AMERICAN MUSEUM Novttates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3060, 17 pp., 15 figures, 2 tables March 25, 1993
Phylogenetic Relationships of Euherbstia with Other Short-tongued Bees (Hymenoptera: Apoidea)
JEROME G. ROZEN, JR.'1
ABSTRACT
The nesting biology of the Chilean Euherbstia information and ofanatomy ofthe mature larvae. excellens Friese (Andrenidae: Andreninae) is pre- Data on nesting biology do not appear helpful at sented, including nest site preference, nest archi- this time in revealing the interrelationships ofthese tecture, provisioning, egg deposition, larval feed- taxa. Cladistic analysis of 24 larval characteristics ing habits, larval defecation, seasonal activity, and suggests that the Euherbstia and the Oxaeidae are daily adult activity. Its egg and mature larva are sister groups which in turn are a sister group of described. The latter is compared with larvae of the Panurginae and Andrena, and that the Sten- Andrena, the Panurginae, the Oxaeidae, and the otritidae is a sister group of the rest, i.e., (Steno- Stenotritidae. The phylogenetic relationships of tritidae + ((Panurginae + Andrena) + (Euherbstia these taxa are considered on the basis ofbiological + Oxaeidae))). INTRODUCTION This paper explores the evolutionary re- mation about the nesting biology ofthis tax- lationships of the monotypic Chilean genus on is now available, and the mature larva can Euherbstia (Andrenidae: Andreninae) with be compared with the immatures of other Andrena, Panurginae, Oxaeidae, and Steno- primitive bees. The first part of the paper tritidae. The primary source of data comes presents the information on nesting and de- from the discovery ofnests ofEuherbstia ex- scribes the mature larva; the second part cellens (Friese) on the southern border of the analyses the new data (particularly with ref- Atacama Desert in Chile. As a result infor- erence to the larva) in an attempt to show
' Curator, Department of Entomology, American Museum of Natural History.
Copyright C) American Museum of Natural History 1993 ISSN 0003-0082 / Price $2. 10 2 AMERICAN MUSEUM NOVITATES NO. 3060 how Euherbstia and the Andreninae are re- antennal sutures shared by the oxaeines and lated to other bees thought now, or in the typical andrenids, a character upon which recent past, to be tied to the Andreninae. Michener (1944) had relied heavily when he The Andrenidae is one of the largest fam- included the oxaeines in Andrenidae. Rozen ilies of short-tongued bees. It is currently di- (1965) placed less importance on this feature vided into two subfamilies, the Andreninae because double sutures forming a triangle be- and Panurginae. Information about the bi- neath each antenna had also been reported ology and immatures of the Andreninae, for the Australian colletid genus Stenotritus which contains only six genera (Michener, (now placed in the Stenotritidae, McGinley, 1986), has been extremely limited. The nest- 1980) and because males ofa few Heterosarus ing biology ofa number ofspecies ofAndrena and Pterosarus (formerly placed in the single has been published (see Batra, 1990, and Mil- panurgine genus Pseudopanurgus) possessed iczky, 1988, for references), but other an- only a single pair ofsubantennal sutures. (We drenine genera have not been so treated ex- now also know that the males ofthe rophitine cept for a paper by Rozen (1992) which genus Penapis has four subantennal sutures.) presented incomplete information for Ancy- I concluded that the oxaeids should be ac- landrena larreae Timberlake and a brief corded family status, a judgment that was mention ofthe nests ofEuherbstia excellens. accepted by most (Hurd, 1979) but not all Of the six andrenine genera, larvae of only specialists (Graf, 1966). Andrena have been described taxonomically Rozen (1964, 1965) was unable, however, (see McGinley, 1989, for references). to demonstrate the relationship of the Ox- Ofall the possible relatives, the Panurginae aeidae with other bee groups but suggested have been, and continue to be, the group most that a tie with the stenotritines should be closely tied to the andrenines. The biology evaluated. This linkage was not supported and immature stages ofthe Panurginae, which when Houston (1975) discovered and de- contains conservatively 35 described genera scribed nests and larvae of Stenotritus pu- as well as a number ofundescribed ones, have bescens (Smith). Synapomorphies between been extensively studied in recent years. See larval Stenotritus and oxaeids could not be Rozen (1989) for references concerning life demonstrated, and the shared features of history information, and McGinley (1989), adults appeared to be convergent, relating to concerning immatures. premating behavior (as was the case with The Oxaeidae, a small, homogeneous fam- Melitturga). ily identified by many autapomorphies, had Hurd and Linsley (1976) extensively re- in the past been linked to the Andrenidae. viewed the literature on many aspects of the Michener (1944) considered it one ofthe three biology ofthe Oxaeidae. Roberts (1973) gave distinct subfamilies of the Andrenidae be- a relatively complete account of the nest of cause the Old World panurgine genus Mel- Oxaea flavescens Klug, and his information itturga shared many similarities with it and agrees closely with my unpublished obser- therefore seemed to be a phylogenetic inter- vations on Protoxaea gloriosa and Mesoxaea mediate between the Panurginae and the Ox- nigerrima (Friese). Larvae of these three ox- aeinae. Rozen (1964) found the mature larva aeid taxa are also remarkably similar (Rozen, of Protoxaea gloriosa Fox to be markedly 1964; Roberts, 1973; unpublished informa- different from that of any known panurgine tion). (that of Melitturga then unknown) or ofAn- McGinley (1980) recognized Stenotritidae drena. Subsequently he (Rozen, 1965) dem- as a family separate from Colletidae, but was onstrated that the larva of Melitturga clavi- unable to establish its affinities with other cornis (Latreille) shared no significant features groups of bees. Hence this family and the with the larva of Protoxaea and was able to Oxaeidae consisted of small assemblages of explain most ofthe adult similarities as evo- genera removed from obvious ties with other lutionary convergences associated with mat- short-tongued bees. Houston (1975, 1984) ing behavior. and Houston and Thorp (1984) have written The only feature that could not be so ex- a number of papers exploring the biology of plained was the presence oftwo pairs of sub- Stenotritidae and describing their larvae, and 1 993 ROZEN: EUHERBSTIA 3
