CHROMOSOME NUMBERS OF COLEOPTERA *
STANLEY G. SMITH
Forest Insect Laboratory, Sault Ste. Marie, Ontario
Received 7.vi.52
THE present is a complete list of the chromosome numbers of
Coleoptera. That in Makino's (zi) An Atlas of the Chromosome
.Jsfumbers in Animals is replete with errors : not in the numbers themselves
—almost without exception, these have been faithfully recorded from
the originals—but in the spelling of the specific, generic, and family
names, as well as in the nomenclature and systematic disposition of
many of the species. j' Considering the difficulties with which Makino
was faced, the coverage of the literature is good, perhaps no more
than 14 determinations by nine authors being omitted.
I had earlier intended to compile a list of some i 6o chromosome
determinations, which I have made as time permitted during the
course of the past eight years, together with others that have appeared
in print, and to publish them as a supplement bringing those given
by Harvey (igi6 and 1920) up to date, but it now appears that a
complete revision of Makino's list is warranted. It is perhaps noteworthy that i r families find themselves represented for the first time
and that the total of cytologically known Coleoptera now reaches 340.
According to Makino's tabulation, this number is surpassed among
Insecta only in the Orthoptera and Heteroptera but in no other
animal group, and constitutes approximately i o per cent, of the
species of animals whose chromosome numbers have been recorded.
The distribution of species, genera, subfamilies, and families by
haploid number of autosomes and by sex-chromosome type is given
graphically in fig. .
In the accompanying tabulation it has been considered advisable
to arrange the species not in alphabetical sequence under families
(a procedure followed by Makino and by Harvey) but so far as is
possible in phylogenetic sequence, since it is only through a logical
grouping based on current concepts of taxonomic relationships that
the full import of the cytological findings becomes evident. The
systematic arrangement adopted herein is that devised by Leng (1920)
on the basis of adult morphology, and his catalogue is also taken as
* Contribution No. 48, Division of Forest Biology, Science Service, Department of
Agriculture, Ottawa, Canada.
t Thirty-three spelling mistakes occur among the total of s 8o determinations tabulated;
45 species are retained in genera from which they are now, or were even at the time of
first publication, removed; and a few, for example Silpha perforata, are listed under the
incorrect family. The majority of these errors and all the obsolete names have been carried
over from the original authors. It is regrettable that cytologists in general have been,
and many still are, remarkably lax in their dealing with taxonomic nomenclature. But
unfortunately many purely typographical errors have arisen de novo during the printing
of both the original Tokyo and the first American editions,
3S
S.
32
G. SMITH
Fio. i.
Coleopteran Sex Chromosomes
XO
Xy
Xy
Xy
Fio. 2.
Neo-XY
XY
CHROMOSOME NUMBERS OF COLEOPTERA
33
the authority for questions of nomenclature. Whereas Leng and his
co-workers were concerned only with Coleoptera of America, north
of Mexico, many of the species listed here occur only outside North
America; the disposition and nomenclature of such exotic species
are those given in the Coleopterorum Catalogus.
With but few exceptions all taxonomic determinations of my
material were made by W. J. Brown, Systematic Entomology, Science
Service; it is a distinct pleasure to record my debt of gratitude to
him for these and for guidance in questions of synonymy.
GENERAL NOTES CONCERNING THE LIST
Papers preceding Stevens (1905) are largely in error or open to
doubt; they have therefore, with the exception of a determination
by Prowazek (see footnote 4'), been omitted but are fully listed by
Harvey (igi6) and Makino (1951). It has not been possible to secure
a few of the papers cited bj Makino; references to these are placed
in parentheses. Unless otherwise stated, undetermined species I
have examined originated from the Lake Kipewa region in the
Province of Quebec. Components marked with an asterisk are
considered as parts of new families by Boving and Craighead (ii).
The first number column gives the somatic chromosomes according
to sex, the second gives the number of autosomal pairs and the sex
chromosomes.
The following symbols are used : "XY ", indistinguishable or
unidentified sex chromosomes; x and y, relatively small sex chromo-
somes; p, r, and c, parachute-, rod-, and centric-associations
neo-XY, derivative, or complex, sex-determining
respectively ;
mechanism (see text fig. 2) ; s, supernumerary chromosome ; Parth.,
Parthenogenetic.
The list has been compiled up to April 1952.
LIST OF CHROMOSOME NUMBERS
I ADEPHAGA': i. CARABOIDEA
i CICINDELIDE'
CICINDELINI
.
Ckindda purpurea
.
.
Dej.
.
repanda
,,
tranqucbarzca
(
20 (3'
.
sexguttata
.
.
.
s. sexguttata Fab.
.
punctulata
scut€llaris lecontei Hald. .
io+Xx
Goldsmith "9
9+XYc
Smith '50
i o+Xx : XXxx
22
-
3
pimcriana4 .
.
22
22 3', 24
.
aneocisconensis
.
(3'
(3
24
2o (3
22 3'
22 (3
22 (3
ca. 20 (3
io--Xy3
io+Xx XXxx
9+Xyc
,o+Xx
...
