it against
dark organic backgrounds and the state of the
swimming hairs on its tarsus is intermediate in
development between those on strictly aquatic
and terrestrial Helophorus species (Angus 1966).
Thus evolution of the tuberculate elytron in the
Cyphelophorus lineage may be one of a number
of adaptations for a semiaquatic existence. The
implication of this argument is that H. coopei
was more aquatic than H. rneighenensis or H.
tuberculatus. It is possibly significant that the
finely punctate, deeply impressed elytral striae
of H. coopei are similar to the striae of the more
aquatic species of Helophorus. The smoother
surfaces on the fossil head and pronotum re-
ferred to H. meighenensis also resemble the
condition seen in aquatic species of Helophorus.
The pronotum of H. meighenensis is of partic-
ular interest for what it implies about Helophorus
evolution. McCorkle (1967) suggested that the
pronotal condition of H. arcticus Brown
(sculpture of upright setate nodes and poor de-
velopment to loss of outer grooves) is primitive,
and that during evolution and diversification
of the genus the outer grooves became better
defined as the setate nodes on the intervals
developed into the flattened granules of many
contemporary species. However, comparison of
the pronotum referred to H. meighenensis with
those of H. tuberculatus suggests an opposite
trend, i.e. upright nodes becoming more pro-
nounced as a result of the breakdown of the in-
tergranular areas and outer grooves tending to
disappear as the individual granules of the outer
intervals become less crowded. If this tendency
is inherent in the whole genus then I would
 672 CAN. I. ZOOL.
expect the condition seen in many Helophorus
species-flattened punctate intervals with deep
well-developed pronotal grooves-to be the
primitive state, and that displayed by both H.
tuberculatus and H. arcticus to be highly derived.
Conclusions such as this are predicated on the
assumptions listed above, but the overriding
assumption of this paper is, of course, that
fossils play a valuable role in construction and
testing of phylogenies. Not only do they mark
real benchmarks in evolutionary history, but
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIVERSITY OF MONTANA on 07/22/19
they often provide surprises (e.g. Coope 1973)
not revealed by the distribution or characters of
extant species. Nevertheless, it is to be expected,
because of the meager fossil record of insects,
that most attempts to decipher evolutionary
history of insect groups will continue to rely
primarily on study of extant species. 'Phylo-
genetic systematics' is often used for such
analyses because it is an evolutionary-based,
logical, step-by-step method of character study
and grouping that allows orderly treatment of
the vast array of morphological characters nor-
mally available to an entomologist. Thoughtful
use of its methodology has resulted in phylo-
genies (e.g. Whitehead's 1972 study of the ground
beetle genus Schizogenius) that agree with
For personal use only.
alternative evidence and are imminently suitable
for testing by fossils, should they become
available. But a rigid interpretation of some of
the principles of phylogenetic systematics may
lead to conclusions that do not reflect reality
(Darlington 1970; Whitehead 1972) and that
seriouslv reduce the value of fossils. Such
practice is to be avoided. At the same time it
should be said that there is good reason for
questioning the way paleontologists have tradi-
tionally explained their fossil evidence (Eldredge
and Gould 1972; Schaeffer et al. 1972), and I
have attempted to consider some of those
arguments in the construction of a phylogeny
for the species of the subgenus Cyphelophorus.
But each case must be treated on the basis of the
facts at hand. and those revealed bv the fossils
and extant ~ L l o ~ h o r uspecies
s to the con-
clusions outlined above. Though the fossils
play a central role in reaching these conclusions,
they do not ensure the validity of the proposed
phylogeny. This can only come if future tax-
onomic work of the genus Helophorus and
study of other fossils fail to refute the hypo-
thesis. In view of the excellently preserved
fossils now being extracted from Tertiary sedi-
VOL. 54, 1976
ments in the Canadian arctic, I can state with
assurance that this report will not be the last
paleontological study to deal with the evolution
of Helophorus or the subgenus Cyphelophorus.
Acknowledgments
The fossils of Alaskan origin discussed above
were collected while the author was partially
sponsored by a research grant from the Boreal
Institute of the University of Alberta. All other
mentioned fossils were collected under the
auspices of the Geological Survey of Canada.
E. C. Becker (Biosystematics Institute, Canada
Department of Agriculture), G. E. Ball (Univer-
sity of Alberta), D. H. Kavanaugh (California
Academy of Sciences), and M. J. D. Brendell
(British Museum) kindly loaned comparative
specimens.- Sergei V. Kiselev (Moscow) helped
by checking localities of specimens housed in
Moscow. Louis Ling (Carleton University)
helped with the scanning electron microscope
photography. Finally I thank sincerely R. B.
Angus, G. E. Ball, and A. Smetana, for their
critical comments on this manuscript and closely
related issues.
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