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Key to Saskatchewan Species of Heteroceridae

(version - February, 2014)

David J. Larson
Box 56, Maple Creek, SK S0N 1N0
[email protected]

Introduction. The family Heteroceridae contains small beetles commonly known as variegated mud-
loving beetles. The name is based on their habitat, moist soil at the margins of water, and the basic
body color of variously developed pale areas on a darker background. Characteristics that define the
family are prominent prognathous (forwardly directed) mandibles that are notched at about the middle
of the outer margin; short, stout antenna of which the apical 5 to 7 segments are elongated on the
anterior side to form a serrate club; legs with tibiae bearing elongate spines on outer face and tarsi
slender and 4-segmented; visible abdominal sternum 1 with a strigate sublateral arc that forms the file
of a stridulatory organ across which a sharp ridge on the upper edge of the hind femur passes. Most
species are densely clothed in water repellent setae although this vestiture is reduced in a few species.
Typically the elytra are dark with three transverse paler markings or fasciae located subbasally,
postmedially and preapically; these fasciae may be broad and continuous, broken into several partly or
fully separate spots or may be largely effaced although even then there may be differentiated setae in
the areas where fasciae typically occur. The lateral and basal margins of the elytra and the pronotum,
usually laterally but sometimes medially also, are often paler in color.

All heterocerids, in both adult and larval stages, burrow in moist soil near water. Adults make
shallow feeding tunnels which are often conspicuous as twisting lines of pushed up soil on bare mud
flats. Adults, and presumably also larvae, feed on organic debris, algae and other small organisms in
the surficial layer of the soil but they may also be predators (Ciegler 2003). In addition to these feeding
tunnels, the beetles also make egg chambers and hibernacula. Eggs are laid in groups in egg chambers
and the larvae tunnel out from the parental tunnel system. Egg guarding by adults has been reported
(Folkerts 1989). The constant presence of stridulatory organs also suggests that sound communication
occurs and may be associated with their gregarious habits for they are often found in local
aggregations. The sexes are generally similar and the best external feature for separating them is the
more strongly developed ridges on the stridulatory arc of the male. The males of some species have the
mouthparts, especially the mandibles and labrum, elongated to various degrees and such males are
referred to as being hypermandibulate. A hypermandibulate male also has a more robust pronotum thus
such males present a very characteristic habitus, but within a species there is an allometric range of
male forms from small female-like individuals to large hypermandibulate individuals with heavy
forebodies,

The habitats occupied by heterocerids are unstable, subject to flooding when levels of adjacent
water rises or to drying out in hot, dry periods. This sort of habitat is maintained by instability which
probably also enhances its productivity of small food organisms. Nevertheless, these are not places
where the beetles can persist for an extended period. Some of the features of heterocerids are obvious
adaptations to their unstable environment. For example, the dense, water-repelling vestiture keeps the
body clean in muddy soils and makes the insects buoyant. When the tunnels are flooded the beetles
bob to the water surface from which they can take wing. The beetles fly readily and some species may
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be taken in numbers at electric light.

Identification and Characters. Horn (1890) well described the attitude of entomologists towards
these “unattractive” beetles that are “not ... held in much regard”, “looked upon as almost
inseparable”, and for which the “small amount of literature devoted to them being practically
inaccessible to nearly all”. In spite of this lament, Horn made an important contribution towards
describing and developing a terminology for the external characters that have proven useful in species
recognition. It was only with the extensive use of male genitalic characters by Pacheco (1964) that
systematics of the American members of the group progressed much past Horn's work. However, as
Horn lamented, the tremendous contribution of Pacheco has been limited by the difficulty in accessing
his work. The following section draws mainly from these two authors.

It is easy to recognize a member of the family Heteroceridae for the species generally look
much the same. This has been explained as “the uniform way in which they live seems to have
favored uniformity in external anatomy (King and Lago 2012, after Charpentier 1968). There are
relatively few easily observed external characters for species identification, these being indicated in the
habitus drawing (Illustration 1). Characters useful for species identification are discussed below.

Illustration 1: Generalized heterocerid body,

ventral view, showing important identification
characters (Modified from Pacheco 1964) Illustration 2: Heterocerid dorsal, showing location
and terminology of color and setal characteristics.

Color: The basic color pattern of the body (Illustration 2) tends to be rather uniform across members of
the family, basically with the head, pronotal disc and elytra dark. Each elytron is crossed by three
transverse, sigmoid pale fasciae located behind the basal margin (basal fascia), behind middle (medial)
and before the apex (apical). The lateral margin of the elytron may be entirely dark, with pale areas
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generally where the fasciae extend to the lateral margin or may be narrowly to broadly pale with the
pale area extending from the shoulder to the elytral apex. The elytral base may be entirely dark, with a
pale spot near the scutellum (juxtascutellar spot) or broadly pale with the pale area extending laterally
to the lateral margin at the humeral angle or sometimes extended posteriorly and merging with the
basal fascia. The discal fasciae may be broad and continuous or broken into several spots and on many
specimens are laterally continuous with a pale lateral margin.

Vestiture and Punctation: Density and form of setae covering the upper body varies between species
but is difficult to describe in other than comparative terms. Setae are of various lengths and color and
recline in various directions. A feature that is subtle yet consistent in some species of Heterocerus
subgenus Lanternarius is a whorl or “cow-lick” in the setae in the dark area just behind the anterior arc
of the medial fascia (Illustration 2).

Habitus: Although heterocerid beetles as a group have a characteristic appearance, there are noticable
differences in body form between some species and groups of species. For example, some have
relatively elongate, cylindrical bodies whereas others are broader and more depressed. The ratio of
length of body (L = single measurement from tip of closed mandibles to apex of elytra)/ maximum
width of body (W = width across widest part of elytra) gives an index of shape, the larger the value, the
more slender the insect. However, in species with hypermandibulate males, intraspecific variation is
much greater than interspecific differences.

Antenna: The antenna typically consists of a large scape (segment 1) and pedicel (segment 2), two
small cylindrical segments forming the funicle and an elongate asymmetrical club composed of 5 to 7
segments each of which is produced anterodorsally. Most species have 11-segmented antenna but
fewer segments occur in members of Microaugyles (10) and Tropicus (9). Although this would seem a
very useful character, it is difficult to count antennal segments on an intact specimen and the antenna
should be removed and examined on a microscope slide. In some species of Augyles the basal one or
two segments of the club are small and only slightly expanded and so are easily overlooked.