,o+Xx
9+Xyc
Stevens '09
Goldsmith 'i9
Stevens 'o6
Goldsmith '19
Smith unpub.
Goldsmith 'r9
Smith unpub.
S. G. SMITH
34
ii CARABIDiE
CARABINAE
Carabus serratus Say
I2---Xy
Smith unpub.
HARPALIN1-BISETOS
BEMBIDIINI
Bembidion carinula Chaud.
bifossulatum Lec.5
...
•
PTEROSTICHINI
Euferonia coracina Newn.
Pacilus lucublandus Say
•
...
•
...
II
II
,,
i8+X
,,
'7 +neoXY
AMARINI
Celia erraiica Sturm
15?+neoXY
'7 +neoXY
Amara impuncticollis Say
AG0NINI
Sericoda obsoleta Say
Agonum sinuatum Dej.6
extensicolle Say
cupripenne Say 6
retractum Lec.6
•
•..
•
...
...
...
Platj'nussp.7
•
DRIPTINI
Galerita bicolor
LEBIINI
Dromius piceus Dej.
•
•
37cr
• 3o
• 25
17
12
+neoXY
+neoXY
,,
'7 +neoXY
Smith '50
i8+X
Smith unpub.
(Yosida '5,)
Xy
12+X
Stevens 'o6
Smith unpub.
HARPALINk-UNISETOSE
CHLENIINI
Chhsnius pennsylvanicus 8
tricolor Dej.
+Xy8
18+X
r8 +X
i6 +Xy
i8 +X
i8 +X
37
pallipes
estivus
latisollis Say
Anomoglossuse marginatus
HARPALINI
'5 +neoXY
(P2 +X)?
Harpalus erythropus Dej.
Agonoderus lecontei Chaud.
ANTHuNI
35
Anthia sexguttata
17 +X
Stevens 'o6
Smith unpub.
(Yosida '51)
Stevens 'o6
Smith unpub.
Stevens 'o6
Smith unpub.
,,
Asana et al. '42
iii DTTISCID4i
COLYMBETINIE
20+" XY"
Agabus confinis Gyll.
Colymbetesfuscus
35-37
Smith unpub.
GUnthert '10
DYTISCIN
Henderson '07
Schafer '07
Debaisieux '09
Schafer '07
Günthert '10
(36-41
- 38
Dytiscus marginal is L.
'40
circunwinctus
38
sp•
38-40
2. GYRINOIDEA
GrRLWDIE
Dineutes americanus Say
.
horni Rbts. .
Ca. 44 c3'
•
...
Ca. 2I
Ca.
2Ii
Smith unpub.
CHROMOSOME NUMBERS OF COLEOPTERA
35
II POLYPHAGA: i. HYDROPHILOIDEA
HTDROPHILID1
Hydrous acuminalus
piceus
30,3'
I4+Xy
30<3'
triangularis Say
Tropisiernu.s lat€ralis Fab.
30(3'
'5ii
14+Xyp
Asana ci al. '42
Arnold 'o8
Smith unpub.
8+Xyp
2. SILPHOIDEA
SILPHID
6+X
6+X
'3
.Jsrecrophoru.s sayi
'3 0
vespilloides Hbst.
Silpha surinamensis Fab.
noveboracensis Forst.
americana
Ca. 12+XYp
26
I 2+Xy,,
I 9+XY
40
ig+X
perforata'0
Stevens '09
Smith '50
Smith unpub.
,,
Stevens 'o6
(Yosida '5')
3. STAPHYLINOIDEA
STAPHYLIXIDIE
STAPHYLININiE
XANTHOLININI
13+XYp
Nudobius cephalus Say
Smith '50
STAPHYLININI
21+Xy
ca. 19+Xyr
Staphylinus violaceus
Philonthus polities L.
Oniholestes cingulatus"
26
Gray.
,,
Creophilus maxillosus villosus
12 +Xy
,,
12+Xyp
ca. 17+Xy
Stevens '09
Smith unpub.
Stevens '09
Smith unpub.
Gray.
ALEOCHARINIE
8+neoXY
Aleochara sp.
Unknown"
"Rove beetle"
.
.
28 3'
Stevens '09
4. CANTHAROIDEA
i LAMP TRID £"
Lucmo'rINx
Lucidota calfornica Mots.
•
corrusca
,,
9+x
L.
•
'9cr
I93',20 g+X
9+x
sp., nr. corrusca L.
Smith and
Maxwell '53
Stevens 'o6, '09
Smith and
Maxwell '53
PH0TININI
Pyractomena
angulata Say
.
borealis Rand.
Photinus consanguineus
.
.
.
sp.
PH0nIRINI
Photuris pennsylvanica 16
pennsylvanka complex
.
.
•
.
•..
19+x
,,
.•.
19 <3', 20
.••
9+X
I93',20 9+X
9+x
Stevens '09
Smith and
Maxwell '53
Stevens '09
Smith and
Maxwell '53
S. G. SMITH
36
ii CLERID
Thanasimu.s dubius Fab.
