119
EXTERX,\L SEX DIFFEREXCES IX STOHED·PRODCCTS COLEOPTERA.
By D. G. H.
HALSTEAD
Agricultural Research {';ouncil, Pe8t lnfe.qtatil)n Lavoratory,
Slough, Bucks.
(Received 26th October 196::.)
External sexual dim<lrphism is common in Coleoptpra and, if not apparent in
the adult, is usually to be found ill the pupa in the form of the genital appendages.
Size and weiaht
of newly
adults mav
""in, an imlication of their "ex but
c
._ emerae,-j
1:":1
....
is not reliable as some dearee ot overlap usually occurs.
The aim of this pape; is to bring togeth~r the klJ()\l"ll eharaeters for sexing the
adults ot stored-products beetles, and for sexihg. the pupae where adult secondary
sexual characters are apparently absent.
.---
Sexual differences in the pupal stage.
The apex of the pupal abdomen may bear two pairs of processes (fig. 14),
one, arising dorsally, representing the lan'al urogomphi. and the other, arising
ventrally, associated with the external genitalia. The former pair is not always
present but the latter pair, the genital apppndages, is u>;ually present and indicates
the sex of the pupa.
In the male, the genib1.1 appemlages are sometime::! represented by only slight
protuberances, as in Ahasver1l8 advena (\Yaltl), bllt generally they take the form
of distinct but short papilliform appendages. either two-segmented or witho1lt
apparent segmentation. The papillae are often convergent: they are never as
protuberant as those of the female.
In the female, with the t'xception of some Tenebrionids (fig. 17), the genital
appendages are always distinct and papilliforlll. The papillae arp, always stron~ly
protuberant and often three-segmented.
Generally they are divergent but in
some species they are pa1'a11",1.
As a rule. therefore, the female pupa may hr rl'adil.v distinguished from the
male by its strongly protuberant and di\'erging genital appendages.
Sexual differences in stored'products Coleoptera.
The sexual differences found in species attacking stored products are tabulated
by families (Table I).
\Vhere figures have been redrawn from publisheu works, reference to the source
is giVim in the legend to the figure.
\
~.
-
-
D. G. H. HALSTEAD.
120
TABLE 1.
Characters for the recognition of sex in stored-products Coleoptera.
Family and species
DERMESTIDAE
Dermestes fri8chii Kug.
D. penWianu,8 Cast.
D. haemorrhoidalis Kust.
D. maculatns Deg.
D. lardaritts L.
D. ater Deg.
Character and its sexual expression
Fig.
Reference
A ' - " p=t~ ~d bru"" of h";"; on tho
4th abdominal sternite :
1
Hinton, 1945
-
Hinton, 194,')
2
Hinton, 1945
J
C present
I
<jl absent
} A large puncture and brush of hairs on the
3rd and 4th abdominal sternites :
<1
Attagenns pellio (L.)
present
-, S absent
Apical segment of antennal club :
o 4 or 5 x as long as
longer
than the combined
length of the two
basal segments
<jl slightly
combined length
of the two basal
segments
-__ A. megatoma (F.)
(= piceus (01.»
Apical segment of antennal club :
<1
Anthrenns verbasci (L.)
3 or 4 X as long as
combined length
of the two basal
segments
<jl about equal in
o globular,
A. !nscus (01.)
o globular, slightly
Apical antenna! segment :
A. mnseorum (L.)
not petiolate
I
about 6 X as long
as the 7th antennal segment
-
4
Hinton, 1945
5
Hinton, 1945
<jl somewhat petio-
late
The 8th (apical) antennal segment:
<1
7
Q 3-segmented,
protuberant,
divergent
convergent
<1
-
strongly
protuberant and nearly
parallel
Pupal genital papillae :
.
6
<jl 3-segmented,
convergent
A.flavipM Lee.
(= vorax Waterh.)
Hinton, 1945
(includes
figure)
length to combined length ofthe
two basal segments
Pupal genital papillae :
".
-
<jl about twice as
long as the 7th
antennal segment
121
EXTERNAL SEX DIFFERENCES IN STORED·PRODUCTS COLEOPTERA.