stenotritid larvae have been made available at the start of the present investigation. I to me for this study. studied it intensively for three days and con- The present availability of data on the bi- tinued thereafter to make casual observations ology and mature larva of Euherbstia com- while pursuing other investigations through bined with similar information about the October 20. I revisited the site on November Panurginae, Oxaeidae, and Stenotritidae in- 15 and 16. vites reconsideration of the possible relat- The site was on an arid mountainside at edness of these taxa at this time. the southern end ofthe Atacama Desert. This region has been called the Coquimban Desert by Peiia (1966) and O'Brien (1971). O'Brien ACKNOWLEDGMENTS characterized the Coquimban Desert as hav- I would like to thank Elizabeth Chiappa, ing "a rich xerophytic flora, with many cacti Luisa Ruz, and Haroldo Toro for showing and shrubby legumes." Prof. Dr. Rodolfo me the site (which was then inactive) in 1989. Gajardo, University ofChile, refers to its veg- I extend my sincere appreciation to Luis E. etation type as desertic shrubby (Pefia, per- Pefia Guzman and Alfredo Ugarte Peina for sonal commun.) and believes that the area their assistance and companionship during contains remnants ofthe old flora that existed field investigations in 1991. Richard H. Kru- in the Atacama Desert. Gajardo suggests that zansky kindly analyzed the soil from the nest- Euherbstia and the related Orphana may also ing area. be relicts. Ground cover in the vicinity ofthe Some of the specimens ofAndrena exam- site (fig. 1) was dominated by low herbaceous ined in this study are on loan from the Essig plants and scattered shrubs. Precipitation, Museum of Entomology, University of Cal- which obviously strongly influences the ecol- ifornia, Berkeley; the Snow Museum, Uni- ogy ofthe area, occurs during the winter, and versity of Kansas, Lawrence; and the USDA flowering coincides with spring warming. Hot Bee Laboratory, Utah State University, Lo- dry summers follow, with no new flowering gan. I would like to thank Ronald J. Mc- on the mountainside. Consequently, adult ac- Ginley for allowing me to examine the larvae tivity of bees is limited to the spring. of Stenotritidae which he plans to describe The pollen-nectar plant of Euherbstia ex- taxonomically by arrangement with Terry F. cellens at the site was not found even though Houston. The late Radclyffe B. Roberts kind- three persons collected in the area for the five ly deposited voucher specimens of larvae of days ofstudy. This fact suggests that the food Oxaea flavescens in the American Museum source was remote from the nesting site. of Natural History, making direct compari- Approximately 20 nests were discovered, sons with that species possible. all restricted to a completely unvegetated, re- Thanks are due Wallace E. LaBerge and cently graded, unpaved road (figs. 1, 2), at Charles D. Michener, both of whom re- the base of a cuesta, far from any source of viewed this manuscript. permanent or semipermanent water (5 km Field work resulting in this report was sup- from the Elqui River). Nests were in the iden- ported by grant number 3844-88 from the tical place where Chiappa had found them National Geographic Society. abundant in 1987; they were scattered in an area about 6 m long and 1.5 m wide on the side ofthe road, where the surface sloped 200 NESTING BIOLOGY from horizontal. The nesting surface was DESCRIPTION OF SITE: The nesting site was completely unshaded and rarely traversed by discovered by Elizabeth Chiappa accompa- motor vehicles. The substrate contained nu- nied by Haroldo Toro and Luisa Ruz in Oc- merous irregular cracks created by the shrink- tober 1987 in an unpaved roadway approx- ing of soil as it dried. These cracks were ob- imately 6 km south ofVicufna, Elqui Province, scured in many places by a thin Chile, elevation 600 m. Although the site was (approximately 1 cm) layer offine loose pow- completely inactive in October 1989 (no dery soil. doubt because ofthe very dry winter), it was Subsurface soil was moist and had a clay fully active when visited on October 16, 1991, texture. Particle size on analysis was found 4 AMERICAN MUSEUM NOVITATES NO. 3060
i.1--: %I
Fig. 1. Nesting site of Euherbstia excellens in unpaved road (arrow), 6 km south of Vicufia, Elqui Province, Chile. to be 30% sand, 26% silt, and 44% clay. The lected between 6:06 and 7:40 p.m. on No- soil was difficult to dig because of its hard, vember 15 and 16 on the flowers ofAdesmia. compact nature and tendency to fracture into Nests ofneither ofthese bees were found, but large chunks. Other areas in the roadway two nests ofParasarus atacamensis Ruz were seemed equally suitable for nesting by Eu- observed nearby but not in the same road herbstia so that it is unclear why the nests and in different soil conditions. were restricted to this limited area. DESCRIPTION OF NESTS: We examined three The cracks in the soil, resulting from its nests with care and encountered numerous clay composition, appeared to be important others from the current year as well as from not only for Euherbstia but also for a number previous years while reaching these three and of other bee taxa. Tapinotaspis sp., a long- while searching at random for immatures. tongued undescribed panurgine genus, and Nests were constructed and occupied by Leioproctus erithrogaster Toro and Rojas and single females, and all nest entrances were in its cleptoparasite, Kelita toroi Ehrenfeld and cracks (fig. 3). Tumuli, always of loose soil, Rozen (New Information) commonly nested were apparently invariably present but were in the roadway and used these cracks as en- often inconspicuous because much of the ex- trees to the subsurface. cavated soil accumulated in cracks rather than In addition, other bees were encountered on the surface. Hence entrances were best in the area but were not detected nesting in the roadway. Chilimalopsisparvula Toro was abundant during the October study, and four cies are currently recognized (Rozen, 1971), the females collected here bridge the coloration differences of the females ofthe andrenine Orphana2 were col- two species and raise the question as to whether the species are distinct. Now that we suspect the vespertine 2 This genus, considered rare, may be vespertine as habits of Orphana, series of males (and more females) suggested by the hours of collection. Although two spe- can perhaps be collected to answer this question. 1993 ROZEN: EUHERBSTIA 5
*£wwhi' i' W;. 't' A<- I ¢-su,;ffi-: - _ \ mw-': > *rx0.jh<'S--_ .' ^ ;.. .~~~ mr :\\ * I -. \S _ I J_..... _-., 'sk, ! 9, 1,F. C. t- i I im--.'Ar c.-,-, A ., .. t. .& 1. k */ W |l~~~~' I rV z... ../Il-11 *LL4 2 ,11 -~~~~~~~~~~~~~~i NetigitoEhesi exc.S7iWhe :op o Pa powd mk n Fig. 2. Nesting site of Euherbstia excellens. White circles of plaster-of-Paris powder mark nest entrances. Fig. 3. Close-up of two nest entrances. 6 AMERICAN MUSEUM NOVITATES NO. 3060 identified by observing returning females as and usually became indistinguishable from they descended into cracks. One conspicuous the substrate. tumulus was about 7 to 8 cm in diameter and Cells were arranged singly (i.e., not in linear less than 1 cm high. Excavated material tend- series) at a depth of 13 to 49 cm. They were ed to accumulate on the downhill sides of elongate ovals, 16.0 mm long (N = 7) and entrances. 