Sutures and lines: The most useful characters for the recognition of major groups of heterocerids is the
presence or absence of the post-coxal lines. The post-mesocoxal line runs across the anterolateral
portion of the metasternum between the posterior rim of the mesocoxal cavity and the metasternal-
metepisternal suture (some authors call this the mesocoxal line but as it occurs on the metasternum and
not the mesocoxa, this term seems inappropriate). The post-metacoxal line occurs on visible abdominal
sternite 1 and runs from the posterior rim of the metacoxal cavity to the hind margin of sternite 1 where
it forms a continuum with the stridulatory arc. The elytral epipleuron of some species has an oblique
line or ridge shortly behind the humeral angle or shoulder. The line starts on the mesal edge of the
epipleuron and runs obliquely outwards and forewards. In some species it crosses all or most of the
width of the epipleuron, in others it may end somewhat short of the lateral margin and in a few is
present only as a short line or notch on the mesal edge.

Stridulatory Organ: A stridulatory organ formed from a file on the lateral portion of the first visible
abdominal sternite occurs on all species. A sharp ridge on the anterodorsal face of the metafemur
passes across this file as the leg rotates. This structure is common to members of the family and no
feature for species recognition has been discovered. However, it is not impossible that species specific
sounds are made and there could be related morphological differences but this has not been
investigated. However, the file is always better developed with coarser teeth on the male than the
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female and this is the only reliable way of sexing most specimens without dissection.

Illustration 3: Male genitalia of Heteroceridae. Major types and the

parts referred to in keys and descriptions.

Male genitalia: The male genitalia are of peculiar structure, the homologies of which are obscure. I
follow the interpretation and terminology of Pacheco (1964) (also Katovich 2002). Within the body
cavity the genital capsule lies above the wishbone-shaped sternite 9 which is narrow anteriorly with the
two arms diverging posteriorly. The dorsal lobe (= aedeagus) is uppermost and articulates basolaterally
with the phallobase which forms the lateral and ventral portions of the capsule. Parameres are fused to
the phallobase and are directed towards the rear of the abdomen. I use the terms distal or posterior for
those structures closest to the posterior end of the abdomen, and proximal or anterior for those closer to
the front end whereas Miller (1994) and King and Lago (2012) reverse this and refer to the end with the
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parameres as anterior and the opposite end posterior. In the genitalic figures, the anterior or proximal
end is uppermost and the posterior or distal end ventral, similar to the position in which the genitalia
would lie in the beetles of the habitus photographs.

Note on Classification. For a long time, North American heterocerids were placed in just one or two
genera (e.g. Arnett 1963). However, Pacheco (1964) reviewed the New World members of the family
and divided it into 5 tribes and 20 genera. Since then there has been conflict in the literature between
adopting Pacheco's system or reverting to the earlier conservative approach of few genera. LeSage
(1991) and Katovich (2002) followed Pacheco, whereas Miller (1996, 1998), King, Starr and Lago
(2011) and Lago and King (2012) recognized only two or three genera and Lago and King (2012)
synonymized 11 of Pacheco's genera with Heterocerus and three with Augyles. However, they then
recognized that there are groups of similar species within these two genera and used an informal and
loosely conceived species group structure. Their implied classification is difficult to apply across the
North American fauna and their dismissal of Pacheco's work looses much that is good and useful. I
prefer a broad generic concept that can be mastered fairly easily by the nonspecialist and for this reason
follow the conservative approach of recognizing only a few genera. On the other hand, a suite of
characters suggests that Pacheco's groups, or at least most of them, are natural units and they certainly
have utility in ordering the species. Thus, I have used Pacheco's names at the subgeneric level, more or
less synonymously with the species groups of other authors.

Key to Genera of Heteroceridae known or likely to occur in Saskatchewan

1. Post-metacoxal lines present. ............................................................... Augyles Schiödte, Key A

1'. Post-metacoxal lines absent. ....................................................................................................... 2

2(1). Post-mesocoxal line absent ........................................................................................................ 3

2'. Post-mesocoxal line present ........................................... Heterocerus Fabricius (in part), Key B

3(2). Antenna 9-segmented (serrate club of 5 segments); size small, L = 2.3 to 3.0 mm; elytra
without trifasciate color pattern, either unifomly reddish brown to almost black or gradually
pale laterally; male genitalia spatulate with proximal third narrow and cylindrical, distal half to
two-thirds broadened ...................................................................................... Tropicus Pacheco
[Three species occur in the US and eastern Canada but none is known from the Canadian
Prairies. Tropicus pusillus (Say) is reported as abundant in the southeastern US, and ranges
north to ON, KS and CO]

3'. Antenna 11-segmented; larger, L = 3.5 to 7.0 mm; elytra patterned, usually dark with 3
transverse fasciae (complete or broken into rows of spots) located basally, medially and
apically, or some specimens with pale areas expanded leaving darker interfasciate spots; male
genital capsule subquadrate to cordate with a short proximal shaft.
........................................................................................... Heterocerus (Neoheterocerus), Key B
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Key A. Genus Augyles Schiödte

The members of this genus are of diverse form but can be easily recognized by the presence of
the post-metacoxal lines on sternite 1. Also, the male genitalia is of a similar form, being of tubular
shape with a trough-like ventral phallobase over which lies the elongate dorsal aedeagus. The aedeagus
distally is reflexed as a tongue-like process that is variously armed with setae, spines or papillae. This
process is usually visible within the body of the genital capsule but it is hinged and can be extended
distally. The distal end of the phallobase is bifurcate.

Size, color and density and type of vestiture vary considerably within the group. Pacheco
(1964, 1978) placed the five prairie species known at the time into three genera: Explorator Pacheco,
Centuriatus Pacheco and Microaugyles Pacheco, which are here treated as subgenera. A more recently
described species, A. blanda Miller, is unplaced.

