Novttates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y

Home , Agapostemon melliventris, Centris atripes, Chelostomoides, Diadasia, Kallstroemia, Kallstroemia grandiflora

AMERICAN MUSEUM Novttates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2546 SEPTEMBER 17, 1974

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IIIrlI.II..I.-.I.,..:I....I.,.: '-I.!.'-, .III..I'. ,.I.,-...".I.,.l..'.;.;.11.,I...,I I...-:".-IIII ..II.,.I.II,.I...I..,.I...1I.., .I..,.,...,. .- ,,- ,I.. ,I .,.Ii II.II.f ,. .:;.I.I.I...I.:IIII.,....I:-.I.I. I.II11I-.,I..,I.I IIII...... II... ':- III..:I--I....;..,.,L,...I IIIIII...I.Ic.,-..II.III.-,,..-.II.....,,... ,..I.iI.....:.:.I.,.. .,II,I. Foraging Behavior of Some Bees and Wasps at Kallstroemia grandiflora Flowers in Southern Arizona and New Mexico

M. A. CAZIER Curator Emeritus, Department of Entomology The American Museum of Natural History Arizona State University, Tempe

E. G. LINSLEY University of California, Berkeley

ABSTRACT one of the most conspicuous annuals on the desert At sporadic intervals in the fall the southwest- landscape. The flowers produce an abundance of ern desert landscape is emblazoned with large, pollen and nectar which are exploited in differ- conspicuous, usually- orange-colored flowers of ent ways by various bees and wasps. Opportuni- the annual Kallstroemia grandiflora (Torrey) ties to observe the flower behavior of a number Gray. These flowers produce an abundance of of these insects arose briefly during the summers pollen and nectar, which is exploited in three of 1970, 1971, and 1972. A summary of these ways by insects belonging to 12 different families observations is presented here. and 46 species, including bees, wasps, flies, and butterflies. LOCALITIES The plant is of special interest from the stand- point of pollination ecology, as both the plant Observations on flower visitors to Kallstroe- and some of the insects have adaptations in mia were primarily made at three sites in the behavior that are not always mutually beneficial. upper San Simon Valley, Arizona-New Mexico as The flowers of K. grandiflora are allogamous for follows: 2 miles east of Portal, Cochise County, most of the daily flowering period and become Arizona, on the lower slopes of the Chiricahua autogamous as the flowers close. Thus, they can Mountains below the mouth of Cave Creek Can- be either cross- or self-pollinated, with the yon (August 31-September 5, 1970; August former evidently favored. The flowers have little 27-September 3, 1971); 2 miles north of Rodeo, or no odor, a color range within "bees yellow" Hidalgo County, New Mexico (August 17-23, (500-650 j,), and a strong ultraviolet reflective pattern on both upper- and under-surfaces. In- 1971) and Apache, Cochise County, Arizona sects are evidently attracted by color and ultra- (August 15-16, 1972). violet reflection, with the latter so arranged as to Kallstroemia grandiflora Torrey form nonreflective, dark target, "nectar guide" areas on both upper- and under-surfaces. The genus Kallstroemia (Zygophyllaceae), One group of bees and wasps gather pollen comprising 17 species native to the New World, and nectar from the top of the flower, effecting has been monographed recently by Porter both cross- and self-pollination in the process. A (1969). The following data on the vegetative and second group of smaller bees and wasps gather floral morphology, reproductive biology, and dis- nectar from within the flower but avoid contact tribution of K. grandiflora have been largely sum- with the sexual portions and are therefore of no importance in pollination. The third group of marized from that publication. honeybees and several other larger bees and Kallstroemia grandiflora like its congeners and wasps, extract nectar from the underside of the the related species of Tribulus (including the flower completely avoiding the sexual parts of South European, naturalized T. terrestris Lin- the plant and therefore play no role in pollina- naeus commonly referred to in the United States tion. Even though the plant gains no direct bene- as "puncture vine") is an annual plant with fit from this last group of insects it nevertheless largely prostrate stems radiating from the root. supplies them with a sepal nectar guide that Unlike T. terrestris and many species of Kall- directs them to the nectaries. It is proposed that stroemia, the stems in K. grandiflora are often these species may be contributing to the econ- ascending. The flowers are borne terminally on omy of the plant by reducing the quantity of available nectar so that the pollinators have to the stems but they appear axillary because of the visit more flowers to get their full nectar supply. stems' sympodial growth habit (D. M. Porter, in litt.). In the Southwest, the plants live in flat INTRODUCTION sandy areas in the Sonoran and Chihuahuan des- Kallstroemia grandiflora (Torrey) Gray, com- erts and develop and blossom following the sum- monly known in the Southwest as Arizona or mer rains. Porter (1969) stated that when plants Mexican poppy, has large, conspicuous, usually are growing under extremely favorable condi- orange-colored (occasionally white or yellow, tions, as in ditches where rains collect, they may D. M. Porter, in litt.) flowers and, at times, it is reach 1 meter in height and several meters in 3 4 AMERICAN MUSEUM NOVITATES diameter. He cited Cannon (191 1) for an excep- of the inhibiting principle until, as the flower tional size of 16 or more square meters. Cannon closes, it is self-fertilized. It is equally possible also reported tap roots reaching 22 cm. into the that the genetic viability of Kallstroemia is main- substrate, a characteristic, Porter elaborated, tained even though cross- and self-pollen are which enables the plant to resume growth follow- introduced onto the stigmatic surface at the same ing the normal growing period if an unseasonable time, because as a rule pollen tubes from differ- rain should occur before the plant succumbs to ent but compatible plants grow much faster than the dry season. do those from the same plant. Even though Pro- The species of Kallstroemia thus far studied toxaea and Campsoscolia females distribute large are not pollinated prior to anthesis but the pollen quantities of both cross- and self-pollen, and ne- is shed and the stigma is receptive as the flower is gate the effect of keeping the anthers away from fully opened. During anthesis the stamens are the stigma for most of the day, cross-fertilization appressed to or very near the spreading petals, of at least a large percentage of the ovules in and the stigma is held erectly above them in the Kallstroemia is probably the rule rather than the center surrounded by the 10 stamens (Porter, exception. 1969). At this point the flower is virgin but There is conflicting information on the floral ready for pollination by insects of various types fragrance of Kallstroemia maxima (Linnaeus) and is openly susceptible to both self- and cross- Hooker and Arnott (Porter, 1969) and therefore pollination by these agents. According to Porter on the function of the nectar as an attractant for (1969), at the end of the blooming period, usu- insect pollinators. Porter said, "I have not de- ally early afternoon, which ends the same day tected any odor in the flowers myself." In K. the flower opens, the filaments curve upward and grandiflora we have detected no prominent odor inward moving the anthers into contact with even in concentrated patches of blooming plants stigma as the petals close and hold them to the or individual flowers but no definitive tests have stigmatic surface. The petals twist around the been made. If, indeed, odor is absent or minimal style as they are closing, helping to insure self- the flower nevertheless seems adequately en- pollination, which becomes a necessity in the ab- dowed with visual stimuli to attract insect pol- sence of insect visitors. linators. Its usual deep orange color seems to be It appears that the flowers of Kallstroemia are well within the visual range known for "bees yel- designed to be primarily allogamous (cross-pol- low" (500-650p), at least in the honey bee. linated) for most of the flowering period and The ultraviolet reflection pattern of the flow- autogamous (self-pollinated) near or at the end er strongly suggests attraction of pollinators by of the period to be safe from no or incomplete visual stimulation once the insect is near or in the fertilization. This plan of pollination, which flower. Except for the extreme basal portion and would insure cross-pollination to some extent, is the heavy veins thereon, the large petals are perhaps negated by the behavior of the female strongly ultraviolet reflective and probably at- Protoxaea and Campsoscolia when they hold the tract from a distance. The pistil, stamens, and stamens against the pistil thereby bringing the pollen grains do not reflect ultraviolet and thus anthers, stigma, and their venters into contact. the center of the flower appears as a dark target, As their body hairs or the scopa undoubtedly "nectar guide," to the insect. On the underside contain foreign pollen soon after flight activities of the flower the petals are strongly ultraviolet begin thiev would be effecting both cross- and reflective, whereas the sepals are not and prob- self-pollination, if Kallstroemia is self-compat- ably serve as nectar guides for the insect visiting ible. It appears that the latter is most probable. this region. These sepals are narrow and pointed According to Porter (1969), Kallstroemia grandi- on their outer extremity, gradually widening to flora and thre,e other species in this genus all set the base, directly above which the unreflecting seed when grown in the absence of pollinators. nectary is situated. Although the nectaries appear Kallstroemia like some other plants, probably to be in close contact with the sepals they are starts out self-incompatible, becoming more and actually attached to the receptacle (D. M. Porter, more self-compatible with time and the decrease in litt.). CAZIER AND LINSLEY: FORAGING BEES AND WASPS 5

