Table Of ContentPHYLOGENY AND REVISION OF
THE ANCHOMENUS CLADE:
THE GENERA TETRALEUCUS,
ANCHOMENUS, SERICODA, AND
ELLIPTOLEUS (COLEOPTERA:
CARABIDAE: PLATYNINI)
JAMES K. LIEBHERR
BULLETIN
OF THE
AMERICAN MUSEUM OF NATURAL HISTORY
NUMBER 202 NEW YORK: 1991
Recent issues ofthe Bulletin may be purchased from the Museum. Lists ofback issues ofthe
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PHYLOGENY AND REVISION OF
THE AN-CHOMENUS CLADE:
THE GENERA TETRALEUCUS,
ANCHOMENUS, SERICODA, AND
ELLIPTOLEUS (COLEOPTERA:
CARABIDAE: PLATYNINI)
JAMES K. LIEBHERR
Associate Professor and Curator
Department ofEntomology, Cornell University
Research Associate, Department ofEntomology
American Museum ofNatural History
BULLETIN OFTHE AMERICAN MUSEUM OFNATURAL HISTORY
Number 202, 163 pages, 77 illustrations, 8 tables
Issued March 14, 1991
Price: $12.75 a copy
Copyright © AmericanMuseumofNaturalHistory 1991 ISSN0003-0090
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Fig. 1. Anchomenuscapensis, new species, holotype male.
CONTENTS
Abstract .......................................................... 5
Introduction .......................................................... 6
Materials and Methods ........................................................ 7
Taxonomic Material ........................................................ 7
Methods .......................................................... 8
Adult Characters ......................................................... 10
Taxonomic Treatment ......................................................... 26
Key to Adults ofSpecies ofthe Anchomenus Clade ........ ..................... 26
Tetraleucus Casey ......................................................... 30
Tetraleucuspicticornis (Newman) ............... ............................ 31
Anchomenus Bonelli ........................................................ 33
Anchomenus dohrnii Fairmaire ................ ............................. 37
Anchomenus dorsalis (Pontoppidan) ............ ............................ 39
Anchomenus cyaneus Dejean ............................................... 42
Anchomenus virescens (Motschulsky) ............ 43
............................
Anchomenus leucopus Bates ................................................. 45
Anchomenus yukihikoi (Habu) .................. 46
............................
Anchomenus quadratus (LeConte) .............. 47
............................
Anchomenus aeneolus (LeConte) ............... 49
.............................
Anchomenusfunebris (LeConte) ............... 52
.............................
Anchomenus capensis, new species .............. 56
............................
Species Discrimination: A.funebris complex ......... ........................ 58
Sericoda Kirby ......................................................... 60
Sericoda quadripunctata (DeGeer) .............. 62
............................
Sericoda lissoptera (Chaudoir) .................. 68
............................
Sericoda obsoleta (Say) ..................................................... 70
Sericoda bogemannii (Gyllenhal) ............... 76
............................
Sericoda ceylonica (Motschulsky) ............... 80
............................
Sericoda bembidioides Kirby . 82
..............................................
Sericoda montana, new species ................ 88
.............................
Elliptoleus Bates ......................................................... 90
Elliptoleus acutesculptus Bates .................. 92
............................
Elliptoleus olisthopoides Bates .................. 96
............................
Elliptoleus corvus, new species ................ 97
.............................
Elliptoleus curtulus Bates . 97
..................................................
Elliptoleus vixstriatus (Bates) . 103
..............................................
Elliptoleus luteipes Csiki ........ ............................................ 106
Elliptoleus zapotecorum, new species ............ 107
............................
Elliptoleus crepericornis Bates . .............................................. 108
Elliptoleus whiteheadi, new species .............. 109
............................
Elliptoleus balli, new species . 110
..............................................
Elliptoleus tequilae, new species ................ 112
............................
Life History Data ......................................................... 113
Cladistic Analysis ofAdults..................................................... 116
Characters 116
..........................................................