Clerus nigripes rufiventris Spin. 18
Trithodes nutalli Kby.
8+Xyp
8±Xy
8+Xyp
Smith '50
Smith unpub.
,,
,. LYMEXYLOIDEA
MICROMALTHIDrE'7
Mkromaltlzus debilis Lec.
io
.
,
20
l0
Scott '36
6. MORDELLOIDEA
i CEPHALOID11
8+xyp
Cepizaloon lepturides Newn.
Smith unpub.
ii MELOID18
MELOINIE
Mylabris pustulata 1G
22
i o+Xy.
9+XYr
2O
9+Xy
g+Xy
9+Xyp
20
Meke sp.
LYTTINE
Epicauta cinerea
pennsylvanica
murina Lec.
2O
Asana et at. '42
,,
Stevens
'09
Smith unpub.
7. ELATEROIDEA
i ELATERITh
PYROPHORINE
PYROPHORINI
4 delocera rectangularis Say
LEPTUROIDINI
Limonius griseus 20
esger Lec.
Ctenicera odireipennis Lec.2'
hieroglyphka Say
propola propola Lec.
i7
8-j-X
Smith unpub.
I7
20
8+X
9+xyp
Stevens '09 20
io+Xyp
xo+X
2I
io+X
appressa Rand.
Io+xyp
mediana Germ.
io+Xyp
I9
I9
splendens Zieg.
ezripennic .sripennis Kby.
a. destructor Brown
appropinquans Rand.
semimetaliwa Wik.
nitidula Lec.
rufopleuralis Fall
arata Lee.
infiata Say
Eanus maculipennis Lec.
estriatus Lec.
Smith '5°
Smith unpub.
193'
193'
•
...
•
...
9+x
9+x
9+x
9+X
9+x
,,
8+X
,,
8+X
8+X
9+XYp
9+Xyp
9+xyp
CHROMOSOME NUMBERS OF COLEOPTERA
37
ELATERIN1E
AGIU0nNI
9+x
Agriotella bigeminata Rand.
Smith unpub.
ELATERINI
Ampedus apicatus Say
meisheimeri Leng
I93'
9+x
9+X
deletus Lee. (?)
sp., nr. deletus Lee..
,,
,,
,,
I o-f-X
9+x
9+x
9+x
sp., nr. rniniipcnnis Lee.
fusculus Lee.
luctuosus Lee.
" Elater
sp. I "
" Elater sp. II "
9+X
,'
Stevens 'o6
,,
MELANOTINI
Melanotus castanipes Payk.
leonardi Lee.
sp., nr. communis Gyll.
sp.
trapezoideus Lee. (?)
I93'
19c3'
9+X
9+x
9+x
Smith unpub.
,,
',
,'
,,
CARDIOPHORINE
Cardiophorusfenestratus Lee.
22 3'
gagates Er.
togatus Horn
convexulus Lee.
I OXyp
Io+xyp
1o+xyp
Io+xyp
,,
,,
ii BUPRESTID
JULODINI
Julodis whithilli
Sternocera nitidicollis 23
l.evigata
24(3'
26(3'
II +neoXY 22
Asana et al. '42
12+fleOXY 22
263'
12+neoXY 22
2! , 22
io+X
,'
CHALCOPHORINI
Chalcop/wra lacustris Lee.
Euchroina gigantea
12+Xy
Smith unpub.
Nichols 'xo
BUPRE5TINI
Dicerca prolongata Lee.
tenebrosa Kby.
Buprestisfasciata Fab..
Melanophila drummondi Kby.
20
20
20
9+XYp
Smith unpub.
i63'
Xy
5±Xyp
,,
i6 (3'
i+Xy
,,
acuminata DeG.
9+xyp
CHRYSOBOTHRINI
Ch7ysobothris dentipes Germ. .
AGRILINI
Agrilus anxius Gory
22 3', 22
liragus Barter and Brown 203', 20
sp., nr. pensus Horn
politus pseudocoryli Fish.
.
sp.
Io+Xyp: XX
9+neoXY : XX
Xy
9+" XY"
Smith '49
Smith unpub.
9+neoXY
Unknown
"Spruce borer sp. I"
"Spruce borer sp. II"
9±Xy
Io+Xy
Stevens 'o6
S. G. SMITH
38
8. BYRRHOIDEA
i DERMESTID
DERMESTINI
Dermestes talpinus Mann.
lardarius L.
signatu.s Lec.
8+Xyp
8+Xyp
8+Xy
ANTHRENINI
8+Xyp
Anthrcnus sp.
Smith unpub.
,,
,,
Smith '50
ii BTRRHID
i8
Cytilus alternatus Say
Xy
Smith unpub.
9. CUCUJOIDEA
i NITIDULIDE
9+xyp
9+xyp
ii CUCUJID
Nitidula bipunctata L.
rufipes L.