TABLE I-cont .
Family and species
Cha.racter and its sexual expression
.
DERMESTIDAE-cont.
Phradorwma tricolor
(Arrow)
Antennal club :
c! indistinctly
6-segmented
Trogoder·rna granari um
Everts
Apical segment of antennal club :
I l?
o elongate
I
<;>
Fig .
Reference
-
Hinton, 1945
(includes
figure)
3
B eal,1956
-
Hinton, 1945
(includes
3-segmented
not elongate
Fusion of antennal segments is common in
both sexes but the above differences can
always be appreC'iated as the places of
fusion are apparent to' s~me degree
,
T. ver8icolor (Creutz.)
Antennal club :
c! 6-8-segmented
I
~
4-segmented
figure)
2
4
5
Figs. 1- 7.- 1, D erme stes frischii Kug ., abdomen ( .3) : ~, Attagenu.s pellio (L.),
antenna (0 . C( ) (Hinton. 1945 p. 307); 3. Trogoderma granarium Everts.
antenna (0 . C( ) (Beal. 1956 p. 561); 4. Anthrenu.s /lISClIS (01.), apical segments
of antenna (0 , C( ) (Hinton. 1945 p. 358): 5. A nthrenlls mllseorll1n (L.) . antenna
(0, 9 ) (Hinton. 19-15 p. 361): 6, Anthr e n !~s verbllsci (L.). terminal segmpnls
of pupal abdomen, ventral ,·iew (0 , C( ); 7, Anthrenu.s jlav<
ipes L ee ., the same.
122
D. G. H. HALSTEAD.
TABLE I-corli.
Character and its sexual expression
Family and species
Fig.
Reference
10
Potter, 1935
.
BOSTRYCHIDAE
Rhyzopertlw d<Jminica
(F.)
Pupal genital papillae :
-
<I
~
convergent,
2-segmented not
protuberant
divergent,
3-segmented and
protuberant
ANOBIIDAE
IFUP", gon"'"
Stegobium paniceum (L.)
p.pill•• ,
o globular. not
Lasioderma serricorne (F.)
-
-
-
':jl protuberant, 3-seg-
protuberant
PTINIDAE
men ted and distmctiy divergent
\
llfezium affine Boield.
___
.-------
Pupal,genital papillae :
'
<I
_--
Gibbium psylioide.Y
(Czenp.)
G. boieldieui Levrat
I}
J
present
I
present
I¥
I
Hinton, 1941
-
Hinton, 1941
-
Hinton, 1941
absent
A brush of long erect hairs arising from a
large, oval. shallow depression on each side
oftlle middle apical region of lith abdominal
sternite :
o absent
-
:jl absent
A median setiferous puncture on the posterior 1/5 of the metasternal disc :
<I
Pseudeurostus hilleri
(Rttr.)
ant, divergent, 3segmented with
the apical segment pointed
A large round puncture bearing an f'lrect
brush of compact hairs on the middle of the
metasternal disc :
<I
TrigonogenituJ globulu8
Sol.
¥ strongly protuber-
slightly protuberant and divergent
but adpressed to
the 9th sternite
for the greater
of
part
their
length, not distinctly segmented,
rounded apically
¥ present
123
EXTERNAL SEX DIFFERENCES IN STORED·PRODUCTS COLEOPTERA.
TABLE I-cont.
Family and species
Character and its sexual expression
Fig.
Reference
Il
Hinton, 1941
PTiNIDAE-COnt.
Niptus hololeucU8 (Fald.)
Pupal genital papillae :
<3 small and globular.
¥ distinctly
protuberant and divergent, apically
pointed. ~egmen
tat ion indistinct
parallel to "onvergent, not distinctly
segmented or
protuberant
Stethomezium squamo8um
Hinton
Pupal genital papillae :
~.~
rJ protuberant,
~
s·t'r()ngly
protuberant. divergent.
3-segmented and
pointed apically
<j2
protuberant. divergent, large 3-segmented and pointed apically
parallel to convergent, 2-segmented and
rounded apically
Tipnus unioolor
(Pill. & Mitt.)
