QL 568 .S7 M35 1999 Scientific Papers

Natural History Museum The University of Kansas 10 December 1999 Number 14:1-55

Phylogenetic Relationships and Classification of the Major Lineages of (), with Emphasis on the Crabronid 1

By

Gabriel A. R. Melo2

Division of Entomology, Natural History Museum The University of Kansas, Lawrence, Kansas 66045, USA

CONTENTS ABSTRACT 2 INTRODUCTION 2 Classification and Phylogenetic Relationships within the Apoidea 2 The Present Study 4 Acknowledgments 4 MATERIAL AND METHODS 4 Selection of Representative Taxa 4 Dissection of Adult Specimens 6 Character Selection and Delimitation 6 Terminology 8 Larval and Behavioral Characters 8 Data Analysis 8 CHARACTERS AND CODES FOR THEIR STATES II RESULTS 23 DISCUSSION 23 Choice of Analytical Method 23 Apoidea and its basal clades 26 Heterogynaidae 34 Ampulicidae 34

'Contribution Number 3234 from the Snow Entomological Division. Natural History Museum, and Department of Entomology, The University of Kansas.

-Present address: Departamento de Biologia, FFCLRP.Universidade de Sao Paulo. Av. Bandeirantes 3900.14040-901, Ribeirao Preto, SP, Brazil.

ISSN No. 1094-0782 © Natural History Museum, The University of Kansas

QL 568 .S7 M3 5 1999 Scientific Papers

Natural History Museum The University of Kansas 10 December 1999 Number 14:1-55

Phylogenetic Relationships and Classification of the Major Lineages of Apoidea (Hymenoptera), with Emphasis on the Crabronid Wasps 1 By

Gabriel A. R. Melo2

Division of Entomology, Natural History Museum The University of Kansas, Lawrence, Kansas 66045, USA

CONTENTS ABSTRACT 2 INTRODUCTION 2 Classification and Phylogenetic Relationships within the Apoidea 2 The Present Study 4 Acknowledgments 4 MATERIAL AND METHODS 4 Selection of Representative Taxa 4 Dissection of Adult Specimens 6 Character Selection and Delimitation 6 Terminology...^ 8 Larval and Behavioral Characters 8 Data Analysis 8 CHARACTERS AND CODES FOR THEIR STATES 11 RESULTS 23 DISCUSSION 23 Choice of Analytical Method 23 Apoidea and its basal clades 26 Heterogynaidae 34 Ampulicidae 34

'Contribution Number 3234 from the Snow Entomological Division. Natural History Museum, and Department of Entomology, The University of Kansas.

; Present address: Departamento de Biologia, FFCLRP.Universidade de Sao Paulo. Av. Bandeirantes 3900.14040-901, Ribeirao Preto, SP, Brazil.

No. 1094-0782 © Natural History Museum, The University of Kansas ISSN 2 Scientific Papers, Natural History Museum, The University of Kansas

(sensu str1cto) + [ (sensu lato) + ] 35 Sphecidae (sensu stricto) 36 Apidae (sensu lato) + Crabronidae 36 Apic/.e (sensu lato) 38 CRABRON.DAE 38 LITERATURE CITED 43 46 APPENDIX (Figures 10-82) /ARD

ABSTRACT The superfamily Apoidea is one of the three major groups of Hymenoptera Aculeata, a small and known being composed of the sphecoid wasps, the , and the Heterogynaidae, poorly were group of wasps. The phylogenetic relationships among the major lineages of apoids investigated six from larval and three using 130 characters from the morphology of the adult , morphology, characters from adult behavior. These 139 characters were analyzed under three parsimony methods: Different equal weighting, implied weighting, and successive weighting. phylogenetic hypotheses were produced by each method (55 trees under equal weighting, four trees under implied weighting, and one tree under successive weighting, for analyses including all 54 exemplar taxa). The results are used to a from implied weighting are favored over those of the other two methods and propose constitute the most basal higher level classification for the Apoidea. Heterogynaidae and Ampulicidae in three apoid clades; however, the position of Heterogynaidae remains ambiguous: implied weight- in the fourth as the sister of the ing-trees, it comes out as the sister group of Ampulicidae and group remaining Apoidea, excluding Ampulicidae. The remaining families recognized and their relation- + five subfamilies of ships are: [Sphecidae (sensu stricto) + [Apidae (sensu lato) Crabronidae]]. Only Crabronidae are recognized: Astatinae, with the tribes Astatini, Eremiasphecini and Ammoplanini; ; (including the genera Dinettes, Laphyragogus, and Xenosphex); , with the tribes Psenini (including the genera Odontosphex and Entomosericus) and Pemphredonini; and the . INTRODUCTION

The superfamily Apoidea is one of the three major since 1974 and new studies, in particular Carpenter's (1986) clades of the Aculeata Hymenoptera (Brothers 1975, Gauld investigation on the (= Brothers' and Bolton 1988, Brothers and Carpenter 1993). A peculiar Bethyloidea). Their results largely support the phyloge- difference exhibited by aculeate females in relation to the netic patterns found in these two previous works, includ- remaining Hymenoptera is their modified ovipositor, no ing the three major lineages of Brothers (1975). The now longer used for laying eggs, but only as a sting to inject widely accepted superfamilial classification for the venom into the host or prey, as well as into potential at- Aculeata proposed by Brothers (1975) is based on the rec- tackers (defensive function). As in many groups of para- ognition of these three lineages, i.e. Chrysidoidea, Apoidea sitic Hymenoptera, females of most aculeate lineages be- and . Chrysidoidea, the basal clade of the have as idiobiont , i.e., upon finding a suitable Aculeata, contains small wasps most of which behave as host, usually concealed in protected places, the female parasitoids or sometimes as cleptoparasites. Vespoidea is paralyzes it with its venomous sting and lays an egg a large assemblage of very distinct aculeate lineages; most on the host surface (Gauld and Bolton 1988). However, are parasitoids, but well-known groups like and so- several lineages of Aculeata departed from this ancestral rial paper wasps are also included, mode of life and have evolved complex nesting and social Classification and Phylogenetic Relationships behaviors to a degree not paralleled by any other group of within the Apoidea insects, except termites. Several aspects of the biology and evolution of the aculeate are and dis- wasps presented The Apoidea is composed of the sphecoid wasps cussed Evans and West-Eberhard (1970), Iwata (1976), by [Sphecidae sensu Bohart and Menke (1976)], the bees and Gauld and Bolton (1988) and Hanson and Gauld (1995). the genus Heterogyna Nagy (the genera Daycatinca and Daya of the re- The phylogeny major aculeate lineages was are treated here as synonyms of Heterogyna; see below), a cently investigated by Brothers and Carpenter (1993). This small and poorly known group of wasps placed in a fam- reevaluated Brothers' comprehensive study mostly (1975) ily of its own (Brothers and Carpenter 1993). Most apoids work, incorporating new characters systems proposed show derived life history traits compared to the ancestral Major Lineages of Apoidea aculeate behavior, with the females exhibiting for Brothers's Sphecoidea. Brothers (1975) also proposed a high degree of parental care. Their host, or better, the an informal division of the Apoidea into two groups, the immature's provisions are now transported and concealed (= Sphecidae sensu Bohart and Menke) and in a pre-existing or especially built cavity. Construction of the Apiformes (bees). a nest before evolved once prey capture apparently only with its Heterogx/na reduced size and particularly its very in the Apoidea (Melo, in prep.). reduced forewing venation remained an enigmatic group The "Sphecidae" of Bohart and Menke forms a large and for a relatively long time since its proposal by Nagy (1969). diverse assemblage of predatory wasps, attacking most This author clearly had very confused ideas about its rela- as as orders, well spiders [see Iwata (1976) and tionships with other Aculeata lineages, since he placed it Bohart and Menke (1976) for prey records]. In the monu- in a large, heterogeneous assemblage combining mental revisionary work of Bohart and Menke (1976), this 'Ampulicidae, and Cleptidae'. Brothers (1975) group was divided into 11 subfamilies: Ampulicinae, placed Heterogyna in his plumariid group based on Nagy Sphecinae, Pemphredoninae, Astatinae, Laphyragoginae (1969). Day (1984), upon gathering material of new spe- (containing only the genus Laphyragogus), Larrinae, cies from Africa, provided convincing evidence that Crabroninae, Entomosericinae (containing only Heterogyna belonged in the Sphecidae sensu Bohart and Entomosericus), Xenosphecinae (containing only Menke, placing it in a separate subfamily. Day (1984) also Xenosphex), Nyssoninae (= Bembicinae; see Menke 1997) described for the first time the females, which are brac- and Philanthinae. Larrinae and Crabroninae have been hypterous and have a very unusual morphology compared treated one in the to under name past (e.g., Evans 1964a) and other sphecoid wasps. The phylogenetic analyses by more recently, Lomholdt (1985) and Menke (1988), among Alexander (1992a) and especially by Brothers and Carpen- others, have advocated such classification. Alternative clas- ter (1993) confirmed Day's placement of Heterogyna, and sifications, based on a division of the aculeate wasps into in the latter work, the genus was assigned to a separate several superfamilies, have recognized a superfamily family, the Heterogynaidae. with the subfamilies of Bohart and Menke Sphecoidea, Alexander (1992a) was the first to investigate the rela- treated as families Krombein 1979). Others have used (e.g., tionships among the major lineages of the apoids using one for bees and but only superfamily sphecoid wasps, modern phylogenetic methods. His study combined two raised all subfamilies to level sphecoid family (e.g., major sets of morphological characters used previously in Finnamore and Michener 1993). Bohart and Menke (1976) determining relationships among sphecoid wasps: Evans' revised all of then genera sphecoid wasps known, provid- larval characters [see reviews in Evans (1959a, 1964a)] and tribal identification as ing subfamilial, and generic keys, Bohart and Menke's (1976) adult characters. One of the well a of the known of the for summary aspects biology major problems of Evans' and Bohart and Menke's works each genus. is their assumption that the major lineages of "Sphecidae" Because of the distinct feeding habits of bees compared could be properly classified without including bees among to other aculeates, including sphecoid wasps, the older them, even after admitting that some lineages of sphecoid Linnaean classifications for the Aculeata always had bees wasps seemed more closely related to bees than to the rest and sphecoid wasps in separate higher categories. The of "Sphecidae". Before Alexander's (1992a) study, sphecoid wasps were usually among a large group of fos- Lomholdt (1982) had already proposed dividing the like the into to sorial wasps (e.g., Shuckard 1837) and the bees, "Sphecidae" two, according him, monophyletic ants, were not recognized as having any clear links to a groups, one uniting Sphecinae + Ampulicinae and the other particular group. Despite relatively earlier recognition of containing all the remaining sphecid subfamilies, forming the sister of the the close relationship between bees and sphecoid wasps his Larridae, which he considered group (Miiller 1872), a formal classification placing these two bees. Alexander's study also provided ample evidence for groups into one superfamily was proposed much later the paraphyletic nature of Sphecidae sensu Bohart and (Handlirsch 1907). Such a classification received strong Menke; however, none of his analyses specifically sup- support from Michener's (1944) study on the relationships ported Lomholdt's phylogeny. The overall results of among bees. Brothers' (1975) study on the phylogenetic Alexander's analyses are inconclusive regarding the rela- relationships within the Aculeata provided reliable evi- tionships among the major groups of Apoidea, especially dence, in terms of shared derived features, for the close because numerous conflicting relations are supported by proximity between bees and sphecoid wasps. Based on the one or more of his analyses. He was well aware of the pre- of his that more phylogenetic tree obtained in his study, he placed bees and liminary status work and concluded much sphecoid wasps in his superfamily Sphecoidea; Michener remained to be done. (1986) has shown, however, that Apoidea is the valid name Scientific Papers, Natural History Museum, The University of Kansas The Present Study sas. He was an outstanding teacher, and I profited im- from an of The present study developed investigation mensely from being his student. I also thank all members the of the tribe the relationships among genera of my dissertation committee, in particular Charles Bohart Menke into Pemphredonini sensu and (1976). Early Michener, Steven Ashe, Deborah Smith and Bryan I realized that their seemed to that study Pemphredonini Danforth for their suggestions to the dissertation manu- be but evaluation of this would diphyletic, proper question script. Michener was very kind for taking me as his stu- a broader of the require investigation relationships among dent after the sudden death of Alexander, and I am very the different Because of the reso- sphecoid lineages. poor grateful for his help and support. Specimens for this work lution obtained Alexander (1992a) when by using mainly were kindly provided by several institutions. I thank in character from authors, I decided to re- systems previous particular Wojciech Pulawski of the California Academy but characters and a peat his study mostly using original of Sciences for providing material of very important taxa, of taxa not of mono- representation requiring hypotheses Robert Brooks for allowing me to dissect numerous speci- used the tribes phyly above the level of genus (Alexander mens from the Entomology Division of the Natural His- Bohart and Menke). For obvious reasons, recognized by tory Museum of The University of Kansas, and Michael received closer attention and better the Pemphredoninae Prentice for allowing me to use his unpublished data on this I am confident that re- representation. Despite bias, the biology and larval morphology of Odontosphex sults obtained here a fair into the represent investigation paradoxus, as well as for calling my attention to the close phylogeny of the major apoid lineages. relationship of Odontosphex and Entomosericus to I also thank Bruce Cutler for all his The preferred phylogenetic hypothesis found by this Pemphredoninae. help I carried in his lab and Antonio study is used to propose a higher level classification for with the work out Marques his Macintosh most the Apoidea. From now on, I will be using the classifica- for making available computer. During tion proposed here, and reference to higher taxa whose of my stay at The University of Kansas, I was financially a from the Brazilian Conselho previous definitions conflict with the ones proposed here supported by scholarship will be marked as such. Nacional de Desenvolvimento Cientifico e Tecnologico (200233/92-0). Thanks to Steven Ashe and Robert Acknowledgments (CNPq) Brooks, I was supported by a curatorial assistantship from The present work is part of my Ph. D. disserta- the Natural History Museum during my last semester. I tion, and as such, I would like to thank posthumously am also grateful to the Department of Entomology and Byron Alexander, first, for agreeing to be my Ph. D. ad- the Panorama Society for providing funds for some of my viser and also for all his support and guidance during my research activities. initial years of graduate studies at The University of Kan- MATERIAL AND METHODS

Selection of Representative Taxa genera, i.e. genera whose taxonomic positions in Bohart and Menke's classification were not supported by Representatives of all major lineages of Apoidea were Alexander's (1992a) study. Besides the taxa listed in Table included in the present study, as well as of some basal lin- 1, material of several other taxa, in particular specimens eages of aculeates as outgroup taxa. Two limiting factors of Apoidea deposited in the insect collection of the Uni- were taken into account when selecting the specific taxa: versity of Kansas, were also examined. More relevant taxa of (adult insects) for dis- availability specimens complete are listed below. section and of published information on larval morphol- Material of the following additional hymenopteran ogy. However, a few important taxa whose larvae are un- taxa not included in the were also known were still included; also, Laphyragogus and analyses completely dissected and examined: Xenosphex were included, despite lack of material for com- Crabronidae.— cockerelli Pate plete dissections (only mouthparts, including oral plate, Astatinae: Ammoplanops and external genitalia were dissected). In some cases, I also (female), AmmopIaneUus wnatilla Pate (female), chose specific genera that were considered previously to Ammoplanellus sp. (female), Dryudella sp. (female); Bembicinae: have a relatively basal position within their respective lin- Alysson melleus Say (female), sp. eages; this is the case for the exemplar taxa of bees, of (male); Crabroninae: Bothynostethus sp. (male), Ectemnius Sphecidae (s.str.), and for most of the Crabroninae and the stirpicola (Packard) (female), Entomognathus texanus Bembicinae. Table 1 lists the exemplar taxa included in the (Cresson) (male), Oxybelus emarginatum Say (female and formal parsimony analyses. Among the Crabronidae, I male), Tn/poxylon frigidum Smith (male); Pemphredoninae: tried to include representatives of all subfamilies recog- Araucastigrnus masneri Finnamore (female), Arpactophilus nized by Bohart and Menke (1976), as well as "problem" sp. (female), Carinostigmus sp. (female), Diodontus atratulus Major Lineages of Apoidea

Table 1 . List of taxa used as exemplars; ingroup taxa arranged accord- Table 1 . Continued to the classification here. F = female; M = male. ing proposed Philanthinae:

48. Aphilanthops frigidus Smith, M OUTGROUP 49. Pliilanthus gibbosus (Fabricius), F, M Bethvlidae: Heterogvnaidae: 50. 1. Epyris sp. (from Brazil), F, M Heterogyna fantsilotra Day, M Pompilidae: Sphecidae (sensu stricto): 51. Chlorion 2. Notocyphus sp. (from Costa Rica), M aerarium Patton, F Rhopalosomatidae: 52. Palmodes rufiventris (Cresson), M 3. Rhopalosoma nearticum Brues, F 53. Podalonia communis (Cresson), M Sapygidae: 54. Stangeella cyaniventris (Guerin-Meneville), F 4. Eusapyga proximo (Cresson), F : 5. Clystopsenella longiventris Kieffer, F Taschenberg (male), Microstigmus nigrophthalmus Melo (fe- Sierolomorphidae: male), cuspidatus Smith (female), 6. Sierolomorpha canadensis Provancher, M lethifer (Shuckard) (female), Polemistus braunsii (Kohl) INGROUP (male), Polemistus dickboharti Menke (female), Spilomena Ampulicidae: subterranea McCorquodale & Naumann (female), Spilomena 7. Ampulex sp. (from Costa Rica), F sp. (female), Stigmus fulvipes Fox (male), Stigmus temporalis 8. Aphelotoma rufiventris Turner, M Kohl (female); Philanthinae: Cerceris Smith 9. Dolichurus sp. (from Costa Rica), F, M rufopicta (fe- Apidae (sensu lato): male). 10. Anthophonda albata (Timberlake), F — Apidae (s.l.). Calliopsis andreniformis Smith (female), 11. Conanthalictus nigricans Timberlake, F Callomelitta 12. Ctenocolletes stnaragdinus (Smith), F antipodes (Smith) (female), Hylaeus sp. (female). 13. carinata Stevens, F Hesperapis Mutillidae.—Myrmosa unicolor Say (male). 14. Lonchopria zonalis (Reed), F — Crabronidae: Pompilidae. Aporinellus fasciatus (Smith) (female). Astatinae: — Evans 15. Astata nevadica Cresson, F Sierolomorphidae. Sierolomorpha nigrescens (male). 16. Eremiasphecium sahelense (Simon-Thomas), F 17. cfr. Pate, F Ammoplanus apache Vespidae.—Eumenes fratemus Say (female). 18. Pulverro mescalero Pate, F, M Braconidae.— 19. Timberlakena yucaipa Pate, F unidentified (female). Bembicinae: Evaniidae.—unidentified species (male). 20. Eembecinus quinquespinosus (Say), F 21. Didineis texana (Cresson), M .—unidentified species (male). 22. Heliocausus larroides (Spinola), F — . Xiphydria sp. (female). 23. spilopterus (Handlirsch), F 24. Nysson rusticus Cresson, F .—Macroxyela ferruginea (Say) (male). 25. Ochlewptera bipunctata (Say), F Crabroninae; Undissected or only partially dissected (e.g., of the taxa were also 26. Dinetus pictus (Fabricius), M terminalia) specimens following 27. Laphyragogus pictus Kohl examined: 28. Mellinus alpestris Cameron, M — Riek (female and 29. Xenosphex timberlakei Williams Ampulicidae. Aphelotoma fuscata 30. Anacrabro ocellatus Packard, F male), Dolichurus corniculus (Spinola) (female), 31. Lindenius columbianus (Kohl), F Paradolichiirus boharti Kimsey (female and male), 32. subita (Say), F Paradolichurus obidensis (Ducke) (female and male), Trirogma 33. amazonica Ducke, F caerulea Westwood (female). 34. Palarus latifrons Kohl, M 35. davisi M Fox, Crabronidae.— Clypeadon laticinctus (Cresson) (male), Pemphredoninae: Entomosericus kaufmanni Radoszkowski (female and male), 36. Odontosphex paradoxus Menke, M E. 37. Entomosericus concinnus Dahlbom, F Eremiasphecium budrysi (Kazenas) (female), longiceps 38. Mimesa cressonii Packard, F, M (Gussakovskij) (female and male), Laphyragogus ajjer 39. Pluto minutus (Malloch), F, M Beaumont (female and male), (Linnaeus) 40. Psenulus Bohart & Grissell, F mayorum (female and male), Mellinus bimaculatus Packard (female), 41. Arpactophilus steindachneri Kohl, F damara Pulawski (female and male), Palarus 42. Diodontus rugosus Fox, F, M Odontosphex 43. Parastigmus huecuvus Finnamore, F variegatus (Fabricius) (female and male), Paracrabro 44. Passaloecus areolatus Vincent, F, M froggratti Turner (female), Tiguipa cfr. fiebrigi (Brethes) 45. inornata F Pemphredon Say, (male), Timberlakena cahuilla Pate (female), Xenosphex 46. Spilomena catamarca Antropov, F Williams (female). 47. Stigmus temporalis Kohl, F xerophilus of Kansas Scientific Papers, Natural History Museum, The University

