Revision and phylogeny of the tribes Curiini LeConte and Plectromerini Nearns & Branham, new tribe (Coleoptera: Cerambycidae: Cerambycinae)

2008·45.7 MB·English

Memoirs
of the
American Entomological Society
Number
47
Revision and Phylogeny of the Tribes Curiini LeConte
and Plectromerini Nearns & Branham, New Tribe
(Coleoptera: Cerambycidae: Cerambycinae)
By
Eugenio H. Nearns & Marc A. Branham
DepartmentofEntomologyandNematology
UniversityofFlorida
P.O.Box110620
Gainesville,FL32611-0620
USA
ABSTRACT—ArevisionandphylogeneticanalysisofthetribeCuriiniLeConteispresented.AphylogeneticanalysisofCuriini
employing31ingrouptaxa,fiveoutgrouptaxa,and39morphologicalcharacterswasconducted.Resultsofthisanalysissuggest
thatthetribeispolyphyleticwithrespecttotheoutgrouptaxachosen.ThegenusCuriusNewmanismonophyleticandstrongly
supportedbyfiveunambiguoussynapomorphies.ThegenusPlectromerusHaldemanismonophyleticandresultssupportinclu-
sionofthemonotypicgenusCuriosaMicheliasamemberofthegenusPlectromerus.ThecladePlectromerus+Curiosaisstrongly
supportedbyfourunambiguoussynapomorphies.Basedontheseresults,thegenusCuriosaissynonymizedandthenewcombi-
nationPlectromerusdominicanus(Micheli)isproposed.ToaddressthepolyphylyofCuriini,thenewtribePlectromeriniNearns&
Branham is proposed and the genus Plectromerus is designated as the type genus. The previous synonymy of the genus Pen-
tomacrusWhitewithPlectromerus issupported,aswellasaprevioustransferofPlectromeruspunctatus (Fisher) to Curius. Eight
new species of Plectromerus are described and illustrated: Plectromerus dezayasi from Nicaragua, Plectromerus giesberti from
Guatemala,PlectromerushovoreifromCostaRicaandHonduras,PlectromerusjosephiandPlectromerusturnboivifromtheDomini-
canRepublic,Plectromerus michelii from theCayman Islands, Plectromerus morrisi from Panama, and Plectromerus thomasifrom
Haiti.Inaddition,sevenspeciesareredescribedandillustrated: CuriuspanamensisBates;Plectromerusdistinctus(Cameron);Plec-
tromerusexisZayas;Plectromerusfasciatus(Gahan);Plectromerusfemoratus(Fabricius);PlectromeruspinicolaZayas;andPlectromerus
serratus (Cameron). The following new synonymies are proposed: Plectromerus costatus Cazier & Lacey = Plectromerus dentipes
(Olivier);and Plectromerus crenulatus Cazier = Plectromerusdistinctus (Cameron). DiagnosesofallknownspeciesofCurius and
Plectromerusaxepresented withnotesondistribution,diversity,andrelationships. NewcountryrecordsarereportedforP.exis
(Jamaica);P.fasciatus(Montserrat);PlectromeruspumilusCazier&Lacey(Cuba);andPlectromerusivappesiGiesbert(Honduras,Ja-
maica).KeystothefourspeciesofCuriusand27speciesofPlectromerusarepresented.Abiogeographicanalysisbasedonthere-
sultsofourphylogeneticanalysissuggeststhatmorebasalspeciesofCuriusandPlectromerusareofAntilleandistributionwhile
morederivedtaxaareofAntillean,CentralAmerican,andSouthAmericandistribution.
REVISIONANDPHYLOGENYOFCURIINIANDPLECTROMERINI
INTRODUCTION eastern Mexico, Central America (Costa Rica,
Guatemala, Honduras, Nicaragua, Panama), Ba-
The longhorned beetle tribe Curiini LeConte hamas, Greater Antilles (Cayman Islands, Cuba,
(Coleoptera: Cerambycidae: Cerambycinae), as Hispaniola, Jamaica, Navassa Island, Puerto
historically recognized, is a medium-sized group Rico), and Lesser Antilles (Grenada, Montserrat,
of Nearctic and Neotropical cerambycid beetles. St. Vincent, British Virgin Islands, U.S. Virgin Is-
As currently defined (Monne 2005), Curiini con- lands). Eight new species of Plectromerus are de-
sistsofthreegenera(CuriosaMicheli, CuriusNew- scribed herein. In addition, a new species ofPlec-
man, and Plectromerus Haldeman) containing 23 tromerus from Barbados (Lesser Antilles), P.
extant and two extinct species. A fourth genus, louisantoini, was described by Dalens & Touroult
Pentomacrus White was synonymized with Plec- (2007) as thismanuscriptwasgoingtopress.
tromerus by Giesbert (1985). A previous phyloge-
netic analysis of Curiini has not been conducted
LiteratureReview
and themonophylyofthetribeisuntested.
Curiini are of predominantly Antillean distri- In his classic work on the Cerambycidae of
butionandshowahighlevelofendemism,witha NorthAmerica, Linsley (1963) redefined the tribe
majority of species occurring in Hispaniola and Curiini based on the two species which occur in
Cuba. Theyalso occurinUSA(eastern, southeast- USA (C. dentatus and P. dentipes) and redescribed
ern,andGreatPlains),Caribbean, andrangefrom the genus Curius based on C. dentatus. Craighead
southeastern Mexico to Venezuela (Hovore 2002; (1923) described and illustrated the larva of C.
