Transactions of the Academy of Science of St. Louis “22! VOLUME XXV TITLE PAGE AND INDEX JANUARY, 1924 AUGUST, 1928 Published Under Direction of in Corea Elv ee JUN 10 i929 PPP PD Academy of Science of St. Louis LIST OF OFFICERS, 1928 2 a. Toone ” -VWice-PRESIDENT .e-eceeeneeenneneensnennene seo Loeb RECORDING GECRETARY 1: .cccccccceeceeeceeneeeeeeesee A. G. Pohlman CoRRESPONDING SECRETARY.........-------.J- I, Shannon ‘PRESIDENT TREASURER. H. E. Wiedemann LIBRARIAN oe ree Cire aii 8 ee H. C. Irish : ) Hermann von Schrenk CONTENTS. TABLE OF CONTENTS Last oF OFFICERS Papers PUBLISHED. January, 1924, to August, 1928: SE W. F. Cuapp.—Three New Species of Teredo, Plates I-II[I.—Issued April, 1924 R. Wauter Mitus.—Mediecal Fads and Fanceies. Lf o —Issued April, 1924 Coke Aueustus G. Pontman.—The Natural History Museum Movement in St. Louis.—Issued April, 1924 Puit Rau.—The Biology of the Roach, Blatta orientalis Linn.—Issued April, 1924 W. F. Cuapp.—Notes on the Stenomorphie Form of the Shipworm, Plates’ IV-V.—Issued January, 1925 J. T. BucHHouz anp E. J. PaumMer.—Supple- ment to the Catalogue of Arkansas Plants, Plates VI-X1I1.—Issued June, 1926... Put Ravu.—The Ecology ot a Sheltered Clay Bank; a Study in Insect Sociology, Plates XIV-XXI—Issued August, 1926.00. CHARLES Rosertson.—Florida Flowers and In- sects.—Issued September, 1927 Pui Rav.—Field Studies in the Behavior of the Non-Social Wasps, Plates XXII-XXXII.— Issued June, 1928. 16 29 121 49 166 i 4, Transactions of the Academy of Science of St. Louis / Volume XXV, No. 1° O.& _ THREE NEW SPECIES OF TEREDO W. F. CLAPP THREE NEW SPECIES OF TEREDO. W. F. Crapp. The material from which the following descriptions were drawn, was removed from test-blocks of wood placed in the water by the Committee on Marine Piling Investigations of the National Research Council. For descriptions of the test-blocks, and also for notes regard- ing the nomenclature used in the descriptions of the various characters of the shell and pallets, and for ref- erences to recent literature on the subject, consult Clapp, 1923, Proc. Bos. Soc. Nat. Hist., vol. 37, no. 2, pl. 3-4, p. 31-38, text fig. 1. I am deeply indebted to Prof. S. C. Prescott of the Massachusetts Institute of Technology, Cambridge, Mass., for laboratory facilities and for other assistance. ; Trans. Acad. Sci. of St. Louts TEREDO (TEREDO) PORTORICENSIS SP. NOV. Plate I, figs. 1-7. Shell subglobular, white, covered with a transparent, colorless: periostracum. The juncture of the anterior with the anterior-median area clearly marked by a broadly curved, slightly incised line. The ventral edge of the anterior area, forming an angle of about 100° with the anterior edge of the anterior-median area. Externally, the anterior area with the usual denticu- late ridges, the ventral posterior portion, with the ridges of about one-half the width of the intervening spaces. Dorsally the ridges are only one-fourth as wide as the spaces between them. Beginning at the ventral posterior edge, there are eight of these ridges to the millimeter, each ridge bearing approximately seventy denticles to the millimeter (fig. 1). The anterior-median area, at a point opposite the ventral edge of the anterior area, is one-quarter of the width of the entire median area. The denticulate ridges on this area, in a line continuous with the ventral edge of the anterior area, average twenty- five to the millimeter. In the type there are eleven of these ridges, which bear the usual broad denticles, there being twenty-eight denticles to the millimeter (fig. 2). The ventral ends of these ridges can be clearly seen continuing as sharp growth lines over the entire median area, but becoming less distinct on the auricle. The middle-median area is milk-white, in contrast to the semi-transparent anterior-median area, and is separated from the anterior-median area by a thin, narrow, trans- parent band. The posterior-median area is milk white and occupies one-half of the entire median area. The _ auricle is semi-transparent, showing more or less irreg- _ ular growth lines, and the periostracum is here thicker — than elsewhere on the shell. : Three New Species of Teredo 3 Internally, in the left valve a short, broad, flat hinge plate, directly beneath the umbone. The blade slightly more than one-half the length of the shell, of nearly uni- form width for its entire length, the lower half reflected posteriorly. The ventral knob large. The internal shelf of the auricle well marked. Pallets (fig. 3) of the type of Teredo navalis Linné. The stalk of about the same length as the blade, and merging into it by a gradual curve. The lower half of the blade white, the calcareous part of the upper half, as seen through the transparent chitinous covering, slightly cupped, the lateral portions extending farther distally than the median. The upper half entirely com- posed of transparent, yellowish horn-colored periostra- eum, which is deeply cupped distally for more than half its length, the outer surface being slightly less deeply cupped than the inner, and with a deep, narrow sinus - at the center, The posterior end of the tube with two short, nar- row, low ridges arising from opposite sides of the in- ternal wall; these ridges continue posteriorly beyond the shelly portion of the tube as sharp points. The type specimen (Mus. Comp. Zodl., 45303) is from San Juan, Porto Rico. Additional specimens from the type locality are also in the U. S. National Museum. The measurements of the type are: Total length of tube 40 mm. Shell, height 3.2 mm., length 3.1 mm. Pallets, length 3.8 mm., divided equally between blade and stalk, width of blade 0.8 mm. Teredo portoricensis is more closely related to Teredo bartschi Clapp than to any other described species. The variation of the shell characters in each species is so great that only very slight constant differences can be “s Trans. Acad. Sci. of St. Louis seen. The normal shell of a mature specimen of T. portoricensis is smaller than that of T. bartschi. The length of the apophysis always proportionately less. The partitions in the tube, while always present, are much lower and shorter than in 7. bartschi. The pallets resemble those of 7. bartschi, but constantly differ from them in having the blade longer and narrower, the june ture of the blade and pallet hardly perceptible, and the basal portion of the blade more gradually expanded. Seen through the periostracum, the calcareous portion of the blade of the pallet of 7. bartschi appears cone shaped, whereas that of T. portoricensis is the opposite, being deeply cupped at the center. In T. portoricensis the periostracum on the outer face of the blade is less deeply cupped than that on the inner, while in 7. bartscht the reverse is true. This species has been found in the test-blocks placed by the Committee on Marine Piling Investigations, at the following locations: Guantanamo, Cuba; San Pedro de Macoris, Santo Domingo; Port au Prince, Haiti; San Juan, Porto Rico; St. Thomas, Virgin Islands; Coco Solo, Panama, and (one specimen) Key West, Florida. At Guantanamo, Cuba, wood placed in the water on April 10th, contained specimens 10 mm. in length on May 10th; 60 mm. in length on June 10th, and 75 mm. in length July 10th. At Port au Prince, Haiti, wood submerged December 1, 1922, contained on January 1, 1923, many 5 mm. spec imens; on January 15th, 30 mm. specimens, and, on Feb- ruary Ist, 30 mm.:specimens with the gills well filled with many fully developed embryos. Wood placed in the water at this location on June 1, 1923, although well filled with several other species of shipworms, contained Three New Species of Teredo 5 no specimens of 7. portoricensis as late as September 3, At San Pedro de Macoris, Santo Domingo, wood sub- merged December 1, 1922, contained 20 mm. specimens on February 1, 1923, many of them with well-developed embryos. At San Juan, Porto Riico, wood placed March 20, 1923, contained on May 30th, 7. portoricensis 40 mm. long with embryos. At St. Thomas, Virgin Islands, wood placed April 1, 1923, contained many-30 mm. specimens on June Ist. At Coco Solo, Panama, wood submerged December 4, 1922, contained many 30 mm. specimens on January 19, 1923, and on February 19th, many specimens with well- developed embryos in the gills. It can be seen from the above records that this species may grow to be 60 mm. in length in a period of two months, or at the rate of approximately 1 mm. a day, and that specimens with a total tube length of but 20 mm. may possess well-developed embryos within the gills. Its rapid growth and early sexual maturity ren- der it one of the species most frequently found in the West Indies, and the destruction caused by it is con- siderable. Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PLATE (All figures reduced *%) Teredo portoricensis Denticulate ridges of the anterior portion. x 100. Denticulate ridges of the anterior-median portion. x 100. Pallets. x 14. Exterior of right valve. x 14. Exterior of left valve. x 14. Interior of right valve. x 14. TRANS. AcaAp. Scr. oF Sit. Louts, Vou. XKV PLATE ) FIG. 4 FIG. 2 FIG. 3 Three New Species of Teredo 7 TEREDO (ZOPOTEREDO) JOHNSONI* SP. NOV. Plate II, figs, 8-15. Shell subglobular, white, covered with a thin, nearly transparent, colorless periostracum. The narrow incised line separating the anterior and the anterior-median areas very slightly curved. The ventral edge of the an- terior area meeting the anterior edge of the anterior- median area in a nearly straight line, forming an angle of approximately 90°. Externally, the anterior area large, with many evenly spaced, denticulate ridges, which are of about the same width as the intervening spaces. There are sixteen of these ridges to the millimeter on the posterior-ventral portion of this area, each ridge bearing one hundred and twenty minute denticles to the millimeter (fig. 8). The anterior-median area occupying, at its widest part, one- third of the entire median area. The denticulate ridges on this area, along a line continuous with the ventral edge of the anterior area, average thirty to the milli- meter. There are twenty-six of these ridges in the type specimen, each ridge bearing approximately thirty-three denticles to the millimeter (fig. 9). The middle-median area is narrow, divided longitudinally into nearly equal halves, the anterior half, with the continuation of the den- ticulate ridges, showing as narrow, diagonally descend- ing growth lines, which curve upward, and become more or less obscure on the posterior half of the middle-median area. The posterior-median area large, occupying more than half of the entire median area, nearly smooth, show- ing only occasional, faint, incised growth lines. The auricle very small, being merely a continuation of the *I take pleasure in naming this species for Mr. A. A. Johnson, As- sistant to the Director of the Marine Piling Investigations Committee of the National Research Council. 8 Trans. Acad. Sci. of St. Louis posterior-median area, with no trace externally of a separating groove or concavity. Internally, a small, square hinge-plate in the left valve. The blade two-thirds of the length of the entire shell, thin, broad dorsally, the middle and ventral portions narrower, its entire length reflected slightly posteriorly. The ventral knob narrow, long, its base extending dor- sally for a considerable distance. The juncture of the anterior with the anterior-median area marked by a nar- row, thickened chord. The juncture of the auricle with the posterior-median area hardly visible. Pallets (fig. 10) with long, stout, opaque white stalks. The blade short, broad, investing a considerable por- tion of the upper part of the stalk in a thin sheath. The outer face convex. The proximal third of the blade eal- eareous, arising from the stalk in an abrupt curve. The middle third swollen, covered with a light horn-colored periostracum. The distal third covered with a dark chestnut colored periostracum with a deep central sinus dividing this portion of the blade into two shallow cups, the outer face considerably more deeply indented than the inner. The inner face of the blade flat, its distal two- thirds covered with a periostracum irregularly streaked with narrow bands of light and dark chestnut. The posterior end of the tube with a long delicate partition (fig. 11) dividing the tube into equal parts. The type (Mus. Comp. Zo6l., 45306) is from Guan- tanamo, Cuba. The measurements of the type are: Shell, height 4.5 mm.; length 4.5 mm. Pallets, total length 4.3 mm.; length of stalk 3 mm.;. width of blade 1.6 mm. The shell of this species is very closely related to that of Teredo clapy Bartsch (Proc. Biol. Soc. Washington, Three New Species of Teredo 9 1923, 56, p. 96). However the apophysis is shorter and narrower, the ventral knob smaller. The entire median area in Teredo johnsoni is always proportionately broader than in Teredo clappi. The range of variation of the denticulate ridges of both species is so great: that no constant difference can be established, but the angle ~ formed at the juncture of the ridges of the anterior area with those of the anterior-median area is always ap- proximately 90° in Teredo johnsoni whereas, in Teredo clapyi it is constantly obtuse, being rarely less than 100°. The pallets are very different from those of any pre- viously described species of Zopoteredo in that they are more nearly truly double-cupped, being in this respect more like Teredothyra. In Teredo somerst the charac- teristic median sinus can be seen only in the pallets of the very young, never persisting in the mature speci- mens. In Teredo clapyi it is frequently obscured or lost entirely in mature individuals, although always present in immature specimens. In Teredo johnsoni the sinus is constant in specimens of all ages. Specimens of this species have been found in the test- blocks placed by the Committee on Marine Piling Inves- tigations at the following locations: Guantanamo, Cuba; Port au Prince, Haiti; Fajardo and San Juan, Porto Rico; St. Thomas, Virgin Islands. This species is comparatively rare and the destruc- tion caused by it very slight. Frew specimens exceed 60 mm. in total length. At Guantanamo, Cuba, a test-block placed in the water on October 1, 1922, contained specimens 20 mm. in length on December 30th. A special block made of shingles, placed in the water in April, 1923, contained several 60 mm. specimens July 7, 1923. At Port au Prince, Haiti, 10 Trans. Acad. Sct. of St. Louis a test-block submerged June Ist and removed August 14th, contained Teredo johnsoni with 30 mm. tubes. At San Juan, Porto Rico, a test-block placed October 1, 1922, contained 30 mm. Teredo johnsoni on January 2, 1923, and wood placed in the water July 1, 1923, con- tained small specimens on August 1, 1923. At Fajardo, Porto Rico, wood submerged from January 9 till April 30, 1923, contained several specimens of this species. At St. Thomas, Virgin Islands, wood submerged four months and removed January 1, 1923, contained sev- eral specimens. any Si bee pgs . Sa es ee ge i ‘ eet Fig ge ", AAV Bie a as i 4 3 Ay Ab itvale ee et RAE bel gs att Aas : - L a yr DT. ‘ 4 oo af 4 ao ye ee ago Lg A atew tae Three New Species of Teredo EXPLANATION OF PLATE PLATE (All figures reduced 3/7) Teredo johnsoni See AOR OT ALLELE OP EONS Denticulate ridges of the anterior portion. x 130. Denticulate ridges of the anterior-median Ia x 130. Pallets. x Posterior portion of the tube. x 13. Exterior of right valve. Exterior of left valve. Interior of left valve. Interior of right valve. bal _ 13. x 13. = 13. 12 Trans, Acad. Sci. of St. Louts TEREDO (ZOPOTEREDO) FULLERI* SP. NOV. Plate III, figs. 16-22. Shell subglobular, small, bluish white, covered with a strong, transparent periostracum. 40 days 1 1 Total. 269 29 42 days 7 3 Thus we see that the average duration of life for the males was 40.2 days, and for the females 43.5 days. This difference is not great, but these few figures show at least that the females are inherently longer lived than the males, a condition that is often found in the insect world. The Biology of the Roach 63 The data gathered in 1916 indicated that July was the natural period of death for the roaches, both those kept for observation and for the population at large. By August 2nd, all those in the jars had died, and simul- taneously the adult roaches at large in the building bad disappeared. The housewife has a justifiable abhorrence of these creatures when they are adult; when thousands of nymphs infest her home, either she sees them not, or she bothers not about them, but the adults are con- spicuous and at once arouse her wrath. Forthwith she hies her to the nearest chemist and gets some good exterminator; two weeks later she will in good faith gladly tell the merits of ‘‘X¥YZ’’ paste, and one cannot convince her that she spread her poison just at the time when the roaches were dying a natural death, and that they would have disappeared at that time of the year regardless of whether or not she spread her paste. Number of egg cases deposited.—The female deposits during her lifetime from one to four egg cases. Fifty- five roaches were confined for the purpose of getting data on oviposition. The following table presents the data. Egg cases. Females. 1 8 2 19 3 19 4 9 The egg cases are not dropped as soon as they are formed, but they are slowly pushed out from the abdo- men, and after they have been carried about for two or — three days, they are finally dropped. I am inclined to suspect that the duration of the period of waiting is determined in greater part by inherent physiological 64 Trans. Acad. Sci. of St. Lous conditions, and in a lesser part to the temporary en- vironment, i. e., the dropping of the egg case is to some degree postponed until a place is reached where it may be deposited amid proper surroundings of moisture, tem- perature, seclusion, etc. Age of females at time of first oviposition—Records were kept of the lapse of time between the date when the female roach went through the final moult and became adult, and the day when she deposited her first egg case. These intervals were 8, 11, 11, 13, 14, 15, 18, 19, 22, and 24 days. Thus the normal females in this series never deposited before they were 11 days old (the 8-day individual was virgin), and one deposited her first egg case at the age of 24 days. Averages, however, are of no significance in this case, since in all probability egg laying is closely connected with mating, and no records were kept of the time that elapsed between mating and oviposition. This oviposition occurred from the third week in May to the middle of June, but no correlation could be detected between the age of the female at first oviposition and earliness or lateness in the season. Records were also kept on the number of days that elapsed between the various egg cases for 35 females. ‘As mentioned above, each fémale deposits during her life-time from one to four cases; these, however, do not follow in rapid succession, but at the following intervals: Days Days Days Roach between between between Number Ist & 2nd 2nd & 3rd 3rd & 4th 1 6 15 — 2 12 ae Se 3 8 oe as 4 11 ae ou The Biology of the Roach 65 Days Days Days Roach between between between Number Ist & 2nd 2nd & 3rd 3rd & 4th 5 10 ie ase 6 9 fe is | 11 RG, aoe 8 6 15 st 9 19 be in 10 6 12 ya 11 11 ie ae. 12 8 16 og 13 8 sds ie 14 8 8 pee 15 8 we ee 16 10 13 ae 17 10 13 ae 18 9 12 mit 19 23 ae -- 20 11 ies a 21 13 8 8 22 13 6 10 23 12 6 11 24 12 5 11 25 6 10 == 26 4 7 -< 27 4 6 -- 28 4 9 aia 29 12 -< a 30 11 =< mer 31 9 -= oe: 32 9 -- “ 33 9 10 14 34 10 14 i 66 Trans. Acad. Sct. of St. Louts These figures indicate that there is no appreciable dif- ference in the time required for the formation of the second, third or fourth egg case, and also that this inter- val is practically the same (except for a few cases) as the time elapsing between the last moult and the first oviposition, as shown on page 63. Both sexes were often kept together in the jars, and it is not known whether mating occurred more than once. The eggs were all fertile, but I suspect that these long lapses between egg cases are probably due to the maturation of the eggs, rather than a necessity of mating for each egg case. An interesting observation from the biological stand- point would be to test the fertility of the eggs in rela- tion to one or more matings. Period of incubation—The period during which egg cases were deposited extended from the latter part of May to the end of July. The period of incubation seems unusually long. At the time of the emergence of the nymphs, all leave the egg case at one time, instead of the alternative of the emergence covering a number of days for each case. The following data on the period of in- cubation were collected on egg cases deposited from May 21st to June 11th: Number of Number of Days egg cases | Days egg cases 45 8 52 6 46 7 53 6 47 3 34 1 48 1 59 5 49 1 56 4 50 3 61 1 _ 8 Total___-54 The Biology of the Roach 67 Thus we see all of these 54 egg cases (exclusive of the last, which might safely be regarded as abnormal), hav- ing a period of incubation varying from 45 to 56 days. The egg cases, as each was deposited, were placed in Jelly glasses and kept in the third floor laboratory. It seems to me that, if temperature has anything to do with hatching, this should have accelerated their development, for it was much warmer in the attic in July than in the cellar where the eggs normally are laid. Emergence of the young from the egg case-—When first I witnessed the hatching of the roaches, I was much surprised to find that their emergence is very similar to that of the mantis, Stagmomantis carolina. The roach egg cases are the well-known purses, the upper edge of which shows a ridge or seam. By one of those miracles which make marvelous every type of birth, this purse or capsule is pressed open at this seam at the moment of the emergence of the young, and the inmates are all released at the same time, after which, the pressure relieved, the capsule again closes and appears outwardly exactly the same as before. If one examines a number of egg cases previous to hatching time, and presses the ends