and Character Selection and Delimitation Heterogynaidae—Heterogyna protea Nagy (female African male) and males of all Heterogyna species, except Most of the characters used in the present study are for H. ravenala Day. derived from the morphology of the exoskeleton of the internal furca, 2nd Dissection of Adult Specimens adult insects, including processes (e.g., was based on phragma). Selection of characters dry, pinned from each of the taxa listed At least one adult specimen as well as on dissected specimens in glycerin, was specimens, in Table 1 and Xenosphex) SZ60 to (except Laphyragogus using a stereoscopic microscope Olympus (up as follows before examination: processed 126X) and incident and transmitted light. The remaining taken from the of immature (1) Soaking in 10% KOH solution overnight; characters were morphology for with a and from the behavior of adult females. (2) Clearing in 3% hydrogen peroxide species stages (larva) dark integument; In order to confirm the glandular nature of two to 50-70% ethanol, then for a few (3) Transfer boiling characters (82 and 83), female specimens of Ammoplanus minutes, followed slow cooling; and americanus by cfr. apache, Passaloecus areolatus, Stigmus Transfer to water and then slow addition of glycerin; were embedded in LR White (4) preserved in Kahle's fixative Transfer to and in (5) storage pure glycerin. resin following the procedures described by Lindley (1992) and In order to avoid excessive clearing of mouthparts and sectioned with a Sorvall Ultra Microtome (MT 5000); before The slides terminalia, these parts were dissected transferring the sections were slide-mounted using Euparal. in ethanol is to remove, from and under a BH-2 to peroxide. Boiling important were observed photographed Olympus the bubbles interference contrast inside the specimen, especially from head, microscope with differential optics. the Partial of the the of produced by peroxide. dismembering It is difficult to explain or justify process was carried out before transferring to pure of those specimen character discovery and subsequent delimitation since at this the is still relatively i.e. glycerin, stage integument characters into states. For complex characters, from the treatment. malleable KOH characters that show a great amount of apparently 1 with one sex listed, the information but are not divisible For species in Table only important cladistir readily and submitted in a manner terminalia of the opposite sex were removed into discrete states ur expressible quantitative described above, so that sex 58 or 68), I tried to detailed to the same procedure specific (e.g., Characters provide be so that the states here characters from this part of the body could examined; descriptions and illustrations, of other and more for three genera, only material congeneric species recognized can be apprehended properly a female of But the of was available: a male of Eusapyga sp., Aphelo- evaluated by other people. problem making a female of decision followed when toma nigricula Riek and Heterogyna protect Nagy. explicit the processes delimiting for these characters still remains. In most The heads of specimens preserved in fixative (or the states complex I included a number of states to match sometimes in alcohol) of the following species were such cases, larger the condition in the different taxa. dissected for examination of the morphology of the more closely present are However, if taken to an extreme and if the states pharynx (see characters 13 and 14): nonadditive, this can make any character useless —Dolichurus (Costa Rica, male). procedure Ampulicidae. sp. for each taxon. — by assigning a different state Apidae (s.l.). Augochlora pura (Say) (female). In some cases, to preserve the informational content — melleus (Bembicinae, Crabronidae. Say it into two Alysson of a complex character, I divided characters, female), Crossocerus (from USA, Crabroninae, male), of a structure sp. one of them representing presence or absence Didineis texana (Cresson) (Bembicinae, female), Diodontus the character or of a particular condition and second Fox (Pemphredoninae, female), Hoplisoides sp. Character 70 and flavitarsis representing its different states (e.g., pairs (from Costa Rica, Bembicinae, female), spp. (female Nysson 71, or 77 and 78). Taxa in which the structure or condition from USA, male from Costa Rica, Bembicinae), Ochleroptera is absent are assigned a question mark for the second (Bembicinae, female), bipunctata (Say) character; this corresponds to treating inapplicable (Fabricius) (Philanthinae, female), Psenulus sp. (from Costa characters as missing data. This can be problematic under Rica, female), speciosus (Drury) but the current Pemphredoninae, certain circumstances (e.g., Maddison 1993), (Bembicinae, male), alini Antropov Spilomena are not able to handle inapplicable Packard computer algorithms female), Stigmus americanus to (Pemphredoninae, characters differentially. One alternative would be have female). (Pemphredoninae, only one character, but this is exactly what was being — female), Sphecidae (s.str.). Isodontia sp. (from USA, avoided in the first place. Sphex ichneumoneus (Linnaeus) (male). Another issue that should be mentioned is the male). to characters Sapygidae.—Sapyga sp. (from USA, treatment given morphometric (approxi- Major Lineages of Apoidea

This is based on mately 25% of the characters used here, not considering assumption presence among members of In most this tribe of a 1st and meristic characters, e.g., Character 4 or 73). cases, disproportional elongate submarginal the wide a quantitative description was adopted to express a relatively separation between lm-cu and the vein most here as in like qualitative nature not easily captured as such; interpreted 2rs-m, especially genera commonly, this qualitative nature involved shape of Diodontus and Pemphredon. structures. For the labrum (Character 1) in example, Ammoplanini.—The presence of two or only one and most is Pemphredonini Ammoplanini relatively submarginal cell in this group seems at first more difficult similar in overall and in a few other attributes, but I shape to explain, because members of the basal lineage, Pulverro tried to this the express similarity mostly by proportions (and Ammoplanops), have a very reduced venation pattern. of the labrum (state 1-1 ); characters 19 and 20 are additional However, I am postulating that the ancestral lineage for examples. this tribe had two submarginal cells as seen in some species Different procedures have been developed to divide of Timberlakenn (and also in Pwtostigmus). This two-celled morphometric characters into more or less objective states condition was created by loss of 3rs-m and M distal to 2rs- of taking into consideration intra- and intertaxon variation m (character 89). These postulated vein losses, instead [see reviews by Stevens (1991) and Thiele (1993)], as well loss of the segment of Rs separating the 1st and 2nd Zelditch cells as for the are as to produce characters representing shape (e.g., submarginal suggested Pemphredonini, close the et al. 1995). I did not employ any of these procedures for inferred from the proximity between lm-cu and as this of in Pulverro and the quantitative characters used here. The characters vein here interpreted segment Rs selected are believed to show very little intraspecific Timberlakenn (see Figs. 20 and 21). Also, the venation pattern variation, since they were chosen exactly for being stable of the Ammoplanini can be easily derived by patterns across at least two representative taxa. However, the limits similar to that of some species of Eremiasphechtm (see Fig. the here. for the different states were usually arbitrary (see e.g., 22) assuming changes postulated 9 or 29) and were defined to circumscribe Characters 8, Dinetus.—The two-celled condition in this genus can two or more taxa to form a natural This thought group. be derived easily from a shortening of the marginal cell has been criticized for its "otential to bias the approach accompanied by loss of 3rs-m and the segment of M distal in favor of ideas of phylogenetic analyses pre-conceived to 2rs-m. The genus Gastrosericus (Crabroninae, Larrini) Stevens 1991, Gift and Stevens 1997). relationships (e.g., has a similar condition, but clearly independently acquired. Submitting these characters to the procedures mentioned Nitela, Lmdenius and Anacrabw.—The presence of above could diminish these potential biases, but I think it only one submarginal cell in these three genera can be would represent little improvement for the quantitative derived from a pattern with two submarginal cells (in characters used here [see also Farris (1990)]. In any case, I which the 2nd cell is petiolate) by loss of M distal to the classified all 139 characters used in the analyses segment of Rs separating the 1st and 2nd submarginal cells accordingly to their nature (see Table 4). This somewhat and loss of the segment of 2rs-m not fused with Rs. Indeed, crude sorting can be used to identify those characters in Encopognatluts [see Fig. 116A in Bohart and Menke (1976)] whose cladistic informational content should be viewed and in some species of Nitela, a clear indication (or the vein with more caution. remains in the case of Nitela) of a petiolate 2nd submarginal In the case of absence of one or more of the veins cell can be seen. delimiting the forewing submarginal cells (see Characters Hesperapis.—The two-celled condition in this genus 87-89), I used relational information to infer the putative is assumed to have occurred by loss of 2rs-m. This has losses. This problem of similarity assessment usually assumption is based on presence of a relatively long 2nd been circumvented by considering only the number of submarginal cell (i.e. 2nd and 3rd fused) and lm-cu cells present [e.g., Alexander's (1992a) submarginal connected to the segment of M delimiting the 2nd cell. In Character 59, or Alexander and Michener's (1995) bees, lm-cu apparently almost always connects to M Character 84]. However, I think this approach can lead to between the segment of Rs separating the 1st and 2nd loss of information, and therefore I tried to introduce a cells and 2rs-m. several submarginal more precise assessment, especially because —This has a somewhat unusual and of these veins. Here Heterogyna. genus lineages of Crabronidae have lost some I use here the same for the very reduced wing venation. I provide more detailed justifications similarity (1985) for H. first assessments made in each case: interpretation given by Day protea: submarginal cell complete, second cell petiolate and —The presence of only two Pemphredonini. (2rs-m but not M). to be distally open present, reaching spectral submarginal cells in this group is considered here I no effort to include known derived from loss of the segment of Rs separating the 1st made especial characters or to look for new and 2nd submarginal cells, i.e. fusion of these two cells. synapomorphic putative 8 Scientific Papers, Natural History Museum, The University of Kansas

found Lin and from the synapormorphies for the following taxa previously by Evans and (1956b), Ctenocolletes, to be monophyletic: Sphecidae (sensu stricto) (Alexander description of the larva of Stenotritus pubescens (Smith) by 1992a), Apidae (sensu lato) (Alexander 1992a, Alexander Houston (1975); these pairs of genera have very similar and Michener 1995) and the Philanthinae sensu Alexander adult morphology and nesting behavior, respectively. for taxa was taken at the level (1992b). Obviously, I paid close attention to any evidence Information outgroup family that could contradict these previous studies. Characters from Evans et al. (1987), except for Scolebythidae and found by Brothers and Carpenter (1993) to support the Sierolomorphidae, for which there is no available monophyly of Apoidea were included (however, several information. as as of them were circumscribed differently), well These six characters were selected from a list of 10 additional relevant characters resolution for providing characters considered of phylogenetic significance by relationships among the outgroup taxa. Evans (1959a). Two of the 10 were omitted because of their The sample of characters used here is not intended to apparent complexity (larval body shape and mandibles); I informative not comfortable them without the represent the result of an exhaustive search for am using examining characters, but only as one of the many samples of specimens. The two others (parietal bands and opening characters that could be extracted from these insects. Some between atrium and sub-atrium of spiracles) were given areas of the body that seem to contain important characters less importance by Evans. Alexander's (1992a) study can the of these four characters were completely ignored. For example, the internal ridges be used to evaluate significance and lamellae associated with the hypostomal bridge in the in a cladistic context. exhibit a amount of variation the taxa head large among Three behavioral characters were incorporated in the but I was unable to this variation sampled, simply organize formal analyses: type of larval food, food relocation and of and in any meaningful way. This region the body many nest construction. All prey records for ampulicids, are worth in future studies. others exploring crabronids, and sphecids were taken from Bohart and Terminology Menke (1976), except for Eremiasphecium taken from Kazenas (1991; prey records for £. budrysi Kazenas), The terms here were morphological adopted mostly Arpactophilus from Matthews and Naumann (1989; prey from Bohart and Menke Michener (1944) or taken (1976), record for A. mimi Naumann), Ammoplanus from Maneval (1942, 1993) and definitions for these terms can Snodgrass (1939; prey record for A. perrisi Giraud) and from Ahrens be found in those works. for However, terminology wing (1948; prey record for A. handlirschi Gussakovskij), characters also 14) was taken from for (see Fig. Day (1988); Laphyragogus from Kazenas (1985; prey record for L. the thoracic from Gibson (1993); for external pleuron, turanicus Gussakovskij), Entomosericus from Kazenas and from Smith Sources for a few additional genitalia, (1970). Alexander (1993; prey record for E. kaufmani terms are in the list of morphological given directly Radoszkowski) and for Odontosphex paradoxus from M. characters. For indication of direction for structures in the Prentice (pers. comm.). Biological information for the I used the convention of an insect with a head, prognathous outgroup taxa was taken from Hanson and Gauld (1995). head, so that the frons is in a dorsal position and the Data Analysis occiput, ventral. Reference is made to metasomal sclerites (Michener 1944), instead of abdominal sclerites, except for Methods incorporating parsimony have won gonocoxites and gonapophyses of female's sting. widespread acceptance among systematists interested in for the various Larval and Behavioral Characters producing phylogenetic hypotheses groups of organisms, especially because parsimony has been Six characters (131-136) derived from the external considered the only criterion that implements Hennig's of the larva for all larval morphology were used. The data auxiliary principle [e.g., Hennig (1966:121)] that the most characters were taken from the literature (Table 2), except preferable tree topology is the one that minimizes the for I also hoc Spilomena; examined larvae of a few additional number of ad hypotheses of homoplasy (e.g., Wiley taxa, like Pemphredon, Psenulus, and Stigmus, as well as of 1981, Farris 1983). Parsimony is usually associated only some taxa not included in the present study, like Sceliphron, with methods that do not assign different weights to the Cerceris, and Megaehile. There is no published information characters being used, i.e. methods in which all characters on the larvae of nine genera of the ingroup: Aphelotoma, are treated equally in terms of cladistic information they Ctenocolletes, Didineis, Eremiasphecium, Laphyragogus, provide. However, parsimony can also be applied under a Xenosphex, Timberlakena, Parastigmus, and Heterogyna. These weighting function. Herein, I employ two distinct methods taxa have missing entries for larval characters in the (implied and successive weighting) that assign differential character matrix, except Didineis, for which the states were weights to the characters based on their degree of taken from the description of the larva of Atysson melleus homoplasy. Major Lineages of Apoidea

for information on larval Table 2. List of ingroup taxa whose larval descriptions were used characters.

Genus 10 Scientific Papers, Natural History Museum, The University of Kansas

ro

i cn r- CN ID

1 CN ID

o

=8 o

ii Si U *

c ,o

-3 c o

>- O £

|2 Major Lineages of Apoidea 11 weighting characters according to their degree of Parsimony under equal weighting.—Under this homoplasy. Unlike successive weighting (see below), the procedure, the characters receive equal weight, and weights are calculated simultaneously with tree search, the therefore one change in a given character has the same different tree topologies being evaluated according to the effect as one change in any other character (except when character weights which in turn are implied by the characters differ in their ordering status). The objective is the This to find distribution of characters on a given topology. the trees that minimize the total number of changes, method seeks trees with the maximum sum of character i.e. trees of minimum length. This method was imple- weights or maximum total fitness, i.e. trees which imply mented by Nona, Version 1.8 (Goloboff 1997b), using the the characters to have, on average, as high a weight as following command sequence: hold*; hold/2; mult*300; max*. The individual character fitness is The data matrix possible. (/) given by was submitted to heuristic search using the function tree bisection-reconnection swapping (mult*), with random addition (300 The trees were I + it + 1 sequence replicates). resulting f.= (k+l) (Sj -m) examined using MacClade. where k is a constant of concavity introduced to regulate under successive — how strongly homoplastic characters are down-weighted Parsimony weighting. Successive was Farris (the higher its value, the lesser the down-weighting), s is weighting originally proposed by (1969); more (1988) has advocated the actual number of steps observed for character i on a recently Carpenter it to and m is the minimum number of applying as a means select among multiple equally given topology, i steps trees. This is an iterative in which possible for character ;' [see Goloboff (1993) for more parsimonious procedure, the character are calculated after each run of tree details]. The higher the degree of homoplasy of a character weights search, the new then to the next run. The in a given topology, the lower its weight will be. weights applied process is stopped when the results of a given iteration The analysis of the present data using the method of are identical to those produced in the iteration immediately implied weighting was implemented by the computer preceding it. The weight for each character is calculated Pee-Wee, Version 2.8 (Goloboff 1997a). Tree search program to the character = according consistency index; this index is was carried out with the command mult* adopting k 3, a function of the amount of homoplasy shown by the the program's default value for the weighting constant. character in a given tree topology. This method was carried The following command sequence was used: hold*; hold/2; * out using PAUP*, Version 4.0 b2 (Swofford 1999). The initial mult*300; max*. The command mult randomizes the order set of most parsimonious trees produced under equal of the taxa in the data matrix, creates a weighted Wagner weighting was used as the starting point. Reweighting of tree, and submits it to tree bisection-reconnection branch- characters was done according to their consistency index, swapping (300 replicates); the command max* does branch- using the maximum value (best fit) with base weight equal swapping on the trees found by mult*, looking for to 10 (weight values not truncate). The data matrix was additional trees with the same fitness. The resulting trees submitted to heuristic search using subtree pruning- were examined using MacClade, Version 3.06 (Maddison regrafting swapping, with random addition sequence (100 and Maddison 1996); character optimizations shown on replicates); also the parsimony settings were adjusted to the illustrated cladograms (Figs. 1, 2, 5 and 6) were carried have branches collapsed when their minimum length was out also using MacClade. zero, which is equivalent to amb- in Nona. The resulting trees were examined using MacClade.

CHARACTERS AND CODES FOR THEIR STATES

The morphological characters are listed according to 1. Labrum: their positions in the insect body, the structures of the head (0) at least one and a half times wider than long. less than a half times wider than listed first, followed by thorax and then abdomen; the lar- (1) one and long, very val and behavioral characters are at the end. When neces- flat (Figs. 34 and 40). sary, I also included (after listing the character and its states) Labral width measured across its base. In taxa with comments on the character or for decisions explanations state (0), the labral apex usually has numerous bristles, made while assigning states to certain taxa. The 54 exem- whereas in taxa with state (1) the labrum has few or no taxa were examined and scored for the 105 plar (Table 1) bristles. Podnlonia has a flat, long labrum, but with several characters. The data list- morphological complete matrix, apical bristles; Ampulex also has an elongate and some- ing all taxa and the state codes assigned to them, is pre- what flat labrum; both were assigned state (1). In Epyris 3. are illustrated sented as Table Morphological characters and Eusapygn, the labrum is vestigial and these taxa were in Figures 10-82 in the Appendix. assigned (?) for characters 1-3. 12 Scientific Papers, Natural History Museum, The University of Kansas

2. Labral apex: 10. Ventral surface of prementum: (0) entire and broadly rounded, or at most slightly emar- (0) continuous with lateral surfaces or separated by ginate in the middle. rounded angles. (1) entire and pointed. (1) separated from lateral surfaces by sharp angles or (2) notched in the middle (Figs. 34 and 40). carinae.

In Arpactophilus steindachneri the apex has several small The ventral surface of the prementum in Podnlonia and teeth, but the labrum is considered notched in the middle. Eusapyga is excavated longitudinally, forming a sulcus The common condition in the genus seems to be a shallow margined by two ridges; Lindenius has a similar condition notch in the middle and small lateral teeth. (although the middle part is slightly elevated). In Diodontus and Passaloecus, the between ventral and lat- 3. A pair of rounded or oval spots on base of labrum: separation eral surfaces is but there are no carinae. These (0) absent. quite abrupt, are all coded as (0). (1) present. 11. Basal of lateral arms of It is not known what those spots represent, but they margins prementum: (0) to ventral surface do not seem to be some type of sensillum; in the cleared approximately perpendicular s over 60 ) 10 and 11). they look like areas where the dorsal and ven- (angle (Figs. specimens, e (1) an acute (45 or smaller) tral surfaces of the labrum are fused, being similar to the slanting, forming angle with ventral surface 12). margins of the labrum. (Fig. (2) lateral arms absent. 4. Female mandibular apex: The lateral arms of the are considered ab- (0) apical plus one dorsal subapical tooth. prementum sent in the bees. In this the basal (articulated (1) apical tooth only, i.e., simple. group, part with the of the anterior thicken- (2) apical plus two or more dorsal subapical teeth. prementum) conjunctival ings of Michener (1944) may be homologous to the lateral Some groups, like Psenulus and some species of arms of the prementum that became detached from the Parastigmus, have a small, more basal tooth along the in- rest of the prementum. Another possibility is that the basal ner margin that it is not taken into consideration here. portion of these thickenings could have originated from a 5. Male mandibular apex: detached part of the stipes and that the lateral arms disap- (0) one dorsal tooth. apical plus subapical peared. (1) apical tooth only. 12. Paramandibular process: (2) apical plus two or more dorsal subapical teeth. (0) absent or very short, well separated from back The number of teeth varies among species of of clypeus. Heterogyna. Males oiH.fantsilotra have two subapical teeth, (1) closing at least 3/4 of mandibular socket, but not whereas in H. protea has only one tooth. Heterogi/na is as- reaching back of clypeus. signed both states, i.e., (0) and (2). (2) reaching back of clypeus, but not fused to it. 6. Subbasal cleft on inner edge of mandible (female): fused to (0) absent. (3) clypeus. In (1) present. some taxa, for example Philanthus, Aphilanthopsis, Palinodes and in This cleft or incision is found in several Crabroninae. Podnlonia, males and females differ the of of the I For an illustration of this character, see Fig. 2 in Pulawski degree development paramandibular process. (1995). chose to use the state found in females, because for most taxa only female specimens were available for complete 7. Outerventral margin of mandible: dissection. In Ampulex females, the mandibular socket is (0) simple. closed by an extension of the gena, and not of the ( 1 ) notched. hypostoma. In the species dissected, the hypostoma gets This is the same as the externoventral notch of Bohart close to but does not reach the clypeus. In the males, how- & Menke (1976). ever, the mandibular socket is closed by a hypostomal pro- 8. Glossa: cess. Ampulex is assigned state (3). (0) less than twice as long as wide. 13. Posterior wall of pharynx (between arms of oral plate): (1) at least twice as as wide. long (0) not expanded. 9. Prementum: (1) forming two bulging sacs (walls usually covered less with (0) than three times as long as apical width. numerous acanthae) (Figs. 29-33). (1) at least three times as long as apical width. Major Lineages of Apoidea 13