According to Porter (1969), the genus Kall- Mexico state line and U.S. Highway 80), Kall- stroemia is "pollinated promiscuously, being vis- stroemia grandiflora was present and blooming in ited by various Diptera and Hymenoptera for large patches on both sides of the road, with scat- pollen, and by these and Lepidoptera for nectar. tered small clusters of Solanum elaeagnifolium Very few reports of the types of insects visiting Cavanilles and Cassia bauhinioides Gray inter- the flowers have been recorded. According to no- mixed. Calliandra schotti Torrey was also grow- tations accompanying herbarium specimens, Kall- ing and blooming abundantly along the road stroemia grandiflora is visited by bees and wasps shoulder. All of these plant species offered com- in Sonora and Colima, Mexico. The only polli- petition for Kallstroemia pollinators. Observa- nators of Kallstroemia to have been positively tions of flower visitors were made on August 17, identified are bees of the genus Perdita. Perdita 22, and 23, 1971. The following summary for pectidis Cockerell in New Mexico, Perdita echi- August 17, 1971, was typical (Mountain Stand- nocacti Timberlake in Arizona and Sonora and ard Time): Perdita euphorbiae Timberlake in Sonora." In 0530. 200C. Kallstroemia flower buds tightly the present paper the number of insects known to closed. visit Kallstroemia is enlarged to 33 species of 0554. 190C. Sunrise. Sky clear. Sun on Kall- bees belonging to five families, nine species of stroemia plants. wasps belonging to four families, two species of 0620. 220C. First bee lProtoxaea gloriosa butterflies belonging to one family, and two spe- (Fox)] working Calliandra for pol- cies of flies belonging to two families. No doubt len. this total of 46 species belonging to 12 families 0625. 220C. Kallstroemia flowers beginning to could be expanded significantly with further open. Pro toxaea females active sampling. nearby taking pollen from Sola- num and Cassia as well as Callian- The change in location from year to year in dra. the present study was dictated by the sporadic 0640. 230C. Many Kallstroemia flowers open appearance of Kallstroemia even though no great ½/2-1 inch but no visitors. Centris variance was observed in the physical environ- atripes Mocsdry now working Sol- ment or other features at these localities. An area anuin along with Pro toxaea. dominated by Kallstroemia one year may have 0700. 230C. Scattered flowers of Kallstroemia no or very few plants for the next year or two now open wide but not reflexed. even though conditions conducive to germination Female Protoxaea approach but do appear similar or identical. Kallstroemia seeds are not enter blossoms. Apparently viable pollen and nectar not yet available. for at least three years, they germinate at 0715. 230C. About one-fourth of Kallstroemia irregular intervals and are difficult to germinate flowers now open wide. Small bees artificially (Porter, 1969). This type of intermit- crawling on stamens but not ob- tency among plants is known to be influenced by taining pollen. germination inhibitors that must be washed away 0728. 230C. Few small bees (Exomalopsis) now before germination occurs and by differences in showing some pollen on legs. Pro- the seed coat, especially in those characteristics toxaea still very active on Callian- effecting permeability. A water soluble inhibitor dra. of germination has been found in the seeds of 0738. 230C. Agapostemon melliventris Cresson other tridentata visiting Kallstroemia but not ob- Zygophyllaceae includingLarrea taining pollen. (DeCandolle) Coville and Tibulus terrestris, ac- 0747. 240C. First Protoxaea female in Kall- cording to Porter (1969), and may also be pres- stroemia flower. No pollen on ent in Kallstroemia. body or legs. Many Protoxaea working Calliandra. Apis mellifera GENERAL OBSERVATIONS Linnaeus extracting nectar from nectaries on top, and at base of, At this site (Portal Road, 2 miles north of sepals on underside of Kallstroemia Rodeo, New Mexico between the Arizona-New flowers. 6 AMERICAN MUSEUM NOVITATES

TABLE 1 RELATIVE ABUNDANCE OF BEES AND WASPS TAKING POLLEN AND NECTAR AT FLOWERS OF Kallstroemia grandifloraa Nectar Nectar Nectar Only Only Only Pollen from from from and Above Beneath Underside Nectar Stamens Stamens of Flower Protoxaea gloriosa 499 5d - 36 Svastra sabinensis sabinensis 29 2d - - Diadasia ochracea 49 - 3d Diadasia megamorpha 29 - - - Exomalopsis solani 49 - 49 Psaenythia mexicanorum mexicanorum 99 - -- - Melissodes montana 19 - Svastra sila - 2d Agapostemon tyleri 39 (2 - carrying foreign pollen) Agapostemon melliventris 29 (I carrying foreign pollen) Xylocopa californica arizonensis - 29 Triepeolus spp. - 29 - - Scolia ardens - 19 Melissode$ rimbus - 29, 1d Apis mellifera - - - 29 Cen trisiatripes - 99 Bembix u-scripta - 2d, 29 Campsoscolia octomaculata texensis - - 29

aTwo miles north of Rodeo, Hidalgo County, New Mexico, August 17, 1971, 0800-1000 MST. 0753. 250C. First Protoxaea carrying Kallstroe- beneath flower (table 1). Female mia pollen. Protoxaea observed resting on sta- 0800. 260C. Protoxaea females still working mens rubbing legs together and re- Calliandra but few now taking pol- moving pollen from scopa; after len from Kallstroemia along with spending about a minute in this Psaenythia m. mexicanorum (Cock- activity she flew off without visible erell), Svastra s. sabinensis (Cock- pollen. erell), Diadasia ochracea Cockerell 1000. 290C. Flowers mostly reflexed and begin- and other bees (table 1). Centris ning to show signs of wilting. Only atripes and Bem bix u-scripta (Fox) visitors were a few nectar seekers. taking nectar from beneath flowers Observations terminated. along with Apis. Male Pro toxaea ro- Apache, Cochise Co., Arizona. Site an old tating on stamens while extracting roadside excavation extending north and south. nectar; underside of body heavily scattered over an area ap- dusted with pollen. Kallstroemia plants 0900. 280 C. Various bees taking pollen but proximately 50 meters wide by 300 meters long. most seeking nectar only, either Observations on flower visitors were made on from above or below stamens or two days in August, 1972. The following sum- CAZIER AND LINSLEY: FORAGING BEES AND WASPS 7 mary is for August 15, 1972 (Mountain Standard niaculata (Say) becoming heavily Time): coated with pollen while rotating 0745. 210C. Sky clear. Cool breeze. Kall- on stamens as they probe nectaries, stroemia flowers on east-facing Tiphiid wasp Myzin um navajo bank mostly wide open, those on Krombein, although as large as flat and west-facing bank mostly Campsoscolia avoids contamina- just beginning to open. No bees tion with pollen by crawling be- evident but substantial number of neath anthers to get to nectaries. Bembix u-scripta flew rapidly 1000. High proportion of flowers now among plants extracting- nectar wilting. Bembicidcs no longer ac- from nectaries on top and at tive. Few nectar seeking bees pres- base of sepals on underside of ent, mostly males. Observations flower. terminated. 0802. First bee. (Agapostemon melliven- Two miles east of Portal, Arizona. Observa- tris) taking pollen. tions were made in an open meadow adjacent to 0805. Several Melissodes t. thelypodii Cockerell collecting pollen. and south of the Portal-Rodeo road. The 0815. Various species of bees now taking meadow was surrounded primarily by Acacia pollen and nectar (table 2), espe- constricta Bentham and Prosopis juliflora cially M. t. thelypodii. Male bees, (Swartz) De Candolle on the north and east and primarily Protoxaea gloriosa, and by Larrea tridentata on the south and west. scolhid wasp Campsoscolia o. octo- These were not in bloom nor were other, less

