The bee genus Chilicola in the tropical Andes : with observations on nesting biology and a phylogenetic analysis of the subgenera (Hymenoptera: Colletidae, Xeromelissinae)

52 Pages·2002·3.6 MB·English

by  Michener, Charles Duncan, 1918-

Papers
Scientific
Museum
Natural History
The University of Kansas
11 July 2002 Number 26:1-47
The Bee Genus Chilicola in the Tropical Andes, with Observations
on Nesting Biology and a Phylogenetic Analysis of the Subgenera
(Hymenoptera: Colletidae, Xeromelissinae)^
MCZ
LIBRARY
By
JUL 2 3 2008
Charles D. Michener
HARVARD
Division ofEntomologi/, Nniural Histon/Museum, and Department ofEcologyand Evolutionan) Bi'Slbg}/,
The Uiiiivrsiti/ofKansas, Lawrence, Kansas 66045
CONTENTS
ABSTRACT 2
RESUMEN 2
INTRODUCTION 3
Acknowledgments 3
ABBREVIATIONS, TERMS, AND METHODS 4
SUBGENERAOFCH/L/COL/l 5
Key totheSubgenera of Ch/l;col4 Found in theAndesNorth ofChile 8
THE GROUPOF CHILICOLA {ANOEDISCELIS) ASHMEADI 8
KeytoMalesofthe Chilicola (Anoedicelis)ashmeadiGroup 11
Chilicola (Anoediscelis) ashmeadiCrawford 11
Ch/l;col4 (Anoediscelis) colombiana newspecies 13
Chilicola (Anoediscelis)misticanewspecies 13
Chilicola (Anoediscelis) venezuelana newspecies 13
Ch/l;col4 (Anoediscelis) wygodzinskyinewspecies 14
Chilicola (Anoediscelis) xanthostoma newspecies 14
Chilicola (Anoediscelis) xanthognatha newspecies 16
Chilicola (Anoediscelis) cooi'eri newspecies 16
Chilicola (Anoediscelis) pedunculata new species 16
'Contribution Number3230 from the Division ofEntomology, Natural History Museum,The University ofKansas. Lawrence,
Kansas66045, USA
©NaturalHistoryMuseum,TheUniversityofKansas ISSN No. 1094-0782
twL»^,.„,^
Ernst
Wu^|H*^. .t r*- -
,.; L'''-
1 ,^:-

Papers
Scientific
Museum
Natural History
The University of Kansas
11 July 2002 Number 26:1-47
The Bee Genus Chilicola in the Tropical Andes, with Observations
on Nesting Biology and a Phylogenetic Analysis of the Subgenera
(Hymenoptera: Colletidae, Xeromelissinae)^
By
JUL 2 3 2008
Charles D. Michener
HARVARD
Division ofEuiotnolo^y, Natural History Museum, and Department ofEcologyand EvolutioiumiBiolbgn/, '
Tlie University ofKansas, Laivrence, Kansas 66045
CONTENTS
ABSTRACT 2
RESUMEN
2
INTRODUCTION
3
Acknowledgments 3
ABBREVIATIONS, TERMS,AND METHODS 4
SUBGENERAOFCH/L/COM 5
Key tothe Subgenera of Chilicola Found in theAndesNorth ofChile 8
THE GROUPOF CHILICOLA (ANOEDISCELIS) ASHMEADI 8
KeytoMalesofthe Chilicola (Anoedicelis)ashmeadiGroup 11
Chilicola (Anoediscelis) ashmeadiCrawford 11
Chilicola (Anoediscelis) colombiana new species 13
Chilicola (Anoediscelis)mistica newspecies 13
Chilicola (Anoediscelis) venezuelana new species 13
Chilicola (Anoediscelis) wycodzinskyinewspecies 14
Chilicola (Anoediscelis) xanthostoma newspecies 14
Ch;ucol4 (Anoediscelis) xanthognatha newspecies 16
Chilicola (Anoediscelis) cooperinew species 16
Chilicola (Anoediscelis) pedunculata new species 16
'Contribution Number 3230 from the Division ofEntomology. Natural Historv Museum. The University ofKansas, Laurence.