Monne & Hovore 2005; Nearns & Ray 2006). The dentatus. Fragoso (1978) illustrated the male and
tribe has traditionally been defined by the pres- female genitalia of C. dentatus in his analysis of
ence of following morphological characters: tribal classification within the subfamily Ceram-
coarsely-faceted eyes; flat, transverse head; and bycinae. Nearns et al. (2005) transferred P. puncta-
strongly clavate femora armed beneath with a tus to Curius based on an examination of the
broad tooth. holotype.
The type genus of Curiini is Curius Newman Plectromerus was first treated by LeConte
which currently contains four species: the type (1873a,b),andthenbyLeConte&Horn(1883)and
species for the genus C. dentatus Newmanknown Leng(1885).Linsley(1963)designatedObriumden-
only from USA (eastern, southeastern, and Great tatumJ.E.LeConteasthetypespecies(=P.dentipes
Plains); C. panamensis Bates known only from (Olivier))andredescribedPlectromerusbasedonP.
Panama (Central America); C. punctatus (Fisher) dentipes. Therehasbeensomeconfusionaboutthe
known only from Cuba (GreaterAntilles); and C. generic attributes ofPlectromerus andPentomacrus
chemsaki Nearns & Ray known only from (Linsley 1963; Micheli 1983; Micheli & Nearns
Venezuela (SouthAmerica) (Fig. 65). 2005), butno previous revisionaryworkhasbeen
The monotypic genus Curiosa Micheli is done. Cameron (1910) described two species in
known only from Dominican Republic (Greater Pentomacrus andprovidedakeyforspeciesinthis
Antilles). genus only. Cazier & Lacey (1952) commented on
The genus Plectromerus currently contains 19 thetaxonomicproblemcloudingthesetwogenera
extant species (P. acunai (Fisher); Plectromerus and included both in a key to 13 species. Later,
bidentatus Fisher; P. dentipes (Olivier); P. costatus Giesbert(1985)statedthatthedifferenceswerenot
Cazier& Lacey;P. distinctus (Cameron);P. crenula- sufficient to justify two genera and thus syn-
tus Cazier; P. exis Zayas; P. fasciatus (Gahan); P. onymized Pentomacrus with Plectromerus. Al-
femoratus (Fabricius);P.femoratusWhite;P. lingafel- though a few recent works mention both genera
teri Micheli & Nearns; P. navassae Nearns & (Piha et al. 2004; Poinar 1992; Vitali 2004; Vitali &
Steiner; P. ornatus Fisher; P. pinicola Zayas; P. Rezbanyai-Reser 2003), revalidation of Pento-
pumilus Cazier & Lacey; P. ramosi Micheli & macrus has not been proposed and we suspect
Nearns; P. serratus (Cameron); P. unidentatus these authors were unaware of Giesbert's syn-
Fisher; P. wappesiGiesbert and two extinctspecies onymy. Several workers provided keys to Curiini
(P. grimaldii Nearns & Branham t and P. tertiarius (Arnett 1973; Turnbow & Thomas 2002; Cameron
Vitali t). Plectromerus are recorded from USA 1910; Cazier & Lacey 1952; Micheli 1983; Vitali
(eastern, southeastern, and Great Plains), south- 2004;Vitali&Rezbanyai-Reser2003).
EUGENIOH.NEARNS&MARCA.BRANHAM
ThemonotypicgenusCuriosawascreatedwith dae. Recent works on the curiines havebeen pro-
thedescriptionofCuriosadominicanaMichelifrom vided by Micheli & Nearns (2005), Nearns (2006),
a single female specimencollected in the Domini- Nearns & Branham (2005), Nearns & Ray (2006),
can Republic. Micheli (1983) stated that this Nearns & Steiner (2006), Nearns & Turnbow
species fit Linsley's (1963) tribal definition with a (2005), Nearns et al. (2005, 2006), Vitali (2004,
fewexceptions,themostsignificantbeingthelack 2006), andVitali&Rezbanyai-Reser (2003).
of coarsely-faceted eyes {Curiosa has finely-
faceted eyes) and the number of antennomeres
Life Historyand HostPlantAssociations
{Curiosa has 10-segmented antennae, all other de-
scribed curiines have 11-segmented antennae). Littlehasbeenpublished aboutthe lifehistory
Only two additional specimens are known to and host plant associations for the majority of
have been collected since Micheli's work, one fe- curiine species. With the exception of C. domini-
male deposited at the National Museum ofNatu- cana, all known curiine species have coarsely-
ral History (USNM) and the other (gender unde- faceted eyes andarethoughttobenocturnal. This
termined) at the Museum of Comparative issupportedbytheobservationthatmanyspecies
Zoology (MCZWeb, 2007). of Curius and Plectromerus have been collected at
The genera of Curiini were first grouped to- lights at night. The finely-faceted eyes of C. do-
getherby LeConte (1873a) who included the gen- minicana suggestthatitmaybediurnal.