Character Polarities ......................................................... 118
3
4 BULLETIN AMERICAN MUSEUM OFNATURAL HISTORY NO. 202
Results .................................. 118
Cladistic Relationships .................................. 118
Character Evolution .................................. 122
Diagnostic Characters .................................. 123
Cladistic Analysis ofLarvae ................................. 124
Tetraleucuspicticornis ................................. 125
Anchomenus dorsalis ................................. 125
Anchomenusfunebris ................................. 128
Sericoda bembidioides ................................. 130
Elliptoleus curtulus ................................. 132
Elliptoleus vixstriatus ................................. 134
Derived Character States .................................. 134
Biogeographic Analysis .................................. 137
Biogeographic Patterns inAnchomenus 137
...................................
Biogeographic Patterns in Sericoda ............... 138
...................
Biogeographic Patterns in Elliptoleus ................. 141
.................
Summary Scenario forAnchomenus Clade .................................. 148
Acknowledgments .................................. 150
References .................................. 150
Appendices .................................. 158
1991 LIEBHERR: ANCHOMENUS CLADE 5
ABSTRACT
Taxa comprising theAnchomenus cladewithin species possess limited powers of dispersal and
the carabid tribe Platynini, subtribe Platyni, pos- exhibitveryrestrictedgeographicranges,whereas
sess the synapomorphy ofa female spermatheca Sericoda species readily fly and possess some of
with a basal reservoir and a long apical filament. thelargestgeographicrangesobservedwithinCar-
As presently constituted, this clade comprises 29 abidae. Thus, relatively less anagenetic change is
speciesarrayedinfourgenera; themonotypic Te- associatedwith speciation inthe less vagile more
traleucus Casey in eastern North America, the endemicElliptoleus, andextensiveanagenesisoc-
HolarcticAnchomenusBonelliwith 10species,the curs during the history ofthe vagile, widespread
Holarctic Sericoda Kirby with 7 species, and the Sericoda species.
Mexican Elliptoleus Bateswith 11 species. Seven First-instarlarvae of6 ofthe 29 speciesarede-
species are newly described; Anchomenus capen- scribed,andacladistichypothesisofrelationships
sis,n.sp.fromBajaCaliforniaSur,Sericodamon- based on six larval characters is proposed. This
tana, n. sp. from Cuba, Elliptoleus corvus, n. sp. hypothesis supports the monophyly ofthe sister
fromMexicostate,Elliptoleuszapotecorum,n.sp. genera Sericoda and Elliptoleus, but is otherwise
from Oaxaca, Elliptoleus whiteheadi, n. sp. from discordantwiththe cladistic hypothesis based on
Guerrero,Elliptoleusballi, n.sp. fromJaliscoand adult characters. Autapomorphies ofthe first-in-
Michoacan, and Elliptoleus tequilae, n. sp. from star larvae ofthe six taxa are listed in order to
Jalisco. Newcombinationsproposedinclude: Te- facilitate comparison of derived states in newly
traleucus picticornis Newman (removed from discovered larvae.