Kan.sas66045. USA
©Natural HistoryMuseum,TheUniversityofKan.sas ISSNNo. 1094-0782
2 Scientific Papers, Natural History Museum, The University of Kansas
SUBGENUSHYLAEOSOMA ASHMEAD 17
KeytotheAndean Speciesoe Hylaesoma 18
Chilicola {Hylaesoma) aequatoriensis Benoist 18
Chiucola (Hylaesoma) involuta newspecies 20
Chilicola (Hylaesoma) umbonata newspecies 21
Chilicola (Hylaesoma) caneinewspecies 21
Chilicola (Hylaesoma) bellinewspecies 23
Chilicola (Hylaesoma) smithpardoi newspecies 24
Noteon ExtralimitalHylaesoma 26
OREOD/SCEL/S NEW SUBGENUS 26
Illustrationsof Oreodiscelis 28
Keytothe Speciesofthe Subgenus Oreod/scel/s 28
Chilicola (Oreodiscelis) espeleticola newspecies 29
NestsofChilicola espeleticola 31
Chilicola (Oreodiscelis) brooksinewspecies 32
CH/L/COL4 (Oreodiscelis) benoistiana newspecies 32
Chilicola (Oreodiscelis) quitensis Benoist 35
CH/i.;coL4 (Oreodiscelis) cuzcoens/s newspecies 36
Ch;l;col4 (Oreodiscelis) bigibbosa new species 36
Chilicola (Oreodiscelis) maculipes newspecies 39
Chilicola (Oreodiscelis) styliventris (Friese) 40
Nestsof Chilicola styliventris 41
Chilicola (Oreodiscelis) transversaria newspecies 41
Chilicola (Oreodiscelis) gibbosa newspecies 42
Chilicola (Oreodiscelis) brzoskai newspecies 43
Chilicola (Oreodiscelis) simplexnewspecies 45
PhytogenyoftheSpeciesof the Subgenus Oreodiscelis 45
LITERATURE CITED 47
ABSTRACT This is a revision ofthe species ofthegenus Chilicola found in theSouthAmericanAndes, Peru
toVenezuela,aboveelevationsof1000m. Inadditiontothefourpreviouslydescribed species known from this
area,23newspeciesare recognized and described. They fall into threegroups: (1)TheC. asliinciidigroup ofthe
subgenus Anoediscelis, for C. ashmcadi (Crawford) and the following new species: C. cokvnbiana, mistica,
veneziielana, un/godziiiskyi, xantliostonia, xaiitho^)wtha, coopcri, pediinctilntn; (2) Subgenus Hylncosoiiia for C.
aequatoriensis Benoistand the following new species: C. involuta, umbonata, canei, belli, siuithpanioi; (3) The new
subgenus Oroediscelis (type species, Chilicola styliventris) for C. styliventris Friese, quitensis Benoist, and the
following new species: C. espeleticola, brooksi, benoistiana, ciizcoensis, bigibbosa, maculipes, transversaria, gibbosa,
brzoskai, simplex. Phylogenetic analyses are presented for the subgenera of Chilicola and for the species of the
subgenus Oroediscelis. Aspects of nesting biology are presented for C. espeleticola and C. styliventris.
Key Words: Chilicola, Anoediscelis, Hylaeosoina, Oreodiscelis, Colletidae, nests. South America, Andes.