These structures are sometimes hard to see in speci- 17. Subantennal sutures: mens treated in KOH, especially when the pharyngeal (0) absent. walls are relatively thin. For this reason, specimens pre- (1) present, not connected to tentorial arms. served in fixative were dissected and examined; due to (2) present, connected to tentorial arms. lack of suitable specimens, most of the material dissected This character applies only to taxa assigned states (0) to belongs species (or even genera) not included in the or (1) for characters 18 or 21. Laphyragogns and Xenosphex phylogenetic analyses. Figures 31 and 33 show cross-sec- have subantennal sutures, but they are assigned (?) because tions of the sacs from material of pharyngeal prepared no internal observations were made. Ammoplanus preserved in fixative. The cleared specimen 18. Distance between antennal socket and clypeus (female): of Bembecinus has what seems to be small expansions on (0) more than one half of socket diameter. the pharyngeal wall, but no fixed material was available (1) one half of socket diameter or less, but not nil. for dissection. It was scored as (?). The male of Heterogyna (2) nil. fantsilotra has a distinct expansion of the pharynx. The 19. posterior wall of the pharynx is dilated, but does not form Epistomal suture, between antennal sockets, in taxa wherein the antennal sockets a pair of sacs, and is continuous with two short expan- are in contact with (female): sions on the upper part of the pharynx. It is assigned (1) clypeus not above despite these differences. (0) transverse median line across antennal sockets 34). 14. of (Fig. Upper part pharynx (at the tip of the oral plate arms): (1) above transverse median line across antennal sock- (0) simple, not expanded. but not to ets, reaching tangent upper rims (Fig. 13). (1) forming a pair of elongate, sometimes very large (2) extending above tangent to upper rims of antennal and branched, diverticula (Fig. 33). sockets (Fig. 14). 15. Apical inflection of clypeus: (0) lateral to tentorial This character to taxa with state (2) in the joining epistomal ridge pit (Fig. 35). applies only character. (1) joining at tentorial pit. preceding (2) mesal to tentorial joining considerably pit (Figs. 13, 20. Tentorial pit (female): 14 and 34). (0) below or level with tangent to lower rims of anten- nal sockets The term "apical inflection" was taken from Roig- (Fig. 13). (1) above to lower rims of antennal sockets Alsina and Michener (1993); see their paper for additional tangent (Fig. illustrations. The segment of the inflection being consid- 14). ered here seems to in most internal to the be, taxa, mem- 21. Distance between antennal socket and clypeus (male): brane that connects the base of the mandible to the inflec- (0) more than one half of socket diameter. tion. In Eremiasphecium sahelense, the joining of the inflec- (1) one half of socket diameter or less, but not nil. tion at the tentorial pits seems to result from an apparent (2) nil. outward displacement of the tentorial and not from pits, 22. Epistomal suture, between antennal sockets, in taxa an of the it is state In expansion inflection; assigned (0). wherein the antennal sockets are in contact with Philanthus, the inflection a lower branch of the tento- joins clypeus (male): rial arm which is broadly fused to- a laminar epistomal (0) not above transverse median line across antennal ridge. In Entomosericus and Mimesa, the apical inflection sockets. joins the epistomal ridge slightly lateral to the tentorial (1) above transverse median line across antennal sock- pits, but these taxa are coded as having state (1). The male ets, but not to rims. of Heterogyna fantsilotra has an internal longitudinal ridge reaching tangent upper each connecting internal rim of the antennal sclerite to the (2) extending above tangent to upper rims of antennal apical inflection of the clypeus. This peculiar condition is sockets. not treated as equivalent to state (2) above. This taxon is This character applies only to taxa with state (2) in the state (0). assigned preceding character. 16. Eye-clypeus contact: 23. Tentorial pit (male): (0) none. (0) below or level with tangent to lower rims of anten- for the diameter of one antennal (1) extending socket nal sockets. or less. (1) above tangent to lower rims of antennal sockets. (2) extending for more than the diameter of one anten- 24. Tentorial II: nal socket. pit (0) situated on epistomal ridge. (1) distinctly placed above epistomal ridge. 14 Scientific Papers, Natural History Museum, The University of Kansas

Schonitzer (1993)]. It differs 25. Internal rim of antennal sclerite: Schuberth and considerably in and the various taxa; I (0) level with internal surface of head or projecting only development position among consider all different forms as homologous. slightly (Fig. 34). the center and most of (1) expanded toward covering 34. Preoccipital carina: antennifer not ex- the socket (portion containing (0) complete (Fig. 44). panded) (Fig. 35). (1) interrupted ventrally. a but distinct carina. (2) expanded and forming short, cylin- (2) interrupted ventrally, but reaching hypostomal antennifer to- der (portion containing expanded (3) interrupted dorsally 36 and 37). gether with rest of rim) (Figs. (4) completely absent. In The term antennifer is taken from Michener (1944). This is the same as occipital carina of Bohart and at the of the rim. does not state (2), the antennifer remains edge Menke (1976). The species of Dolichurus dissected a but it is state (1) based 26. Pedicel attachment: have preoccipital carina, assigned on a female of D. corniculus. (0) basically centric. 35. Periforaminal (1) eccentric (Figs. 38 and 39). depression: (0) absent. 27. Socket on apex of scape with an eccentric pedicel: (1) present (Fig. 44). (0) entirely membranous. This character concerns a distinct on the (1) sclerotized in the center (Fig. 38). depression It is more dorsally, as with state (1) for occipital region. usually developed This character applies only to taxa well as and in some taxa, it is marked dorsally the character. laterally, preceding and carina. The an- and laterally by a marginal sulcus /or 28. Sexual in number of antennomeres: dimorphism terior dorsal portion of the pronotum seems to fit in this (0) none. with a depression; in the taxa well-developed marginal male with 13 and female with 12 antennomeres. (1) sulcus, one could imagine that head and pronotum are of the 29. Female 2nd flagellomere: locked together when the anterior dorsal margin (0) at least 3X longer than pedicel. pronotum is inside the marginal sulcus. In Entomosericus, than this taxon is con- (1) less than 2X longer pedicel. the depression is weakly indicated, but state This character is used to characterize the relative length sidered as having (1). of the antenna. The female of Heterogyna protea has an un- 36. Cervical sclerite: state the fact usually long pedicel; it is assigned (0) despite (0) absent. than the its 2nd is not 3X 1 ) that flagellomere longer pedicel. ( present. 30. Facets of compound eyes (female): 37. Posterolateral angle of pronotum: uniform in size. from state 1). (0) approximately (0) evenly rounded (or modified differently than ones. (1) frontal facets much larger remaining (1) reduced dorsally above and anterior to differenti- 31. Facets of compound eyes (male): ated spiracular operculum. (0) more or less uniform in size. This character corresponds to character 35 of Brothers than ones 39). (1) frontal facets much larger remaining (Fig. and Carpenter (1993). 32. Inner orbits of compound eyes (female): 38. Ventral angle of pronotum: or more or less (0) straight slightly concave, parallel. (0) scarcely exceeding base of procoxa.

1 ) below. ( concave, diverging (1) greatly produced mesad and closely approaching below. (2) more or less straight, diverging its counterpart midventrally. (3) convex, below. diverging This character is being used to indicate the distinct (4) sinuate (upper portion concave, lower portion con- condition found in all Apoidea, irrespective of the small vex), strongly converging below. variation found among them in how closely the two ven- (5) below. straight, converging tral halves of the pronotum approach each other. Brothers (6) concave, converging below. and Carpenter's (1993) assignment of a distinct state to of area of female: 33. Integument paraocular bees is unjustified, since a similar condition to that found median of frons. (0) not differentiated from more part in bees is present in several other Apoidea. (1) with a area, sometimes distinct and specialized very 39. Pronotal collar: a fovea 40-43). forming (Figs. (0) anterior edge rounded, or a collar not differenti- the surface for release This specialized area represents ated from anterior portion of pronotum. of the secretions of an underlying epidermal gland [see (1) delimited anteriorly by a transverse carina (some- Major Lineages ofApoidea 15

distinct reduction of the times interrupted in the middle) (Fig. 45). posterior of the medial portion basisternum in s.l. and in relation to (2) delimited by a carina only laterally. Apidae Crabronidae the other Apoidea. In Clystopsenella and Eusapyga, the me- 40. Pronotum (internally): dial portion is not strongly pointed, but they are assigned (0) without lateral ridges. state (0). (1) with a pair of lateral, oblique ridges (converging of anteriorly). 45. Apophyseal arms prothoracic endosternum: (0) 48 and 49). Ammoplanus and Hesperapis have only a pair of weak separate (Figs. (1) fused a 50) carinae (very short in Hesperapis); both are coded as (1). In (forming bridge) (Fig. the some taxa, e.g., Astata, Ctenocolletes and Stangeella, 46. Bases of apophyseal arms of prothoracic endosternum in of ridges are continuous the middle. The male (internally): but the female of A. Aphelotoma rufiventris has no ridges, (0) not connected by divergent plates (Fig. 48). in where nigricula has a distinct external sulcus the place (1 ) connected by two continuous, divergent plates origi- the ridge is situated; this taxon is assigned state (1). nating at base of furcasternum (broadening dor- are also based Laphyragogus and Xenosphex assigned (1) sally) (Fig. 49). on external examination (presence of a sulcus in only In Palarus, these plates are very broad and close half and a line in Laphyragogus). Xenosphex of the coxal cavity. In Didineis and Ochleroptera, the plates

41 . Outer ventral posterior corner of prothoracic episternum: are apparently absent, but I assume that the plates fused (0) not differentiated from rest of episternum. completely to the furcasternum (the medial line in the (1) more or less protuberant in lateral view, lateral ca- furcasternum is absent, contrary to what occurs in groups rina of episternum not differentiated. where the plates are originally absent). (2) as (1), but lateral carina of above the episternum, 47. Internal divergent plates of prothoracic endosternum: a distinct lamella. protuberance, forming (0) separate from furcasternum by a medial ridge. 42. Short sulcus dorsal to outer ventral posterior corner of (1) partially (dorsally) or completely fused to prothoracic episternum: furcasternum, medial ridge absent at least dorsally (0) absent. (Fig. 50) (1) 47). present (Fig. This character applies only to taxa with state (1) for A correspondingly short segment of the anterior mar- the preceding character. this it does gin of the pronotum fits inside sulcus, although 48. Fore basitarsus: not show modification in relation to the re- any particular (0) apex not modified, or only with a short apical lobe. mainder of the anterior This sulcus is margin. present only (1) with a distinct apical lobe, extending at least to half in Heliocausits and Ochleroptera. the length of second tarsomere. 43. Prothoracic basisternum: The condition described in state (1) is found only in (0) lateral segments of posterior edge oblique, converg- females of Laphyragogus and in most species of ing in the middle. Eremiasphecium [see Figs. 118-123 in Marshakov (1976)]. for small (1) posterior edge basically straight, except 49. Foretarsal rake (females): medial lateral corners projection, pointed. (0) absent. (2) as (1), but lateral corners rounded (basisternum very (1) present, bristles longer than diameter of basitarsus. small). of (2) present, bristles as long as or shorter than diameter This structure shows considerable variation among the basitarsus. taxa examined, making difficult the delimitation of dis- 50. Socket of foreleg spur: crete characters and states. This character is used to recog- (0) broadly connected to basitarsal socket. nize what seems to be a distinct morphology found in the (1) narrowly connected to basitarsal socket and away Bembicinae examined. from tibial apex. (2) as (1) or even farther from tibial and, and spur 44. Medial portion of prothoracic basisternum: apex socket almost or closed 51). (0) at same level as rest of basisternum, strongly pointed completely (Fig. the socket is some- posteriorly (Fig. 46). In Lyroda and Laphyragogus, spur (1) declivous in relation to anterior portion (sometimes what closed, but it is situated near the tibial apex. Both or only slightly), rounded weakly pointed posteri- taxa are coded (0). orly. 51. Leg form of female: The comment for character 43 also applies here; how- (0) all similar, slender and generalized (or modified dif- ever, the present character is being used to represent the ferent! v from state 1). 16 Scientific Papers, Natural History Museum, The University of Kansas

(1) all femora inflated and fusiform although midfemur only along its dorsal half; the depressed anterior margin often less so; tibiae and tarsi fairly slender. of the mesepisternum would be a modification of the itself in to a 52. Claws: mesepisternum response modified pronotum (ventral mesad). The (0) with a subapical or at least one subbasal tooth. angles greatly produced prepectus in is narrow, but it has a distinct fovea (1) simple, without subapical or subbasal teeth. Rlwpalosoma very also present in Ensapx/ga and Sierolomorpha. 53. Notauli: 58. (0) indicated externally by a sulcus and internally by a Mesepisternal ridge: (0) anterior of ridge. complete, reaching edge mesepisternum from midline. (1) indicated externally by a line and internally by a away body's (1) midline ridge. complete, reaching body's ventrally. (2) ventral of but ab- (2) indicated only externally by a sulcus. reaching portion mesepisternum, sent from middle of (3) vestigial. mesepisternum. (3) restricted to ventral of (4) no indication (absent). portion mesepisternum (ab- sent laterally). Laphyragogus and Xenosphex are assigned (1) based on (4) restricted to lateral portion of mesepisternum (ab- external examination only. sent ventrally). 54. carina: Supra-alar (5) absent. (0) absent or if present, not meeting tegular ridge This is an internal ridge present laterally and/or ven- (preaxilla open anteriorly; Fig. 25). trally on the mesepisternum (Fig. 52). In most cases it is (1) curving down anteriorly and fused to the anterior marked externally by a sulcus (see next character). In segment of the tegular ridge (preaxilla closed off Dineius and Lyroda, the ridge is interrupted ventrally and anteriorly; Fig. 26). only a small segment is present along the anterior edge of The terms carina and were supra-alar tegular ridge the mesepisternum; both taxa are coded as (0). In taken from Michener (1944). This carina has also been Bembecinus, the mesepisternal ridge is vestigial since only named scutal Menke (1988). flange by a very short segment is present below the subalar fossa; 55. Setal patch on anterior segment of tegular ridge: however, it is still coded (4). (0) present. 59. Mesepisternal sulcus: absent. (1) (0) complete, reaching anterior edge of mesepisternum This setal patch seems to be a proprioreceptor field away from body's midline.

I midline and is present in all aculeate outgroup taxa studied. It is (1) complete, reaching body's ventrally (Fig. 53). absent from a braconid, evaniid, trigonalyid and a (2) restricted to ventral portion of mesepisternum (ab- xiphydriid examined. Judging from Ronquist and sent laterally). Nordlander (1989), it is also apparently absent from (3) restricted to lateral portion of mesepisternum (ab- . This might be a synapomorphy for Aculeata. sent ventrally). (4) absent. 56. Oblique scutal carina: (0) absent. Sometimes the sulcus is only weakly indicated ven- In (1) present. trally. Ctenocolletes, the sulcus is only weakly indicated on the upper part of the mesepisternum; it is (3). 57. Prepectus: assigned 60. Omaular sulcus: (0) not immovably fused to the mesepisternum. absent. (1) immovably fused to mesepisternum, suture be- (0) tween them not obliterated. (1) present (Fig. 54). 61. Omaular carina: (2) as (1), but suture completely obliterated. (0) absent. I recognized only three states for the taxa included (1) present. here, taking into consideration only the degree of fusion to the mesepisternum. Brothers (1975) assumed that the This carina corresponds to the "omaulus" of Bohart Menke prepectus in Apoidea extended completely across the an- and (1976). Omaulus here is used to designate the area of the terior margin of the mesepisternum, being fused in the mesepisternum where its anterior and lateral surfaces meet. When both a carina midline, as well as fused to and forming the depressed and a sulcus are present, the carina is anterior to the sulcus anterior margin of the mesepisternum. Considering the always (Fig. 54). condition in basal Chrysidoidea and Vespoidea, an alter- 62. Interfurcal muscle: native interpretation for the Apoidea would be a prepectus (0) present. not contiguous medially and fused to the mesepisternum (1) absent. Major Lineages of Apoidea 17

of this is Absence muscle considered a synapomorphy (1) present (Fig. 18). for Apoidea by Heraty et al. (1994). a male of Only 66. Mesepisternum and metepisternum: Sierolomorpha canadensis and of Dolichurus sp. were exam- (0) not fused laterally. ined using the technique described by these authors. The (1) fused laterally; suture mostly obliterated. remaining taxa were assumed to have the groundplan con- In most taxa, this fusion is restricted to the lower lat- dition postulated by Heraty et al. (1994). The loss of this eral parts of the mesepisternum and metepisternum. muscle is probably correlated with the ventral fusion of 67. Medial the meso- and metathoraces in Apoidea (condition not used portion of mesometepisternal suture (between here as an independent character). midcoxae): (0) visible 59). 63. Arms of meso- and metathoracic furca: clearly (Fig. (1) obliterated 23, 24, 60-62). (0) fused. mostly (Figs. In all the and (1) not fused or only weakly fused (separate in KOH Apoidea, mesepisternum metepisternum are fused The of this area is cleared specimens; Figs. 15 and 16). ventrally. morphology very complex and variable, making character delimitation dif- Together with loss of the interfurcal muscle, Heraty et ficult. Brothers and (1993) consider loss of al. (1994) considered fusion of the arms of the meso- and Carpenter any sulcus between meso- and metepisterna, ventrally, as part metathoracic furca a synapomorphy for the Apoidea. In of the Apoidea groundplan. However, in Ampulicidae and two apoid groups, however, the arms are not immovably Heterogyna, the suture is clearly visible (the two lateral fused and become separated after KOH treatment. This halves converge forward) and the midcoxal sockets are condition was found in Ampulicidae and in the crabronid small and widely separated; also the mesal articulation is genera Astata, Eremiasphecium and Pulverro (as well as closer to the lateral articulation than to the body's mid- Ammoplanops). In the Ampulicidae, this probably repre- line. The coxal sockets are large and the suture is mostly sents a plesiomorphic condition, while in those crabronid obliterated only in the remaining apoids. The expansion genera, it is clearly a secondary derived condition. In the of the coxal sockets was apparently accompanied by a cleared specimen of Astata nevadica, the metafurcal arm posterior expansion of the mesokatepisternum, forming a has a cup-like expansion and the mesofurcal arm has a broad flap covering the sockets medially (Fig. 23); concomi- callus-like structure (finely fibrous) in the region where tantly, the suture is directed posteriorly (in lateral view; they are supposed to be fused (Fig. 15). The lateral arm of Fig. 24). Brothers and Carpenter (1993) also inappropri- the metafurca has also an additional cup-like expansion ately consider this condition as part of the Apoidea on its tip and it is weakly attached to the larger cup-like groundplan (see their Fig. 11 and state 57-2 in their Ap- expansion projecting from the endophragmal (= upper pendix IX). metapleural) apophysis. Dissection of a specimen of Astata and the have a some- sp. (Costa Rica) preserved in alcohol showed that the two Eremiasphecium Ammoplanini what distinct condition. Their is not furcal arms are firmly attached to each other and that the metepisternum pro- in the middle as a keel continuous with the cup-like expansions of the metafurcal arms are covered jected strong (see 62); there is a transverse line that with tendon-like material (finely fibrous and whitish). The mesepisternum Fig. looks like the this line does not KOH treatment probably breaks down this material, caus- mesometepisternal suture; seem to be to the suture and it is a ing the separation of the furcal arms. In the cleared speci- homologous probably structure to these taxa. these mens of Pulverro and Eremiasphecium sahelense, the lateral unique Despite modifications, are coded as state (1). arms of the metafurca are broad (laminar) and also sepa- they having rate the from mesofurcal arms (Fig. 16). Apparently the 68. Medial flap of mesokatepisternum and condyle of me- two cup-like expansions observed in Astata fused together sal midcoxal articulation: and became one large expansion entirely covering each (0) flap well developed and broadly continuous in the lateral arm of the metafurca in these taxa. middle; portion containing condyle not or only 23 and 60). 64. Upper margin of discriminal lamella (segment poste- slightly projecting (Fig. rior to furcal arms): (1) flap well developed, interrupted in the middle by a close to midline of but (0) narrow, as broad as remainder of lamella. deep cleft; condyle body, not situated at of (1) expanded, forming a horizontal lamella perpendicu- apex flap projection. (2) well in the middle a lar to vertical portion. flap developed, interrupted by deep cleft; condyle situated at apex of flap projec- This character applies only to taxa assigned state (1) tion and close to midline of 61 and 62). for character 67. body (Fig. (3) flap narrow, interrupted in the middle by a deep 65. of second Pseudophragma phragma: cleft; condyle situated at apex of flap projection and (0) absent (Fig. 17). well separated from midline of body (Fig. 53). 18 Scientific Papers, Natural History Museum, The University of Kansas

This character applies only to taxa assigned state (1) tachment of the membrane connecting the base of the coxa for the preceding character. This region probably repre- to its thoracic socket. It is assumed here that a basicoxite is sents the fusion of the true katepisternum with the tro- absent from the Apoidea and that the mesocoxal carina, chantin [see discussion in Gibson (1993)], since it seems therefore, does not mark the limit between the basicoxite more parsimonius to assume that the mesal articulation in and a disticoxite. the is to the trochantinal ar- Hymenoptera homologous 71. Mesocoxal carina 2: ticulation of other insects, and not a new articulation. (0) weak, sometimes restricted to upper half or indi- 69. Medial of portion metepistenum: cated only by a ridge (Fig. 55). (0) a keel and fused well or less narrow, forming strong narrowly (1) defined, more uniform throughout (Fig. to medial portion of mesokatepisternum (anterior 56). portion of keel extending through vertical medial (2) conspicuously enlarged, becoming a lamella toward portion of mesokatepisternum) (Fig. 59-61). lower half (Fig. 57). (1) narrow, a carina, to vertical forming perpendicular This character applies only to taxa assigned state (1) medial of (anterior portion mesokatepisternum in the preceding character. portion of carina not extending through vertical 72. Basal part of mesocoxa: medial portion of mesokatepisternum) (Fig. 62). or less (0) more continuous with rest of coxa (Fig. 55-57). (2) narrow, but not forming a strong keel or carina, nar- (1) forming a narrow pedicel (coxa pedunculate) 58). rowly fused to mesokatepisternum. (Fig. 73. Number of mid tibial (3) wide and flat, broadly fused to mesokatepisternum spurs: two. (mesometepisternal suture transverse or V-shaped (0) (1) one. in ventral view) (Fig. 23). (4) wide and flat, broadly fused to mesokatepisternum 74. Hind coxal socket: (mesometepisternal suture indistinct) (Fig. 53). (0) closed by membrane only. (1) closed a narrow sclerotized This character applies only to taxa assigned state (1) by bridge connecting for Character 62 (ventral fusion of meso- and metathora- the propodeum to the metakatepisternum. (2) closed mos an of the medial ces). Xenosphex is assigned (3) despite the fact that the by enlargement por- tion of the r" medial portion of its metepisternum is mostly vertical. "'-katepisternum (Fig. 23).

