Taxonomic notes about the Middle American genus Cyrtolaus Bates (Coleoptera: Carabidae: Pterostichini)

10 Pages·1991·4.4 MB·

by  G E Ball

PROC. ENTOMOL. SOC. WASH.
93(2), 1991, pp. 230-239
TAXONOMIC NOTES ABOUT THE MIDDLE AMERICAN GENUS
CYRTOLAUS BATES (COLEOPTERA: CARABIDAE: PTEROSTICHINI)
George E. Ball
Department ofEntomology, University ofAlberta T6G 2E3, Edmonton, Alberta, Can-
ada
Abstract.—Basedon featuresofadultsandonchorological considerations, 2 newspecies
of Cyrtolaus {s. str.) are described from Oaxaca, Mexico: C. oaxacanus (type locality—
Sierra Zempoal, 10.4 km S. Totontepec), and C whiteheadi (type locality—Sierra Madre
de Oaxaca, 1 1.7 km E. Huautla de Jimenez). Character states and localities ofspecimens
C
notrecordedpreviouslyofthe Chiapan-Guatemalan subiridescensWhiteheadand Ball,
C. ricardo Whitehead and Ball, C.furculifer Bates, and C. grumufer Whitehead and Ball
confirm generally the published treatments ofthese taxa.
Key Words: new species, phylogeny, biogeography, Mexico, Coleoptera, Carabidae
In August, 1965, toward the beginning of be ranked as subtribes ofPterostichini, and
the 14-month period that Don Whitehead that Cyrtolausshouldgoin itsownsubtribe,
and I spent in Mexico, we found in a cloud Cyrtolaina,whichweproposed. Thelastde-
forest in central Chiapas 2 carabid adults of cision was based on both the seeming in-
rather remarkable form (heavy-bodied, but termediate position of Cyrtolaus in the
withstrikinglylongmandibles, narrowhead, Pterostichini, anditsseemingrelictstatus—
andmarkedlyvaultedelytra)thatwethought the genus seemed not to have close phylo-
represented some aberrant species of the genetic relatives.
platynine genus Colpodes {sensu latissimi). Liebherr (1986) discovered that 2 Antil-
Someyearslater, afterstudyingmany ofthe lean species, described originally as mem-
types that H. W. Bates had described in bers ofColpodes, in fact represented an un-
writingthecarabidpartoftheBiologiaCen- describedgenus, which he named Barylaus.
trali-Americana, we concluded that our Further, Liebherrdetermined thatBarylaus
specimens were members ofthe genus Cyr- and Cyrtolaus, though phylogenetically iso-
tolaus Bates (1882: 99). We described this lated, were endemic New World represen-
speciesand fourothersas new, in a revision tatives of the pterostichine subtribe Coe-
oiCyrtolaus (Whitehead and Ball 1975). In lostomina, a group that had been thought
that treatment, we combined in a single ge- to be confined to the Old World (except for
nus Ithytolus Bates and Cyrtolaus, ranking an introduced speciesofCoelostomusin the
each group as a subgenus. Antilles). Thus,thenameCyrtolainaWhite-
Because the range of elytral features of head and Ball, 1975 is ajunior synonym of
Cyrtolaus {sensu lato) seemed to bridge the Coelostomina. Also, Liebherr argued that
range ofelytral features ofthe tribes Ptero- the platynines should be ranked as a tribe,
stichini and Agonini (= Platynini), we con- andthusequivalenttothe Pterostichini, and
cluded that this genus belonged in neither not as a component ofthe latter.
tribeintherestrictedsense,thattheyshould I regard as elegant Liebherr's discovery
VOLUME 93, NUMBER 2 231
of the subtribal relationships of Cyrtolaus detailed examination suggested otherwise,
andBarylaus. However, onthebasisofcho- the specimens were regarded as taxonomi-
rological considerations, I do not accept his cally distinct, and were ranked as species.
conclusion thatBarylausis moreclosely re- For specific ranking of C. oaxacanus, new
lated to the geographically distant Afro- species, primacy was given to chorological
tropicallineageofCoelostomina thantothe features. See below for details.
more proximal Middle American Cyrto- Measurements.—Thefollowingmeasure-
mentswere madeaspartofthe descriptions
laus.