8.0-9.2 mm in maximum diameter (median Within the cracks, entrances were open, 9.0 mm; N = 9). Rounded at the rear, they and main tunnels descended following a me- appeared somewhat asymmetrical in that the andering path first through the crack itself ceiling was perhaps slightly more vaulted than and then into the substrate. Main tunnels were the floor. Cell orientation varied greatly, with generally open; apparent occasional short the long axis tilting from 35 to 850 (median plugs may have been created either by the 500; N = 19) from horizontal, the front end female or by our own excavation accidentally always higher than the rear. allowing soil to fall into the tunnel. Circular Cell walls (as distinct from linings) were in cross section and 6.0 to 7.0 mm in di- not distinguishable in texture, hardness, or ameter, tunnels were unlined, and their walls color from the substrate. Cell linings were were rather rough and completely absorbent shiny, and covered the entire surface of the to water droplets when tested. Microscopic cell except for the closure. When the substrate examination ofthe walls showed distinct im- was allowed to dry, the cell lining appeared prints of the females' pygidial plates, indi- semitransparent and waxlike in that it could cating that females tamped the walls repeat- be scraped with sharp forceps leaving a shiny edly during excavation. trough and a ribbon of scraping. Further- Apparently females excavated main tun- more, the lining could not be peeled from nels to their lowest level before starting cell fragments of moist cell wall as the wall was construction. Cells, connected by short side broken. However, this material was not wax- tunnels to the main tunnel, were then exca- like in that, when a piece of cell wall was vated, provisioned, and closed one by one in heated on a hot plate to 700°F, the lining did a generally ascending order. Hence, lower cells not melt but rather it charred (as does paper tended to contain the oldest immatures, and at the same temperature). When a piece of cells at the upper levels were most recently dried cell wall with attached lining was sub- constructed and provisioned. However, merged in water, the wall fell apart and sep- groups ofcells in a nest at approximately the arated from the lining. However, a thin layer same depth not infrequently were found to of very fine soil continued to adhere to the have a slightly older immature above a youn- lining, suggesting that the lining material ger one. This might be explained ifsome side (presumably a secretion) had slightly im- tunnels sloped more than others or if a fe- pregnated the cell wall and had bound to- male, after filling a side tunnel, constructed gether fine soil particles as it was applied by the next one at a slightly greater depth be- the female. The lining was completely wa- cause the main tunnel had not been com- terproof when tested with a droplet ofwater. pletely filled to the level ofthe last construct- The inner cell surface was quite smooth and ed cell. shiny although some small pebblesjutted into Side tunnels leading to cells were short, as the lumen ofa few cells. All such protrusions evidenced by cells being closely grouped were covered by the lining. Some cells also around main tunnels. Distances between cells exhibited faint impressions presumably made in a group measured 2, 2, 2.5, 2.5, 3, 3, and by the female's pygidial plate. 5 cm for a sample ofseven pairs ofcells from In addition to the lining, fresh cells bore a a number ofnests. Side tunnels were the same liquid over most of the inner surface except diameter (approximately 6.5 mm) as main near the closure. This liquid imparted a high tunnels except they narrowed at the cell en- gloss that was more reflective than that ofthe trance to about 5.0 mm. Their walls bore cell lining alone. In some cells it beaded into pygidial impressions like those of main tun- small flattened bubbles as if an oil were in- nels. Side tunnels were filled after cell closure volved in addition to a water-soluble liquid. 1993 ROZEN: EUHERBSTIA 7 When cells dried after excavation, the liquid distinct, rough, and uncoated. They varied disappeared. The source and significance of considerably in width from cell to cell so that this liquid are unknown. However, the thin some cells had as few as three coils, and oth- veil of mold (described below) on the inner ers as many as five, although most had four surface ofcell closures and cell walls adjacent to five coils. Diameter ofthe inner surface of to closures (surfaces without the liquid) might cell closures ranged from 5.2 to 6.5 mm (me- suggest that the liquid is a mold inhibitor. dian 6.0 mm; N = 1 1). These figures are great- Cell closures were distinct spirals on the er than the diameter of the cell entrance be- inner surfaces which varied from being slight- cause most closures extend into the cell ly concave to having the center coils consid- beyond the narrowest part of the cell en- erably more recessed than the outer ones so trance. that surfaces were more strongly concave. The All except very fresh cells exhibited a mass fill of the side tunnel fused with the closure ofwhite, interwoven, fine mold hyphae cov- in most cases, so that the presence of a con- ering the inner cell closure and extending sev- cave outer surface is uncertain. One closure eral millimeters onto the cell wall and into may have had a distinct outer surface in which the cell lumen. This mass partly obscured the case the closure was 3.0 mm thick in the mid- spiral structure of the closure but was not dle. In any event, closures were unlike the sufficiently dense to hide the pattern com- elongate, loosely filled ones ofAncylandrena pletely. larreae (Rozen, 1992) and the double or mul- PROVISIONING AND DEVELOPMENT: Larval tipartitioned closures of Stenotritidae and provisions ofEuherbstia excellens were mealy Oxaeidae.3 The coils ofthe spiral closure were moist, grayish orange masses formed into flattened spheres. The spheres had a height 3 Houston (1984, 1987) and Houston and Thorp (1984) (minimum diameter) of 5.3 to 6.0 mm (me- recognized and carefully described cell closures of the dian 5.5 mm; N = 9) and a horizontal di- Stenotritidae as consisting of one or more outwardly ameter (maximum diameter) of 7.0 to 7.6 concave septa of soil ("false cell bases") in the lateral in mm (median 7.2 mm; N = 10). One pollen front of a plug of soil closing the cell mouth. The inner with a of surface of the plug bore a concave spiral pattem (char- mass was slightly elongate height acterisitic of many groups of bees); its outer surface was 5.7 mm, greater horizontal diameter of 7.2 identical to the concave septa in front of it. Between the mm, and the other horizontal diameter of6.7 outer surface and the septum in front of it, the lateral mm. The flattened spheres were dull on the was filled with loose soil and gravel, as were spaces be- surface and lacked a coating of either nectar tween septa ifthere was more than one septum. Because or waterproof material. Because the masses such closures had not been observed before, Houston were sweet to taste, nectar is thought to be (1987) believed them to be restricted to the Stenotritidae. the liquid that held the pollen together. The I now believe that the elongate cell closure of Ancylan- flattened spheres were positioned toward the drena (Rozen, 1992) may be a homologous behavioral rear of the cell with the plane of the maxi- feature. Furthermore, on examining the cell closures of Protoxaea gloriosa Fox