era Curius and Plectromerus in "Group IV, the Host plant associations for P. dentipes, a com-
Curii" and placed the tribe before the Obriini monly collected species found in USA (eastern,
Mulsant. LeConte (1873a) also provided a de- southeastern, and Great Plains) have been pro-
scription of the unifying characters for the tribe. vided by Linsley & Chemsak (1997) and Ree
In his Coleopterorum Catalogus, Aurivillius (2003). In general, curiines are collected by beat-
(1912) listed Curiini and included the genera ing dead twigs and branches of various trees
Curius,Plectromerus,andPentomacrus.Leng(1920) (Giesbert 1985; Ree 2003; Zayas 1975). Plec-
andBlackwelder(1944)placedCuriinibeforeGra- tromeruspinicolaisreportedtohaveemergedfrom
ciliini Mulsant. Arnett (1973) placed Curiini be- cut pine branches (Zayas 1975), P. fasciatus has
tween Ibidionini Thomson and Hyboderini Lins- beenreared fromgirdledInga ingoides (L.C. Rich.)
ley. Linsley (1963) and Downie & Arnett (1996) Willd. (Fabaceae) branches (Chalumeau &
placed Curiini between Ibidionini and Obriini. Touroult 2005b), and P. ramosi has been reared
Recent literature placed Curiini between Calli- from Eugenia near ligustrina (Sw.) Willd. (Myr-
diopini Lacordaire and Graciliini (Turnbow & taceae) branches (Micheli & Nearns 2005). Fe-
Thomas 2002; Monne & Hovore2003; Peck2005). males of C. dentatus were collected with
Thesedifferencesinclassificationdemonstratethe pheromone-baited traps in Illinois (Lacey et al.
need for a comprehensive phylogenetic analysis 2004). Life history and host plant associations are
ofgroup. not well understood for the majority of curiine
Early workers provided brief, unspecific de- species andmeritfurtherstudy.
scriptionsofnewspeciesandillustrationswereei- Sexual dimorphism in pronotal and/or
ther missing or of poor quality (e.g., Bates 1885; prosternal punctation has been noted in species
Fabricius 1792; Newman 1840, 1841; Olivier 1790; from several cerambycine tribes (e.g., Casey 1912;
White 1855). Improved workbeganwithGahan's Dusham 1921; LeConte 1873a; Linsley 1963; Mer-
description of Pentomacrus fasciatus in 1895. Ga- mudes & Napp 2000, 2004; Micheli & Nearns
han (1895) also recognized that White (1855) and 2005; Monne & Napp 2005; Nearns & Steiner
other workers overlooked Fabricius' description 2006). Nearns & Ray (2006) included the presence
with regard to Pentomacrusfemoratus Fabricius {- ofmale-specificpheromoneglandporesasamor-
Plectromerusfemoratus). Othernotable workers in- phological character for C. chemsaki. Histology
cludeFisher,Linsley,andZayas.Fisherwasapro- and SEM studies ofthree cerambycine species re-
lific worker who described five new species of vealed thatmale-specificpuncturescontaingland
curiinesfrom1932to 1947. Zayas (1975) described pores that are pheromone release sites (Iwabuchi
two Cuban species and provided illustrations to 1986; Nakamuta et al. 1994; Noldt et al. 1995).
alldescribedCubancuriines exceptP. ornatus and Nearns & Ray (2006) identified male-specific
P. pumilus inhisrevisionofthefamilyCerambyci- glandpores (rounded,elevatedtubercleswithcir-
REVISIONANDPHYLOGENYOFCURIINIANDPLECTROMERINI
cular median impressions) on the pronota and comm.) and another undetermined Plectromerus
prosterna of C. chemsaki (Fig. 2c), C. dentatus, C. fossil is deposited in the American Museum of
panamensis, and C. punctatus and suggested that Natural History (No. DR-10-1857). Finally, two
volatile pheromones mayplay a role inthe repro- fossilPlectromerus aredepositedintheprivatecol-
ductive behavior of these species. In addition, lection of F. Vitali (FVPC), identified by Vitali
Nearns & Ray (2006) identified male-specific (2006) as the second knownspecimens ofP. tertia-
gland pores with a different structure on the rius andP.grimaldii.
prosterna of P. dentipes. Volatile pheromone pro-
duction by curiine species is supported by the
presence of C. dentatus in traps baited with syn-
theticpheromone(Laceyetal. 2004).Arecentmor- MATERIALSANDMETHODS
phologicalsurveybyRayetal. (2006) usedSEMto
A
identifymale-specificglandporesin49additional phylogenetic species concept is applied in
cerambycine species, suggesting gland pores are this study. Species are defined as the smallest ag-
an informative morphological character that pro- gregation of populations (sexual) or lineages
videsinformationaboutnaturalhistory. (asexual) diagnosablebyauniquecombinationof
character states incomparable individuals (sema-
phoronts) (Nixon&Wheeler 1990).
Fossils
Approximately 800 specimens from the fol-
Dominican amber is renowned for its well- lowingentomologicalcollectionswerestudied:
preserved and highly diverse insect inclusions.