As knowledge ofrelationships of Cyrto- oftaxa and to compare with those reported
laus was increasing, so was additional ma- by Whitehead and Ball (1975: Tables 1-3).
terial accumulating ofthe species. Below, I HL = length ofhead from base ofdorsal
reportandanalyzethe newinformation thus condyle of left mandible to poste-
acquired,toseekvalidationofthestructural rior margin ofeye;
andchorologicallydiagnosticfeaturesoftaxa ML = length of left mandible, from apex
previously described, and to make known to base ofdorsal condyle;
taxa not known previously. PL = length ofpronotum along mid-line,
from anterior to posterior margin;
Material and Methods EL = length ofelytra, along suture, from
Material basal ridge to apex.
suTbhgiesnusstuCdyyrtoislabuass,ecdololnect3e6daidnultthse oMfextih-e LeSnigztehw(aSsBLe)xparensdseisdtahseStsaunmdaorfdiHzLed+BoPdLy
can states of Oaxaca and Chiapas, and in + EL. Two ratios were computed: ML/PL
and PL/EL.
southwestern Guatemala. Four-letter cod-
Not reported in detail are values for the
ens used in the text to indicate sources or
ratio Antennomere 3: width/length. Values
depositories ofstudy material are:
are between 0.26 and 0.32, thus well within
CASC Department ofEntomology, Cal- the range reported previously (0.26-0.38)
ifornia Academy ofSciences; for Cyrtolaus {s. str.), excluding C. lobipen-
UASM DepartmentofEntomology, Uni- nis. These relatively low values indicate the
versity ofAlberta; and relatively elongate antennae that are char-
USNM Department of Entomology, acteristicofthemostderivedlineageofCyr-
United States National Museum tolaus.
—
ofNatural History. Chorology. Localitiesreportedareinthe
followingmountainranges: SierraMadrede
Methods OaxacaandSierraZempoal, intheMexican
Recognition.—Specieswereidentifiedus- state ofOaxaca (see outline map ofthe re-
ing the key and illustrations in Whitehead gion in Ball and Roughley [1982: 364, Fig.
bayndexBaalmlin(1a9t7i5o)n,oafntdhiedmenatlietygewnaitsalcioan,fwihrimcehd n63o]r)t;hwaensdterMneCshaiapCaesnt,raaln,d SiinerrcaenMtraadlreandde
Chiapas, in southeastern Chiapas and
were prepared using standard techniques.
southwestern Guatemala (see outline map
Specimens the combination of features of
which did not fit the couplets and regarded ofregion in Whiteheadand Ball [1975: 606,
initially as taxonomically distinct entities Figs. 22, 23]).
were examined in detail. If such examina- Key to Species of Cyrtolaus Bates,
tion showed them to be distinctive in few Based on Characteristics of Adults
and varied features, the specimens were in- This key is modified from the one pre-
cluded as members ofthe described species sented by Whitehead and Ball (1975: 598),
towhich eachwas mostsimilar. Ifthe more andincludesallknownspeciesoiCyrtolaus.
,
232 PROCEEDINGSOFTHE ENTOMOLOGICALSOCIETY OF WASHINGTON
01. Pronolum with anterior marginal bead bate,lateralmarginsbroadlyrounded,not
complete; postero-lateral setae near pos- sinuate; anterior angles produced. Geo-
terior angles. Elytron with plica clearly graphical range—central Guatemala
. .
developed, epipleuron interrupted C. spinicauda Bales
("crossed"); interneurs impunctate. Ab- 07'. Elytron with interneurs hardly punctate
dominalsternumVIIwith2setaeinmale, todistinctlyso. Pronotum with hmdan-
4 in female Subgenus Ithytolus Bates gles various; lateral margins less round-
C. orizahae Bates ed, orsinuateposteriorly(Figs. 2, 4);an-
or. Pronotum without anterior marginal teriorangles not produced 08
bead; postero-lateral setae distinctly an- 08(07'). Geographical range—Sierra Madre de
teradposteriorangles(Figs. 2-4). Elytron Chiapas, southeastern Chiapas and
with epipleuron interrupted or not; in- southwestern Guatemala. Elytron with
terneursmoreorlesspunctate(Figs. 5A- apical declivity gradual (Fig. 7B), apical
7A).AbdominalsternumVIIwith4setae spine longeror shorter(Fig. 7\)
in male, 6 to 8 setae in female C. ricardo Whd. & Ball, p. 238.
Subgenus Cyrtolaus 02 08'. Geographical range—Mesa Jentral of
02(01'). Elytron with epipleuron interrupted by Chiapas, or mountains ofOaxaca. Ely-
plica. Labrum with 4 marginal setae tron with apical declivity various; apical
. .