70. Mesocoxal carina 1: 75. Socket of mesal articulation on hindcoxa:

(0) absent. (0) away from ventral surface of hindcoxa (distance to ventral surface at least one half of distance between (1) present (Fig. 55-57). socket of mesal articulation and condyle of lateral This carina runs from the lateral articulation obliquely articulation). across the coxa to the ventral articulation with the tro- (1) on ventral surface of hindcoxa or very close to it chanter posteriorly (Michener 1981). In some taxa, for ex- (Fig. 27). ample Aphilanthops and Philanthus, it is present only ba- 76. Hind coxal carina: sally on the rest of the coxa being indicated by a rounded (0) absent. to 1 ridge; contrary Alexander ( 992a), I assigned state ( 1 ) to (1) well these taxa (but see next character). present, developed but not forming a lamella. (2) present, very strong, forming a lamella (Fig. 63). Michener (1981) assumed, without further argumen- This hind coxal lamella is in the tation, that the basal groove on the coxa of the Apoidea present Ammoplanini. In several taxa in this tribe, the lamella is anteri- represents the separation of the basicoxite from the rest of stronger and sometimes absent the coxa (Michener's disticoxite) and that the mesocoxal orly posteriorly, forming a spine- like carina is present only in those taxa whose basal groove projection. has been 77. Paired lobes on inner side of hind coxal displaced distally, i.e. taxa in which an enlarge- apex: ment of the "basicoxite" and a correlated reduction of the (0) subequal in size and separated by a narrow cleft. dorsal lobe "disticoxite" occurred. Michener's terminology and ho- (1) large, usually forming a spatulate pro- ventral lobe small or absent mology assessments were also adopted by Johnson (1988) cess, (Fig. 64). for the rest of The examined here Hymenoptera. Xyelidae Eretniasphecium and the Ammoplanini are assigned has a distinct basal suture, to the state the fact that their certainly homologous (1) despite coxal apices are practi- basicostal suture of other insects as defined by Snodgrass cally straight, without any lobes (see Fig. 63). In (1993), a short basicoxite. delimiting very The other Hy- Ochleroptera, the lobes are subequal in size, but the dorsal examined have no such basicoxite and the one is it is menoptera spatulate; assigned state (1). In Heterogyna, basicostal suture seems to to correspond the region of at- Clystopsenella and Sierolonwrpha, the lobes are very small Major Lineages of Apoidea 19

thickened in and separated by a shallow notch in the female and prac- (1) dorso-ventrally (Figs. 72-74), dry speci- state mens con- tically nil in the male; these taxa are assigned (0). postero-medial portion conspicuously vex ventrally. 78. Dorsal apical process on inner surface of hindcoxa: thickened is (0) relatively small, trochanter without any conspicu- A pterostigma present in members of the ous depression. subtribes Pemphredonina and Stigmina of the In (1) well developed, articulating with a basal depres- Pemphredonini. Stigmas, the pterostigma is greatly en- sion on inner surface of trochanter, apical edge of larged and both dorsal and ventral surfaces possess a dis- tinct field 68-71 these there depression not delimited by a weak crest (Fig. 64). micropore (Figs. ); under fields, (2) as (1) or even more developed, apical edge of is a thick glandular epidermis (Figs. 72 and 73). This gland trochanterical depression delimited by a weak crest. is probably present in all Stigmina, because most of them also have a distinct field on the (3) nil (Fig. 63). micropore pterostigma. Also a somewhat diffuse field is in This character applies only for taxa assigned state (1) micropore present Diodontus 75). The of Passaloecus is simi- in the preceding character. These modifications of the coxa (Fig. pterostigma swollen, but no tissue was detected 74). and trochanter are probably related to the truncation of larly glandular (Fig. the femoral apex, since they are more developed in taxa 84. Width of forewing costal cell: with such modified femora (see next character), like (0) at least as wide as width of vein C.

Entomosericus and Odontosphex (also Bothynostethus, (1 ) linear, narrower than vein C. and a few other crabronid taxa). These Cerceris, Oxybelus Maximum width measured perpendicular to costal structures allow the femur to be locked in one probably margin of wing. Dinctus was assigned state (0), but its con- position. dition is somewhat intermediate between the two states. 79. Apex of hind femur (females): Lapln/ragogus is assigned (?) because its vein C is unusu- (0) unmodified. ally slender. truncate. (1) broadened, 85. Pterostigma width: with an (2) apical spatulate process. (0) less than or subequal to length of prestigma (Sc + R 80. Basitibial plate (females): distal to Rs). (0) absent. (1) at least one and a third the length of prestigma. (1) present. Maximum width measured perpendicular to costal 81. Hind tibial bristles: wing margin. (0) present (at least one). 86. Marginal cell: (1) absent. (0) longer than pterostigma (measured along vein C) 19). These bristles correspond to relatively large and (Fig. (1) shorter than 20-22). spiniform setae, in contrast to the fine and usually shorter pterostigma (Figs. a cell than setae also present on the hind tibia. Despite the large and Some species of Astata have marginal longer have state plumose setae on the hind tibia of bees, bristles are con- the pterostigma, but the remaining Astatini (1), that Astata does not sidered absent from this taxa; their plumose setae prob- including A. nevadica. Considering seem to be the most basal within the tribe, the rela- ably correspond only to enlargedTine setae. lineage tively long marginal cell in some of its species might rep- 82. Gland on posterior surface of hind tibia: resent a derived condition. A positive correlation between (0) absent. body size and relative length of the wing cells has been (1) 65-67). present (Figs. documented for the Hymenoptera (Danforth 1989). Since of this was inferred from it is The presence gland initially Astata contains the largest forms in the tribe, expected field taken from Finnamore the distinct micropore [term that they have longer cells as well. (1995)] on the surface of the tibia (Figs. 65 and posterior 87. Segment of Rs separating 1st and 2nd submarginal cells: 66). Its was confirmed with histological section- presence (0) present. for 67). This is ing only Ammoplanus (Fig. gland possibly (1) absent. absent from Timberlakena, because there is no micropore The characters of venation for Eremiasphcciuin for a transverse darker area in forewing field on its hindtibia, except be- were taken from £. hmgicepis and not from £. sahelense, the region where the field is situated in the other cause this latter species has some reductions that are prob- It is assigned (?), because a histological Ammoplanini. ably not part of the groundplan for the genus [see also Fig. reveal the of the study might presence gland. 184G in Bohart and Menke (1976) for the wing venation of 83. Pterostigma: £. schmiedeknechtii Kohl]. For Hetcrogyna, wing characters (0) flat, not conspicuously thickened. taken from male of H. protea. 20 Scientific Papers, Natural History Museum, The University of Kansas

88. Forewing 2rs-m: Nitela and Timberlakena are assigned (?) because their venation is reduced and of the veins (0) present. hindwing very parts (1) absent. involved are lacking. 2A: Present only as nebulous vein in Heterogyna. 98. Hindwing vein (0) indicated at least as a short on basal 89. Forewing M (distal to 2rs-m) and 3rs-m: spur portion of 1A. (0) present. (1) absent. (1) absent. 99. Hindwing clavus (= plical lobe): Present only as spectral veins in Sierolomorpha. (0) indicated posterodistally by moderate incision. 90. CuAl and 2m-cu: Forewing notch. (1 ) indicated by short incision or only a shallow (0) present. (2) not indicated posterodistally on wing margin. (1) absent (discal cell 2 absent). 100. Jugal lobe: Present as spectral veins in Sierolomorpha. only (0) absent. Rs to 2r-rs) and 2rs-m: 91. Forewing (anterior (1) small to moderately long and indicated by dis- (0) separated by Rs (segment distal to 2r-rs) anteriorly tinct incision. 19 and 21). (Fig. (2) large and not indicated by an incision on wing 2nd cell (1) touching anteriorly (i.e., submarginal margin (fused to clavus). pointed anteriorly). 101. Metapostnotum I: (2) fused anteriorly (i.e., 2nd submarginal cell petiolate; (0) transverse, depressed and distinct mesally be- 20 and 22). Figs. tween metanotum and propodeum (or shortened). and Anacrabro are (?) for this Nitela, Lindenius assigned (1) strongly expanded posteromesally to form character because some of the veins involved are absent, "propodeal triangle." but there is good evidence that their reduced venation 102. II (propodeal enclosure): with a 2nd sub- Metapostnotum pattern is derived from lineages petiolate (0) restricted to dorsal surface of propodeum, apex marginal cell. rounded. 92. M and CuA: Forewing (1 ) extending as a narrow triangle onto posterior sur- distal to cu-a. (0) diverging face of propodeum, but for less than half the length at cu-a. (1) diverging of the posterior surface. basal to cu-a. (2) diverging (2) extending as a narrow triangle for more than half 93. Forewing M + CuA (distal to cu-a): the length of the posterior surface, but not reach- (0) subequal to or shorther than cu-a (Fig. 21 ). ing posterior apex of propodeum. 19 as a narrow to (1) longer than cu-a (Fig. and 20). (3) extending triangle posterior apex of This character applies only to taxa assigned states (0) propodeum. or (1) in the preceding character. This character applies only to taxa assigned state (1) in the character. It is used to an 94. Forewing M (basal to Rs): preceding being represent progressive shortening of the and a (0) gently curved or straight (Figs. 19, 20 and 22). apparent propodeum simultaneous relative elongation and posterior narrowing (1) strongly bent (Fig. 21). of the metapostnotum within Apoidea. 95. Forewing vein CuA2: 103. Third mesosomal phragma: (0) present, reaching vein 1A. (0) forming a transverse, vertical flange, continuous (1) much reduced or absent, not reaching vein 1 A. or narrowly interrupted in the middle. 96. C: Hindwing (1) forming only a narrow medial, transverse flap, (0) present. situated at the apex of expanded metapostnotum. (1) absent. (2) as (1), but flap longitudinal. In Heterogyna, Epyris and Clystopsenella, it is not pos- (3) as (1), but forming a spine-like projection. sible to determine if the vein along the costal margin of (4) absent or indistinctly fused to metapostnotum (ex- the hindwing represents only Sc+R or a fusion of C with cept sometimes for presence of a longitudinal ca- Sc+R. They are assigned (?). rina). 97. Hindwing M: 104. Triangular posterior extension of metapostnotum: (0) diverging from CuA before or at cu-a (Fig. 19). (0) flat or forming a broad, shallow sulcus. a (1) diverging from CuA after cu-a. (1) forming narrow, deep sulcus (Fig. 76). Major Lineages of Apoidea 21

112. Adult female silk This character applies only to taxa assigned states (1) glands: to (3) for character 102. (0) absent. (1) present, associated with tergum VI. 105. Metasomal petiole: (2) present, associated with sterna IV and V (0) absent, tergum I and sternum I not immovably See Melo (1997) for more details on the structure, func- fused, suture between them clearly visible. tion and taxonomic distribution of these (1) present, tergum I and sternum I fused anteriorly, glands. portion of tergum I forming petiole very reduced, 113. Female pygidial plate (tergum VI): suture between tergum I and sternum I along peti- (0) absent. ole mostly obliterated. (1) present. I I not scler- (2) present, tergum and sternum fused, 114. Sixth metasomal sternum (females): in suture between them ites subequal size, clearly (0) similar to other segments, except for troughlike visible. vertical side walls. an tube Dolichurus has what looks like a very short petiole and (1) elongate, forming exposed tapering is coded as state (1). through which sting is exserted. 115. of female sternum VI 106. Lateral line on tergum I: Apex medially: as a carina. (0) (truncate, rounded or (0) present, sometimes marked weak simple slightly emarginate), or less continuous with lateral of (1) absent. more portions the apex. 107. Lateral carina on base of tergum I (dorsal to lateral (1) forming a medial lobe (Fig. 79). line): (2) with two pointed projections separated by a deep (0) absent. V-emargination (Fig. 80). (1) present, basal portion simple. (3) denticulate (Fig. 81). (2) present, basal portion protuberant. 116. Seventh metasomal tergum of female I: 108. Medial longitudinal ridge on base of sternum I: (0) partly exposed and evenly sclerotized. (0) absent. (1) hidden under tergum VI and considerably (1) more basally present, developed desclerotized. 109. Second sternum posteromedially: 117. Seventh metasomal tergum of female II: or less (0) slightly convex to flat, basal portion more (0) two broad, lateral plates connected anteriorly by at same level as remainder of sclerite. a sclerotized bridge. (1) strongly convex, basal portion distinctly in a dif- (1 ) as (0), but bridge displaced toward posterior mar- ferent level in relation to remainder of sclerite, gin of tergum. surface separating these two portions almost ver- (2) lateral plates narrow (not sclerotized dorsad to spi- tical 28). (Fig. racles), connected by a narrow, but strongly scle- is used to the distinct condi- This character recognize rotized bridge. Palarus and males of tion found in the Ampulicidae. (3) as (2), but lateral segments of bridge forming a II modified transverse and Heliocaimis have sternum (with 90° angle with dorsal segment. from the condition found in thick keels), but differently (4) forming two separate lateral plates (hemitergites), two taxa are state (0) to the Ampulicidae. These assigned connected by membrane only. avoid states for them. creating autapomorphic This character applies only to taxa assigned state (1) 110. Basal portion of lateral gradulus of sternum II. in the preceding character. not modified, or absent. (0) simple, gradulus 118. Female hemitergites VIII: laminar and directed inward, a (1) forming special- (0) narrowly connected dorsally by a sclerotized ized surface with the differentiated articulating bridge. of the lateral of sternum I posterior portion edge (1) connected by membrane only. 77). (Fig. 119. Female gonapophyses VIII in lateral view: on sternum II: 111. Anterior lateral epidermal gland (0) strongly curving downward. to lateral (0) situated mesal gradulus. (1) gently curving downward. situated lateral to lateral 78). (1) gradulus (Fig. (2) straight. Taxa in which the lateral gradulus is absent, as well as (3) curving upward. coded is those in which the gland could not be detected, were Ctenocolktes is assigned (?) because its sting very their as (?). reduced (gonapophyses widely separated at bases). 22 Scientific Papers. Natural History Museum, The University of Kansas

120. Articulation within gonocoxite IX of female: 131. Larval integument: (0) absent. (0) with minute spicules or smooth. (1) present. (1) with dense, short spicules. (2) with seta-like acanthae. 121. Male tergum VII: dense, conspicuous (0) entire. In the descriptions of the larva of Mimesa bicolor, Janvier (1) with lateral lobes. (1956) makes no reference to the integument; Mimesa was coded as (?). 122. Male cerci (tergum VIII): 132. Position of larval anus: (0) present. (1) absent. (0) terminal, directed caudad. (1) ventral, directed ventrad. 123. Male sternum VII: preapical, 133. Larval antennal (0) partly exposed. papillae: (0) absent (sometimes the orbits are but (1) completely hidden under sternum VI. protuberant, there are no 124. Apex of male sternum VIII: papillae). (1) present, usually well developed and conspicuous. (0) continuous with disk or forming a broad and short 134. Larval maxillae: medial lobe (broader than or as broad as long). (0) directed mesad associated with (1) forming a medial lobe or projection longer than apically, closely labium and broad, but less than 3x longer than wide, relatively hypopharynx. broad. (1) projecting apically as large, free lobes. 135. Larval (2) medial projection more than 3x longer than wide, galea: broad, sides diverging distally. (0) large, subequal in size to maxillary palpus. (1) less than half the size of (3) as (2), but narrow and sides parallel or converg- small, maxillary palpus. (2) absent. ing distally. (4) with two long, lateral spiniform projections. 136. Larval spinneret: (5) forming three long, spiniform projections. (0) a transverse slit.

(1 ) with each at the end of a 125. Lateral margin of projection of male sternum VIII: paired openings, projec- tion. (0) entire. (2) absent. (1) serrate (Fig. 82). (2) with short, thick bristles. 137. Provisions for larvae: (Blattodea, Phas- 126. Apical margin of male sternum VIII: (O)Orthopteroids Mantodea, matodea and (0) entire. Orthoptera). (1) (1) denticulate. Thysanoptera. (2) (Heteroptera and Homoptera) 127. Posterior edge of gonobase foramen (ventrally): (3) immature Holometabola. (0) close to bases of gonocoxites. (4) adult Holometabola. (1) widely separated from bases of gonocoxites. (5) pollen. 128. Volsellae: (6) Araneae. (0) clearly differentiated from gonocoxites. Besides preying on aphids, Nitela is also known to prey (1) fused to small or largely gonocoxites, usually on Psocoptera, but this state was not included because, sometimes apparently absent. among the exemplar taxa, it would be present only in this 129. Gonapophyses of male genitalia dorsally: genus. Liiidenius and Arpactopliilus are assigned more than (0) connected by membrane only or by a sclerotized one state. Ni/sson and Eusapn/ga are coded (?) since they are bridge, but not forming a distinct tube. cleptoparasites. (1) completely fused, forming a tube. 138. Relocation of larval food: (0) absent. Various forms of sclerotized bridges are found among the taxa (1) analyzed, making difficult the recognition of dis- present. crete states. is Chlorion is This character used to recognize the distinct assigned both states. Epi/ris females are condition found among several Crabroninae. known to relocate their prey, but this is an exception for 130. of bethylids; it is assigned (0). Apicoventral edge gonapophyses of male genitalia: (0) without teeth. 139. Construction of a nest before obtaining larval food: (1) with numerous short teeth. (0) absent. Sometimes, these teeth are very inconspicuous, like (1) present. those in Mellimis [see illustrations in Menke (1996)]. Chlorion and Podalonia are assigned both states. Major Lineages of Apoidea 23

RESULTS

Analysis of the complete data matrix resulted in four quency distributions for the character weights determined most fit when 1 cladograms applying implied weighting. by implied weighting-1 tree and by the successive weight- The four differ in the of tree are shown in it cladograms mainly position ing-1 Figure 7; can easily be perceived Heterogyna: three of them have Heterogyna as sister group how stronger use of the consistency index in successive to is shown in 1 whereas the fourth Ampulicidae (one Fig. ), weighting down-weighs the homoplastic characters com- has it as the sister group of the remaining Apoidea, ex- pared to the weighting function employed by implied cluding Ampulicidae (Fig. 2). A strict consensus tree for weighting. these four cladograms is shown in Figure 3. Under equal Table 5 presents information on length, fitness (sensu weighting, 55 most parsimonious trees were found (one is Goloboff) and some statistics (consistency, rescaled con- shown in Fig. 5a), and a strict consensus tree for the 55 sistency and retention indices) for the resulting trees (see trees is shown in Figure 4a. Successive weighting produced also Figs. 1, 2, 5 and 6). Unweighted length for trees pro- only one most parsimonious tree 6). This tree is simi- (Fig. duced under implied and successive weighting was ob- lar to the trees the implied weighting regarding position tained from Nona; total fitness for trees produced under of the main lineages. equal and successive weighting were obtained by import- Under implied weighting, removal of Lapln/ragogus ing the trees produced by Nona and PAUP* into Pee-Wee and from the data matrix resulted in one and then the Xenosphex only executing command fit. The total fitness for tree (not shown), which was identical to the tree in Figure the 55 trees produced under equal weighting varies from 1 regarding the composition and relationships among the 695.5 to 697.0. main lineages. Analysis of the partial data matrix under Only unambiguous changes were plotted in the cla- trees (one is shown in equal weighting produced eight Fig. shown in 1 5 6. dograms Figures , 2, and Some of the am- 5b), and a strict consensus tree for the trees is shown eight biguous changes could have been resolved on a one to one in 4b. Under successive of the Figure weighting, analysis basis, especially for characters with a low number of alter- data matrix resulted in one most partial only parsimoni- native optimizations, but for several characters the num- ous tree, which is identical to one of the trees produced ber of possible optimizations is so numerous that choos- under equal weighting (Fig. 5b). ing any of them would be extremely arbitrary. Also, no The character the weights implied by topologies of restrictions in relation to character irreversibility were implied weighting-1 tree 1 and implied weighting-2 tree, imposed; however, changes in a few characters, as for ex- and the final weights for succesive weighting-1 and suc- ample in Characters 73 and 122, probably should have been cessive weighting-2 trees are listed in Table 4. The fre- optimized as irreversible. DISCUSSION

Choice of Analytical Method differentially weighted before reaching the stage of data Considering that the different parsimony methods analysis, although this is not always realized. The process produced conflicting arrangements involving the major of character selection, for example, is a form of attributing lineages being investigated, one could manifest no prefer- different weights to the potential characters, the ones se- ence for any of the specific results and then make use of lected receiving weight one and the ones left out receiving only those components common to the different arrange- weight zero. Not considering cases where potential char- ments. Nevertheless, I prefer to argue in favor of using acters are overlooked, what is assumed to be random varia- one of the parsimony methods over the others and then tion among the exemplar taxa is usually neglected as a make use of all components present in the resulting tree source of informative characters. The problem of selecting (or set of trees) produced by application of this method. genes showing an amount of sequence divergence "ap- for taxa in a The major contrast among the methods used here in- propriate" resolving relationships among molecular data Simon et volves use or not of differential weights for the characters study using (e.g., Graybeal 1994, al. 1994) is another of to the during tree search. Character weighting has always been example weighting equivalent initial differential selection of characters. a controversial issue (e.g., Kluge 1998), despite the frequent morphological assumption that characters are not all equally informative The main reason why characters are assumed to be et al. Goloboff (e.g., Farris 1983, Swofford 1996, 1993). Ex- uneven regarding their phylogenetic informational con- plicit a priori weighting of morphological characters has tent is because they can evolve at different rates. In a study been strongly criticized because of its intrinsically subjec- involving a large number of taxa, the characters selected tive and arbitrary nature; however, characters are usually will certainly have changed at different rates during the 24 Scientific Papers, Natural History Museum, The University of Kansas

* snuiiiairtj 7 7 7 7 7 ; t f 7 7 7 7

X3L]d$0U8X-+-*.