together with the fingers in an attempt to open this seam, he will find them so tightly sealed that the case will mash under the pressure before this ridge will open. At hatch- ing time, however, they will yield readily, and at the slightest pressure will open beautifully, revealing the two halves of the case, each with the row of about eight mummy-like organisms, perfectly white except the eye- spots, which are black, and all arranged in two precise rows, heads up. In about three minutes after opening the case, all will have struggled out, and they are at once extremely active. This reveals the fact that when the egg case is deposited it has no opening, and only at the 68 Trans. Acad. Sci. of St. Louis time of emergence does the opening gradually form. One often wonders, upon picking up an empty egg case, how it is possible for the sixteen young to have emerged without leaving a break in the case. The mechanics is extremely interesting. As in the Carolina mantis, all the eggs are deposited with the head nearest the opening. The eggs, being produced practically at one time, or at least within forty-eight hours, all develop alike. In this mummy stage preceding emergence, the caudal end of the body tapers, while the greatest development is about the head. Now with all of the enlargement occurring at the fore part of the body, which is uppermost, it is easy to see how sixteen enlarging fore parts would mechani- eally and gradually open the exit, and simultaneously all would slide out en masse and automatically the closure would come together again. Some writers think that the young secrete a liquid which dissolves the cement in the seam of the capsule. This process is not merely an emergence, but also a real hatching, for in the process each insect issues from its individual egg shell and also from the dotheca enclos- ing a number of eggs. One often finds the thin, white caps that cover the forepart of the insect caught in the seam of the empty case, where the emerging nymphs have left them behind. In addition to this cap each mummy is completely enveloped in a thin, brown, papery bag, the egg shell. It is very pretty to see, when one splits the egg case in two parts, the two rows of heads facing the opening, wriggling and writhing out of this paper shell, and to see the two rows of heads, face to face, gradually make their appearance. I have said that at this stage the insect is completely white save the black eye-spots, but I have discovered with the aid of the microscope that the three teeth of each mandible are brown and seem to be The Biology of the Roach 69 thoroughly chitinized, the only heavily chitinized part - about the insect. When this mummy is removed from this bag, the head is inflexed, the legs, antennae and mouth-parts pressed flat against the body, and the segments very distinct—in fact, the whole organism looks like head and segments, and not at all like the roach it will be in five minutes. Even at this stage the two cerci are very prominent. The roach works itself out of the egg shell by a series of contortions, comes out through the opening and there sheds a white membranous cap, which, no doubt, is for the protection of the head. This frees its legs, and then the newly born roach clumsily walks away. This behavior is very much like that described for the praying-mantis.” Another irregular piece of semi-chitinous covering hangs from the tip of the abdomen, and this, I think, is the shed skin that bound the legs and antennae close to the body. With a few hard jerks this is left behind and the roach, for only now does it really appear as a roach, scampers briskly away. If the egg case is not artificially opened, it indeed seems like magic to see almost simul- taneously all heads protrude through the opening, struggle for an instant to get the limbs free, kick the hind legs loose from the adhering skin, and again simul- taneously scamper away in all directions, all in less time than it takes to tell. In opening many egg cases I found all of them fertile and in the same stage of development. In one case, how- ever, amid healthy looking organisms, were two eggs opposite each other, hard and brown, dead or infertile. Number of young in each egg case.—Notes were made on the number of nymphs that hatched from each of 43 egg cases. This number varied from 12 to 18, as follows: *Trans. Acad. Sci. St. Louis, 22:1-57. 1913. 70 Trans. Acad. Sci. of St. Louts No. in each Number of No. in each Number of egg case egg cases egg case egg cases 12 2 16 15 13 5 1% 3 14 6 18 oe 19 10 Total____43 It is at once apparent that the mean number at which nature aims is 16, and her deviations from that are not significant; even the mortality in embryo is surprisingly small. Food of nymphs and adults—During the period of observation an attempt was made to keep a record of food materials which they accepted. Soon, however, the list became so voluminous that it was apparent that it would be more brief and simple to list the things which they did not eat. This dwindled almost to nothing. They showed a preference for the softer portions of the food before them, and left the hard or chitinized portions, but otherwise they ate everything of a vegetable or animal nature, either fresh or decaying, which came in their way. Cockroaches are undoubtedly attracted to their food by the sense of smell located in the antennae. Lloyd Morgan * tells us that if their antennae be extirpated or coated with paraffin, the roach no longer rushes to food, and takes little notice of and will sometimes even walk over blotting-paper moistened with turpentine or benzo- line, which a normal insect cannot approach without agitation. Enemies.—Dermestes larvae often infest the egg cases of the roach. Some large frogs of the edible variety SAnimal Life and Intelligence, p. 257. 1891. The Biology of the Roach 71 escaped from a tank in the cellar one night; when they were recaptured and dressed the next day, the contents of the stomachs revealed an astonishing number of roaches in the large nymph stage. In nature the habitats of the frogs and roaches are so remote that this relation- ship can hardly be expected to exist. Some common gray lizards kept in the laboratory also readily accepted the roaches as food. ‘A neighbor told me that her chickens greedily ate the roaches which she had caught in traps. Woods’ says that the cockroach is a favorite food of many animals and the hedgehog is so partial to it that one of these animals is sometimes kept in the kitchen for the express purpose of destroying these pests. A young friend who is a nature student informs me that he fed roaches of this species to a pet garter snake. Roaches seem to require a larger supply of water than do most species of insects, and they drink it with an evi- dent enjoyment that is pleasant to see. Their food often becomes mouldy, and fungus growths of various colors, black, yellow, green or gray, cover it, but this seems not to affect them in any way, and no increase in the mor- tality could be noticed at these times. Very often, too, their food, especially the starchy materials, becomes covered with myriads of mites, Tyroglyphus lintneri Osborn (identified by Mr. H. E. Ewing), but even these do not affect the health or the mortality of the roaches in the jars. Method of concealing the egg cases.—The color of the egg case as it leaves the abdomen is almost white. Some- times the exposed part of the case becomes dark while it is being carried, so a newly deposited case may have every shade from white through a reddish pink to a red- ‘Insects at home, p. 237. 1872. 72 Trans. Acad. Sci. of St. Louis dish brown. Eventually all change entirely to a reddish brown. Although the adults and nymphs of the roach may appear in abundance about the house, the egg cases are rarely seen. The reason for this was discovered in the habits of those kept for close observation in the Jabora- tory. In the great majority of cases, the mother very deftly conceals the egg case, either by depositing it in a erevice or by carrying earth and debris, sometimes for considerable distances, and with this and her saliva, daubing the new case until it is quite hidden from view. This is so stealthily done that I have never been able to eatch her in the very act, but the result leaves no doubt that the act is deliberately and purposely done. Of course the insects in their native habitat have every chance to conceal the egg cases beyond all chance of our ever finding them; even in glass jars, with a limited amount of material with which to work, they succeeded so well that in a few cases I myself did not discover them until I had injured them in handling. A large number of females carrying egg cases were at different times placed in the jars, together with a small amount of earth and rubbish, just to see what would be their course of action. Out of 90 egg cases thus de- posited, 36 were deposited in some place which afforded some concealment, as crevices in bark, the cells of old wasp nests, under a bit of paper or, in a few cases, in slight depressions which, I was sure, the mother roach had made in bark or clay, expressly to fit and receive her dotheca. Many, if not the majority, of these were also daubed. Of those which were dropped on the bottom of the jars (this open space formed by far the greater part of the available space), 38 were well hidden by having been daubed with dirt and rubbish stuck on with saliva, The Biology of the Roach 73 and only 16 were left uncovered. It is only reasonable to surmise that the unnatural conditions of their life at this time and my frequent interruptions may have hin- dered the normal functioning of some of the mothers in so delicate a task as this, and while the mothers in con- finement concealed only 82 per cent of the egg cases, in their own environment and undisturbed, they would show a greater per cent of efficiency. As suggested above, if the egg cases had been dropped by mere chance, by far the greater part of them would have been on the floor of the jars. In many cases they were found crammed into crevices that must have been really dif- ficult of access, and in more than one instance, three or four lay end to end in the same cranny in the bark, when nooks were scarce. In some cases we know posi- tively that the mud must have been carried at least three inches. I repeat the assertion that the act of plastering the egg cases must be deliberately done. While these mothers were never caught in the act of this work, we have the interesting details of this behavior of an allied Species, Periplaneta americana Linn., by V. R. Haber.” He says that at 2:50 a. m., the female began to scar and roughen the surface of a cardboard in the cage. She chewed and munched at the upper surface of the paste- board until she had made quite an appreciable dent, not dropping the pasteboard on the bottom of the cage, but mixing the bits with a secretion from the mouth until all became a damp mass. At 3:30, she crawled forward over the scar with her abdomen bent anteriorly and ventral- ward, probing about with the protruding dotheca until she located the sear which she had just made. After sev-_ eral unsuccessful attempts at placing the egg case in the 10Ent. News 31:190-193. 1920. 74 Trans. Acad. Sct. of St. Louis scar, because each time it rolled off, it finally fell and bounced to the far side of the cage, whereupon she promptly ran down and located it on the floor. She cleaned it with her mouth-parts, coated its exposed sides and ends with a secretion from her mouth and then picked up loose bits of trash, with which she covered it. She even went so far as to try to cover it with a piece of paper. Girault” also found this habit of covering the egg case with mud or bits of wood prevalent for Peri- planeta australasiae Fab. The mating habits of the roach.—In the proper sea- son, a sudden switching on of the light often reveals a pair of roaches in copulo; they are motionless, and are united with their heads in opposite directions. The mated pairs are found on vertical walls, as well as on horizontal surfaces, but always in the same position. While I have seen many mated pairs, and have seen some attempts at mating, it still remains a mystery to me how this position is accomplished; it must be through some intricate movements. Mating always or nearly always occurs in darkness, and it is most difficult to observe the manoeuvers of these shy creatures, although by timing the experiments and by switching on and off the lights, one can get good records of the wooing. The ardor of the courting male is not cooled when the lights are turned on, but the modest little mate usually walks away and breaks up the party. In the preliminary movements the male gets directly in front of the female, but turns so his head is away from her (so that the two are in single file), with the rear tip of his body directly in front of her face. He then walks backwards, pushing his body underneath hers and at the same time lifts his wings and holds them vertically, NEnt. News 26:54. 1915. The Biology of the Roach 75 thereby exposing the dorsal segments of his abdomen. Sometimes a feeble attempt is made to flutter these ver- tical wings in a way that suggests coquettishness. When he has partly inserted his body under her, she slowly walks on top of his back, touching or feeling the seg- ments of his abdomen with her jaws and palpi. The cerci of the male are protruding and his clasping organs ex- tended and opening and closing like a pair of tongs, in his attempt to grasp or feel segment after segment as she crawls forward over him or as he crawls backward under her (sometimes it is difficult to say just which is really happening). These movements continue, and just when one expects a culmination, the female breaks away or the male makes a false step and the pair separate. There must be some very interesting contortions occur- ring before he makes the connections and faces about completely from backing under the female to facing in the opposite direction, and it is probable that many attempts are made before this intricate task is accom- plished. I hope some day, with better technique in the way of light regulation and mirrors, to be able to get the details. There is actual courtship on the part of the males, and there is actual rivalry between them, as will be seen from the following experiment. On June 2, at 11:35 p. m, I placed 9 chaste males about two weeks old (since they had become adult) with a virgin female. Five minutes later, upon switching on the lights, I found the female had crouched under a bit of bark, and at the end of ten minutes she had already mated with one of the males, and several of the others, walking high up in the air with legs outstretched like stilts (an extremely unusual position which might be a way of trying to attract the attention of the female), were pestering the pair. One 76 Trans. Acad. Sci. of St. Louis male in particular seemed to be biting the pair apart, and when I examined the spot after he had left I found that he had bitten away a large portion of the wing of his rival. I examined them at intervals for an hour, and found the pair still in copulo; the next morning they were separated, but there must have been some rivalry between the males, for two others had the wings badly bitten away. In another cage containing one female and four males, similar rivalry occurred; the female was pas- sive while the males ‘‘walked on stilts’? and protruded claspers. After some little display, three of these males retired and left her to the other suitor, which pursued her for fifteen minutes before he attempted to back under her. C. L. Turner” says the process of copulation in the cockroach is so rapid that the details cannot be followed; it can only be said that the male shoves his body under that of the female, and accomplishes the transfer of sper- matozoa in a few seconds. All those which I observed were much more deliberate, and in certain instances sev- eral hours were spent in actual mating. I have recorded* very similar courtship behavior for the wild roach, Parcoblatia pennsylvanica DeGeer, but here in the only case observed the female assumed the aggressive role. During the night males often go through mating ma- noeuvers with other males. Ezxudation.—In working with adults one soon notices at times a gathering of a thick, milky jelly on the ventral surface that seems to ooze out at the lines of demarca- tion of the last two segments. This is an almost trans- parent, slime-like substance of sufficient gumminess to 22Ann. Ent. Soc. Amer., 9:122. 1916. 18Trans. Acad. Sci. St. Louis., 24:57. 1922. The Biology of the Roach 77 hold itself together for many days, and is very notice- able on the black backs of the roaches. Just exactly where it comes from or what its function is, I do not know, but it probably oozes out from pores in thin chitin at the joints of segments and probably functions in some way in the mating or reproductive processes. This ma- terial was never seen to gather on the bodies of the males or on the mated females, but only on the virgin females. It is not wholly accurate to say that the mated females never showed this secretion; I should say, rather, that this material did not appear on the bodies of hundreds of females caged together with males, but at the same time in a group of virgin females that were segregated, almost all of them had varying amounts of this slime that exuded from the dorsal folds of the last few segments. Miscellaneous activities—Roaches are negatively phototactic and positively thigmotactic. Writers gen- erally agree that roaches are gregarious, but it seems to me that their gregariousness has more or less been thrust upon them; being negatively phototactic and posi- tively thigmotactic, and with large numbers of them seek- ing the same environment in limited space and hiding places, one can see how easy it is for the roach to appear as gregarious, but there probably is no more social ten. dency in the life of the roach than in any other member of the Orthopterous group. While the roaches are dark-loving creatures, there is a behavior that indicates a response which is so aptly called differential sensibility. This can be tested by sud- denly lighting up a roach infested cellar. The creatures run frantically until they come within the shadow of a post; there they stop short, and may easily be picked up. They seem certain of safety when in this darkness, 78 Trans. Acad. Sct. of St. Lows although what they respond to is not real darkness, but only a less light condition. In the same way, I have seen roaches escape into crevices; when their bodies touched the crack they were satisfied, even though they were ex- posed to full view in the light; their condition of positive thigmotropism satisfied their feelings in the matter of protection from danger, and they did not venture deep into the crevice. Here, however, it seems that light and sight were factors that prevented me from picking them up, for when attempts were made they crawled deeper into the crevices, whereas in the dark shadow on the floor an attempt was seldom made to escape because the roach could not see my approach. Roaches are very sensitive to odors or air conditions. One may handle the jar without disturbing the inmates if the jar is covered, or one may even remove the glass cover and come very near to look in if the breath is deflected, but to breathe, even gently, down into a jar at once creates a panic among the roaches. This simple test shows them to be susceptible to odor or air vibra- tions more than to sight. Their coming out of hiding does not seem to be regu- lated entirely by the quietness of the place. On many occasions I have found, when a house had been closed for several hours, roaches were out to feed as early as 8:30 or 9 o’clock; on other occasions they may be found out foraging when the room has been darkened only a few minutes, when the noises of the city have by no means subsided. On the other hand, during the night when everything is quiet and one expects them to be out in abundance, one may find few or none. It seems they feed as early in the evening as possible and then retire. The Biology of the Roach 79 I have never seen male roaches use their wings for flying. I have thrown them up in the air; they seem to have no control over the fall; they did not even open the wings to act as a parachute, but turned many somer- saults in the air and hit the earth with a thump. The males can, however, make their escape by backward pro- gression as rapidly as by traveling forwards. % 7 ot ee A 5) hen 3 Transactions of the ease - Science of St. Louis ae ntitgy Volume XXV, No. 5 we NOTES ON THE STENOMORPHIC FORM OF THE SHIPWORM W. F. CLAPP | vy eS Gur BORg, Issued January, 1925 eS e Wy . [ RB ‘ NOTES ON THE STENOMORPHIC FORM OF THE SHIPWORM. W. F. Crapp. The term stenomorph has been given by Dr. Bartsch (Bartsch, 1923, p. 330) to those specimens of shipworms which are abnormal in being dwarfed, although they may be sexually mature. Dr. Bartsch discovered the fact that this form is frequently found in small pieces of wood and inferred that it is caused by the cramped habitat of the individual. A study of the stenomorphic specimens found in the test-blocks placed by the Committee on Marine Piling Investigations of the National Research Council (Clapp, 1923, p. 31) has added further evidence of the conditions which cause this peculiar form. The shell of the stenomorphic specimen (pl. 4, fig. 1) may generally be distinguished from the normal speci- men of the same species by its smaller size and greater shell thickness, as well as by the proportionately greater number of denticulate ridges on both the anterior and the anterior-median areas of the shell. The internal blade or apophysis is greatly broadened and strength- ened (pl. 4, fig. 2). There is, however, no sharp dividing line between the stenomorph and the normal Specimen of the same species. Many large specimens, with tubes 250 millimeters or more in length, show traces of stenomorph characters in those portions of the shell last secreted. There is a well-graded series between these large, nearly