C. lohipennis Bates spine short (Fig. 5A) 09
02'. Elylral epipleuron not interrupted. La- 09(08'). Pronotum with posterior angles rectan-
brum with 6 marginal setae 03 gular. Elytron with interneurs distinctly
03(02'). Elytron with microsculpture of trans- punctateondisc;slopeofelytraldeclivity
verse lines, surface faintly iridescent; in- moreabrupt.Geographicalrange—Mesa
terneursgrossly punctate; intervals cari- Central, Chiapas
nate; apex spined or not, spines shorter C. hrevispina Whd. & Ball
or longer(Figs. 6A, B) 09'. Pronotum with posterior angles acute.
C. subiridescensWhd. & Ball, p. 237. Elytron with interneurs very obscurely
03'. Elytron with microsculpture mesh pat punctate on disc; apical declivity sloped
tern isodiametric; apex spined 04 gradually (Fig. 5B) 10
04(03'). Headwith2(singlepair)or3supraorbital 10(09'). tLaurmgerrel(atSiBvLelymoshroerttehra(nM1L1/.0PLmm0).6;6-p0r.o7n4o)-.
setigerous punctures 05
Male with apex ofmedian lobe pointed
04'. Head with 4 (2 pairs) supraorbital setig-
erous punctures 07 (Fig. IA). Geographical range—Sierra
Zempoal (vicinity of Volcan Zempoal-
05(04). Elytron with interval 2apically moreor
tepetl, Oaxaca)
lBe)s.sGfleato,gnroatphdiicstailnctrlayngsewo—llHeunau(tFiigas.P5lAa,- C. oaxacanus, new species, p. 232.
teau, Oaxaca 10'. Smaller(SBLlessthan 10.0mm);prono-
C. whitehcadi, new species, p. 236. tGeuomgrraeplhaitciavlelyralnogneg—erSi(erMrLa/MPaLdr0e.60d).e
05'. Elytron with interval 2 swollen apically.
Geographical range—Mesa Central and Oaxaca(near Valle Nacional, Oaxaca) .
Sierra Madre de Chiapas, Chiapas, and C. newioni Whd. & Ball
Guatemala 06
06(05'). Elytron with interval 3 markedly raised
apically. Disc ofpronotum with micro- New Taxa, New Locality Records, and
sculpture normal, surface without seri- Variation in Diagnostic Features
ceousluster. Geographical range—Sierra
Madre de Chiapas, Chiapas and Guate- Cyrtolaus oaxacanus, New Species
mala C.furculifer Bates, p. 238. (Figs. lA-C)
06'. Elytron with interval 3 not raised api-
cally. Disc ofpronolum with microlines Type material.—HOLOTYPE male and
dense,surfacewithsericeousluster.Geo- ALLOTYPE female, each labelled: MEX.
graphical range—western part of Mesa OAXACA 10.4 km./ s. Tolontepec cloud/
Central, Chiapas
07(04'). ElytronwithC.ingtreurmnuefuresrgWrhosds.ly&puBnacltla,tep.. 238. f1o9r7es9t:7t9r-e3e8f;erMnsE/XiIn CliAtteNr 2E4X8P0.m.1/97J9u/neJ.1S7.,
Pronotum with hind anglesrounded, lo- Ashe, G. E. Ball/ and D. Shpeley/ Collec-
VOLUME 93, NUMBER 2 233
Fig. 1. MalegenitaliaofCyrtolausoaxacanus. newspecies: A, median lobe, apical portion, dorsalaspect; B
andC,median lobe, with internal saceverted—B, leftlateral aspect; C, rightlateralaspect. Scalebar= 1.0 mm.
Legend(forfeaturesofinternal sac): 1, 2,and 3, spinosesclerites 1, 2. and 3, respectively;C-scl, C-sclerite; V.I.,
ventral lobe.
tors. The holotype is in USNM, and allo- The 2 pairs of supraorbital setae distin-
type in UASM. guish adults ofC. oaxacanus from C. white-
Type locality.—Mexico, Oaxaca, Sierra headi, the third species o^Cyrtolaus known
Zempoal, 10.4 km S. Totontepec. from Oaxaca, the adults ofwhich have only
Derivation ofspecific epithet.—The Lat- a single pair ofsuch setae.
inized adjectival form of "Oaxaca," the The virtually impunctate elytral inter-
name ofthe Mexican state that includes the neurs, angulate elytral apices and isodia-
type locality ofthis species. metricmesh patternoftheelytradistinguish
Recognition.—The type specimens re- adults of C. oaxacanus from those in the
semble closely the holotype of C newtoni mountains of Chiapas and Guatemala.