Key to species of Augyles Schiödte known or likely to occur in Saskatchewan

1. Small (L = 2.2 to 3.1 mm), narrow (L/W = 2.47 to 2.57) species; antenna 10-segmented (i.e.
serrate club with 6 segments) ..................................................................................................... 2

1'. Larger (L = 3.0 to 5.5 mm), broader (L/W = 1.96 to 2.46) species; antenna 11-segmented but
basal 1 or 2 segments of club may be similar to funicle segments ........................................... 3

2(1). Pronotum with basal bead very faint to obsolete laterally; epipleural carina absent or at most
weakly indicated; mesosternum between mesocoxae narrow, its width subequal to width of a
coxa measured along same line; dorsal surface finely and densely punctate, pubescence fine,
dense and suberect; elytra with diffuse fasciate markings, pronotum darkened subbasally each
side of paler median line; L = 2.5 to 3.1 mm; L/W = 2.47 to 2.57.
.......................................................................................Augyles (Microaugyles) mundulus (Fall)
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Genus Subgenus Species Author CU DG MG PL BP BS

Augyles Microaugyles mundulus (Fall) 0 1 0 0 0 0
H: Specimens conform well with Fall's (1920) description of the species in characters other
than color. Fall described the species as “flavo-testaceous throughout, elytra with a small,
vague fuscous shade behind the scutellum”. Prairie specimens have at least a vague
suggestion of fasciate elytral markings and the pronotum is darkened, especially sub-
basally each side of the paler medial area. Pacheco's (1964) illustration shows an
intermediate color pattern. Fall did not mention the number of antennal segments, the
character used by Pacheco (1964) in defining Microaugyles.
The pale specimens of A. mundulus were described from El Paso, Texas. Subsequent
records include CA, CO, NM, TX (Pacheco 1978) and AB (Bousquet 2013),
Specimens have been collected on bare sandy-clay banks of the Milk River, AB, and the
Frenchman R., SK.

Illustration 4: Augyles
(Microaugyles) mundulus

Illustration 5: Augyles (Microaugyles)

mundulus, male genitalia. A - genital capsule
dorsal; B - aedeagus; C - phallobase; D -
genital capsule ventral with sternite 9.

2'. Pronotum with basal bead distinct laterally; epipleural carina well developed and extending
across at least inner half and usually most of epipleural width; mesosternum between
mesocoxae broader, its width greater than width of a coxa measured along same line; elytron
with dual punctation consisting of relatively coarse punctures separated by about their own
diameters and interspaces minutely punctulate, pubescence of elytra fine, short, yellowish and
recurved, darker and erect on pronotum; elytra with irregular, often interrupted, pale fasciae of
the usual type but varied in development; pronotum piceous with sides and median line paler; L
= 2.2 to 3.0 mm; W = 1.0 to 1.3 mm. ...........................Augyles (Microaugyles) moleculus (Fall)

Genus Subgenus Species Author CU DG MG PL BP BS

Augyles Microaugyles moleculus (Fall) 0 0 0 0 0 0
H: This species was described from Aweme, MB, and subsequently recorded from WA,
WY, CO, NE, KS, WI, IL (Pacheco 1978, Miller 1996)) and AB, SK and PQ (Bousquet
1991, Bousquet et al. 2013). I have not seen specimens. The key characteristics given
above are from the original description and Pacheco (1964). The antenna are reported to
be 10-segmented (Pacheco 1964, Miller 1996).
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3(1). Maxilla with galea broad, flattened with apical margin capped with a dense brush of golden
setae; body stout, broadly ovate, L/W = 1.95 to 2.2; pronotum and elytra with short,
appressed, flattened and scale-like golden setae and with or without fine suberect setae; elytra
with trifasciate pattern usually well developed with distinct orange markings which on some
specimens may be expanded to cover much of disc, pronotum piceous with midline and lateral
margins pale. .............................................................................................................................. 4

3'. Maxilla with galea narrower, apically with a row of about 5 to 10 apically curved yellow setae;
body shape various but generally less robust; dorsal surface with both appressed and distinct
erect setae; elytron and pronotum color various ........................................................................ 5

4(3). Prosternum in lateral aspect with ventral margin of midline and prosternal process more or less
straight and in same plane; dorsal surface with short, appressed golden scale-like hairs and
devoid of sparse, fine, semierect setae; smaller on average, L = 3.4 to 3.9 mm; L/W = 1.96 to
2.17. ............................................................................. Augyles (Centuriatus) compactus (Fall)
Genus Subgenus Species Author CU DG MG PL BP BS
Augyles Centuriatus compactus (Fall) 0 1 0 1 0 0
H: This is the most distinctive species of heterocerid in the fauna. The usual dense
covering of setae is represented by short, adpressed golden scale-like setae which are
even smaller and sparser on the ventral surface. The only elongate erect setae, other than
those of the legs and mouthparts, are on the frons, a pair between the eyes, a pair on the
disc of the pronotum and one pair and two pairs respectively on visible sternites 4 and 5.
The galea of the maxilla is broad and flattened and bears a brush of stout golden setae, a
characteristic of the subgenus. The aedeagus has the same pair of apical hooks and
reflexed setose tounge-like structure as shown by A. blanda.
These beetles have been found on sparsely vegetated, sandy creek banks, 0.5 to 2 m from
the water line. Specimens were collected by splashing water on the bank and flooding the
insects out. Unlike other heterocerids, specimens did not fly when flooded, rather they ran
up the bank or dug back into the sand, and in terms of color and movement are reminiscent
of dermestid beetles.

Illustration 6: Augyles
(Centuriatus) compactus.

Illustration 7: Augyles (Centuriatus)

compactus, male genitalia. A - genital capsule
dorsal; B - aedeagus dorsal; C - phallobase.
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4'. Prosternum in lateral aspect with prosternal process strongly arcuate and not forming a straight
line with ventral margin of precoxal prosternum; dorsal surface of body with short appressed
golden setae but also with fine semierect hairs (may have to be examined in correct light or
profile to be visible); larger on average, L = 3.5 to 5.5 mm.
...................................................................... Augyles (Centuriatus) auromicans (Kiessenwetter)

Genus Subgenus Species Author CU DG MG PL BP BS

Augyles Centuriatus auromicans (Kiessenwetter) 0 0 0 0 0 0
H: This species has a wide distribution, being reported from BC to the Atlantic coast and
south to Texas (Pacheco 1964) and AL (King & Lago 2012). Pacheco (1964) indicated AB
and MB records but no SK record. The records by Bousquet (1991), Hooper & Larson
(2012) and Bousquet et al. (2013) could be based on A. compactus and need confirmation
as I have not seen SK specimens.
Fall (1937) described this species as differing from A. compactus as follows: larger on
average, short erect hair present on the elytra, with faint elytral striae in addition to sutural
stria, pronotum less rounded and narrowed anteriorly, and the prosternal process
conspicuously arched in lateral aspect.
Kaufmann (1987) has described behavior and mating characteristics of the species.