normal specimens and those typical stenomorphs of the same age which possess tubes only ten millimeters long. In the typical stenomorph the auricle is entirely lost, being overlaid by the an- 82 Trans, Acad. Sci. of St. Louis terior-median area, since the denticulate ridges of this area completely cover the entire median area and also the auricle to its posterior edge. The anterior area is also proportionately large, the denticulate ridges of that area being added to its ventral edge until it is nearly as large as the anterior-median area both in length and in height. In a young normal specimen of Teredo navalis, the ridges of the anterior area are approximately one-fourth the width of the intervening spaces (pl, 4, fig. 3). In a typical stenomorphic speci- men of the same species and of the same size, the den- ticulate ridges of the anterior area are so closely crowded that the spaces between the ridges are reduced to mere threads (pl. 4, fig. 4). This dwarfed form, while the same age as a large normal individual, may be but one-tenth of its size. The embryos in the gills of a sexually mature stenomorphic specimen are much less numerous than are the embryos in a normal specimen of the same species, but in all other respects the embryos are identical. At many of the places where test-blocks have been placed by the Committee on Marine Piling Investiga- tions, these stenomorphic specimens are frequently found in the wood, and since the blocks used in 1922 were small (2 inches by 4 inches), it was thought that the abnormality was caused by the cramped quarters. As Dr. Bartsch states (loc. cit.), they are frequently found in laths and other small pieces of wood, but that some factor other than the size of the wood enters into the result, is shown by the fact that laths and small pieces of wood frequently contain large and perfectly normal specimens. For example, Teredo navalis causes considerable damage to the wooden lobster pots in the Notes on the Stenomorphic Form of the Shipworm 83 waters south of Cape Cod. The laths of which the lobster pots are made are often completely riddled in a few weeks. The number of stenomorphic specimens in these laths is no greater proportionately than one finds in heavy piling and the average size of a larger number of specimens removed from the laths is the same as that of specimens removed from big timber. ‘Also, large blocks, made up of many shingles bolted more or less tightly together (Mark, 1924, p. 266, footnote 3, p. 268, fig. EZ), were placed at fifty or more stations from Massa- chusetts to Texas. In some of these shingle blocks, stenomorphic specimens occurred, but in the majority the specimens were not stenomorphic, but nearly or en- tirely normal. Because of the fact that the average shipworm will endeavor to avoid crossing a large crack or open area in the wood, as long as progress is possible in some other direction, many of the shipworms in the shingle blocks made more or less of an effort to remain within a single shingle, the effort depending entirely upon how firmly the shingles were bolted together. When the specimens entered the thicker end of the shingle and advanced with the grain of the wood toward the thinner end, this effort to refrain from crossing to the adjacent shingle, resulted in a peculiarly abnormal form, in which the tubes are excessively attenuate. Many of these specimens were found which had attained a tube length of 400 millimeters in three months, but, owing to the thinness of the wood, had increased the diameter of the tube very slowly. The anterior ends of many of these long tubes were frequently but five milli- meters and occasionally only four millimeters in diameter. The shipworm is able to contract its body into one-half of its normal length. The body is firmly attached to the wall of the burrow only at the extreme 84 Trans. Acad. Sci. of St. Louis posterior end. In an effort to locate thicker wood an individual may occasionally withdraw its shell and the anterior portion of its body from the anterior portion of the tube, forming a new tube in a different direction. At the point of separation from the original tube a diaphragm is formed completely sealing the abandoned tube. This operation may be repeated several times by a single individual (pl. 5, fig. 3). Tubes of normal speci- mens of the same species which are 400 millimeters in length are approximately twelve millimeters in diameter at the anterior end. This abnormal form which occurs in thin wood, however, possesses none of the characters of the stenomorph. The thickness of the shell, arrange- ment of the denticulate ridges and the proportionate sizes of the anterior and anterior-median areas, being the same as in normal specimens, the abnormality caused by the thickness of the wood can be seen only in the excessively narrow form of the tube. The large proportion of abnormal specimens with attenuate tubes and the small number of true stenomorphs found in the shingles, would tend to show that the stenomorph characters are not caused by the size of the wood. Also, many of the small test-blocks used in 1922 con- tained in one portion of the block many hundreds of stenomorphic specimens, while in another equally large portion of the block there were only relatively a few normal specimens of the same species. For example, small test-blocks from Galveston, Texas, contained for several weeks during the summer of 1922, in one-half of the block, hundreds of specimens of stenomorphic Bankia gouldi, few being more than twenty millimeters in length, whereas in the other half of the block there were always two, three, or more large normal 200-300 Notes on the Stenomorphie Form of the Shipworm 85 millimeter specimens of the same species. Furthermore, at the same locality the increase in the size of the test- blocks in 1923 to 4x4x6 inches did not change the result in the slightest degree, the proportion of stenomorphs being as great and the average size of the stenomorphic specimens being the same in the large blocks as in the small. Also, several of the numerous specimens of piling received from various localities on the Atlantic Coast, some of which were 18 inches or more in diameter, showed the same stenomorph form, not separable from those found in the small test-blocks. From the above facts it is evident that the size of the wood is not necessarily the deciding factor in the production of the stenomorphs, for if it were, steno- morphs would not occur in large piles where plenty of wood remained unoccupied. In a great majority of the pieces of wood containing stenomorphic specimens which I have examined, I have found that the conditions at the time of the attachment to the wood of the embryo, were apparently very favorable. Many embryos entered the wood at the same time, frequently thirty or forty to the square centimeter. A large majority survived, but many specimens, owing to the fact that some of their neighbors entered the wood slightly earlier, or were more active in boring than they, soon found themselves unable to advance in any direction, for the reason that they were completely surrounded by the tubes of other specimens. The shipworm will never break through the partition of wood, however thin it may be, which sepa- rates it from a neighboring tube, whether the occupant of that tube be living or dead. These confined specimens continued to live for as long a period as those more fortunate individuals which were not checked in their 86 Trans, Acad. Sci. of St. Louis growth. Since it was not possible for them to increase the length or diameter of the tube or the size of the shell, the only noticeable activity was in the regular addition of denticulate ridges and an increase in the thickness of the shell, this peculiar growth rendering them almost unrecognizable. In view of the fact that the shipworm is supposed to be able to obtain more or less nutriment from the wood into which it bores (Dore and Miller, 1923), (Harrington, 1921), it is interesting to note that the stenomorphic specimen, at least, is not dependent upon the wood for food, for living specimens of steno- morphic Teredo navalis have been found with a total tube length of only ten millimeters and yet with a known age of ten months. A normal Teredo navalis during the active season, will form a ten-millimeter tube in less than two weeks time. That the animal of the steno- morph should continue to produce denticulate ridges when there is no more available wood upon which the denticles can be used, is even more interesting. I wish to express my appreciation for the assistance which has been given to me in my study of the shipworms at the Massachusetts Institute of Technology, through the kindness of Professor Samuel C. Presscott. Ex- cellent laboratory facilities, photographic apparatus, etc., have been freely placed at my disposal. I am also indebted to Mr. Nelson M. Fuller for the excellent photographs and for other assistance in my work. TrANS. Acap. Sci. oF St. Lours, Voit. XXV Pirate IV 1G 2 Notes on the Stenomorphic Form of the Shipworm EXPLANATION OF PLATE Piate IV External view of both valves of a ee specimen Teredo navalis. Actual size .94 mm Internal view of both valves of a stenomorph specimen Teredo navalis. Actual size .94 m External view of h valves of a normal specimen : .3 mm. ew of bot Teredo navalis. Actual size 1 External view of left valve of a stenomorph specimen Bankia gouldi. Actual size 2.7 m 87 of of of Pallets of stenomorph Teredo navalis shown in fig. 1. Actual i 5 mm. 88 Trans, Acad. Sci. of St. Louis EXPLANATION OF PLATE PLaTe V Fig. 4 secon Scene many stenomorphic specimens of Teredo lis from Warren, R. I. Fig. 2. aa showing attack of normal Teredo navalis from Fall River, Mas Fig. 3. Tube of a ee gouldi in a shingle block. TRANS. Acab. Sci. oF St. Louis, Vot. XXV PLATE V Notes on the Stenomorphic Form of the Shipworm 89 BIBLIOGRAPHY 1921. Harrington, C. R.—A note on the physiology of the shipworm Teredo norvegica). The Biochemical Journ. Vol. XV. No. 6, 741, fig. 1. pp. 736- 1923. Bartsch, Sg aes ali a new term in taxonomy. Science N. S. Vol. 57, p. 330. 1923. bea tok —— Pere ew species of Teredo from Florida. Proc. Bos <. Net. Hist. Vol. 37, No. 2, pp. 31-38, pl. 3-4. 1923. Dore, W. H. and R. C. Miller—The digestion of wood by Teredo navalis. Univ. of Cal. Publ. Zool. Vol. 22, pp. 383-400, pl. 18. 1924. Mark, E. L.—Marine borers in Bermuda. Proc. anes Acad. Arts and Sci. Vol. 59, No. 11. Coy Rm : at Transactions of the Academy of Science of St. Louis Volume XXV, No. 6 SUPPLEMENT TO THE CATALOGUE OF ARKANSAS PLANTS J. T. BUCHHOLZ and E. J. PALMER Se <9 ae < » Issued June, 1926 19.19%) Py SUPPLEMENT TO THE CATALOGUE OF ARKANSAS PLANTS.* J.T. Bucunouz ano EB. J. Paumen. More than thirty-six years have elapsed since the pub- heation of a Catalogue of the Flora of Arkansas. We are presenting here the first supplement to the List of Arkansas Plants by Branner and Coville (35). Many plant names have been changed since this publication, but we are not attempting to include names in this list where mere changes in synonymy are involved, except in a few instances where these changes are such as would not be included with the synonymy given in current botanical manuals. Our supplementary list of plants presented here consists almost exclusively of additions to the for- mer catalogue and includes also the species reported by various other botanists, both in publications appearing since 1890, and from reports rendered privately to the Department of Botany of the University of Arkansas within the last six years. One or more collection numbers are given where pos- sible. However, collections made by J. T. Buchholz were usually not numbered, and when thus marked, all num- bers used by this observer are below 1000. Nearly all of these specimens are in the Herbarium of the Uni- versity of ‘Arkansas. Collection numbers of Arkansas plants by E. J. Palmer are those given with numbers above 4000, unless otherwise stated, and duplicate speci- mens of many of these plants are in the Herbarium at Fayetteville. Most of the early collections by Palmer *Research Paper No. 22, Journal Series, University of Arkansas. (91) 92 Trans. Acad. Sci. of St. Lows are to be found in the Herbarium of the Missouri Botan- ical Garden, while a complete set of collections of all ligneous species with Palmer numbers is to be found in the Herbarium of the Arnold Arboretum. The flora of Arkansas has been largely neglected by botanists during the last thirty-six years. Those who have collected within the State have usually been re- warded by the discovery of many new and rare species of plants. It is safe to predict that as this flora becomes better known very many more additions will be made, and with a more careful and critical study of the material many new species and varieties will be added. The Catalogue of Plants by Branner and Coville (34) included 1610 ferns and seed plants. A small reduction must be made from this list for plants erroneously re- ported or those now represented under new names among our 534 additions. However, it may safely be estimated that the known ferns and seed plants of Arkansas now comprise about 2100 species. The earliest naturalists to visit and report on the vegetation of the area included in this state, of which there are available records, were Bradbury (1) in 1809-11, Schooleraft (2) in 1818-1819 and Nuttall in 1819. All of these early explorers gave valuable and interest- ing accounts of the geography and vegetation, but the celebrated botanist Nuttall described the vegetation in greatest detail and published valuable contributions in- cluding the descriptions of many new species of plants (4, 5, 7). Nuttall’s Journal of Travels Into ‘Arkansa Territory (3) is considered a classic in the literature of travel, and this together with his Collection Towards a Flora of the Territory of Arkansas (7), are noteworthy Supplement to Catalogue of Arkansas Plants 93 contributions to the flora of the West. His Flora of Arkansas was never completed, but he mentioned many plants in his Journal of Travels, and described others in his Flora and other contributions (4, 5). The works of Bradbury, Schoolcraft and Nuttall constitute the earliest reliable records on the flora of Arkansas. In 1834 Featherstonehaugh traveled through Missouri and Arkansas from the Missouri to the Red River. Though his report (6) was largely geological, he made frequent mention of many species of plants observed and the types of vegetation encountered. In 1859 Professor Leo Lesquereux made a study of the fossil and recent flora of the state for the Second State Geological Survey, and gave a description of the botani- cal features of the northern and northwestern counties and a catalogue of Arkansas plants (8), including those observed and reported by Nuttall. As Lesquereux did his work during October-December and covered very exten- Sive territory, he probably did not assemble an herba- rium. To this catalogue Butler added a list (9) of over a hundred species in 1877. Prof. F. L. Harvey published many short articles snd notes in the Botanical Gazette (10-15, and 17-29) between 1880 and 1885, including a more comprehensive treatise on the Ferns of Arkansas (17-18) and one on the arboreal flora (32). Harvey began to assemble an herbarium. A large portion of his collection is still preserved at the University of Ar- kansas, and many of his duplicates were exchanged with other herbaria. Professor R. Ellsworth Call, who was engaged in a Study of the geology of Crowley’s Ridge, prepared two 94 Trans. Acad. Sci. of St. Louts special reports on the native forest trees of this region (33, 37). He probably did not assemble any collections. J. Francis Williams made a special study of the timber of the Magnet Cove region (38) and reported a definite relation between the vegetation and the underlying igneous rocks, confirming a feature previously mentioned by Featherstonehaugh, but apparently not observed by Harvey (21). In the Annual Report of the Arkansas Geological Sur- vey for 1888 Branner and Coville (35) published the most recent comprehensive catalogue of Arkansas plants, a publication which appeared in 1891. This also in- cluded a discussion of the general botanical features of the state (35). Coville made some collections which are on deposit in the U. S. National Museum, but the bulk of the work done for the plant list seems to have been based on field notes. The Harvey collection at Fayetteville was worked over by Professor Simonds in preparing the Branner and Coville catalogue. An extensive collection made by Mr. E. N. Plank, who lived at Decatur, Arkansas, for many years, was acquired by the New York Botanical Garden. The Plank collec- tion contained many Arkansas plants, but he seems not to have published a list of these except in a single brief paper: ‘‘Concerning the Plants of Southwestern Ar- kansas’’ (41). The material collected by Plank and in- cluded in this report has been consulted at the herbarium of the New York Botanical Garden. In an ecological study, Dr. S. M. Coulter, (43) re- corded a number of swamp species new to Arkansas. Similarly Dr. Roland M. Harper (47-49) has made records of a number of species characteristic of our Supplement to Catalogue of Arkansas Plants 95 prairie and the coastal plain region of southeastern ‘Ar- kansas, containing names new to the state. More re- cently Dr. Harper and Rev. H. E. Wheeler sent us speci- mens representing unreported species, as credited to them. Others contributing data, specimens, or assist- ance in finding favorable collecting stations are: Mr. Ralph Shreve of Farmington, Prof. D. Demaree of Hen- drix College, Conway; Mr. W. W. Ashe of the Forest Service; staff members of the Agricultural Experiment Station, Fayetteville; some of the county agricultural agents and a number of former students of the Univer- sity of Arkansas. These miscellaneous unpublished records are included here with the name of the observer. From these, from some of our own publications, and for the most part from our other unpublished records, this list of additions to the catalogue of Arkansas plants has been prepared. A list of publications pertaining entirely or in part to the flora of Arkansas is given below. This is represented by a complete collection of these publica- tions, nearly all bound in eight volumes deposited in the University of Arkansas Library. PTERIDOPHYTA. Polypodiaceae: Polypodiwm virginianum L. (formerly reported as P. vulgare L.) Fayetteville, Washington Co.; Jasper, Newton Co.; Magazine Mountain, Logan Co., probably the southwestern limit of the species. Pteridium aquilinum var. pseudocaudatum Clute. Savoy, Washington Co. Pellaea glabella Mett. Bentonville (near Bella Vista), Benton Co. | 96 Trans. Acad. Sci. of St. Lowis Asplenium platyneuron forma serratum (KE. 8. Miller) R. Hoffman. On chert and limestone rocks, with the species. Polystichwm achrostichoides forma incisum (Gray) Gilbert. With typical form, chert hills, Northwest Arkansas, 7 Dryopteris spinulosa (Muell.) Kuntze. On north slope, Magazine Mountain, Logan Co. (D. M. Moore). Woodsia scopiina D. C. Eaton. On sandstone ledges, north side of Magazine Mountain, Logan Co., near the summit. Dennstaedtia punctilobula (Mich.) Moore. On par- tially shaded damp sandstone cliffs, north side of Magazine Mountain, Logan Co. 24142. Ophioglossaceae: Ophioglossum Engelmanni Prantl. 5376 Fulton, Hempstead Co., clay barrens; also in glades and rocky barrens of the Ozark region. Botrychium obliquum var. tenuifolium (Underw.) Gilbert. 24008 McNab, Hempstead Co. Botrychium obliquum var. dissectwm (Spreng-) Clute. Farmington, Washington Co. Low woods, rare, occurs with the species. Equisetaceae: Equisetum hyemale var. intermedium Fat. fide J- H. Schaffner. On calcareous deposits near spring, Jasper, Newton Co. Supplement to Catalogue of Arkansas Plants 97 Equisetum praealtum Raf. (E. robustum (A. Br.) A. 'A. Eaton.) Includes the plants formerly referred to E. hyemale. Along sandy river banks. Isoetaceae: Isoetes Butleri Engelm. Little Rock, on ‘‘Pu- laskite,’’ Fourche Mountain; Eureka Springs, Car- roll Co. (Bush). Isoetes melanopoda J. Gay. In springy ground, top of Magazine Mountain, Logan Co.; 24921 Hot Springs, Garland Co.; Farmington, Washington Co. MONOCOTYLEDONAB. Sparganiaceae: Sparganium americanum Nutt. Prairie View, Wash- ington Co.; 10598 Washington, Hempstead Co. ; 8358 Ashdown, Little River Co. Alismaceae: Sagittaria longirostra (Micheli) J. G. Sm. Fayette- ville, Washington Co.; 10599 Washington, Hemp- stead Co. Sagittaria papillosa Buch. 25043 Hazen, Prairie Co. Graminae: Erianthus strictus Baldw, Fulton, Hempstead Co. (4386 Greenman). Paspalum distichum L. Fulton, Hempstead Co. (4387 Greenman). Paspalum circulare Nash. 6363 Beaver, Carroll Co. 98 Trans. Acad. Sci. of St. Louis Paspalum elatum L. C. Rich. Corning, Clay Co. Paspalum laeviglume Scribn. 26210 Cotter, Baxter Co. Paspalum Muhlenbergii Nash. 6093 Corning, Clay Co.; 6370 Beaver, Carroll Co.; 8175 Fayetteville, Washington Co. Panicum Boscii Poir. 8426 Benton, Saline Co. Panicum commutatum Schultes. 25015 Little Rock; 24880 Hot Springs, Garland Co. Panicum Helleri Nash. Eureka Springs, Carroll Co. Panicum huachucae Ashe. 24827 Magazine Moun- tain, Logan Co. Panicum linearifolium Seribn. 5611 Eureka Springs, Carroll Co.; 25178 Shirley, Van Buren Co. Panicum lucidum Ashe. 25169 Shirley, Van Buren Oo. Panicum mutabile Scribn. & Smith. 24872 Hot Springs, Garland Co. Panicum polyanthes Schultes. 25053 Hazen, Prairie Co. Panicum Scribnerianum Nash. 5558 Eureka Springs, Carroll Co. Panicum tennesseense Ashe. Westfork, Washing- ton Co. Panicum Werneri Scribn. 5610 Eureka Siena: Carroll Co.; 25024 Hazen, Prairie Co. Echinochloa colona (L.) Link. 26727 McNab, Hempstead Co. Supplement to Catalogue of Arkansas Plants | 99 Digitaria Ischaemum Schreb, (Digitaria humi- fusa Rydb.) 4564 Eureka Springs, Carroll Co. Digitaria villosa (Walt.) Ell. Arkadelphia, Clark Co. (fide H. R. Rosen). Axonopus furcatus (Fluegge) Hitche. 