smeied 7 7 i 7 . 7

9 JjSSS"7 "-"7 ^

epojAn 77?T 7777' t

-' ®

' 3 33 o ^ 5 2 "^ 3 r ' " eiaiiN T77 i T i 7 7 T 7 7 7

snjuepun 7^7^1 ® ©

sn6o6ejALjde-)-

>

;.-i: ,[! , _e- sniauig T7 7"'

eiuoiepod 7777777 snqiue|iu|d-^

© sdOLjiueiiqdv © m ejaidojamoQ-

snsneooi|an 777

r ^v99*"? t?Ti , ,OrSai

'. - : 1

14-1 17-1 33-1 46-0 49-0 50-1 56-0 58-5 60-1 70-0 79-2 96-1 99-2 102-0 127-1 Major Lineages ofApoidea 25

euA6oja)8H V 7 7/ £-5

eiaiiN 7 7 V V" ? ? 7 7 7 7 T T T 7

sniuapun 7 7T?f?7' i '77T7

epojA") 777777 26 Scientific Papers, Natural History Museum, The University of Kansas evolutionary history of the clades involved, and therefore CEpyris will be appropriate to resolve relationships at distinct hi- Clystopsenella Rhopalosoma erarchical levels. In this case, characters should not com- Sierolomorpha basis in a unless pete on an equal parsimony analysis, Eusapyga someone had a way to guarantee that the characters used Notocyphus were always represented proportionally to the weights Heterogyna Aphelotoma deserve. But, as this cannot be done, and because there they Ampulex is a of no way to know priori what the rates change for tE Dolichurus these characters are, one has to rely on other types of in- Conanthalictus formation to formulate a weighting scheme. Lonchopria Ctenocolletes in The amount of homoplasy exhibited by a character Anthophorula a given topology has been suggested as the only type of Hesperapis Mellinus information that can defensibly be considered an appro- Dinetus criterion for characters priate weighting differentially Xenosphex (Goloboff 1993). As already explained above, the two Laphyragogus weighting methods employed here, implied and succes- Palarus Lyroda sive weighting, use the degree of homoplasy of the char- Plenoculus acters in their functions. fundamental dif- weighting One Anacrabro ference between implied and successive weighting is that CLindenius in the former method calculation of the appropriate Nitela Astata weights is carried out simultaneously with search for the Eremiasphecium most trees, whereas in the latter the parsimonious method, Pulverro weights are calculated after each search run, in an itera- Ammoplanus tive process. This reliance on tree topologies obtained pre- Timberlakena Odontosphex viously to weight computation, in particular for the start- Entomosericus makes successive less desirable as a ing point, weighting Pluto weighting method when compared to implied weighting Mimesa (Goloboff 1993). The influence of the starting trees is well Psenulus illustrated here by the discrepant results produced by suc- CArpactophilusSpilomena cessive weighting for the complete and partial

(Laphyragogus and Xenosplwx excluded) data matrices (com- CParastigmusStigmus pare Figs. 5b and 6). Diodontus Passaloecus One associated with both and suc- problem implied Pemphredon cessive weighting is that each character is assigned a fixed Nysson weight to be applied for any type of change within that Bembecinus character Didmeis (see e.g., Horovitz and Meyer 1995), as well as Hoplisoides for all sections of a given topology. If different types of changes within the same character occur at different rates CHeliocaususOchleroptera or if the rates vary among the different clades being CAphilanthopsPhilanthus sampled, then these weighting procedures will be insensi- Chlonon tive to the distinct sources of This potential homoplasy. Stangeella type of problem is probably less relevant for studies using Palmodes morphological characters than for those using molecular Podalonia data, for example DNA sequence information. Fig. 3. Strict consensus tree of the four most fit trees produced Taking into consideration the issues discussed above, under implied weighting {complete data matrix). the results obtained under implied weighting (see Figs. 1- are 3) being favored over the results of the other methods relationships among the clades given family status here here employed (equal and successive weighting; see Figs. are shown in Figure 8. 4-6). The tree shown in Figure 1 will be used throughout Apoidea and its basal clades the remainder of the present work as a basis to discuss the phylogenetic relationships among the taxa involved, and The monophyly of the Apoidea, under its present com- also to propose the classification being adopted here. The position, was already demonstrated by Brothers and Car- Major Lineages of Apoidea 27

Epyris -c Epyris Clystopsenella -cClystopsenella Eusapyga Eusapyga Sierolomorpha cSierolomorpha Notocyphus Notocyphus -c Rhopalosoma c Rhopalosoma -Heterogyna Heterogyna -Mellinus Mellinus Astata Bembecinus Eremiasphecium Hophsoides Pulverro Didineis Ammoplanus Nysson Timberlakena c; ^ Heliocausus Bembecinus Ochleroptera Hophsoides Astata Didineis Eremiasphecium Nysson Heltocausus Pulverro

Ochleroptera Ammoplanus Timberlakena Arpactophilus -dSpilomena Arpactophilus Diodontus c Spilomena Passaloecus Diodontus Pemphredon Passaloecus Parastigmus Pemphredon cStigmus Parastigmus Anthophorula -c Stigmus Hesperapis Dinetus Conanthalictus Palarus Ctenocolletes Plenoculus c— Lonchopria Lyroda Laphyragogus Nitela — Palarus Anacrabro — Dinetus Lindenius — Xenosphex Anthophorula — Plenoculus Hesperapis — Lyroda Ctenocolletes — Nitela Conanthalictus

Lonchopria CAnacrabroLindenius Odontosphex — Odontosphex Entomosencus Entomosencus Pluto Pluto Mimesa Mimesa Psenulus Psenulus Aphilanthops Philanthus CAphilanthopsPhilanthus c Chlonon Chlonon I -\ Stangeella Stangeella —| Palmodes — Palmodes I Podalonia I— Podalonia Aphelotoma - Dolichurus Ampulex -Ampulex Dohchurus B - Aphelotoma

— tree of the 55 — Fig. 4. A. Strict consensus most parsimonious trees produced under equal weighting (complete data matrix). B. Strict consen- trees sus tree of the eight most parsimonious produced under equal weighting (partial data matrix).

In the seven penter (1993). present work, unambiguous (3) pronotum with a pair of lateral, oblique ridges (40-1 ); changes support the monophyly of the Apoidea (Branch (4) prepectus immovably fused to mesepisternum, su- l,Fig.l): ture between them completely obliterated (57-2); (1) pronotum with posterolateral angle reduced above (5) interfureal muscle absent (62-1); to form spiracular lobe (37-1); (6) metapostnotum expanded posteromesally (2) ventral angle of pronotum considerably produced "propodeal triangle" (101-1); mesad(38-l); (7) larval food relocated (138-1). 28 Scientific Papers, Natural History Museum, The University of Kansas

smnipiioQ 7 7 V 7 T "i

sdomueiigdv- va|nrimy 7 7 7 V f 7 7 .

sn|nuasd-+ "7 BLUO)0|3LjdV-*

omid-

snouasoiuoju3_^?^^:

xagdsoiuopo TTTfTT

ojqejoeuv-

- - - • 7 g t 9 7 9 v o e oi o -" s ' ^ - u

epojX-] 7T?71

xagdsouax-

sniauig f f 7 V V ;

sn6o6ejALjdB-i_

eudoijouoi ,j?f 9^ ,

saianooouaio

snpiiemueuoo 7 V?

sidBjadsai-i 7/7 VTT * i * * » * t

eiruogdoifluv 7 7 V 7 7 i v" H

snuj6|is_ i-*-! T9-79T.S I £«SSS-

sntuGiisejEcjl

u opaj L)d lu aj 7^7777

snoaotBssed-

EU3LUO|ldS_

sniiLjdopBdjy 7?-££

eua>|B|jaquji_i.

9^TTT * * * H QniiP|Hnimny 7 7 ? f f

LunosLjdSBiujajg.

BiBisy ??T'T T-79T999T-rT^^^A

ejaidojamoo 7 7 7 , T 7 7 f'f 7 T 7 7 7

~ . . -: 9 ? t t i A J . i on(MO'-?o-r uossAn TTTTTTTTTTT

r< 9 7 T " ^ i - t

sapouj|ed3

B||336ueis-I- Major Lineages of Apoidea 29

3 O 5, snifluenMd 3Z2SSssss

— s sdou,iue|iu,dv (0

S2£5

co- omid -

snouasouioiu3 -

13

snpnemuGuoQ - c .: saiaiioaouajQ 60 T-T--"H>SSft«»--- sidejads3H

&

sniuapun —— 13 C ojqejoeuv 3

777? e|a ,, N 3 73 - O BpOiA")

- T V lib t£ & S -- Or.

\ i s i £

O "' X snm6iis i "T79T9-:A fc Z. « 2 c sniuSiisejej

sniuopoia—' BO C uopajLjdujScj 3 a - EU3U10|ldS 1 snundopedjv- §•

EU8i|e|jaquJ!j_ -++

snueidoaimv -

3 "tAo JioN - ujnio8L|dseiiu3J3

- *SSg BLuoiO|aL|dv

ejaidojaiqoo -

snsneoot|9H-

uossAn - = £$$8 i '--'litl jjoooAc

S3plOSI|dOH - Vjj:::'

snuioaqiuag - ' :zss — - sapoujied t t t 9 ,y°n

- CO euA6oja)9H 30 Scientific Papers, Natural History Museum, The University of Kansas

eja»doj3|ij30

uossAn

eujoioiaijdv —

srunipnoa

xaindwv-

euaneijaquuij.

" snueidoujiuv »• ?

e aa6u e -£* 1 1 is Major Lineages of Apoidea 31

Table 4 sive weight! ing their qu; 32 Scientific Papers, Natural History Museum, The University of Kansas

Table 4 continued

Character Major Lineages of Apoidea 33

Apidaes.1. Evans (1964a)

- Crabronidae Bembicinae

Philanthinae Sphecidae s.str. Astatinae

Heterogynaidae Pemphredoninae

Mellininae • Ampulicidae Crabroninae

Bohartand Menke (1976)

Apidae s.l. Pemphredoninae

Crabronidae Astatinae

Laphyragoginae Sphecidae s.str. Crabroninae

Heterogynaidae Larrinae

Ampulicidae Entomosericinae

Fig. 8. Cladograms summarizing the relationships among the Philanthinae clades status here. 1 Apoidea given family See Fig. for component taxa. Ampulicidae corresponds to branch 4; Heterogynaidae, branch 5; branch Bembicinae Sphecidae (sensu stricto), 6; Crabronidae, branch 8; and Apidae (sensu lato), branch 9 in Fig. 1. Xenosphecinae (3) mesocoxa subdivided by a broad sulcus into large basicoxite and disticoxite and mesocoxal cavities This — Implied large and approximated or narrowly separated study weighting medially [193-0].

The 1 st and 3rd characters above are not valid because Bembicinae Ampulicidae has basically the same morphology as Philanthinae Heterogyna. The second synapomorphy depends on Heterogyna having a parasitoid life style, with no reloca- tion of is prey; however, nothing known of its biology. Pemphredoninae Brothers and Carpenter (1993) have suggested that because Astatinae of their brachypterous condition, females in this group are unlikely to move their prey from where it is captured and Crabroninae paralyzed. However, considering that Ampulicidae relo- cate their prey on foot, instead of carrying it on flight as Fig. 9. Relationships among the subfamilies of Crabronidae ob- most of the do, it is then that remaining apoids possible tained under implied weighting. Cladograms presented by previous behaves in the Heterogyna same way. Also, Day (1984) authors are also shown for comparison. speculated that the modifications of the female's tergum VI and sting gonocoxites could be adaptations for prey transport. 34 Scientific Papers, Natural History Museum, The University of Kansas

In contrast to Brothers and Carpenter's study, the was described by Antropov and Gorbatovskii (1992). Noth- present analyses did not support placement of ing is known of the biology of these wasps. Large num- Heterogynaidae as the most basal clade in Apoidea, its bers of males have been taken with Malaise traps and one position varying depending on the analyses. In the implied female specimen was collected in a yellow pan trap (Day weighting cladograms, it comes out either as the sister 1984). of or as the sister of the remain- group Ampulicidae group Argaman (1986) proposed a new genus, Daycatinca ing Apoidea, excluding Ampulicidae. (type-species: Heterogyna fantsilotra), for the three Unambiguous changes supporting placement of Madagascan species. Antropov and Gorbatovsky (1992), Heterogynaidae as sister group to Ampulicidae are (branch apparently unaware of Argaman's paper, proposed the 2, Fig. 1): subgenus Daya (type-species: Heterogyna madecassa) for the (1) distance between antennal sockets and clypeus nil four African species. These two genus-group names are (18-2 and 21-2); treated here as synonyms ofHeterog\/na, especially because (2) presence of an omaular carina (61-1); recognition of more than one generic name for such a small (3) forewing M+CuA (distal to cu-a) longer than cu-a group seems completely unnecessary and also because (93-1); proposal of these names was not based on phylogenetic (4) jugal lobe absent (100-0). studies. Recognition of Daycatinca or Daya would prob- make (sensu stricto) or vice- Except perhaps for presence of an omaular carina, the ably Heterogyna paraphyletic versa. characters supporting this relationship are relatively weak. The fourth character, jugal lobe absent, probably does not Ampulicidae represent a synapomorphy for this putative clade, because This (branch 4, 1) has been treated presence of a lobe in Dolichurus (as well as in Trirogma) in group Fig. usually as a of the Ampulicidae can hardly be considered a true reversal. subfamily "sphecid" wasps (e.g., Leclercq 1954, Evans 1964a, Bohart and Menke 1976) or sometimes as a The alternative position of Heterogynaidae as sister separate family of "sphecoid" wasps (e.g., Evans 1959a). group of [Sphecidae (sensu stricto) + [Apidae (sensu lato) In Alexander's (1992a) study, the two tribes of Bohart and + Crabronidae]] is supported by (based on Fig. 2): Menke's classification, Dolichurini and Ampulicini, were (1) labrum at least one and a half times wider than consistently grouped together, forming a monophyletic long (1-0); group. He presented the following putative synapo- (2) arms of the meso- and metathoracic furca immov- morphies for them (see his Table 7): ably fused (63-0); (1) pitted transverse basal sulcus on scutellum; (3) hindwing clavus indicated by moderate incision (99-0). (2) subalar line a very prominent carina or flange; Likewise, the evidence for this placement of Heterogyna (3) posterior margin of 'metasternum' distinctly bi- is rather weak, except for Character (2) above. Even in this lobed, lobes diverging apically; case, however, the fusion of the furcal arms in Heterogyna (4) metasomal sternum 2 swollen at base, with a trans- and in (sensu stricto) + (sensu lato) + Sphecidae [Apidae verse sulcus and/or carina; Crabronidae] might be a parallelism, because loss of the (5) male with fewer than seven visible metasomal seg- interfurcal muscle is (Character 62-1) already part of ments. Apoidea groundplan and the reduced size, as well as fe- Characters (2), (3) and (5) above do not seem to be valid male brachyptery, in Heterogyna could have favored this fusion. synapomorphies for Ampulicidae. In Sphecidae (sensu stricto), the subalar line is also a prominent flange (the outer Heterogynaidae margin of the tegula rests on it when the wings are in re- pose); a weaker or vestigial carina can also be seen in some This family contains only seven described species, all Crabronidae. Also, the of the ventral placed here in the genus Heterogyna (see below). Before morphology portion of the posteriorly (Character 3 above) in Day (1984), only the type species, H. protea Nagy from the metepisternum does not seem to be different Greek island of Rhodes, was known. Day (1984) described Ampulicidae particularly from that of (sensu of some four new species from Africa and provided a key for iden- Sphecidae stricto), Crabronidae, or even of The distinct tifying all five species. He also described for the first time Heterogyna. metepisternal morphol- of is not found in the other ex- the putative females of this genus. The type specimen of ogy Ampnlex Ampulicidae amined. the 5th H. protea was redescribed by Day (1985). Argaman (1986) Regarding character, the condition de- scribed to described a new species (based on a female specimen) from above seems be restricted to males of Ampnlex, Israel and also provided a key for all species known at because males of Dolichurus and Aphelotoma have seven that time. More recently a new species from Turkmenia visible metasomal segments; nevertheless, enlargement of Major Lineages of Apoidea 35

the first three metasomal a distinct reduc- segments and (1) mesepistemal ridge present (58-4); tion of the distal are characteristic of medial remaining segments (2) portion of mesometepisternal suture mostly Ampulicidae. obliterated (67-1); (3) basal of midcoxa In the present work, Ampulicidae always came out as a part continuous with rest of coxa, not monophyletic group; it is supported by the following un- pedunculate (72-0); on (4) females construct a nest before larval food ambiguous changes (based optimizations shown on Fig. 1): obtaining (139-1). (1 ) apophyseal arms of prothoracic endosternum sepa- rate (45-0); Presence of a mesepistemal sulcus (see Character 59) notauli indicated (2) externally by a sulcus (53-0); should also be considered a synapomorphy for this clade. (3) of second pseudophragma phragma absent (65-0); As this character was divided into several states, there is (4) hindwing M diverging from CuA before or at cu-a no unambiguous optimization for the states used at this (97-0); branch. If the character had been coded only as either ab- (5) second sternum convex sent or then strongly posteromedially present, presence of a sulcus would certainly (109-1); be shown as an unambiguous change for this branch. (6) of of male apicoventral edge gonapophyses geni- Someone could argue that the sulcus is not independent talia with short teeth (130-1). from the internal ridge and therefore should not be used The assignment of this condition for the notauli (Char- as a separate character. The only reason why this was done is because the in these two characters acter 2 above) as a synapomorphy for Ampulicidae is prob- changes are not per- correlated (Table 3). Bohart and ably an artifact of having Pompilidae (represented by fectly Menke (1976) con- sidered the Notocyphus) coming out as sister group of Apoidea in the presence of a mesepistemal sulcus as part of the of their because implied weighting analysis. Notaulus indicated externally groundplan Sphecidae, they assumed that a remnant of this sulcus was by a sulcus is plesiomorphic for apocritans; among the sometimes present in the small of outgroup used here, this state is present in , Ampulicidae. However, segment a ventral sulcus in the Scolebythidae, Sierolomorphidae and Sapygidae. The con- present anteriorly mesepisternum of most could be the anterior dition in Ampulicidae is certainly not a reversal. The sixth Ampulicidae remnant of a long Such a character is also found in Sphecidae (sensu stricto) and a hypersternaulus. long and continuous hyper- few sternaulus can be seen in Crabronidae (e.g., Mellinus and Xenosphex). Its opti- Aphelotoma. mization on the tree that this condition arose sev- implies Johnson (1988) considered a midcoxa broadly con- eral times however, its in these independently; presence nected and open to the mesothorax as part of the basal that it be of the lineages suggests might part apoid groundplan for the Aculeata, but as demonstrated by been lost in ground plan, having Heterogynaidae, Apidae Sharkey and Wahl (1992) and Brothers and Carpenter lato) and in most Crabronidae. (sensu (1993), the ancestral condition for the aculeates is a midcoxa In Bohart and Menke's (1976) tribal classification for with a narrow, pedunculate basal portion, similar to that their Ampulicinae, Ampulex is placed in its own tribe, found in Heterogyna and Ampulicidae. whereas the remaining genera are placed together in a sec- A sister-group relationship between Sphecidae (sensu ond tribe, the Dolichurini. The results obtained here in the stricto) and Ampulicidae has been suggested by several implied weighting analyses contradict this classification, authors (e.g., Lomholdt 1982, Alexander 1992a, Ohl 1996a). because Ampulex and Dolichurus form consistently a mono- Two of the three supposed synapomorphies presented by phyletic clade. This relationship is supported by a rela- Lomholdt (1982) are simply plesiomorphies, as correctly tively strong synapomorphy, supra-alar carina not meet- interpreted Ohl (1996a). The third one, ing the tegular ridge (54-0). In the remaining Apoidea, in- by namely pres- ence of a is not valid because a scle- cluding Aphelotoma, the supra-alar carina is fused to the "propodeal sclerite", rotized the hindcoxal sockets is not anterior segment of the tegular ridge, a condition also bridge closing present in the and and also the present in Pompilidae. It seems improbable that the pecu- ampulicids Aphelotoma Doliehurus, liar mesoscutal morphology seen in Ampulex and condition in the others (Ampulex and Trirogma) does not Doliehurus (as well as in Trirogma and Paradolichurus) arose seem to be homologous to that of Sphecidae (sensu stricto) twice independently. (see Character 74). Of the four supposed synapomorphies presented by Alexander (1992a), only two were considered Sphecidae (sensu stricto) + [Apidae (sensu lato) + possible synapomorphies by Ohl (1996a). One of them, Crabronidae] female metasomal sternum VI forming an exposed taper- In the present study, the monophyly of this lineage ing tube (see Character 114), cannot be considered a (branch 3, Fig. 1) is strongly supported by the following synapomorphy because a similar condition is found in the synapomorphies: outgroup. The second character, "penis valves" with small 36 Scientific Papers, Natural History Museum, The University of Kansas

