Observations on the Nesting Biology of Tetralonia ...

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 58(4), 1985, pp. 686-700

on the Nesting Biology of Tetralonia hamata with a Description of itsMature Larva

Observations Bradley

(Hymenoptera:

Anthophoridae)

Eugene R. Miliczky1 Section of Faunistic Surveys and Insect Identification, Illinois Natural History Illinois 61820 Survey, Champaign, abstract: lonia site

hamata in east

4 days.

on the bionomics results of field and laboratory observations are presented. Nests in June of 2 bees were discovered Bradley at their nests were central Illinois. Observations of female behavior

During

The

this

time

T.

hamata

females

were

observed

foraging only and cell structure

on

of Tetra 1982

at a

made

on

red clover

are described. Both nests were excavated and nest pratense). (Trifolium on larval development The mature larva and the structure of the is presented. Information cocoon are described and illustrated. Parasites of T. hamata were Meloe americanus (Co nomadine bee (Anthophoridae). and a parasitic of the biology Aspects le?ptera: Meloidae) of T. hamata

are compared

with

those

of other

known

eucerine

bees.

Members inhabit both the Old and New of the bee genus Tetralonia Spinola to large size bees occur in North America and 54 species of these medium Worlds Timberlake of Tetra (Hurd, 1979, as Synhalonia). (1969) stated that members are vernal, rarely found later than June, and visit a broad lonia (as Synhalonia) from composites. but rarely collect pollen range of plant families Although are large, conspicuous of the North American many insects, knowledge species of floral visitation Rust and Clement consists predominantly records. However, It (1977) reported briefly on a single nest of T hurdi (as Synhalonia hurdi). with pollen from Collinsia of 11 cells provisioned consisted (Scroph sparsiflora nest structure. Linsley et al. (1963) found but they did not describe ulariaceae) in the vicinity nests of T speciosa of its host plant, Oenothera (as S. speciosa) The nesting substrate was loose sand and nests reached (Onagraceae). pallida on female foraging behavior was given more 1 meter. Information than depths of of nest structure was not included. but a description have been studied in greater detail. Malyshev A number of Old World Tetralonia on a central and southern European T malvae, the of life history (1930) reported on floral rela He information sites, adult phenology, nesting species. presented a detailed immature and structure and cell gave stages, provisions, tionships, account and structure. A second species, T dentata, was of nest construction infor discussed (1969) presented briefly and earlier literature reviewed. Rozen on the nests, cells, and immatures and a nomadine of T. minuta mation parasite. of South Africa. Finally, The bees were nesting in a grassland area in the Republic and Mohamed Mohamed (1970) studied the bionomics (1973, 1974) and Wafa of broad bean (Vicia in Egypt where it is an important pollinator of T. lanuginosa nest These authors provide data on nest site characteristics, faba, Leguminosae). and cell structure, and the life cycle of T. lanuginosa. on the biology of Tetralonia The purpose of this paper is to present information

1 Present Accepted

address:

27576

for publication

Anchor

Place

10 October

NW, 1984.

Poulsbo,

Washington

98370.

This content downloaded from 35.10.204.136 on Tue, 10 Jun 2014 14:23:00 PM All use subject to JSTOR Terms and Conditions

687

VOLUME 58, NUMBER 4

at the nesting site, hamata gathered during 1982. It is based on field observations excavation observations of immature of 2 nests, and laboratory stages. Infor on adult female behavior, nest and cell structure, behavior and devel mation mature of this bee is The and the larva of opment immatures, presented. parasites and illustrated. is described Study

Site and Methods

Forest Preserve Tetralonia hamata was studied at the Piatt County (Lodge west The nests were near 2.5 Piatt Illinois. miles of White Heath, County, Park), the western end of a long, narrow field about 10 m by 100 m whose long axis ran car path ran the length of the field. The car path was east-west. A little-used mowed but at the time of the study it had not been recently cut and periodically some vegetation had reached a height of 0.5 m or more. Both nests were located between the tire tracks of the car path. The field was grown over with various grasses and forbs and there were no bare areas of significant size. The northern edge of the field was bordered by a wild rose thicket (Rosa sp.) beyond which was a plantation of pines. South of the field were patches of wild rose, blackberry (Rubus sp.), and sumac (Rhus typhina), as well as scattered trees. Farther south the ground was low and moist with vegetation trees and shrubs. of deciduous primarily consisting seen entering when the female bees were their Both nests were discovered was found on 18 June, #2 burrows after returning from foraging trips. Nest #1 on 21 June 1982. Observations at one or both of female behavior were made nests on 18, 21, 22, and 23 June during part or all of each day and were recorded on cassette tapes. A watch with a sweep second hand was used to time bee activities to the nearest 5 second interval. Air temperature about 0.3 m above ground in the bee under observation after left the sun was taken periodically, usually just at the bees which her nest on a foraging trip. This was the approximate height on general weather conditions foraged on red clover (Trifolium pratense). Notes were made when the temperature was taken. became Both nests were excavated after their occupants inactive. A block of all the cells from nest #1 was brought into the laboratory soil containing intact were made on the to allow more accurate measurement. Detailed observations of 3 larvae from nest #2, 1 of which was followed from egg hatch development to post defecating larva. Less detailed notes were made on the larvae from nest #1. All larvae were held at room temperature (ca. 22?C) during these observations. Four larvae (in their cells) from nest # 1were held at room temperature throughout 4 larvae (also in their cells) from nest #2 were overwintered the winter while in a covered, 15 cm below ground outside (to simulate natural glass jar about to determine probable stage. conditions) overwintering Results active season: over most Tetralonia hamata is distributed of the eastern In east United States and as far west as Colorado central Illinois (Hurd, 1979). the species is active principally and 74 female and June. during May Forty-five area in male the Natural from this Illinois collection specimens History Survey bear collection dates ranging from 11 May (1911) to 13 July (1924) for females and 11May (1911) to 15 June (1936) for males (1 male was labelled July 1902).