10594 Wash- ington, Hempstead Co. Setaria viridis (L.) Beauv. Holcus halepensis L. (Sorgum halepense Pers.) Naturalized in waste places and widely distributed. Sporobolus canovirens Nash. 6908 Harrison, Boone Co. Sporobolus cryptandrus (Torr.) Gray. 6055 Corning, Clay Co. Sporobolus pilosus Vasey. 4374 Eureka Springs, Carroll Co. | Sporobolus vaginaeflorus Torr. Fide Standley and Hitchcock, from specimens by County Agr. Agent. Agrostis Elliottiana Schultes. Central Arkansas, from District Agr. Agent. Eragrostis glomerata (Walt.) Dewey. El Do- rado, Union Co. Bromus secalinus L. Warren, Bradley Co. Bromus tectorum L, Fayetteville, Washington Co. Sphenopholis pallens (Spreng.) Seribn. 25168 Shirley, Van Buren Co. Glyceria septentrionalis Hitche. 25046 Hazen, Prairie Co, Trans. Acad. Sci. of St. Louts Elymus arkansanus Scribn. & Ball. North Ar- kansas, District Agr. Agent. Elymus australis Scribn. & Ball. Fayetteville, Washington Co.; 8126 Benton, Saline Co. Elymus glabriflorus (Vasey) Scribn. & Ball. Lake Village, Chicot Co. Cyperaceae: Cyperus compressus L. 14641 McNab, Hempstead Co. Cyperus cylindricus (Ell.) Britton. Fulton, Hemp- stead Co. (fide J. M. Greenman). Cyperus ferax Rich. 26747 Fulton, Hempstead Co. Cyperus Houghtonti Torr. Fayetteville, Washington Co. Cyperus hystricinus Fernald. 14657 McNab, Hemp- stead Co. Cyperus lancastriensis Porter. 8124 Benton, Saline Co. Cyperus pseudovegetus Steud. 6061, 8026 Fulton, Hempstead Co. Cyperus rivularis Kunth. 4565 Eureka Springs, Carroll Co. Cyperus speciosus var. ferruginescens (Boeckl.) Britton. Fayetteville, Washington Co. Eleocharis Torreyana Boeckl. Lonoke Co.. woods of second bottoms. Fimbristylis Baldwiniana Torr. Stuttgart, Ar- kansas Co. (H. R. Rosen). A troublesome weed of rice fields. Supplement to Catalogue of Arkansas Plants 101 Fimbristylis castanea var. puberula (Michx.) Britton. 5531 Eureka Springs, Carroll Co.; Hazen, Prairie Co., and Stuttgart, Arkansas Co., where a troublesome weed in rice fields. Fuirena hispida Ell. 8099 Gifford, Hot Springs Co. Fuirena simplex Vahl. 6002 Cotter, Baxter Co.; 8370 Ashdown, Little River Co. Scirpus debilis Pursh. Lonoke Co., Portland clays (rice land soil). Rynchospora corniculata (Lam.) Gray. 8070 Arka- delphia, Clark Co.; 10520 Doddridge, Miller Co. Rynchospora cymosa Ell, South Arkansas, District Agr. Agent. Rynchospora gracilenta Gray. 8095 Gifford, Hot Springs Co. Rynchospora macrostachya Torr. 6071 Corning, Clay Co. Rynchospora Plankii Britton. Benton Co. (type locality) ; Washington Co. Carex arkansana Bailey. Arkansas River Valley. Carex Bicknellii Britton. 25060 Hazen, Prairie Co. Carex cherokeensis Schwein. 27104a Conway, Faulk- ner Co.; Pulaski Co., Roland M. Harper. Carex communis, Bailey. 26930 Magazine Mountain Logan Co.; 27098 Jasper, Newton Co. Carex crus-corvi Schuttlw. Corning, Clay Co. Carex debilis var. Rudgei Bailey. 27142 Conway, Faulkner Co.; 24938 Lonsdale, Garland Co. 102 Trans. Acad. Sct. of St. Louis Carex festucacea var. brevior (Dewey) Fernald. 27107a Conway, Faulkner Co. Carex glaucodea Tuckerm. 25069 Hazen, Prairie Co.; 24874 Hot Springs, Garland Co. Carex gynandra (Schwein.) Boott. 24927a Lons- dale, Garland Co. Carex hormathodes var. Richiit Fernald. 27007 Tontitown, Washington Co. Carex intumescens Rudge. 24990 Fulton, Hemp- stead Co. Carex Jamesit Schwein. Fayetteville, Washington Co.; 5581 Beaver, Carroll Co. Carex Joorii Bailey. 24004 Fulton, Hempstead Co. Carex awe Michx. 24747 Fayetteville, Wash- ington C Carex laxiflora var. gracillima Boott. 24746 Fay- etteville, Washington Co.; 24452 Hot Springs, Gar- land Co. Carex laxiflora var. latifolia Boott. 6038 Rush, Marion Co. Carex leptalea Wahlenb. 27132 Lofton, Hot Springs Co Carex louisianica Bailey. 27106 Conway, Faulk- ner Co.; Lonoke Co., heavy clay soil in woods. Carex lupulina var. pedunculata Bailey. 6047 Corn- ing, Clay Co. Carex Meadii Dewey. 5533 Eureka Springs, Carroll Co. Supplement to Catalogue of Arkansas Plants 103 Carex Muhlenbergii var. enervis Boott. 5552 Kureka Springs, Carroll Co.; 24815 Magazine Moun- tain, Logan Co. Carex oligocarpa Schkuhr. 27036 Goshen, Wash- ington Co.; 24849 Magazine Mountain, Logan Co. Carex platyphylla Carey. 26929 Magazine Moun- tain, Logan Co. Carex Sartwellii Dewey. Lonoke Co. Carex scirpoidea Michx. 24928 Lonsdale, Gar- land Co. Carex echinata var. angustata (Carey) Bailey. 27105a Conway, Faulkner Co. Carex stricta var. decora Bailey. 24745 Fayette- ville, Washington Co. Carex tetanica Schkuhr. 27041 Eureka Springs, Garland Co. Carex torta Boott. 26898 Lawrence, Garland Co. Commelinaceae: | Tradescantia brevicaulis Raf, Fayetteville, Wash- ington Co. On sandstone, cap rock of high hills. Tradescantia hirsuticaulis Small. Hot Springs, Garland Co. (Roland M. Harper). Dry woods near summit of West Mountain. Tradescantia occidentalis (Britton) Smyth. 25067 Hazen, Prairie Co.; Lonoke, Lonoke Co. Tradescantia reflexa Raf. England, Lonoke Co. Commelina communis L. (C. nudiflora Gray). Texarkana, Miller Co. 104 Trans. Acad. Sci. of St. Louis Juncaceae: Juncus aristulatus Michx. Hazen, Prairie Co. (Roland M. Harper). Juncus brachycarpus Engelm. 25065 Hazen. Prairie Co.; Jonesboro, Craighead Co., and 8S. E. Arkansas. (Roland M. Harper.) Juncus Dudleyi Wiegand. 5995 Cotter, Marion Co. Luzula campestris var. bulbosa A. Wood (Jun- coides bulbosum (Wood) Small). Most of our plants, probably all, belong to this variety or species. N. W. and central Arkansas. Liliaceae: Allium stellatum Ker. West Fork, Washington Co.; 4370 Eureka Springs; Sulphur Springs, Ben- ton Co. Iilium superbum L. Wet meadows, Fayetteville, Washington Co. (C. Chandler). Stenanthium robustum Watson. Fayetteville, Wash- ington Co.; 8205 Brentwood, Washington Co. Toxicoscordion Nuttallii (Gray) Rydb. 24929 Lons- dale, Garland Co.; Magazine Mountain, Logan Co.; Rudy, Crawford Co. Trillium pusillum. Michx. Valley of Osage Creek and Monte Ne region, Washington and Benton Co.; War Eagle Creek Valley, Madison Co. Occurs in cherty soil in woods (rare). Trillium Underwoodii. Small. Near Batesville, Independence Co. (‘‘Cherty slope near Polk Bayou’’—Roland M. Harper). Smilax hispida Mihl. 10558 Cotter, Marion Co.; 22207 Van Buren, Crawford Co. Supplement to Catalogue of Arkansas Plants 105 Dioscoreaceae: Dioscorea villosa var. glabra. Lloyd. 5945 Cot- ter, Marion Co. Amaryllidaceae: Hypoxis hirsuta (L.) Coville (H. erecta L.) 25027 Hazen, Prairie Co. Also frequent in Ozark region. Iridaceae: Iris cristata Ait. (Formerly reported as I. verna L.) Cherty hills in woods north and west of Fay- etteville, Washington and Benton Co.; similar situa- tions Madison and Carroll Cos.; top of Magazine Mountain, Logan Co. A form with white flowers has been collected at Farmington and Hot Springs. Iris foliosa. Mackenzie & Bush. Prairie Grove and Farmington, Washington Co. Rare and local (Ralph Shreve). Sisyrinchium albidum Raf. 25061 Hazen, Prairie Co.; Fayetteville, Washington Co. Sisyrinchium furcatum Bicknell. 25050 Hazen, Prairie Co. Sisyrinchium graminoides Bicknell. Fayetteville, Washington Co. Common throughout the state. Orchidaceae: Cypripedium pubescens Willd. A specimen sent from Red Star, Newton Co., compared with material at the Herbarium of the New York Botanical Gar- den was found to be typical for leaf and size; 24993 McNab, Hempstead Co. 106 Trans. Acad. Sci. of St. Louis Spiranthes ovalis Lindl. (Ibidium ovale (Lindl.) House). Conway, Faulkner Co. Spiranthes vernalis Engelm. & Gray. Fayette- ville, Washington Co.; 8146 Ozark, Franklin Co. Corrallorrhiza Wisteriana Conrad. Fayetteville, Washington Co.; Fulton, Hempstead Co. Pogonia verticillata (Willd.) Nutt. 26894 Law- rence, Garland Co. DICOTYLEDONAE. Salicaceae: Populus balsamifera var. virginiana (Henry) Sarg. Common Cottonwood. Formerly Populus deltoides Marsh. (P. monilifera Ait.) Distribution general along streams. Salix cordata Muhl. J ohnston, Washington Co. Salix nigra var. altissima Sarg. Tall black wil- low. 5394 Fulton, Hempstead Co. Saliaz longifolia Muehl. Sand Bar Willow. 24978, McNab, Hempstead Co.; 25013, Little Rock, Pulaski Co.; 25085, Helena, Phillips Co. Salix longipes var. Wardii (Bebb) Schneider. Ward’s willow. Fayetteville, Washington Co.; Eureka Springs and Beaver, Carroll Co.; Cotter, Marion Co. Salix longipes var. Wardii x nigra. 20500 Eureka Springs, Carroll Co. Leitneriaceae: Leitneria floridana Chapm, Cork tree. 4792 Moark, 6074 Corning, Clay Co.; St. Francis River swamps (S. M. Coulter). Supplement to Catalogue of Arkansas Plants 107 Juglandaceae: Carya alba var. ficoides Sarg. 20990 and 22198 Van Buren, Crawford Co. Carya alba var. ovoidea Sarg. 29441 McNab, Hempstead Co. Carya alba var. subcoriacea Sarg. The common form in Southern and Eastern Arkansas. Carya Buckleyi var. arkansana Sarg. 20991 Van Buren, Crawford Co. (type locality) ; 20638 Fulton, Hempstead Co.; 10524 Doddridge, Miller Co. Com- mon throughout the state. forma pachylemma Sarg. Fulton, Hempstead Co. (type locality). Carya cordiformis var. latifolia Sarg. 8219 Fay- etteville, Washington Co.; 20671 Cotter, Marion Co. Carya leiodermis Sarg. 8953 Fulton, Hempstead Co.; 26653 Helena, Phillips Co.; 26491 Mt. Nebo, Yell Co.; 26424 Magazine Mountain, Logan Co. Carya ovalis var. obovalis Sarg. Northern Ar- kansas. Carya ovalis var. obcordata Sarg. 22267 Fulton, Hempstead Co. Carya texana DC. Bitter Pecan. Van Buren, Crawford Co., bottom lands of Arkansas River (G. M. Brown), X Carya Brownii Sarg. Van Buren, Crawford Co., bottom lands of Arkansas River. A hybrid be- tween Carya cordiformis and C. pecan. X Carya Brownii var. varians Sarg. Van Buren, Crawford Co. (G. M. Brown). Carya alba x Buckleyi arkansana. 20989, 22210 Van Buren, Crawford Co. 108 Trans. Acad. Sci. of St. Lowis Betulaceae: Corylus rostrata Ait. Beaked Hazelnut. 26692 Jonesboro, Craighead Co. Fagaceae: Castanea alnifolia var. floridana Sarg. 413 Me- Nab, Hempstead Co.; 8440 Benton, Saline Co.; 26536 Hardinville, Faulkner Co. (C. Margaretta Ashe). Castanea ozarkensis Ashe. Ozark Chinquapin. (C. arkansana Ashe). War Eagle, Madison Co. (type locality); Benton, Washington, Carroll and other northern and western counties. (Formerly re- ferred to C. pumila.) Fagus grandifolia var. caroliniana Fern. & Rehd. Beech. 8076 Gum Springs, Clark Co.; Pettigrew, Madison Co., and along Mulberry Creek in Madison and Franklin Cos. The forma mollis Fern. & Rehd. has also been found at Helena, Phillips Co. Quercus arkansana Sarg. Arkansas Oak. 5383, 7188, 8043, ete. McNab, Hempstead Co. Locally com- mon in sand hills northwest of Fulton. Quercus borealis Michx. Coll. by W. W. Ashe in Ozark National Forest near Lurton, Newton Co. Quercus borealis var. maxima Ashe. Northern Red Oak. (Formerly erroneously called Q. rubra.) 6360 Beaver, 20510 Eureka Springs, Carroll Co.; 14300 Cotter, Marion Co.; Conway, Faulkner Co.; Fayetteville and Savoy, Washington Co., and in all of the northern, western and central counties. Supplement to Catalogue of Arkansas Plants 109 Quercus coccinea Wang. Scarlet Oak. Jones- boro, Craighead Co.; reported by R. M. Harper from near Forrest City, St. Francis Co. Quercus Durandii Buckley. Durand’s Oak. 5378, 6818, etc. Fulton, 8050 McNab, Hempstead Co. Quercus nigra var. heterophylla (Ait.) ‘Ashe. 23995 Texarkana, Miller Co.; 24011 McNab, Hemp- stead Co.; 24312 Shirley, Van Buren Co. Quercus palustris Muench. Pin Oak. 6069 Corning, Clay Co.; 21000 Van Buren, Crawford Co.; Conway, Faulkner Co.; Russellville, Pope Co.; Clarendon, Monroe Co. Reported by Harvey but omitted by Branner & Coville. Quercus Phellos L. Willow Oak. Conway, Faulk- ner Co.; Pine Bluff, Jefferson Co.; ‘Alma, Crawford Co.; Clarksville, Johnson Co. Reported by Harvey but omitted by Branner & Coville. Quercus prinoides Willd. Shin Oak, Prairie Oak. Benton Co., margins of prairies (F. L. Harvey). Quercus obtusa Ashe. Water Oak. Southern Arkansas. Q. rhombica Sarg.) Quercus rubra L. Southern Red Oak. (Formerly Quercus triloba Michx., Q. digitata Sudw., Q. falcata Michx.) Common throughout except in parts of Ozark region. Quercus rubra var. leucophylla Ashe. Little Rock, Pulaski Co.; Fulton, Hempstead Co.; Stutt- gart, Arkansas Co.; Conway, Faulkner Co. Quercus rubra var. pagodaefolia (Ell.) Ashe. Coastal Plains, S. E. Arkansas (fide Roland M. Harper). 110 Trans. Acad. Sci. of St. Louis Quercus Shumardu Buckley, Spotted Oak. (For- merly reported as Quercus texana.) Fulton, 20635 McNab, Hempstead Co.; Dermott, Chicot Co.; Sa- voy, Washington Co.; 6009 Cotter, Marion Co. Quercus Shumardii var. Schneckii (Britton) Sarg. Spotted Oak. 8182 Fayetteville, Washington Co.; Benton Co.; Madison Co.; 4453 Eureka Springs, Carroll Co.; 8949 Fulton, Hempstead Co. Quercus stellata Wang. Post Oak. (Quercus minor Sarg.) General distribution. Listed by Harvey but omitted by Branner and Coville. Quercus stellata var. araniosa Sarg. Sand Post Oak. 22454, 8985, Texarkana, Miller Co.; 8439 Ben- ton, Saline Co. Quercus stellata var. Margaretta Ashe. Southern Arkansas. Quercus velutina var. missouriensis Sarg. Ben- ton Co.; Washington Co.; Austin, Lonoke Co.; Con- way, Faulkner Co. Hybrid Oaks: X Quercus Bushii Sarg. (Q. marilandica x Q. velutina.) 20509 Eureka Springs, Carroll Co.; Fayetteville, Washington Co. (Mock St. and also on Lindell Ave.) X Quercus Rudkinii Britton. (Q. marilandica x Q. Phellos.) 10504, 23999, ete. Fulton, Hempstead Co. X Quercus Schochiana Dieck. (Q. palustris x Q. Phellos.) 27096 Dover, Conway Co.; Faulkner Co. (D. Demaree), Supplement to Catalogue of Arkansas Plants 111 X Quercus subfalcata Trel. (Q. Phellos x Q. rubra.) 22466, 24013, etc. Fulton and McNab, Hemp- stead Co.; Cove Creek, Faulkner Co. (D. Demaree). Quercus Durandii x stellata. 12659, 20715, ete. Mc- Nab, Hempstead Co. Quercus nigra x Shumardii. 22301 McNab, Hemp- stead Co. Quercus nigra x rubra. 23139, 24876, ete. Hot Springs, Garland Co. Ulmaceae: Ulmus serotina Sarg. 6944, Jasper, Newton Co.; 21013 Van Buren, Crawford Co.; 26603 Magnet Cove, Hot Springs Co.; 22277 McNab, Hempstead Co. Celtis laevigata var. texana Sarg. Southern Hackberry. 412 McNab, Hempstead Co.; 20491 Eureka Springs, Carroll Co. Common throughout the state. Celtis occidentalis var. canina (Raf.) Sarg. Fay- etteville, Goshen, Washington Co.; 14329 Cotter, Marion Co. Celtis occidentalis var. crassifolia. (Lam.) Gray. Rough leaved Hackberry. 20672 Cotter, Marion Co.; 22208, 21012 Van Buren, Crawford Co. Celtis pumila var. georgiana (Small) Sarg. Savoy, West Fork, Washington Co. Moraceae: Broussonetia papyrifera (L.) Vent. (P apyrius papyrifera (lL). Kuntze). Cultivated, occasionally escaped. Paper Mulberry. 412 =: Trans. Acad. Sct. of St. Louis Aristolochiaceae : Asarum reflezum. Bicknell. Johnson, Goshen, Kessler Mountain, West Fork, Washington Oo. Frequent in Ozark region. Aristolochia reticulata. Nutt. 8058 McNab, Hemp- — stead Co.; 8384 Ashdown, Little River Co. Reported in Nuttall’s Arkansas Flora but omitted from all subsequent lists. Polygonaceae: Eriogonum longifolium Nutt, 14319 Cotter, Marion Co. In sandy soil. Eriogonum hirsutum Nutt. North Arkansas and Missouri border region, confined to caleareous rocks. Polygonum densiflorwm Meisn. (Persicaria per toricensis Small.) St. Francis River, Mississipp! Co. (S. M. Coulter). Chenopodiaceae : Chenopodium Botrys L. 14304 Cotter, Marion Co. Amaranthaceae: Froelichia gracilis Moq. Fayetteville, 8215 Brent- wood, Washington Co. Nyctaginaceae: Boerhaavia erecita L. 8065 Arkadelphia, Clark Co.; 24257 Hot Springs, Garland Co. Supplement to Catalogue of Arkansas Plants 113 Caryophyllaceae: Cerastium brachypodum (Engelm.) Robinson. Short Stalked Chickweed. Fayetteville, Washington Co. Waste fields. Cerastium semidecandrum L. Fayetteville, Wash- ington Co. Stellaria media (L.) Cyrill. Common Chickweed. Waste places over entire state, sometimes flowering during all months of winter. Saponaria Vaccaria L. Cow herb. Conway, Faulk- ner Co., near tunnel (H. E. Wheeler). Silene nivea (Nutt.) Otth. Fayetteville, Washington Co., escaped from cultivation. Portulacaceae: Talinum calycinum Engelm, 4563 Eureka Springs, Carroll Co.; 26215 Cotter, Baxter Co. Talinwm parviflorum Nutt. Thin soil above sandstone cap rock of waterfall near Winslow, Washington Co.; Fourche Mountain, Little Rock, on *‘Pulaskite’’ rocks. Nymphaeaceae: Cabomba caroliniana Gray. Near Blytheville, Mississippi Co.; St. Francis River swamps (S. M. Coulter). Ranunculaceae: Anemone caroliniana Walt. Fayetteville, Wash- ington Co.; Conway, Faulkner Co. 114 Trans. Acad. Sci. of St. Louis Anemone decapetala Ard. Along sandstone cliffs, south slopes, near base of Magazine Mountain, Logan Co. Delphinium Nortonianum Mackenzie & Bush. Fay- etteville, Washington Co. Delphinium Treleasei Bush. 27038 Eureka Springs, Carroll Co. Delphinium Penardii Huth. 5585 Beaver, Car- roll Co. Frequent in barrens and on prairies of Ozark region. Ranunculus hispidus Michx. 5597 Eureka Springs, Carroll Co. Thalictrum texanum (Gray) Small, 7151 Fulton, Hempstead Co. Clematis versicolor Small. (Viorna versicolor Small.) 6352 Beaver, Carroll Co.; 8226 Westfork, Fayetteville, Washington Co. Clematis reticulata Walt. (Viorna reticulata Small.) 10508 Columbus, Hempstead Co. Clematis paniculata Thunb. 20498 Eureka Springs, Carroll Co. Introduced along rocky creek. Anonaceae: Asimina parviflora (Michx.) Dunal. Crowley’s - Ridge north of Helena, Phillips Co. (fide Roland M. Harper.) Menispermaceae: Cocculus carolinus (L.) DC. (Epibaterium car- olinum Britton). 6005 Cotter, Marion Co.; 22193 Van Buren, Crawford Co.; Fayetteville, Washing- ton Co. Common throughout the state. Supplement to Catalogue of Arkansas Plants 115 Calycocarpum Lyoni (Pursh) Nutt. 25153 Helena, Phillips Co.; 26541 Cove Creek, Faulkner Co.; 24290 Shirley, Van Buren Co. Fumariaceae: Corydalis campestris (Britton) new comb. (Cap- noides campestre Britton). Benton Co. (Type locality). Cruciferae: Coronopus didymus (L.) J. E. Sm. 24989 Fulton, Hempstead Co.; Conway, Faulkner Oo. A trouble- some pasture weed. Sisymbrium altissimum L. (Norta altissima Brit- ton). 8158 Ozark, Franklin Co.; Washington Co. Erysimum inconspicuum (Wats.) MacM. North- west Arkansas (HE. N. Plank). Camelina sativa (L.) Crantz. Gold of Pleasure. Fayetteville, Washington Co. Naturalized from Europe. Dentaria laciniata Muhl. 24855 Magazine Moun- tain, Logan Co. Cardamine pennsylvanica Muhl. 24458 Hot Springs, Garland Co.; Fayetteville, Washington Co. Cardamine parviflora L. 7176 Fulton, Hempstead Co.; 24451 Hot Springs, Garland Co.; Conway, Faulkner Co.; Fayetteville, Washington Co. Droseraceae: Drosera annua E. L. Reed. Conway, Faulkner Co., Campus of State Teachers College. 116 Trans. Acad. Sci. of St. Louis Saxifragaceae: Sazifraga texana Buckl. (Micranthes texana Small). Conway, Faulkner Co.; Fayetteville, Washington Co. Heuchera arkansana Rydb. (May be the H. villosa of Harvey.) On cliffs, Northwest Arkansas. . Heuchera hirsuticaulis (Wheelock) Rydb. Shaded north slopes, on Boone chert hills, Washington Co. Heuchera macrorhiza Small. Fayetteville, Wash- ington Co. Heuchera parviflora Bartl, Springdale, Wash- ington Co.; Shirley, Van Buren Co. On moist cliffs. Flowers in October. . Philadelphus pubescens Schrad. Cove Creek, Faulk- ner Co.; 6934 Jasper, Newton Co.; 6033 Rush, Marion Co.; Shirley, Van Buren Co.; Magazine Mountain Logan Co. Hydrangea cinerea Small. 6959 Heber Springs, Cleburne Co.; 25160 Helena, Phillips Co.; 25205 Shirley, Van Buren Co. Ribes Cynosbati L. Prickly Gooseberry. (Gross- laria Cynosbati Mill.). 242 Rich Mountain, Polk Co.; Magazine Mountain, Logan Co. Kibes: missouriense Nutt. Gooseberry (Grossul- aria missouriensis Coville & Brit.) 5572 Beaver, Carroll Co. Hamamelidaceae: Hamamelis macrophylla Pursh. Witch-hazel. 249 Rich Mountain, Polk Co.; 8441 Benton, Saline Co.; 22296 McNab, Hempstead Co. Supplement to Catalogue of Arkansas Plants 117 Hamamelis vernalis Sarg. Spring-blooming Witch- hazel. 6032 Rush Marion Co.; 12649, Mena, Polk Co.; Goshen, Washington Co. Common along rocky streams of Ozark region. Hamamelis vernalis var. tomentella Sarg. 20494 Eureka Springs, Carroll Co. Rosaceae: Physocarpus intermedius (Rydb.) Schneider. Nine Bark. (Opulaster intermedius Rydb.) 4385 Eureka Springs, Carroll Co.; 5573 Beaver, Carroll Co..; 5982 Cotter, Marion Co. Probably the Neillia opulifolia Benth. & Hook., observed by Lesquereux and by Har- vey, belongs here. Malus angustifolia (Ait.) Michx. Narrow-leaved Crabapple. 20719 Arkadelphia, Clark Co. Malus iowensis var. Palmeri Rehd. 10581 Wash- ington, Hempstead Co. Crataegus. Crus-galli group: Crataegus bellica Sarg. 5335, 5336 Fulton, Hemp- stead Co, (type locality). Crataegus Bushii Sarg. 5344 Fulton, Hempstead Co. (type locality); 277 Rich Mountain, Polk County; 24531 High Point, Garland Co. € Crataegus erecta Sarg. 21017 Van Buren, Craw- ford County. Crataegus Palmeri Sarg. Fayetteville, Washington Co. Crataegus paradoxa Sarg. 20993 Van Buren, Craw- ford Co. 118 Trans. Acad. Sci. of St. Louis Crataegus pilifera Sarg. 5540 Eureka Springs, Carroll Co.; Van Buren, Crawford Co. Crataegus subpilosa Sarg. Eureka Springs, Car- roll Co. (type locality). Crataegus triwmphalis Sarg. 5371, 6825, Fulton, Hempstead Co. (type locality). Crataegus palliata Sarg. Fulton, Hempstead Co. (type locality). Crataegus villiflora Sarg. 5519, 5522. Eureka Springs, Carroll Co.; 20992, 22205 Van Buren, Crawford Co. | Crataegus. Punctatae group: Crataegus fastosa Sarg. Fulton, Hempstead Co. (type locality). Crataegus secta Sarg. 19028 Sulphur Springs, Benton Co. Crataegus sordida var. villosa Sarg. 20709, 22266 McNab, Hempstead Co. Crataegus sucida Sarg. 4387 Eureka Springs, Carroll Co.; 8179 Fayetteville, Washington Co. Crataegus verruculosa Sarg. 24453 Hot Springs, Garland Co. Crataegus. Virides group: Crataegus amicalis Sarg. Fulton, Hempstead Co., (type locality). Crataegus blanda Sarg. 20712 McNab, Hempstead Co.; 24496 Hot Springs, Garland Co. Crataegus micrantha Sarg. Fulton, Hempstead Co. (type locality). Supplement to Catalogue of Arkansas Plants 119 Crataegus velutina Sarg. Fulton, Hempstead Co. (type locality). Crataegus viridis L. 4789 Moark, Clay Co.; 5345 Fulton, Hempstead Co.; 8134 Little Rock, Pulaski Co.; 8221 Greenland, Washington Co. The collec- tions credited to Harvey as C. arborescens Ell. prob- ably belong here, since this is the species occurring in the Fayetteville region. Crataegus. Intricatae group: Crataegus Harveyana Sarg. Eureka Springs, Car- roll Co. (type locality); 26849 Hot Springs, Gar- land Co. Crataegus leioclada Sarg. 4388, 5539, 5547, Eureka Springs, Carroll Co. Crataegus neobushii Sarg. 5546 Eureka Springs, Carroll Co. Crataegus villicarpa a 5525 Eureka Springs, Carroll Co. Crataegus padifolia var. incarnata Sarg. 6026 Cotter, Marion Co. ; 26611 Magnet Cove, Hot Springs Co. Crataegus. Tenuifoliae (?) group: Crataegus lacera Sarg. Fulton, Hempstead Co. (type locality). Crataegus. Molles group: Crataegus arkansana Sarg. Type (at Arnold Arboretum) from seed collected at Newport, Ar- kansas, 120 Trans. Acad. Sci. of St. Louis Crataegus brachyphylla Sarg. 432, 8975, 9393, 9392, 10607, 16333, 20711, McNab, Hempstead Co. (type locality). Crataegus invisa Sarg. 5397, Fulton (type lo- cality), 6879, 7193, 7196, McNab, Hempstead Co. Crataegus induta Sarg. Fulton, Hempstead Co. (type locality). Crataegus limaria Sarg. a Hempstead Co. (type locality). Cartaegus transmississippiensis Sarg. 8419 Cot- ter, Marion Co. (type locality). Crataegus. Dilatatae group: Crataegus coccinoides Ashe. Farmington, Washing- ton Co. Crataegus. Uniflorae group: Crataegus trianthophora Sarg. 6970 Heber Springs, Cleburne Co.; 26535 Cove Creek, Faulkner Co.; 26885 Lawrence, Garland Co.; 14322 Cotter, Marion Co.; Fulton, Hempstead Co. (B. F. Bush 5690) ; Baker Springs, Howard Co. (J. H. Kellogg). Crataegus. Brachyacanthae group: Crataegus brachyacantha Sarg. & Engelm. 