TABLE 2 RELATIVE ABUNDANCE OF BEES AND WASPS TAKING POLLEN AND NECTAR AT FLOWERS OF Kallstroemia grandifloraa Nectar Nectar Nectar Only Only Only Pollen from from from and Above Beneath Underside Nectar Stamens Stamens of Flower Melissodes thelypodii thelypodii 259 lid Agapostemon tyleri 79 I Svastra sabinensis sabinensis 69 3d Ptilothrix sumichrasti 39 Psaenythia mexicanorum mexicanorum 39 Diadasia diminuta 29 Halictus tripartitus 29 _9_ Pro toxaea gloriosa 1 4d Triepeolus sp. 39, 2d Campsoscolia octomaculata texensis 89 Diadasia ochracea 10d Pseudopanurgus aethiops 29 Andrena accepta 29 Coelioxys sp. 19, id Nomada sp. id Myzinum navajo 219 Myzinum frontale 19 Centris atripes Id, 59 Bembix u-scripta 30d, 169

aApache, Cochise County, Arizona, August 15, 1972, 0800-1000 MST. 8 AMERICAN MUSEUM NOVITATES numerous shrubs. The principal plants competing which are abundantly covered with large orange for pollinators were Baileya multiradiata Harvey pollen grains and the stigma, which is covered and Gray, Gutierrezia microcephala (De Can- with stigmatic fluid, in contact with the ab- dolle) Gray, Eriogonum abertianum Torrey, E. dominal venter and hind legs. fasciculatum Bentham, Allionia incarnata Lin- As soon as the bees' hold on the style and naeus, Zinnia pumila Gray, Lepidium montanum stamens is secured, the mouthparts are inserted Nuttall, and Pectis papposa Harvey and Gray. into the nearest nectary for from a fraction of a Observations were made in connection with second to about one second, presumably depend- another study and centered on Protoxaea glori- ing on the amount of nectar present. From one osa, whose females worked Kallstroemia flowers or two to all five nectaries are visited and on much longer in the day than at any other site en- several occasions six mouthpart insertions are countered. Flower visitors other than female Pro- made, the first nectary being overlapped. After toxaea, although less diverse than at the Rodeo each nectary is sampled the bee shifts its hold on site, included the following: females ofMelissodes the style and stamens, shaking them rather vio- t. thelypodii, Svastra s. sabinensis, Diadasia lently, as its body moves around to a position ochracea, D. megamorpha Cockerell, Mellissodes from which the mouthparts can contact the next (E.) montana Cresson, Megabombus (M.) penn- nectary. This shifting and shaking motion of the sylvanicus sonorus (Say) taking pollen; females bee in rotating its position completely around of Xylocopa califomica arizonensis Cresson, Me- the inside of the flower would account for the lissodes limbus La Berge, Triepeolus sp., and Sco- large amounts of loose pollen found throughout lia ardens Smith and males of Protoxaea gloriosa, much of the body of the bee and occasionally in Svastra s. sabinensis, Svastra sila (La Berge), and the bottom of the corolla. In Protoxaea each Diadasia ochracea taking nectar while on top of stop in the rotary movements is made at the the stamens; females ofExomalopsis solani Cock- juncture between two petals beneath which is a erell extracting nectar while beneath the anthers nectary. These are small, elongate (rectangular), and Apis mellifera (hundreds of workers) and fe- dark green, spongy-looking masses of tissue situ- males of Centris atripes, Pterocheilus arizonicus ated middorsally at the base on the elongate, nar- R. Bohart, and Monobia texana (Cresson) ex- row, hairy sepals issuing from beneath the area tracting nectar from nectaries on top of and at where the petals overlap. the base of the sepals on the underside of the By late morning the movements of females on flower. Protoxaea pollen collectors in the early exposed flowers are very rapid because most of afternoon were captured as follows: (September these blossoms have already been visited. Very 1 and 2, 1970; Mountain Standard Time): 1304, often the bee then samples only one, two, or 1314, 1328, 1334, 1343, 1347, and 1357. No three of the nectaries before flying to the next females were evident between 1357 and 1430 flower. When an obscured, previously unvisited when observations were terminated. flower is found, either inside the plant or beneath another plant in the shade, females slow their flight and make a more complete visitation. BEE AND WASP BEHAVIOR We did not observe honeybees taking pollen WHEN COLLECTING FROM FLOWER TOPS from Kallstroemia grandiflora, even in areas The behavior of bees collecting pollen and where they were abundant, as at the Portal local- nectar in flowers of Kallstroemia grandiflora var- ity. However, Porter (1969) made the following ies according to the size of the insect. Females of observations of honeybee behavior in flowers of Protoxaea gloriosa alight in the upturned flowers a related species: "When a honeybee lands on a which droop downward with the bees' weight, flower of Kallstroemia maxima it thrusts aside sometimes bending 180 degrees so that the bee is the stamens, while standing over them and the working upside down. As they land in the flow- style in order to reach the nectar at the bases of ers, facing downward, they gather the style and the sepals. In this way pollen is transferred from all 10 stamens into the venter, holding them the anthers to the bee, and from the bee to the there with all six legs. This places the anthers, stigma. The bee circles clockwise on the flower CAZIER AND LINSLEY: FORAGING BEES AND WASPS 9

3 4 // FIGS. 1-4. 1. Male of Protoxaea gloriosa extracting nectar from underside of Kallstroemia grandiflora flower. 2. Worker of Apis mellifera extracting nectar from underside of flower. 3. Female of Protoxaea gloriosa extracting nectar and collecting pollen while rotating with stamens clasped between legs. 4. Male Protoxaea gloriosa extracting nectar while standing on stamers and rotating. until it has sampled all the nectaries, then moves in either the initial visit or upon return of the bee on to another flower, not uncommonly returning to a flower previously visited." later to the original flower, either to repeat the The wasp Campsoscolia o. octonaculata ac- previous performance or to reject the flower. The tually holds the stamens together under the ab- bee, therefore, may act not only as an agent in domen, in the manner of larger bees when gather- cross-pollination, but also may effect self-pol- ing pollen, and in the process the wasp becomes lination by transferring pollen from anthers to heavily coated beneath with pollen, even though stigma in the same flower. This is accomplished it has no scopa and is sparsely pilose. 10 AMERICAN MUSEUM NOVITATES