RESUMEN El presente trabajo es una revision de las especies del genero Chilicola que se encuentran en los
AndesSuramericanos desde Peru hasta Venezuela porencima de los 1000 msnm. Adicionalmentea la especie
deMesoamerica y lasquatroespeciesconocidaspara elarea,seestablecen y describen 23especiesnuevas. Las
especies se distribuyen en tres grupos: (1) el grupo ashmeadi del subgenero Anoediscelis, para C. ashmeadi
(Crawford) y las siguientes especies nuevas: C. colombiana, mistica, venezuelana, wygodzinskyi, xantliostoma,
xanthognatha,cooperi,pedunculata;(2)subgeneroHylaeosomaparaC.aequatoriensisBenoistylassiguientesespecies
nuevas: C. involuta, umbonata,canei, belli,smithpardoi; (3) el nuevo subgenero Oroediscelis (especie tipo, Chilicola
styliventris)paraC.styliventrisFriese,C.quitensisBenoist,y lassiguientesespeciesnuevas:C.espeleticola,brooksi,
benoistiana, cuzcoensis, bigibbosa, maculipes, transversaria, gibbosa, brzoskai, simplex. Se presentan analisis
filogeneticospara lossubgenerosdeChilicolaypara lasespeciesdelsubgeneroOroediscelis,ademasdealgunos
aspectos de los habitos de nidificacion para C. espeleticola y C. styliventris.
Palabras claves: Chilicola, Anoediscelis, Hylaeosoma, Oroediscelis, Colletidae, nidos, Suramerica, Andes.
Bee Genus Chiucola in the TropicalAndes
INTRODUCTION
The genus Chilicola consists of small, slender, largely
blackbees (Fig. 1) ranging from Santa Cruz Province, Ar-
gentina,andAsien,Chile,northtothestatesofTamaulipas
andJaliscoinMexicoandSt. Vincentin theLesserAntilles.
In much of this wide range, species of Chilicola are few
andspecimensrarelycollected. Theliteratureshowsthem
tobeabundantanddiverseonlyinChile,whichhasafauna
of32knownspecies,placed insevensubgenera (Toroand
Moldenke, 1979;Toro, 1986). Michener(1995)reduced the
number ofChilean subgenera to four, but the diversity is
clearly substantial.
Onlythreespeciesfrom theAndesnorthofChilehave
been described (Benoist, 1942, two species from Ecuador;
Friese,1908,onespeciesfromPeru);afourthspeciesfound
inColombia wasdescribed fromCentralAmerica. Itnow
appears, however, that there is a rich fauna in the Andes
ofPeru,Ecuador,Colombia,andVenezuela,i.e.,inthePima Fig. 1. Oiilicohi(Orocdiscclis)espdcticolaMichener,male(photo
and Paramo Nordandino and adjacent provinces recog- by M. S. Engel).
nized by Morrone (2001). The presentpaperconcerns the
27 species known from these mountains. Because these Anocdiscelis in Chile; the second ranges from the Andean
beesaresmall, because there are relatively few beecollec- region and Brazil and Bolivia, north to Mexico; the third
tors, and above all because Andean weather is so often group, Owediscelis, is limited to the Andean region north
notfavorableforflightbyinsectslikebeesthatareusually ofChile. Thus, exceptpossibly for the C. ashmeadigroup,
in their nests when the weather is cold, windy, cloudy, theAndean fauna does notconsist ofa northern montane
foggy, and rainy, collecting with a net from flowers is of- extension of the rich, temperate zoneChilean andArgen-
tennota successfulwayofsampling theAndean Chilicola tine fauna. At least the Andean Hylaeosoma presumably
fauna. Many more species will no doubt be found, for were derived from lowland tropical antecedents.
several ofthose described herein are known from a single Hi/lacosouia exists in the lowland tropics and was present
locality and one or a few specimens. Benoist (1942) ap- intheDominicanRepublicanddoubtlesselsewhereinthe
pears tohavebeen the firstperson to realize that they can tropics at least as early as Miocene times; Chilicola
be obtained from pithy stems in Ecuador (regardless of {Hylaeosoiua) gracilis Michener and Poinar (1997) and C.