deposited in the American Mu- mum diameter approximately parallel to the seum ofNatural History, I found that they too routinely long axis of the cell. possessed concave septa-in one case as many as 11 Eggs and larvae rested on top ofthe pollen septa in addition to the outer surface of the plug! This mass (i.e., the surface opposite to that which matter becomes even more complex because in the same unpaved road where Euherbstia excellens nests, a new new panurgine, brood cells are horizontal or nearly so, panurgine genus with an extremely long tongue nested but the Protoxaea cells are vertical. Cell walls of Sten- and its cell closures consisted of three to four septa (in- otritidae, Ancylandrena, and Protoxaea are thick and cluding the outer surface ofthe plug). Hence what seemed harder than the substrate and in all ofthese three genera at first to be a behavioral autapomorphy of the Steno- the cell wall extends forward surrounding the lateral where tritidae is actually also found in the Panurginae and Ox- the concave septa are found but no farther. Because of aeidae (but presumably not in Euherbstia). Further in- the hard walls surrounding the septa, I think ofthe septa vestigations into other groups ofbees should be pursued and the spiral plug together as the components of the in order to understand the evolutionary implications of cell closure. The cell wall of the new panurgine is thin this feature. so that whether it extends forward to surround the septa In the case of Stenotritidae, Ancylandrena, and the is uncertain. 8 AMERICAN MUSEUM NOVITATES NO. 3060 was in contact with the cell floor). One egg, Emerging adults exited from the cells 3.1 mm long and 0.7 mm in maximum di- through the cell closure. Vacated cells were ameter, was attached to the pollen mass only filled with soil, and the fecal material re- by its posterior end while its more blunt an- mained at the rear ofthe cell as did the larval terior end, raised from the food mass, pointed integument which could easily be recognized toward the cell closure. It was white, and its by the sclerotized and distinctive spiracular chorion was nonreticulate and shiny. tubercles. Young larvae fed on the pollen-nectar mass SEASONAL AND DIURNAL ACTIVITIES: All immediately beneath their heads, and, as they immatures encountered at the last visit to the grew, more and more ofthe front ofthe sphere site (November 15 and 16, 1992) were post- disappeared. Their position in relation to the defecating larvae so that there is no doubt food mass did not change, so that, even when that this species has at most a single gener- one third or one fifth of the food mass re- ation per year. Furthermore, a significant part mained, the large larva rested on the re- of the population may take two (or more) maining food (which appeared as a crescent years to mature because during the October seen in lateral view) even though the abdom- study we found many postdefecating larvae inal apex touched the rear cell floor. Larvae obviously remaining from the previous year did not reorient to rest on their dorsa on the and no pupae (which meant that none ofthese cell floor, as do many panurgines. With most larvae were destined to develop further dur- (but certainly not all) Panurginae, larvae that ing the current year). Lack oftotal population have eaten approximately halfthe food move emergence in any one season might have an onto their dorsal tubercules, cradle the food adaptive advantage for bees living in the vi- masses on their venters, and bring their heads cinity of Vicunia because of the uncertainty upward toward the food to complete inges- of winter precipitation from one year to the tion. next. Flowering of food plants apparently is Larval feeding ofEuherbstia excellens pro- not a dependable annual event, as indicated gressed rapidly. Nesting activity (nest con- by the absence of Euherbstia nesting in Oc- struction, foraging, egg-laying, and larval tober 1989. growth) was at the maximum during the ini- Postdefecating larvae that developed in tial investigation in mid-October, and by No- 1991 remained slightly active even two vember 16 all larvae encountered at the site months after reaching that stage. They con- had completed feeding, had defecated, and tinuously (even when not touched by forceps) had taken on all aspects of postdefecating rotate the abdominal apex very slowly by forms including darkening and hardening of slightly flexing the middle segments of the the integument and pigmentation of the spi- abdomen. With one specimen, the complete racular tubercles. rotation took 20 seconds. In the brood cell Fecal material was deposited as a single such activity apparently results in a slow curved thin mass appressed to the rear and shifting ofthe larval position or even rotation one "side" of the cell. The curvature of the of the entire larva. The advantage (if any) of mass conformed to the curvature of the cell. such activity is unknown. After this obser- Because of the uniformity of this placement vation was noted on 15 larvae from the cur- in many cells, it seems likely that the "side" rent season, I examined three postdefecating of the cell was actually the ceiling, as is usu- larvae remaining from the previous season ally the case with panurgines. Feces-laden cells and found them completely inactive. Surviv- were not observed in situ so that further study ing larvae from the current season had be- is required. come inactive by July, nine months after def- Larvae ofEuherbstia do not spin cocoons, ecating, but I did not observe when they had and as described below do not have the an- reached that stage. In any event muscular ac- atomical structures typical of cocoon-spin- tivity ceases before the following season of ning larvae. Postdefecating larvae had a rigid emergence. This prolonged period ofslow ab- integument, so that when punctured or sev- dominal flexing by postdefecating, hibernat- ered in two, their bodies retained their shape. ing larvae is uncommon in the Apoidea; most 1993 ROZEN: EUHERBSTIA 9 bee larvae when diapausing become inactive duced into acute, toothlike projection (figs. within a matter ofdays upon becoming post- 10-13) (not strongly produced, obtuse, Roz- defecating forms. Indeed, the term "postde- en, 1973: figs. 8, 10, except in Andrena4 sp. fecating larva" was originally coined to de- from Chiapas, Mexico5); swelling on outer scribe overwintering, completely inactive surface ofmandible pronounced (figs. 10, 12, larvae after they had voided their fecal ma- 13) (small, inconspicuous, Rozen, 1973: fig. terial and assumed (at least in the cases of 8, except in Andrena spp. from Chiapas, noncocoon-spinning larvae) integumental Mexico, and Lusignan, France); labiomaxil- changes that apparently help insure against lary region greatly recessed (fig. 8) (only mod- water loss. erately recessed, Rozen, 1973: fig. 6); cardo Adults ofEuherbstia excellens were active and stipes not sclerotized, vaguely pigmented during the warm part ofthe day. Provisioning (more sclerotized); spiracular tubercles pres- for a few females started before 10:00 a.m. ent (figs. 4, 6) (absent, Rozen, 1973: figs. 3, and still continued into late afternoon, but 4,7); body without dorsolateral tubercles (figs. peak activity seemed to be around midday. 