-.·~~pal genital papillae :
Ptinus sexpunctatu8 Panz.
J
Metasternum:
J
P. tectus Boield.
convergent, small
and appearing globular (much as P.
tectus, fig. 12)
broader, with the
median
longitudinal line narrow
and
extending
from the base to
the middle of the
metasternum
¥ narrower, with the
median
longitudinal line broad
and confined to
basal fifth
Xo secondary sexual characters occur in
the adult but adult sexing can be achieved,
without injury, by squeezing out the genitalia on the day of emergence
12
Pupal genital papillae :
rJ globular
P.fur (L.)
<j2
3-segmented,
protuberant,
divergent
<3 (i) Body elongate, ¥ (i) Body obovate
suboarallel
side'ct
(ii) Pronotum
with a median
hump
(ill) Antennae
long
(ii) Pronotum
without a
median hump
(iii) Antennae
relatively
short
-
Hinton, 1941
124
D. O. H. HALSTEAD.
TABLE I-cont.
Character and its sexual expression
Family and species
Fig. Reference
PTI.lHDAE-cont.
Ptinua clauipu Panz.
~
(= hirteUua Sturm)
= ;
P. puaillua Sturm
Ii) Body elongate, ~ Ii) Body obovate
subparallel
sided
(ii) Eyes smaller
(ii) Eyes large and
prominent
(iii) Antennae
(iii) Antennae
shorter
long
'i1 (i) Body ovate
0 (i) Body slightly
elongate
Iii) Middle and
hind tibial
spurs long
and curved
-
Hinton, 1941
-
Hinton, 1941
(ii) Middle and
/'
hind tibial
spurs short
and straight
,
............
,Ir-I
·r
CLERIDAE
~.-
Nu,robia rufipea (Deg.)
-
.~
~~---------------I-----------~----------I--TROGOSITIDAE
Tenebroidea mauritanicus
(L.)
CUCUJIDAE
Cryptole8tea species
-
Hind tarsi:
ci 4-segmented
-~
I
,
'i1 5-segmented
In addition to the basic separation on tarsi,
the antennae and mandibles are useful in
C. t'~rcicus (Grouv.)
C. pusilloide8
(Steel & Howe)
C. pusillU8 (Schonh.)
C. ugan.dae
Steel & Howe
C. Jerrugineua (Staph.)
C. capensis (Waltl)
E=~=~':
~:~boU'
J
Lefkovit~h,
1959
2/3 <he
length of the body
the body
} Extemal mandibular tooth :
J ($ present
-
I
-
Lefkovitch,
1959
c;? absent
EXTER.'IAL SEX DIFFERENCES IN STORED·PRODUCTS COLEOPTERA.
11
14
Figs. 8-14.- 8. Oryzaephilus surinamensis (L.), hind leg (0, <;? ) ; 9. Ahastlerus adtlena (Waltll, terminal segments of pupal abdomen , ventral view
<0, <;? ); 10, Rhy zo perthv. dominiea (F.), the same (Potter, 1935 p. 463);
11, Ptin·us sexpunctatus Panz., median area of metasternum (0 , <;? ); 12.
Ptinus teet us Boield., terminal segments of pupal abdomen. ventral view
(0. <;? ); 13. N ec robia rujipes (Deg.). median th ird of elytron (0, <;?); 14,
T enebroides mauritanieus (L.), terminal segment s of pupal abdomen , ventral
view (0, ~ ).
125
TABLE I-cont.
Family and species
Character and its sexual expression
Fig.
Reference
SILVANlDAE
Aha8veru8 advena (Waltl)
Pupal genital papillae:
<i very small, hardly
~
9
-
protuberant, 3-segmented, divergent
visible
Cathartus quadricollis
(Ouer.)
<3 (i) Pronotum
~ (i) Pronotum
-
-
Oryzaephilw,
8urinamenai8 (L.)
6 Hind co~l process
~
8
-
elongate
(ii) Hind tibiae
with a tooth at
the apex
quadrate
(ii) Hind tibiae
without a tooth
at the apex
Hind coxal process
not conical and
conical and hind
femora
with
tooth
O. merCldor (FBUV.)
hind
"(
felllOra
without
B
tooth
,
MYCETOPHAGIDAE
;:;-"
Mycetophagu.'J
} Front tarsi :
quadriguttatU.'J (Mull.)