688

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

I first observed this species at Piatt County Forest Preserve on 29 May 1980 and saw at that time I only males but on 3 June both sexes were active. On my first on that day visit in 1982 (13 June) I saw only females. The specimen collected had the entire apical wing margins frayed, indicating she had been active for some here reported on died of natural causes between time. Both individuals 21 and 30 June. Thus it appears that this species was active from the latter half of May until the end of June in 1982. flower preferences: is polylectic Tetralonia hamata and has been recorded in 67 36 of families families are genera plants visiting (Hurd, 1979). Twenty-four a 7 2 families and the 5 families genera, represented by single genus, by remaining are represented 4 3 5 6 and by (Cruciferae), (Compositae), (Labiatae), (Rosaceae), was II (Leguminosae) T this hamata observed genera. During study foraging the most abundant plant in bloom at the only on red clover (Trifolium pratense), as a time. Folsom (1922) reported on the efficiency of T hamata (as T. dilecta) were in of red clover. Other bloom the repens, pollinator plants legumes Trifolium T. hybridum, Melilotus and the composites Rudbeckia alba, and M. officinalis hirta and Erigeron sp. The abundant wild rose (see above) was no longer in bloom by the time of the study. bees: Four days of observations activity of female at the 2 nests indicated individual bees may forage continuously that during suitable weather for the entire were similar on each of the 4 days and ranged from 21?C day. Air temperatures to 23?C at the start of observations (ca. 0900) and reached daily highs of 28?C to 29?C. June 18 was mostly cloudy, the other 3 days were largely sunny, and winds were light all 4 days. 7 foraging trips between On 18 June female #1 made 1140 and 1530 and had were terminated. she collected begun another trip when observations Presumably in well with the pollen color blended the pale, pollen on each trip. However, to verify its presence in ochraceous, scopal hairs and it was not always possible the scopa during the brief period of time between first sighting the bee and when she entered the nest. at nest #1 were resumed on 21 June at 0900. This bee was first Observations seen at 0934 just below her nest entrance when air temperature in the sunlight was 23.5?C but the nest itself was partly shaded by vegetation. Female #1 left her nest #2 and between nest at 0943:30 on her first foraging trip. At 1043 I discovered 9 foraging trips and female #2 made 8. this time and 1500 female #1 made from 0915 to 1640. The bee On 22 June nest #1 was observed continuously were terminated on her first foraging trip at 0939:30 and observations departed she began her thirteenth for trip at 1637:40. She thus foraged continuously than 7 hr. to observe nest #2 for the entire day commencing at On 23 June I intended to nest #1. 0900. By 1020 female #2 had not appeared and I shifted observation Female #1 returned from a foraging trip at 1047:15 but the number of previous the last at 1656:30. 9 subsequent She made trips completing trips is unknown. not finished for the day. She had begun another trip by 1715 and had presumably was 31 min 56 sec #1 timed female The mean 33 for foraging trips length of with a range of 22 min 30 sec to 51 min 45 sec. Mean trip length for female #2 was 28 min 24 sec (n = 7) with a range of 23 min 55 sec to 36 min 55 sec. Time 10 sec (n = 33) for female spent in the nest between foraging trips averaged 4 min

after more


689

VOLUME 58, NUMBER 4 shortest was 2 min and her than any other. The mean for longer 15 sec. 15 sec to 6 min of 4 min #1 was observed Female leaving she made an orientation departures #1. Her

than 3 min longest, 9 min 20 sec, was more 10 sec (n = 8) with a range bee #2 was 5 min

her nest on foraging trips 38 times. On 7 flight, 2 of which were quite lengthy (ca. 5 the nest or its immediate 10 sec). Both of these occurred after I had disturbed The other 5 orientation flights were shorter (less than 5 sec). Usually vicinity. her female #Ts departures were as follows: On several occasions when I watched cm away a series of short, sharp buzzes could be heard leave from only 25-30 the nest. The buzzes began 2 to 3 min after her return from within emanating 45 sec later she appeared near the entrance and from the previous trip. About a time, occasionally at to 1-2 her head from side crawl out, steps began moving to side as if "looking around". Finally she emerged completely, took flight, and a partial circle of the nest vicinity. flew off down the car path often after making When leaving the nest for the first time on a given day the length of time between at the nest entrance and departure first appearance is probably more protracted as it was on 21 June (see above). The returns of female #1 to the nest were swift and direct. She suddenly appeared in the vicinity of the nest, quickly above down near the entrance, and crawled inside. She it, dropped to have she from this 6 times. Twice pattern only departed noticeably appeared a some difficulty entrance nest the after disturbance of the locating vicinity. The other 4 times she flew into the vicinity of the nest and landed on the ground or some vegetation. a minute, to the nest and Within she flew directly however, generally localized