4, 5) (with more or less transverse tubercles Many males were observed a few meters from on anterior segments; on abdominal seg- the nesting area searching the ground for ments these tubercles distinctly transverse and crevices, presumably their nighttime loca- appearing as elevated caudal annulets in lat- tion, around 3:00 p.m. Females almost cer- eral view, Rozen, 1973: fig. 3). tainly spend the night in their nests. A few The Panurginae as larvae are a diverse females seen searching the ground late in the group. Although many of their larvae have day for convenient cracks in which to spend been described (McGinley, 1989), others are the night were probably individuals that had still unknown. The mature larva ofEuherbs- either completed their nests or had not yet tia excellens can probably be separated from started nesting. most if not all Panurginae by the following PARASITISM: No cuckoo bees were associ- (panurgine features in parentheses) but this ated with any nests of Euherbstia excellens. listing is not necessarily complete: Labral tu- Cells were heavily infested by meloids. Bom- bercles apical, rounded (figs. 7,8) (arising from byliids, while common in the area, did not labral disc, usually acutely pointed); mandib- appear to attack Euherbstia nests in that no ular cusp produced into acute, toothlike, den- larvae were recovered from any of the cells. ticulate projection (figs. 10-13) (not pro- DESCRIPTION OF MATURE LARVA 4 These species include the following: (Callandrena) Figures 4-14 accepta Viereck, (Thysandrena) bisalicis Viereck, (Scrap- DiAGNosIs: The mature larva ofEuherbstia teropsis) morrisonella Viereck, (S.) imitatrix Cresson, can be from available larvae of (Tylandrena) halli Dunning, (T.) perplexa Smnith, (Leu- distinguished candrena) erythronii Robertson, (L.) placida Smith, Andrena by the features listed here. Char- (Euandrena) caerulea Smith, (Melandrena) flexa Mal- acteristics ofAndrena are given in parenthe- loch, (Larandrena) miserabilis Cresson, and two un- ses with figure numbers referring to Rozen identified species ofAndrena, one from Lusignan, France, (1973). Vertex not produced and without and the other from San Cristobal de las Casas, Chiapas, swelling on each side above antenna (fig. 8) Mexico. In addition, the published descriptions and/or (produced on each side and in many cases illustrations of larvae of other Andrena species listed in with distinct swelling above antenna, Rozen, McGinley (1989) seem to agree. 1973: figs. 5, 6); antenna on vague swelling (fig. 8) (on pronounced swelling, Rozen, 1973: 5The mandible of this species is similar to that of fig. 6); epistomal ridge between anterior ten- Euherbstia excellens both because of its produced, den- tate cusp and because the outer swelling is large and torial pits well developed (fig. 7) (absent, Roz- closer to the mandibular apex than base. In other respects en, 1973: fig. 5); mandible robust at base, the larva agrees with the characteristics ofAndrena pre- slender and tapering apically (figs. 11, 13) sented here, and the cuspal teeth are substantially larger (tapering throughout its length to slender apex, than those of Euherbstia. Unfortunately, no adults are Rozen, 1973: fig. 9); mandibular cusp pro- associated with these specimens. 10 AMERICAN MUSEUM NOVITATES NO. 3060 R TUBERCLES 1.0 r OMAL RIDGE L PALPUS \K\ PROJECTION \/ INNER APICAL / \ 14 10 t 1 CONCAVIW 13U Figs. 4-14. Mature larva ofEuherbstia excellens. 4. Postdefecating larva, lateral view. 5. Predefecating larva, lateral view. 6. Abdominal segments VIII-X, posterior view (drawn to larger scale than figs. 4, 5). 7, 8. Head, frontal and lateral views. 9. Right half of head, ventral view (drawn to larger scale than figs. 7, 8). 10-13. Right mandible, dorsal, adoral, ventral, and outer views, respectively. 14. Spiracle, side view. Scale refers to figures 4 and 5. 1993 ROZEN: EUHERBSTIA I1I duced, obtuse, or, if so produced as in ilar to that found in oxaeids (Rozen, 1964: Melitturga [Rozen, 1965: figs. 9, 10, 13], pro- fig. 3) except area around pits more strongly jection not denticulate); spiracular tubercles recessed and pits larger; postoccipital ridge, present (absent); body without dorsolateral hypostomal ridge, pleurostomal ridge, and tubercles (most segments with conical dor- epistomal ridge (including section mesad of solateral tubercles). anterior tentorial pits) well developed; epis- The following description follows closely tomal ridge nearly straight between anterior the format adopted in Rozen and Michener tentorial pits, not arching upward; median (1988). Comparisons with oxaeids were made longitudinal thickening of head capsule only after examination oflarvae ofProtoxaea glo- faintly developed near postoccipital ridge, riosa, Oxaea flavescens, and Mesoxaea ni- absent below. Parietal bands distinct. Anten- gerrima, all in the collection ofthe American nal prominences only vaguely developed; each Museum of Natural History. Larvae of the antenna a small, low convexity with approx- following stenotritids, kindly loaned by R. J. imately four sensilla; antennal disc small. La- McGinley, were also examined comparative- brum (figs. 7, 8) bearing two rounded tuber- ly: Ctenocolletes nicholsoni Cockerell, C. or- cles (swellings) apically, similar to those of densis Michener, and Stenotritus greavesi Andrena accepta (Rozen, 1973: figs. 5, 6) and (Rayment). much more rounded than the acutely pointed LENGTH: Postdefecating larva 12-13 mm. labral tubercles ofoxaeids (Rozen, 1964: figs. HEAD (figs. 7, 8): Integument of head cap- 2, 3); labral apex nearly straight or shallowly sule without long setae but with scattered sen- emarginate, not cleft or deeply, narrowly silla some of which are bristlelike when seen emarginate as in Oxaeidae; median part of under high magnification; head integument labrum bearing very fine setiform spicules; of postdefecating larva moderately pigment- sides of labrum as well as epipharyngeal sur- ed except following more darkly pigmented: face with pronounced, nonsetiform spicules; clypeus, labrum, mandibles, mandibular co- epipharynx vaguely produced medially and ria, and apices of labiomaxillary parts; that without large median preoral lobe as found of predefecating larva faintly pigmented ex- in Oxaeidae. cept for more darkly pigmented mandibles Mandibular base robust; mandibular apex (but not mandibular coria), their apices very (figs. 10-13) slender, curved, simple (i.e., darkly pigmented. ending in single point), its inner surface di- Head size of postdefecating larva (figs. 4, rected adoroventrally (contrasting with slen- 5) small by comparison with rest of body. der but straight bladelike mandibular apex of Vertex (fig. 8) not greatly produced on each oxaeids, the inner surface ofwhich is directed side above antenna so that in profile top of ventrally while the upper serrated edge is di- head sloping in contrast to that of Andrena rected adorally); upper apical edge distinctly accepta Viereck (Rozen, 1973: fig. 6); front- serrate; lower apical margin smooth or with