315 Texarkana, Miller Co.; 8386 Ashdown, Little River Co. Crataegus. Macracanthae group: Crataegus carrollensis Sarg. Eureka Springs, Carroll Co. (type locality). Crataegus hispidula Sarg. 5542 Eureka Springs, Carroll Co. Supplement to Catalogue of Arkansas Plants 121 Crataegus mollicula Sarg. 6023 Rush, Marion Co. Crataegus pudens Sarg. 20998, 22204 Van Buren, Crawford Co. Crataegus tomentosa lL. This species, reported by Branner & Coville, is found in ‘Arkansas, but the variety punctata Gray (C. punctata Jacquin), reported also by them, probably does not occur here. Crataegus aestivalis (Walt.) T. & G., C. coccinea L., C. flexispina (Moench) Sarg. and C. subvillosa Schrad., listed for Arkansas by Branner & Coville, very probably are not found here and should be dropped from the state plant list. C. cordata Ait. (=C. phaenopyrum (L. f.) Medic.) and C. crus-galli L. reported by Harvey may occur in the northern counties. Crataegus hybrids: : X Crataegus notha Sarg. A hybrid between C. apifolia and C. brachyphylla. 8974, 20646, 20716 McNab, Hempstead Co. Alchemilla arvensis Scop. Lady’s Mantle. 23133 Hot Springs, Garland Co. Potentilla recta L. Fayetteville, Washington Co. Prunus mexicana Watson. Mt. Nebo, Yell Co.; Rich Mountain, Polk Co.; 7149 Fulton, 16329 McNab, Hempstead Co.; 20474 Eureka Springs, Carroll Co.; 20668 Cotter, Marion Co. Prunus mecicana var. fultonensis Sarg. 12663 Ful- ton, Hempstead Co. (type locality). 122 Trans. Acad. Sci. of St. Louis Prunus mexicana var. polyandra Sarg. Fulton, Hempstead Co. Prunus Munsoniana Wight & Hedrick. Fulton, Hempstead Co. Prunus wnbellata Ell. 8109 Gifford, Hot Springs Co. ; 10597 Washington, Hempstead Co. Prunus umbellata var. tarda (Sarg.) Wight. 105395 Arkadelphia, Clark Co. Prunus angustifolia var. varians Wight. 7148 Ful- ton, Hempstead Co. Rosa Eglanteria L. (Rosa rubiginosa L.) Escaped from cultivation, Fayetteville, Washington Co. Rosa Lyonii Pursh. 24957 Hot Springs, Garland Co.; 24994 McNab, Hempstead Co.; 25055 Mesa, Prairie Co.; 25103 Helena, Phillips Co. Rosa Palmeri Rydb. 8067 Ashdown, Little River Co. Rosa setigera var. tomentosa T. & G. 22294 Mc- Nab, Hempstead Co. ; 5978 Cotter, Marion Co. ; 20507 Eureka Springs, Carroll Co. Rosa subserrulata Rydb. 8159 Ozark, Franklin Co. ; 10531 Ft. Lynn, Miller Co.; 6960 Heber Springs, Cleburne Co.; 8137 Little Rock, Pulaski Co. Rosa texarkana Rydb. Type collected by Eggert, Texarkana, Miller Co. Rubus occidentalis L. 5574 Beaver, Carroll Co. Common throughout Ozark region. Rubus ostryaefolius Rydb. 16315 Texarkana, Mil- ler Co. Supplement to Catalogue of Arkansas Plants 123 Rubus Andrewsianus Blanchard. Fulton, Hemp- stead Co. The common high blackberry throughout the State. Rubus rubrisetus Rydb. 327 Texarkana, Miller Co.; 20727 Gum Springs, Clark Co. Leguminosae: Amorpha croceolanata Watson. (A. fruticosa var. croceolanata Schneider). 948 Cove Creek, Faulkner Co.; 23051, 26863 Hot Springs, Garland Co.; 26918 Magnet Cove, Hot Springs Co. Amorpha nitens Boynton. 23148, 26862 Hot Springs, Garland Co.; 26915 Magnet Cove, Hot Springs County. Amorpha tennesseensis Shuttl. 20659 Cotter, Marion Co.; 24251 Hot Springs, Garland Co.; Van Buren, Crawford Co. (G. M. Brown). Amorpha glabra Desf. 24187 Magazine Moun- tain, Logan Co.; Hot Springs, Garland Co.; Rich Mountain, Polk County. Baptisia sulphurea Emgelm. Little Rock, Pu- laski Co. (H. E. Wheeler). Cassia Medsgeri Shafer. Fayetteville and West- fork, Washington Co.; 6885 Eureka Springs, Car- roll Co.; Cotter, Marion Co. Frequent in Ozark region and perhaps throughout the state. Cladrastis lutea (Michx.) K. Koch. Kessler Mountain, near Farmington, Washington Co.; Mag- azine Mountain, Logan Co.; Shirley, Van Buren Co.; 8408 Cotter, Marion Co. 124 Trans. Acad, Sci. of St. Louis Lathyrus latifolius L. Perennial Pea. Fayette- ville, Washington Co. Escaped from cultivation. Lespedeza procumbens Michx. Fayetteville, Wash- ington Co.; Eureka Springs, Carroll Co. Medicago arabica Huds. Spotted Medick. Fay- etteville, Washington Co. (R. H. Austin). Escaped from cultivation. Medicago lupulina L. Black Medick. Fayette- ville, Washington Co. Introduced. Melilotus officinalis (L.) Lam. Yellow Sweet Clover. Washington Co., waste places, introduced and becoming common in many parts of the state. Melilotus alba Desv. White Sweet Clover. In- troduced and becoming abundant all over the state. Desmodium paniculatum var. pubens T. & G. 4475 Eureka Springs, Carroll Co. Petalostemum villosum Nutt. 5994 Cotter, Marion Co. Phaseolus polystachyus (l.) B. S. P. Wild Kid- ney Bean. 8396 Cotter, Marion Co.; 10592 Wash- ington, Hempstead Co. Sesbania macrocarpa Nutt. Fulton, Hempstead Co. (4405 J. M. Greenman). Stizolobium Deeringianuwm Bort. Velvet Bean. McNab, Hempstead Co. (4436 J. M. Greenman). Wistaria macrostachya Nutt. Fulton and Me- Nab, Hempstead Co. Strophostyles umbellata (Muhl.) Britton. 12659 McNab, Hempstead Co. Supplement to Catalogue of Arkansas Plants 125 Strophostyles helvola (L.) Britton. Fayette- ville, Washington Co. Trifolium hybridum L. Alsike Clover. Fayette- ville, Washington Co. Introduced. (R. H. Austin). Trifolium dubium Sibth. Fayetteville, Washing- ton Co.; Eagle Mills, Ouachita Co. Introduced. Vicia angustifolia (L.) Richard. 24510 Hot Springs, Garland Co. Introduced. Vicia sativa L. Fayetteville, Washington Co. Introduced. Vicia tetrasperma (L.) Moench. 5395 Fulton, Hempstead Co. Introduced. Geraniaceae: Erodium cicutarium (L.) L’Hér. University of Arkansas campus, Fayetteville (A. D. Oxley). Geranium pusillum Burm, f. University of Ar- kansas campus, Fayetteville. Oxalidaceae: Oxalis Brittonae Small. England, Lonoke Co. Oxalis texana (Small) Fedde. Introduced in favored places or as weed in greenhouses, Oxalis Langliosii (Small) Fedde. Fayetteville, Washington Co. Oxalis interior (Small) Fedde. Benton Co. (type locality) (E. N. Plank). Linaceae: Linum medium (Planch.) Small. Hazen, Prairie Co. (Roland M. Harper). 126 Trans. Acad. Sci. of St. Louis Zygophyllaceae: Tribulus terrestris L. Ft. Smith, Sebastian Co.; 6094 Corning, Clay Co. Rutaceae: Ptelea polyadenia Greene. 5923, 5921 Cotter, Marion Co.; 6957 Heber Springs, Cleburne Co. Zanthoxyllum americanum Mill. Savoy, Wash- ington Co. Meliaceae: Melia Azedarach L. Escaped from cultivation, Fulton, Hempstead Co. Euphorbiaceae: Andrachne phyllanthoides (Nutt.) Muell. Arg. 5528 Eureka Springs, 27053 Berryville, Carroll Co.; Magazine Mtn., Logan Co.; Hot Springs, Garland Co. Croton Engelmanni var. albinoides Ferg. Ful ton, Hempstead Co.; Texarkana, Miller Co.; Ash- down, Little River Co. Crotonopsis elliptica Willd. 6962 Heber Springs, Cleburne Co., also common in barrens of Ozark re- gion. Euphorbia arkansana Engelm. & Gray. (Tithy- malus arkansanus Kl. & Garcke). Clark Co. (C. Woolsey). Euphorbia commutata Engelm. (Tithymalus com- mutatus Kl. & Garecke). Eureka Springs, Car- roll Co. (C. Woolsey). Supplement to Catalogue of Arkansas Plants 127 Euphorbia Nuttallii Engelm. (Chamaesyce Nuttallii Small). 6333 Beaver, Carroll Co. Tragia ramosa Torr. 4382 Eureka Springs, Carroll Co. Anacardiaceae: Cotinus americanus Nutt. American Smoke-tree, Chittamwood. 5971, 10556 Cotter, Marion Co.; Van Buren, Crawford Co. (G. M. Brown). Rhus trilobata Nutt. Polecat-bush. (Schmaltzia trilobata Small). 4384 Eureka Springs, Carroll Co.; 4754, 8411 Cotter, Marion Co. Common on rocky bluffs of Ozark region. Schmaltzia serotina Greene also occurs in the northern part of the state and may be distinct. Rhus quercifolia (Michx.) Steud. Poison Oak. 258 Rich Mountain, Polk Co.; 24810 Blue Mountain, Logan Co.; 25052 Hazen, Prairie Co. Aquifoliaceae: Ilex caroliniana (Walt.) Trel. 12661, 14648 Mc- Nab, Hempstead Co.; 23060 Hot Springs, Garland Co.; Cove Creek, Faulkner Co. Aceraceae: Acer floridanum (Chapm.) Pax. McNab, Hemp- stead Co.; Crowley’s Ridge (Roland M. Harper). Acer leucoderme Small. 24508, 26474, ete. Hot Springs, Garland Co.; Baker Springs, Howard Co. (J. H. Kellogg). Acer saccharum var. glaucum (Pax.) Sarg. 14313 Cotter, Marion Co.; 20488 Eureka Springs, Carroll Co. The common Sugar Maple of the state. 128 Trans. Acad. Sci. of St. Louis Acer saccharum var. Rugellii (Pax.) Rehd. Eureka Springs, Carroll Co.; near Hindsville, Madison Co. Acer rubrum var. Drummondii (Hook. & Arn.) Sarg. (Acer Drummondii Hook. & Arn.) 7164 Ful- ton, Hempstead Co.; St. Francis River swamps (S. M. Coulter). Acer rubrum var. tridens Wood. 8364 Ashdown Little River Co.; Fulton, Hempstead Co. Acer Negundo var. texanum Pax. 6977 Heber Springs, Cleburne Co.; 8072 Arkadelphia, Clark Co.; 14639 Cotter, Marion Co.; 22261 Fulton, Hempstead Co. The form latifolia Sarg. is also found at Fulton. Hypocastanaceae: Aesculus discolor var. mollis Sarg. Red-flowered Buckeye. (Aesculus austrina Small). 8078 Gum Springs, Clark Co.: Austin, Lonoke Co.; Fulton, Hempstead Co. Common over the state except in parts of the Ozark region. : Aesculus glabra var. micrantha Sarg. Fulton, Hempstead Co. (type locality). Aesculus glabra var. pallida Kirch. 8203 Wins- low, Washington Co. Aesculus glabra var. leucodermis Sarg. (Reported by Harvey as A. glabra). Fayetteville, Washington Co.; 4442 Eureka Springs, Carroll Co. X Aesculus Bushii Schneider. Fulton, Hemp- stead Co. (type locality). (B. F. Bush). A supposed hybrid between A. glabra var. leiodermis and A, discolor var. mollis. Supplement to Catalogue of Arkansas Plants 129 Sapindaceae: Sapindus Drummondit Hook & Arn. 5949 Cot- ter, Marion Co.; Fulton, Hempstead Oo.; Savoy, Washington Co. Rhamnaceae: Ceanothus ovatus Desf. 21015 Van Buren, Crawford Co.; Farmington, Washington Co. Ceanothus pubescens (T. & G.) Rydb. North Arkan- sas, exposed limestone ledges. Rhamnus lanceolata Pursh. 5931, (17239) Cotter, Marion Co.; 20495 Eureka Springs, Carroll Co. Vitaceae: Ampelopsis cordata Michx. False Grape. Fayette- ville, Washington Co.; 5933 Cotter, Marion Co. Parthenocissus quinquefolia var. hirsuta (Don.) Planch. 216 Huntsville, Madison Co.; 22189, 22209. Van Buren, Crawford Co. Parthenocissus quinquefolia var. Saint-Paulii (Koehne & Graebn.) Rehd. 5909 Cotter, Marion Co. ; 22269 Fulton, Hempstead Co. Vitis cinera Engelm. Winter Grape. 5932 Cotter, Marion Co. Common throughout state. Vitis Linsecomii var. glauca Munson. Post Oak Grape. 5554 Eureka Springs, Carroll Co.; 22191 Van Buren, Crawford Co.; 228 MeNab, Hempstead Co.; Fayetteville, Washington Co. Vitis rotundifolia Michx. Muscadine. Conway, Faulkner Co.; 257 Rich Mountain, Polk Co.; For- rest City, St. Francis Co.; 6839 Columbus, Hemp- stead Co.; 8470 Gifford, Hot Springs Co.; 6956 Heber Springs, Cleburne Co. 130 Trans. Acad. Sci. of St. Louis Vitis palmata Vahl. 6052 Corning, Clay Co.; 10526 Ft. Lynn, Miller Co.; 12667 McNab, Hemp- stead Co. Tiliaceae: Tila caroliniana Mill. 16336 McNab, Hempstead Co. Tilia caroliniana var. rhoophila Sarg. 8074 Gum Springs, Clark Co. Tilia floridana Small. (Tilia pubesceus Ait., re- ported by Coville, may belong here). 7204, 9401 Me- Nab, Hempstead Co.; 20512 Eureka Springs, Car- roll Co.; 20657 Cotter, Marion Co.; 20729 Gum- Springs, Clark Co. Tilia floridana var. hypoleuca Sarg. 5943 Cot- ter, Marion Co.; Kessler Mountain near Farming- ton, Washington Co. Tilia nuda Sarg. Fulton, Hempstead Co. Malvaceae: Callirhoe alcaeoides (Michx.) Gray. University of Arkansas campus, Fayetteville. Callirhoe Bushii Fernald. Baldwin, Savoy, Wash- ington Co. Callirhoe triangulata (Leavenworth) Gray. 5977 Cotter, Marion Co. Hibiscus incanus Wendl. 466, 16321 Fulton, Hemp- stead Co. Hibiscus lasiocarpus Cav. 8365 Ashdown, Little River Co.; 26638 Helena, Phillips Co. Malvastrum angustatum Gray. 6910 Harrison, Boone Co. Supplement to Catalogue of Arkansas Plants 131 Hypericaceae: Ascyrum multicaule Michx. 22289 McNab, Hemp- stead Co. Hypericum apocynifolium Small. Hazen, Prairie Co. and §. E. coastal plain. (Roland M. Harper). Hypericum pseudomaculatum Bush. Fayetteville, Washington Co.; 5957 Cotter, Baxter Co.; 8155 Ozark, Franklin Co. Hypericum oklahomense E. J. Palmer. Cove Creek, Faulkner Co. Cistaceae: Lechea villosa Ell. 8122 Malvern, Hot Springs Co. Helianthemum rosmarinifolium Pursh. 8162 Piney, Johnson Co. Violaceae: Viola eriocarpa Schwein. Johnson and Elm Springs, Washington Co. (V. pubescens Ait, of Branner and Coville, probably belongs here). Viola fimbriatula J. FE. Sm. 420 and 7153 Me- Nab, Hempstead Co. Viola Lovelliana Brainerd. Mena, Polk Co. (HK. Brainerd). : Viola missouriensis Greene. Osage Creek, Wash- ington Co.; Benton Co.; 5393, 7165, Fulton, Hemp- stead Co. Viola primulifolia L. Washington, Hempstead Co.; West Mountain near Hot Springs, Garland Co. Viola sororia Willd. Fayetteville, Washington Co.; 4782 Moark, Clay Co. 132 Trans. Acad. Sci. of St. Louis Viola triloba Schwein. (This may be Harvey’s ‘var. cordata). Mt. Comfort, Washington Co.; 5374, 7150, 9390 Fulton, Hempstead Co. | Viola triloba var. dilatata (Ell.) Brainerd. Mena, Polk Co., West Fork, Washington Co.; (Westville, Okla. E. Brainerd). Viola papilionacea Pursh. 24465 Hot Springs, Garland Co.; Fayetteville, Washington Co.; Mena, Polk Co, (KE. Brainerd). Viola striata Ait. Fayetteville, Washington Co. Viola viarum Pollard. Along White River. (H. Brainerd). Lythraceae: Cuphea petiolata (L.) Koehne, (Parsonsia petiolata Rusby). 4423 Eureka Springs, Carroll Co. Onagraceae: Oenothera linearis Michx. (Kneiffia linearis Spach). Fayetteville, Washington Co.; 23218 Top of Maga- zine Mountain, Logan Co.; 5404 Fulton, Hemp- stead Co. Oenothera missouriensis Sims. (Megapterium mis- souriense Spach). Eureka Springs, Carroll Co. Oenothera arenicola (Small) Coker. (Kneiffia arent- cola Small). 25038 Hazen, Prairie Co. Cornaceae: Cornus alternifolia L. Sylamore, Stone Co. (W. W. Ashe), also in Washington Co. Cornus stricta Lam. 6090 Corning, 10591 Moark, Clay Co.; 12649a Mena, Polk Co.; 8363 Ashdown, Little River Co.; 16324 Fulton, Hempstead Co. « Supplement to Catalogue of Arkansas Plants 133 Umbelliferae: Ammoselinum Butleri (Engelm.) Coult. & Rose. Credited to State in Coulter & Rose, Monograph of Umbelliferae. Angelica villosa (Walt.) B. S. P. 6024 Rush, Marion Co. Chaerophyllum texanum Coult. & Rose. 5409 Fulton, Hempstead Co. Daucus Carota L. Along roadsides and railroads, Fayetteville, Washington Co. Erigenia bulbosa (Michx.) Nutt. West Fork, Washington Co. Eryngium prostratum Nutt. 6048 Corning, Clay Co.; 8166 London, Pope Co. Hydrocotyle bonariensis Lam. Coastal Plain of Arkansas. (Roland M. Harper). Ptilimnium Nuttallii (DC.) Britton. 8027 Ful- ton, Hempstead Co.; 8164 Piney, Johnson Co. Ptilimnium capillaceum (Michx.) Raf. 26682 Jonesboro, Craighead Co. Sium suave Walt. (Sium cicutaefolium Schrank). St. Francis River swamps. (S. M. Coulter). Torilis Anthriscus (L.) Bernh. Fayetteville, Wash- ington Co. (Miss C. Chandler). Zizia aurea (L.) Koch. 5559 Eureka Springs, Carroll Co. Zizia cordata (Walt.) DC. 5557 Eureka Springs, Carroll Co. 134 Trans. Acad. Sci. of St. Lows SYMPETALAE., Ericaceae: Monotropa Hypopitys L. (Hypopitys americana Small). Prarie View, Washington Co., densely wooded north slopes of chert hills. Rhododendron oblongifolium (Small) Millais. Aza- lea, Honeysuckle. (Azalea oblongifolia Small). 6932 Jasper, Newton Co.; 6973 Heber Springs, Cle- burne County; 10505 Columbus, Hempstead Co.; 10515 Doddridge, Miller Co. Rhododendron rosewm (Loisel.) Rehd. Azalea, Honeysuckle. 260 Rich Mountain, Polk Co.; 26466 Meaford, Garland Co.; Batesville, Independence Co. ; Cove Creek, Faulkner Co. Rhododendron canescens (Michx.) G. Don (Azalea canescens Michx.). Yellville, Marion Co. Vaccinium arboreum var. glaucescens (Greene) Sarg. (Batodendron glaucescens Greene). 6942 Jasper, Newton Co. Vaccinium vacillans Kalm. Fulton, Hempstead Co. and occasional through Ozark region, but much less common than the following variety. Vaccinium vacillans var. crinitum Fernald. 5556 Eu- reka Springs, Carroll Co.; 5989 Cotter, Marion Co. Common throughout Ozark region. Primulaceae: Dodecatheon brachycarpa Small. Wheeler, Wash- ington Co. Steironema intermedium Kearney. Fayetteville, Washington Co. Supplement to Catalogue of Arkansas Plants 135 Sapotaceae: Bumelia lycioides (L.) Pers. 25137, 26649 Helena, Phillips Co.; 26725 McNab, Hempstead Co. Ebenaceae: Diospyros virginiana var. platycarpa Sarg. Large- fruited Persimmon. 20669 Cotter, Marion OCo., also in northwestern counties. Styracaceae: Halesia monticola var. vestita Sarg. Silver-bell Tree. 255 Rich Mountain, Polk Co.; 6978 Heber Springs, Cleburne Co.; 24926 Hot Springs, Garland Co. Oleaceae: Forestiera acuminata var. vestita E. J. Palmer. Miller Co. (type by Bush); Fulton, Hempstead Co. ; Van Buren, Crawford Co. Ligustrum vulgare L. Privet. 14321 Cotter, Marion Co. Escaped from cultivation. Fraxinus pennsylvanica var. lanceolata (Borck.) Sarg. Green Ash. 6073 Corning, Clay Co.; 5942 Cotter, Marion Co.; 8080 Arkadelphia, Clark Co.; Fayetteville, Washington Co. Common along streams throughout state. Fraxinus profunda Bush. Pumpkin Ash. 6067 Corning, Clay Co.; St. Francis River swamps (S. M. Coulter), According to Harvey, Fravinus nigra Marsh (P. sambucifolia L.) is found in ‘Arkansas. This is prob- ably an error which has been repeated by others. 136 Trans. Acad, Sci. of St. Louis No specimens are to be found in any of the Amer- ican herbaria and Harvey left no specimens in the University of Arkansas collections. Gentianaceae: Bartonia virginica (L.) B. S. P. 10586 Washing- ton, Hempstead Co.; 26476 Lonsdale, Garland Co. Centaurium texense (Griseb.) Fernald. (Erythraea texensis Griseb. Northern Arkansas border counties. Gentiana flavida Gray. (Dasystephena flavida Britton.) Magazine Mountain, Logan Co. Apocynaceae: Amsonia ludoviciana Vail. Fayetteville, Wash- ington Co.; Lonoke, Lonoke Co. Apocynum cannabinum var. pubescens (R. Br.) DC. 6092 Corning, Clay Co. Vinca minor L, Periwinkle or Myrtle. Wash- ington Co. Escaped locally from cultivation. Asclepiadaceae : . Asclepias stenophylla Gray. 8397 Cotter, Marion Co. Asclepias Sullivantii Engelm. Fayetteville, Wash- ington Co. Asclepias perennis Walt. Corning, Clay Co. Asclepias purpurascens L. 12648 Mena, Polk Co. Acerates floridana (Lam.) Hitche. 25018 Hazen, Prairie Co. Convolvulaceae: Cuscuta cuspidata Engelm. 16322 Fulton, Hemp- stead Co. Stylisma humistrata (Walt.) Chapman. 8111 Gif- ford, Hot Springs Co. : Supplement to Catalogue of Arkansas Plants 137 Polemoniaceae: Phlox bifida Beck. War Eagle Creek near Hunts- ville, Madison Co. Hydrophyllaceae: Nemophila phacelioides Nutt. Wheeler and Johnson, Washington Co.; Hot Springs, Garland Co. Boraginaceae: Onosmodium occidentale Mackenzie. 8039 Fulton, Hempstead Co.; Baxter Mountain, Washington Co. Onosmodium subsetosum Mack. & Bush. 6041a Northfork, Baxter Co. Verbenaceae: Verbena Drummondii (Lindl.) Baxt. Savoy, Wash- ington Co. Verbena Lambertit Sims. Son’s Chapel, Wash- ington Co. Labiatae: Satureja glabella (Michx.) Briquet. (Clinopodium glabellum Kuntze). West Fork, Washington Co., on calcareous outcrops. Lamium purpureum L. Fayetteville, Washington Co. Introduced. Monarda citriodora Cerv. Sulphur Springs, Ben- ton Co. Perilla frutescens (L.) Britton. 4437 Eureka Springs, Carroll Co.; Monte Ne, Benton Co.; Crow- ley’s Ridge, near Forrest City, St. Francis Co. (Roland M. Harper). 138 Trans. Acad. Sci. of St. Lowis Salvia azurea Lam. 451 McNab, Hempstead Co. Salvia lanceifolia Poir. 4485 Eureka Springs, Car- roll Co. Solanaceae: Datura Metel L. Egger, Polk Co. (R. H. Rosen). Introduced. Physalis missouriensis Mack. & Bush. 6955 Fulton, Hempstead Co. Bouchetia anomala (Miers) Britt. & Rusby. Sum- mers, Washington Co., prairie glades. Solanum elaeagnifolium Cav. 338 Texarkana, Miller Co.; Fayetteville, Washington Co. Solanum Torreyi Gray. 23009 Pinnacle, Pulaski Co. Serophulariaceae: Agalinis fasciculata (Ell.) Raf. 8464 Gifford, Hot Springs Co.; 8472 Gum Springs, Clark Co. Buchnera elongata Sw. Prairies of Arkansas, coastal plain (Roland M. Harper). Aureolaria grandiflora var. serrata (Torr.) Pen- nell. (Dasystoma serrata Small) 10603 Washing- ton, 8977 McNab, Hempstead Co. LIinaria Cymbalaria (L.) Mill. Kenilworth Ivy. Fayetteville, Washington Co. Occasionally escaped. Paulownia tomentosa (Thunb.) Steud. 10527 Ft. Lynn, Miller Co.; Monticello, Drew Co.; Hot Springs, Garland Co. Escaped from cultivation. Pentstemon arkansanus Pennell. 