Oryz&philus surinamensis
13-14
7 elements"
Smith '50
Smith unpub.
McMullen '28
iii EROTflIDf
8+neoXY
Triplax thoracica Say
Smith unpub.
iv COCCLVELLIDiE
COCCINELLINiE
HYPERASPINI
Brachyacantha ursina Fab.
SYNONYCHINI
20
Synonyc/za grandis Thunb.
20
Cxlophora i?uequalis24
1'7
Aiolocaria mirabilis 25
7+neoXY
Smith unpub.
9+XYr
9±Xyr
Yosida '44, '46
(Yosida '49, '5!)
(T. and Y. '37)
8+X
120
(Yosida '49, '51)
COCCINELLINI
lengi 20
Coleomegilla maculata
Timb.
9+XYp
Smith unpub.
9+Xyp
Yosida '44
Smith '50
Smith unpub.
Hippodamia tredecimpunctata L. 20
.
t. tibialis Say.
20
.
Coccinella trfasciata L.
transversoguttata Fald.
bruchii 28
I\4uIs.26
.
crotchi Lewis .
.
axiridis Pallas27
.
Adalia bipunctata
i8
.
.
20
.
i6
.
20
L.
,,
,,
9+xyp
9+xyp
Cleis hudsonica Csy.
.
{
2 d'
Anisocalvia duodecim-maculata
Gebi.
quatuordecimguttata L.
,,
,,
Propyl&zjaponica
Anatis mali auct.
.
20
T 28
i8
8+Xyr
9±Xyr
I 7+XYp
1 7+XYr
9+XY
9+xyp
5+" XY"
5+"XY"+2S
9+xyp
(T. and Y. '37)
Yosida, '44
,,
Li '40
Yosida, 4
Stevens 'o6
Smith unpub.
,,
9+xyp
9+XYr
Yosida'44
8+neoXY
Smith unpub.
9+Xy
(Yosida '49, '51)
CHROMOSOME NUIfBERS OF COLEOPTERA
39
EPILACHNIN
5 i8
Epilachna borealis
1,8 , i8
20
nipponka Lewis '
vigintioctomaculaa
Mots.3°
20 d'
18
viginhioctopunctata Fab.
8+Xy
Stevens 'o6
9+XYr
Yosida '44
(T. and Y. '37)
Yosida '48
8+"XY"
Xy
Xy
20
pustulosa Kôno 32
chrysomelina F. x capensis
Various undetermined species 20
,,
Xy
i8
Thunb.
Unknown
Hoy 'i8
Strasburger '36
9+Xy
Stevens '09
, o. TENEBRIONOIDEA
i ALLECULID1E
Isomira quadri.striala Couper .
20
9+XYp
Smith '50. '52b
ii TEJfEBRIOJUD2E
ER0DI1NE
Erodius nitidisollis Sol.
edmondi Sol.
Leptonychus curvicornis Peyer..
20
20
20
9+Xyr
Guénin '5ib
9+XYr
9+Xyr
TENTYRIIN)E
20
20
Mesostema angustata prasa- 20 d'
9+XYr
9+XYr
9+XYr
Guénin '50
Guénin '5ia
Genus nr. Hylocrinus Csy. 20
9+XYp
Smith '52b
7+Xy
Smith '52b, '53
Tentyria mucronata Stev.
subcosta Sol.
hariana Koch.
(Texas)
,6
ZoPherus haldemani Salle
ELENOPHORIN1E
Elenophorus collaris L. .
AKIDINIE
Akis bacarozo Schrank
26 d'
.
i6
7+neOXY
i8
8+Xy
PIMELIIN
Pimelia bipunctata Fab.
i8 3'
serviL,i Sol.
aranacea Sol.
grandis la/es/i Sen.
angulata lestei Peyer
sub quadrata valdani Guer.
Ocnera hispida Forsk
Prionot/jeca corona/a 01.
BLAPTIN)E
Blaps waltli Seidl.34
,,
GuCnin '5,a
,,
8+Xyr
8+Xyr
8--Xy5
8±Xyr
•
gigas L.
mucronata Latr.
mortisaga L.
lath jfera Marsh.
•
Opatrinus aciculatus Lec.
Guénin '50
8+XYr
•
DIAPERINJE
Diaperis bole/i L.
8+XYr
8+Xyr
lusitanica Hbst.
PEDIN1N
I 2Xy
.
•
•
34cr
135 '
lI9,20
35,38
36c,38
37,38
'5+3 XY
'5+4 XY
8+2 XY : 4 X
'5+4 XY : 8 X
16+3 XY :6 X '
'6+3 XY :6 X
17-1-2 XY :4 X
Nonidez '25
Nonidez '24, '20
Guénin '49
,,
20
9+Xyp
Smith '52b
'4cr
6+neoXY
GuCnin '50
S. G. SMITH
40
ULOMIN
.
20
.
.
x8
Alobates pennsylvanica DeG. .
.
Scolobates calceratus Fab.
.
.
Upis ceramboides L.
.
Tenebrio obscurus Fab.