These ancient resins formed from extinct Hy- AMNH American Museum of Natural His-
menaea L. (Fabaceae) trees from the mid-Miocene, tory,NewYork,NewYork,USA
approximately 17-20 MYO, and have yielded a BMNH The Natural History Museum, Lon-
richfaunaofover400families and 1500speciesof don,UnitedKingdom
insects (Grimaldi 1996; Grimaldi & Engel 2005). CMNH Carnegie Museum of Natural His-
However, specimens of the beetle family Ceram- tory, Pittsburgh,Pennsylvania,USA
bycidae are not especially common in Dominican DHPC Daniel Heffern Private Collection,
amber. Linsley (1961) observed that although ce- Houston,Texas,USA
rambycid fossils were known from various parts EFGC Edmund F. Giesbert Collection,
of the world, they were generally not well stud- Gainesville (atFSCA),Florida,USA
ied. Approximately two dozen species of ceram- EMEC Essig Museum of Entomology, Uni-
bycidsweredescribedfromcompressionfossilsof versity of California, Berkeley, Cali-
the Florissant (Meyer 2003) and a cerambycid in fornia,USA
Dominican amber has been described by Martins ENPC Eugenio H. Nearns Private Collec-
&Galileo (1999). tion, Albuquerque, New Mexico,
Atleasteight fossil Plectromerus specimens are USA
known from Dominican amber. Vitali (2004) de- FDZC Fernando de Zayas Collection, Ha-
scribed the first curiine fossil, P. tertiarius, from a vana,Cuba
partial specimen included in Dominican amber FSCA Florida State Collection of Arthro-
(Fig. 18b).Nearns&Branham(2005)describedthe pods,Gainesville,Florida,USA
second curiine fossil P. grimaldii, from Dominican FTHC Frank T Hovore Private Collection,
amberandprovidedadditionalnotesontheholo- Santa Clarita, California,USA
typeofP. tertiarius. Poinar (1992) states thatspeci- FVPC Francesco Vitali Private Collection,
mens of Plectromerus have been identified from Genova, Italy
Dominican amber. Evans & Bellamy (1996: plate IESC Instituto de Ecologia y Sistematica,
41) illustrated a well-preserved Plectromerus fossil Havana,Cuba
which unfortunately was unavailable for study INBio Instituto Nacional de Biodiversidad,
duringthecourseofthis research (G. Poinar,pers. Santo Domingo de Heredia, Costa
comm.). Two additional Plectromerus fossils inex- Rica
cellent condition are deposited in the private col- JAMC JulioandCharynMicheliPrivateCol-
lection of E. Morone, Italy (D. Grimaldi, pers. lection,Ponce, PuertoRico,USA
EUGENIOH.NEARNS&MARCA.BRANHAM
—
JEWC James E. Wappes Private Collection, Specimen Preparation. Male specimens were
SanAntonio, Texas,USA preparedfordissectionbyrelaxingtheminhotwa-
LSAM Louisiana State Arthropod Museum, terforonehour.Theaedeaguswasextractedusing
BatonRouge, Louisiana,USA a technique described by McDermott & Buck
MCZ Museum of Comparative Zoology, (1959).Theapexofa#0insectpinwasbentforming
HarvardUniversity,Cambridge,Mas- a tinyhook. This was inserted into the abdominal
sachusetts,USA opening and carefully retracted so that the hook
MNDR Museo Nacional de Historia Natural, caught the aedeagus, which is gentlypulled from
Santo Domingo,DominicanRepublic the abdomen. Using this technique, the aedeagus
MNHN
Museo Nacional de Historia Natural, was extracted withoutdamaging the exoskeleton.
Havana,Cuba Theextractedaedeaguswasthenpreparedusinga
MNRJ Museu Nacional, Universidade Fe- techniquedescribedbyLingafelter(1998)wherein
deral do Rio de Janeiro, Rio de Ja- itwasplacedin 10%KOHsolutionandheatedfor
neiro, Brazil 10 minutes. This procedure removed tissues that
MZSP Museu de Zoologia, Universidade de wouldotherwiseobscurethestructures.Itwasob-
KOH
SaoPaulo,SaoPaulo, Brazil served thatleavingtheaedeagusin solution
REWC Robert E. Woodruff Private Collec- longer than 30 minutes caused excessive clearing
tion,Gainesville, Florida,USA ofthestructures and distorted the setaeatthetips
RFMC Roy F. Morris Private Collection, oftheparameres.Theclearedaedeaguswasplaced
Lakeland, Florida, USA inawatchglass containing95% ethylalcohol and
RHTC RobertH. Turnbow,Jr. PrivateCollec- the tegmen was separated from the median lobe
tion,Ft. Rucker,Alabama,USA usingforcepsandacamelhairbrushunderaNikon
SDPC Sergio DevesaPrivate Collection, San SMZ800 stereo dissection microscope. A tempo-
Vicente, Spain raryslidewaspreparedtoviewthetegmenundera
TAMU Texas A&M University, College Sta- compoundmicroscope.Thetegmenwasplacedon
tion,Texas, USA a glass well-slide with the well filled with 95%
UCRC University of California Entomology ethylalcoholandcoveredwithacoverslipandpo-
Research Collection, Riverside, Cali- sitionedbycarefullyslidingthecoverslipoverthe
fornia, USA well, rotating the tegmen into the correct align-
USNM NationalMuseumofNaturalHistory, ment. The temporary slide was viewed under a
Smithsonian Institution, Washington, Nikon Eclipse E600 compound microscope with
DistrictofColumbia,USA 2x PlanApobrightfield, 10x DIC PlanApo, and
WIBF West Indian Beetle Fauna Project, 20X DIC Plan Apo compound objectives fitted
Michael A. Ivie, Bozeman, Montana, withadrawingtube.