Typhaea steroorea (L.)
j <3 3-segmented
I
~
-
Hinton, 1945
-
Hinton, 1945
4-segmented
LATHRIDIlDAE
ConinomU.'J nodifer
(Westw.)
Hind tibiae :
6 with a tooth
Enicmus minutU.'J (L.)
I
without a tooth
Pupal genital papillae:
<3 apparently with-!
out papillae
NITIDULIDAE
CarpophilU.'J obsoletus
Erichs.
C. ligneU.'J Murr.
C. dimidiatU.'J (F.)
C. hemipterua (L.)
C. maculatU.'J MUIT.
~
~
-
-
-
Dobson,
1954
oval, apically
pointed
,
l
6th abdominal sternite :
)
ventral
and visible,
5th sternite with a
very deep round
emargination to
accommodate 6th
r1 6 entirely
c;1 not visible,
5th sternite
rounded apically
127
EXTERSAL SEX DIFFERENCES IN STORED-PRODUCTS C()LEOPTERA.
TABLE I-cont.
Family and species
Character and its sexual expression
Fig.
Reference
CRVPTOl'HAGlDAE
Cryptophagus specie8
Hind tarsi:
_____________ I_
J_4_-_S_e_gm_e_n_t_e_d_ _ _1
~ 5-~egmented
-
Coombs &
\V oodroffe,
1955
TENEBRIONIDAE
Tenebrio ObSCUI"U8 F.
~
T. moliWr L.
17
Pupal genital papillae :
¥ divergent and con-
slightly divergent
at apices, small
and inconspicuous
spicuous, flattened,
somewhat
chitinised, not papillate
Fowler (1891) separates the sexes on the
degree of curvature of the front tibia, but
the author found this too variable to be
useful
17
Pupal genital papillae :
J
Pribolium con/Mum
Duv.
<s
paraIlfll, fused at
base, small and
inconspicuous
'i' identical with
(il All elytral striae
abbreviated before
apex
r;:' (i) Elytral striae 4
& 6 and 7 & 3
meeting Itt apices
and entire
(ii) Anterior femora.
without a setiferous puncture
those of
T.obscurus
(ii) Anterior femora
with a sub-basal
setiferous puncture
on the ventral side
T. castaneum (Hbst.)
The elytral striae uRed in the separation of
the sexes in T. conju8um can also be used
here but the difference is not so apparent
and considerable pract.ice is necessary. The
setiferous puncture is. however, more easily
seen in this species
Sub-basal setiferous puncture on anterior
femur:
J present
I
~ absent
19
Hope. 1953
Hinton, 1942
-
lHinton, 1942
(includes
figure)
128
D. G. H. HALSTEAD.
TABLE I-cont.
Family and specie8
TE)lEHRIO:>IDAE--cont.
ratzebwrgi
(Wissm.)
P. subd.epre88U8 (W011.)
PalOTUS
I
Cnaracter and its sexual expression
Pupal genital papillae :
ci
Reference
-
-
16
-
'i2 conspicuous. pro-
small, globular
and inconspicuous
Gnathoeent8 maxillosu8
(F.)
Fig.
tuberant and divergent apically
A ' tusk-like' dorsal tooth on each mandible. and head with two median prominences:
rS present
"-
, 'i! absent
I
G. t»rnutua (F.)
--- ~
A large, tapering dorsal tooth on each mandible, and head with two prominences :
~
r1 present
SiWphagu8 Iwlolepwides
(Cast.)
AlphitobiU8laevigatU8 (F.) }
Mid-tibiae:
I¥
-
-
25
Surtees, 1961
absent
I
'i? absent
Ja apical
with one spur of I 'i? with both apical
pair turned
spurs straight
inwards
J
¥ absent
A brush of hairs at the centre of the labium:
a present
A. diaperinU8 (panz.)
I
Anterior region of head thickened and excavated. Top of head with two black,
shining ridges :
a present
LatheticU8 C¥rYzae Waterh.