BEE AND WASP BEHAVIOR WHEN as discussed later. As Kallstroemia evidently pro- EXTRACTING NECTAR FROM duces large amounts of nectar, the "pirating" ac- WITHIN THE FLOWER tivities of these species can best be considered as Smaller bees and a few wasps such as the ti- being commensalistic in that the plant is un- phiidMyzinum navajo (tables 1, 2) crawl into the harmed and the insects are benefited by the nec- flower over the petals and beneath the anthers, tar. If these insects were using ultraviolet light among the filaments, and thus reach the nectaries they would be attracted to the highly reflective with minimal contamination by pollen, except surface of the petals or perhaps just to the orange for that which has been shaken into the corolla color of the petals or both. Upon landing on the by the activities of the larger bees and wasps and petal the insect may be attracted to the nec- through which they have to walk to get to the taries by olfactory stimuli out of the range of nectaries. They do not come into contact with human detection that would guide them down into the stigmatic surface and therefore are of no im- the corolla and to the nectaries, which are on the portance in pollination. Their small size makes it edge and beneath the nonreflective central area. difficult if not impossible for them to manipulate BEE AND WASP BEHAVIOR WHEN the sexual portions of the flower, and as their EXTRACTING NECTAR FROM weight does not bend the flower they have no THE UNDERSIDE OF THE FLOWER need to hold onto these plant structures. Because the smaller bees avoid the main source of pollen Honeybees and several of the other larger bees and do not collect "waste" pollen in the corolla, and wasps that seek only nectar from Kallstroe- even though pollen is a necessary food for the mia, extract it from a position beneath the flow- larvae of at least four of the listed species, it er (tables 1, 2). They approach the flower from appears that Kallstroemia pollen may also have beneath, landing on the stem, which they grasp some undesirable characteristic. Also, this partial with all six legs. They move quickly up to the avoidance of pollen contamination may be a logi- flower, pull the sepal down with one or both cal step in the evolution of complete avoidance, front legs, and insert the mouthparts into or onto

FIGS. 5, 6. 5. Male of Diadasia ochracea extracting nectar from a position underneath the stamens of Kallstroemia grandiflora. 6. Female of Myzinum navajo extracting nectar from a position underneath the stamens. CAZIER AND LINSLEY: FORAGING BEES AND WASPS I1I the nectary that lies middorsally at the base of It is well known that pollen-collecting honey- the sepal. They move around the stem to get to bees, if offered a choice of pollens, show a pref- the other nectaries but are not nearly so quick in erence for some over others. Levin and Bohart completing their visit as species that rotate to the (1955) have demonstrated that this is so even nectaries from a position astride the stamens when the pollens are separated from the flower within the flower. However, this procedure does structures, color patterns, and perfumes by allow the bees and wasps to avoid contamination which they are normally surrounded. They found with Kallstroemia pollen, which clings abun- that pollen color, reflectivity, moisture content, dantly and persistently to body hairs. protein, and sugar ingredients all appear to have These species have no role in pollination, no influence on attractiveness (they did not un- being thieves of the nectar supply which they equivocally eliminate odor as a factor). Even in- may deplete thereby discouraging those larger side the hive, bees show a preference among pol- bees and wasps that do pollinate while gathering lens of individual plant species when offered a nectar. However, if the nectar supply is merely choice and eat some and reject others (Synge, reduced in volume the pollinators may be forced 1947; Doull, 1966). to visit and pollinate more flowers in order to In hundreds of observations we failed to see fulfill their nectar requirements. Thus the rob- even one honeybee learning by experience. This bers could be an asset rather than a liability to indicates that there is something more than ran- the plant which through its ultraviolet reflection dom foraging activities on Kallstroemia. Even the of the underside of the petals assisted by the new foragers recruited to Kallstroemia appear to sepal nectar guide, actually seems to encourage have been "instructed" not only that the hive these activities. needs nectar and not pollen but also the best way The pirates of the pollination world, the to collect from this source. That the bees have honeybees, have "leamed" this shortcut to the been recruited to Kallstroemia indicates that the nectaries as they have learned similar shortcuts in nectar is good and that at least some preferential other plants to avoid disagreeable features (al- recruiting has been accomplished during dancing falfa) or to take advantage of more highly con- and Nasanov scent gland exposure. Perhaps infor- centrated nectar (cotton). In Kallstroemia they mation on how to gather the nectar is trans- may have learned by being "rewarded" by the mitted by the scouts and foraging bees when abundance of easily available nectar and by being they return to the hive. We can, of course, only "penalized," if they go inside the flower, by the speculate on how this behavior was originally large, sticky pollen grains which, for some rea- learned. The most obvious suggestion lies in the son, they do not seem to utilize for larval food. fact that the flower advertises its undersurface It is well known that some honeybee foragers both in color and ultraviolet reflection in addi- collect only nectar, some only pollen, and others tion to having the sepal nectar guide. Further- both pollen and nectar from the same plant. This more, the underside of the flower is exposed is the case if both the pollen and nectar are satis- when the larger insects land on top causing the factory to the honeybee but does not seem to flower to bend as much as 180 degrees. Wind apply here as all (hundreds) of our observations action may also be responsible for some exposure indicated complete avoidance of the inside of the of the undersurface. Only once did we see a Kallstroemia flowers and therefore the pollen. honeybee taking nectar from a flower that was Many experiments, especially with economic already bent over by a Protoxaea bee. However, crops, indicate that the pollen itself can deter- repeated exposure to the overturned flowers, the mine the bees' tendency to select it, irrespective reward of easily available nectar, and the lack of of the ease with which it can be collected. In- sticky pollen may have fixed the visual image versely this means a selective avoidance of some pattern in the bee to an extent that it has be- pollens, and as it has been shown that bees do come communicable. How the other two regular not select pollens for their nutritional value, it and consistent visitors (solitary) to the underside may be simply a matter of mechanical contam- of the flowers, Centris atripes and Bembix u- ination that makes it undesirable. scripta, learn as maintain their fixed behavior 12 AMERICAN MUSEUM NOVITATES pattern in this respect is difficult to understand tar. At this site Solanum elaeagnifolium was pres- unless each new generation relearns the pattern ent in scattered patches, and the earliest female suggested for the honeybee or mimics the honey- nectar visitors to Kallstroemia carried pollen bee in this behavior. That mimicry may occur is from Solanum. suggested by the fact that in all the areas where A favorite nectar source for both sexes of Psa- we have encountered male Protoxaea in abun- enythia in the surrounding area is Asclepias sub- dance, they consistently visited the inside of the verticillata (Gray) Vail. flower except at the only site where honeybees Perdita (Macroteropsis) echinocacti Timber- were abundant near Portal. However, no honey- lake. Recorded by Timberlake (1954) from flow- bees were present at Apache, where large num- ers of Kallstroemia grandiflora in the Santa Cata- bers of Bembix u-scripta were extracting nectar lina Mountains, Pinal County, Arizona, and near from the underside of the flower. Nogales, Santa Cruz County. Perdita (Perdita) pectidis Cockerell. Three fe- BEE AND WASP VISITORS TO males from Kallstroemia maxima at Las Cruces, KALLSTROEMIA Dona Ana County, New Mexico, have been reported by Timberlake (1960). According to Por- FAMILY ANDRENIDAE ter (in litt.), K. maxima does not occur in the Andrena (Pterandrena) accepta Viereck. Fe- western United States. The Timberlake record males were found in small numbers at Apache, represents either K. parviflora Norton (probably) Arizona, feeding at the nectaries by crawling or K. californica (S. Watson) Vail. under the stamens. They were not uncommon in Perdita (Epimacrotera) euphorbiae Timber- the vicinity of Kallstroemia plants taking pollen lake. Males and females have been collected at from Helianthus annuus (Linnaeus). As empha- flowers of Kallstroemia and Euphorbia near Nav- sized by La Berge (1967), this species is an oligo- ajoa, Sonora Mexico (Timberlake, 1960). We en- lege of Compositae. Of 219 females with flower countered no Perdita at Kallstroemia in our records examined by him, 218 were taken from study sites. Compositae and of these, 194 were from Helian- Pseudopanurgus aethiops (Cresson). Occasion- thus. The species is widespread, ranging from ally found taking nectar from beneath the sta- eastern California and Oregon to New Jersey, mens of Kallstroemia. In our experience the spe- Georgia, Arizona, and northern Mexico. Their cies is an oligolege of Compositae. We have taken presence at flowers of Kallstroemia was un- it locally gathering pollen from Baileya pleniradi- doubtedly opportunistic. ata Harvey and Gray, Heterotheca psammophila Psaenythia (Protandrena) mexicanorum mexi- Wagenknecht (Porter, in litt.), Verbesina enceli- canorum (Cockerell). Females of this subspecies oides (Cavanilles) Bentham and Hooker, and Heli- have been reported by Timberlake (1955) from anthus annuus. flowers of Kallstroemia grandiflora at Tucson and Cortaro, Arizona, and from Solanum rostra- FAMILY OXAEIDAE tum Dunal near Patagonia, Arizona. We have also recorded it previously taking pollen from So- Protoxaea gloriosa (Fox). This large bee was lanum rostratum and S. elaeagnifolium near by far the most abundant and consistent visitor to Douglas, Arizona (Linsley and Cazier, 1963). Kallstroemia in the areas sampled. It is a polylectic At the Rodeo, New Mexico locality, both species, usually active from sunrise to midday, sexes were relatively numerous, taking nectar taking pollen from Solanum, Cassia, Calliandra, from within the flower working over or between and Larrea during the first two hours of flight the stamens and as a result usually having some (Linsley and Cazier, 1972). However, at flowers loose pollen on body and legs. Some females, of Kallstroemia, which do not open and present however, had accumulations of pollen on the legs pollen until an hour or more after sunrise, pollen that did not appear to be accidentally acquired collecting females may be active in small num- and others definitely had Kallstroemia pollen bers taking pollen at noon or shortly thereafter, packed into the tibial scopa and mixed with nec- particularly on cool overcast days. Females take CAZIER AND LINSLEY: FORAGING BEES AND WASPS 13 nectar from within the flower, as described ear- takes pollen from a variety of plant sources lier. Males were usually found taking nectar from but is not known to do so from Kallstroemia. within the flower, in which case they became Megachile (Pseudocentron) sidalceae Cock- heavily dusted beneath with pollen. However, at erell. Kallstroemia is among the wide variety of the Portal site, where honeybees were abundant, plant genera reported as flower records for this some but not all males were seen taking nectar species by Butler (1965). He did not record the from the underside of the flower. plant species or whether the bee was taking The pollens of several of the plants preferred pollen. by this species are fine. Pollen grains of Kallstroe- Chalcidoma (Chelostomoides) adelphodonta mia removed from females averaged 66.7,u (range (Cockerell). Recorded from Arizona by Butler 63.4,u-68.8,u) (Linsley and Michener, 1962) (see (1965) at Allionia and Kallstroemia without fur- also Martin and Drew, 1970), those from So- ther data. lanum 25.6, (23.5, - 27.0g) (Linsley and Cazier, Coelioxys sp. Both sexes of this parasitic 1963). megachilid were found at the Apache site in Kall- stroemia flowers taking nectar from beneath the stamens. FAMILY HALICTIDAE Halictus (Seladonia) tripartitus Cockerell. A FAMILY ANTHOPHORIDAE widespread polylectic species. A few females Exomalopsis (Phanomalopsis) solani Cock- were observed at the Apache site crawling on the erell. Females visit Kallstroemia flowers primarily anthers of Kallstroemia and gathering pollen. for nectar, crawling beneath the stamens to ob- Agapostemon melliventris Cresson. A wide- tain it. The species is polylectic. Timberlake spread polylectic species. Roberts (1972) (1947) recorded it from a diverse group of flow- mapped its range as primarily Great Basin and ering plants but its pollen collecting habits are the desert areas of southwestern United States not well known. and northern Mexico, but also extending over Nomada sp. This parasitic bee was found ex- most of the Great Plains. We found it in Kall- tracting nectar from a position beneath the sta- stroemia flowers but only taking nectar. When mens ofKallstroemia at the Apache site. carrying pollen, it proved to be from another plant Holcopasites caliopsidis carinatus Linsley. A source. male recorded by Hurd and Linsley (1972) from Agapostemon tyleri Cockerell. This species is flowers of Kallstroemia hirsutissima Vail at El more restricted in its distribution than A. melli- Salto, San Luis Potosi, Mexico. This parasitic bee ventris. Maps by Roberts (1972) indicate that the probably crawls beneath the stamens to reach the two are sympatric primarily in the eastern part of nectaries, avoiding contact with pollen. the range of that species. Like A. melliventrns, Ptilothrix sumichrasti (Cresson). This species females we observed were carrying foreign pollen was among the bees collecting pollen from K. on the legs, although the body was contaminated grandiflora at the Apache site. Butler (1967) with Kallstroemia pollen. recorded females taking pollen from cultivated Nomia tetrazonata tetrazonata Cockerell. Rib- cotton (Gossypium) in the Santa Cruz and Gila ble (1965) recorded three females from Kall- river valleys, as well as from Kallstroemia (pre- stroemia grandiflora but did not identify the cise locality not stated). In these localities both locality or whether they were taking pollen. We sexes were abundant and mating was observed on did not encounter the species in our study sites. the latter flowers. He also recorded visits ofPtilo- thrix to the related Tribulus terristris (puncture- vine) but did not state whether they were taking FAMILY MEGACHILIDAE pollen. In Mexico the females take pollen from Megachile (Acentron) albitarsis Cresson. The Ipomoea spp. (Linsley, MacSwain, and Smith, female of this widespread southern species was 1956) and probably also in southem Arizona. recorded by Cockerell (1908) from Kallstroemia They occur in areas of summer rain and require as M kallstroemiae, new species. The species moisture for nest construction. At the site near 14 AMERICAN MUSEUM NOVITATES