USA
Label redundancy among specimens is mini- PHYLOGENETICANALYSIS
mized by not repeating identical localities within
Introduction
a species treatment in Specimens Examined sec-
tions. Holotypeandallotypelabeldataarequoted Although the family Cerambycidae has re-
verbatim, and placed in quotes. Character state ceived significant taxonomic attention over the
observations of specimens were made using a lastcentury, few phylogenetic analyseshavebeen
Nikon SMZ800 stereomicroscope with 20X eye- conducted to date on the group as awhole. Napp
pieces equipped with a drawing tube. Habitus (1994) employed adult and larval morphological
photographs were produced with a Microptics characters to test the phylogenetic relationships
Digital Lab XLT photography system, an Auto- among subfamilies. She found Oxypeltidae and
Montage Pro system, and a Nikon Coolpix 995 Disteniidae to be distinct from Cerambycidae.
with an Optem microscope adapter. For detailed Also, she proposed two monophyletic subfamily
studyofsexuallydimorphicpronotalandproster- groups within the Cerambycidae: Parandrinae +
nal punctation, specimens were imaged with a Prioninae, and Spondylinae + Lepturinae +
JEOL JSM-5510LV Scanning Electron Microscope Aseminae + Cerambycinae + Lamiinae (Napp
operated at 1.5kV. 1994).Lingafelter(1998) conductedageneric-level
REVISIONANDPHYLOGENYOFCURIINIANDPLECTROMERINI
phylogenetic analysis ofthetribe Elaphidiini (Ce- prothorax; five characters (16 states) fromtheely-
rambycinae)employingmorphologicalcharacters tra and scutellum;ninecharacters (24 states) from
and implied weighting parsimony. He included the metafemora and metatibiae; four characters
99 taxa and 70 characters and found that mono- (11 states)fromthemesosternum;andonecharac-
phyly of the tribe Elaphidiini was weakly sup- ter (fourstates) frommale genitalic structures.All
ported by three characters (Lingafelter 1998). characters were run as non-additive and un-
Monne conducted a phylogenetic analysis on the weighted (Table 1).
genus Coccoderus Buquet (Cerambycinae: Torneu- The following is a description of the morpho-
tini) based on31 morphologicalcharacters and 12 logicalcharactersandtheirstatesusedinthephy-
taxa and found thatmonophyly ofthe genus was logenetic analyses. Morphological characters
supported by five characters (M.L. Monne 2005). werecodedfrombothmalesandfemales(because
Monne & Napp (2005) conducted a generic-level little sexual dimorphism is present) unless indi-
cladistic analysis of the tribe Torneutini (Ceram- cated otherwise. Character and character state
bycinae) based on 72 morphological characters numbersrefertodatacodedinthedatamatrixfor
and 31 ingroup taxa and found the tribe tobe pa- each taxon (Table 1). The data matrix was con-
raphyletic. The monophyly ofCuriini and genera structed and edited using the program WinClada
withinthe tribehasnotpreviouslybeentested. (Nixon 1999-2002). Inapplicable data was coded
asmissingdata (Strong&Lipscomb 1999).
IngroupTaxa
The ingroup taxa in our analyses consisted of 1. Eyeshape.
31 species of Curiini, including two fossil species = ovate (Fig. 37a).
fromDominicanamber,P. tertiariusandP.grimaldii 1 = ovate-emarginate (Fig. 37b).
(Table 1). Due to specimen damage and obscured 2 = subreniform (Fig. 37c).
viewsinthefossilP. tertiarius,28of39morpholog- State2isthegeneralconditioninPlectromerus.
ical characters were coded as missing for this
taxon. This resulted in a considerable amount of 2. Longestantennomere(s).
missing data (72%) for this taxon which severely = scape.
degradedresolutionduetothetaxon'stendencyto 1 = third.
groupwithothertaxainmultiple,disparateplaces 2 = fifth.
3 = eleventh.
because ofambiguities incoding (Table 1) (Nixon
1996).Therefore,thistaxonwasbothincludedand 4 = third and fifthlongest.
excluded from analyses to examine the effects of This character has been used historically to
doingso (e.g.,Miller2005). separate Curius and Plectromerus. Curiosa domini-
cana is unique among curiines for having the
OutgroupTaxa scape as the longest antennomere. State 2 is the
general condition in Plectromerus. In Curius, the
Atotaloffiveoutgroup taxawasselectedfrom thirdislongest(C. dentatus,C.panamensis,C.punc-
tribes traditionally near Curiini in the subfamily
Cerambycinae (Nixon & Carpenter 1993). Obrium tatus) orthethirdisequaltothefifth (C. chemsaki).
maculatum (Olivier) (Obriini) was chosen as the
3. Scapewith long, suberectorerectsetaeon dorsal
root taxon. In addition, the following outgroup
surface.
taxa were included: Coscinedes gracilis Bates (Cal- = absent.
lidiopini); Parommidion extricatum Martins (Calli- 1 =present.
diopini); Hypexilis pallida Horn (Graciliini); and
Perigracilia delicata Knull (Graciliini) (Table 1). 4. Baseofscapewith depression on dorsalsurface.