to-back length of head capsule short com- a few irregularities; cusp strongly developed pared to height as is characteristic ofandren- into acute, toothlike projection; cuspal sur- ids, not elongate as in oxaeids (Rozen, 1964: face and inner apical concavity strongly, fig. 3). Tentorium of predefecating larva evenly denticulate; dorsal surface bearing moderately developed (i.e., not robust) but spicules; outer surface with pronounced complete including dorsal arms; anterior ten- swelling bearing numerous bristles. Labio- torial pits moderately low on face; posterior maxillary region (fig. 8) greatly recessed, ex- tentorial pits below junction of postocciptial tensively fused basally. Maxilla distinct api- ridge (i.e., posterior thickening of head cally, its apex produced mesially and with capsule6) and hypostomal ridges, connected some spicules; galea not evident; palpus to postoccipital thickening by mesoventral moderately large, about as long as basal di- extension of thickening, this condition sim- thickening of the head capsule, it is the postoccipital 6 Although this ridge, arching dorsally over the fora- ridge, identified as such by the posterior tentorial pits at men magnum, has usually been called the posterior it lower extremities. 12 AMERICAN MUSEUM NOVITATES NO. 3060 ameter, rounded apically; cardo and stipes posteriorly somewhat farther than dorsum. perhaps visible as faintly pigmented areas but Anus a transverse slit positioned apically not as sclerites. (rather than dorsally or ventrally), with faint Labium (figs. 8, 9) inconspicuously divided but distinct lips. Spiracles moderate in size, into prementum and postmentum, distinct on distinct sclerites forming top of spiracular apically; labial palpus rounded apically, sim- tubercles; those of abdominal segment VIII ilar in appearance to, but somewhat smaller slightly smaller than others; peritreme flat; than, maxillary palpus. Salivary opening ap- atrium not projecting beyond spiracular pearing as a curved slit but in actuality open- sclerite; atrial wall without denticles or teeth ing essentially a small transverse slit beneath but finely ringed; primary tracheal opening small median swelling below hypopharyngeal with collar, circular (i.e., not slitlike as in groove, so that folds of integument around Oxaeidae, Rozen, 1964: figs. 5, 6); subatrium this swelling give appearance of curved slit consisting of approximately eight chambers. (in Andrena accepta, salivary opening similar Male postdefecating larva ventrally bearing except labium slightly more projecting on each short, transverse, median cuticular invagi- side of opening so that curvature of folds nation toward rear ofabdominal segment IX; seems more pronounced; in Protoxaea glo- female postdefecating larva ventrally with riosa, opening also similar except swelling pairs of paramedian cuticular scars on ab- not bordered above by hypopharyngeal dominal segments VII, VIII, IX, each pair groove which is absent medially in all Ox- closer together than proceeding one. aeidae and integumental folds around swell- MATERIAL STUDIED: 28 mature larvae, 6 ing pronounced only below). Hypopharynx a km south of Vicuiia, Elqui Province, Chile, single lobe (not bilobed) with surface spicu- October 17, 18, and November 16, 1991 (J. late laterally and adorally, similar to that of G. Rozen, A. Ugarte). Andrena accepta; hypopharmgeal groove deep, clearly separating hypopharynx from base of labium. PHYLOGENETIC RELATIONSHIPS BODY: Integument of postdefecating larva BASED ON BIOLOGIES AND rigid, thick, pigmented, of predefecating lar- MATURE LARVAE va normally soft, thin, unpigmented; integ- ument without setae but with scattered sen- BIOLOGIES: Data presented here are not re- silla particularly noticeable below anus and vealing in sorting out the affinities of Eu- at summits ofbody segments; some areas mi- herbstia with other short-tongued bees be- nutely spiculate; postdefecating larva with cause most aspects of nesting biology appear low, rounded spiracular tubercles that are to be plesiomorphic or not significantly dif- sclerotized and pigmented; predefecating lar- ferent from the nesting biologies ofthe other vae with these tubercles apparently sclero- taxa. These aspects include nest configura- tized but unpigmented; atrial walls of both tion, cell orientation and shape, spiral cell predefecating and postdefecating larvae closure, appearance ofcell linings, shape and darkly pigmented. Body (figs. 4, 5) without position of provisions, and lack of cocoons. dorsolateral tubercles; mesothorax and meta- Euherbstia, the other andrenines, and the thorax dorsally divided into cephalic and Stenotritidae differ from the Oxaeidae in that caudal annulets; abdominal segments at most the latter has vertical cells (also true for some indistinctly so divided; body without tuber- Andrena) and provisions that are semiliquid cles except for those supporting spiracles (figs. and unshaped. These features presumably are 4-6); intersegmental lines deeply incised on autapomorphies of the oxaeids, and do not postdefecating larva except between abdom- illuminate the affinities of that family to the inal segments VIII and IX, and IX and X (fig. other taxa. 4); venter of abdominal segment IX not pro- MATURE LARVAE: Because ofthe uncertain duced; segment X attached to IX centrally; relationships of the involved taxa to one an- segment X without dorsal ridges or plates; other, a phylogenetic analysis was attempted venter of abdominal segment X projecting to determine the interrelationships of Eu- 1993 ROZEN: EUHERBSTIA 13 TABLE 1 TABLE 1-(Continued) Explanation of Characters of Mature Larvae of Euherbstia, Andrena, Panurginae (Protandrena), veloped antennal prominences are uncom- Oxaeidae, and Stenotritidae Used in Phylogenetic mon in bee larvae, their presence was consid- Analysis ered apomorphic. (Plesiomorphy = 0; apomorphy = 1, 2) 5. Labral tubercles low, rounded, positioned ap- icolaterally on labrum (0); acutely pointed, 0. Vertex in lateral profile evenly curved (fig. 8) arising abruptly from labral disc (Rozen, 1970: (0); strongly angulate (Rozen, 1970: fig. 14) or fig. 10) (1). Larvae of many different groups even bulbous above each antenna (1). The of bees (e.g., Melittidae, Colletidae, Halicti- plesiomorphic condition is found widely dae, Anthophoridae) characteristically have among bee families; the apomorphic condi- rounded apicolateral labral swelling bearing tion is restricted to Andrena and the primitive sensilla, so that this condition is thought to Panurginae. A reversal of the apomorphic be plesiomorphic. Distinct, sharp-pointed tu- condition occurs in the higher panurgines, but bercles on the labral disc presumably arose the plesiomorphic condition exists in Protan- independently in the Panurginae, Oxaeidae, drena and such other primitive members of Euryglossa (Colletidae), and Nomadinae. the subfamily as Liphanthus, Pseudopanur- 6. Labral apex entire or weakly broadly emar- gus, Pterosarus, Metapsaenythia, Rhophitu- ginate medially (fig. 7) (0); deeply cleft me- lus, Psaenythia, and Anthemurgus. dially (Rozen, 1964: fig. 2) (1). The latter is a 1. Front-to-back head length normally short (fig. unique apomorphy of the Oxaeidae. 