the weather), and Robert W. Brooks of the University of (H.) elcctwdoniinica Engel (1999) were based on fossils in
Kansas recognized the same thing in the Venezuelan Dominican amber. The subgenus does not occur in tem-
Andes, where he collected many specimens used in this perate austral South America; its southernmost known
study. This collecting method should greatly increase localitiesarerecordedbelowinanoteonextralimital(non-
knowledgeofthisgenusinthefuture,although somespe- Andean) species at the end of the account of species of
cies may nest principally orexclusively in burrows made Hi/laeosoiiia. The origin of the Andean endemic subgenus
by beetles in wood or woody stems, rather than in pithy Owediscelis is notknown.
stems. Traps thatcan function whenever there isa period
Acknowledgements
ofsunny weather also would be worth trying.
Allspecimensusedinthepresentpaperwereobtained I am especially indebted toDr RobertW. Brookswho
at or above the elevation of 1,000 m. The only lowland collected many of the specimens ofAndean Chilicola and
species known from Peru to Venezuela are Chilicola who assembled much oftheborrowed material forstudy.
(Hylaeosoma)stciwcephalaBrooksand MichenerfromAma- Healsoprovided thenestsand informationonnestingbi-
zonian Colombia and an undescribed speciesprobably of ology oftwospecies. Itwasouroriginal intenttopublish
the C. ashiiicadi group (males not known) from Talara in this study jointly, but pressure of other responsibilities
northern coastal Peru. made his furthercontributions impossible.
Andean Chilicola can be easily divided into three Dr. Michael S. Engel made the photographic illustra-
groups: (1) the C. ashiiieadi group usually included in the tions and the analyses of my character matrices with
subgenus Anocdiscelis, (2) the subgenus Hylacosoina, and NONA.
(3) the new subgenus Owediscelis. The firstgroup ranges The cooperation of many museum curators and col-
from the Andean region to Mexico, with other species of lectorswho loaned specimensis muchappreciated. They
Scientific Papers. Natural History Museum, The Univershv oi Kansas
arelistedinthesectiononAbbre\iations.Termsand Meth- Finally, Dr. J. S. Ashe, Curator-in-charge of the Ento-
ods. Roy S. Snelling and Bryan N. Danforth were espe- mology Division, University of Kansas Natural History
cially helpful in arranging for the disposition of impor- Museum and BiodiversityResearchCenter, kindl\-permit-
tant material. ted my use of space and facilities.
ABBREVIATIONS, TERMS, AND METHODS
—
T. tergum, followed by the number of a metasomal Davis = Department of Entomology, University of Cali-
tergum—; thus Tl is the first tergum of the metasoma. fornia, Davis, California 95616, U.S.A. [L. Kimsey].
S. sternu—m (asexplained forT). Ithaca = Department of Entomology, Cornell University,
Alveolus . antennal socket. — Ithaca, New York 14853, U.S.A. [E. R. Hoebeke].
Distal stigmal perpendicular. an imaginary line LaMolina = MuseodeEntomologia, UniversidadNacional
acrossthefrontwing(Fig.3)tliroughtheapexofthestigma Agraria La Molina,nearLima,Peru [ClorindaVergara,
and at a right angle to thecostal margin ofthe wing. The C. Rasmussen].
descriptionsbelow record where this line leaves the mar- Lawrence =Entomology Division and Snow Entomology
ginalcell,inrelationtothesecondsubmarginalcell,i.e.,in Collection,Unix'ersityofKansasNaturalHistoryMu-
relation to the anterior end of the first or of the second seum, Lawrence, Kansas 66045, U.S.A.
submarginal crossvein. Lima = Museode Historia Natural, UniversidadNacional
Measurements and illustrations of the hind tibia of Mayor de San Marcos, Lima, Peru [G. Lamas, C.
males of the subgenus Oroediscelis appear repeatedly in Rasmussen].