entered. Female #2 differed slightly in her entering and departure behavior. On each of her 8 observed departures she made an orientation flight (less than 5 sec or not nest whether the had been When disturbed. duration) returning she usually hovered for 2-3 sec about 0.3 m above the nest before landing and entering. nest location and structure: Both nests were dug in level ground amid and were very effectively The nests were more concealed. than 15 m vegetation of apart. The soil, when dry, was light tan in color and consisted predominantly silt and clay-sized particles. When the nests were excavated the soil was moderately moist from the surface to below the level of the cells and was sufficiently compacted to make possible the removal of a large block containing all the cells in nest #1. A thin layer of organic matter was present on the surface and a rather dense growth in the top 2-4 cm, some of which grew through the upper parts of roots occurred of the burrows. Below this level the soil was largely free of roots, stones, and other debris. the remnants of tumuli, surrounded both nest entrances but Soil, apparently at had been compacted rain the time of the this study both by study. Throughout entrances were open with no evidence that the burrows were closed for the night. The structure of the open part of the burrow was similar in each nest (Figs. 1, 2). After penetrating the soil vertically for a short distance each burrow bent abruptly and gave rise to a short, horizontal section in nest #1 (Fig. 2) and a short, obliquely in nest #2 (Fig. 1). Both burrows section then bent vertically and descending to depths of about 20 cm with little meandering descended before they began to bend toward the horizontal. After levelling off and reaching maximum depths of 23 cm (#1) and 25 cm (#2) both burrows ascended #1 at a Nest ended slightly. cell but the cell and final 2 cm of the burrow had been filled with soil (the cell



690

Ob aO

n^r

eO

1-3. Figs. most recently view showing filled

d?? f0Ok Oh cO

? Letters

09 JO

Nests

cells.

correspond section of main purposes.

hamata. 2. Nest

1. Nest #1. Vertical

#2. Vertical

section

and 2 burrow through main #1. Plan burrow. 3. Nest

through main to lower, horizontal

section

burrow. section of main of cells and their relationships as explained in text. Dashed soil lines in Figs. 2, 3 indicate terminal, same point in burrow cell. Point x in Figs. 2, 3 designates burrow and aborted Scale (= 1 cm) is the same for all figures.

arrangement

a-k

for reference

of Tetralonia

constructed

to cells

having apparently been aborted for some reason). Female #1 was found dead near a of her nest. Nest #2 ended in an open, unprovisioned the bottom cell while an egg, was located about 2 cm away at a slightly greater second cell, containing a large #2 was also found dead in her burrow and in addition depth. Female carabid beetle was found in the lower section of the nest. The beetle was alive that it had killed or fed upon the bee. but there was no indication in diameter 8-9 mm their Both burrows were roughly circular, throughout at their entrances. Some sections of the burrow lengths, and were not constricted marks made by the mandibles walls showed distinct grooves, presumably during smoother The remainder of the walls had somewhat excavation. surfaces, possibly of the bees. having been worn down by the movements In nest #1 (Fig. 3) the 2 most recently provisioned Both nests were multicellular. cells (a, b) were about 1 cm apart but were well separated (by about 20 cm) from a second group of 9 cells. Six cells (c-h) of the second group contained older larvae of the soil showed that but the other 3 cells (i, j, k) were filled with soil. Removal these cells were in good condition with their waterproof linings intact. No remains near the bottom of each of cocoons or other signs of bees were present. However, in diameter, cell was an earth-filled hole, 5mm or more indicating that something 1982 an adult male Meloe 10 November had dug its way out of each cell. About det. J. K. Bouseman) Leach (Cole?ptera: Meloidae, americanus emerged from the cells c-k. Since the contents of cells c-h were accounted block of earth containing on the contents of 1 or more of cells i, j, for, the Meloe developed presumably


691

VOLUME 58, NUMBER 4

Figs.

4, 5.

egg placement Letters section.

Cell

cocoon

and

and

of Tetralonia

of provisions. to the following:

level

hamata.

5. Cocoon

section Vertical, showing longitudinal structures. associated Vertical, longitudinal to feces and cocoon below lining adheres

4. Cell. and

B?cell A?feces, E? line indicates lower limit of fecal material, D?dashed layer of cocoon, upper region of inner layer of cocoon, upper region of outer layer of cocoon, F?thickened, thickened, in are sandwiched between cell lining and outer layer of cocoon G?outer layer of cocoon, H?feces refers to both figures. this region. Scale (=2 mm) dashed

refer

line, C?inner

and Pinto and Selander and k (see also section on parasites). (1970) and Mayer or more 2 will bee that o? Meloe utilize various Johansen species (1978) report larvae and/or their provisions during development. to be determined of nest #2 did not allow the cell arrangement My excavation as that of nest #1. However, as accurately it appeared to be similar. The 2 newest cells (shown in Fig. 1)were close together. Well separated from these 2 cells were in a block of soil about the size the other 6 cells in the nest. They were contained to build cells in the of a fist. Had this bee lived longer she may have continued ones. newest 2 of the vicinity sec in longitudinal and were elongate-ovoid Cells were oriented subvertically tion (Fig. 4) and circular in cross section. They were wider in the lower half, with = a mean maximum the upper half tapered to of 9.7 mm diameter (n 7), while = 18.4 a mean diameter Cell at the cell closure of 6.8 mm (n length averaged 5). mm (n = 4) from the lower end to the level of the juncture between cell wall and cell closure. The walls were very smooth, slightly shiny, and lined with a thin, to the cell wall beaded up and 45 A water of applied drop waterproof coating. of a spiral arrangement min later it was not absorbed. The cell closure consisted of soil particles with about 4 rows to the cell radius and was concave on the side = facing the cell. Cell depth from ground level to cell closure averaged 24.2 cm (n