Apache the females were nesting in open hard- gloriosa this is one of the most abundant visitors packed soil in the vicinity of the Kalistroemia to Kallstroemia flowers in the San Simon Valley. plants. Both sexes were present in numbers at all three Diadasia ochracea (Cockerell). Females ofDia- principal study areas, especially at Apache, tak- dasia ochracea were found collecting pollen from ing nectar from within the flower. All but a few Kallstroemia at the Rodeo site and males were of the females were also actively taking pollen. observed taking nectar from positions beneath Melissodes (Eumelissodes) montana Cresson. the stamens at both the Rodeo and Apache study MAelissodes (E.) montana was regarded by La areas. Habits of the species have been described Berge (1961) as an oligolege of Compositae, and by Linsley and MacSwain (1957) under the name most of our collections confirm a preference for "olivacea." These authors reported several spe- plants of this family. Locally we have found fe- cies of Sphaeralcea as pollen hosts in various males taking pollen from Gutierrezia sarothrae parts of its range, a genus to which D. ochracea (Pursh) Britton and Rusby and Heterotheca psam- has been generally believed to be oligolectic. mophila However, in our study area near Por- Sphaeralcea is usually evident at both of these tal females were found taking pollen from Kall- sites but relatively few plants were in bloom stroemia. when Kallstroemia was sampled. In a discussion Melissodes (Eumelissodes) limbus La Berge. of floral constancy in Diadasia, Linsley and Mac- This species ranges from central Arizona through Swain (1958) described a stress situation in southwestern Arizona and western Texas to which D. australis californica Timberlake, usually south-central Mexico. According to La Berge restricted to pollen of cactus, turned to Phacelia (1961), it is an oligolege of Compositae, prefer- in the absence of its normal host. ring plants of the genera Verbesina,Haplopappus, Diadasia diminuta (Cresson). Diadasia diminu- and Baileya. Females have been observed extract- ta, like D. ochracea, generally regarded as an ing nectar from Kallstroemia at the site near Ro- oligolege of Sphaeralcea (Linsley and MacSwain, deo inside the flower but beneath the stamens so 1957, 1958), was found taking pollen from Kall- as to avoid contact with the pollen. stroemia at the Apache site under what may have We have found it locally taking pollen from been conditions of stress. Further studies of fe- Haplopappus gracilis (Nuttall) Gray and Baileya male behavior in each of these species should be pleniradiata Harvey and Gray. made to clarify the extent of flower specificity. Melissodes (Tachymelissodes) sonorensis La Diadasia megamorpha Cockerell. Two females Berge. Although not encountered by us, La Berge of this species were taken from flowers of Kall- (1963b) recorded males and one female from stroemia at the Rodeo site as they were gathering Kallstroemia grandiflora south of Navajoa, So- pollen. We did not encounter it on other plants nora, Mexico. locally and are not familiar with its floral rela- Svastra (Epimelissodes) sabinensis sabinensis tionships. (Cockerell). This species occurs in south-central Melissodes (Melissodes) thelypoddi thelypoddi Arizona and southwestern New Mexico and has Cockerell. This subspecies occurs from southern been recorded by La Berge (1956) from plants of California through Arizona and New Mexico to several genera of Leguminosae, Compositae, and southeastern Texas and northern Durango (La Malvaceae as well as from Kallstroemia grandi- Berge, 1956). As would be expected it takes pol- flora. We found both sexes at Kallstroemia at the len from wide variety of plants, including culti- Portal, Arizona, Apache, Arizona, and Rodeo, vated Gossypium (cotton) (Butler et al., 1960) New Mexico, sites. Most females were taking and Carthamus (safflower) (Butler, Werner, and pollen. Levin, 1966). Cockerell (1905) recorded the Svastra (Epimelissodes) sila (La Berge). We male from Kallstroemia at Mesilla Park, New have found this species locally visiting Com- Mexico, under the name Melissodes kallstroe- positae for pollen, particularly Baileya pleni- miae, new species, and La Berge listed Kallstroe- radiata andHeterotheca psammophila, host plants mia grandiflora among the flower records but also recorded by La Berge (1963a). This is a without reference to pollen. Next to Protoxaea southwestern desert species occurring from CAZIER AND LINSLEY: FORAGING BEES AND WASPS 15 southeastern Arizona to Baja California (La and probably would do so at the Portal site Berge, 1958). Males were taken at the Portal under appropriate conditions, such as pollen study area feeding on nectar from within the stress, when foraging for the first time, or when Kallstroemia flowers, rotating on the stamens. the colony needed it. Centris atripes Mocsary. This species collects Austin (1972) recorded their behavior at flow- pollen locally from Solanum (Linsley and Cazier, ers of the related Tribulus cistoides Linnaeus as 1963) and was active on Solanum elaeagnifolium follows (text references to illustrations omitted): at the Rodeo and Portal sites. However, along "Most of the Honey Bees visiting the flowers with other Solanum pollinators it tums to other approach in the 'normal' bee fashion described plants for nectar. Females were abundant at Kall- by Meeuse (1961) and Faegri & Pijl (1966). stroemia in all three study sites taking nectar from Instead of landing on the stamens and stigmas as beneath the flower, numerically being second they should, however, they circle around the only to honey bees at the Portal and Rodeo sites. flower and land on the outside of the calyx and Xylocapa califomica arizonensis Cresson. Fe- corolla. After landing they separate two petals' males collect pollen locally from a variety of with their front legs, insert their tongues, and sip plant species but were not observed doing so nectar. They continue this procedure around the from Kallstroemia. They occasionally visit the flower until they have collected nectar between flowers for nectar and their weight causes the all the petals. Once a flower has been utilized, blossoms to invert, requiring the bee to cling to the bees fly to another and repeat the process. the stamens for support. In the process they Several flowers are usually visited before the bee become liberally dusted beneath with pollen disappears. which they transport from flower to flower. "Other Honey Bees approach and land within They were taken at the site near Rodeo. the corolla in the 'normal' manner. In so doing Megabombus (M.) pennsylvanicus sonorus they position their bodies over the stigma and (Say). The bumblebee M. pennsylvanicus sonorus stamens. Normally they insert their tongue be- is one of the most ubiquitous and conspicuous tween the petals into the nectary, sip nectar and flower visitors in the San Simon Valley area turn around the flower to drink nectar from (Linsley, 1960). It is active throughout the day, other nectaries. They dust much of their body working early blooming plants such as Solanum with pollen as they turn. After the nectar has before sunrise (Linsley, 1962) and late aftemoon been collected, they often brush the stamens or early evening flowers such as Mentzelia (Lins- with their front legs to gather pollen. ley and Hurd, 1959). However, we found it tak- "After nectar has been taken and pollen gath- ing pollen from Kallstroemia only rarely and ered with the front legs, the bees may fly to only at the site near Portal. The status of this another flower to repeat the process. It is com- species has been recently discussed by Milliron mon, however, for an individual to brush pollen, (1973). He showed its range as extending from fly off and hover in front of the flower, return to central California to southern Baja California, brush more pollen, and hover again. This may be south-central Mexico, southem Arizona, New done three or four times before the process is Mexico, and Texas. Kallstroemia is not included repeated at another flower. among the many flower records reported by him. "Those individuals which exhibit this hovering Apis mellifera Linnaeus. Honeybees were very behavior usually have their pollen baskets full. abundant at the site east of Portal (an apiary was The bees with pollen in the pollen baskets have nearby), scarce near Rodeo, and absent at little or no pollen adhering to the hairs of their Apache. At no time during the three summers of legs and body. Bees which have no pollen in their observations on Kallstroemia were individuals baskets rarely hover, and their bodies and legs seen inside the flower taking either pollen or nec- usually are dusted with pollen." tar. All the hundreds of worker bees at the Portal site took nectar from the underside of the flower. However, honeybees do work Kallstroemia 'This is not the case in Kallstroemia as observed by flowers for pollen, as attested by Porter (1969) us. 16 AMERICAN MUSEUM NOVITATES