5(3). Body mainly piceous to black except with pale anterolateral angles of pronotum and faint to
sharply defined yellow fasciae on elytra; elytra bristly with long, suberect setae and an under-
layer of decumbent setae; elytron surface shiny with large conspicuous punctures, its lateral
margin with a fringe of elongate setae; L = 3.1 to 3.8 mm, L/W = 2.15 to 2.32.
......................................................................................... Augyles (Explorator) canadensis (Fall)

Genus Subgenus Species Author CU DG MG PL BP BS

Augyles Explorator canadensis Fall 1 0 1 0 0 0
H: Three specimens examined, collected on patches on bare organic silt or clay along a
beaver pond, a slow creek and a large prairie river.

Illustration 8: Augyles
(Explorator) canadensis Illustration 9: Augyles (Explorator)
canadensis, male genitalia. A - genital capsule
dorsal; B - aedeagus lateral showing reflexed
apical projection; C - phallobase; D - genital
capsule with sternite 9, ventral.
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5'. Color paler, much of elytra and legs testaceous to reddish-brown, pronotum of many specimens
with medial line broadly paler than lateral areas of disc; elytra with short, appressed golden
setae plus much sparser suberect longer setae; lateral margins of pronotum and elytra only
sparsely fringed with erect setae; L = 3.1to 3.5 mm, L/W - 2.3 to 2.5.
............................................................................................... Augyles (subgenus?) blanda Miller
Genus Subgenus Species Author CU DG MG PL BP BS
Augyles unplaced blanda Miller 0 1 0 0 0 0
H: Miller (1994) did not assign this species to a subgenus but he did state it was closely
related to A. canadensis. Also, he described the antenna as 10-segmented. The
specimens I have examined have 11-segmented antenna although segments 5 and 6 are
smaller and less asymmetrically expanded than usual so one could easily get a lower
segment count if only the well defined club segments were considered. It is chiefly in the
coarse punctation of the elytra that there is a similarity to A. canadensis. In color, reduced
erect setae of the dorsal surface of the body, and structure of male genitalia, this species
resembles A. auromicans and A. compactus, but it lacks the brush of setae on the galea.
Most of the specimens examined by Miller (1994) were collected on the shores of muddy
creeks and rivers. I have seen specimens from muddy banks of the lower reaches of the
Red Deer River, Alberta.

Illustration 10: Augyles blanda

Illustration 11. Augyles blanda, male

genitalia. A - genital capsule, dorsal; B -
aedeagus dorsal with apical process extended;
C - genital capsule, ventral; D - sternite 9.
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Key B Genus Heterocerus Fabricius

All Saskatchewan species that lack the post-metacoxal lines on sternite 1 belong to this genus.
Pacheco's (1964) classification would have the these species in five genera, based largely on the
presence or absence of the post-mesocoxal line and structure of the male genitalia. There are two
major forms of male genitalia within this group, that of the subgenus Dampfius in which the phallobase
has distally elongated lateral arms and its basal piece is prolonged distally, and the form shown by the
rest in which the phallobase is widest across proximal lateral angles and the parameres are small pale
lobes on the distal margin of the basal piece.

Key to species of Heterocerus Fabricius known or likely to occur in Saskatchewan

1. Post-mesocoxal line absent. ...................................................... Heterocerus (Neoheterocerus), 2

1'. Post-mesocoxal line present. ....................................................................................................... 6

2(1). Elytral surface rather shiny, with two or three evidently impressed longitudinal impressions or
striae lateral to sutural margin, convex ridges between striae conspicuously coarsely and usually
roughly punctate on basal half; pronotum with anterolateral angle and usually a narrow lateral
margin yellow; elytron with dark areas of disc expanded laterally and contacting or only
narrowly separated from lateral margin, especially the medial and preapical dark bands; male
genitalia with phallobase relatively broad, distal width at level of basal piece only slightly
narrower than its medial width; median plate with dark sclerites each side of distal tip;
parameres in lateral view curved ventrally, in ventral view median edges diverge slightly and
outer margin concave; smaller, L = 3.5 to 4.1 mm; L/W = 2.29 to 2.50 (n = 11,
hypermandibulate male not seen). (Note: - Pacheco 1964 gives L = 4.3 to 5.7 mm)
................................................................... Heterocerus (Neoheterocerus) sandersoni (Pacheco)
Genus Subgenus Species Author CU DG MG PL BP BS
Heterocerus Neoheterocerus sandersoni (Pacheco) 0 1 1 1 1 0
H: Specimens of this species can usually be recognized by the evident elytral striae and
coarsely punctate interspaces. However, many members of other species of the subgenus
also have more or less distinct striae but when present the punctation of the elytral disc is
more regular and finer.
A species of southeastern Canada and eastern US, west to MB, ND and MT (Pacheco
1964). Recorded from SK by Miller 1996 and Bousquet et al. 2013. Uncommon in SK.
Specimens labelled as being collected in pitfall trap (3), in flooded stream bed and on a
sticky trap on elm (indication of flight). No specimens were collected at light although King
and Lago (2013) report it as very common at light in se US.

Illustration 12. Heterocerus

(Neoheterocerus) sandersoni
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Illustration 13: Heterocerus (Neoheterocerus) sandersoni, male genitalia. A - genital

capsule, dorsal; B - aedeagus; C - phallobase; D - genital capsule, ventral.

2'. Elytral surface lacking longitudinal impressions or if present the convex ridges between striae
have punctation similar in density and size to that on remainder of disc; pronotum and elytra
with color various; phallobase narrower, lateral margins distinctly constricted and sinuate
distally, width at level of basal piece 0.8 or less medial width; median plate without distinct
dark sclerites each side of distal tip; parameres in lateral view flat, in ventral view outer margin
straight to slightly convex. ...................................................................................................... 3

3(2). Phallobase with ventral plate in distal third distinctly broadened and subangulate then concavely
narrowed to rounded apex. .......................... Heterocerus (Neoheterocerus) fatuus Kiesenwetter
Genus Subgenus Species Author CU DG MG PL BP BS
Heterocerus Neoheterocerus fatuus Kiesenwetter 0 0 0 0 0 0
H: King and Lago (2012) give additional characteristics for the species as: color dark
brownish-red; pronotal disk dark without pale margins; elytral fasciae usually broken into a
series of spots; L = 4.2 to 5.0 mm, hypermandibulate males rare.
Recorded from SK by Bousquet et al. (2013). However, Pacheco (1964) and King and
Lago (2012) indicate the distribution as being mainly coastal, along the Gulf coast of central
Mexico and the coasts of Mississippi, New York and New Brunswick. King & Lago (2012)
report it as being collected near saline and freshwater bodies of water. Without
examination of the genitalia this species is very difficult to identify and the prairie records
are probably due to misidentification. Nevertheless, the occurrence of a coastal species
at the margins of saline lakes is a possibility that should be investigated.