the following pages. The measurements were made on Logan = Bee Biology and Systematics Laboratory, Utah
the upper outer view with the preapical brush of hairs State University, Logan, Utah 84322, U.S.A. [T. L.
characteristicofmostspeciesofthissubgenusinfullview, Griswold].
the crests on the under surface being out of sight. For il- London=TheNaturalHistoryMuseum, LondonSW75BD,
lustrations, however, the tibia, is tilted to show the sum- England, U.K. ]C. Taylor].
mit of the innercrest visiblebehind the outer crest. Los Angeles = Natural History Museum of Los Angeles
ForadditionalcommentsontheillustrationsofOroediscelis, County, Los Angeles, California 90007, U.S.A. [R. R.
see the accouiitofthat subgenus. Snelling].
Facial proportions were obtained by dividing the Lyme Regis = M. Cooper collection, L\-me Regis, Dorset,
length(apexofclypeus tosummitofvertex in facial view) England, U.K.
by the width (greatest distance between apparent outer Maracay = Museo del Instituto de Zoologia Agricola,
limitsofeyes in facial view), usinganeye-piece microme- Universidad Central de Venezuela, Maracay 2101A,
ter in a dissecting microscope. Venezuela ]]. L. Garcia, L. J. jolly].
Forewing length was measured with thesameequip- Medellin=MuseoEntomologico"FranciscoLuisGallego,"
ment from the base of the enlarged base of vein R to the Unixersidad NacionaldeColombia,Medellin,Colom-
wing apex. bia jGilberto Morales-Soto].
For all illustrations divided by a vertical line, the left New York = American Museum of Natural History, New
hand side is dorsal, the right hand side ventral. York, New York 10024, U.S.A. G. Rozen].
[J.
Following is a list of collections and museums con- Ottawa =CanadianNationalCollectioninCentreforLand
taining material that I used, the cities by which they are and Biological Resources,Ottawa,Ontario,Canada JL.
identified (in italics) in thetext,and inbrackets, thenames Dumochel].
of the persons who facilitated my use of the specimens. Paris= Mu.seum National d'HistoireNaturelle,75005Paris,
Aarhus = Claus Rasmussen collection, to be placed in France [J. Casevitz-Weulersse].
Aarhus, Denmark. PCAM-Austin = Programa Cooperati\'o sohre la
Berkeley = Essig Entomological Museum, University of Apifaima Mexicana, specimens at Central Texas
California, Berkeley, California 94720, U.S.A. [Cheryl Melittological Institute, .Austin, Texas78731, U.S.A.
Barr]. [J. L. Neff].
Berlin = Museum fiir Naturkunde der Humboldt- Sao Paulo = Museu de Zoologia, Unixersidaele de Sao
Universilat, Berlin, Germany [F. Koch]. Paulo, Sao Paulo, Brazil [R. F. Brandao].
Bogota Humboldt = Instituto Alexander von Humboldt, San Francisco = California Academy of Sciences, Golden
Santa Fe de Bogota 2 DC, Colombia [F. Fernandez]. GatePark,San Francisco94118,U.S.A. [VV.J. Pulawski].
Bogota Univ. = Departamento de Biologia, Universidad Washington = National Museum of Natural History,
Nacional deColombia, Santa Fede Bogota,Colombia Smithsonian Institution, Washington, D. C. 20560,
(G. Nates Parra, and V.H. Gonzalez-B.j. U.S.A. [R.J. McGinley].
.
Bee Genus Chiucola in the TropicalAndes
In listing collecting localities, I have given provinces important, because of occasional duplication of place
ordepartments in italics. When these were not on labels, names within a country. Bracketed names are my inter-
thesenamesareinbrackets. Thisissometimespotentially pretations, and therefore possibly incorrect.
SUBGENERAOFCHILICOLA
Michener(1995,2000)divitiedtheXeromelissinaeinto Table2. Charactersand theirstatesforsubgenericanalysis.
two tribes, Xeromelissini and Chilicolini, the last for two
genera, Chilicola and Xeiiochilicola. Michener and Rozen 1 innerorbits: nearlystraight(0);slightlyemarginateatupper
third or fourth (1).