HYMENOPTERA SCOLIOIDEA and southern Colorado through Texas, New FAMILY TIPHIIDAE Mexico, and Arizona to northern Mexico (Brad- ley, 1928a). When visiting Kallstroemia flowers Myzinum navajo Krombein. Myzinum navajo for nectar, the females either enter the flower was described from material representing various and hold the stamens together with the legs like a localities in Arizona and also Sonora, Mexico bee and as they rotate from nectary to nectary (Krombein, 1938). No flower records were re- the venter and legs become heavily coated with ported. At the Apache, Arizona, site it was the pollen (Apache site), or on the site near Rodeo second most abundant wasp visiting Kallstroemia they collected nectar from the underside of the flowers for nectar, Bembix u-scripta being the flower. most abundant. Unlike the latter species, which extracts nectar from the underside of the flower, HYMENOPTERA VESPOIDEA M. navajo crawls beneath the stamens to reach FAMILY VESPIDAE the nectaries. It plays no significant role in Kall- stroemia pollination. Monobia texana (Cresson). This species, rang- Myzinum frontale Cresson. A single female of ing from Texas to Arizona (Bequaert, 1940), lo- this species was taken from a flower of Kallstroe- cally visits a variety of flowers for nectar, includ- mia at the Apache site along with females of M. ing Baccharis and Eriogonum. At Kallstroemia navajo. Like individuals of that species, it had we have observed females taking nectar only crawled beneath the stamens to reach the nec- from the underside of the flower, in the manner taries and was uncontaminated with pollen. used by Bembix and Pterocheilus at the site near Krombein (1938) reported the species as oc- Portal. curring from Kansas and Colorado to Texas, New Pterocheilus arizonicus R. Bohart. This south- Mexico, Nevada, Arizona, and southern Cali- western species was found in small numbers at fornia. Recorded flowers visited include: Poly- the site near Portal taking nectar from the under- gonum and Eriogonum (Polygoniaceae), Meli- side of Kallstroemia flowers in 1970 and again in lotus (Leguminosae), Chilopsis (Bignoniaceae), 1971. We have observed a related species, P. pedi- and Solidago, Baccharis, and Pluchea (Com- cellatus R. Bohart, chewing its way into the base positae). of the long tubular flowers of Gilia longiflora (Torrey) George Don to gain access to the nec- FAMILY SCOLIIDAE taries.