Illustration 14: Heterocerus

(Neoheterocerus) fatuus,
phallobase, ventral.
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3'. Phallobase with lateral margins of ventral plate slightly and evenly curved, subparallel or
converging distally, but not evidently broadened and subangulate. ............................................ 4

4(3). Phallobase with lateral margins of ventral plate evenly tapered and converging to a distal point;
color dark brown to piceous, pronotum with lateral margin dark, shortly pale anteriorly or
narrowly pale along entire length; elytra with fasciae usually broken into spots, lateral margin
narrowly pale but on many specimens with a lateral infuscation between basal and median
fasciae; relatively small, L = 3.5 to 4.5 mm, hypermandibulate males not seen.
.......................................................................... Heterocerus (Neoheterocerus) infrequens Miller

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Neoheterocerus infrequens Miller 0 1 1 0 0 0
H: Described from near Roche Percee, SK (Miller 1996). Two species of Neoheterocerus ,
infrequens and angustatus Chevrolat, have the ventral plate of the phallobase evenly
tapered distally. Miller (1996) states they differ in that the point of the plate meets the basal
plate of the phallobase in infrequens and does not meet it in angustatus. In some SK
specimens the point is short of the basal piece and thus the specimen should be
angustatus. H. angustatus is a species of the Caribbean and the se US and it is likely that
either infrequens is variable in this phallobase character or the two forms are conspecific.
The limits between infrequens and gnatho are obscure for a range of median plate shapes
occur iwithin the province and it may prove that the infrequens state is just the end in a
continuum.
The species is well named as it is uncommon in collections. Habitat data include mud
puddle, sandy lake shore, muddy creek, shore of river. Collections are from grassland
areas, especially in drier areas. I collected a specimen on the clay bank of a roadside ditch
at Old-Man-on-His-Back Reserve.

Illustration 15: Heterocerus

(Neoheterocerus) infrequens.

Illustration 16: Heterocerus

(Neoheterocerus) infrequens, male
genitalia. A - genital capsule,
dorsal; B - phallobase, ventral.
 14

4'. Phallobase ventral plate with lateral margins subparallel for much of length, narrowed distally
to a blunt point which is separated from the basal plate of the phallobase. ................................ 5

5(4). Male genitalia with right distal margin of dorsal plate (aedeagus) relatively deeply notched,
medial edges of parameres diverging slightly towards their apicies; phallobase with ventral plate
narrower, lateral margins more or less straight medially; color of body darker but highly varied,
pronotum with dark area covering entire disc, with anterolateral angle pale, or with lateral
margin partly or entirely pale but if entirely pale, pale margin usually narrow but on some
specimens wide and conspicuous; elytra with dark portion of pattern predominant; L = 4.1 to
6.4 mm, hypermandibulate males common. .... Heterocerus (Neoheterocerus) gnatho LeConte

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Neoheterocerus gnatho LeConte 1 1 1 1 0 0
H: This species has been recorded from SK by Pacheco (1978), Bousquet (1991), Hooper
& Larson (2012) and Bousquet et al (2013).
King and Lago (2012) point out that although color, especially that of the pronotum, may be
useful to distinguish between gnatho and pallidus, both species vary greatly and only
characters of the male genitalia are trustworthy for separating them. However, even the
genitalic characters are subtle.
Pacheco (1964: Fig. 494) shows H. gnatho to be a northern and western species,
occurring mainly in northern US and southern Canada as well as west of the Rocky
Mountains whereas H. pallidus is southern and eastern. However he records both species
in SK. Saskatchewan experience supports this pattern for gnatho is abundant and
widespread whereas pallidus is less common and occurs mainly in the south.
The vast majority of heterocerids in collections belong to this species and H. (Lapsus)
fenestratus. Habitats from which specimens have been collected include organic mud, clay
and fine sand at the border of fresh and saline ponds as well as the margins of streams.
Many series have been collected from UV light.

Illustration 17: Heterocerus

(Neoheterocerus) gnatho,
male genitalia exserted.

Illustration 19: Heterocerus

Illustration 18: Heterocerus (Neoheterocerus) gnatho,
(Neoheterocerus) gnatho, male genitalia
male genitalia: A - genital capsule, dorsal; B - aedeagus,
with aedeagus reversed to sinistral position:
dorsal; C - phallobase, ventral; D - genital capsule, ventral.
A - genital capsule, dorsal; B - aedeagus,
dorsal; C - phallobase.
 15

5'. Male genitalia with right distal margin of dorsal plate more shallowly notched; parameres
proximate and parallel to one another; phallobase with ventral plate broader, its lateral margins
usually distinctly convex; body color paler with yellow portion of pattern more extensive,
pronotum with lateral margins broadly yellow and dark area of disc not contacting lateral
margin, elytra of many specimens with pale pattern predominant; L = 4.5 to 6.9 mm,
hypermandibulate males common. ...................... Heterocerus (Neoheterocerus) pallidus Say
Genus Subgenus Species Author CU DG MG PL BP BS
Heterocerus Neoheterocerus pallidus Say 1 1 1 1 0 0
H: See comments above under H. gnatho regarding the separation of the two species in
SK. In many populations of H. gnatho specimens have the lateral margins of the pronotum
completely bordered by yellow but the yellow border is usually narrower and the dark
maculations of the elytra more extensive than in H. pallidus.
The largest and palest specimens, and thus the specimens I most confidently assign to this
species, were collected on silty sand at the margin of gravel pit pools in a river valley.
Specimens have been found in a variety of moist sites beside both standing and running
water, including at the edges of saline lakes,

Illustration 20: Heterocerus

(Neoheterocerus) pallidus,
hypermandibulate male.

Illustration 22: Heterocerus (Neoheterocerus) pallidus, male

genitalia: A - genital capsule, dorsal; B - aedeagus, dorsal; C -
phallobase, ventral; D - genital capsule, ventral.