692


10; range 22-27 cm). After cell closure an unknown length of the burrow leading to it was backfilled traced. tightly with soil so that its course could not be precisely due to the considerable distance between the cells and the vertical part However, of the main burrow and the close association of the cells, it seems unlikely that a separate lateral was constructed from the vertical part of the burrow to each of a long lateral was excavated the cells. Possibly at its and the first cell constructed end. After provisioning the first cell, the bee may have started digging in a different a short distance back from it using the loosened direction soil to close the cell at the end of and backfill the lateral. The next cell would have been constructed the newly excavated section of the burrow. Such a behavior would result in a tight the time spent in excavation and backfilling. group of cells and would minimize observations on immature stages: The amber to dirty orange provisions the lower lh (6-7 mm) of the cell (Fig. 4). They had a faint, sourish odor occupied detectable from a short distance examined away (5-6 cm). One provision mass, surface. A free layer of liquid prior to egg hatch, had a smooth, gently concave was not present but the provision mass was quite moist and its concave, meniscus like surface may have been due to a high nectar :pollen ratio. The white, gently end curved, 4 mm long egg was placed atop the provision mass with its posterior near the cell wall and the slightly broader anterior end near the cell's center. The concave so that only its ends contacted side of the egg faced downward the mass was as set in The rather the it survived egg place provision firmly (Fig. 4). and transport. of excavation rough handling The succeeding notes on larval development follow the progress of larva A (nest to the form. from the egg #2) They are supplemented by obser postdefecating on 2 other larvae (B, C) from the same nest which vations eclosed prior to and indicated 4 larval instars as did those shortly after excavation. My observations Dr. Jerome and Mohamed for Tetralonia of Wafa (1970) lanuginosa. However, true I have overlooked the first instar indicated that G. Rozen may (pers. comm.) o. obliqua was to reference made larva. In his paper on Svastra (Rozen, 1964) was which of short its the larviform duration, embryo (and embryonic cuticle) this to be the true first larval the chorion. He now believes living largely within stadium and S. o. obliqua therefore, has 5 instars. If this is also true for Tetralonia the first instar it should be born in mind hamata that, in the following discussion, in fact be the second, etc. Few, if any, other eucerine bees have had their would the number of instars. studied in sufficient detail to determine larval development in Coelioxys 5 larval instars were shown to be present However, sayi and C. He that Baker found the first octodentata instar, which (1971). by (Megachilidae) mandibles of the second and third stadia, had been lacks the huge, sickle-shaped to short duration, and the fact due its delicacy, overlooked by previous workers the chorion. within that it remains almost completely on 21 or 22 June, with oviposition Cell A was probably shortly provisioned on 21 June but inactive was #2 because female thereafter, actively provisioning 2 days later (23 June) and was found dead in her nest on 24 June. The cell was on 25 June with its provisions and egg intact. On 26 June several small discovered not that had been beads of moisture present the previous day were noted (10-12) on the surface of the egg. These were perhaps an indication of the initial stages on 27 time 1400 had hatch June of eclosi?n. by which definitely begun by Egg of contractions the anterior end of the chorion had split open. Slight muscular


VOLUME 58, NUMBER 4

693

the larva's anterior end were visible, perhaps representing its efforts to shed the of the chorion. By 0900 on 28 June eclosi?n was complete remainder and the in the provisions. It was larva was lying with the right half of its body submerged near was and the with cell wall its curled body slightly positioned actively feeding. As it fed the larva swung its anterior end dorso-ventrally and from side to side its anterior end in the provisions) while open (sometimes completely submerging bidentate mandibles. 1615 the larva's ing and closing the lightly sclerotized, By surface appeared position had changed slightly and while most of the provision's an area of liquid with a number of small, from its original condition, unchanged to the larva. Similar areas formed air bubbles had formed adjacent suspended, near larvae B and C as well. Larva A molted to the second instar prior to 0830 on 29 June and at that time was lying with its dorsal surface closely paralleling, the cell wall. It was pointed counterclockwise but not touching, and still had the in the provisions. in this orientation, It remained right half of the body submerged as did larvae B and C, almost to the completion of feeding. About lh of the surface mass was very watery in appearance of the provision the by this time while as de remainder was undisturbed. The larva's feeding movements continued scribed above although pauses seemed more frequent. Gentle waves of muscular contractions and were probably in coursed along its body almost continuously in propelling or moving volved the larva through the provisions food through the the dorsal surface in gut or both. By 1540 on 29 June larva A was lying with contact with the cell wall. It had progressed around the circumference of the cell about 1.5 times and in the process had formed a channel in the provision mass to the cell wall. This channel was largely filled with adjacent liquid which was but free of in the region around the larva's suspended pollen except cloudy mostly head. The central part of the provision mass was not as deeply covered with liquid and formed a sort of "island". As the larva continued feeding and circling around in width and the central "island" decreased the cell the channel increased in size as the larva increased in girth. The larva had molted to the third instar by 0850 on 1 July and by 1300 on 3 July it had molted to the fourth, and final, instar. Increase in size was now rapid and the larva had nearly completed feeding by 6 was At this time the larva near with the oriented head and tail the cell bottom July. and the middle of the body looped up toward the top of the cell. The larva had completed feeding by 0920 on 7 July and had its head near the top of the cell, its tail near the bottom. All 3 larvae, after the completion of feeding, continued to scrape the mandibles this behavior against the cell walls for 1-2 days. Whether was to scrape up any food remnants or to perform some other task is unknown. The larvae did scrape areas of the cell wall that had never been in contact with the provisions. An interesting feature of late fourth instar larvae was their ability to change position in the cell. Such mobility is probably for efficient important and cocoon defecation spinning. By 9 July larva A was well into the defecation rather sticky or viscous, and dark olive green or process. Feces were quite moist, were in blackish color. about the xh of the cell as green upper They deposited masses. 2 Defecation about for each of the larvae and elongate required days some silk may have been spun before defecation was complete. The feeding pattern of Tetralonia hamata larvae agrees quite closely with that o. described by Rozen for Svastra (1964) obliqua and Wafa and Mohamed (1970) for T. lanuginosa. However, Wafa and Mohamed (1970) found that each instar