'i2 absent
Anterior lateral regions of head produced
into a pair of horns in front of the eyes :
<1 present
AlphitophagU8 bifasciatus
(Say)
I
-
- Hafeez, M. A.,
& Chapman.
G. (personal
communication)
18
Hewlett.
1958
EXTERNAL SEX DIFrEREXCES IN STORED·PRODUCTS COLEOPTERA.
129
19
Figs. 15-19. - 15, Gnathocer~ cornutuB (F.), head (0, 9); 16, GnathoceTus maxillosus
(F.l. head (d', 9 ); 17 . TenebTio obscurU8 F. , terminal segments of pupal abdomen.
ventral view (d' a, 9), T. molito., L . . the same (d' b); 18, Alphitobius la evi,]atlls (F.),
apical tibial spur (d', 9 ) (Hewlett, 1958 p. 144); 19, Tribolium con!lwlln Duv .• apex of
elytron (d'. «' ) (Hope, 1953 p. 265).
(L 1052)
B
(
130
D. G. H. HALSTEAD.
TABLE I-cont.
Character and its sexual expression
Family and species
CuRCULIONIDAE
Sitophilua granariU8 (L.)
<3 (i) Rostrum com-
<j?
paratively rough ;
shorter and wider
than that of the ~
(ii) Abdominal
sterrutes 5 & 6
projecting downwards
S. Q1"yzae (L.)
~Rostrum :
S. zeamaia Motsch.
J<3and
distinctly shorter
wider than
(i) Rostrum
smooth and shining, somewhat
longer and thinner
than that of the 0
(ii) Abdominal
sternites 5 & 6
not bent downwards
Fig.
Reference
20
'j? distinctly
longer
and narrower than
that of the 0 ,
smooth and shining
that of the- ~ ;
rough
....
ANTHRIBIDAE
Araecerus jaaciculatU8
(Deg.)
------
Pygidiurn :
o vertical,
not distinctly visible dorsally
BRUCHIDAE
Callo8obruchus chinett8"i8
o Antennae
(L.)
pectinate
C. rhodeBianus (Pic)
<3 Antennae strongly
serrate
C. phwleoli (Gylh.)
d' (i) Antennae
strongly serrate
(ii) Pygidium without dark
patches
(Elytra usually
brown with ~hite
scal~)
<;jl
23
Mohamed
Taber el
Sayed, 1940
21
Southgate,
1958
inclined, distinctly
visible dorsally
<i' Antennae
serrate
if' Antennae weakly
serrate
<i' (i) Antennae
weakly serrate
(ii) Pygidium with
two dark
patches, one on
each side of the
mid-line
(Elytra usually black
with patches of
brown that are
covered with white
scales forming a distinct pattern)
22
Southgate,
1958
'1.
131
EXTERNAL SEX DIFF&RENCES IN STOB-ED.PRODUCTS COLEOPTERA.
I
d
22
21
23
24
Figs. ~0-24.- 20, SitophiLus granarius (L.). posterior region of abdomen (side view), head
and. rostrum (r5. C;); 21. Callosob.,.uc}l/Us chinensis (L.), antenna (0, C;) (Southgate,
1958 p. 593): 22, Callosobruchus Thodesianus (Pic), antenna. (0. C;) (Southgate. 1958
p. 593); ~3. AraeceTus fascicutatus (Deg.). posterior region of abdomen, side view (o , C;)
().:[ohamed Taher el Sl1yed, 1940 p. 136); 24 , Ca ry edon gonagra (F.), posterior region of
abdomen. side view (0. <;? ) (Davey, 1958 p. 401).
132
D. G. H. HALSTEAD.
TABLE I-cont.
Family and apecies
Character and its sexual expresaion
Fig.
Reference
-
Southgate,
Howe &
Brett, 1957
-
Southgate,
Howe &
Brett, 1!J57
BRUCHIDAe---cont.
Oallosobruehus analis (F.)