8) (0); elongate (Rozen, 1964: fig. 4) (1). Among 7. Labrum nonspiculate medially on dorsal sur- short-tongued bees, the apomorphic condi- face (Rozen, 1970: 13) (0); bearing fine spic- tion is found only in the Oxaeidae. ules medially (Rozen, 1973: fig. 5) (1). Fine 2. Posterior tentorial pit at junction of postoc- spiculation on the dorsal surface ofthe labrum cipital ridge and hypostomal ridge as seen in is an uncommon phenomenon among bees lateral view (Rozen, 1970: fig. 14) (0); below and hence was judged apomorphic. junction of postoccipital ridge and hyposto- 8. Epipharynx without basal swelling (0); with mal ridge (Rozen, 1964: fig. 3) (in Euherbstia, distinct basal swelling (Rozen, 1964: fig. 4) this condition is best seen in ventrolateral view (1). The latter is a unique apomorphy of the rather than lateral view, fig. 8) (1). Among Oxaeidae. short-tongued bees, the apomorphic condi- 9. Outer surface of mandible with conspicuous tion is found only in Oxaeidae and Euherbstia tubercle which in some cases extends across and would seem to be a strong synapomorphy. dorsal surface of mandible forming a distinct However, there may be a correlation between line distally (figs. 10, 12, 13) (0); without a 1) the position of the pits relative to the hy- tubercle or with very small one (Rozen, 1973: popharyngeal ridge and 2) the fusion of the figs. 8, 10) (1). The polarity ofthese character elements of the labiomaxillary region (char- states is problematical. At one time, the large acter 13) and reduction of the maxillary scle- tubercle on the outer surface of the mandible rites (character 15).a It is unknown whether appeared to be an autapomorphy of the Hal- there is a functional relationship between these ictidae (Michener, 1953; McGinley, 1987). features, but the weight to be placed on the However, the presence of this feature in Eu- position of the pits as a synapomorphy is re- herbstia, the Oxaeidae (though somewhat re- duced. duced), and in Ctenocolletes but not Steno- 3. Epistomal ridge between anterior tentorial pits tritus in the Stenotritidae (hence coded "?" in complete (fig. 7) or nearly so (0); absent be- table 2), as well as in most but not all halictids tween pits (Rozen, 1973: fig. 5) (1). Multiple suggests that it is a primitive feature that has losses ofthe midsection ofthe epistomal ridge been lost in numerous different lines. seem more likely than the midsection evolv- 10. Mandibular apex normal so that inner apical ing de novo time and again. Hence the pres- surface adoral (figs. 10-12) (0); bladelike, ro- ence of the midsection was considered ple- tated so that dorsal apical edge adoral (Rozen, siomorphic. 1964: figs. 10-12) (1). The latter state is found 4. Antennal prominence weak (fig. 8) (0); strong only in the Oxaeidae. (Rozen, 1973: fig. 6) (1). Because strongly de- 11. Mandible with subapical tooth (Rozen, 1970: figs. 11, 12) (0); without subapical tooth (figs. 10-12) (1). The polarity of this character is aA appears to similar correlation exist in the Noma- uncertain. The inner apical dinae among the long-tongued bees (Rozen, 1966). angle of the sem- 14 AMERICAN MUSEUM NOVITATES NO. 3060 TABLE 1 -(Continued) TABLE 1 -(Continued) itruncated mandibular apex (Rozen, 1964: figs. herbstia), and its anatomical structure and 10, 12) of the Oxaeidae is tentatively inter- functional basis are not understood at this preted as representing a subapical tooth. The time. Its significance as a synapomorphy is differently shaped mandibular apex in the questionable. Stenotritidae makes interpretation ofits char- 18. Hypopharyngeal groove deep (fig. 7) (0); ab- acter state uncertain (coded "?" in table 2). sent (Rozen, 1964: fig. 2) (1). This groove is 12. Mandibular cusp without strong projection present in most groups ofbees, suggesting that (Rozen, 1973: figs. 8-10) (0); with strong pro- it is plesiomorphic; rarely is it absent. Among jection (figs. 10-13) (1). The latter condition the groups being considered, it is an autapo- in Euherbstia may or may not be homologous morphy of the Oxaeidae. with a similar condition in Ctenocolletes and 19. Spiracles not on tubercles (Rozen, 1973: fig. perhaps Stenotritus. It is probably not ho- 3) (0); on tubercles that on postdefecating lar- mologous with the cuspal projection of Mel- vae become pigmented and sclerotized (figs. itturga (Rozen, 1965: figs. 8-13). 3-6) (1). Because sclerotized spiracular tuber- 13. Labiomaxillary region produced (0); recessed cles on larvae of other short-tongued bees are (Rozen, 1973: fig. 6) (1); greatly recessed (fig. unknown, their presence is considered apo- 8; Rozen, 1964: fig. 3) (2). None of the taxa morphic. under consideration spins a cocoon, and 20. Dorsolateral body tubercles either low and therefore their larvae have more or less re- transverse or absent (figs. 4, 5) (0); present on cessed labiomaxillary regions. As discussed most segments as distinct transverse tuber- under character 2, the apomorphic states of cles, which arise from caudal annulets (Rozen, this character may be linked with the apo- 1973: fig. 1) (1); present on most segments as morphic states of characters 2 and 15. The conical tubercles (caudal-cephalic annula- plesiomorphic condition is characteristic of tions not clearly defined on abdominal seg- cocoon-spinning larvae. ments) (Rozen, 1970: fig. 9) (2). In many 14. Maxilla spiculate at least apically (Rozen, groups of bees, caudal body annulets are el- 1973: fig. 5) (0); completely nonspiculate evated dorsally and in some cases these ele- (Rozen 1964: figs. 2-4) (1). Although the po- vations appear as low transverse tubercles. larity ofthis feature is uncertain, it seems more Hence this condition appears to be plesiom- likely that spiculation is lost rather than re- orphic. Among the taxa under consideration gained. The latter is an autapomorphy of the distinct, well-defined transverse tubercles are Oxaeidae. found in only Andrena (although this condi- 15. Cardo and stipes well sclerotized (0); evident tion has arisen presumably de novo in such though weak (1); faint so as to be virtually groups as the Halictinae and Nomiinae). Con- nonexistent (fig. 8) (2). Strongly developed ical tubercles are far less common and among maxillary sclerites are a primitive wasplike short-tongued bees occur (presumably as sep- feature; the near loss ofany suggestion ofthem arate evolutionary events) only in the Pan- in Euherbstia and the Oxaeidae is derived. urginae and Rophitinae. This character is However, because such evolutionary losses treated as additive, the assumption being that might well happen independently, they should pronounced transverse tubercles are a pre- not be considered a strong synapomorphy. cursor to conical tubercles. 16. Labium divided into a well-defined premen- 21. Spiracular atrium moderately or strongly pro- tum and postmentum (0): weakly divided into duced above body wall (Rozen, 1973: fig. 7; prementum and postmentum (figs. 8, 9) (1); 1964: figs. 5, 6) (0); not produced above body not divided (Rozen, 1964: fig. 3) (2). The ple- wall (figs. 6, 14) (1). The latter appears to be siomorphic condition is characteristic of co- an autapomorphy of Euherbstia. coon-spinning bee larvae. 