20
Tribolium castanewn Hbst.
.
confusum Duval
destructor Uyttenb. .
,,
,
z8
TENEBRIONIN
molitor .
,,
L.
.
.
i8
i8
...
20 d'
20
,
20
.
.
20
20
Smith '52a, b
g+Xyp
Smith '52b
9+Xyp
g+Xyr
9+Xy
Guénin '50
8+neoXY : XX
8+neoXY : XX
8+neoXY
(9+XYr
. 20 d'
.
g+Xyp XX
g+XYp
L
pkipes Hbst. .
.
20
.
9+XYp
Stevens o536
Guénin '50
Smith '50, '51
'52b
Smith '52b
iii LAGRIID
Arthromacra €znea Say .
6+neoXY
.
Smith '50, '52b
iv MELAXDRrID1f
* PENTHINI
.
.
Penthe obliquala .
Fab. .
,,
,,
...
.
7+neoXY
Stevens '09
Smith '52b
9+Xyp
9+Xyp
Smith '50, '52b
Smith '52b
7+Xy
MELANDRYTNI
20
Serropalpus substriatus Hald. .
Dircea quadrimaculata Say .
...
ii. SCARABAOIDEA
i SCARABID
COPRIN
Pharusus vindex MacL.37
.
igneus MacL.
APHODIIN
APH0DITNI
.
2
. I2
.
Aphodiusfimelarius L. .
...
5+" XY"
5+" XY"
Hayden '
9+Xyp
Smith unpub.
EUPARUNI
.
At&nius spretulus Hald.
Ca. 20
*GEOTRUPIN
Xy
.
.
.
.
Diploto.xLs sp. (Texas)
Phyllophaga gracilis 38
.
fusca 38
.
.
delata
.
.
.
20
20
20
,,
sp., crcnulata group.
.
ca.
20
.
.
20
Geotrupes balyiJek.
MELOLONTHIN1E
SERICINI
Serica tristis Lec.
20
MELOLONTHINI
trISttS
20
RUTELINE
,
,
20
20
20
g±Xyp
Smith '50
9+Xyp
Smith unpub.
Shaffer '20
9+Xy
g±Xy
g+Xy
...
20
Smith unpub.
9+Xy
Shaffer '20
8-j-Xy
Yosida '49b
ANOMALINI
Popilliajaponica Newn.38
Anomala corpulenta Mots.
.
rufocuprea Mots.
.
.
. i8
8+Xy
8+Xy
CHROMOSOME NUMBERS OF COLEOPTERA
RurauNi
,
20 , 20
Pelidnota punctata4°
Catalpa lanigera .
DYNASTINIE
20
Oryctes nasicornis L.41
.
12
.
20
20
d
41
9+Xy
9+Xy
Shaffer '20
5+" XY"
Prowazek '02
9+Xy
9+Xyp
9+Xy
Stevens '06
Smith unpub.
Yosida '49b
9+Xyp
g+Xy
Yosida '49b
CETONIINE
CETONIINL
.
Euphoria inda
,, L.
,,
..•
•
Glycyphana fulvistemma Mots. 20 d
TRICHIINI
...
Trichiotinus assimilis Kby.
.
Tridiiusfasciatus L.
20 d'
.
Smith unpub.
ii LUCAXID
9+X
Psalidoremus inclinatus4t
(T. and Y. '37)
iii PASSALIDI
,,
26
-
Popilius disjunctus
Ill.
,,
.
26
,
26
12+Xy
12+neoXY
Shaffer '17
Smith unpub.
12. CERAMBYCOIDEA
i CERAMBrCID
CERAMBYCINE
ELAPHInIONINI
9+Xyp
Elaphidion, prob. parallel us
Newn.
Smith unpub.
LEPTURINI
io+Xyp
Acrnaops proteus Kby.
CLYTINI
Cyllene robinia"
LAMHNAi
•..
g+Xy
Stevens '09
•..
9+Xyp
g+Xyp
g+Xyp
Smith '50
Smith unpub.
9+Xy
(Yosida '51)
g+Xyp
Smith unpub.
M0N0cHAMINI
Monoc/jamus notatus Drury
scutellatus Say
marmorator Kby.
.
Pterolophia caudata
.
...
...
20
GRAPHISUR.INI
...
Graphisurue obsoletus OIiv.
PHYTOtCITNI
Oberea trip unctata H
...
...
Swed.
,,
,,
mandarina Fab.
affinis Leng
Xy
9+Xyp
9+Xyp
9±Xyp
Stevens '09
Smith unpub.
TETRAOPINI
Tetraopes tetrophthalmus 46
,,
fernoratus
Forst.
.
.
20
20 d'
.
.
.
20
...
g+Xy
9+XY
9+Xyp
g+XY
Stevens '09
Snyder '34
Smith unpub.
Snyder '34
S. G. SMITH
42
ii CHRTSOMELID
*DONACIIN
•..
...
'3+Xyp
i3+Xyp
Smith unpub.
.