694


of T. lanuginosa lasted considerably longer than did those of T. hamata. Whereas the time elapsed between and hatch of feeding in T hamata was egg completion on the order of 10-12 days (for each of 3 larvae), the equivalent period for T was 7-12 around 35 each Time with instar spent in days lanuginosa lasting days. was and cocoon in T. defecation also much These spinning longer lanuginosa. in rearing conditions differences may have been due to differences (both sets of were conducted or adaptation to the different in the laboratory) observations in which and therefore different climatic the 2 locations, conditions, geographic species

occur.

as their cells had been of the above 3 larvae produced normal cocoons to permit observation. broken open to various degrees Five larvae, whose cells were undisturbed cocoons. normal The spun by excavation, general appearance and structure of these cocoons was as follows (Fig. 5). The cocoon was closely to the cell wall except near its upper end where a 1-3 mm gap separated appressed the cocoon from the cell closure. This space was largely filled with feces. Apart structure from its upper region the cocoon was a delicate, that de thin-walled formed under slight pressure. The upper region, however, was considerably thicker None

and tougher. to consist of 3 discrete The cocoon appeared layers. The outermost layer had a fine, non-fibrous texture, was tan or light brown in color, and did not cover the cell lining upper part of the structure. This was probably part of the waterproof to the cocoon proper to pull away from the cell walls which adhered sufficiently when the cocoon was removed from the cell. Thus this layer was a product of the adult female bee, not the larva. It was present on each of the 5 cocoons examined and fecal material was sandwiched between it and the upper region of the cocoon a similar situation for Svastra o. obliqua. (see Fig. 5). Rozen (1964) described The cocoon proper had an outer and inner layer. The outer layer consisted of a series of crisscrossing, thickness. The fibers light brown, silken fibers of variable were embedded transparent, in, or connected together, by a very thin, delicate, structure so that the outer layer of the cocoon was free of gaps. It membranous was sometimes to gently tease the outer layer into an undetermined possible thin and flexible except at number of sublayers. The outer layer was uniformly in thickness. At the upper end a series the upper end where it greatly increased and gave the structure a coarser and more rigid of layers was more apparent Fecal material often intruded into the free spaces of the outer parts composition. of the upper The inner

region. except at its upper end, was a single, homoge layer of the cocoon, neous, brown, translucent sheet, rather shiny on its inner surface. At magnification of a fibrous nature (except at the upper end) up to 50 x there was little evidence save for a faint streaking that was visible the sheet. The inner layer throughout was uniformly thin, except above where a roughly circular region, about 3 mm to as much as 0.5 mm (10 times or more the in diameter, increased in thickness thickness of the rest of the inner layer). This area lay below the thickened region of multiple of the outer layer and was likewise quite stiff. It consisted layers of fibers with the layers more closely appressed than those of coarse, crisscrossing the periphery of the circular region the the thick part of the outer layer. Toward fibers fused with the lower part of the inner layer and lost their separate identities. This would indicate that the inner layer was laid down as separate, silken strands


695

VOLUME 58, NUMBER 4

6. Postdefecating larva of Tetralonia hamata. Mature Figs. 6-12. frontal view. 8. Head, lateral view. 9-11. Right mandible, dorsal, ventral, Scale for Fig. 6 = 1mm.

7. Head, lateral view. larva, 12. Spiracle. and inner views.

which fuse together before they harden to form a homogeneous sheet, except at strands remain separate. the upper end where the individual stage: overwintering Four larvae from nest #1, held at room temperature 1male) by 15 from the time of nest excavation, reached adulthood (3 females, cold period is not required for trans that an obligatory 1983, indicating February to the adult. The 4 larvae (nest #2) overwintered outside were unearthed formation on 28 February 1983. The winter had been extremely mild up to this time. One an adult male but the other 3 contained cell contained These healthy prepupae. the prepupa may be the normal over results indicate that under field conditions stage but further replication would be desirable. wintering parasites: bees in the genera N?mada field observations parasitic During were and present in the area but were (Anthophoridae) Coelioxys (Megachilidae) not seen in the immediate nest. of either cell G from nest #1 However, vicinity the larva of a parasitic bee in the subfamily Nomadinae contained (Anthophor as a defecating larva when mold began to grow in the cell idae). It was preserved to have died. Using and it appeared the key to nomadine larvae in Rozen (1966) it has been tentatively identified as the genus Triepeolus.