22. Atrial wall smooth (Rozen, 1973: fig. 7) (0); 17. Salivary opening not associated with a curved ridged (fig. 14) (1); spinose (Rozen, 1964: figs. groove descending from hypopharyngeal 5, 6) (2). Because both smooth and orna- groove (0); opening at lower extremity of mented atrial walls are found in many bee curved fold or groove extending either from groups, the polarity of these states is uncer- hypopharyngeal groove or, in Oxaeidae which tain, and that character was coded nonaddi- has no such groove, from hypopharyngealV tive. labial surface above (1). The apomorphic con- 23. Primary tracheal opening circular (fig. 14) (0); dition exhibits considerable variability (as in- slitlike (Rozen, 1964: figs. 5, 6) (1). The latter dicated in the description of the larva of Eu- condition is unique to the Oxaeidae. 1993 ROZEN: EUHERBSTIA 15 TABLE 2 dae are sister groups and that they in turn are Data Matrix for Analysis of Relationships of Eu- a sister group ofAndrena and the Panurginae herbstia, Andrena, Panurginae (Protandrena), Ox- (as represented by Protandrena). The mono- aeidae, and Stenotritidae on Basis of Larval Char- phyly of the Andrena-Panurginae-Euherbs- acteristics tia-Oxaeidae clade is supported by a single (Character codings given in table 1) synapomorphy (17), the presence ofa curved Character states groove in which is found the salivary opening identified and used 11111 11111 2222 (a character first by Taxon 01234 56789 01234 56789 0123 McGinley, 1981). As stated in table 1, this character state exhibits considerable vari- Ancestor 00000 00000 0?000 00000 00?0 ability, and its anatomical structure and func- Euherbstia 00100 00100 01120 21101 0110 tional basis are not understood at this time. Andrena 10011 00101 01010 11100 1000 Protandrena 10011 10001 00010 12100 2000 These facts suggest that it may not be reliable. Oxaeidae 01100 11010 10021 22111 0021 Therefore, the analysis was performed again, Stenotritidae 00000 0000? 0? 110 12000 0000 with this feature coded inactive. Three most parsimonious cladograms resulted (length 27, ci 77, ri 60): herbstia excellens, Andrena, the panurgines, 1. (Ancestor + (Euherbstia + Oxaeidae) oxaeids, and stenotritids based on mature lar- + (Stenotritidae + (Andrena + vae. Footnote 4 lists the species of Andrena Protandrena))). whose larvae were available. The Panurginae 2. (Ancestor + (Andrena + Protandrena) are a large diverse group so that the larva of + (Stenotritidae + (Euherbstia + one exemplar, Protandrena verbesinae (Tim- Oxaeidae))). berlake),7 was examined because of its large 3. (Ancestor + Stenotritidae + ((Andrena size and because the genus is relatively prim- + Protandrena) + (Euherbstia + itive within the Panurginae (Ruz, 1986) and, Oxaeidae))). therefore, its connections to nonpanurgine With the exclusion of character 17, the re- taxa might be more obvious than those of lationship of the Stenotritidae to the other highly specialized panurgines. In the Oxaei- taxa became ambiguous, but the sister-group dae, larvae ofProtoxaea gloriosa, Oxaeafla- linkage between Andrena and the Panurginae vescens, and Mesoxaea nigerrima were used, and between Euherbstia and the Oxaeidae as well as were larvae ofthe stenotritids Cten- ocolletes nicholsoni, C. ordensis, and Steno- persisted. Nonetheless, the synapomorphies tritus (2, 132, 152, and 19; 0, 3,4, 9, and 201) uniting greavesi. each of these two groups are not particularly Table 1 discusses the larval features used in the analysis, and table 2 is the character strong as explained in table 1. Furthermore, matrix. The the appearance de novo of the apomorphic analysis, using James S. Farris' states of characters 7 and 1 1 in Andrena and Hennig86 Version 1.5 program (Farris, 1988), Euherbstia raises the possibility that they was executed with the ie command. Auta- might actually be synapomorphies. pomorphies (1, 6, 8, 10, 14, 18, and 21) were In last analysis, I am not convinced of the coded inactive, and character 22 was coded phylogenetic relationships suggested by the nonadditive. The result was a single, most parsimonious cladogram (fig. 15), length 28, study of larval characters. At the same time, ci 78, ri 62. the hypothesized linkages may be accurate, and they do offer the following avenues of This tree (fig. 15) suggests that, on the basis of larval study: (1) Adult characters should be rein- features, Euherbstia and the Oxaei- vestigated to test the results of the present study. (2) Larvae of other andrenine taxa 7 The larva (and pupa) ofthis species was misidentified in Rozen (1970) and later in McGinley (1981) as Psae- (Megandrena, Ancylandrena, Alocandrena, nythia bicolor Timberlake. However, the species attri- and Orphana) need to be found and exam- bution to Protandrena bicolor (Timberlake) (as Psae- ined to see what bearing they have on the nythia biciolor Timberlake) in Rozen (1967) regarding proposed relationships. (3) Although biolog- nesting activity and nest architecture was correct. ical characteristics of Euherbstia and other 16 AMERICAN MUSEUM NOVITATES NO. 3060 12 162 Stenotritidae Andrena Protandrena Euherbstia 2 132 152 19 156 8 10 14 162 18 23 Oxaeidae Fig.15. The single most parsimonious cladogram ofEuherbstia, Andrena, Protandrena, the Oxaeidae, Stenotritidae, and a hypothetical ancestor, based on characters ofmature larvae. Solid squares are unique character states; open squares are parallel states. Character 11 assumes that the loss of the mandibular subapical tooth is apomorphic, and the undivided labium (character 16) is depicted to have evolved three times from a weakly divided one. All autapomorphies were coded inactive, and character 22 was excluded because its polarity was uncertain. For further explanation, see text. taxa have not been helpful in defining affin- Stenotritidae). Rec. West. Australian ities within this group, new life history fea- Mus. 11: 153-172. tures should be identified and additional taxa 1987. Fossil brood cells of stenotritid bees of Andreninae should be investigated bio- (Hymenoptera: Apoidea) from the in Pleistocene of South Australia. Trans. logically search ofuseful comparative data; R. Soc. South Australia 3: 93-97. REFERENCES Houston, T. F. and R. W. Thorp 1984. Bionomics ofthe bee Stenotritusgreave- Batra, S. W. T. si and ethological characteristics of 1990. Bionomics of a vernal solitary bee An- Stenotritidae (Hymenoptera). Rec. West drena (Scrapteropsis) alleghaniensis Australian Mus. 11: 375-386. Viereck in the Adirondacks ofNew York Hurd, P. D., Jr. (Hymenoptera: Andrenidae). J. Kansas 1979. In K. V. Krombein et al., Catalog of Entomol. Soc. 63: 260-266. Hymenoptera in America North of Farris, J. S. Mexico, 2: 1741-2209. Washington, 1988. Hennig86 Reference, Version 1.5. D.C.: Smithson. Inst. Press. Available from J. S. Farris. 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Phylogenetic-taxonomic significance of menoptera-Andrenidae). Ph.D. thesis, last instar of Protoxaea gloriosa Fox, Univ. Kansas, 312 pp. with descriptions offirst and last instars Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.