8143 Little Rock, Pulaski Co.; 24556 Hot Springs, Garland Co.; 25167 Shirley, Van Buren Co. Supplement to Catalogue of Arkansas Plants 139 Scrophularia neglecta Rydb. 12687 McNab, Hemp- stead Co. Veronica arvensis L. Fayetteville, Washington Co. and throughout the state. Veronica serpyllifolia L. Withrow Springs, Madi- son Co. Veronica Tournefortii C. C. Gmel. Fayetteville, Washington Co., University of Arkansas campus. Orobanchaceae: Orobanche ludoviciana Nutt. (Myzorrhiza ludo- viciana Rydb.) Kingdon Springs, Marion Co. (Roy Morrow). Rubiaceae: Cephalanthus occidentalis var. pubescens Raf. Fulton, Hempstead Co.; Hot Springs, Garland Co. Galium. tinctorium L. 6083 Corning, Clay Co. Houstonia lanceolata (Poir.) Britton. Lee Co. (C. Woolsey). | Caprifoliaceae: Lonicera japonica Thunb. Japanese Honeysuckle. Escaped in all parts of state and often becoming troublesome. Naturalized from Asia. Sambucus canadensis var. submollis Rehder. 22295 MeNab, Hempstead Co. Viburnum affine Bush. Ozark region, on rocky cliffs. Dipsacaceae: Dipsacus sylvestris Huds. Teasel. Washington and Benton Counties. Introduced in waste places. 140 Trans. Acad. Sci. of St. Louis Cucurbitaceae: Cucurbita foeditissima H. B. K. (Pepo foetidts- sima Britton) Washington and Benton Counties. Echinocystis lobata Michx. Micrampelis lobata Greene) Washington Co. Lobeliaceae : Lobelia spicata var. hirtella (Ell.) Gray. 4407 Kureka Springs, Carroll Co. Campanulaceae: Sphenoclea zeylanica Gaertn. Lonoke, Lonoke Co.; Stuttgart, Arkansas Co. An introduced annual weed of rice fields. Compositae: Artemisia annua L. 4732 Cotter, Baxter Co. Aster lateriflorus (L.) Britton. 26670 Helena, Phillips Co. Aster ericoides L. 4510 Eureka Springs, Carroll Co. Aster ericoides var. pilosus (Willd.) Porter. 6812 Fulton, Hempstead Co.; 6883 Eureka Springs, Carroll Co. Aster salicifolius Lam. 8981 Fulton, Hempstead Co. Aster subsessilis Burgess. Benton County (type locality, E. N. Plank). Aster Tradescantu L. 4491 Eureka Springs, Carroll Co.; Moark, Clay Co. Bidens discoidea (T. & G.) Britton. 26733 Me- Nab, Hempstead Co. Supplement to Catalogue of Arkansas Plants 141 Brauneria pallida (Nutt.) Britton. 25022 Hazen, Prairie Co.; Savoy, Washington Co. Common in Ozark region. Cacalia similis (Small) new comb. (Mesadenia similis Small.) Benton Co. (type locality, E. N. Plank); Savoy, Washington Co. Chrysopsis mariana (L.) Ell. Southeast Arkan- sas (County Agric. Agent). Chrysopsis microcephala Small. 6847 Columbus, Washington Co. Cirsium horridulum Michx. 24977 Fulton, Hemp- stead Co. Coreopsis lanceolata var. villosa Michx. (C. crassifolia Ait.) 26889 Lawrence, Garland Co.; 27056 Berryville, Carroll Co. Coreopsis pubescens Ell. Savoy, Greenland, Wash- ington Co.; 4490 Beaver, 6350 Eureka Springs, Carroll Co.; Harrison, Boone Co. Eupatorium incarnatum Walt. 4766 Cotter, Marion Co. ; 14626 Texarkana, Miller Co. Erigeron quercifolius Lam. Screeton, Prairie Co. (Roland M. Harper). Gaillardia lutea Greene. 10596 Washington, Hemp- stead Co. Helenium campestre Small. Little Rock, Pulaski Co. (type locality, coll. by Hasse, 1885); Conway, Faulkner Co. (H. E. Wheeler). Helianthus divaricatus L. 12631 Mena, Polk Co. 26444 Magazine Mountain, Logan Co.; 29445 MeNab, Hempstead Co. 142 Trans, Acad. Sci. of St. Louis Helianthus heterophyllus Nutt. 6979 Heber Springs, Cleburne Co. Hymenopappus carolinensis (Lam.) Porter. 27052 Berryville, Carroll Co. Lactuca canadensis L. Fayetteville, Washington Co. Lactuca scariosa var. integrata Gren. & Godr. Common over entire state. Liatris squarrosa var. intermedia (DC.) T. & G. 23061 Hot Springs, Garland Co. Matricaria suaveolens (Pursh.) Buchenau. Fay- etteville, Wheeler, Washington Co. Introduced. Mikania scandens (L.) Willd. St. Francis River Swamps (S. M. Coulter). Parthenium hispidum Raf. (Parthenium repens Eggert). 5591 Beaver, 27059 Berryville, Carroll Co. Pluchea petiolata Cass. 314 Texarkana, Miller Co.; 4756 Cotter, Marion Co. ; Verbesina virginica Iu. 274 Rich Mountain (at summit), Polk Co. Lepachys columnaris (Sims) T. & G. Prairie Grove, Washington Co.; 8024 Fulton, Hempstead Co.; 8216 Benton, Saline Co. Rudbeckia Brittonii Small. Fayetteville, Wash- ington Co. Senecio obovatus var. rotundus Britton. 7201 McNab, Hempstead Co.; 26866 Hot Springs, Gar- land Co.; 24444 Little Rock, Pulaski Co. Supplement to Catalogue of Arkansas Plants 143 Senecio plattensis Nutt. Prairie, near Hazen, Prairie Co. (fide J. M. Greenman). A noxious weed. Silphium Gatesii Small. 8048 McNab, Hemp- stead Co. Solidago amplevicaulis T. & GQ. 434 MeNab, Hempstead Co. Solidago caesia var. axillaris ae! Gray 6930 Jasper, Newton Co. Solidago celtidifolia Small. 8461 Gifford, Hot Springs Co. Solidago arguta Ait. 26430 Magazine Mountain, Logan Co. Solidago hispida Muhl. 6912 Harrison, Boone Co.; 26464 Lonsdale, Garland Co. Solidago juncea Ait. 547 McNab, Hempstead Co.; 4401, 4368 Eureka Springs, Carroll Co. Solidago latifolia L. (Solidago flexicaulis L.) 4554 Hureka Springs, Carroll Co.; 4779 Cotter, Marion Co. Solidago Lindheimeriana Scheele. 20482 Eureka Springs, Carroll Co. Solidago pendula Small. Benton Co. (Type locality, collected by E. N. Plank.) Spilanthes americana var. repens (Walt.) A. H. Moore. (Spilanthes repens. Michx.) 6085 Corning, Clay Co. Sonchus asper (L.) Hill. Lonoke and Arkansas Counties in rice fields; Conway, Faulkner Co.; Fay- etteville, Washington Co. Trans. Acad, Sci. of St. Louis Sonchus oleraceus L. Fayetteville, Washington County and elsewhere (a common weed). Tragopogon porrifolius L. Fayetteville, Wash- ington Co. (as a weed). Taraxacum erythrospermum Andrz. Conway, Faulk- ner Co. Vernonia missurica Raf. 300 Texarkana, Miller Co. Xanthium glabratum (DC.) Britton. 316 Tex- arkana, Miller Co. Trans. Acap. Sci. oF St. Louis, Vor. XXV Puate VI Supplement to Catalogue of Arkansas Plants 145 EXPLANATION OF PLATE Puiate VI Upper: Scouring Rushes (Equisetum suppers along railway em- ankment near Palarm, Pulaski Co. . E. Wheeler.) Lower: Arrow Leaf a latifolia) border of pond near Con- way, Faulkner 146 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE Pate VII Upper left: Short Leaf Pine (Pinus echinata) in characteristic stand, about 25 years old. Butterfield, Ark. Upper right: Same growing on quartzite hills along Ouachita River, Garland Co. Lower eco Fringe Tree or Old Man’s Beard (Chionanthus virginica) of Magazine Mountain, Logan Co. Lower taki, Southern Buckthorn ere lycoides), Crowley's Ridge, near Helena, Phillips Co. Trans, Acap. Sci. or St. Louis, Vou. XXV PLate VII inital. Trans. Acapb. Sci. oF Str. Louis, Vout. XXV Puate VIII Supplement to Catalogue of Arkansas Plants 147 EXPLANATION OF PLATE Prate VIII Left: Texan Saxifrage hi Saxifraga texana) probably the earliest spring flow % wer, appearing late in February, Conway, Faulkner Co, x Middle: A very rare Wake Robin (Trillium pusillum) appearing in il in oak woods on flinty soil. Tontitown, Washington Co. Right: False Aloe (Agave virginica) flowering in June, near Fayette- ville, Washington Co. 148 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE Puiate 1X Upper left: Indian Plantain (Cacalia tuberosa) near Imboden. Upper right: ad Poison (Amianthium muscaetoxicum) near Imboden, Lawre o. (H. E. Wheeler.) Lower opie ae hay of the Ozark Chinquapin (Castanea ozarkensis) ar F ington, Washington Co. Lower ree Bark of Ozark Chinquapin, same tree as above. TRANS. AcabD. Sci. oF Sr. Louis, VoL. XXV PLaTe IX Trans, AcaD. Sci. oF St. Louis, Vor. XXV Piate X Supplement to Catalogue of Arkansas Plants 149 EXPLANATION OF PLATE PuatTe X Upper left: Blue-fruited Hawthorn (Crataegus brachyacantha) near Texarkana, Miller Co. Upper right: Bark of same. Below: Field of Spider Lilies (Hymenocallis occidentalis and Iris) ina marsh near Palarm, Pulaski Co. (H. E. Wheeler.) 150 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PiatTe XI Upper left: Honey Locust (Gleditsia triacanthos) near Helena, Phil- lips Co. Upper right: Bark of same. Lower grin White Ash (Fraxinus americana) growing from opening trunk of Satiow Post Oak (Quercus encter Fulton, Hemp- Co, Lower right: Azalea (Rhododendron canescens) in April near Hot Springs. Pirate XI ry eee, 4 uls, Vi I s. Acapb. Sci. oF Sr. TRAN = ‘yy Puate XII XXV OL. rT Sci. oF St. Louis, \ TRANS. ACAD. Supplement to Catalogue of Arkansas Plants 151 EXPLANATION OF PLATE Priate XII Upper left: Blazing Star (Liatris elegans) and Button Snakeroot Eryngium yuccaefolium) in a meadow near Palarm, Pulaski Upper right: Chicasaw Plum igeiaas angustifolia) growing on loess s, Helena, Phillips Lower ets ogi Leaf Pine Feet echinata) West Mountain, Hot Lower right: French Mulberry ( Callicarpa americana) a shrub with br -epubers lavender colored berries in autumn. Helena, Phillips 152 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PuatTe XIII Cone Flower (Brauneria pallida) in a meadow in May, shington Co. Upper right: Red Buckeye Shin discolor var. mollis) flowering in May, Central Arkans Lower left: Prairie a 8 ee purpureum) in June, North- west Arkansa Upper left: Wa Lower right: Rough ee [Mecente scabra) Arkansas border and Southwestern Missouri in ‘LG dO ‘IDG “GVOY “SNVUL — iq ~ ~ ~ in a ~ <=) ~ = ie AXX LIX SLV Id css ad ~ es 18. 19, Supplement to Catalogue of Arkansas Plants 153 BIBLIOGRAPHY ON FLORA OF ARKANSAS. (Chronological) Bradbury, John. Travels in the interfor of North America in the years ae _ 1811. Liverpool, 1817. (Mentions Several plan om “Upper Louisiana” and Great Prairie, probably ria Arkansas, seen on trip down Mis- sissippi Rive - Schoolcraft, Henry R. Journal of a tour into the interior of telat and Arkansas. (1818-1819). London (Phillips). Nuttall, Thos. Journal of travels into Arkansas territory dur- ing mo year 1819, with tigger observations on the manners of the aborigines . 236. Phila. 1821. rnin cae of some new species of plants recently introduced into the gardens of Philadelphia from the Arkansas territory. Journal Philadelphia Academy o Science. 2:114-23; Phila. 1821. (Includes description of twelve species of Arkansas ane Ss.) ~————— Description of two new genera of the natural order Cruciferae. Journal Philadelphia Academy of Science, 5:132-35. 1825. (Described: Selenia aurea Nutt. and Strepthanthus maculatus Nutt.) Plates 6-7. Featherstonehaugh, G. W. Geological report of an examination made in 1834 of the elevated country sorcenn the MiIs- rie and Red Rivers. Exx-Doc. No. 151, 23d Cong, 2nd Vol. IV. 97 pages. Washington (Gates and 35. - Nuttall, Thos. Raiegeconsy towards a of the territory of Arkansas, Trans. Am. Phil. eg oan Series) 2:139-202. Phila. "1887 - Lesquereux, Leo. Botanical and paleontological report on the geological state survey of Ar as. Rep. Second Geol. in of Ark. pp. 295-399. Phila. 1860. Butler, Geo. D. Additions to the flora of Arkansas. Bot. Gaz. 2:104. 1877. Harvey, F. L. Asplenium Bradleyi. Bot. Gaz. 5:15, 1880. Some Arkansas ferns. Bot. Gaz. 5:39. 1880. Early plants. Bot. Gaz. 5:56. 1880. (Coulter, J. M.) Southwestern plants. Bot. Gaz. 5:84. Harvey, F. L. Notes from Arkansas. Bot. Gaz. 5:91-3, 1880. Distribution of Nymphaceae in Arkansas. Bot. Gaz. : 80. 1880. Warder, John A. Notes from Arkansas, Bot, Gaz. 6:188. 1880. Harvey, F. L. Ferns of Arkansas. Bot. Gaz. 6:189-90. 1880. Ferns of Arkansas. Bot. Gaz. 6:213-15. 1880. —————— Some Arkansas Trees. Bot. Gaz. 6:215. 1880. oo or ow ~1 i) oo . Williams, J. te Trans. Acad. Sci. of St. Louis Leavenworthia in S. W. Missouri and N. W. Arkansas. . Gaz. 6:230. 1880. The flora of Arkansas. Am. Nat, 15:388-89. 1881. Forest Notes. Bot. Gaz. 6:273. 1881. A second Spring in Arkansas. Bot. Gaz. 7:12. 1882. Selenia aurea. Bot. Gaz. 7:57. 1882. Forestry Notes. Bot. Gaz, 8:355-56. 1883. Carya myristicaeformis, Bot. Gaz. 9:195. 1884. Dioclea Boykinii. Bot. Gaz. 9:196. 1884. Notes on forest trees. Bot. Gaz. 10:279-80. 1885. Some abnormal Rudbeckias. Bot, Gaz. 10:296. 1885. . Sargent, . S. Report on the forests of North America Sees rth th Mexico). 10th U. S. Census (Arkansas) 9:543-4 Washingt. 1884. Map of Arkansas rene the distribution of the forests. 10th U. S. Censu . Harvey, & L. Arboreal flora of Arkansas. — Jour, Forestry. 1:413-24 and 451-58. June-July. . Call, R. Ellsworth. Notes on the native forest trees of eastern Arka nsas. Proc. Ia, Acad. Sci. 76-85. 1887-89. (Branner, J. C.) Flora of Arkansas with additions by W. a Blatchley, 1889, and index; also notes of others added a various ti Manuscript in University of ” arkansas library. (No date. . Branner, J. C. and F. V. Coville. A list of the plants of Arkansas. 18 Annual Report for 88. Arkansas Geol. Survey, 4:155-242. Little Rock. 1891. . Coville, F. V. Notes on the Botany of Arkansas. Annua al port for 1888 Arkansas Geol. Survey, 4:243-252. ens Rock. Call, R. sg irdhe ac Notes on the forest trees of the Cromer? Ridge region. Annual Report for 1889 Arkansas Geo Survey, 2:183-202. Little Rock. Distribution of timber and its relation to underlying =P formations. (Magnet Cove — 2 kaunas Report for 1889, Part 2. (Igneous Rocks kansas Geo Garris for 1890. 2:167-170. Little ee Harvey, F, Bat Varieties of Ranunculus abortivus L., Bull. Torrey . Club. 19:93-94. March, 1892. Bush, B. F. Bot of some southern swamps. Garden and Forest 10: 51416, Dec. 1897. Plank, E. N. Concerning the plants of southwestern Arkansas. Plant World. 2:45-47. 1898-99. : Olmstead, F. E. A I plan for the forest lands near Pine Bluff, U. S. Bureau of Forestry Bull. 32. Washiantan. 1902. we wo - Ashe, W. W. Further notes on woody plants. 1925. Supplement to Catalogue of Arkansas Plants 155 - Coulter, S. M. Ecological comparison of some typical swamp rea. Fifteenth Annual Report Missouri Botanical Gar- den 39-71. 1904. - Record, Samuel J. The forests of Arkansas. For. Quart. 5:296- 301. S. 1907. The forest resources of Arkansas. Mo, Pac, Land Office, Little Rock. 1910. - Howell, A. H. Birds of Arkansas. Bull. No. 38, Biological Sur- ., Washington. 1911. (Contains several botanical notes.) - Harper, Roland M. Botanical evidence of the age of certain ox- bow 6 1912 lakes, Science, N. S. 36:760-761. Nov. 29, sae notes on the _ coastal saabae of Arkansas. Plant World, 17:36-48. Feb. 1914. Some undescribed prairies in northeastern Arkansas. Plant World 20:58-61. Feb. 1917. Sargent, C. S. Notes on North American trees. II. Carya. Bot. Gaz. 66:229-258. Sept. 1918. - Palmer, E. J. The forest flora of the Ozark region. Journal Arnold 1921. rboretum 2:216-32. Apr. he Red River forest at Fulton, Arkansas. Journal avede Arboretum 4:8-33. Jan. 1923. Ashe, W. W. Further notes on the trees and shrubs of south- eastern United States. Bull. Torr. Bot. Club. 50:359- 363. Notes on woody plants. Jour, Mitchell Soc. 50:43-48. Aug. 1924. - Palmer, E. J. The ligneous flora of Rich Mountain, Arkansas and Oklahoma. Journal Arnold Arboretum 5:108-134. Apr. 1924. Am, Fern Jour. Two interesting ferns from Arkansas. 14:39-41. Apr.-Jun. 1924. Buchholz, John T. Notes on Arkansas Pteridophytes. Jour. 14:33-38. Apr.-Jun. 1924. Am. Fern - Shreve, ean W. Iris cristata. Flower Grower 11:301. Aug. Flower Grower. 11:385- Native irises of the Ozarks. 386. Oct. 1924 Buchholz, Rages T. and Wilbur R. Mattoon. Common forest trees f Arkansas. Bull. Agr. Extension Division, University (Little Rock Office.) ae godine No. 180. Nov. 1924. Torreya, 25:10-11. Jan.-Feb. a 3 2 j bs e Transactions of the Academy of Science of St. Louis He Volume XXV, No. 7 Se THE ECOLOGY OF A SHELTERED CLAY BANK; A STUDY IN INSECT SOCIOLOGY PHIL RAU . THE ECOLOGY OF A SHELTERED CLAY BANK; A STUDY IN INSECT SOCIOLOGY. Pum Rav. A. Introduction. B. Interrelations of animal life. (a) Fauna of the unit. (b) Pioneer life. (c) Renters. (d) Visitors. (e) Parasites. C. Relation of population to environment. (a) Relation to temperature. (b) Relation to light and sunshine. (c) Relation to cold, cloudiness and darkness. (d) Relation to rain. (e) Relation to plant life. (£) Death by violence and natural death. (g) Relation to soil conditions. D. Concluding remarks. 158 Trans, Acad. Sci. of St. Louis (A) INTRODUCTION. “‘Tiving things are real things—but their reality 1s in their interrelations with the rest of nature, and not in themselves.’’—Brooks. The ecologist, when he wishes to study the interrela- tions of a fauna to a restricted area, usually selects a virgin plot, uncontaminated and undisturbed by the hand of man. The area which I have studied is unique in that it has been created by man. These pages will reveal that the invertebrate population readily responded to this man-made habitat and lost no opportunity to utilize il to advantage. Despite the fact that the clay bank here re- ferred to was artificially built, the attraction to it of the life about the area for nesting and other purposes was legitimate, and this made the place a biotic unit of un- usual interest. I have given to the unit a layman’s study of ecology, made without the technical and refined instrumental ex- aminations and prepared without the use of new and complex terms so often introduced into a study of this kind, which, in my opinion take commonplace phenomena and place them beyond the understanding of the layman. By trying to become familiar with complex and unusual terms, one often loses the thread of the story of the in- terrelations of the inhabitants. In other words, this paper is, I hope, analogous to the study of the stars through an opera glass, instead of a study by mathe- matical formulae. The ecological unit under discussion lay in a little valley at the station of Wickes, twenty-two miles south of St. Louis on the Iron Mountain Railroad. The Mera- The Ecology of a Sheltered Clay Bank 159 mec River empties into the Mississippi about one-third of a mile northeast of this point. This region is attrac- tive to fishermen and as a result about a dozen club- houses have been built near the station. Some of these cottages were built upon posts or stilts, thus exposing the earth beneath them. Two such houses were situated on a northward slope, but there was no perceptible life in the earth under them. This was probably due to their shaded condition and the friable nature of the earth there. In the construction of another of these houses, a quantity of clay had been excavated from the cellar and had been shoveled under the porch to save the expense of hauling it away. In the twelve years that this heap of subsoil had lain there, it had become packed to a very hard consistency. The house faced the east where it received the benefits of the morning sun. The porch was four to five feet above the ground, and thus admitted a flood of light in the morning and midday, but excluded all the afternoon sun, and afforded protection from the weather. This bank of hard-packed yellow clay was about eighteen feet long, and varied from thirty to thir- ty-six inches in height. The wood-boring inhabitants of the porch, the life in the clay bank, the insects among the rubbish on top, the occupants of the rambler-rose stems in front of the bank, and the occasional visitors all consti- tuted a legitimate biotic unit, the subject of this study. (See Fig, ie In May, 1917, when the unit was first discovered, it was noted that the northern half of the clay bank was riddled With holes, while the south half was not so, but at the extreme south end a few turrets of a wild bee were found. Figs. 3 and 4 show the south and the north half of the clay bank respectively, and the enormous amount of 160 Trans. Acad. Sci. of St. Louis activity which occurred in the north half in contrast to that in the south half will be seen by the contrast of the great number of holes in the one to the lack of them in the other. Before the summer was far advanced, the cause for the occupancy of one portion of the bank and not of its opposite half was clearly apparent. ‘A glance at Fig. 2 will show how the central and south rose- bushes had been trained to overshadow the southern por- tion of the clay bank, while the third bush had been so bent as to leave the northern half exposed to the eastern sun.* This shows beautifully the relation of sunlight to the activities of Hymenoptera. The loose appellation usually applied to them, ‘‘Children of the Sun,’’ is here seen to have truthful significance. It is generally known that bees and wasps love the sunlight, but a glance at Figs. 3 and 4 will forcibly emphasize this impression. While this clay bank was small and unimposing, its riddled portions indicated that many bees and in all probability their parasites were potentially present, and for an ecological unit for intensive study nothing better could be wanted. Accordingly, frequent visits were made to the spot from June 5 to October 3, 1917, and several trips were made there for comparative data dur- ing the summers from 1918 to 1921. When one looks at Fig. 2 and thinks of the life, love and death tragedies enacted within the limits of an area of a few square feet, one can see very clearly a literal example of the statement by Adams: ‘‘ We may profitably compare an association of animals in a given habitat to a play upon the stage. The environment corresponds to *The photograph for figure 2 was taken in October, when most of the leaves had fallen, and so does not give an adequate idea of the density of the shade. The Ecology of a Sheltered Clay Bank 161 the stage, the dominant members of the association cor- respond to the leading characters; the secondary species, always present, to the essential but subordinate charac- ters. The individual animals adjust themselves to one another, especially to the dominant forms, and to the environment, as the personalities in the play adjust them- selves to the dominant characters, to one another, and to the environment.. In both groups some individuals are dominant, some used and useful, some tolerated, others pick up the crumbs, still others are predatory or parasitic, and all must be mutually adjusted to one another and to the environment.’’”* How this classification of various types fits into what we here record, the reader may presently see for himself. (B) INTERRELATIONS OF ANIMAL LIFE. (a) Fauna of the unit. If all the bank’s a stage, and all the six and eight- legged creatures merely players, it is fitting that we should give now a cast of characters, in the order of their importance, and follow the careers of the dominant ones for the five years, 1917 to 1921. Early in the work it was seen that the many insects did not use the clay bank and the environs in the same way, but they were easily classi- fiable into four distinct groups. Group 1. Pioneers. The permanent dwellers in the clay bank and environs, in a general way the pioneers. Group 2. Renters. The insects less hardy and more ease-desiring than the pioneers; those which rented or appropriated the abandoned dwellings of the pioneers. They might be called squatters. *Guide to the Study of Ecology, p. 47. 