Illustration 21: Heterocerus

(Neoheterocerus) pallidus,
female.
 16

6(1). Elytral fasciae each composed of 2 or 3 more or less separated longitudinally stretched spots,
the innermost spot of the subbasal fascia extending anteriorly to basal margin just lateral of
scutellum, or if basal margin of elytron without pale spots (some melanistic individuals),
medial fascia with the two innermost spots strongly elongate in shape; pronotum piceous to
black with lateral margin pale mesad to bead; legs strikingly bicolored, femora mainly
yellowish, remainder piceous or tibiae diffusely pale medially; L = 3.4 to 4.5 mm; L/W =
2.12 to 2.52; hypermandibulate males occur but usually mandibles only slightly
elongated. ........................................................Heterocerus (Lapsus) fenestratus (Thunberg)

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lapsus fenestratus Thunberg 1 1 1 1 1 1
H: This is an abundant and easily recognized species. The typical color pattern as
described above is distinctive. However there are melanistic as well as unusually pale
individuals that may pose difficulty in recognition. Specimens with expanded pale markings
on the elytra resemble some specimens of Neoheterocerus but the presence of the post-
mesocoxal line in Lapsus is distinctive. Dark specimens may resemble some specimens
of Lanternarius but in Lapsus the elytral punctation and setal arrangement is uniform and
there is never a cow-lick in the setae posterior to the medial fascia, whereas there is
usually an evident cow-lick pattern in Lanternarius. Also, the elongate triangular shape of
the male genitalia is unlike that of any other species.
This is the only Holarctic species in the fauna. It was previously treated as H. (L.) tristis but
has been shown to be conspecific with the Palaearctic H. fenestratus (Mascagni 1993, King
& Lago 2012).
Widely distributed and one of the two (with H. gnatho) most abundant heterocerids in the
province. Specimens occur on various types of open, moist soil near both standing and
running water. Most specimens examined were collected at light.

Illustration 23: Heterocerus

(Lapsus) fenestratus.

Illustration 24: Heterocerus (Lapsus) fenestratus, male genitalia: A -

genital capsule, dorsal; B - aedeagus, dorsal; C - phallobase, ventral;
D - genital capsule, ventral.
 17

6'. Elytral fasciae various, if broken into two or more spots, spots not strongly longitudinally
stretched and inner spot of basal fascia not extended anteriorly to basal elytral margin, base
of elytron with color various, entirely dark, pale with pale area diffusely extended posteriorly, or
some specimens with a distinct isolated juxtascutellar spot. .....................................................7

7(6). Smaller (L = 2.7 to 4.5 mm), narrow species; color paler, dorsal surface pale reddish brown to
rufo-piceous, pronotum never black, darkest basally each side of middle, lateral margins pale
and usually medial line and along anterior and basal margins paler than darkest area of disc;
elytron usually with pale markings on disc, fasciate markings present but may be poorly
delimited on paler specimens; elytron with punctation and vestiture dense and uniform over
disc; phallobase with lateral arms directed posteriorly on either side of distally extended basal
piece ....................................................................................................Heterocerus (Dampfius), 8

7'. Larger (3.7 to 6.1 mm) and usually broader species; color darker, pronotum rufo-piceous to
black medially and not evidently paler along medial line but pronotum may be paler than dark
color of elytra; elytron ground color rufo-piceous to black, base with or without pale
juxtascutellar spots, fasciae various, absent to well developed, limits of pale spots usually well
defined; phallobase with lateral angulate protrusions proximad of middle.
.................................................................................................... Heterocerus (Lanternarius), 13

8(7). Male genitalia in ventral aspect with lateral arms of phallobase directed distally with the apex of
each outwardly curved to an acute point. .................................................................................. 9

8'. Male genitalia in ventral aspect with lateral arms directed distally but their apex not recurved
outwardly to an acute point ..................................................................................................... 10

9(8). Male genitalia in ventral aspect with median plate of phallobase gradually narrowed and curved
upwards in distal third; lateral arm of phallobase curved into an outwardly directed acute point
far short of level of apex of median plate; parameres membranous and fused, their bilobed apex
prolonged distally beyond tip of median plate; L = 3.0 to 4.2 mm.
..................................... ................................ Heterocerus (Dampfius) collaris Kiesenwetter

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius collaris Kiesenwetter 0 0 0 0 0 0
H: Brown to reddish-brown; elytra fasciate but pale areas may be enlarged and their limits
poorly defined; basal margin pale.
Recorded from SK by Bousquet (1991), Hooper and Larson (2012) and Bousquet et al.
(2013), all likely in error. Specimens that I have seen identified as H. collaris have proven
to be other species. Both Pacheco (1978) and Miller (1988) indicate this is an eastern
species that does not occur in the northern Great Plains and therefore I delete it from SK
list.

Illustration 25: Heterocerus

(Dampfius) collaris, male
genitalia.
 18

9(8). Male genitalia in ventral aspect with median plate of phallobase abruptly narrowed and ending
in an acute point short of distal level of apicies of lateral arms of phallobase; parameres
membranous, small and pointed and reaching slightly beyond tip of median plate but not
attaining level of apex of lateral arms; L = 3.2 to 4.5 mm .
.............................................................................. Heterocerus (Dampfius) undatus Melsheimer

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius undatus Melsheimer 0 0 0 0 0 0
H: Color brown to reddish brown; elytron with trifasciate pattern but limits of pale fasciae
usually poorly defined.
Recorded from SK by Bousquet (1991), Hooper and Larson (2012), and Bousquet et al.
(2013) probably from misidentification. Pacheco (1978) and Miller (1988) record it as an
eastern species that extends west to MB. Deleted from SK list.

10(8). Male genitalia with lateral arms of phallobase extending distally well beyond apex of
parameres; dark reddish-brown with bright pale area on basal margin of elytra, this spot usually
isolated from lateral margin and basal fascia; L = 3.5 to 4.0 mm.
..........................................................................................Heterocerus (Dampfius) subtilis Miller

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius subtilis Miller 0 0 0 0 0 0
H: This species is not known from SK. It is an eastern species but has been recorded
from MB at Winnipeg and Tp 17 Rge 29 W 1 (near St. Lazare, close to SK/MB border)
(Miller 1988).

Illustration 26: Heterocerus

(Dampfius) subtilis, male
genitalia: A - dorsal; B -
ventral.

10(8). Lateral arms of phallobase not extending distally beyond apicies of parameres. .................... 11
 19

11(10).Lateral arms of phallobase as long as apicies of parameres; aedeagus with median plate truncate
posteriorly. ....................................................... Heterocerus (Dampfius) inciertus (Pacheco)

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius inciertus (Pacheco) 1 1 0 0 0 0
H: Identification based on rather unsatisfactory key given by Miller (1988). The genitalia
do not exactly conform to the drawings of Pacheco (1964) for H. inciertus so this
identification needs to be checked. Pacheco (1964) recorded the species from AB and MB
but there are no previous SK records. Specimens collected along stream banks on mainly
bare sand or sandy-clay soil.