...
i5+Xy
i5+Xyp
Stevens 'og
Smith unpub.
Cryptocephalus venustus Fab. .
Ca. 22 <3'
Donacia hirticollis Kby.
.
subtilis Kunze
.
*CRIOCERIN
Leina trilineata .
auct.
,,
,,
.
.
32
CRYPTOCEPHALIN
*EUMOLPIN
.
.
Fab.
.
...
...
.
33-34<3', 36
.
.
36
Chrysochus auratus
*CHRYSOMELIN
Ca.
i 2+Xy.
Stevens '09
i 6--Xy
z 6+XX : 4 X
1 7+Xy
17+X : XX
Stevens '09
I2Xy
Smith unpub.
CHRYSOMELINI
Labidomera clivicollis 48
Leptinotarsa signaticollis
decemlineata °
.
Stãl.
,,
.
.
.
Say
.
Calligrap/za similis51 .
.
.
philadeiphica L.
multipunctata bigsbyana Kby.
Chrysolina exanthematica
<3'
35 3', 36
...
...
...
17+X
...
i z+Xyp
23 3'
Wied.5'
Chrysomela interrupta auct. in 34 3'
part
sp. .
.
.
.
. 34?
...
...
...
lapponica 52
tremu1e auct. .
Mclasoma snca H
populi 53
.
.
.
.
.
32
Phytodecta americana Schffr. .
.
.
.
variabilis
24
<3'
ii +X
i
Wieman '10
Stevens 'o6
Guénin and
Scherler '52
Smith unpub.
Stevens '09
Smith unpub.
ii
Yosida '4
i fiH-Xyp
Smith unpub.
16+Xyp
i 6+Xy
Stevens '09
i
i6+Xy
Smith unpub.
(Yosida '51)
1 5+XYr
PHYLLODECTINI
*GALERUCIN
ArysINi
Trirhabda virgata
.
,,
Kby.
.
i
,
28 28
30 <3', 30
30 3'
.
.
canadensis
...
Galerucella vaccinii Fall
cavicollis Lee.
nymp/useL.
.
.
Xy
24 <3'
.
OIDINI
Rhaphidopalpafemoralis 54
.
.
...
...
.
...
DIABROTICINI
Diabrotica duodecimpunctata H
.
soror5 .
.
.
Acalymma vlttata
.
.
.
.
i4+Xy
i 6+Xyp
Smith unpub.
,,
28+Xy
(Yosida '49, '51)
i6+Xyp
g+X+o-.-4s
9+X+o —4s
!2I 3', 22
...
io+XX
+neoXY
32 <3', 32
I5+neoXY
24 <3'
1 i+Xy.
AGELASTICINI
Agelastica caculea Mots.59
Stevens 'o6
19-21
(21
Smith unpub.
Galán '31
I3+Xy
19, 20
LTJPERINI
Luperus discrepans Baly
Luperodcspreusius Mots.68
i+Xy
Stevens 'o8
Hoy 14, i8
Yoida '4
,,
'49a
,,
'44
CHROMOSOME NUMBERS OF COLEOPTERA
43
HALTICINIE
BLEPHARIDINI
.•
Blepharida rhoLs .
HALTICINI
Hallica subplicata Lec.
.
chalybea
Argopus punclipennis 61
23
.
22
...
*HISPIN
...
.
.
i6
Chelymorpha cassidea 63
.
22
.
Deloyala guttata"
Plagiometriona clavata 64
Agroiconota aurichalcea 64
.
Anoplitis ineequalis auct.6'
62
Chalepus dorsalis
22
i8
.
.
Stevens 'o6
ii +X
Smith unpub.
Stevens '09
(Yosida '49, '5')
io+X+y 66
24--Xy
B+Xyp
7+Xyp
,
*CASSIDIN
i5+Xyp
i0+Xyp
8+Xy
Xy
io±Xy
22
Smith '50
Stevens 'o6
Nowlin 'o6
Stevens '09
Nowlin 'o6
iii BRUCHID1
AMBLYCERINI
Zabrotes subfasciatus Boh.
26
.
,
26,
12+X+y : XX+s Minouchi '35
27
BRUCHINA
59 d', 20
Callosobruclius maculatus 65
Acanthoscelides obtectus Say 66 20
9+X
9+" XY"
Braucr '28
Bushnell '36
I 3. CURCULIONOIDEA
i PLATTSTOMIDIE
Euparius marmoreus Oliv.
...
.
io+Xyp
Smith '52C
ii CURCULIONID
OTIORHYNCHINE
POLYDRUSINI
Polydrusus pilosus Gredl.67
.
...
io+Xy
Suornalainen
...
io+XX
Suomalainen
'40a
Suornalainen
undatus F.66 .
.
.
mollis Strom. .
.
.
22
.
33
22, Parth.
SCIAPHILINI
Sciaphilus asperatus Bonsd.
Strop/zosomus capitatus DeG.69 Ca. 20 ', 22
.
34
io+X : XX
3I-35 Parth.