1913. = 162 Trans. Acad. Sci. of St. Louis Group 3. Visitors. The insects and animals which dropped into the community accidentally, or in quest of shelter or food. They very often influenced the inhabi- tants, the members of groups 1 and 2, in two ways: by eating them or by becoming food for them. Group 4. Parasites. This contains the names of parasites whose hosts are listed in groups 1 and 2. (b) Pioneer life on the clay bank. The pioneers, those which blazed the trail, those which first came from elsewhere and discovered the clay bank and homesteaded it, and made it easy for others follow- ing to live near them or in their old abodes, or to para- sitize their children—these are first considered, and are listed in order of their importance. With this pioneer life as a foundation for a community, the other chapters that follow will point out their interrelations, sometimes simple, sometimes indifferent and sometimes complex. The carpenter-bee, Xylocopa virginica. Drury.” In the Spring of 1917, the place was visited at in- tervals to fix the date of the appearance of the first life. The carpenter-bee, Xylocopa virginica, was the first to appear. On June 15, perhaps a dozen of the insects were at work enlarging the tunnels in the wooden rafters of the porch above the clay bank. They were all doing precisely the same thing at this time, enlarging the holes and kicking out the sawdust. The flickering streams of golden grains falling in the sunlight from sources un- seen would have attracted the attention even of persons unaccustomed to observations of this kind. Judging from *All of the material has been identified by expert entomologists, ose names appear in brackets along with the specific name of the i; The Ecology of a Sheltered Clay Bank 163 the piles of sawdust here and there on the ground, this work must have been going on for a good many days. Hence the emergence of the bees was simultaneous, but whether from hibernation or from the pupal stage I do not know, since I could not ascertain just how this species spends the winter, unless I used an ax on the rafters, and this I had not the immoral courage to do. However, it was plain from the first that no new tunnels were being made, but that old ones were being enlarged. The activities of the bees in tunneling and pollen-gath- ering increased, although not their number, up to July 16, when they almost completely disappeared. This was probably the end of the first generation of 1917, regard- less of whether they had hibernated or had emerged as new adults in the spring. On July 30, other adults appeared—probably the first of the second generation. On this first day, two females were at work; on the next day four were in evidence, and the numbers continued to increase during the next few days, until by August 20 they were more abundant and seemed more industrious than the population earlier in the season. They were so intensely busy cutting out the tunnels, and they threw down the yellow dust to the ground in such abundance that the pits of the ant-lion larvae beneath suffered complete obliteration. They were seen in the same activity and numbers up to Sep- tember 7 (with certain pauses in their work due to mete- orological conditions described elsewhere). The three nights before September 12 were very cold; the insects about the unit were dwindling in numbers, and those which still lived were slow to come out of their burrows. That day it was 2 o0’clock in the afternoon before the first carpenter bee made its appearance, but her sisters re- 164 Trans. Acad. Sci. of St. Louis mained within, in either temporary or final rest. Again on September 23 only one carpenter-bee was seen. At this date all the other life was proportionately reduced, as was to be expected amid unfavorable weather condi- tions. A visit to the spot ten days later, October 3, re- vealed several dead specimens of these bees and other species about the bank, but no living ones. In all prob- ability they were either dead in their burrows or closely huddled together, ready to hibernate for the winter. In the year 1918, as I shall relate elsewhere, all the more important species in the unit appeared earlier than in the previous year. In 1917, the first carpenter-bees were sawing their tunnels on June 15; in 1918, on May 28 not only were they at work, but they had already com- pleted their burrowing and were busily gathering the yellow pollen from the wild roses growing some distance away. The big shining black bee, heavily laden with bright yellow pollen, made a conspicious bit of color, and was as pretty as the blossom itself. An examination showed that no new tunnels had been made, but the old ones from the last year had been again used. My next visit on June 28 found no carpenter-bees out; this was probably the period between the two genera- tions as was July 30 in 1917, when the first generation was late by a corresponding margin of time in getting a start. On July 17, when it seemed time to expect the second generation, I visited the colony, and found one lone carpenter-bee at work; July 31 likewise found only one bee out. All of the evidence so far in 191% indicated a marked reduction in the population of carpenter-bees. An abun- dance of shedding-skins of the silver-winged parasite, Argyromoeba tigrina, adhered to the wooden rafters The Ecology of a Sheltered Clay Bank 165 about the bees’ tunnels; hence it is probable that this parasite was largely responsible for the small number of bees, as well as of the mud wasps, Monobia quadri- dens, and the grass-carrier wasps, Chlorion (Isodontia) auripes, which occupied the old tunnels of the carpenter- bees. The details given later under the subject of para- sites will show an enormous increase at that time in the number of adults of the A. tigrina parasite, So the record for the second generation continued dis- mal; on August 31, barely two bees were busy, and by the middle of September none at all were to be seen. During the summer of 1919 no observations could be made, so we do not know by what struggles or turns of fortune Xylocopa again came into her own, but in the following spring, on May 27,* 1920, they were once more fairly abundant. Their emergence had probably just occurred when I arrived upon the scene, for they were not tunneling, neither were they carrying pollen, but both sexes were in a gleeful mood of courtship. Several days later others were often seen refresh- ing themselves on the blackberry blossoms, while sev- eral returning females were so heavily laden with green pollen that much of it would spill in a delicate shower as they tried to gain entrance to the home. During the week of June 13 to June 19, eight mothers were at work on their nests. Since only these eight remained to work, out of the large number in the courtship dance, it leads one to suspect that a very large proportion of the latter group were males. During the days of June 28th and 29th, I kept a very careful watch and was startled by the revelation which met my eyes; during the two hours *In 1922 they were flying about as early as April 7. One newly dead bee was taken from a spider’s web. 166 Trans. Acad. Sci. of St. Louis from five to seven p. m., 35 adult Xylocopa returned to the colony and quietly crept into their burrows! This certainly shows how an observation must be made from every possible angle before one may dare to be satisfied with an answer. Fourteen of these were cap- tured and examined; twelve were females and two were males. The presence of the males made me wonder whether they were the remainder of the first generation, or the forerunners of the second. I cannot imagine just what was their business abroad which kept them away from home all day thus, day after day, at a time when they were not gathering provisions of any kind for the nest. May it be that they were out seeking food to gorge and fatten themselves before beginning the arduous tasks of nidification? From this time on the population again waned, until by the middle of September they were seldom seen. Long after the others had disappeared from view, one lone mother was seen going into her burrow on October 13. Just as the mining bees make way for numerous other species by bequeathing to them their old burrows, 8o Xylocopa makes it possible for two species of wasps, Chlorion (Isodontia) auripes and Monobia quadridens to inhabit the locality. For details of the behavior of these sub-tenants and for details of parasitism on the carpenter-bee, and for the relation of the bee to weather conditions, see later pages. The mining bee, Anthophora abrupta Say [S. A. Rohwer]. The most important of the pioneers getting a foothold in the clay bank were two species of mining bees, one supplementing the other beautifully in the point of time. The Ecology of a Sheltered Clay Bank 167 The first to appear each year was Anthophora abrupta, and when its life cycle had run its course before the middle of the summer, the white-banded bee, Entechnia taurea, made its appearance. Other species of bees ap- peared from time to time, but none reached the impor- tance that these two species attained.* On June 25, 1917, it was found that several specimens of the mining-bee, Anthophora abrupta (Fig. 5), had emerged from their winter sleep in the depths of the clay bank, and were busily coming in and going out of their burrows. Some of these activities were directed to burrows which had mud chimneys at the openings, and some to others which had none. Some of these bees came in heavily laden with pollen; others had their gullets full of water; others were kicking out loose moist earth from their burrows and letting it fall to the ground below. Some were working within the burrows, and at intervals would come backing out with a ball of soft mud under the chin, toss it back to the hind pair of legs, and with only these appendages would fashion the wonderful little chimney which crowned the entrance; others were fre- quenting the puddle of water in a wagon rut near by, getting the water with which to moisten the hard yellow clay. Quite probably all these bees had emerged within a few days prior to June 25, and they were already at work in various stages of nest-building or provisioning. June 28 was my next visit, and this was after a heavy Shower. There was little activity among the bees; most of them were waiting indoors until the sunshine should warm up the bank, but several had learned to make a *In 1919, a third and new species of mining bee, Anthophora a oi ractically the same life cycle, and ap- peers at the’ odes S cant a few days later than a. 168 Trans. Acad. Sci. of St. Louis lucky turn out of this misfortune; instead of going some distance to the roadway puddle for their water, a dozen bees with their long tongues protruding were lapping up the drops of water from the vegetation nearby. Frequent visits showed the activity of the A. abrupta bees to be the same without change in numbers up to July 12. Thus the entire population lived practically simultaneously, without individual variation in time. During the four days from July 12 to 16, the number of workers dwindled alarmingly from day to day. This sud- den dropping off led me to believe that I was witnessing the death and the end of the first generation. I hoped that the second generation would come soon enough and be large enough to give me material for certain homing studies without the danger of exterminating them. The next few days saw the demise of the last few survivors; but my anticipation of a second generation was in error, for no others appeared that year. Furthermore, in the years following I found only one generation of these bees each year, and that with a remarkably short life eycle. In this 1917 brood, the duration of adult life was less than thirty days, the first bees emerging a few days before June 25, and the last ones dying about July 16. While daily visits during the following week proved that A. abrupta had entirely disappeared, these daily visits also made it conspicuously clear that another species of turret-building mining bee, Entechnia taurea, had emerged from the bank and was busily engaged in nest building activities. And in precisely the same way in the following years it was observed that just as soon as A. abrupta had finished her labors and gone the way of all flesh, then the white-banded bees appeared upon the scene. The interrelations of the two species of bees The Ecology of a Sheltered Clay Bank 169 in point of numbers from year to year show very fine adjustment. As one species increased progressively from year to year, the other decreased. The details of this phenomenon are given on later pages. While 4. abrupta appeared about June 25 in 1917, a visit on May 28, 1918, showed this species already out and active. Early emergence this year affected most Species inhabiting the bank. Later I shall discuss the relation of this emergence to meteorological conditions. But even at this early date, the A. abrupta bees must have emerged five or six days previously because the work on their chimneys was in many cases far advanced or completed. Having been born a month earlier in the year than their parents of the preceding year had been, the population had the same number of days to their adult life and lived, builded, and many had died before June 28, Then, within three days after their disappear- ance came the white-banded bees again, following with all the precision of clock-work! The interesting item was that Anthophora abrupta had increased in numbers from 22 nest-building mothers in 1917 to 92 in 1918.* Not all of the 92 mothers had nests with turrets; only 52 had full-sized chimneys, 15 with half-sized ones and 25 nested in burrows devoid of chimneys. This increase was, as we shall later show, very decidedly at the ex- pense of the white-banded bees whose numbers this year rapidly decreased. What should we expect in the fol- lowing year, and what was the cause of the change in dominance? *It , that any accretions in numbers from ene tu this cis We A. abrupta were progeny from the t Since the latter species diminished in the bank from year to year, is evident that no new stock took up their abode there. 170 Trans. Acad. Sci. of St. Louis In 1919, circumstances were such that I could pay my first visit to the clay bank only on September 6. There- fore, I could not get their date of emergence, bunt they had left for me a clearly-written record of their abun- dance on the face of the bank. I counted 232 turrets,” which indicated that at least that number of mothers had been at work. Since a small portion of the mothers used the burrows without turrets, and since some may have been broken down, this is a conservative number. I am sure that none were from the previous year, for I had harvested the entire lot in the autumn for my cabinet and study. A recapitulation of the figures is of interest: in 1917, 22; 1918, 92; 1919, 232. In 1920, the first visit to the clay bank was made on May 28. Until 9:45 a. m. no bees appeared; then the sun warmed the bank and I saw two A. abrupta flying before the bank and entering numerous holes, one after the other. During the next fifteen minutes I sat before the bank and saw three others emerge from the burrows and dart away on the wing. The white faces of all five proved that they were males. A shrill voice within a burrow caused be to rivet my attention to a certain spot, where soon a sixth white face made its appearance. So here I was at hand for the first time to see the males emerge! There were no females about. There occurs in this species that which we see in so many species of insects—the priority of the emergence of the males. About noon the first female was seen going into burrow after burrow, and a little later a second one was doing precisely the same, while the males began to go away to *This had been an especially wet year and the bees could get water for digging. In dry years, as in 1922, hardly any of them made turrets. The Ecology of a Sheltered Clay Bank 171 the blackberry blossoms, or to rest listlessly upon the nearby vegetation. The females moved about with an air of hesitation, of uncertainty. All this showed that the emergence of this species had probably begun on that day, so, judging by the last year’s population, we had reason to expect that the next two days would show an exodus of great magnitude. Unfortunately, I could not be present during the next few days. On June 13, a very different type of activity was ap- parent. Hundreds of turrets were already completed and hundreds of busy mothers were flying in and out of their nests. In two weeks wonders had occurred. The males had come and gone—not one was to be seen. Judging by the progress of the work each activity must have been practically simultaneous for the whole group. Thus, all nests were in the same stage of construction, indicating that the work had begun simultaneously; hence probably emergence and mating had occurred for all within a day or two. The amount of work which they had already done was enormous, such as could only have been accom- plished by their toiling as they do from sun to sun. At 5:30 a. m., one day I saw some of them beginning their work, and at 7:30 p. m., when it was almost too dark in the shadows to find their burrows, they came home and crept in. Later in the season they were not so in- dustrious early and late on cool or cloudy days. : It is their habit to carry water in the gullet, with which to reduce the hard yellow clay to workable mud. When one takes a bee in a test-tube, it disgorges the water on the glass in its futile fury. On one occasion, several bees in a test-tube disgorged so much water that they became 172 Trans. Acad. Sci. of St. Louis quite drenched and made their elegant pubescence all sodden. The notable feature was that this species had again increased enormously. Their distribution over the face of the clay bank was interesting: at the extreme south end were 10 turrets, in the central part, 275, and at the extreme north end, 390, making a total of 675 in three rather distinct groups. Besides this, more than a hun- dred bees had nests with no turrets and during the week I saw about 75 more chimneys constructed. This may indicate that each mother may make more than one nest in a season.* Many of the bees were bringing in pollen, others were building, and yet many more than three- fourths were at this time coming in apparently empty- handed. I think that they moisten the pollen with nec- tar, and this would mean many trips which to an ob- server would appear empty. Then again each cell is varnished inside to a certain depth which gives it a smooth glossy finish. This is no doubt for the purpose of making the brood cup impervious to water; at least when one drops water on the inside from a pipette, the drops retain their shape, and if the entire cup is filled, it retains the water for a long time. On the con- trary, if a drop be placed on the outside of the cell, it immediately spreads and makes mud out of the mass. This varnish is probably made from plant resin that is carried at the time when they seem to be coming home empty handed. This bee, and the two other species of mining bee, do more than merely make a hole in the . ground and fill one end with food; they actually make mud honey-cells which ean be taken from the burrow in- *There may be dull seasons, e. g. cold or dry, like sacs when it would “eg impossible for a female to make more than on The Ecology of a Sheltered Clay Bank 173 tact (Fig. 11), and all three species varnish the interior with the waterproof substance already referred to. As noted above, they customarily work from early till late. On June 17, 1918, when the temperature* dropped from 98 degrees to 79 degrees Fahrenheit, their activities were greatly lessened. Less than a dozen as- sumed activity at all. I tried to find where they got their pollen. On a per- simmon tree some two hundred yards away, with its blossoms already deteriorating I found many bees, in- cluding this species, getting nectar. The two popula- tions, bees and blossoms, waned at the same time. In 1921, notes on the relation of this species to the persim- mon blossoms were made, and details are given else- where. What other sources of food supply they may have had I did not learn. From May 28 to June 13 of the year 1920, they had built turrets to the enormous number of 675, a substan- tial increase over the number for the previous year. On July 21, only a week later, these short lived creatures were almost gone; only two live females could be found, the remnant of the recently noisy and mighty throng, and dead specimens of A. abrupta were scattered all about. In 1921, Man was again a factor in regulating the abundance of Anthophora life; the tenants of the “‘Ham and Bud’? club-house unwittingly caused the death of hundreds of these little creatures, by merely going away and leaving the door of the screened porch open. The bees, in their circuitous flights, entered this open door at the head of the stairway (Fig. 8, point x), and Were not able to find it again to make their way out; thus the screening which was intended to keep insects out, ee *U. S. Meteorological Summary for June, 1918. 