Whitehead and Ball. However, they are Males are distinguished by details of the
larger (Table 1), and the pronotum is rela- genitalia (Figs. 1A-C): apex ofmedian lobe
tively shorter, as reflected by higher values acute (Fig. lA); internal sac with sclerite 1
forthe ratio ML/PL(Table 2). These 2 taxa hardly evident (Fig. IB), and spinose scler-
occur in different but adjacent mountain ites 2 and 3 small, and widely separated
ranges(C oaxacanusin theSierraZempoal, from one another (Figs. IB, C).
at 2400 m; C. newtoni in the Sierra Madre Description.—Adultswithcharacterstates
de Oaxaca, at 1700 m). ofsubgenus Cyrtolaus, and following. Data
234 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON
Figs. 2-7. PronotaandleftelytraofspeciesofCyrtolaus. 2-4, Pronotum, dorsalaspectof: 2, C. whiteheadi,
new species; 3, C. subtridescens Whitehead and Ball; 4, C. ricardo Whitehead and Ball. 5-7, Left elytron. A,
dorsal aspect, B. outline ofleft lateral aspect, respectively, of: 5, C. whiteheadi, newspecies; 6, C subiridescens
Whiteheadand Ball; and 7, C. ricardoWhitehead and Ball.
VOLUME 93, NUMBER 2 235
Table 1. Data about variation in Standardized Body Length (SBL—mm) among samples ofthe species of
Cyrtolaus{s. str.).
236 PROCEEDINGS OFTHE ENTOMOLOGICAL SOCIETY OFWASHINGTON
Table 3. Data about variation in values for the ratio PL/EL among sample ofthe species ofCyrtolaus {s.
str.).
VOLUME 93, NUMBER 2 237
1975; MEX. EXP. 1975/ G.E. Ball &./ H.E. Chorological affinities.—The locality for
USNM
Frania/ collectors. Holotype in and this species is the northernmostforthe sub-
paratype in UASM. genus Cyrtolaus. It is 80 km west of the
Type locality.—Mexico, Oaxaca, Sierra locality ofthe next nearest species (C. new-
Madre de Oaxaca, 1 1.7 km E. Huautla de toni) and is on the western rim ofthe Rio
Jiminez. Grande basin, which is probably one ofthe
Derivationofspecificepithet.—Latinized major barriers for montane taxa in eastern
genitive form of the surname of the late Mexico.
—
Donald R. Whitehead, in whose honor and Phylogenetic relationships. Evidently,
C C
memory this species is named. this species is related to newtoni and
Recognition.—Adultsofthisspecieshave oaxacanus, based on the derived form of
a single pair ofsupraorbital setae, a feature the posterior angles of the pronotum, and
distinguishing them from all other species on geographical proximity but disjunct dis-
ofCyrtolaus, except C.furculifer Bates and tribution ofthe three species.
C
grumuferWhitehead and Ball. Adults of
Cyrtolaus subiridescens Whitehead and Ball
the latter 2 species have the apical portions
ofone or more elytral intervalselevated {cf. (Figs. 3, 6A, B)
—
Whitehead and Ball 1975: 600, Figs. 12, Material examined. Sixteen specimens,
C
13A, B), whereas in whitehead!, the api- as follows. All localities are in Mexico, in
cal portion ofinterval 2 is not more convex the state ofChiapas, on the Mesa Central.
thanisthebasalportionofthisinterval(Figs. Municipio Mitontik, 1 1 km N. Tzonte-
5A, B). huitz, on road to Pueblo Viejo, 1981 m,
Description.—With features ofsubgenus X.22.1976 (7 males, 3 females-CASC).
Cyrtolausand following. See Tables 1-3 for MunicipioTenejapa, TenejapaCenter, 2042
data about variation in Standardized Body m, III.15.1975 (male-CASC). Municipio
Length and in values forratios ML/PL and Las Margaritas, 48 km NE Las Margaritas
PL/EL. on road to Campo Alegre, 2134 m,
Head, antennae and mouthparts average X.25.1976 (2 males, 2 females-CASC).
for subgenus Cyrtolaus. MunicipioLasRosas,VillaLasRosas, 1371
Pronotum. Form as in Fig. 2, anterior m, X.30.1976 (female-CASC).