Characters andTheirStates = absent(Fig. 38a).
1 = shallow (Fig. 38b).
A total of 39 morphological characters was 2 - deep (Fig. 38c).
coded (19binary,20multistate).Tencharacters(27 This character is present in only three species:
states) were coded from the head, including eyes P.femoratus, P.josephinew species, and P. turnbowi
and antennae; 10 characters (30 states) from the newspecies, allfrom theGreaterAntilles.
EUGENIOH.NEARNS&MARCA.BRANHAM
5. Length ofthirdantennomerecompared tofourth. 13. Pronotum,anteriorportion ofdiskstrongly
= about 1.3 times longerorless (Fig. 39a). shining (glossy).
1 = about 1.5 times longer (Fig. 39b). = absent.
2 = about 1.7timeslongerormore (Fig. 39c). 1 = present.
6. Length offifth antennomerecompared tofourth. 14. Pronotum,dorsalsurface.
= about 1.3 times longerorless (Fig. 40a). = microsculptured with sparse, shallow
1 = about 1.5 times longer (Fig. 40b). punctation (Fig. 44a).
2= about 1.7timeslongerormore (Fig. 40c). 1 = granulose (Fig. 44b).
Curius dentatus and C. punctatus have the fifth 2 =punctatewithglabrousareas (Fig. 44c).
antennomereequal to oronlyslightly longer than 3 = microsculptured with punctures inter-
fourth. The fifth antennomere is about twice as spersed (Fig. 44d).
long as fourth in C. chemsaki and C. panamensis 4 =heavily,evenlypunctate (Fig. 44e).
and is a good character for separating those State 1 is thegeneralconditioninCurius.
speciesfromthe othertwo species inthe genus.
15. Pronotum,dorsalsurfacesetae.
7. Antennaeannulate. =withoutscattered, long, suberectsetae.
= absent (Fig.41a). 1 = with sparsely scattered, long, suberect se-
1 =present (Fig. 41b). tae.
State 1 is the general condition in all Curius 2=withdenselyscattered,long,suberectsetae.
species but is also present in two Plectromerus
species (P. dentipes andP. exis). 16. Pronotum ornamentedwithdistinct "inverted Y"
marking.
8. Antennomeres 6-10flattened. = absent(Fig. 45a).
= absent. 1 -present (Fig. 45b).
1 =present. The presence ofthis character is a synapomor-
State is the general condition in Curius. State phy of Curius and is not present in any known
1 is thegeneralconditioninPlectromerus. Plectromerus species.
9.Antennomeres 6-10producedexternallyatapices 27. Pronotalsides.
on outermargins. =nearlyparallel, slightly inflated (widest) at
= absent (Fig. 42a). middle (Fig. 46a).
1 = present (Fig. 42b). 1 = widest area distinctly behind middle (Fig.
State is the general condition in Curius. State 46b).
1 isthegeneralconditioninPlectromerus. 2 = evenly rounded, nearly cylindrical (Fig.
46c).
10.Antennal length. 3 = sides tuberculateorprotuberate (Fig. 46d).
= less than 1.3 times aslongasbody. 4= globose, sidesbroadlyrounded (Fig. 46e).
1 = 1.3timeslongerthanbodyormore. State 2 is the general condition in Curius; state
is thegeneralconditioninPlectromerus.
11. Pronotum with long,suberectsetaeanterolaterally.
= absent. 18. Pronotalconstriction.
1 =present (Fig. 43a). = somewhat evenly constricted at apex and
State 1 is the general condition in both Curius base (Fig. 47a).
andPlectromerus. 1 = slightly more constricted atbase than apex
(Fig.47b).
12. Pronotum,sub-medial, basal thirdofdiskwith 1-4 2 = slightly more constricted atapex thanbase
long,suberectsetaearisingfrom deeppuncture. (Fig. 47c).
= absent. 3 = stronglyconstricted atbase (Fig. 47d).
1 -present (Fig. 43b). State is the general condition in Curius; state
State 1 is the generalconditioninPlectromerus. 1 isthegeneral conditioninPlectromerus.
REVISIONANDPHYLOGENYOFCURIINIANDPLECTROMERINI
19. Pronotaldisk. 26. Prosternalprocess betweenprocoxae.
= withoutcalli (Fig. 48a). -thin,about0.1 timeswidthofprocoxalcav-
1 = with one slightly raised, median callus at ity (Fig. 51a).
about the center, withtwo slightly raised, subme- 1 = moderate in width, about 0.2 times width
dial calli slightly anterior to center, and two ofprocoxalcavity (Fig. 51b).
slightly raised, submedial calli slightly posterior 2 = wide, about 0.3 times width of procoxal
to center (Fig. 48b). cavity (Fig. 51c).