is treated here under teeth on ventral edges (Character 130-1), was optimized metapostnotum and propodeum this char- here as having arisen independently in Sphecidae (sensu Character 102. In the present analysis, however, this not the of (sensu stricto) and Ampulicidae. As was suggested above, acter does support monophyly Apidae for the + Crabronidae (see branch 7, 1 ) because Mellinus, condition might be part of the groundplan Apoidea. lato) Fig. which occupies a relatively basal position in the Sphecidae (sensu stricto) Crabronidae, has a condition similar to that of the basal Palinodes and Podalonia The name Sphecidae is used here in its narrow sense, apoids and because [Sphecidae of Bohart and have a condition similar to that of some corresponding to the subfamily Sphecinae (sensu stricto)] fur- Menke's (1976) classification. As explained in Material and crabronids. In any case, this character would require to include charac- ther to have its better evaluated. Methods, no particular effort was made study significance could the of this ters that potentially support monophyly The second synapomorphy used by Lomholdt (1982), came out in group. Despite that, Sphecidae monophyletic i.e. subbasal claw tooth absent, is interpreted differently several all analyses, being supported by unambiguous here. The subbasal tooth present in most Sphecidae (sensu Alexander (1992a) and Ohl changes (see branch 6, Fig. 1). stricto) is considered only a displaced, but still homolo- list several other for tooth (1996a) putative synapomorphies gous, subapical tooth (see Character 52). Asubapical stricto). Ohl's however, is com- and its absence in Sphecidae (sensu study, is part of the aculeate groundplan his of a and promised by assumption sister-group relationship Heterogyna represents an autapomorphy (Brothers (sensu stricto) and it is in between Sphecidae Ampulicidae. Judg- Carpenter 1993: see also branch 5 of Fig. 1); present no from ing by his list of exemplar taxa, representatives Ampulicidae and Apidae (sensu lato). Crabronidae or Apidae (sensu lato) were included in his A close relationship between Apidae (sensu lato) and Therefore, some of his supposed synapomorphies analysis. Crabronidae was also found by Alexander (1992a). Indeed, for (sensu stricto) could be plesiomorphies Sphecidae Crabronidae, as defined here, always comes out in shared with Crabronidae + Apidae (sensu lato). Alexander's analyses as a paraphyletic assemblage in re- the lin- The phylogenetic relationships among major lation to Apidae (sensu lato) (his Apiformes) and also some- were evaluated eages of Sphecidae (sensu stricto) recently times in relation to Ampulicidae. Taking into consideration The results of his studies confirm is not by Ohl (1996a). basically only Alexander's analyses in which Crabronidae Bohart and Menke (1976), those in which the arrangement proposed by paraphyletic to Ampulicidae and also only the that the Menke is re- the with exception genus Stangeella character polarization was determined by parsimony from (= their and it is the fol- moved Sceliphrini Sceliphronini) analyses (his analyses 5A, 5B, 6B, 7A, 7B, 9B, 10), of the clade + Ammo- placed as the sister group Sphecini lowing characters support, in at least one of these seven In the the the all the mem- philini. present study, relationships among analyses, the monophyly of a group including are con- four exemplar genera of Sphecidae (sensu stricto) bers of Crabronidae and Apidae (sensu lato) (numbers of the characters used gruent with Ohl's results. Most by from Alexander's list of characters): 17-1, 19-1, 25-0, 27-0, and it is Ohl (1996a) are from the male genitalia possible 32-1, 33-1, 38-1, 41-1, 47-1, 51-0, 67-2, 68-1, 80-1, 82-1, 83-1, of additional character that detailed investigation systems 84-1, 85-1, 88-1. Characters 17, 27, 33, 68, 80, 82, 83 and 85 alternative could produce phylogenetic arrangements. were not used in the present work and will not be dis- Another shortcoming of Ohl's study is that he did not sub- cussed. mit his statements to a test primary homology congruence Alexander's Characters 32-1 [Character 70-1 here] and the (e.g., parsimony) and therefore putative synapo- 51-0 [113-1] were also found here to support the mono- morphies presented by him are not derived from second- phyly of Crabronidae + Apidae (sensu lato) (see list of ary homology statements [sensu Pinna (1991)]. The rela- synapomorphies below); 25-0 [58/59] and 41-1 [49-1; am- tionships among the four genera of Sceliphrini restricted biguous in some optimizations], however, were found to to the Neotropical region were studied by Ohl (1996b). His the monophyly of Sphecidae (sensu stricto) + results also confirm Bohart and Menke's arrangement for support [Apidae (sensu lato) + Crabronidae]; 38-1 [52-1] and 88-1 these four genera. [136-1] were found to support the monophyly of + Apidae (sensu lato) Crabronidae Crabronidae (see below); 19-1 [53-1] supports a larger of the or, in some cases, a Lomholdt (1982) was the first one to suggest a sister- group, including part outgroup clade containing all Apoidea excluding Ampulicidae; 84-1 group relationship between Apidae (sensu lato) and was found to different clades within Crabronidae (his Larridae). Of the two putative synapo- [133-1] support is the Crabronidae, on the 67-2 [98-1 ] morphies he presented in support of this hypothesis, only depending analyses; condition in all taxa used, as well as in one, a shortened propodeum with an elongated meta- outgroup and and therefore of the postnotum, seems tenable. This modification of the Ampulicidae Heterogyna, presence Major Lineages of Apoidea 37 vein 2A was as in optimized arising de novo the ingroup crabronids, for example Odontosphex, Astata, Mellinus and taxa which 47-1 have it; [108-1] was treated here differ- the Philanthinae, but they have been lost at least three times ently. (see branches 12 and 24 and Nysson in the Bembicinae). These sacs have not been In all of my analyses, the bees [Apidae (sensu lato)] paired previously described and their function is unknown. the a also consistently grouped with a large assemblage formed Among Apoidea, larval with by all sphecid subfamilies of Bohart and Menke's classifi- spinneret paired openings is also unique to this clade. This is cation, with the exclusion of their Ampulicinae and the only synapomorphy provided by Lomholdt (1982) to the of his Larridae. Sphecinae, i.e., the Crabronidae. This relationship was support monophyly No case of reversal the Crabronidae is strongly supported by several characters, including the among known for the sec- ond or fourth for the following unambiguous changes (based on optimizations synapomorphies, except presence of a small subbasal tooth (or in the shown on Fig. 1): teeth) female's claw in some of (1) female antennae shortened (29-1); species Crabroninae [see genera Liris and Kohliella in Bohart and Menke in (2)medial portion of prothoracic basisternum (1976); Liris, the unique parallel teeth seems to be derived from the carinae the declivous in relation to anterior portion (sometimes parallel on ventral side of the claws]. only slightly), rounded or weakly pointed posteri- orly (44-1); Despite this relatively weak support, the recognition of (3)base apophyseal arms of prothoracic of a monophyletic Crabronidae seems well founded. Its endosternum connected (internally) by two con- supposed paraphyly in relation to Apidae (sensu lato) tinuous, divergent plates originating at the base of found in some of my analyses and also by Alexander furcasternum (46-1); (1992a) seems spurious. Alexander's results indicated a mesocoxal carina (4) present (70-1 ); possible sister-group relationship between Apidae (sensu dorsal lobe (5) on inner side of hindcoxal apex large, lato) and the Philanthinae (sensu Alexander, 1992a, b). This a usually forming spatulate process, ventral lobe relationship was consistently supported by both groups small or absent (77-1); subantennal sutures. sharing However, this similarity is VI (6) metasomal tergum of female with a pygidial probably superficial because in bees the subantennal su- plate (113-1); ture represents the line of attachment of the dorsal sheet (7) metasomal sternum VI of female not forming an of the anterior tentorial arm to the frons (Roig-Alsina and exposed tapering tube (114-0); Michener,1993), whereas in the Philanthinae the sutures (8) male cerci absent (122-1). have no connection to the tentorium. Also, subantennal sutures similar to those of the Except for absence of male cerci, all the unambiguous Philanthinae are present in some Crabroninae and some changes listed above seem to represent valid synapo- Bembicinae, Sphecidae (sensu morphies for Apidae (sensu lato) + Crabronidae. Males in stricto) (see Character 17). Some of my analyses, based on and successive several taxa of Crabronidae possess cerci and as discussed equal weighting of characters, resulted in a with bees more below (see section on the Astatinae), presence of cerci in paraphyletic Crabronidae, closely related to these crabronids should not be treated as a reversal, but Crabroninae (excluding Mellinus; Figs. 4 and 5). Rea- sons for not the results these two only retention of a plesiomorphic condition. Changes for favoring produced by methods were character 122 should have been optimized as irrereversible. parsimony discussed above. Contrary to Alexander's (1992a) results discussed Biogeographic patterns also contribute to reinforce the of a between above, most of my analyses supported a monophyletic hypothesis sister-group relationship Apidae (sensu lato) and Crabronidae. In both these several Crabronidae (branch 8 in Fig. 1). Four unambiguous taxa, changes were found to support this clade: lineages usually regarded as older basal clades are re- stricted or exhibit in deserts of the tem- (1 ) posterior wall of pharynx forming two bulging sacs to, higher diversity, (13-1); perate zone [for bees, see Michener (1979); for Crabronidae, (2) claws simple, without subapical or subbasal teeth one can cite the genera Odontosphex, Entotnosericus, Dinetus, (52-1); Xenosphex, Heliocausus, Pseudoscolia and the Astatinae as a (3) hindwing M diverging from CuA before or at cu-a whole]. Such a pattern is not apparent within the Sphecidae (97-0); (sensu stricto). (4) larval spinneret with paired openings, each at the Apidae (sensu lato) and Crabronidae could easily be end of a projection (136-1). treated under only one name, because the degree of diver- for a somewhat similar condition Except found in gence between them seems comparable to that present Heterogyna, the first character is unique to this clade; these among the subfamilies of Crabronidae. However, the bees are well pharyngeal expansions developed in several have traditionally been considered distinct from the rest 38 Scientific Papers. Natural History Museum, The University of Kansas of the Apoidea and such classification would be consid- sification is very similar to that proposed by Evans (1964a), ered confusing and probably of little use. based on larval morphological characters. It is worth not- ing that larvae of some of Crabronidae Dinetus, Apidae (sensu lato) genera (e.g., Entomosericus) were only recently described, and larvae are the intentional exclusion of characters known still unknown for others Despite (e.g., Eremiasphecium, Laphy- to support the monophyly of bees, except for Character 80 rngogus, Xenosphex), and therefore were not included in and a few character states unique to bees, this clade was Evans' studies. Evans' classification also differs from the present in the results of all analyses and in all cases, sup- one proposed here regarding the placement of the genus ported by a large number of synapomorphies (branch 9, Mellinus: Evans (1964a) recognized a monotypic subfam- Fig. 1). These putative synapomorphies are not listed or ily for Mellinus, the Mellininae; herein, I take a more con- discussed here; a discussion of the monophyly of bees and servative approach and include Mellinus in the a more complete and detailed list of their synapomorphies Crabroninae. In Evans' phylogeny, Mellininae is the sister can be found in Alexander and Michener (1995). group of his Larrinae, an arrangement that corresponds with the one here in which Mellinus is The various lineages of bees have been traditionally exactly preferred the basal clade of the Crabroninae. the classified in several separate taxa accorded family status However, position of Mellinus within the Crabronidae differs (e.g., Michener 1944, Roig-Alsina and Michener 1993, considerably the Alexander and Michener 1995). Such a classification was depending on analysis, an indication that additional studies reveal a different from the one fa- justifiable while the bees were considered the sole compo- might position vored here. nents of the superfamily Apoidea. However, with the in- clusion of other non- taxa in the Apoidea and the rec- Except perhaps for the Crabroninae, the monophyly ognition of only three superfamilies for the aculeate wasps of each of the five subfamilies recognized here seems well- (Brothers 1975), treating the higher groups of bees at fam- supported. In contrast, the relationships among the sub- families indicated in 1 ily level would leave the whole clade without a formal Figure can be considered very name and would be inconsistent with the higher-level clas- weakly supported. Two of the internal basal branches in- sification of the other aculeate clades. For these reasons, side the Crabronidae are each supported by only one un- some authors have adopted a classification in which all ambiguous change. The support for the branch leading to branches 19 + 20 is bees are treated under one family, the Apidae (e.g., even more suspicious considering that Lomholdt 1982, Gauld and Bolton 1988, Gauld and Hanson character 22 was considered inapplicable [i.e. coded (?)] for all the 1995, Griswold et al. 1995). The recognition of a monophyl- members of the Psenini (branch 22). At this point, etic Crabronidae as the sister group of bees, as proposed the relationships indicated should be taken as very tenta- here, strongly supports such a classification. The use of tive and a perhaps better representation would be to con- Spheciformes (= Sphecidae sensu Bohart and Menke) and sider the subfamilies forming a polytomy, except perhaps Apiformes (bees), informal divisions of the Apoidea pro- for Bembicinae and Philanthinae. Figure 9 summarizes the posed by Brothers (1975), should be avoided because relationships among the subfamilies indicated by implied Spheciformes, as it has been demonstrated, is paraphyletic weighting, as well as shows cladograms previously pro- in relation to Apiformes. posed for these groups by Evans (1964a) and Bohart and Menke (1976). In the present work, an investigation of the relation- In the the ships among the higher-level lineages of the Apidae (sensu following subsections, monophyly and com- of the five crabronid subfamilies are in lato) was avoided. This problem was recently studied by position discussed Roig-Alsina and Michener (1993) and Alexander and detail, particular attention being given to taxa whose place- ments are in conflict with Michener (1995). Nonetheless, the arrangements found those proposed by Bohart and here Menke (1976). Discussion of the here under implied weighting for the five bee representa- arrangements found for the more distal taxa within each tives do not contradict the possible arrangements found subfamily is avoided by Alexander and Michener (1995). because, in most cases, taxonomic representation is rather inadequate. Crabronidae Astatinae.—The subfamily Astatinae (branch 20) is The family Crabronidae (branch 8, Fig. 1), as defined defined here to include the tribe Astatini, the genus here, includes all the taxa classified at the subfamily level Eremiasphecium and the subtribe Ammoplanina of Bohart by Bohart and Menke (1976), with the exception of their and Menke's (1976) classification (each one of these groups Ampulicinae and Sphecinae. Only five crabronid subfami- is treated here as a tribe, i.e. Astatini, Eremiasphecini and lies are here recognized: Astatinae, Bembicinae, Ammoplanini). The discovery of this clade was somewhat Crabroninae, Pemphredoninae and Philanthinae. This clas- surprising because its members seem superficially distinct Major Lineages of Apoidea 39 and to a lesser extent because they were placed in totally projection and well separated from body's midline separate taxa in Bohart and Menke's classification. In their (68-3); classification, Astatinae contained the tribes Astatini and (5) use of Thysanoptera as larval food (137-1). Dinetini the Dinetus), (containing only genus In relation to the fourth synapomorphy, considering Eremiasphecium was part of the Philanthinae and that Pulverro has a distinct state (68-2) than the one indi- was of the The mono- Ammoplanina part Pemphredonini. cated above (68-3), one can suspect that the similar condi- of the Astatinae is the un- phyly supported by following tion in Eremiasphecium, Timberlakena and Ammoplanus could ambiguous changes: have evolved independently from a condition similar to to (1) tentorial pit of female situated above tangent that of Pulverro. These two tribes also share a similar mor- lower rims of antennal sockets (20-1); phology for the medial portion of the metepisternum (2) lateral arms of meso- and metathoracic furca weakly (Character 69-1 ) and practically lack the dorsal apical pro- fused (63-1); cess of the hindcoxa (78-3). However, these similarities cell shorter than (3) forewing marginal pterostigma (86-1 ); were not considered synapomorphies because of their male cerci (122-0). (4) present ambiguous status; the conditions for these characters Conditions (1) and (3) above are not unique to this present in Astata and in the immediate outgroups prevent clade; among the exemplar taxa, character state 20-1 was the optimization process from reaching any unambiguous also found to be a synapomorphy for the Spilomenina statements. (branch 24) and an for Nitcla and Dinetus, autapomorphy A sister-group relationship between Eremiasphecium whereas 86-1 is an for Dinetus. The 2nd autapomorphy and the Ammoplanini was recently postulated by Kazenas is to the Astatinae, it re- synapormorphy unique although (1991) when describing his new genus Taukumia [subjec- verses within the as a Ammoplanini, probably consequence tive junior synonym of Eremiasphecium; see Pulawski of a reduction in these are the body size; wasps among (1992)], although he assumed that Ammoplanini was part smallest 2 to 3 mm in The fourth Apoidea, only length. of the Pemphredonini and that Eremiasphecium was a mem- condition above cannot be considered as a valid synapo- ber of the Philanthinae; also he does not present any list of because it is that once lost the cerci morphy, highly unlikely characters that would support his hypothesis. Pulawski would be In the cerci are regained. Hymenoptera, quite (1992) revised Eremiasphecium and considered a possible in be considered reduced and most groups can vestigial. relation with Ammoplanini unfounded. The two charac- Absence of cerci in females is a for the ac- synapomorphy ters mentioned by him to dismiss this relationship, how- uleate (Brothers and and Hymenoptera Carpenter 1993), ever, have no bearings to this problem: absence of cerci no case of reversal is known. Given the condition vestigial and the 3rd submarginal cell in Ammoplanini. Presence of of the cerci in males of aculeate it seems Hymenoptera, cerci would certainly be simply a plesiomorphy (contrary to assume that their loss is irreversible. reasonable to Pulawski's assumption, males of most genera of If Dinetus had not been included in the analysis, some- Ammoplanini do have cerci) and loss of the 3rd submar- one looking only at these synapomorphies could run the ginal cell is an autapomorphy of Ammoplanini. that this is shown to be of risk of arguing genus easily part The monophyly of Ammoplanini is supported by nu- the Astatinae, as Bohart and Menke. originally proposed by merous unambiguous changes (see branch 28 in Fig. 1). But the carried out here show that the simi- analyses clearly Some of these changes (85-1 and 117-4), as well as some of larities of Dinetus to the Astatinae are the result of conver- the changes in the branch leading to Timberlakena and gence. Ammoplanus (1-1, 2-2, 33-0 and 81-1) also occur as part of or Astatini (represented by Astata in the cladograms) is the groundplan of the Pemphredonini (branch 21) only in the sister group to the rest of the Astatinae. The sister-group of the Spilomenina (branch 24). The similarities wing width relationship between Eremiasphecini and Ammoplanini characters, namely increase in pterostigma (85-1), and of (branch 27) is supported by: absence of the veins Cul and 2m-cu (90-1) presence and at (1) eye-clypeus contact extending for more than one only two submarginal cells, between Ammoplanini antennal socket diameter (16-2); least some of Pemphredonini were probably important in these two (2) epistomal suture, between antennal sockets, situ- Bohart and Menke's decision to maintain groups ated above median line across antennal sockets (19-1); together (absence of Cul and 2m-cu occurs in Spilomenina + (3) setal patch on anterior segment of tegular ridge and also in the branch leading to Parastigmus Stigmus, absent (55-1); but it is not shown because its optimization is ambiguous;

I these instead (4) medial flap of mesokatepisternum narrow, inter- would favor the repeated loss of two veins, then rupted in the middle by a deep cleft, and condyle of loss in the ancestor of Pemphredonini and reap- of mesal midcoxal articulation situated at tip of flap pearance in the branch leading to Diodontus, Pemphredon 40 Scientific Papers, Natural History Museum, The University of Kansas and Passaloecus). As shown by the analysis, these similari- close association of Ochleroptera with Heliocausus, sup- ties between Ammoplanini and Pemphredonini are the ported here by a considerable number of unambiguous result of convergence, probably in relation to reduction of synapomorphies (see branch 17), seems well founded. It body size in these lineages. Danforth (1989) showed that is interesting to note that males of some species of small species of Hymenoptera tend to have disproportion- , a group closely related to Ochleroptera, exhibit ately large pterostigmata, but it is worth noting that the derived morphological features similar to those typical of Crabroninae apparently does not follow this trend; very Heliocausini males, as for example eyes large and strongly small species in this group, like some species of Belomicrus, converging above, thorax somewhat spherical and have the pterostigma as reduced as that of its larger rela- metasomal sternum II with a strong, keel-like protuberance. tives. Also, insects in loss of veins among general, wing The close relationship found here between Hoplisoides seems to be correlated with reduction in size, strongly body and Didineis in the implied weighting analysis also de- but this trend has not been documented. properly serves to be discussed. Didineis together with the genus Bembicinae.—Bohart and Menke (1976) recognized Alysson has traditionally been placed in a separate tribe seven tribes in their subfamily Nyssoninae (= Bembicinae). (or subfamily depending on the classification) distinct from rest of the I included in this study representatives of six of these seven the Bembicinae (e.g., Evans 1966, Bohart and tribes. One of them, Bohart and Menke' s Mellinini (con- Menke 1976, Krombein 1979, 1985), and usually consid- ered a relictual basal taining only Mellinus), is considered here the basal clade somewhat and group (e.g., Evans of the Crabroninae. The other five were found to form a 1966, Bohart and Menke 1976). [More recently Krombein monophyletic group (branch 16), being supported by: (1985) described a new genus, Anahjsson, from Sri Lanka; (1) eye-clypeus contact extending for the diameter of judging from the diagnostic characters, recognition of one antennal socket or less (16-1); Anahjsson probably makes Alysson paraphyletic, but this (2) internal divergent plates of prothoracic endo-ster- needs to be evaluated by a phylogenetic analysis.] How- num partially (dorsally) or completely fused to ever, I suspect that the often-assumed plesiomorphic ap- furcasternum, medial ridge absent at least dorsally pearance of the species in this group is in reality the result (47-1); of several derived modifications of the (3) omaular carina present (61-1); groundplan. The absence in Alysson and Didineis of the (4) hindwing clavus indicated by short incision or only "oblique scutal carina", a feature whose presence is heavily a shallow notch (99-1); weighted as diagnostic of the Bembicinae, could be the (5) medial longitudinal ridge on base of sternum I result of an elongation and narrowing of the whole body. present (108-1). Alysson and Didineis resemble four genera, Eogorytes (not examined), and Psaimnaletes, assumed The two main lineages (branches 17 and 18) found here , Oryttus to form a within the for the six bembicine taxa do not seem to be spurious, as monophyletic group Gorytini (Nemkov and The one could suppose because of somewhat different relation- Lelej 1996), especially Lestiphorus. spe- cies in these six have in common females with elon- ships compared to that indicated by the current classifica- genera fore in which the fifth tarsomere and arolia are tions. The two representatives of Bohart and Menke's gated legs, than those of the other an- Gorytini, Hoplisoides and Ochleroptera, did not form a mono- conspicuously larger legs, long tennae (in the slender and basal phyletic clade. Ochleroptera was found to be most closely particular long and no sulcus (at least on the related to Heliocausus (branch 17), whereas Hoplisoides is flagellomeres) mesepisternal of the Future most closely related to Didineis. This is not a surprising upper part mesepisternum). phylogenetic studies should take into consideration these rela- result considering that Bohart and Menke had already in- putative dicated the polyphyletic nature of their Gorytini (see their tionships. — Fig. 155). Also, as Alexander's (1992a) study did not evalu- Crabroninae. The Crabroninae (branch 10) is defined ate the monophyly of the tribes he used, one cannot as- here to include the genera Mellinus, Dinetus, Laphyragogus sume that his Gorytini is monophyletic. and Xenosphex, and the subfamilies Crabroninae and Larrinae of Bohart and Menke (1976). In Bohart and Nemkov and Lelej (1996) analyzed the phylogenetic Menke's classification Mellinus was considered the basal relationships among the genera of Bohart and Menke's in of their (= Bembicinae), Dinetus was Gorytini. Despite some weaknesses their study [e.g., lineage Nyssoninae in had assuming monophyly of the tribe, or using only charac- their Astatinae, and Laphyragogus and Xenosphex ters listed in Bohart and Menke (1976)], they also found each its own subfamily. Bohart and Menke (1976) treated that Ochloptera was very weakly associated with the rest their Larrinae and Crabroninae as separate subfamilies "for of Gorytini. However, they kept this genus (together with practical considerations". These two taxa have been treated in in the in Clitemnestra) the Gorytini, as the most basal clade. The under one name past (e.g., Evans 1964a), and also Major Lineages of Apoidea 41