angles obtuse, narrow, protruded slightly; Taxonomic notes.—In body size (SBL)
posteriorangles subspinose, protruded pos- and in proportions (ML/PL and PL/EL) all
teriorly;lateralareaselevated,morebroadly of the specimens are within the ranges re-
so posteriorly; lateral margins slightly sin- ported previously (Tables 1-3; cf. White-
uate posteriorly. head and Ball 1975: Tables 1-3). The ma-
Elytra. Inform, as in Figs. 5A, B, vaulted. terialfrom MitontikandTenejapaistypical
Humerus narrowly rounded; lateral margin inallrespects,asdeterminedbycomparison
sinuate preapically, apex with short spine with paratypes ofC. subiridescens.
about opposite termination of interval 2. The specimens collected at Las Margar-
Intervals slightly convex; intemeurs finely itas and Villa Las Rosas, however, are dis-
and sparsely punctate on disc, punctures tinctive in having fairly long apical elytral
larger on apical declivity. spines (Figs. 6A, B). In pronotal form (Fig.
Legs. Average for subgenus Cyrtolaus. 3) and in details ofthe male genitalia, these
Habitat.—The type material was found specimens are typical of C. subiridescens.
indamp leafHtteronagentle slope in cloud The difference in elytral spination, in the
forest. genus as a whole, is important phylogenet-
Geographical distribution.—Known from ically, and one might be tempted on this
the type locality only. basis to propose taxonomic recognition, at
-
238 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
the subspecies level. The localities, how- 1975: 610). The range of morphological
ever, are within the known geographical variation is extended slightly, but not tax-
C
range of subiridescens (Whitehead and onomically significantly so.
Ball 1975: Fig. 23). Under this circum-
stance, I doubt that the difference in elytral Cyrtolausfurculifer Bates
spinationisindicativeofanythingmorethan Material examined.—Five specimens, as
minor intraspecific geographical variation.
follows. All were collected on the north-
Cyrtolaus ricardo Whitehead and Ball eastern slopes of the Volcan Tacana, in
(Figs. 4, 7A, B) southeasternChiapas, Mexico. UnionJuar-
— ez, Barranca Providencia, montane tropical
Material examined. Eleven specimens,
forest, in leaf litter, 1500 m, XII.15.1975
as follows. The localities are in the Sierra (male and female, UASM). Union Juarez,
(cMMsaiaenddvdTrelaneecdamAenamalCe,ehrsiNi,acEpafa.sesl,moMapoEeln,Xe—It1h8CUe3O0AP,aSmcM,iC)fhiX.icIaIvp.Gea2Urs0s.A.aTn1Vt9oE7lo-6-f FmUMiu,AnxScbXMaaI)lIM.,.u2x6bU.1an14li96,o73e5nxm(pJ2,iulaefrXeeoImzf,Iam.lo2eausg1rl-.id1cUy9uA7lwt6SouMroa(d)lf,.eml1aa4Rln3ied0o,,
MALA. San Marcos Sivinal, cloud forest, 1720 m, Xn.14-26.1975 (set of elytra
2621 m, XII.31.1976 (male, 2 females- UASM).
UASM).
Taxonomicnotes.—Thespecimensnoted
Taxonomic notes.—Tables 1-3 provide
above, mostofthemcollectedin ornearthe
data about variation in Standardized Body
Barranca Providencia, from where White-
Length and in values for the ratios ML/PL
head and Ball (1975: 611) reported this
andPL/EL. The specimenscollected on Ta- species,aretypicalmembersofC. furculifer,
cana are indistinguishable from the type se- asshownbyStandardized Body Lengthand
ries, which wascollected in the samegeneral proportions (Tables 1-3). The individual
caroesa.SiTvihneals,poecnitmheensslcooplelsecotfedVoatlcSaannTMaajru-- collected at 1720 m extends the known ver-
tical range ofthis species upwardbyca. 200
mulco, adjacent to and within a few hours m, but still beneath the known altitudinal
walkfromTacana,aresimilartotheTacana
range ofC. ricardo (see comments for that
material in pronotal form (Fig. 4), and in
species, above).
details of the male genitalia (2 males dis-
sected). In proportions, the 2 groups differ Cyrtolaus grumufer Whitehead and Ball
slightlyinvaluesfortheratioML/PL(Table
2). Also, the apical spines of the elytra of Material examined.—One female, from
theSan Marcosspecimensare shorter(Figs. Chiapas, Mexico, in the Mesa Central: Cer-
NW
7A, B). ro Baul, slope, west of Rizo de Oro,
Elevation.—The new material collected 1768 m, X.2.1979(CASC).