BARYNOTINI
Barynotus rnarens F.
.
obscurus F.
.
.
...
.
.
...
BRACHYRHININI
Brachyrhinus geniculatus
Germ7°
infiatusGyll. .
.
i. salebrosus Boh.
.
arcticus O.F. .
.
22
'oa, b
Parth.
THYLACITINI
melanogrammus FOrst.
'oa, b
551
Parth.
Parth.
Suomalainen '
Suomalainen
'oa, b
...
i o±Xy : XX
Suomalainen '47
...
...
io+Xy
io+Xy
Mikuiska 'g
io+Xy : XX
Suornalainen
'4O, '45
S. G. SMITH
armadillo Rossi
.
.
.
.
•
•
bcukatus F. .
•
. 11
fuscipes Ohv.
...
(22
...
•
...
••
morzo F.
multipunctatus F.72 .
.
•..
.
.
.
...
dubius Strom.73
.
•
...
repkius
.
.
...
niger F.
Boh. .
xo+Xy : XX
io+Xy : XX
xo-f-Xy
io+Xy
I i o+Xyp : XX
1 Io+Xy
xo+Xy
(zo+Xy : XX
- i o+Xy
-33r Parth.
Parth.
xo+Xy
.
.
.
•
...
...
...
.
...
austriacus F. .
.
.
corvus Boh.
Parth.
.
sensitivus Scop.
.
equestru Richt.
gemmatus Scop.
.
.
...
Icollari Gyll. .
.
.
obsidianus Boh.
obtusus Boh.74
pupillatus Gyll.
.
.
.
.
.
...
...
saiwis Strom..
.
.
singularis L.
.
.
•
sulcatus F.
•
.
.
liguslici L.
.
.
•
ovatus L.
.
•..
.
...
Ca.
chrpops Hbst.
TRACHYPHLEINI
Trachjphkzus bifoveolaluc Beck.
i o+XX
o+Xy
io+Xy
I
Ui o+Xy : XX
!-33, Parth.
i o+Xy
Parth.
...
...
33 Parth.
(34 Parth.
...
Suomalainen
Suoma1inen
'oa, b,
Suomalainen '7
Mikulska '49
I Suomalainen '7
1.Mikulska '4o
Suomalainen '47
Mikuiska '49
io--Xy
...
...
Mikuiska '49
,,
Suomalainen '47
Mikuiska
Suomalainen '47
Suomalainen
'4oa, b,
xo+Xy
Suomalainen '7
32 Parth.
33 Parth.
33-35, Parth.
30-34, Parth.
i o+XX
32-33,
Parth.
Seiler ,'
47
Suomalainen
'4oa, b,
Suomalainen '47
Suomalainen
'oa, b
EPISOMINI
Episomus turritus"
: XX
(io+Xy :XX
30
33
i o+Xy,
Mikulska 'g
Suomalainen 'j
Mikuiska 'z
331 Parth.
scaber L.
Suomalainen '7
.••
.
9,,
(T. ari.d Y. '7)
io+Xyp
Smith '50, '52C
Smith '52c
CURCULIONIN1E
SITONINI
.
.
.
.
.
.
.
.
congener D.S. and M.
.
22
.
22 <3'
Sitona lepidus Gyll.76
clindricollis Fâhr.
..•
...
HYPERIM
Hypera punctata •
Xy
HYLOBIINI
Hylobius abietis L.
i o+Xyp
...
i o+X
io+Xyp
Stevens '09
Suomalainen
'4oa, b
Smith '52c
CURCUUONINI
Curculio obtusus Blanch.
Smith unpub.
CHROMOSOME NUMBERS OF COLEOPTERA
45
ANTRONOMINI
...
Anthonomus scute1laus Gyll. .
*LENDRIN
12
Sitophilus granarius L.'7
oryzes L."
.
,,
.
24 , 24
112
II , 12
.
22
22
Smith '526
z3+Xyp
5+" XX"
Inkmann '33
Smith '52C
i z+Xyp
s+"XX"
5X .XX
Tiegs and
Murray '38
Gunson'45
io+neoXY : XX
Smith '526
SCOLYTOIDEA
SCOLTTIDIE
HYLESININ1E
Dendroctonus engelmanni Hopk.
...
Hylurgops pinfex Fitch
.••
i4+Xyp
i4+Xyp
...
i4+Xy
IPINA
Ips pini Say
.
.
.
Smith '526
Footnotes.—Quotations are from the original papers unless otherwise stated:
Boving and Craighead (1931) regard this suborder as being logically considered
most recent. "Cicindcllicbe" in Makino (1951).
"uulgaris" ; not X1X9-Y1, contra
Collected from the north shore of Lake Michigan,
Makino (1951).
"primeriana ".
although it would not be expected east of Manitoba and Nebraska.
Playnus sens Lang.
"Palatinus" in Makino (1951). 8 "Chalenius" in Makino (1951) ; probably an
incorrect determination (Smith, 1950).