696


As stated earlier, an adult male Meloe americanus emerged from nest #1 some In to 2 10 November time prior larvae were taken from 1982. Meloe addition, cells (b, d) in this nest. On 3 July a large larva was pulled from cell d. It had consumed the bee larva (a large fourth instar) and was digging its way out of the bottom of the cell. The second larva was medium sized and was found dead in in this cell but had begun cell b on 9 July. A good deal of provisions remained to mold. Thus I believe that 5 of 11 cells in this nest were probably destroyed by and a sixth by a parasitic bee. None cells from nest meloids of the 7 provisioned #2 was parasitized. description larva: of the mature larva of Tetralonia The mature hamata and spiracular structure and is of typical eucerine appearance. Head, mandibular, those form conform with of other known eucerines. quite closely general body The following description larvae the is based on 7 mature, and follows preserved format of Rozen (1965). head on maxillae with a few setae?primarily and (Figs. 7, 8): Integument dorsal labrum of of and and surfaces maxillae labium; epipharyngeal apical, apices remainder of head cap spiculate; mandibular apices darkly pigmented; minutely and hypostomal sule nearly unpigmented articulations except for mandibular are arms and and which Anterior tentorial posterior ridges lightly pigmented. dorsal arms rudimentary; tentorial bridge complete and well developed; posterior well and pleurostomal thickening of head capsule, hypostomal ridges moderately between but evident anterior tentorial epistomal pits developed; weakly ridge of head capsule well developed in median developed; longitudinal, thickening and not reaching epistomal half, weakening posterior anteriorly ridge; parietal bands weak. Antennal low, the papillae about as high as their basal prominences in well developed. Mandible diameter. Labral tubercles moderately (Figs. 9-11) dorsal and ventral views with robust, nearly square base and much narrowed, from base slightly bidentate, apical region; apical concavity with ridge extending more than 3Away to apex; dorsal plane of apical concavity narrow; a distinct, hairlike spicules elongate cusp ventrad of base of ridge; several dorsally projecting, outer in with 3 base of of of mandible dorsad present ridge; (aboral) edge region or 4 minute from base to apex, each tubercle bearing tubercles about lh distance an elongate seta. Maxillae sclerotic with subtruncate, adorally produced weakly of into mesad divided apices; galeae present elongate palpi. Labium projecting, more and the somewhat prementum prementum postmentum, sclerotic; salivary present as transverse, lips; apical slit with well developed, projecting opening labial palpi about as long as maxillary palpi. body: ro larva (Fig. 6) gently C-shaped, postdefecated Recently moderately bust. Body annulations similar to other eucerines. Integument lacking spicules and setae; without distinct sclerites except as follows; thoracic each segments on plates dorso-laterally elongate, bearing pair of transversely lightly sclerotized es of terminal, abdominal caudal annulets; much sclerotized segment similarly atrium (Fig. pecially dorsal and lateral regions, ventral region less so. Spiracular in about 7 circumferential rows; outer 12) with very small denticles arranged these somewhat subatrium also with denticles, larger than those of atrium; atrium not projecting above body wall; peritreme large; primary trach?al opening without similar to those of other eucerines. Form of tenth ab collar; spiracle generally


VOLUME 58, NUMBER 4

697

dominai

segment

and

robustior

(Rozen,

1965).

apical

placement

of anus

resembling

that of Melissodes

Discussion Tetralonia hamata and 3 Old World congeners have been studied in sufficient the genus and with other eucerines. The Eucerini detail to allow comparison within 15 of which occur in America of genera, north of is a large, diverse assemblage but the attention Mexico species have received detailed (Hurd, 1979). Several certain majority, including several entire genera, remain unknown. Nevertheless and affirm the close relationship traits have been observed among the repeatedly genera from a behavioral standpoint. construct Eucerines oriented, subvertically elongate cells with their lengths ap as in Svastra twice the greatest width. The cells may be parallel-sided proaching o. obliqua (Rozen, minuta (Rozen, 1969) or they may be 1964) and Tetralonia in the region of the cell closure as in their lower halves with a narrowing wider in T. lanuginosa It is noteworthy that 2methods 1974) and T. hamata. (Mohamed, are common within of cell construction the tribe. In the first case a cavity, larger than the finished cell, is excavated. Then soil, probably mixed with a strengthening is applied to the cavity walls and smoothed into the shape of the finished secretion, a to In the second is smooth the cell walls. cell. Finally applied waterproof lining case a cavity, in the shape of the completed cell, is excavated and its walls smoothed of further soil. The cell walls may be impregnated with a as are more often resistant than the surrounding soil. agent strengthening they Then a waterproof may be found within a genus. lining is applied. Both methods constructs Svastra o. obliqua for example, cells with a built-in wall (Rozen, 1964) whereas S. sabinensis does not (Rozen, appear to 1983). The known Tetralonia a built-in wall although Mohamed construct cells without that (1974) indicated a small amount of clay may have been applied to the cell walls of T. lanuginosa a smoother to provide surface. The cell closure usually consists of soil particles in a more or less well-defined concave on the An interesting variation side. of spiral arrangement, cell-facing this was noted in T. lanuginosa in which the closure consisted (Mohamed, 1974) of a spiral of camphor-tree leaf particles mixed with saliva. in the nature of the waterproof Some variation has been indicated cell lining. as waxlike It has been described for some species rustica (Clement, [Melissodes in others [Svastra sabinensis 1973), M. agilis (Parker et al., 1981)] and non-waxlike and Jones (1980) indicated that the cells ofMelissodes (Rozen, 1983)]. Buchmann were not lined with an adult secretion. The cell linings of Tetralonia persimilis minuta as (Rozen, 1969) and T lanuginosa (Mohamed, 1974) were described waxlike but that of T hamata did not appear so. Likewise Malyshev (1930) the cell lining of T. malvae as a silklike varnish or lacquer to distinguish described it from the waxlike and Macropis. lining of Anthophora, Melitta, Most eucerine bees construct cells singly although Rozen (1969) found that some Tetralonia minuta cells and in others linear series of 2. placed singly Cell depth and arrangement varies greatly within the Eucerini. Nests may be rustica (Clement, shallow, simple structures with but a single cell as inMelissodes M. and and persimilis 1973) Jones, 1980). Peponapis (Buchmann pruinosa (Ma without