the is more recent works (e.g., Lomholdt 1985, Menke 1988). The present analyses, however, Lapln/ragogus unambigu- valid name for the taxon including these two groups is ously placed in the Crabroninae (see branch 13). Four Crabroninae [see discussion in Menke (1993)]. synapomorphies support placement of Laphyragogus as the most basal branch of the distal crabronines (based on Bohart and Menke (1976) suggested that the genus opti- mizations shown in Fig. 1): Xenosphe.x was remotely, but most closely, related to male mandibles with tooth Mellinus. However, as these authors themselves acknowl- (1) apical only (5-1); (2) female mandibles with a subbasal cleft on their in- edged, their list of similarities between Xenosphex and ner (6-1); Mellinus involved only shared plesiomorphies. In edge (3) mid tibia with one spur (73-1); Alexander's (1992a) study, Xenosphex shows no consistent (4) of male fused association with any other taxon. In the present study, gonapophyses genitalia completely a tube (129-1). Xenosphex always came out as one of the basal lineages of dorsally, forming the Crabroninae. This position is supported by several The subbasal cleft in the mandibles of females (as well unambiguous changes (see branches 10-12), among them as in males in some groups) is present in several distal a notch on the outerventral margin of the mandible (7-1), crabronine taxa and is usually associated with a correspon- the a strong, lamella-like mesocoxal carina (71-2), the loss of dent notch (or notches) on each side of apical margin of 8 in also the paired expansions on the posterior wall of the phar- the clypeus [see e.g., Fig. Lomholdt (1985); Fig. ynx (13-0) and a linear forewing costal cell (84-1). The 2 in Pulawski (1995)]. These modifications of the mandible lateral are to in taxa considered morphology of the oral plate, with the arms strongly known occur only traditionally the taxa converging posteriorly, is also found only among mem- as members of the Crabroninae (among repre- bers of this lineage. The peculiar morphology, in compari- sented here, the inner notch is present in Lyroda, Palarus, son to the more distal crabronines, of the mesepisternum Plenoculus and Anacrabro, besides Laphyragogus). and in (medial of metepisternum ventrally Xenosphex flap The condition in Laphyragogus for two characters ex- narrow, anterior of mesokatespisternum portion hibiting synapomorphic change in branches 11 and 12, re- vertical not leveled with metepisternum medially, spectively, can be considered a derived divergence from seems to have been modified in mesepisternum) conjunc- what is present in the more distal crabronines: (1) the tion with the of the mesocoxa. The male enlargement geni- mesocoxal carina (Character 71) is not particularly strongly talia has a not generalized morphology (gonapophyses developed as in most distal crabronines, but its morphol- fused and volsella differentiated from the dorsally clearly ogy is somewhat reminiscent of their condition, especially like Dinetus and Mellinus, and unlike the dis- gonocoxites), at the region near the ventral articulation with the tro- tal crabronines. The larva of is unknown, but I Xenosphex chanter; (2) the relatively broad forewing costal cell (char- it to a anus would expect have ventral, preapical (132-1), acter 84) could be considered as an artifact due to the way as do all other Crabroninae. One modification interesting the character states were defined, because in Laphyragogus, observed in the of the male of X. timberlakei (the the vein C is unusually slender. The volsellae (Character male of examined) is the of lat- only Xenosphex presence 128), although clearly differentiated from the gonocoxites, eral transverse sulci at the bases of T4-7 (each lateral sul- seem to possess an intermediate morphology between a cus extends almost to the middle of the Exami- tergum). more generalized condition, as found in Dinetus and nation of KOH-cleared VI and VII showed hundreds terga Xenosphex, and the more specialized condition found in of chitinous ducts associated with each sulcus (these are most distal crabronines, because they are largely fused to ducts of unicellular It would probably epidermal glands). the gonocoxites. be worth checking to see if males of the other two species Some changes considered ambiguous in the optimiza- of Xenosphex also possess such sulci. tions can be taken as additional evidence supporting place- has been considered of Laphyragogus traditionally part ment of Lapln/ragogus in the Crabroninae. The Beaumont but Bohart and the Crabroninae (e.g., 1959), mesepisternal sulcus in Lapln/ragogus is very similar to that Menke (1976) placed it in its own subfamily based on some of most crabronines (reaching the anterior edge of wing features, the relatively generalized male genitalia, mesepisternum away from the body's midline). Although the peculiar mouthparts and the morphology of the tho- also found in other crabronid subfamilies, the elongate rax in the area adjacent to the midcoxae. They indicated a mouthparts and the putative use of holometabolous insects somewhat intermediate position between their Astatatinae as prey (Lepidoptera; see Kazenas 1985) suggest crabronine and Larrinae. Except for the specialized mouthparts, which taxa. Indeed, the somewhat specialized galeal comb, blunt is reminiscent probably could be considered autapomorphies, the remain- formed by numerous short, bristles, the condition in Palarus. seems to ing features used by Bohart and Menke do seem to sug- of present Laphyragogus a somewhat relictual and that gest exclusion of Laphyragogus from the Crabroninae. In represent specialized lineage 42 Scientific Papers, Natural History Museum, The University of Kansas

of this the has strongly diverged from the other distal crabronines, a monophyly group, especially periforaminal This is a structure not observed in other hypothesis somehow supported by its possession of sev- depression. unique eral unusual features, for example (Da proepisternum al- taxa included in this study. most sunk inside the (2) a completely pronotum, strongly The monophyly of the tribe Pemphredonini is sup- area and a reduced enlarged occipital correspondingly ported by several synapomorphies (see branch 21). Two in the pronotal collar, (3) enlarged mouthparts, particular main clades where found within this tribe, one represented broad (4) broad connection between the meso- cardines, by Spilomena and Arpactophilus [branch 24; the subtribe and and (5) basal metasomata, apically expanded Spilomenina as defined in Menke (1989)] and the other tarsomeres of the female the basitarsus foreleg, especially containing the subtribes Stigmina, as redefined in Menke also found in (such modifications of the tarsomeres are (1989) and Pemphredonina, as defined in Bohart and a most species of Eremiasphecium, probably convergence Menke (1976) (branch 23). The monophyly of Spilomenina due to in because these bear nesting sand, expansions large is very strongly supported, with 14 unambiguous synapo- rake bristles). morphies listed for branch 24. Some of these changes, how- The somewhat heterogeneous composition proposed ever, should not be considered part of the subtribe here for the Crabroninae might be seen by some as an in- groundplan, because they are not present in some of its a of restricted to dication of an artificial, rather than a natural, monophyl- members [e.g., group species Australia, of with one described (as a see etic group, especially because of the inclusion several, only species Spilomena; and does not silk highly divergent basal lineages. However, except perhaps McCorquodale Naumann 1988), possess for Mellinus, this heterogeneous composition seems rather glands (Character 112; see Melo 1997) and has a pygidial these conditions to be an indication of a relatively old clade, in which several of plate (Character 113); seem for this and not reversals as one its basal lineages did not go extinct. This pattern seems plesiomorphies group, in The clade also to support a basal position for the subfamily as a whole could suspect (Melo, prep.)]. containing within the Crabronidae. Stigmina and Pemphredonina is supported by fewer, but as well. The Pemphredoninae.—This subfamily (branch 19) is de- strong synapomorphies supposed sister-group between and fined here to include the genera Odontosphex and relationship Stigmina Spilomenina suggested Bohart and Menke (1976) was not the Entomosericus, and the tribes Psenini and Pemphredonini by supported by is a well-defined of Bohart and Menke (1976), except for their Ammoplanina. parsimony analyses. Stigmina monophyl- etic [Parastigmus + Stigmus; see also Finnamore Two tribes are recognized: Pemphredonini (branch 21), group (1995)], whereas only two support the which corresponds to Bohart and Menke's Pemphredonini synapomorphies of without their Ammoplanina, and Psenini (branch 22), monophyly Pemphredonina. which includes Odontosphex, Entomosericus and Bohart and The close relationship of Odontosphex and Menke's Psenini. Each one of these two clades is well-sup- Entomosericus to Bohart and Menke's Psenini can also be was in the ported in most analyses, but they formed together a mono- considered somewhat surprising. Odontosphex Philanthinae and Entomosericus had its own in phyletic clade only under implied weighting and under subfamily successive weighting (analysis of the complete data ma- Bohart and Menke's classification. The psenine wasps with

I look distinct from the more trix). Despite this relatively weak support, opted for hav- their metasomal petiole quite ing both clades under one subfamily, instead of treating robust and non-petiolate Odontosphex and Entomosericus. each as a separate subfamily, to emphasize their probable However, they share several derived characters (see sister-group relationship and at the same time, to avoid synapomorphies for branch 22), including a somewhat an unnecessary disruption of the traditional classification. specialized articulation between sterna I and II (Character 110). One subdivision for this tribe would be the The following synapomorphies were found to support possible of three subtribes, the basal clade Odon- unambiguously the monophyly of the Pemphredoninae: recognition Odontosphex) and the sister (1) apical inflection of clypeus joining epistomal ridge tosphecina (containing only subtribes Entomosericina (containing only Entomosericus) considerably mesal to tentorial pit (15-2); and Psenina (= Psenini of Bohart and Menke's classifica- (2) occiput with periforaminal depression (35-1); between Entomosericus (3) well-defined mesocoxal carina (71-1); tion). The sister-group relationship and Psenina, as well as the monophyly of Psenina, also The last character can barelv be considered a synapo- are well-supported (see branches 25 and 26, respectively). morphy for this group, because presence of a mesocoxal Philanthinae.—The Philanthinae (branch 15) was re- carina, which was probably well-defined when first origi- redefined Alexander (1992a) to include four nated, is part of the Apidae (sensu lato) + Crabronidae cently by only groundplan (see Character 70). The first two synapo- of the six tribes attributed to it by Bohart and Menke (1976). Alexander's the affinities of the two excluded morphies seems to provide the strongest evidence for the In study, Major Lineages of Apoidea 43 tribes, Eremiasphecini (containing only Eremiasphecium) Alexander (1992b) indicated that the only reliable and Odontospheciiu (containing only Odontosphex), re- synapomorphy for the Philanthinae is the presence of a unsolved. In a mained subsequent paper, Alexander clypeal brush in the males. Although in the present study (1992b) analyzed the relationships within the Philanthinae representatives of only two genera (Philanthus and and found evidence for recognition of only five monophyl- Aphilanthops) were included and no effort was made to etic genera: Philanthinus, Philanthus (including Trachypus), evaluate the monophyly of this subfamily, the unambigu- Pseudoscolia, Cerceris (including Eucerceris), Clypeadon and ous changes shown in figure 2 for the branch leading to Aphilanthops. the these two genera (branch 15) can be considered as pu- tative synapomorphies for the Philanthinae. LITERATURE CITED

L. E. 1948. handlirschi an ex- of British vol. Ahrens, [Solitary wasps Ammoplanus Guss., Insects, 6, Part 4. Royal Entomological Society of Lon- terminator of Thysanoptera], Entomologicheskoye Obozreniye 30:88 don, London, pp. 1-60. [in Russian]. Evans, H. E. 1957. Studies on the larvae of digger wasps (Hymenoptera, Alexander, B. A. 1992a. An exploratory analysis of cladistic relationships Sphecidae). Part III: Philanthinae, Trypoxyloninae, and Crabroninae. within the with superfamily Apoidea, special reference to sphecid Transactions of the American Entomological Society 83:79-117. wasps (Hymenoptera). Journal of Hymenoptera Research 1:25-61. Evans, H. E. 1958. Studies on the larvae of digger wasps (Hymenoptera, Alexander, B. A. 1992b. Acladistic analysis of the subfamily Philanthinae Sphecidae). Part IV: Astatinae, Larrinae, and Pemphredoninae. (Hymenoptera: Sphecidae). Systematic Entomology 17:91-108. Transactions of the American Entomological Society 84:109-139. B. A. and D. 1995. Alexander, C. Michener. Phylogenetic studies of the Evans, H. E. 1959a. Studies on the larvae of digger wasps (Hymenoptera, families of short-tongued bees (Hymenoptera: Apoidea). Univer- Sphecidae). Part V: Conclusion. Transactions of the American Ento- sity of Kansas Science Bulletin 55(ll):377-424. mological Society 85:137-191. A. V. and V. V. 1992. H. E. 1959b. Antropov, Gorbatovsky [Heterogyna polita, new spe- Evans, The larvae of the Ampulicidae (Hymenoptera). Ento- cies (Hymenoptera, Sphecidae): the first representative of the sub- mological News 70:57-61. family Heterogvnainae in the fauna of the USSR]. Byulleten' Evans, H. E. 1964a. The classification and evolution of digger wasps as Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii suggested by larval characters (Hymenoptera: Sphecoidea). Ento- 97:53-59 [in Russian]. mological News 75:225-237. Argaman, Q. 1986. of Heterogynaidae (Hymenoptera: Evans, H. E. 1964b. Further studies on the larvae of digger wasps (Hy- Aculeata). Israel Journal of Entomology 19:7-12. menoptera: Sphecidae). Transactions of the American Entomologi- Asis, J. D., S. F. Gayubo and J. Tormos. 1993. The larva of Psenulus schencki, cal Society 90:235-299. with some remarks on the larval characters of Pemphredonini (Hy- Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand menoptera: Sphecidae: Pemphredoninae). Annales de la Societe Wasps. Harvard Univ. Press, Cambridge (Massachusetts). Entomologique de France (N.S.) 29:55-60. Evans, H. E. 1971. The larva of Heliocausus larroides (Hymenoptera, S. F. 1997a. Asis, J. D., Gayubo, and J. Tormos. Description of the mature Sphecidae). Psyche 78:166-168. larvae of several species of Psenulus, with comments on larval char- Evans, H. E. and C. S. Lin. 1956a. Studies on the larvae of digger wasps acters in the genus (Hymenoptera: Sphecidae). Annales de la Societe (Hymenoptera, Sphecidae). Part I: Sphecinae. Transactions of the Entomologique de France (N.S.) 33:197-203. American Entomological Society 81:131-153. Tormos and S. F 1997b. of the E. Lin. Asis, J. D., J. Gayubo. Description mature Evans, H. and C. S. 1956b. Studies on the larvae of digger wasps larva of the wasp Dinetus pictus with phylogenetic implications for (Hymenoptera, Sphecidae). Part II: Nyssoninae. Transactions of the the tribe Dinetini (Insecta: Hymenoptera: Sphecidae). Journal of American Entomological Society 82:35-66. 242:179-183. H. D. A. W. T. R. Zoology Evans, E., Smith, W. Middlekauff, Finlayson and J. de. 1959. Le Beaumont, J. genre Laphyragogus Kohl (Hym. Sphecid.). Re- McGinley. 1987. Order Hymenoptera. In F W. Stehr (ed.), Immature vue Suisse de Zoologie 66:723-734. Insects. Kendal/Hunt Publ. Co., Dubuque, pp. 597-710. R. M. and E. E. Grissell. 1972. habits and larva of Pulverro E. Bohart, Nesting Evans, H. and M. J. West-Eberhard. 1970. The Wasps. University of monticola (Hymenoptera: Sphecidae). Pan-Pacific Entomologist Michigan Press, Ann Arbor. 48:145-149. Farris, J. S. 1969. A successive approximations approach to character Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World. Uni- weighting. Systematic Zoology 18:374-385.

of California S. I. versity Press, Berkeley. Farris, J. 1983. The logical basis of phylogenetic analysis. In N. Platnick D. 1975. classification of V. in vol. 2. Brothers, J. Phylogeny and the aculeate Hy- and A. Funk (eds.), Advances Cladistics, Columbia Uni- menoptera, with special reference to Mutillidae. University of Kan- versity Press, New York. pp. 7-36. 1990. in sas Science Bulletin 50(ll):483-648. Farris, J. S. Phenetics camouflage. Cladistics 6:91-100. D. 1993. of Brothers, J. and J. M. Carpenter. Phylogeny Aculeata: Finnamore, A. T 1995. Revision of the world genera of the subtribe Chrysidoidea and Vespoidea (Hymenoptera). Journal of Hy- Stigmina (Hymenoptera: Apoidea: Sphecidae: Pemphredoninae), menoptera Research 2:227-304. Part 1. Journal of Hymenoptera Research 4:204-284. 1986. Cladistics of the A. C. D. 1993. In H. Carpenter, J. M. Chrysidoidea (Hymenoptera). Jour- Finnamore, T. and Michener. Superfamily Apoidea. nal of the York 94:303-330. the Iden- New Entomological Society Goulet and J. T. Huber (eds.), Hymenoptera of World: An 1988. tification to Carpenter, J. M. Choosing among multiple equally parsimonious Guide Families. Agriculture Canada Publications, Ot- cladograms. Cladistics 4:291-296. tawa, pp. 279-357. Danforth, B. N. 1989. The evolution of hymenopteran wings: the impor- Gauld, I. D. and B. Bolton. 1988. The Hymenoptera. Oxford Univ. Press, tance of size. Journal of Zoology 218:247-276. Oxford. Day, M. C. 1984. The enigmatic genus Heterogyna Nagy (Hymenoptera: Gauld, I. D. and P. E. Hanson. 1995. The evolution, classification and Sphecidae; Heterogyninae). Systematic Entomology 9:293-307. identification of the Hymenoptera. In P. E. Hanson and 1. D. Gauld Day, M. C. 1985. Redescription of Heterogyna protea Nagy (Hymenoptera: (eds.), The Hymenoptera of Costa Rica. Oxford Univ. Press, Oxford, 10:125-128. Sphecidae; Heterogyninae). Systematic Entomology pp. 138-156. M. C. 1988. In W. R. S. D. Day, Spider Wasps (Hymenoptera: Pompihdae). Gayubo, F, J. Asis and J. Tormos. 1992. A new species of Palarus and R. R. Askew Handbooks for the Identification Dooling (eds.), Latreille from Spain with a comparative study on nesting behavior 44 Scientific Papers, Natural History Museum, The University of Kansas