on Volcan Tacana extends the known alti- Taxonomic notes.—This specimen is
tudinal rangeofC. ricardodownwardbyca. within the morphological range ofthe type
C
900 m. However, this species and fur- material (Whitehead and Ball 1975: 612) as
culifer, the only other species of Cyrtolaus shown by Standardized Body Length and
inhabitingtheslopesofthisvolcano,remain body proportions (Tables 1-3). It is also
allopatric, though it seems likely that their within the reported vertical range (1524-
rangesare (orwere) in contact, at least mar- 1829 m), and from northwestern Chiapas.
ginally. However, Cerro Baul is about 160 km from
Conclusion.—The new material is within theonlyareareportedpreviouslyforC. gru-
the general geographical range that we mufer. This is a marked but not surprising
ascribed to C. ricardo (Whitehead and Ball range extension for this species.
VOLUME NUMBER
93, 2 239
Notes about Phylogenetic Aspects Breedlove. It is a distinct pleasure, also, to
A reconstructed phylogeny ofthe species acknowledge the continuing efforts of Dr.
ofCyrtolaus was offered by Whitehead and Breedlove, adistinguished systematistinthe
Ball (1975: 615, Fig. 25). The newdata sug- Academy's Department of Botany, to en-
gest that the Oaxacan species (C oaxacan- rich the entomological holdings of his in-
us, C. newtoni, and C. whiteheadi) form a stitution.
clade apart from a Chiapan-Guatemalan Finding specimens ofCyrtolaus is both a
complex (C brevispina, C. ricardo, C. spini- challenge and a delight. I am grateful to H.
cauda, C. furculifer, and C grumufer). E. FraniaandD. Shpeley,whoaccompanied
The longer apical elytral spines reported me on the trips when the new Oaxacan
C species of Cyrtolaus were taken, and who
for 2 samples of subiridescens render C
collectedthe specimensof whiteheadiand
equivocal the position ofthistaxon. Ifthese
spineswere developed independently ofthe C. oaxacanus, respectively. I thank also
Chiapan-Guatemalancomplexnotedabove, Henry Frania, J. Belicek, and D. R. Procter
for their efforts to obtain material of Cyr-
C. subiridescens could continue to occupy a
tolaus during Christmas time sojourns on
relatively basal position, as postulated pre-
viously by Whitehead and Ball (1975). If, the chilly slopes ofVolcan Tacana.
The careful illustrations for this contri-
however, the longer spines are indicative of
butionwerepreparedbyDianeHollingdale.
relationship (and their absence from most
C D. Shpeley, Assistant Curator ofthe Strick-
specimens of subiridescens a loss), then
land Museum, provided the curatorial as-
this species could be placed with the Chia-
aplatne-rGnautaitveemoanlacnhcooromlpolgeixc.alI fgarvoourndtsh.e lTathtiesr sisFtuanncdeinognfwohrifciheldIadnedpemndu.seum work and
forpublicationcostswasprovidedbyGrant
resolution postulatesa monophyletic Oaxa-
A- 1399, Natural Sciences and Engineering
can assemblage and a monophyletic Chia-
Research Council ofCanada.
pan-Guatemalan assemblage, separated
from one another by the Isthmus of Te-
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the relationships ofthe Oaxacan species of like taxa ofsouthern Mexico: Classification, and
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newtoni and C. whiteheadi, their relation- dae: Ptewstichus). Transactions ofthe American
Entomological Society 108: 315-399.
ships to one another and to C. oaxacanus Bates, H. W. 1882. Insecta, Coleoptera, Carabidae,
will remain equivocal. Consequently, it Vol. 1, Part 1, pp. 40-152, plates iii-v. In God-
seemsprematuretopursuethistopicfurther man,F. D.,andO. Salvin,eds., BiologiaCentrali-
at this time. Americana. 1879-1911.Coleoptera, 7 volumesin
17 parts. London.
Acknowledgments Liebherr, J. K. 1986. Barylaus, new genus (Coleop-
tera: Carabidae) endemic to the West Indies with
I take pleasure in thanking D. H. Kava- Old World affinities. Journal of the New York
naugh, Department of Entomology, Cali- Entomological Society 94: 83-97.
fornia Academy of Sciences, for making Whitehead,D.R.andG.E.Ball. 1975. Classification
of the Middle American genus Cyrtolaus Bates
availableforstudymaterialo^Cyrtolausthat (Coleoptera: Carabidae: Pterostichini). Quaes-
was collected in Chiapas by Dennis E. tiones Entomologicae 11: 591-619.