the addition


698


thewson, 1968) and Florilegus condignus 1966) construct (LaBerge and Ribble, more nests of 4 or 5 cells while other species regularly include 10 or complex more and unique cells per nest in interesting arrangements. See, for example, Melissodes analis (Rozen, 1974), and Xeno agilis (Parker et al., 1981), Thygater 0.5 m, other glossa strenua (Bohart, 1964). Nest depth in these species approaches construct multicellular nests species' nests reach 1 m. The known Tetralonia 1930) to nearly 70 ranging in depth from less than 20 cm (T. malvae, Malyshev, cm (T. lanuginosa, Mohamed, 1974). in eucerines is near the center of the upper surface of the provision Egg placement mass with only the extremities the food. An exception was noted for contacting and Melissodes Jones, 1980) in which only 1 end (probably persimilis (Buchmann contact the pro The eggs of Tetralonia the posterior) was set in the provisions. at both ends. visions The unshaped provision mass is placed in the bottom of the cell filling, usually, V4 to Vi of the interior although Malyshev malvae (1930) noted that Tetralonia filled Vi to % of its cells with food. A 1-2 mm deep layer of liquid (probably mass in a number of species. nectar) has been noted on top of the provision in has fermentation of been noted several instances the Furthermore, provisions o. obliqua this resulted in a significant increase in their volume and in Svastra was noted in of the upper region of the provisions 1964). Liquefaction (Rozen, M. and rustica (Clement, and Melissodes Chemsak, 1973) (Thorp pallidisignata larva began feeding. Since both authors noted a 1964) shortly after the young odor this change may have been due to larval feeding and/or strong fermenting and a fermenting and Mohamed Wafa fermentation. (1970) noted gas bubbles no such odor in the of T. lanuginosa. Rozen found odor in the provisions (1969) T. and odor cells of of minuta the faint of cells provisioned newly provisioned T that the of malvae stated T hamata has been noted. Malyshev provisions (1930) were very moist or pasty with a layer of liquid on top. He also noted a liquefaction in their volume such that they sometimes and an increase of the provisions a fermenting the top of the cell. He did not mention odor, however. approached the feces about the upper lh or so of the cell and upon, larvae deposit Eucerine or shortly before, completion of defecation begin to spin a cocoon. The cocoon seems similar to those structure and that of T hamata is a complex, two-layered of the difficulty and because of other eucerines. Certain details may differ, however, a comparative a structure of such in interpreting written complex descriptions study of eucerine cocoons might prove interesting and revealing. above show a considerable The traits discussed among degree of uniformity known eucerines with respect to nest and cell structure, provisioning, egg place to the evo Such traits have probably been important ment, and larval behavior. so Yet no variation. success have with and little the of group persisted lutionary for example, is highly variable 2 species are alike in all details. Cell arrangement, choice of nest site, nest depth, and nest density also differ and floral relationships, Svastra o. obliqua (Rozen, from species to species. Some species nest communally, and this 1959) for example, composita (Hurd and Linsley, 1964) and Melissodes deserves further study. The large number of unstudied behavior species and the interest in this group of interesting several unstudied genera should spur continued bees. and beautiful


699

VOLUME 58, NUMBER 4

Acknowledgments E. LaBerge, Donald W. Webb, and like to thank Wallace The author would 2 for Natural and reviewers the Illinois of John K. Bouseman History Survey was on comments This made the their constructive manuscript. possible study to Dr. Wallace E. Grant #NSF DEB 82-07173 Science Foundation by National LaBerge. Literature and 1971. Development Entomol. Soc. 44:225-235.

J. R.

Baker,

Kansas

sexual

Cited of

dimorphism

larvae

the bee

of

J. W.

Folsom,

P. D., B. D.

1979.

Jr. Burks

In K. V. Krombein, in America

Apoidea.

Superfamily

a description Entomol.

Ann.

Observations 1959. Jr., and E. G. Linsley. with notes on Tucker and its parasites, Entomol. News 70:141-146. menoptera: Apoidea). composita W.

and D. W.

E.,

P. D. Hurd, Jr., D. R. of Mexico, pp.

Soc.

with phoridae), Ann. Entomol.

Ribble.

a description Soc. Amer.

1966. Biology of its larva, and 59:944-950.

and Smith, 1741-2209.

North

on

P. D.,

LaBerge,

persimilis

(Say), with 46:516-525.

and Melissodes.

by Tetralonia

of Hymenoptera (eds.), Catalog Institution Press, Washington.

Smithsonian Hurd,

ofMelissodes

Soc.

Entomol.

(Hymenoptera:

15:181-184.

Amer. Hurd,

J. Kansas

Anthophoridae). of red clover

(Hymenoptera: 1922. Pollination

J.