Illustration 27: Heterocerus

(Dampfius) inciertus.

Illustration 28: Heterocerus (Dampfius)

inciertus, male genitalia: A - genital capsule,
dorsal; B - aedeagus, dorsal; C - phallobase,
ventral.

11'. Lateral arms of phallobase shorter than apicies of parameres. ................................................. 12

12(11).Aedeagus with median plate bluntly rounded distally; reddish-brown with a bright pale area on
basal margin of elytron, on some specimens expanded to lateral margin or posteriorly to contact
basal fascia; L = 4.0 to 4.5 mm. ...................................... Heterocerus (Dampfius) unicus Miller
 20

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius unicus Miller 0 1 0 0 0 0
H: Recorded from nr. Willow Bunch (Tp 5 Rge 28 W 3) SK by Miller (1988). I have not
seen specimens.

Illustration 29: Heterocerus

(Dampfius) unicus, male
genitalia.

12'. Aedeagus with median plate sharply pointed distally; darker species with pale area on base of
elytra. .... ................................................................... Heterocerus (Dampfius) mexicanus Sharp

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Dampfius mexicanus Sharp 0 0 0 0 0 0
H: Not recorded from SK but with a wide distribution in the western US, occurring north to
MT, SD and ON.

13(7). Elytron base entirely dark, lacking juxtascutellar pale spot; pronotum either piceous to black
with at most anterolateral angle paler, or entire pronotum distinctly paler than elytra and with
pale lateral margins. ................................................................................................................. 14

13'. Elytron base with a pale juxtascutellar spot; pronotum black with lateral bead and usually
margin adjacent to bead, yellow; propleuron yellow; elytron with fasciae broad, sharply
delimited and contacting lateral margin; epipleuron yellow; legs mainly yellow; L = 4.5 to 5.5
mm (hypermandibulate males reported from eastern North America have not been seen from
SK) ............................................................. Heterocerus (Lanternarius) mollinus Kiesenwetter
 21

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lanternarius mollinus Kiesenwetter 1 1 1 1 0 0
H: SK specimens are darker in color than those described from se US by King & Lago
(2012), but the general elytral pattern and structure of the male genitalia are similar
indicating they are conspecific.
King & Lago (2012) consider this to be the most common species in the eastern United
States, where it can dominate light trap collections. They also report it as common near
rivers and ponds of all sizes.
The species is uncommon in SK where I have seen five locality records, all in the south.
Specimens have been collected from the banks of Battle Creek, both within and
downstream of the Cypress Hills. They were taken by splashing bare yellowish clay or
sandy-clay banks with water to flush the beetles out.

Illustration 30: Heterocerus

(Lanternarius) mollinus.

Illustration 31: Heterocerus (Lanternarius) mollinus, male

genitalia: A = genital capsule, dorsal; B - aedeagus, dorsal; C -
phallobase, ventral; D - genital capsule, ventral.

14(13).Elytral epipleuron with an oblique subhumeral carina posterior to shoulder (carina may vary in
development from a small ridge on mesal margin of epipleuron to a strong ridge that extends
across the entire width of the epipleuron). ................................................................................ 15

14'. Elytral epipleuron lacking an oblique subhumeral carina. ........................................................ 16

15(14). Elytral epipleuron with subhumeral carina strongly developed and extending almost entire
width to lateral margin; elytron lateral margin at shoulder microserrate with fringing setae each
set on a sharp spinose projection; body dorsally with dense adpressed yellow setae and more
sparse and rather fine and inconspicuous erect brown setae; elytral setae not conspicuously
whorled behind medial fascia; elytral fasciae conspicuous yellow to orange and well developed;
body large, L = 4.8 to 6.4 mm (larger specimens hypermandibulate males)
................................................................................. Heterocerus (Lanternarius) gemmatus Horn
 22

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lanternarius gemmatus Horn 0 0 0 0 0 0
H: Recorded from BC, AB and SK by Bousquet (1991). I have seen specimens from BC
and southwestern AB (Waterton) but am not able to confirm SK records. All SK specimens
with a subhumeral carina seem to be H. (L.) brunneus.
Specimens have been collected in BC and AB from muddy banks of backwaters and
embayments of low gradient streams.

Illustration 32: Heterocerus

(Lanternarius) gemmatus.
Illustration 33: Heterocerus (Lanternarius) gemmatus, male
hypermandibulate male.
genitalia: A - genital capsule, dorsal; B - aedeagus, dorsal; C -
phallobase, ventral; D - genital capsule, ventral.

15'. Elytral epipleuron with subhumeral carina various, often shorter than half width of epipleuron;
shoulder margin of elytron somewhat granulate but setae not clearly arising from spinose
projections; dorsal vestiture with decumbent yellow setae sparser, with denser erect piceous
setae and body appearing more bristly, most specimens with decumbent elytral setae behind
anterior arc of medial fascia evidently whorled or with a cowlick; elytral fasciae obsolete or
poorly defined, often only evident through presence of pattern of yellow setae over area
typically occupied by fasciae; smaller, L = 4.1 to 4.8 mm (hypermandibulate males not seen)
......................................................... Heterocerus (Lanternarius) brunneus Melsheimer (in part)
 23

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lanternarius brunneus Melsheimer 1 0 0 0 0 1
H: This mainly western species has been recorded from BC and NT to MB and MT. I have
seen only a few specimens from western SK (Cypress Hills, Lake Athabasca). All SK
specimens examined had a more or less well defined subhumeral epipleural carina, a
character that occurs in other populations but less consistantly. The presence of this carina
could be the basis of the SK records of H. (L.) gemmatus in the province.
Single specimens have been collected at the margins of streams draining the south sides
of the Cypress Hills.

Illustration 34: Heterocerus

(Lanternarius) brunneus.
Illustration 35: Heterocerus (Lanternarius) brunneus (Kitamat,
BC), male genitalia: A - genital capsule, dorsal; B - aedeagus,
dorsal; C - phallobase, ventral; D - genital capsule, ventral.