(1999), however, described the genus Geodiscelis, which 2. Anteriortentorialpit: notextended(0);extendeddownward
combines characters of the two tribes. They therefore did asdeepshininggroovealongepistomalsuturenearlytoapex
not recognize tribes in the subfamily. ofclypeus (1).
Toshed lighton thegenus Chilicola asa whole, and to 3. Head ofmale: not more than 1.2 times as long as wide (0);
show relationships and justification for the new Andean more than 1.5 timesas long aswide (1).
subgenus, a preliminary phylogenetic study was made, 45.. FMaacliaalrfsopvaecea:oflfienmeaarle(:0);abbsreonatde(0r);twhealnl-ldoenfgin(1e)d; (l1o).nger than
using representatives of all genus-group taxa including broad (2).
those that were synonymized by Michener (1995, 2000). 6. Clypeusofmale: dark (0);with yellow (1).
The species used in this study of Chilicola, with current 7. Clypeusoffemale: dark (0);withyellow (1).
generic and subgeneric names, are listed in Table 1. The 98.. FFaarraaooccuullaarraarreeaassooffmfeamlael:e:dadrakrk(0)(;0)w;iwtihthyeylellolwow(1)(.1).
outgroups were Xenochilicoln, Xeromelissa, and Geodiscelis, 10. Faraocular lobe: absent (0); extending downward into
species 18 to 20 in Table 1. clypeus (1).
The characters, characterstates, and their codes used 11. Face: withoutdepressionextendingdorsolaterallv froman-
in the study are listed in Table 2. Table 3 is a matrix that tennal socket (0); with suchdepression (1).
shows the distribution of the character states among the 12. Labrum: broaderthanlong (0);aboutas longasbroad (1).
13. Last antennal segment of male: normal (0); much reduced
taxa. (1);a merenub so thatantenna appears tobe 12-segmented
AnalysiswasbyperformedbyMichaelS. Engel.Char- (2).
acterstatesweretreatedasnonadditive(notordered).The 14. Pronotum: with dorsal surface small (short) and at same
datamatrixwasconstructedinWinclada(Nixon,1991)and level as scutum (0); very short and below level of scutum,
otfheNaoniamly(sGioslwobaosffm,a1d9e93)u.siTnwgothmeirme/i;*maanldlmeangxt*hctorememsawnedrse 15. emExpetidseitnedurimnnagplabgrertlodooevwcel:sicvreioxtbtoauelsndg(ir1)no.govteol(1o)w.erpartofthorax(0);not
produced through Winclada (Length 84, Consistency In- 16. Basal area ofpropodeum: about as long as metanotum (0);
dex 52, Retention Index 64), using unambiguous optimi- longerthanmetanotum(1);abouttwicelengthofmetanotum
zations only. Figure 2 is a strict consensus tree based on (2); about three times lengthofmetanotum (3); shorter than
metanotum
these two. (4).
17. Stigma with marginsbasal tovein r: divergingapically (0);
Table 1. Species used in phvlogeneticanalysisofthesubgenera paralell (1).
ofChilicola. 18. Stigma with margin within marginal cell: convex (0);nearly
straight (1).