Coelioxys.

strenua

on the biology of Xenoglossa and larval morphology 1964. Notes G. E. 40:174-182. Entomol. Pan-Pacific Apoidea). on the nesting biology 1980. Observations S. L., and C. E. Jones. Buchmann, Pan-Pacific Entomol. 56:200-206. Ckll. Anthophoridae). (Hymenoptera: rustica of'Melissodes S. L. 1973. The nesting biology Clement, (Eumelissodes) Bohart,

of the larva

genus

the nest-site

of Florilegus remarks on

of Melissodes

behavior use

the communal

entrances

of nest

(Hy

Antho condignus (Hymenoptera: in alfalfa pollination. its importance

1963. Comparative and P. H. Raven. E. G., J. W. MacSwain, behavior of bees and Ona II. Oenothera bees of the Great Basin. Univ. Calif. Pub. Entomol. 33:25-50. graceae u. malvae Rossi S. J. 1930. Lebensgeschichte der Tetralonia Zeitschr. Morph. (Apoidea). Malyshev, Okol. Tiere. 16:541-558. Linsley,

J. A.

Mathewson, Mayer,

Nest

1968.

predator 28:1-22. M.

Mohamed,

of the alkali I.

1973.

-.

bee, Nomia

Some

tera: Anthophoridae). 1974. Nest plan

and

construction

Bionomics

Soc.

of Tetralonia

58:97-102. Egypte and G. Parker, F. D., V. J. Tepedino, bee, Melissodes {Eumelissodes)

Ent.

the eastern

of

Soc.

of Meloe Cockerell

melanderi

Egypte

lanuginosa

niger Kirby

bee,

(Cole?ptera:

(Hymenoptera:

lanuginosa 57:251-258.

Klug

cucurbit

Peponapis

41:255-261.

of Tetralonia

characteristics

Bull.

life history Entomol.

J. Kansas

pruinosa (Hymenoptera: Apoidea). 1978. D. F., and C. A. Johansen.

Halictidae). nesting

Klug

(Hymenoptera:

a

Meloidae) Melanderia

sites

Anthophoridae).

(Hymenop Bull.

Soc.

Ent.

Pinto, Rozen,

E. Bohart.

J. D., and R. B. Selander. classification of the New World

species. of Svastra

1964. The biology J. G., Jr. its larvae (Apoidea, Anthophoridae). larvae of the Anthophoridae 1965. The

of -. -. -.

1981.

Notes

on

J. New York agilis Cresson. 1970. The bionomics of blister Illinois

beetles

Biol. Monogr.

obliqua obliqua Amer. Mus. Nov.

of a common

the biology Entomol.

Soc. of

sunflower

89:43-52.

the genus Meloe

and

a

42:1-222.

(Say), with 2170:1-13.

a taxonomic

description

Eu part 1. Introduction, (Hymenoptera, Apoidea) Amer. Mus. Nov. 2233:1-27. (Anthophorinae). of the Anthophoridae part 2. The Nomadinae. (Hymenoptera, Apoidea) Amer. Mus. Nov. 2244:1-38. notes on the bee Tetralonia 1969. Biological minuta and its cleptoparasite, histrio Morgania transvaalensis Proc. Entomol. Soc. Wash. 71:102-107. Anthophoridae). (Hymenoptera: cerini, and Centridini 1966. The larvae

-.

1974. New

York

Nest

of the eucerine biology Entomol. Soc. 82:230-234.

bee Thygater

analis

(Hymenoptera,


Anthophoridae).

J.


700 -.

1983. York

Nesting Entomol.

of the bee Svastra biology Soc. 91:264-268.

R. W., and S. L. Clement. Collinsia Fisher sparsiflora R. W., and J. A. Chemsak. Thorp,

sabinensis

(Hymenoptera,

1977. of the self-compatible Entomophilous pollination and Meyer. Entomol. Soc. 50:37-48. J. Kansas 1964. on Melissodes observations Biological pallidisignata

Rust,

Pan-Pacific Entomol. 40:75-83. Anthophoridae). to the systematics 1969. A contribution of North American Univ. Calif. Pub. Entomol. 57:1-76. (Hymenoptera, Apoidea).

menoptera: P. H. Timberlake, Wafa,

A. K.,

and M.

I.Mohamed.

The logical

following Society

1970. Bull.

Apoidea-Anthophoridae).

J. New

Anthophoridae).

individuals for the year

The

Soc.

Ent.

of Tetralonia

life-cycle

MEMBERS

Bldg. Survey

George W. Byers of Entomology Department Kansas of University 66045 Kansas Lawrence, R. Enns

of Entomology Department of Missouri University MO 65211 Columbia, L. Fischer of Entomology State University Michigan East Lansing, MI 48823 Roland

Department

Gordon W. Frankie 201 WellmanHall of California University Berkeley, CA 94720 Dr. Richard Department Smithsonian Washington, Leslie

C. Froeschner of Entomology Institute, USNMNH D.C. 20560

K.

Johnson of Zoology Department The University of Iowa

Iowa City, IA 52242 L. C. Woodruff Road 2 Westwood Kansas Lawrence,

species

of Synhalonia

Klug

(Hymenoptera:

of the Kansas

1985.

John K. Bouseman 172 Natural Resources Illinois Natural History 607 E. Peabody IL 61820 Champaign,

Wilbur

(Hy

54:259-267.

Egypte

are SPONSOR

lanuginosa

species

66044


Entomo