Agonum Bonelli); Anchomenus virescens Mot- The biogeographic patterns within the individ-
schulsky (removed from Chlaeniomimus Seme- ual genera are analyzed using cladistic biogeo-
now, 1889,asthatgenericnameisanewsynonym graphic methodology. Theentire cladeis hypoth-
ofAnchomenus Bonelli, 1810); Anchomenus yu- esized as Eocene in age, with the basal taxon
kihikoi (removed from Agonum [Nipponanchus] Tetraleucus isolated from Old World progenitors
Habu, 1978, as that subgeneric name is a new ofthe rest ofthe clade due to amphi-Atlantic vi-
synonymofAnchomenus); Sericodaceylonica(re- cariance.Anchomenusisdivisableintotwoclades;
moved from Agonum Bonelli); and Sericoda lis- fourNewWorldspeciesdistributedalongthePa-
soptera(removedfromAnchomenusBonelli).Spe- cificcoastfromAlaskato BajaCalifornia,andsix
cies-level names placed into synonymy within OldWorldspeciesdistributedfromJapantowest-
Anchomenus, followed by their respective senior ern Europe. Beringian vicariance in the Miocene
synonyms, include: Dohrni Diana Sahlberg = isthelikelycauseofthispattern. Sericodaspecies
dohrnii Fairmaire, discophorus Chaudoir = dor- exhibitacollectiveHolarcticdistribution,withone
salisPontoppidan, cyaneus asturicus Heinz = cy- speciesfoundinmontaneregionsofsoutheastAsia
aneus Dejean, gracilicollis Jakowleff = virescens and Indomalaya. Beringian vicariance has oc-
Motschulsky.Batenus?borealisMotschulskyisre- curred at least twice during the diversification of
movedfromsynonymyunderSericodabogeman- the seven species leading to divergence of: (1) S.
nii,andisrecognizedasajuniorsynonymofAgon- ceylonica from its sistergroup ofS. bembidioides
umconsimileGyllenhal.Lectotypesaredesignated plus S. montana, (2) the sister species S. boge-
for Anchomenus dohrnii diana Sahlberg, Ancho- manniiandS.obsoleta.Thesevicarianteventsare
menus virescens Motschulsky, A. leucopus Bates, hypothesizedtohave occurredinlate Mioceneor
A.funebrisLeConte,SericodalissopteraChaudoir, Pliocene times. Elliptoleus was probably isolated
S. ceylonica Motschulsky, Elliptoleus acutesculp- in Mexico from its more northern sister group,
tusBates,E.olisthopoidesBates,E.curtulusBates, Sericoda,inlateMiocene. Cladisticbiogeographic
E. vixstriatusBates, E. luteipesCsiki, andE. cre- analysis allows derivation ofa fundamental area
pericornis Bates. cladogramforthetemperate humidmontane for-
A cladistic hypothesis ofthe phylogenetic re- est habitats occupied by Elliptoleus species. The
lationshipsamongthespeciesisformulatedbased Sierra Madre Occidental is hypothesized as the
on 65 potential synapomorphies ofadults, repre- sister area of the Transvolcanic Sierra plus the
sentedby 54 characters. ThegeneraSericodaand moresoutherlySierradeOaxacaandSierraMadre
Elliptoleusarehypothesized assistergenera, with del Sur. Biotic connections ofthe Transvolcanic
Anchomenustheir sistergroup. Tetraleucusis the Sierra with montane regions to the south are via
outgroup forthe rest ofthe clade. The amount of a filter bridge along the Atlantic versant, running
character evolution is heterogeneous across the from Pico de Orizaba to the Sierra de Oaxaca.
clade,withsignificantlylessanageneticchangeper Divergence events at the species level are geo-
internode of the cladogram within Elliptoleus graphicallyassociatedwithrivervalleys, orregions
compared to its sistergroupSericoda. Elliptoleus ofgeological uplift near major volcanoes.
6 BULLETIN AMERICAN MUSEUM OFNATURAL HISTORY NO. 202
INTRODUCTION
Generic concepts and the application of logenetic relationships within Platynini in
generic names within Platynini have varied general, andallowdiscrimination ofthegen-
substantially in previous classifications. Bo- era Agonum, Anchomenus, and Platynus in
nelli (1810) proposed three of the generic particular.
namesmostoftenconfused orvariously syn- This study presents a taxonomic revision,
onymized: Agonum, Anchomenus, and Pla- cladisticanalysis,andcladisticbiogeographic
tynus. Overtime,eachgenushasbeentreated analysis ofall species-level taxain genera re-
as distinct; Agonum and Anchomenus have latedtoAnchomenus. Thisgroup maybede-
been treated as subgenera ofPlatynus (Au- fined monophyletically based on a synapo-
douin and Brulle, 1834; Habu, 1973), Pla- morphous configuration of the female
tynusandAgonum havebeen treated as sub- spermatheca, which is composed ofa basal
genera of Anchomenus (Erichson, 1837), reservoir and long apical filament (e.g., figs.