and larvae in the genus (Hymenoptera: Sphecidae). Annals of the Liege. of 85:26-33. Entomological Society America Lindley, V A. 1992. A new procedure for handling impervious biological Gibson, G. A. P. 1993. Groundplan structure and homology of the pleuron specimens. Microscopy Research and Technique 21:355-360. in Hymenoptera based on a comparison of the skeletomusculature Lomholdt, O. 1982. On the origin of the bees (Hymenoptera: Apidae, of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Mem- Sphecidae). Entomologica Scandinavica 13:185-190. oirs of the Entomological Society of Canada 165:165-187. Lomholdt, O. 1985. A reclassification of the larrine tribes with a revision Gift, N. and P. F. Stevens. 1997. Vagaries— in the delimitation of character of the of southern Africa and Madagascar (Hy- states in quantitative variation An experimental study. System- menoptera: Sphecidae). Entomologica Scandinavica, Supplement atic Biology 46:112-125. 24:1-183. Goloboff, P. A. 1993. Estimating character weights during tree search. Maddison, W. P. 1993. Missing data versus missing characters in phylo- Cladistics 9:83-91. genetic analysis. Systematic Biology 42:576-581. Goloboff, P. A. 1997a. Pee-Wee, Version 2.8. Computer program distrib- Maddison, W. P. and D. R. Maddison. 1996. MacClade: Analysis of Phy- uted by the author, New York (NY.). logeny and Character Evolution, Version 3.06. Sinauer Associates, Goloboff, P. A. 1997b. Nona, Version 1.8. Computer program distributed Sunderland (Massachusetts). by the author, New York (N.Y). Maneval, H. 1939. Notes sur les hymenopteres (6e. serie). Annales de la Graybeal, A. 1994. Evaluating the phylogenetic utility of genes: a search Societe Entomologique de France 108:49-108. for genes informative about deep divergences among vertebrates. Marshakov, V G. 1976. Digger wasps of the genera Eremiasphecium Kohl, Systematic Biology 43:174-193. Ammoplanus Gir., Ammoplanops Guss. and Anomiopteryx Guss. (Hy- Griswold, T, F. D. Parker and P. E. Hanson. 1995. The bees (Apidae). In P. menoptera, Sphecidae) in the USSR and Mongolia. Entomological E. Hanson and I. D. Gauld (eds.), The Hymenoptera of Costa Rica. Review 55:115-125. Oxford Univ. Press, Oxford, pp. 650-691 . Matthews, R. W. and I. D. Naumann. 1 989. Nesting biology and taxonomy Handlirsch, A. 1906-1908. Die Fossilen Insekten und die Phylogenie der of Arpactophilus mimi, a new species of social sphecid (Hymenoptera: Rezenten Formen. Wilhelm Engelmann Verlag, Leipzig. Sphecidae) from northern Australia. Australian Journal of Zoology Hanson, P. E. and I. D. Gauld. 1995. The Hymenoptera of Costa Rica. 36:585-597. Oxford University Press, Oxford. McCorquodale, D. B. and I. D. Naumann. 1988. Anew Australian species Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, of communal ground nesting wasp, in the genus Spilomena Shuckard Urbana. (Hymenoptera: Sphecidae: Pemphredoninae). Journal of the Aus- B. D. C. 1994. Heraty, J. M., J. Woolley and Darling. Phylogenetic implica- tralian Entomological Society 27:221-231. tions of in the mesofurca and mesopostnotum Hymenoptera. Jour- McGinley, R. J. 1981. Systematics of the Colletidae based on mature lar- nal of Hymenoptera Research 3:241-277. vae with phenetic analysis of apoid larvae. University of California Horovitz, I. and A. Meyer. 1995. Systematics of New World monkeys Publications in Entomology 91:1-307. (Platyrrhini, Primates) based on 1 6S mitochondrial DNA sequences: Melo, G. A. R. 1997. Silk glands in adult sphecid wasps (Hymenoptera, a comparative analysis of different weighting methods in cladistic Sphecidae, Pemphredoninae). Journal of Hymenoptera Research analysis. Molecular Phylogenetics and Evolution 4:448^156. 6:1-9. Houston, T F. 1975. Nest, behaviour and larvae of the bee Stenotritus Menke, A. S. 1988. Pison in the New World: a revision (Hymenoptera: pubescens (Smith) and behaviour of some related species (Hy- Sphecidae: Trypoxylini). Contributions of the American Entomo- menoptera: Apoidea: Stenotritinae). Journal of the Australian Ento- logical Institute 24(3):1-171. mological Society 14:145-154. Menke, A. S. 1989. Arpactophilus reassessed, with three bizarre new spe- Iwata, K. 1976. Evolution of Instinct: Comparative Ethology of Hy- cies from New Guinea (Hymenoptera: Sphecidae: Pemphredoninae). menoptera. Amerind Publishing Co., New Delhi. Invertebrate Taxonomy 2:737-747. H. Janvier, 1928. Recherches biologiques sur les predateurs du Chili. Menke, A. S. 1993. Crabroninae vs Larrinae (Sphecidae). Sphecos 26:6-7. Annales de Sciences Nafurelles (Zoologie), lOe. Serie, 11:67-207. Menke, A. S. 1996. Neotropical Mellinus: a review (Hymenoptera: Janvier, H. 1956. Observations sur deux predateurs chasseurs Sphecidae). Memoirs of the Entomological Society of Washington d'homopteres (Hym. Sphegidae). Annales de la Societe 17:125-141. Entomologique de France 124:195-208. Menke, A. S. 1997. Family-group names in Sphecidae (Hymenoptera: Janvier, H. 1962. Recherches sur les hymenopteres nidifiants aphidivores. Apoidea). Journal of Hymenoptera Research 6:243-255. IV Le genre Diodontus (Curtis). Annales des Sciences Naturelles Michener, C. D. 1944. Comparative external morphology, phylogeny, and (Zoologie), 12e. Serie, 4:489-516. a classification of the bees (Hymenoptera). Bulletin of the American Johnson, N. F. 1988. Midcoxal articulations and the phylogeny of the or- Museum of Natural History 82(61:151-326. der Hymenoptera. Annals of the Entomological Society of America Michener, C. D. 1979. Biogeography of bees. Annals of the Missouri Bo- 81:870-881. tanical Garden 66:277-347. Kazenas, V. L. 1985. The prey of Laphyragogus turanicus Gussakovskij Michener, C. D. 1981. Comparative morphology of the middle coxae of (Hymenoptera: Sphecidae). Sphecos 10:17. Apoidea. Journal of the Kansas Entomological Society 54:319-326. Kazenas, V L. 1991. [Anew genus Tnukumin (Hymenoptera: Sphecidae) Michener, C. D. 1986. Family-group names among bees. Journal of the from Kazakhstan]. Zoologicheskii Zhurnal 70:156-158 [in Russian]. Kansas Entomological Society 59:219-234. Kazenas, V L. and B. A. Alexander. 1993. The nest, prey and larva of Miiller, H. 1872. Anwendung der Darwinschen Lehre auf Bienen. Entomosericus kaufmani Radoszkowski (Hymenoptera: Sphecidae). Verhandlungen des Naturhistorischen Vereines der preussischen Journal of Hymenoptera Research 2:221-226. Rheinlande und Westphalens 29: 1-96. A. G. 1998. Kluge, Sophisticated falsification and research cycles: conse- Nagy, C.G. 1969. A new taxon of the family Latreille (Hym., for quences differential character weighting in phylogenetic sys- Aculeata). Entomologische Mitteilungen aus Zoologischen Museum tematics. Zoologica Scripta 26:349-360. Hamburg 64:7-12. K. V. 1979. In K. P. P. Krombein, Superfamily Sphecoidea. V Krombein, D. Nemkov, G. and A. S. Lelej. 1996. Phylogenetic relationships and clas- Hurd, D. R. Smith and B. D. Burks (eds.), Catalog of Hymenoptera sification of the digger wasps tribe Gorytini (Hymenoptera: in America North of Mexico. Smithsonian Institution Press, Wash- Sphecidae, Nyssoninae). Far Eastern Entomologist 37:1-14. ington, DC. pp. 1573-1740. Ohl, M. 1996a. Die phylogenetischen Beziehungen der Sphecinae (Hy- Krombein, K. V 1985. Biosystematic studies of Ceylonese wasps, XV: A menoptera: Apoidea: "Sphecidae") aufgrund morphologischer of the monograph Alyssoninae, Nyssoninae, and Gorytinae (Hy- Merkmale des Exoskellets. Zoologische Beitrage (Neue Folge) 37:3- menoptera: Sphecoidea: Nyssonidae). Smithsonian Contributions 40. to Zoology 414:1^3. Ohl, M. 1996b. The phylogenetic relationships within the Neotropical 1954. Leclercq, J. Monographie systematique, phylogenetique et Podiinae with special reference to Podium Fabricius (Hymenoptera: zoogeographique des hymenopteres crabroniens. Lejeunia Press, Apoidea: "Sphecidae"). Deutsche entomologische Zeitschrift 43:189-218. Major Lineages of Apoidea 45

Pinna, M. C. C. 1991. Concepts and tests of homology in the cladistic Shuckard, W. E. 1837. Essay on the Indigenous Fossorial Hymenoptera. paradigm. Cladistics 7:367-394. Shuckard, London. 1897 F. A. B. H. Liu P. 1994. Pulawski, W. J. 1992. A review of Eremiasphecium Kohl, (Hy- Simon, C, Frati, Beckenbach, Crespi, and Hook. and menoptera: Sphecidae). Entomofauna 13:397-406. Evolution, weighting, phylogenetic utility of mitochondrial gene 1995. Gastrosericus 1839 and a of conserved chain reac- Pulawski, W. J. The wasp genus Spinola, (Hy- sequences compilation polymerase menoptera: Sphecidae). Memoirs of the California Academy of Sci- tion primers. Annals of the Entomological Society of America 87:651- ences 18, vi+173 pp. 701. Roig-Alsina, A. and C. D. Michener. 1993. Studies of the phylogeny and Smith, E. L. 1970. Evolutionary morphology of the external insect genita- classification of long-tongued bees (Hymenoptera: Apoidea). Uni- lia. 2. Hymenoptera. Annals of the Entomological Society of America 63:1-27. versity of Kansas Science Bulletin 55(4):123-173. Ronquist, F. and G. Nordlander. 1989. Skeletal morphology of an archaic Snodgrass, R. E. 1942. The skeleto-muscular mechanisms of the honey cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica bee. Smithsonian Miscellaneous Collections 103 (2):1-120. Scandinavica Supplements 33, 60p. Snodgrass, R. E. 1993. Principles of Insect Morphology. Cornell Univ. 1957. of the larva of Ithaca. of 1935 Rozen, J. G. External morphological description Press, [Reissue original edition]. with other P. F. 1991. Character Exomolopsis chionura Cockerell, including a comparison Stevens, states, morphological variation, and phylo- anthophorids (Hymenoptera: Apoidea). Annals of the Entomologi- genetic analysis: a review. Systematic Botany 16:553-583. cal Society of America 50:469-475. Swofford, D. D. 1999. PAUP*. Phylogenetic Analysis Using Parsimony (* of the bee tribe and Other Methods). Version 4.0. Sinauer Sunderland Rozen, J. G. 1984. Comparative nesting biology Associates, Exomalopsini (Apoidea, Anthophoridae). American Museum (Massachusetts). P. D. Novitates 2798:1-37. Swofford, D. L., G. J. Olsen, J. Waddell and M. Hillis. 1996. Phyloge- of the netic Inference. In D. M. C. Moritz and B. K. Mable Rozen, J. G. 1993. Nesting biology and immature stages rophitine Hillis, (eds.), bees (Halictidae) with notes on the cleptoparasite Biastes Molecular Systematics. Sinauer, Sunderland (Massachusetts), pp. (Anthophoridae) (Hymenoptera: Apoidea). American Museum 407-514. Novitates 3066:1-28. Thiele, K. 1993. The Holy Grail of the perfect character: the cladistic treat- 1974. of ment of data. Cladistics 9:275-304. Rozen, J. G. and R. J. McGinley. Phylogeny and systematics morphometric Mellitidae based on the mature larvae (Insecta, Hymenoptera, Wiley, E. O. 1981 . Phylogenetics. Wiley & Sons, New York. Apoidea). American Museum Novitates 2545:1-31. Zelditch, M. L., W. L. Fink and D. L. Swiderski. 1995. Morphometries, der and characters as Schuberth, J. and K. Schonitzer. 1993. Vergleichende Morphologie homology, phylogenetics: Quantified Fovea facialis und der Stirnseitendriise bei Apoidea und Sphecidae synapomorphies. Systematic Biology 44:179-189. (Hymenoptera, Aculeata). Linzer biologische Beitrage 25:205-277. B. 1992. Cladistics of the Sharkey, M. J. and D. Wahl. (Hymenoptera). Journal of Hymenoptera Research 1:15-24. 46 Scientific Papers, Natural History Museum, The University of Kansas

of 12. of Figs. 10-22. 10-Prementum of male of Mellinus alpestris, lateral view. ll.-Same, female Diodontus rugosus. -Same, female Mimesa cressoni. 13.-Clypeus and antennal sockets of female of Stigmus temporalis, frontal view. 14.-Same, female of Arpactophilus sp. 15.-Furcal arms of = female ol Astata nevadica, dorsolateral view; abbreviations: AA = anterior arm, PA posterior arm. 16.-Same, male of Pulverro mescakro, anterodorsal view. female of Mimesa abbreviation: = 19- 17.-Second phragma of male of Dolichurus sp., lateral view. 18.-Same, cressoni; PP pseudophragma. Pulverro 21. Timberlakcna 22. Wings of Aphelotoma rufiventris showing wing venation terminology. 20.-Forewing of mescalero. -Same, yucaipa. -Same, Eremiasphecium budrysi. Major Lineages ofApoidea 47

0.5 mm — Figs. 23-24. 23. Posterior portion of mesosoma of female of Chlorion aerarium, ventral view; abbreviations: C = condyle of mesal articulation of mid coxa, S = suture = between mesepisternum and metepisternum, F medial flap of mesokatepisternum. — = = 24. Same, lateral view; abbreviations: AMP anteroventral metapleural pit, HC hind coxa.

— = = Figs. 25-28. 25. Mesoscutum of male of Dolichurus sp., lateral view; abbreviations: SC supra-alar carina, TR = — = tegular ridge; scale 0.3 mm. 26. Same, male of Mimesa cressoni; scale 0.3 mm. 27.—Hind coxa of male of Dolichurus = = — sp., antero-ventral view; abbreviation: S socket of mesal articulation; scale 0.2 mm. 28. Metasoma of male of = Dolichurus sp., lateral view; note modified sternum II (S2); scale 0.5 mm. 48 Scientific Papers, Natural History Museum, The University of Kansas

— Figs. 29-35. 29. Paired sacs (S) of the pharynx (showed attached to labio-maxillary complex) of female of in = — Odontosphex paradoxus (specimen cleared KOH), frontal view; scale 0.5 mm. 30. Same as Fig. 29; note numerous acanthae sac scale = 0.2 31.-Section of of female of covering walls; mm. head Ammoplanus sp. (cfr. apache) showing sacs = — pharyngeal (S), oblique sectioning (anterior to the left); scale 0.1 mm. 32. Same as Fig. 31; note thick epidermis = (E) forming the sac wall and numerous acanthae occupying lumen (L); scale 0.05 mm. 33. —Expansions (E) of the upper pharynx of female of Philanthtis gibbosus (material preserved in Kahle's fixative), frontal view; note also pharyn- = geal sacs (S); scale 0.5 mm. 34. —Internal view of lower frons, clypeus, and labrum (soft tissues and most of tentorial arms and = paramandibular processes removed) of female of Diodontus flavitarsis; abbreviations: AI apical inflection of TP = anterior tentorial = — clypeus, pit; scale 0.3 mm. 35. Internal view of lower frons and clypeus (soft tissues and most of tentorial = arms removed) of female of Ochleroptera bipunctata; abbreviations: AI apical inflection of clypeus, = = TP anterior tentorial pit; scale 0.3 mm. Major Lineages of Apoidea 49

— female of Figs. 36-43. 36. Ventral internal view of the protruding antennal sclerite (left) of PhUanthus gibbosus; = 37. — section of of of americanus scape (right) still attached to socket; scale 0.3 mm. Sagital head female Stigmus show- = — ing the protruding antennal sclerite (AS); base of scape (S) inserted into socket; scale 0.1 mm. 38. Base of antenna of female of Diodontus flavitarsis showing eccentric insertion of the pedicel into the socket at the apex of the scape; the = — central portion of the membrane covering the socket is sclerotized (S); scale 0.3 mm. 39. Head of male of Stignuis = — americanus showing the distinctly enlarged frontal facets of the compound eyes; scale 0.5 mm. 40. Head of female of = = — as 40. Diodontus virginianus (Rohwer), frontal view; abbreviation: FF facial fovea; scale 0.5 mm. 41. Same Fig. = Close-up view of the elongate and shallow facial fovea (FF); scale 0. 1 mm. 42. —Vertex of female of Stigmus americanus, dorsal view; abbreviations: FF = facial fovea, SMF = secondary micropore field; scale = 0.1 mm. 43. —Close-up view of the facial fovea of female of Pidverro mescalero, frontal view; scale = 0.02 mm. 50 Scientific Papers, Natural History Museum, The University of Kansas

— = Figs. 44-50. 44. Head of female of Passaloecus monilicornis Dahlbom, posterior view; abbreviations: PD = = — periforaminal depression, PC preoccipital carina; scale 0.5 mm. 45. Pronotum of female of Passaloecus monilicornis, dorsal view; abbreviation: TC = transverse carina; scale = 0.3 mm. 46.—Prothoracic basisternum of female of Ckhrion = aerarium, ventral view; scale 0.5 mm. 47. —Prothoracic episternum of female of Ochleroptera bipunctata, lateral view; = = abbreviation: LS lateral sulcus; scale 0.2 mm. 48. —Internal view of prothoracic endosternum of male of Dolichurus = — abbreviation: = scale = 0.2 sp., anterior view; scale 0.2 mm. 49. Same, male of Mimesa cressoni; DP divergent plates; mm. 50. —Same, female of Didineis texana; scale = 0.2 mm. Major Lineages of Apoidea 51

— fore tibia of Mimosa cressoni and closed socket scale Figs. 51-58. 51. Apex of male of showing displaced spur (S); = = 0.2 mm. 52. —Internal view of the mesepisternum of female of Passaloecus monilicomis; abbreviation: MR mesepisternal = — for left ridge; scale 0.2 mm. 53. Thorax of female of Ammoplanus sp. (cfr. apache), ventral view (legs removed, except = mid coxa); abbreviation: MS = mesepisternal sulcus; scale 0.3 mm. 54.—Mesosoma of male of Stigmas amoricanus, = = 0.5 55. — coxa of male lateral view (legs removed, except for coxae); abbreviation: OS omaular sulcus, scale mm. Mid = = — of Mellinus crabroneus (Thunberg), dorsolateral view; abbreviation: C midcoxal carina; scale 0.2 mm. 56. Same, male of Mimosa cressoni; scale = 0.2 mm. 57. —Same, female of Plenoculns davisi, lateral view; scale = 0.2 mm. 58. —Same, = male of Dolichurus sp., ventral view; scale 0.2 mm. 52 Scientific Papers, Natural History Museum, The University of Kansas

Figs. 59-62. 59. —Posterior portion of mesosoma of male of Dolichurus sp., postero-ventral view (legs removed); = = abbreviations: S suture between mesepisternum and metepisternum; C condyle of mesal articulation of mid coxa; = scale 0.5 mm. 60. —Same, male of Mimesa cressoni; scale = 0.3 mm. 61.—Mesosoma of female of Stigmus americanus, = ventral view (legs removed, except for right mid coxa); abbreviation: C condyle of mesal articulation of mid coxa; = — scale 0.5 mm. 62. Posterior portion of mesosoma of female of Pulverro mescalero, posteroventral view (legs removed); = = abbreviations: S suture between mesepisternum and metepisternum; C condyle of mesal articulation of mid coxa; scale = 0.3 mm. Major Lineages of Apoidea 53

— Figs. 63-69. 63. Hind coxa of female of Pulverro mescalero showing dorsal, crest-like lamella (L), inner view; = scale = 0.1 mm. 64.—Same, male of Mimesa cressoni, inner view; abbreviation: UL upper lobe; scale = 0.2 mm. 65.— Hind tibia of female of Pulverro mescalero, inner view; MF = micropore held; scale = 0.1 mm. 66.—Same, female of = — Ammoplanus sp. (cfr. apache); scale 0.1 mm. 67. Cross-section of hind tibia of female of Ammoplanus sp. (cfr. apache); = = = abbreviations: GE glandular epidermis, MF micropore field; scale 0.05 mm. 68.—Pterostigma of female of Stigmus = = — americanus, dorsal view; abbreviation: MF micropore view; scale 0.2 mm. 69. Same as Fig. 68; close-up view of = micropore field; scale 0.05 mm. 54 Scientific Papers, Natural History Museum, The University of Kansas

' *

72 73

— = Figs. 70-75. 70. Pterostigma of female of Stigmus americanus, ventral view; abbreviation: MF micropore field; = = . — — scale 0.2 mm. 71 Same as Fig. 70; close-up view of micropore field; scale 0.03 mm. 72. Cross-section of pterostigma of female of Stigmus americanus showing the distinctly developed epidermal glands (dorsal and ventral glands); scale = — = — 0.1 mm. 73. Same as Fig. 72; close-up view of glandular tissue; scale 0.05 mm. 74. Cross-section of pterostigma = of female of Passaloecus areolatus (material not stained); scale 0.05 mm. 75. —Close-up view of micropores (M) form- ing a diffuse micropore field on the dorsal surface of the pterostigma of the female of Diodontus rugosus; note absence = of pores on upper half of illustration; scale 0.01 mm.

QL566.S7 M3S 1999

I'ln log( ml I, ucl (Lis Harvard MCZ Library MDIUIi.

3 2044 062 474 572 Major Lineages of Apoidea 55

w

of scale Figs. 76-82. 76.—Mesosoma of male of Mimesa cressoni, posterior view; note deep sulcus (S) separating the two halves the propodeum; = = 0.3 mm. 77. —A. Posterior apex of sternum I of male of Mimesa cressoni, dorsal view; SF specialized surface. B. Anterior apex of sternum II = — showing specialized portion of lateral gradulus (G), ventral view; scales 0.1 mm. 78. Anterior apex of sternum II of female of Diodontus virginianus, = = = ventral view; G gradulus; GL glandular integument (contours indicated by dotted line); scale 0.1 mm. 79.—Posterior apex of sternum VI of = = — female of Diodontus virginianus, ventral view; ML medial lohe; scale 0.1 mm. 80. Posterior apex of metasoma of female of Pulverro mescalero, = 81.— of ventral view; note modified apex of sternum VI (S6); scale 0.1 mm. Same, female Ammoplanus sp. (cfr. apaclie); note denticulate apex of = = sternum VI (S6); scale 0.1 mm. 82.—Base of apical projection of sternum VIII of male of Stigmas americanus, lateral view; scale 0.02 mm. NATURAL OF KANSAS

The Univer: istory, beginning with Volume 1 i; Shorter research papers formerly "he University of Kansas Natural ;r 180 in Decem- berl996. TheM isas Natural His- tory Museum be iber 68 in Febru- ary 1996. Monog v Museum were initiated in 1970 , liversity of Kan- sas Science Bulle nnued with Vol- ume55inl996. 1 I in a new series entitled Scientific y of Kansas, be- gun with Numbe i continue as an outlet for longer c Ml manuscripts are subject to criti al acceptance is at the discretion c

The publicatic composed us- Microsoft and are ing Wo DEMCO INC 38-2931 tputer __ „. ..Uiua3 1 1 printed by The Ur y mting services.

Institutional libraries interested in exchanging publications may obtain the Sci- entific Papers, Natural History Museum, The University of Kansas, by addressing the Exchange Librarian, The University of Kansas Libraries, Lawrence, Kansas 66045- 2800, USA. Available back issues of The University of Kansas Science Bulletin may be purchased from the Library Sales Section, Retrieval Services Department, The University of Kansas Libraries, Lawrence, Kansas 66045-2800, USA. Available is- sues of former publication series, Scientific Papers, and Special Publications of the Natural History Museum can be purchased from the Office of Publications, Natu- ral History Museum, The University of Kansas, Lawrence, Kansas 66045-2454, USA. Purchasing information can be obtained by calling (785) 864-4450, fax (785) 864- 5335, or e-mail ([email protected]). VISA and MasterCard accepted; include ex- piration date.

Series Editor: William E. Duellman

Printed by The University of Kansas Printing Services Lawrence, Kansas