Formerly placed in Hydrophilus. 1 Not in
Carabid, contra Makino (1951). 11 "Listotrophus cingulatus "
Not necessarily genus
Staphylinus (v. Makino, 1951). ' All species listed have post-reductional division of the
X-chromosome; not Cantharid,
contra Makino (5952).
16
14 "Ellychnia ".
"can-
"Photinus pennsylvanicus "
Boving and Craighead
sanguineus" (Makino, 195!).
(293!), with other authors, consider this to be in a third, and most primitive, suborder,
the Archostemata. The species listed has haploid parthenogenetic males. ' E. permogloanwa
is not given in Stevens (1909), contra Makino (195!). 19 " Mylabiris" (Makino, 1951).
20 "Limoneus" = "Ljnoneus" (Makino, 1951) ; not Stevens, 'o6, contra Makino (1951).
21 Species complexes in the genus Ctenieera separated according to Brown's arrangement
to an autosome".
as given in the supplements to Leng"(1920).
"XO; X attached
"
U " bruckil
"
" Calophola
Ithone
"S. nitidicolis "•
(Makino, 195!).
27 "Harmonia axyridis ".
"Calvia 14-gutiata L."
(Makino,
"niponica".
1951).
30 " = nipponica"
Contra Makino (1951). 32 "pustlosa ".
Species
(Makino, 1952).
has pre.metaphase stretch of autosomes and precession of sex chromosomes. 84 "walt/u"
36
Not
Stevens
of many cytologists, including
Makino (,g,). ' Contra Makino (195!).
"
'o6, contra Makino (1951). 57 " carnifex (L.) ". 38 " Lachnosierna ". " " Poppilia
40 "Pelidonata"
41
Compare
this
complement
with
(Makino, 195,).
(Makino, 1951).
syn. Passalus cornutus Fab.
those of Phan€eus spp. 42 "Paslidoremus" (Makino, 1951).
" Obera
" ylene robinia "
Sex chromosomes as identified
U "tetraophthalmus ".
18
"
by Snyder are incorrect for the former species and, therefore, possibly for the latter.
' "signatkolis" in Makino (1951). 0 "Doryphora ".
Doryphora clivicolis ".
= Chrysomela L. u "Rhaphidopolpa" in Makino
" Chrysomela " "Lina laponica ".
This, the eastern subspecies, is D. undecimpunctata lwwardi Barber. 5 This,
(1951).
the western subspecies, is D. u. undecimpunctata Mann.Formerly placed in Diabrotica.
= Luperus auct. = "Luperoides" (Makino, 1951). 29 "carulea 60 It is not improbable
that this is an XO species, the" y" not associating with the X because it is a supernumerary
chromosome. The generic name is alternatively Altica. 61 "A. orientalis " 62 Formerly
65 "Bruchus quadriplaced in Odontota Chev. 63 syn. C. argus Licht. 64 " Coptocycla
"
maculatus "
68 Not
in Curculionid, contra Makino (5951).
"
87 Suomalainen
uses the
European variant Polydross,s. 68 syn. tereticollis DeG. • syn. Strophosomus rufipes capitatus
DeG. =
Strophosmus
in Makino (1951). 70 American authors refer to the genus as
Brachyrhinus. Europeans use Otiorhynchus (= " Otiorrhynchus"). 71 A variety of clavipes
Bonsd. ' syn. irritans Hbst. syn. nodosus 0. Mull. syn. graniventris Mill. " "turitus"
76 = fiavescens Marsh. nec. F.
syn. Calandra.
S. G. SMITH
46
SUMMARY AND ABSTRACT
Approximately 340 species and subspecies of Coleoptera are listed
of which about one-half are determinations made by the author.
The number of component genera totals i the number of families
3. The commonest sex-determining mechanisms are Xy, neo-XY,
and XO as follows :—
Xy : 193 species in 112 genera in 24 families
neo-XY : 29
12
,,
,,
24
XO: 6o ,,
,,
30
9
The distribution of species, genera, subfamilies, and families by
haploid number of autosomes and by sex-chromosome type is given
graphically. Parthenogenesis occurs in two families; compound sex
chromosomes are found in a third and possibly a fourth.
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ADDENDUM
Since this list was compiled, reprints of three papers have been received:
Virkki (Ann. Zaol. Soc. Zoo!. Bot. Fenn. " Vanamo ", 14, 1-104, 1951) provides
information on 30 additional species in the Scarabnid and new determinations
for Trichius fasciatus L. (2nIg, contra Yosida herein) and Oiyctes nasicornis L.
(2fl = 18 contra Prowazek, v. footnote 42 herein) ; Guénin (Rev. suisse Zoo!., 59, 277-
282, 1952) reports that four European species of Cicindela have XXXY : XXX/XXX
sex-determining mechanisms; and Yosida (Papers Co-ord. Comm. Res. Genet., 3,
41-49, 1952) gives determinations for another carabid and another chrysomelid,
besides giving Silpha perforata Gebler as 2n=2o and 21 (contra Yosida, 1951, v.
Makino, 1951).