Platynus has been treated as a subgenus of 189-197, 199). This configuration is distinct
Anchomenus with Agonum a distinct genus both from that observed in Agonum, where
(Casey, 1920),andAnchomenusandPlatynus thespermathecapossessesanapicalreservoir
have been treated as subgenera ofAgonum and long basal duct, and from that in Pla-
(Csiki, 1931; Lindroth, 1966). Likewise, the tynus, where the spermatheca possesses an
tribal name has varied from Platyniens (Au- apical reservoir and a short basal duct (Lie-
douin and Brulle, 1834) to Anchomenites bherr, 1986). Three genera cladistically re-
(Laporte, 1834) to Agonidae (Hope, 1838), latedtoAnchomenusare: Tetraleucus Casey,
with various later authors adopting varia- Sericoda Kirby, and Elliptoleus Bates. Seri-
tions on any ofthese three tribal names. The coda and Elliptoleus are sister genera, and
majorcause ofthe confusion has been alack liveinmontaneandborealhabitats.Sericoda
ofapplication oftype species to the generic hasaHolarcticdistribution,withoutlierpop-
concepts, lack of attention to the concepts ulations entering the Old World tropical
adopted by the first revisers of the group montaneregions, andElliptoleusisrestricted
(Habu, 1973), and a lack ofdetailed exami- tothemountainsofMexiconorthoftheIsth-
nation ofthe attributes ofthe type species to mus ofTehuantepec. The ranges ofSericoda
determine generic limits and relationships. speciesarequite large-someare circum-bo-
This work continues a line ofresearch in real or overlapping-and the taxon-area re-
Platyninithatisaimedatelucidatingthephy- lationships for this genus do not result in a
logeneticrelationships ofplatynine taxaona highly predictive biogeographic hypothesis.
worldwide basis. The foundation for sound Conversely, Elliptoleus speciesarelargelyal-
taxonomy is careful examination of taxo- lopatric,withnomorethan2ofthe 11 species
nomic characters, rigorous analysis ofthose sympatric in any area of endemism. Their
characters as a means for proposing hypoth- taxon-area relationships permit proposing a
esesofphylogeneticrelationships,andproper fundamentalareacladogramformontanear-
application of nomenclatural procedures to easofMexico.Anchomenusisthesistergenus
construct a classification that most closely to these two, and is distributed across the
represents phylogenetic relationships. PalearcticandalongthePacificcoastofNorth
InaworklargelybasedonNorthAmerican America. Tetraleucusisamonospecificgenus
taxa (Liebherr, 1986), I reported differences that forms the outgroup to the rest of the
hypothesizedtobefundamentalinthefemale clade,andisfoundineasternNorthAmerica.
reproductive tract configuration among var- The age ofthe clade asawhole isconsidered
ioustaxaofPlatynini. Thesedifferenceswere to be Eocene, as the outgroup status for Te-
first discovered by Schuler (1963). Habu traleucus is most parsimoniously explained
(1978) used some ofthese characters as the by its isolation from Eurasian progenitors of
basisforhisclassificationoftheJapanesepla- the rest ofthe clade due to thebreakdown of
tynine fauna. The differences within the fe- faunal connections between Europe and
maletractformthebasisforelucidatingphy- Americaduringthattime. Subsequently, fau-
1991 LIEBHERR: ANCHOMENUSCLADE 7
nal connections and fragmentations across the 29 species are represented in a separate
Beringia played an important role in the di- cladisticanalysis. The results ofthis analysis
versification ofAnchomenus and Sericoda. are compared to those obtained from the
Larvae representing the 4 genera, and 6 of study ofadult characters.