2 = with one moderately raised, median callus
at about the center, with two moderately raised, 27. Procoxalcavities.
submedialcallislightlyanteriortocenter,andtwo = closed (Fig. 52a).
moderately raised, submedial calli slightly poste- 1 =narrowlyopen,nearlyclosed (Fig. 52b).
rior tocenter (Fig. 48c). 2 =widelyopen (Fig. 52c).
State is thegeneralconditioninCurius. State2isthegeneralconditioninthecuriines.
20. Males with sexuallydimorphicprothoracic 28. Prosternalprocess betweenprocoxae.
punctuation. =nearlyflat,notdeclivous (Fig. 53a).
= absent (Fig. 49a). 1 = graduallydeclivous (Fig. 53b).
1 =present(Fig. 49b-f). 2 = abruptly declivous (Fig. 53c).
The presence of sexually dimorphic, protho- State is the general condition in Curius; state
racic punctation in male cerambycines has been 1 isthegeneralconditioninPlectromerus.
notedbyseveralworkers.
29. Mesosternum with broad mesocoxalprocess with
21. Scutellum surfacegranulose. lateralextensions into mesocoxae.
= absent. = absent.
1 =present. 1 =present (Fig. 54).
This character is present in two species of
Curius (C. chemsakiand C. panamensis). 30. Metafemoralarmature.
=no toothpresent (Fig. 55a).
22. Scutellum with densesetae. 1 =withone sharp tooth (Fig. 55b).
= absent. 2 =withtwosharp teeth (Fig. 55c).
1 = present. State 1 is the general condition for Curiosa,
Curius, andPlectromerus.
23. Elytraldiskconcave medially, subsuturally.
= absent. 31. Ifmetafemurarmedwith onesharp tooth, then
1 = shallow. tooth withserrations on posteriormargin.
2 = shallow tomoderatelydeep. = absent (no serrationpeaks) (Fig. 56a).
1 = feeblyserrate (small, indistinctpeaks) (Fig.
24. Elytralapices. 56b).
=broadlyrounded (Fig. 50a). 2 = moderately serrate (moderate sized) (Fig.
1 = narrowly rounded (slightly constricted or 56c).
pointed) (Fig. 50b). 3 = strongly serrate (deep, distinctpeaks) (Fig.
2 = subtruncate / rounded (Fig. 50c). 56d).
3= stronglytruncate,straightacross (Fig. 50d).
4= stronglytruncate,concaveacross (Fig. 50e). 32. Metafemurwith long,erectsetae.
5 = outer margins with large, acute spine (Fig. = absent.
50f). 1 = present (Fig. 57a).
6 = inner margins forming a blunt, curved
point (Fig. 50g). 33. Metafemora: distalportion distinctlydarkerthan
basal.
25. Elytralsetae. = absent.
=withoutscattered long, suberectsetae. 1 = present (Fig. 57b).
1 = withscattered long, suberect setae. State 1 is thegeneralconditioninCurius.
EUGENIOH.NEARNS&MARCA.BRANHAM
34. Basal (non-clavate)portion ofmetafemora male specimens were available for dissection for
compared to metafemoralclub. C. dominicana, P. acunai, P. femoratus, P. giesberti
= distinctlylonger (Fig. 58a). new species; P. grimaldii, P. josephi new species, P.
1 = aboutequal (Fig. 58b). michelii new species, P. serratus, P. tertiarius, P.
2 = distinctlyshorter (Fig. 58c). thomasinew species, and P. unidentatus. This char-
State 1 is the general conditioninPlectromerus. acterwascoded asmissingfor thesetaxa.
35. Metafemoralshape. PhylogeneticAnalysis Methods
= gradually enlarged from base, not pedun-
Phylogenetic analyses were performed using
culate-clavate (Fig. 59a).
parsimony as the optimality criterion imple-
1 =pedunculate-clavate (Fig. 59b). mentedinTNT 1.0 (Goloboffetal. 2005) usingtra-
State is thegeneralconditioninCurius. ditional (heuristic) search. The commands "50
replications" and "25 trees saved per replication"
36. Metatibialshape.
were used to find the most parsimonious trees.
= nearly straight (Fig. 60a).
Consistency Index (CI) and Retention Index (PJ)
1 = moderatelysinuate (Fig. 60b).
werecalculatedinWinClada. Bremersupportval-
2 = strongly sinuate (Fig. 60c). NONA
ues (Bremer 1994) were calculated using
(Goloboff1995)byreadingintheconsensus ofthe
37. Lengthofmetatibia in relation to metafemur. most parsimonious cladograms and using the
1 == salbioguhttleyqsuhaolrtleern,gatbho(uFtig0..761tai)m.eslength(Fig. commands "hold 1000; sub 1; find*", "hold 2000;
sub3;find*","hold4000;sub5;find*",and"bsup-
61b).
port 10." Bootstrap support values were calcu-
2 = distinctly shorter, about 0.5 times length NONA
lated in as implementedby WinClada us-
(Fig. 61c).
ing 1,000 replications, 10 search replicates, and
five starting trees per replicate without doing a
38. Metatarsus withfirst tarsomereabout twiceas
"max*" search and saving the consensus of each
longas second.
replication.
= absent (Fig. 62a).
1 =present (Fig. 62b).