1. C. (Anoediscelis)ashmcadi (Crawford) 19. Distal stigmal perpendicular: crossing near middle of sec-
2. C. (Anoediscelis)herbsti(Friese) ond submarginal cell (0); crossing near base ofsecond sub-
3. C. (Stenoediscehs) inermis (Friese) marginal cell (1); crossing near apex ofsecond submarginal
4. C. {Chilicolas.str.) mbriventrisSpinola cell;crossingbasal to second submarginal cell (3).
5. C. {Chilioediscelis)patagonicaToreand Moldenke 20. Hindtibialspurs: slenderandalmoststraight(0);strongand
6. C. (Hylaeosomn) mcxicannToreand Michener curved (1).
7. C. {Hylaeosomn)aeqnatoricnsis Benoist 21. Claws of female: bifurcate (0); with inner ramus rudimen-
8. C. {Oediscelis) ivnialis (Philipps) tary (1).
9. C. {hiloprosopii) solerviceiisiToroand Moldenke 22. Hind basitarsus of male: simple (0); with ventral swelling
10. C. {Oediscelisca) dalmcidai Moure basally or medially (1).
11. C. {Heteroediscelis) mavidaToroand Moldenke 23. Hindfemurofmale: notoralittleswollen(0);stronglyswol-
12. C. {Prosopoides)firosopoidcs (Ducke) len (1).
13. C. {Pseiidoscelis) rostrata (Friese) 24. Hind tibia of male: longer than femur, when folded reach-
14. C. {Owediscelis)espeleticola Michener ingbase oftrochanter unless swollen so thatitcannotdo so
15. C. {Owediscelis) brzoskaiMichener (0);shorter than toas longas femur (1).
16. C. (unassigned) hahiiiHerbst 25. SI ofmale:withlargetubercleorprojection(0);withoutsuch
17. C. (unassigned)^v;i/f(['m'r; Moure a projection (1).
18. XcnochUicola mamignaToro & Moldenke 26. Hind tibia of male: unmodified (0); strongly swollen dis-
19. Xeromelissa wilmattaeCockerell tally(l).
20. Geodiscelis iriegacephala Michenerand Rozen 27. Hind trochanter of male: simple (0); with ventral angle or
protuberance (1).
Scientific Papers, Natural History Museum, The University of Kansas
Table2. Continued
28. S4tifmale: simple (0); with twotuberclesorprojections(1). 34. Malegonoforceps: with ventral, mesial angleor lobedistal
29. S8 ofmale, distal process: slender, deeply bitid (0); greatly tovolsella (0); withoutsuchanangleor lobe (1).
broadenedmedially,bifid(1);truncateorcmarginate,ifbroad- 35. Thoraciclength(measuredfromposteriormarginofpronotal
ened,broadestatapex (2);slenderand pointed apically (3). lobe to metasomal articulation): little more than thoracic
30. S7ofmale, apical lobes: distinct from body ofsternum (0); height (lateral view) (0); nearly 1.5 times thoracic height, or
unrecognizably fused tobodyofsternum (1). more (1).
31. S7ofmale: withfourapical lobes(0);withonepairoflobes 36. Topofhead: notgreatlydeveloped abovesummitsofeyes,
reduced sothatonly twoareeasily recognizable (1). margin in facial view extending mesiad from slightlyabove
32. Apexofpenisvalve: withtwomembranousappendages(0); summits of eyes and upper ocular tangent usually passing
with one such appendage (1); without such appendages (or throughoraboveanteriorocellus(0);stronglyde\'elopedabove
they areminute) (2). summitsofeyes,margininfacial viewcontinued upward be-
33. Malegonostylus: recognizablydistinctfromgonocoxite(0); yondeyesbeforebendingmesiadandupperoculartangentin
indistinguishably united withgonocoxite(1). astrictlyfacial view usuallybelowanteriorocellus(1).