MATERIALS AND METHODS
TAXONOMIC MATERIAL MSUC Michigan State University, East Lan-
sing; R. L. Fischer
This study was based on approximately MSUM Moscow State University Zoological
7035 adult specimens obtained through the Museum, Moscow; N. B. Nikitsky
courtesy of41 institutional collections and 4 MZEL Museum of Zoology and Entomology,
private collectors. Institutional codens used University of Lund, Sweden; R. Dan-
inthetaxonomictreatment, andcooperating ielsson
curators are listed below. NHMB NaturhistorischesMuseum,Basel,Swit-
zerland; M. Brancucci
AMNH American Museum ofNatural History, NIAJ National Insititute ofAgro-Environ-
NewYork; Lee H. Herman mentalSciences,TsukubaCity;S.Yosh-
ANSP Academy ofNatural Sciences, Philadel- imatsu
phia; D. Azuma, D. Otte NMNH National Museum of Natural History,
BEMI Barr Entomological Museum, Univer- Smithsonian Institution, Washington,
sity ofIdaho, Moscow; F. Merickel D.C.; T.L. Erwin
BMNH BritishMuseum(NaturalHistory),Lon- ODA Oregon Department ofAgriculture, Sa-
don; N. E. Stork lem; R. L. Westcott
BPBM Bernice P. Bishop Museum, Honolulu; OHSU OhioStateUniversity,Columbus;C. A.
G. A. Samuelson Triplehorn
CAS California Academy of Sciences, San ORSU OregonStateUniversity,Corvallis;J.A.
Francisco; D. H. Kavanaugh DiGiulio and J. D. Lattin
CISC California Insect Survey, University of ROM RoyalOntarioMuseum, Toronto; D. C.
California, Berkeley; J. A. Chemsak Darling
CMNH Carnegie Museum of Natural History, RSM Naturhistoriska Riksmuseet, Stock-
Pittsburgh; R. L. Davidson holm; P. Lindskog
CNC Canadian National Collection, Ottawa; UASM University ofAlberta Strickland Muse-
Y. Bousquet um, Edmonton; G. E. BallandD. Shpe-
CUIC Comell University Insect Collection ley
FDPI Florida Dept. of Plant Industry, State UAZ University of Arizona, Tucson; F. G.
Collection of Arthropods, Gainesville; Werner
R. E. Woodruff UCD University of California, Davis; R. 0.
FMNH Field Museum ofNatural History, Chi- Schuster
cago; A. F. Newton, Jr. UCR UniversityofCalifornia,Riverside; S. I.
IRSU Irkutsk State University, Irkutsk; V. G. Frommer
Schilenkov UCVM UniversidadCentraldeVenezuela, Ma-
JBWM J. B. Wallis Museum, University of racay; L. J. Joly T.
Manitoba, Winnipeg; R. E. Roughley UKSM University of Kansas Snow Museum,
JECW JamesEntomological Collection, Wash- Lawrence; J. Pakaluk
ington State University, Pullman; R. S. UMMZ University ofMichigan Museum ofZo-
Zack ology, Ann Arbor; M. F. O'Brien
LACM Los Angeles Co. Natural History Mu- UNAM Universidad Nacional Autonoma de
seum; Los Angeles, CA; C. L. Hogue Mexico, Instituto de Biologia, Mexico
MCZ MuseumofComparativeZoology, Har- City; H. Brailovsky A.
vardUniversity,Cambridge, MA;D.G. UNH University of New Hampshire, Dur-
Furthand S. R. Shaw ham; D. S. Chandler
MNHP Museum National d'Histoire Naturelle, UVM University ofVermont, Burlington; R.
Paris; T. Deuve and H. Perrin T. Bell
MPM MilwaukeePublicMuseum,Milwaukee, ZMUH Zoological Museum, Helsinki; H.