Scolia (Trichoscolia) ardens Smith. This large, FAMILY SPHECIDAE conspicuous wasp occurs throughout much of southwestern United States and northern Mexi- Bembix u-scripta (Fox). This was the most co. Females are not uncommon in the San Simon abundant wasp observed taking nectar from Kall- Valley, especially at flowers of Baccharis. When stroemia flowers, particularly at the Apache, Ari- taking nectar from Kallstroemia they position zona, site. The species differs markedly from astride the stamen and the venter becomes dusted other members of the genus both by the pos- with pollen. One female was taken at the site session of functional ocelli and the fact that the near Rodeo. females hunt their dipteran prey and provision Campsomeris (Campsomeris) tolteca (Saus- their nests only in the dusk, around and after sure). A female was captured while taking nectar sunset (Evans, 1960). It occurs in southwestern from Kallstroemia at the site near Rodeo. The United States and Mexico but is generally con- venter was heavily dusted with pollen. The spe- sidered to be rare or localized. cies occurs in southern California, Arizona, west- Both sexes take nectar from beneath Kall- ern New Mexico, and Mexico (Bradley, 1928b). stroemia flowers and the males in particular be- We have taken it locally visiting flowers of Bac- gin patrolling the plants before the flower buds charis. begin to unfurl, attempting to force their mouth- Campsoscolia octomaculata texensis (Saus- parts into the nectaries. Males were already very sure). This species occurs from western Texas active at the Apache site by 0745 when observa- CAZIER AND LINSLEY: FORAGING BEES AND WASPS 17 tions were initiated. Although our observations tion pattern, their obvious nectar guides, and were made in mid-August and Evans's observa- their copious supplies of both pollen and nectar tions were made in May, it seems likely that Kall- make the flowers prime targets for insect visitors stroemia nectar may have been providing energy even though there is no evident floral fragrance. for the morning "sun dance" at a nearby nest site Forty-six species belonging to three orders and as described by Evans (1960): "Each morning we 12 families of insects are herein listed as regular entered the area about 0800, and at this time the visitors to the flowers but not all use the flowers males were already engaged in their 'sun dance.' in ways directly beneficial to the plant. Some Each male flew rapidly in loops, figure eights, species gather pollen and nectar from inside the and irregular patterns only 2-6 cm. above the flower and pollinate the flower in the process. sand surface, much as in Bembix cinerea (Hand- Others visit the inside of the flower for nectar lirsch) (Evans 1957). By 1030 some reduction of only; some visit the inside of the flower but males could be noted, and by 11 15 each day the avoid the anthers and stigma and a fourth group last male had disappeared. By 1500 a few males avoids the inside of the flower and its reproduc- were once again active, and from 1600-1900 each tive parts and gains access to the nectaries from day there was a second though somewhat less underneath the flower. The nonpollinators that populous 'sun dance.' " If there was an evening visit the inside of the flower use the same visual "sun dance" near the Apache site, Kallstroemia cues as do the pollinators (orange, ultraviolet- could not have provided nectar for it. reflecting petals and a nonreflecting target area over the nectaries), whereas the ventral visitors MISCELLANEOUS INSECT VISITORS are supplied with their own guides (orange, ultra- LEPIDOPTERA violet reflecting undersurface of petals and elongate, wedge-shaped, nonreflecting sepals that At the Apache, Arizona site two species of point to the nectaries) and are therefore encour- skipper butterflies were present in small numbers aged by the flowers to avoid contributing, at taking nectar from the underside of the flowers. least directly, to the primary reproductive func- These were Pholisora catullus (Fabricius) and Ce- tion for which the flower was evolved, pollina- lotes nessus (Edwards). According to D. M. Mac- tion resulting in fertilization and seed set. Neil (in litt. ), the larvae of the former feed princi- Males among the insect visitors to Kallstroe- pally on various Chenopodiaceae and Ama- mia are interested only in nourishment and are ranthaceae and those of the latter on Malvaceae. involved in only the three categories of nectar Representatives of these families were present at collecting behavior. Females, on the other hand, the site. Pholisora catullus is a widespread North are not only responsive to the needs of their own American skipper; C. nessus, a southwestern energy supply but also to the need for storage of species. pollen and nectar for the consumption of their DIPTERA larval offspring. All but the honeybee store individual cells in the ground with various mixtures Diptera were seen at flowers of Kallstroemia of pollen and nectar and may, depending on the at each of the study sites but usually in small species and the stage in stocking the cell, be numbers. Exceptions were the ubiquitous syr- gathering pollen and nectar or one or the other. phid Heliophilus latifrons Loew and small bom- In the social honeybee, the determination on byliids of the genus Phthiria. The former brush which flowers they forage and what commodity the stamens as they extract nectar and frequently they gather is dictated by the hive, either by the have small amounts of pollen on the underside. behavior of the sister bees and scouts or by the Phthiria approach the nectaries below the sta- immediate food requirements of the hive. As in mens and avoid contamination with pollen. all of our observations of the honeybee at Kall- stroemia, workers gathered nectar only, it seems SUMMARY AND DISCUSSION reasonable to assume that the primary if not sole The brightly colored flowers of Kallstroemia activity of the hive or hives was that of honey grandiflora, their conspicuous ultraviolet reflec- manufacture and storage at this time of year, 18 AMERICAN MUSEUM NOVITATES

August 17. Also, it may be assumed that the sug- Linsley MacFarlane in obtaining field samples. ar content of Kallstroemia nectar was satisfac- Identifications of flower visitors were made or tory, as the flowers were visited frequently even confirmed by Dr. P. H. Timberlake (bees), Dr. though they were in competition with flowers R. M. Bohart (vespids, scoliids), Dr. K. V. Krom- of other plants. bein (tiphiids), Dr. E. I. Schlinger (flies), and Dr. Why the honeybee and several other species J. Powell (butterflies). Miss Celeste Green made approach the flower from the underside where it the accompanying sketches in part from photo- takes them longer to reach the nectaries, and how graphs by Mrs. Marjorie Statham Favreau. Dr. they learn this special technique is more difficult Duncan M. Porter kindly read the manuscript and to understand. If indeed Kallstroemia pollen is offered numerous helpful suggestions of botani- undesirable, as we have already suggested, it may cal importance. be responsible for the obvious avoidance of the LITERATURE CITED pollen and for the incorporation of such information into honeybee language, including infor- Austin, Daniel F. mation on the new approach system. However, it 1972. Interaction between Apis mellifera is is in (Hymenoptera: Apidae) and Tribulus by no means certain that the pollen any cistoides (Zygophyllaceae). Rhodora, way repellent. The grains range in size from vol. 74, pp. 117-123. 63.4,u-68.8,, average 66.7,i, which is well within Bequaert, J. the size range of insect transmitted pollens. That 1940. Synopsis of Monobia de Saussure, an it is adhesive may or may not be a disadvantage American genus of solitary wasps. Rev. but this appears to make it more easily carried Ent., vol. 11, pp. 822-842, 3 figs. and to minimize drying. However, there may be a Bradley, J. Chester more subtle, undesirable character that may ac- 1928a. A revision of the New World species of count for the fact that some species shake or Trielis a subgenus of Campsomeris (Hy- brush the pollen off after visiting a flower and menoptera: Scoliidae). Trans. Amer. or Ent. Soc., vol. 54, pp. 195-214. others partially completely avoid it. This type 1928b. The species of Campsomeris (Hymen- of behavior among the solitary bees and wasps is optera-Scoliidae) of the Plumipes even less expected unless they mimic the behav- group, inhabiting the United States, the ior of the honeybee or respond to the ventral Greater Antilles, and the Bahama Is- nectar guides frequently enough to fix the behav- lands. Proc. Acad. Nat. Sci. Phila- ior pattern. Further evidence that the pollen in delphia, vol. 80, pp. 3 13-337, 2 figs., itself is useful lies in the fact that 11 different 1 p1. species of bees are collecting and utilizing the Butler, George D., Jr. pollen in their economy. 1965. Distribution and host plants of leaf-cut- Although three other species of Kallstroemia ter bees in Arizona. Arizona Agr. Exp. have been shown to be intermittent in their Sta. Tech. Bull. no. 167, 19 pp. germination, it is by no means certain that seeds 1967. Biological observations on Ptilothrix sumichrasti (Cresson) in southern Ari- of Kallstroemia grandiflora germinate at irregular zona. Pan-Pacific Ent., vol. 43, pp. intervals. Whole areas may be covered with these 8-14, 1 table. poppies in the appropriate season of one year Butler, G. D., Jr., F. E. Todd, S. E. McGregor, and have none for the next several seasons, which and F. G. Werner strongly suggests an overriding environmental 1960. Melissodes bees in Arizona cotton control. Further studies are necessary to deter- fields. Arizona Agr. Exp. Sta. Tech. mine which factor or factors are responsible for Bull. no. 139, 11 pp. the sporadic appearance of this plant on the des- Butler, G. D., Jr., F. G. Werner, and M. D. Levin ert landscape. 1966. Native bees associated with safflower in south central Arizona. Jour. Kansas ACKNOWLEDGMENTS Ent. Soc., vol. 39, pp. 434-436. Cannon, W. A. We gratefully acknowledge the assistance of 1911. The root habits of desert plants. Carne- Juanita M. (Mrs. E. G.) Linsley and Mrs. Joan gie Inst. Washington, publ. no. 131. CAZIER AND LINSLEY: FORAGING BEES AND WASPS 19