Illustration 36: Heterocerus

(Lanternarius) brunneus (SW SK),
male genitalia: A - aedeagus,
dorsal; B - phallobase, ventral.
 24

16(14).Pronotum light reddish-brown, paler in color than darker brown elytra; male genitalia with
medial plate of phallobase with a broad, square attachment to body of phallobase; L = 3.8 to
4.7 mm (from King & Lago 2012). .................. Heterocerus (Lanternarius) sinuosus (Pacheco)

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lanternarius sinuosus (Pacheco) 0 0 0 0 0 0
H: Pacheco (1964) shows records for BC, MB and MT; King and Lago (2012) describe the
distribution as southern Canada and the US west of the 100th meridian. Recorded from YT,
NT and BC to ON (including SK) by Bousquet et al. (2013). I have not seen SK specimens.
The species should be readily recognized by the pale color, especially that of the pronotum,
and the male genitalia in which the medial plate is broadly attached distally to the main
body of phallobase, a condition that does not occur in other species of Lanternarius. The
phallobase is more or less evenly narrowed distally and the parameres are narrowly
separated with their mesal margins more or less parallel.

Illustration 37: Heterocerus

(Lanternarius) sinuosus, male
genitalia: A - genital capsule,
dorsal; B - phallobase, ventral.
(From King & Lago 2012).

16'. Pronotum with disc piceous to black, as dark or darker in color than elytra, at most with
anterolateral angle paler; male genitalia with basal plate of phallobase narrowed
distally. .......................................................................................................................................17

17(16). Body dorsally with erect setae longer and appearance somewhat shaggy; elytra with pale spots
generally small and poorly defined, obsolete on many specimens and evident mainly due to
paler setae occurring on typically fasciate portions of disc; male genitalia with basal piece of
phallobase subparallel and parameres narrowly separated.
................................ Heterocerus (Lanternarius) brunneus Melsheimer (in part, see couplet 15)

17'. Body with erect setae shorter and more uniform giving a more trim appearance; elytra with
fasciae usually broken into a series of spots but limits of spots well defined; male genitalia with
phallobase distinctly widened distally and parameres separated basally by a broad semicircular
notch; L = 3.7 to 4.5 mm, hypermandibulate male not seen.
........................................................................... Heterocerus (Lanternarius) parrotus (Pacheco)

Genus Subgenus Species Author CU DG MG PL BP BS

Heterocerus Lanternarius parrotus (Pacheco) 1 1 1 1 1 1
H: Color various but generally pattern of elytron more strongly developed than on
brunneus; elytron base lacking juxtascutellar spot, fasciae variously developed, on many
specimens broken into spots, sometimes spots obscure or obsolete; lateral margin and
epipleuron dark; legs usually somewhat bicolored, piceous with reddish areas medially on
femora and sometimes also tibia. Most specimens with the post-medial elytral cowlick well
developed.
This species has been recorded from BC and NT to NB. It is widespread in SK but not
abundant for specimens are usually taken singly or only short series. Specimens have
been found on the muddy banks of creeks, small dams as well as in lake wash and at light.
 25

Illustration 39: Heterocerus (Lanternarius) parrotus, male

genitalia: A - genital capsule, dorsal; B - aedeagus, dorsal; C -
phallobase, ventral; D - genital capsule, ventral.

Illustration 38: Heterocerus

(Lanternarius) parrotus.

Checklist of Saskatchewan Heteroceridae

(Note: - species names that are struckout have been recorded from SK but I have not been able to
confirm the record and the recorded distribution suggests the record was due to misidentification.
Therefore they are deleted from the SK list)

Tribe Augyliini
Augyles Schiødte
Heterocerus auctorum
Littorimus deGozis
Augyles (Centuriatus) auromicans (Kiessenwetter)
Augyles (Centuriatus) compactus (Fall)
Augyles (subgenus?) blanda Miller
Augyles (Explorator) canadensis (Fall)
Augyles (Microaugyles) moleculus (Fall)
Augyles (Microaugyles) mundulus (Fall)

Tribe Heterocerini
Heterocerus Fabricius
Heterocerus (Dampfius) collaris Kiesenwetter
Heterocerus (Dampfius) inciertus (Pacheco)
Heterocerus (Dampfius) undatus Melsheimer
Heterocerus (Dampfius) unicus Miller
Heterocerus (Lanternarius) brunneus Melsheimer)
 26

Heterocerus (Lanternarius) gemmatus Horn

Heterocerus (Lanternarius) mollinus Kiesenwetter
Heterocerus (Lanternarius) parrotus (Pacheco)
Heterocerus (Lanternarius) sinuosus (Pacheco)
Heterocerus (Lapsus) fenestratus Thunberg
(= Heterocerus tristis Mannerheim)
Heterocerus (Neoheterocerus) gnatho LeConte
Heterocerus (Neoheterocerus) infrequens Miller
Heterocerus (Neoheterocerus) pallidus Say
Heterocerus (Neoheterocerus) sandersoni (Pacheco)

Acknowledgements

I am most grateful to Dr. Paul Lago, Department of Biology, University of Mississippi, for obtaining a
copy of Pacheco's 1964 paper, and for permission to reproduce Illustrations 12 and 35.

Dr. Cory Sheffield, Curator of Entomology, Royal Saskatchewan Museum, Regina, provided a loan of
specimens, mainly collected by the late R. Hooper, in his collection.

References

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Washington. 1112 pp.

Arnett, R.H., M.C. Thomas, P.E. Skelley and J.H. Frank. 2002. American Beetles, Volume 2. CRC
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Bousquet, Y. (Ed.). 1991. Checklist of Beetles of Canada and Alaska. Research Branch Agriculture
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Bousquet, Y., P. Bouchard, A.E. Davies and D. Sikes. 2013. Checklist of beetles (Coleoptera) of
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Fall, H.C. 1920. New Coleoptera. IX. Canadian Entomologist, 52: 211 - 216.

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 27

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(Coleoptera: Dryopoidea). Systematic Entomology. 36: 435–445.

LeSage, L. 1991. Family Heteroceridae. Pp. 169 - 170 (In Bousquet 1991).

Miller, W.V. 1988. Two new species of Heterocerus from North America, with notes on related
species (Coleoptera: Heteroceridae). Coleopterists Bulletin 42: 313 - 320.

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Miller, W.V. 1996. Heteroceridae. Pp. 723-726. In Downie, N.M. and R.H. Arnett (eds.). The beetles
of northeastern North America. Volume 1. Sandhill Crane Press. Gainesville, FL. 880 pp.

Pacheco, F. 1964. Sistematica, filogenia y distribucion de los Heteroceridos de America (Coleoptera;

Heteroceridae). Monografias del Colegio de Post-graduados, No. 1, Escxuela Nacionial de Agricultura,
Chapingo, Mexico, Vol. 1, 155 pp., 501 figs.

Pacheco, F. 1978. A catalog of the Coleoptera of America north of Mexico, family Heteroceridae.
United States Department of Agriculture. Handbook 529-47. 8 pp.
28