WI; G. R. Noonan Silvferberg
8 BULLETIN AMERICAN MUSEUM OFNATURAL HISTORY NO. 202
CBOC Carlos Bordon, Maracay, Venezuela base ofscutellum along sutureto apex ofleft
DHKC David H. Kavanaugh, CaliforniaAcad- elytron.
emy ofSciences, San Francisco Genitalia were prepared by boiling speci-
PMOC Pierre Morvan, Clamart, France mens in distilled water with mild detergent
RENC Robert E. Nelson, Colby College, Wa- for30minutes,followedbydissection.Entire
terville, ME femaleabdomenswereremovedandallowed
Therevision ofthe speciesofAnchomenus to clear overnight in cold 10% KOH. The
is based primarily on specimens available in female reproductive tract, defensive glands,
most North American collections, supple- andhindgutwere then acidifiedinweak ace-
mentedbymaterialfromthemajorEuropean tic acid, dissected, and stained using Chlor-
collections. Knowledge of the geographic azol Black suspended in cellosolve. Defen-
ranges ofOld World species based on spec- sive glands and associated abdominal
imens I have examined has been augmented sclerites, and the reproductive tract and as-
by authoritative references (e.g., Jeannel, sociated sclerites were slide mounted sepa-
1942; Lindroth, 1945; Habu, 1978). have rately in glycerin. Male terminalia were
I
examined the types of all North American soaked overnight in 10% KOH, acidified in
species names in this genus, and have des- aceticacid, andthendissected. The aedeagus
ignated lectotypes where appropriate. I have was generally removed from its associated
attempted to examine types of European tergal apodemal ring, and parameres slightly
names onlywhenthe specieswas considered distended to remove associated membranes
valid in Csiki (1931). Many ofthe names of from the surface ofthe median lobe. The ae-
older European authors have lain in synon- deagal internal sacwas manually everted us-
ymy since Gemminger and Harold (1868), ing modified minuten nadeln in at least one
and their status is not a possible source of specimen ofeach taxon. After examination,
contention. allgenitalicpreparationswerestoredwiththe
Revision ofthe genera Tetraleucus, Seri- specimens in plastic genitalia vials.
coda,andElliptoleusisbasedonaworldwide Flight wings were examined in situ by re-
searchforappropriatespecimens,andknown laxing specimens and raising the elytra. Ve-
ranges reflect specimen availability. I note nation was examined by detaching the wing,
where I have examined types, andhave des- flatteningitinwateronamicroslide, remov-
ignatedlectotypes whereappropriate. Where ing the water with a paper towel, and then
I have not examined type specimens, I rely covering the wing with glycerin in a tempo-
onLindroth(1966)andDarlington(1971)for rary mount. Wings were drawn using a mi-
interpretation ofspecies identifications. croslide projector.
Scanning electron microscopy was used to
illustrate various features, andto explore the
METHODS
finerfeatures ofthe femalegonocoxae. Spec-
CharacterAnalysis. Adult specimens were imenswererelaxedinboilingwater,dissected
examined by stereomicroscope from 8 to in alcohol, and then dehydrated using a 10%
125 x, using fiber-optics ringlight illumina- step series from 70% to pure ethanol. Struc-
tion. Drawings were made using an ocular tures were mounted on double-sided scotch
grid. External characters are described and tape,coatedwithgoldorgold-palladium,and
analyzed using standard nomenclature. The examined using an Amray 1000 scanning
ocular ratio, used to quantify head shape, is electron microscope at 5 kV. After exami-
definedasthewidthofheadacrossthewidest nation, the gold coated parts were reasso-
portion oftheeyes, dividedbythenarrowest ciatedwithdissectedspecimensbymounting
widthacrossthevertexbetweentheeyemar- on card points.
gins. Bodysizewasquantifiedusingthestan- Larvae were prepared using the methods
dardizedbodylength, i.e., thesumof: (1)the ofGoulet(1977), andexaminedusingphase-
distancefromapexofrightmandibleinclosed contrastcompoundmicroscopyat 100-400x.
position to cervical collar, (2) the median Taxonomnic Methods. A diagnostic com-
length of pronotum, (3) the distance from binationandfulldescriptionareprovidedfor