Cockerell, T. D. A. 1967. A revision of the bees of the genus An- 1905. Descriptions and records of bees. I. drena of the Western Hemisphere. Part Ann. Mag. Nat. Hist., ser. 7, vol 16, pp. I. Callandrena (Hymenoptera: Andreni- 216-224. dae). Ibid., vol. 7, pp. 1-316, figs. 1908. Descriptions and records of bees. IVII. Levin, M. D., and G. E. Bohart Ibid., ser. 8, vol. 1, pp. 259-267. 1955. Selection of pollen by honeybees. Doull, K. M. Amer. Bee Jour., vol 95, pp. 392-393, 1966. The relative attractiveness to pollen- 402. collecting honeybees of some different Linsley, E. G. pollens. Jour. Agr. Res., vol. 5, pp. 9-14. 1960. Ethology of some bee and wasp-killing Evans, Howard E. robber flies of southeastern Arizona 1957. Studies on the comparative ethology of and western New Mexico (Diptera: Asi- digger wasps of the genus Bembix. New lidae). Univ. California Publ. Ent., vol. York, Comstock Publishing Co. (Corn- 16, pp. 357-392, pls. 48-55. ell Univ.), 248 pp. 1962. The colletid Ptiloglossa arizonensis 1960. A study of Bembix u-scripta a crepus- Timberlake, a matinal pollinator of So- cular digger wasp. Psyche, vol. 67, pp. lanum (Hymenoptera). Pan-Pacific Ent., 45-61. vol. 38, pp. 75-82. Faegri, F., and L. van der Pijl Linsley, E. G., and M. A. Cazier 1966. The principles of pollination ecology. 1963. Further observations on bees which New York, Pergamon Press, pp. 1-248. take pollen from plants of the genus Hurd, P. D., Jr., and E. G. Linsley Solanum. Pan-Pacific Ent., vol. 39, pp. 1972. Parasitic bees of the genus Holcopasites 1-18, 6 figs., 2 tables. Ashmead (Hymenoptera: Apoidea). 1972. Diumal and seasonal behavior patterns Smithsonian Contrib. Zool., vol. 114, among adults of Pro toxaea gloriosa pp. 1-41, 16 figs., 1 table. (Hymenoptera, Oxaeidae). Amer. Mus. Krombein, Karl V. Novitates, no. 2509, pp. 1-25. 1938. Studies in the Tiphiidae, II. A revision Linsley, E. G., and P. D. Hurd, Jr. of the Nearctic Myzininae (Hymen- 1959. Ethological observations on some bees optera Aculeata). Trans. Amer. Ent. of southeastern Arizona and New Soc., vol. 64, pp. 227-298. Mexico (Hymenoptera: Apoidea). Ent. La Berge, W. A. News, vol. 70, pp. 63-68. 1956. A revision of the bees of the genus Me- lissodes in North and Central America. Linsley, E. G., and J. W. MacSwain Part I (Hymenoptera, Apidae). Univ. 1957. The nesting habits, flower relationships, and parasites of some North American Kansas Sci. Bull., vol. 37, no. 2, pp. species of Diadasia (Hymenoptera: An- 911-1194, 129 figs. thophoridae). Wasmann Jour. Biol., vol. 1958. Notes on the North and Central Ameri- 15, pp. 199-235, 4 figs. can bees of the genus Svastra Holmberg 1958. The significance of floral constancy (Hymenoptera: Apidae). Jour. Kansas among bees of the genus Diadasia Ent. Soc., vol. 31, pp. 266-273. (Hymenoptera: Anthophoridae). Evolu- 1961. A revision of the bees of the genus Me- vol. pp. 219-223. lissodes in North and Central America. tion, 12, Part III (Hymenoptera, Apidae). Univ. Linsley, E. G., J. W. MacSwain, and R. F. Smith Kansas Sci. Bull., vol 42, no. 5, 1956. Biological observations of Ptilothrix su- 283-663, 125 figs. michrasti (Cresson) and some related 1963a. Records of the genus Svastra in North groups of Emphorine bees (Hymenop- America with the discription of a new tera: Anthophoridae). Bull. Southern species (Hymenoptera: Apidae). Jour. California Acad. Sci., vol. 55, pp. Kansas Ent. Soc., vol. 36, pp. 52-56. 83-101. pls. 25, 26. 1963b. New species and records of little known Linsley, E. G., and C. D. Michener species of Melissodes from North Amer- 1962. Brief notes on the habits of Protoxaea ica (Hymenoptera: Anthophoridae). (Hymenoptera: Andrenidae). Jour. Kan- Bull. Univ. Nebraska State Mus., vol. 4, sas Ent. Soc., vol. 35, pp. 385-389. pp. 227-242. figs. 20 AMERICAN MUSEUM NOVITATES

Martin, P. S., and C. M. Drew sas Sci. Bull., vol. 59, pp. 437-590, 1970. Additional scanning electron photo- 228 figs. micrographs of southwestern pollen Synge, A. D. grains. Jour. Arizona Acad. Sci., vol. 6, Jour. pp. 140-16 1, figs. 1-72. 1947. Pollen collection by honeybees. Meeuse, B. J. B. Animal E-col., Vol. 16, pp. 122-138. 1961. The story of pollination. New York, Timberlake, P. H. the Ronald Press, pp. 1-243. 1947. A revision of the species of Exomalop- Milliron, H. E. sis inhabiting the United States (Hy- 1973. A monograph of the Westem Hemi- menoptera: Apoidea). Jour. New York sphere bumblebees (Hymenoptera: Api- Ent. Soc., vol. 55, pp. 85-106. dae; Bombini) II. The genus Mega- 1954. A revisional study of the bees of the bombus subgenus Megabombus. Mem. genus Perdita F. Smith, with special ref- Ent. Soc. Canada, vol. 89, pp. 81-237. erence to the fauna of the Pacific Coast Porter, D. M. (Hymenoptera: Apoidea). I. Univ. Cali- 1969. The genus Kallstroemia (Zygophyl- fornia Publ. Ent., vol. 9, pp. 345-432, laceae). Contrib. Gray Herbarium, Har- figs. 1-72 vard Univ. no. 198, pp. 41-153, figs. 1955. Notes on the species of Psaenythia of 1-14, maps 1-9. North America. Boll. Lab. Zool. Gen. Ribble, D. W. Agr. "Filippo Silvestri," Portici, vol. 1965. A revision of the banded subgenera of 33, pp. 398-409. Nomia in America. Univ. Kansas Sci. 1960. A revisional study of the bees of the Bull., vol 45, pp. 277-359, 32 figs., 5 genus Perdita F. Smith, with special ref- tables. erence to the fauna of the Pacific Coast Roberts, Radclyffe B. (Hymenoptera: Apoidea). IV. Univ. 1972. Revision of the bee genus Agapostemon California Publ. Ent., vol. 17, pp. (Hymenoptera: Halictidae). Univ. Kan- 1-156. 3 3