5 9 16 17 18 19 23 26 35
010113110 Xeromelissa wilmattae
1 12 21 29 30 31 33
113 110 Geodiscelis megacephala
Prosopoidesprosopoides
2 3 1 Pseudoscelis rostrata
3 5 18 24
6 22 26 28 29 32 12 11 Oroediscelis espeleticola
r-C>-»0 • • O- —
c 1 1 1 1 Oroediscelis brzoskai
Stenoediscelis inermis
Anoediscelis ashmeadi
Anoediscelis herbsti
—
3
Hylaeosoma mexicana
Hylaeosoma aequatoriensis
Xenochilicola mamigna
Oediscelisca dalmeidai
UNASSIGNED gutierrezi
Heteroediscelis mavida
- UNASSIGNED hahni
6 8 13
Idioprosopis solervicensi
1
iJ 16
Oediscelis vernalis
2 1 —
20 Chilicola rubriventris
15 21 2i 25 26
110 10 Chilioediscelispatagonica
Fig. 2. Preliminary phylogeneticstudyofthesubgenera ofChilicola; Xeromelissaand Geodiscelisareoutgroups. This is a strict
consensus tree based on two minimum-length trees found by NONA (Length 84, CI 52, RI 64). Black dots represent unique
changes,whitedots(circles)representhomoplasiouschangesthatoccurelsewherein thistree. Numbersabovedotsarecharacter
numbers; numbersbelow dotsshow thestatearisingat that point.
Bee Genus Chilicola in the TropicalAndes
Table3. Characterstatesofrepresentative taxa used in thesubgenericanalysis.
Characters 1-18
Taxon 10 11 12 13 14 15 16 17 18
C. (Anoediscelis)nshiiieiuii
8 Scientific Papers, Natural History Museum, The University of Kansas
I regard this analysisaspreliminarybecausesomany 2. Head above antennal alveolus w^ith depression (some-
cladesaresupportedbyonlyoneortwocharacters.Anew times very weak) extending up toward ocellocular re-
analysisbased on moretaxaand morecharactersisneces- gion; S8 ofmale with apical process deeply bifid (as in
mm
sar\'before the resultsshould be used asabasis forclassi- Figs. 14c, 15c);body length4.0 to5.5 butas littleas
mm
fication. Inparticular,Ihesitatetoplace/4H0t'(f;srt'//swithin 3.0 in C. sniithpardoi Hylaeosoma
Hylaeosoma. However, recognition of the new subgenus Head without depression above antennal alveolus; S8
Oroediscelis is well supported. ofmale with apical process truncate (as in Figs. 5b, 6c);
mm
KeytotheSubgener.a of Chilicola Known From theAndes body length 3.0 to 3.75 Anoediscelis
North ofChile
TheGroupof Chiucola {Anoediscelis) ashmeadi
1. Distal stigmal perpendicular crossing or near to first This group consists of nine species and ranges from
submarginal crossvein or at leastbefore middle ofsec- central Mexico (states of Jalisco and Puebla) to Peru. It
ond submarginal cell (Fig. 3a);malarspaceonethirdas agrees with the subgeneric characters as indicated by the
long as broad or more (Fig. 4d); S4 ofmale with pairof keysanddescriptionsofToroand Moldenke (1979),using
tubercles or projections; hind tibia and usually Anoediscelisinanarrowsense,andofMichener(1994, 1995,
basitarsus ofmale swollen and modified... Oroediscelis 2000), recognizing the subgenus in a broader sense, ex-
Distal stigmal perpendicular crossing near middle of cept that the stigma is sometimes as long as the costal
second submarginal cell or more distally (Fig. 3b-d); margin of the marginal cell. The following are characters
malar space linear (= absent) (Fig. 4a-c), rarely about of the species group:
one third as long asbroad; S4 ofmale simple or nearly Body length 3.0 to3.75 mm; forewing length2.4 to2.9
so; hind tibia and basitarsus ofmale slender, notmodi- mm. Coloration: Black, clypeus of male usually largely
fied 2 yelloworwith yellow area, faceotherwiseblack (orrarely
a
Fig.3. ForewingsofC/«7ico/rtwithdistalstigmal perpcndicuLirsindiciitodIn broken lines, a,C. {Oroniisceli$)espclctiivlaparatype;
b, C. (Hi/laeosoma) nearcniici (female); c, C. {Hylaeosonia) ncquntoriciniis; d, C. (Aiiocdixclis) xoiitlio^iintlw paratype. (I'hotus by M. S.
Engel.)