Separation ill possible only by using
cuticular colour of the pygidium :
~
O. maculatus (F.)
th~
+pygidium with a
pygidium with a
"ide median testaceous (i.e.,
brownish-yellow)
strip or ' V •-shaped
area
narrow median
testtlceous strip
In freshly emerged specimens the
¥ usually
has a dilltinct large spot on each elytron
whilst the ~ is without dilltinct large spots
but there is a wide range of intermediate
forms
'_
~
Oaryedon gonagra (F.)
O. cassiae (Gylh.)
_______
<1 not visible from
"
24 Davey, 1958
IPygidium :
above
(gonagra)
!
C; distinctly visible
fl'omabove
I
"·-~·=:;----;;'_-----------I---------+---------I--
Bruchidius atrolineatus
(Pic)
J Antennae
S' Antennae
26
serrate
flabellate
!
AoonthOilcelides obtectus
(Say)
Zabrotea .mbJasciatua
(Bah.)
Pygidium:
<3 vertical, only partly
seen from above
(as in fig. 23<1)
¥ oblique,
3 Elytra "ariegated
Q General eJytral pubescence black but
with patches of
grey and fawn pubescence
in full
view from above
(as in fig. 24 '!')
with
a
distinct
small area of white
pubescence
surrounded by a verY
narrow circle of
fawn pubescence
-
oj
I
I
EXTERXAL SEX DIFFERE:s'CES
IN
STORED-PRODUCTS COLEOPTERA.
133
26
25
Figs _ :25-:26.- 25, A l philophagu8 bijascinf us (S a~-) , dorsal view of hea u (3, 9 ) (Surtees,
1961 p . 112); 26, Br.u chidius alrolineatu.s (Pic), antenna (3 , 9).
Summary .
The known external characters . and some new ones, for separating the sexes
of pupae and adults of stored -prod ucts Coleoptera a re described and in many cases
fig ured .
Acknowledgements.
Thanks are due to the follow ing members of t h e Pest Infestation Lab o rator~:
Mr. G. E. vVoodroffe and Dr. R. W . Howe for ad vice and ::\Ir. J. Hammond for
photographic assistance.
References.
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in the United States ,yith description of a new species (Coleoptera :
Dermestidae) .-Ann. ent. S oc. A 'me r. 49 pp. 559-566.
COOMBS . C. W. & \\"OODROFFE. G. E. (1955). A revision of the Briti"h species of
L'ryptophaUl/s (Herbst) (Coleoptera: Cryptophagidae) .-Trans . R. ('nt.
Soc. Land . 106 pp. 237-282.
DAVEY , P. yf. (1958). The groundnut bruchid Carycdol1 gO ll agra (F.) .-Bllll.
cnt. Res. 49 pp. 385-104.
DORso::-;- , R. ::\1. (1954) . The species of Carpoph ilus Stephens (Col. , Nitidulidae)
associated with stored products .-Bldl. en.t. R es . 45 pp. 389-402.
FOWLER. W . W. (1891). TEe Colenptera of the Britis h I slands . Vol. V.L ondon, Reeve .
HEWLETT, P. S . (1958) . Secondary sexual characters in Alphitobi1LS la ev igattls
(F.) and .-1. diap e-rinll8 (Panz .) (Col., Tenebrionidae).-Ent. mall . -'l a.g .
94 p. 144.
H rxTox , H. E . (194 1r. Th e Ptinidae of ecollom ic im portance.-Bull. ent. Res .
31 pp . 331-381.
Hr:s'TOx . H . E. (1942).
Secondary sexual characters of T riboli·um.-Natllrc.
LOlld. 1019 pp . 500-501.
134
HI~TO~, H.
D. O. H. HALSTEAD.
E. (1945). .\ monograph of the beetles associated with stored pro·
ducts. Vol. 1.-443 pp. London. Brit. ~lus. (Nat. Hist.).
HOPE, J. A. (1953).
A simple method for sexing the confused flour beetle.Nature, Lond. 171 pp. 265-206.
LEFKOVITCH, L. P. (1959). A revision of the European Laemophlocinae (Cole·
optera: Cucujidae).-Tran8. R. cut. Soc. Land. 111 pp. 95-118.
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