Results
The presence ofthis character is a synapomor-
phy ofCurius. Phylogeneticanalysesof36taxa (includingthe
fossil P. tertiarius) and 39 morphological charac-
39. Malegenitalia. ters produced 19 most parsimonious trees of
=paramereswithmorethanthreeshortsetae length 180. The strict consensus of 19 most parsi-
projectingfromtips oflaterallobes (Fig. 63a). monious trees is poorly-resolved within the Plec-
1 = parameres with two long setae projecting tromerus + Curiosa clade (Fig. 73). Phylogenetic
fromtipsoflaterallobes (Fig. 63b). analyses of 35 taxa (excluding the fossil P. tertiar-
2 = parameres withmore thanthree long setae ius) and 39 morphological characters produced
projectingfromtipsoflaterallobes. two mostparsimonious trees (L = 180, CI = 40, RI
3 =paramereswiththreeshortsetaeprojecting = 61). The strict consensus of two most parsimo-
from tips oflaterallobes. nious trees supports the genus Curius as a mono-
Several authors have employed characters of phyletic group thatischaracterizedby fiveunam-
the parameres (e.g., Entwistle 1963; Fragoso 1978; biguous synapomorphies (Fig. 75). The genus
Franceschini2002; Komiya & Nylander2005; Lin- Plectromerus is supported by four unambiguous
gafelter 1998; Marques & Napp 2003; Mermudes synapomorphies and is monophyletic with the
& Napp 2004; Micheli & Nearns 2005; M.L. monotypic genus Curiosa as a derived member of
Monne 2005; Monne & Napp 2005; Veiga-Ferreira thePlectromerus clade. As historicallydefined, the
1964). The morphological characters present in tribe Curiini is polyphyletic due to four outgroup
the parameres of male Curiini were not found to taxa (Coscinedes gracilis, Hypexilis pallida, Parom-
be useful for species identification. However, the midion extrication, and Perigracilia delicata) being
numberand lengthofsetae atthetips ofthepara- placedbetween the Curius and Plectromerus + Cu-
mereswere useful as a generic-levelcharacter. No riosa clades (Fig. 75). Bremer support values
10 REVISIONANDPHYLOGENYOFCURIINIANDPLECTROMERINI
Table 1.Datamatrixfor36taxaand39morphologicalcharactersusedincladisticanalysisofCuriusand
Plectromerus. Inapplicablecharacterstates aremarkedwith'-' andunobserved characterstateswith'?'
10 20 30
Obriummaculatum 121000100 0111210330 0001010000 0-11200010
Coscinedesgracilis 130000011 0000400410 7001302000 0-00210103
Hypexilispallida 130001010 1000330120 7000101210 0-00200010
Perigraciliadelicata 130001000 1000100120 0001100210 0-00200002
Parommidionextricatum 130000011 0110300300 1011411211 0-10100012
Curiuschemsaki 210022110 1100101200 1101601201 1101021110
Curiusdentatus 110010110 0100101200 1001001201 1101200010
Curiuspanamensis 140022110 1100101200 1101101201 1101100110
Curiuspunctatus 110010110 0100101200 1002101201 1001200210
Plectromerusacunai 220022011 0111200010 1002501211 110110010?
Plectromerusbidentatus 220022011 0111200010 1002511221 2701101201
Plectromerusdentipes 220022111 0111200010 1002301211 1201111201
Plectromerusdezayasinewspecies 220022011 0110000011 1002001211 1301112101
Plectromerusdistinctus 220012011 0111310010 1002111211 1200112201
Plectromerusdominicanus 001011011 0110310010 7010012111 101111000?
Plectromerusexis 120012111 1100300010 0002201211 1101010201
Plectromerusfasciatus 221022011 0110310010 1001331211 1110100001
Plectromerusfemoratus 220222010 1110300410 0001001211 110021000?
Plectromerusgiesbertinewspecies 220012011 0110000010 7002001211 130011220?
Plectromerusgrimaldiit 220022011 011727001? 7001271211 100010000?
Plectromerushovoreinewspecies 220022011 0110300010 1002001211 130011220?
Plectromerusjosephinewspecies 220122011 0110300011 1002101211 110111120?
Plectromeruslingafelteri 220022011 0110300010 1002001211 1101111101
Plectromerusmicheliinewspecies 220012011 0110310010 7001111211 110011120?
Plectromerusmorrisinewspecies 021012011 0111310010 1002011111 1311112201
Plectromerusnavassae 221012011 0110310010 1001111211 1210211101
Plectromerusornatus 220012011 0110300010 1002001211 1100210101
Plectromeruspinicola 220012011 0111200010 1002001211 1100200101
Plectromeruspumilus 220012011 0111200010 1002001211 1100111001
Plectromerusramosi 220022011 0110200010 1002101211 1300101201
Plectromerusserratus 220022011 0110300010 1002101211 130011220?
Plectromerustertiariust 727722011 0??????01? ????0????7 1?????????
Plectromerusthomasinewspecies 220002011 0111010021 7001001211 100111010?
Plectromerusturnbowinewspecies 220122011 0111300010 1001201211 1100200001
Plectromerus unidentatus 220012011 0111300010 1000001211 110011120?
Plectromeruswappesi 221012011 0110310010 1002211211 1210212201