174 Trans. Acad. Sci, of St. Louis kept these in. Within the porch was a pitiful sight; in the two sunniest corners of the porch was literally a drift of dead bees. In their desperate attempts to es- eape, they had directed all their efforts against these two points of maximum sunlight, and even while I watched, at the southeast corner in the bright sun were one hun- dred and fifty mothers trying frantically to escape. It was with a feeling of satisfaction for the privilege of doing reparation for the wrongs done by my kind that I picked these up one by one, and liberated them. It seems strange, and yet it is logical, that these crea- tures could not find their way out. In the first place they had entered by chance, instead of direct quest, which would entail flight of orientation and the formation of memory images of the open door. Second, these are sun-loving creatures, following the greatest intensity of light, so it is easy to see how this factor would guide them in their efforts to escape. This failing is common not alone to Anthophora abrupta, but to other insects as well; several specimens each of Arotes amoenus, Cress. (R. A. Cushman), Megarhyssa lunatrix, Fab. (R. A. Cushman), and Tabanus lasiophthalmus, Macq., were also found there. A humming-bird had been trapped in just the same way, and its little carcass too lay in the sunniest corner, with its delicate bill thrust through the wiring, showing its frantic efforts to escape. An actual count of these dead bees gave me 18 males and 703 females. This, plus about 300 females then at work, shows the enormous proportions to which this population had grown. Furthermore there should be added to this total some 60 specimens from this group that had been lost in certain of my experiments. One wonders what would have happened to the bank in the The Ecology of a Sheltered Clay Bank 175 following year if all these mothers, over 1000, had been left alive to propagate the proportionate population. It seems hardly possible that the clay bank could have held them (and we know that crowded housing condi- tions lead to ill health and ill morals). Would they, under such pressure, have disseminated over other areas? At the end of the season the results of this catastrophe could be seen, for on June 24, 1921, we could count only 175 nests, as compared with the 675 counted for June 13, 1920. The white-banded bee was now present, but in shame- fully reduced numbers. One can hardly say that the A. abrupta in ever increasing numbers had simply crowded out the white-banded bees, because there was so much unusued space about the clay bank, and the two species occurred at different times, so they could not have been in competition for the same flowers. It was almost wholly a question of parasites, and this is treated elsewhere. Other details on the life history of this bee, and the re- lation of the species to moisture, light, etc., are given in later pages.* White-banded Mining-bee, Entechina taurea Say [J. C. Crawford].** The preceding pages show how the mining-bees, An- thophora abrupta, rose in five years from a colony of few individuals to a great and important population. We have also seen how J. abrupta emerges early 1n sum- mer, does its life-work in about thirty days and is done he: logical aspects of the life Wess or ths ee othe Oe camerien species of bees and i : however, will be treated biologically in other papers to follow. It is, often difficult to draw the line between the two aspects of the study. **Mr. Rohwer writes that this bee is now known as (Entechnia) Melitoma taurea Say. 176 Trans. Acad. Sci. of St. Louis for that year. These pages will show that each year immediately after this, the second mining-bee, Entechnia taurea, comes upon the scene and continues the mining activities. The bees were observed for five years, and their numbers ran practically in inverse ratio to those of A. abrupta; as the one increased, the other decreased. These white-banded bees (Fig. 7) emerged from the clay bank in 1917 on about July 16, and took up their work on the spot. This species likewise made turrets over its burrows, but they were a little smaller in size and of finer texture than those of A. abrupta. Like A. abrupta, these bees did not seal their burrows, either at the end of the chimney or the mouth of the burrow.* The males of this species do not seem to emerge before the females; or at least, if they do, they do not die earlier, for on July 311 saw a lot of frolicsome males nea at the north end of the bank. The activities of the white-banded bees ne throughout the summer. They never swerved from their self-restricted nesting areas, two spots at the extreme ends of the clay bank. By August 14, the burrows of these bees were becoming somewhat abundant, and the dancing males had now spread themselves to the south end, where they buzzed and danced about the busy fe- males coming home heavily laden with pollen. While some of the latter burrowed horizontally into the face of the bank, others dug vertically into the flat top. They seem to have a preference for the top, since about ninety per cent of the nests were there. These bees, like their predecessors, also carried water to moisten the hard clay *One scoring finds the turrets sealed at the orifice, but in all the specimens of ‘bis kind which I examined, the burrows been usurped poses used by the wasp, Tryporylon clavatum. The Ecology of a Sheltered Clay Bank 177 to make it workable; then they carried out mouthful after mouthful of soft mud in the form of pellets. These were used in the construction of the chimney until that was of adequate size, after which the surplus material was kicked aside. By August 20 the males had disbanded and were no longer seen about the premises. They had not gone far, however, but were around on the other side of the house, Testing, one each in the wild morning-glory blossoms which they had found there. They quietly occupied the floral cups for hours at a time, serving no good purpose that I could see, but only usurping the rightful place of the pollen-hunting mother, and often even fighting des- perately to keep her ont—a perfect dog-in-the-manger. At the end of August, despite several days of rain, fe- males were still at work bringing in loads of pollen. By September 3, several cold nights had caused them to lay aside their virtue of early rising. They were abundant and actively engaged at this late date. By the 6th or 7th, however, only about a dozen bees were still alive, or at least in evidence about their business, some bringing in pollen and others actually excavating. On September 12, with the preceding night very cold, no bees were to be seen at work in the morning; at noon, one crept out, and by three o’clock, two more ventured. But even at this late date, eight males were seen as before, huddled in the cup of the morning-glory flowers, where they had Spent the night. These flowers had not closed for the night,* and even at ten a. m., these males were in a tor- aan ight, *It is unusual for morning-glory flowers not to close at n but this actually happened late in the season when the —— were Cold and dewey. These flowers which remained open gave lent shelter to the bees. 178 Trans. Acad. Sci. of St. Louis pid, dejected condition, heavily covered with dew. On September 23 it was quite apparent that these insects were being rapidly reduced in numbers by death. Several dead ones were picked up from where they had fallen beneath the mouth of their burrows, and there was no way of ascertaining how many more dead remained within. It is natural to think that females of such species as this continue to work to the last, but on the occasion of my visit on October 3, there were three mother bees that were no longer exerting themselves on home cares, or in any way trying to work, but appeared to be simply idly waiting to be overtaken by death. In- deed they lingered on beyond their just day, for even their food-plants, the pollen bearing flowers, had almost all ceased to bloom.” While I was unsuccessful in determining the number of nests made by each mother, the total number of nests for the whole colony from year to year is of interest. In 1917, there were 62 nests; in 1918, 55; in 1919 the number went down to 37, and in 1920, 49; in 1921, 51 and 1922, 40. When one sees the enormous population of Anthophora in 1921, many of which were trapped and perished, and compares it with the 1921 Entechnia population, one sees a vast difference in their rate of increase; in the five years residence, one species has increased phenome- nally, while the other has barely held its own. The dates of emergence for the various years will be treated under climate, and other details are elsewhere given in these pages. *At even a later date than this, however, on October 17, 1919, one was seen to enter its ‘ene at Hematite, Mo., several miles south of the clay bank. The Ecology of a Sheltered Clay Bank 179 The Mining-bee, Anthophora (Antheméessa) raui.* (S. A. Rohwer]. In 1917 I noticed among the turrets of Anthophora ab- rupta, three which turned upwards, as indicated by ar- rows in Fig. 9. The next year, there were five of these in the mass, and, since this showed a very distinctive digression, I concluded that this might be an advanta- geous variation, and began forthwith to philosophize upon the laws of variation, heredity, and survival of the fittest, with the deepest solemnity. The reason why the reader is mercifully spared this is that, when mak- ing a later visit to the place, June 13 to 19, 1920, I was Shocked to find about 90 of these upturned turrets, seg- regated into three distinct colonies, and bees, which greatly resembled A. abrupta, but still showed distinct differences, industriously taking care of them! In all characteristics excepting the upward turn, the turrets were precisely like those of A. abrupta. Since these bees had habits of digging, carrying water, and pro- visioning their nests with pollen, all of which were so very similar to the way of A. abrupta, it was not surpris- ing that they proved to be taxonomically very nearly re- lated to the latter species.** Thus the great increase in the population in 1920 came practically as a saltation, for in 1918 they were Practically nil. This sudden increase seems entirely pos- sible, however, even though this species has the same Short adult life eycle that A. abrupta has for I am of the opinion that this bee makes more than one nest in a “Described in Proc. Ent. Soc. Wash. 25: Sa 1923, by Mr. Rohwer, who Rebbe that it is closely allied to sodal brupta, **In 1922, this species a peared three sas later than A. @ ated a few spaulava sues, seen a week after A. abrupta had all off. 180 Trans, Acad. Sci. of St. Louis season, and has three or four cells to each nest. By the third week in July, the population was noticeably wan- ing, and before the end of the month had almost dis- appeared. When these bees make their turrets, they use a process which I have not yet been able fully to analyze. The bee walks out backward, and when the tip of the abdo- men touches the rim of the chimney, she applies her pellet of mud. At first this is quite crumbly, but soon one sees that in some way water passes over it, making it more plastic and adhesive. One can see the water as it spreads over the mud, but cannot see where it comes from. It does not seem to come from out of the ab- domen and if it comes from the mouth one cannot see how it is passed to the tip of the abdomen. One load of mud brought from within makes from one-fourth to one- third of a ring. ‘As mentioned before, the only perceptible differences between the chimneys of these bees and those of A. abrupta, are that these turn upward instead of project- ing from the bank horizontally, and these are usually solid tubes while those of A. abrupta have a gap or fis- sure, of unknown significance, extending the full length of the tube on its upper side.* Sometimes this upward turning chimney goes into pretty curves, but it never fails to turn upward. This distinction is unusually short and clear for the two species; I have never yet seen an A. abrupta go into one of these vertical turrets, nor have I ever seen an A. raui enter the horizontal ones, although I have often seen both species go into holes that had no turrets. *Out of several hundred turrets of A. ebrupta, only twelve lacked this split. The Ecology of a Sheltered Clay Bank 181 My earlier notes show that in 1917 there were only three of these upturning turrets, which indicated that this species was newly arrived then. As early as June 18, 1921, I counted 40 such turrets, with A. rawi coming in and out. They were not scattered over the whole bank, but were distributed in groups; this probably means that the bees build very near to the spot where they are born. Fig. 10 shows a colony of 20 upturned turrets in a small area (for this picture a wad of cotton was - placed in each vertical chimney, to better distinguish them from their neighbors), and Fig. 6 shows a mother twice natural size. I accidently broke off the turret of one nest about an inch long, while it was in course of construction. The bee coming out with the next load of mud did not seem in the least surprised, but without turning about to ex- amine the condition, she placed her load at the base and began on a new turret. On June 18th ten new turrets Were just begun. The following record of one typical bee’s work gives an idea of the manner of making the turrets. This bee entered a shallow old hole and kicked out a small quan- tity of loose dust; after some minutes of this, she left and returned presumably with water. She then began to bring out loads of mud and started to make her turret. The face of the bank here was uneven and her first tine loads went to fill the depression, first above the orifice and then below it, until it was level and ready to re- ceive the chimney, which she continued in the same man- ner. A trip for water usually consumed two minutes. In leaving the nest for water she came out head first, having turned around in the tunnel; in coming out with mud, she always emerged backwards and applied her 182 Trans. Acad. Sci. of St. Louis mortar. Between 11:10 and 2:30, the burrow was dug and the chimney built up to a height of % inch; from that time until five o’clock, nothing more was done. Like A. abrupia, these bees build very definite cups of mud within the burrows, in which to oviposit. Fig. 11 shows a group of such cells—some of them opened to show the pupae within, while Fig. 12 shows the pupae , in two stages of development. The interior wall of this mud cup is varnished with some substance which renders it waterproof, a condition we have already described for A, abrupta. Halictus bee, Halictus (Chloralictus) zephyrus Sm. [J. C. Crawford]. A small burrow which went down vertically on the top of the bank first attracted my attention in 1917. It contained a pair of these Halictus bees. One of them, probably the male, closed the aperture of the burrow with his head, and prevented his consort from entering until it suited his whim to admit her. This was the only nest of this species found that year, and it now seems likely that in taking the pair for identification, I exter- minated this species in this locality, for in the two years following none appeared. The species must have been reinstated, however, be- cause in 1920, three burrows were again discovered, and in 1921, a dozen nests were there. They dug their bur- rows horizontally into the face of the clay bank near the base. Below each burrow was a little mound of pulver- ized earth. It was at length discovered that several adults occupied one gallery, and the nesting cells branched off from either side of this gallery (Fig. 13). The Ecology of a Sheltered Clay Bank 183 Rach of these cells, or pockets, contained an egg attached to a small pellet of yellow pollen. At first I thought that it was always the head of the male which plugged the doorway; but as I later learned that each burrow was really an apartment-house occupied by many adults, I have been in doubt whether the one at the door may not have been one of the feminine oceu- pants. This common passage way is larger underground than is the doorway, which is just wide enough to admit one bee at a time. One nest which I opened had a tunnel in the hard yellow clay nine inches long and three- sixteenths of an inch wide, excepting at the narrow en- trance. Pockets on either side of this tunnel had young in all stages of development, from minute creatures cling- ing to a little ball of pollen, to pupae already deeply pig- mented. Besides these there were a half-dozen adults, possibly mothers or adult sisters. After the nest had been dug up, eight bees returning home congregated before the ruins. The adults as they emerge nidify in the same tunnel and gradually extend it. While these are solitary bees they are very neighborly, and this habit of community dwellings seems to point in the di- rection of socialization. Indeed it almost seems to be a link between the solitary and the social habit of bees. Near the end of the season, October 2, 1920, and again September 28, 1922, at Wickes, Mo., I witnessed a new phase in the life of this species. Thousands of these little bees were executing a sun-dance on the sunny hilltop, where the grass was closely mowed. They were in a great many groups of a few bees to several hun- dred, while a few groups had several thousand. Some groups were close together and some isolated, some dane- ing over the short grass and others over the barren olay 184 Trans. Acad. Sct. of St. Louis spots. Thousands were taken in the net, but most of them escaped through the large mesh. They kept near to the ground and moved rapidly; they were so small that individuals could not be followed. Some burrows in the grass were plugged with the heads of the guards and in some the females were seen to enter. From: the size of those in the dance, I suspected them to be males, and the behavior indicated courtship. Little Carpenter-bee, Ceratina calcarata Rob. [J. C. Crawford]. A good many stems of the red rambler roses had been eut back, and each year dozens of these contained nests of these bees and their parasites, but in so far as I could see there was no relation between these bees and the other inhabitants of the clay bank, They went afar for their pollen, and neither preyed upon the others, nor were preyed upon; in fact, in every way they held them- selves aloof from their neighbors. The caterpillar wasp, Ancistrocerus fulvipes Saussure. Among the pioneers in the clay bank should be con- sidered the caterpillar huntress, Ancistrocerus fulvipes, since she almost never used the old burrows for nest- ing, but dug her own tunnels among those of the min- ing bees. In 1917, this species was seen about the clay bank as early as June 17, when three adults were out. By July 7, this number had increased to a dozen, and by the 16th, neither their number nor their industry had waned, although by this time, their companions Anthophora abrupta, Trypozylon clavatum and the chalcid parasite Monodontomerus had disappeared, either temporarily oF permanently for the year according to species. For sev- The Ecology of a Sheltered Clay Bank 185 eral days these mothers had been digging, carrying mud or bringing in caterpillars. On July 30 I returned after a few days absence and found the numbers of this wasp materially reduced, so I suspected that this was about the time for the passing away of the first generation. Thereafter, a careful watch up to August 10 revealed no trace of this citizen of the colony. On August 14, one wasp, probably the first of the second generation, was found at work. Up to ‘August 20, this one and another were the sole representatives of their tribe, and on Au- gust 30, three were out. Although September 3 and 4 were almost sunless, yet 7 individuals were active; this was the maximum number of the second generation Seen at any one time; from September 6 to 23, only two or three at a time were out at work. On the last visit of the year, on October 3, the day was gray and chill, but it found two of these mothers still at work, one plugging her burrow with mud and the other bring- ing in caterpillars, both plodding faithfully on toward the completion of their work, oblivious to the fact that it Was now only a matter of days or perhaps hours until for them the sun would shine no more. In 1918 this wasp, like the other species described above, emerged and began its work much earlier than in 1917; on May 28, they were already out and busy. In 1919, no records were made of their emergence, but in 1920, when the bank was visited on May 27, they were again found at work. Their appearance is correlated - With temperature conditions in later pages. They ap- peared in slightly greater numbers in 1918 than in 1917. On July 17, about the same number were at work; on July 31, none at all; hence this is evidently approxi- mately the dividing line between the two generations. A 186 Trans, Acad. Sct. of St. Louis visit just a month later showed a partial return of the population. No notes were made for 1919. In 1920, as already stated, May 27, found these wasps already alive. Their activity continued throughout June and the most of July. I was unable to visit the grounds then, at the date when the depletion of the first generation was ex- pected, but on September 2, A. fulvipes were more abundant than ever; 13 individuals were seen at one - time, the greatest number yet seen at work simultane- ously, which indicated that they were on the increase. None were in evidence on October 3. These notes, while not based on large numbers, are probably sufficient to show an increase in numbers, an earlier date of emerging when the weather is warmer as with the other inhabitants of the bank, and a strong indication of two generations a year. These wasps, like the mining bees, went elsewhere than the bank for the food for their young and them- selves; hence the relation that this species had to the community as a whole was in bequeathing their discarded tunnels to the other inhabitants who might want them, and in occasionally falling prey to other insects. One such instance was observed on June 28, 1920, when a bug, Reduvius personatus L. (W. L. McAtee), which was amply protected in coloration by having its sticky body covered with dust, and which was half concealed in a crevice, was found feeding upon this wasp. This eumenid was elsewhere seen to nest in tunnels in logs; for an account of its life history, see ‘‘ Wasp Studies Afield.’’ The Ecology of a Sheltered Clay Bank 187 The caterpillar wasp, Ancistrocerus unifasciatus Sauss. [S. A. Rohwer]. Three mothers of this species were seen carrying green caterpillars into cracks of the old building above the clay bank, on May 28, 1921, where they were evi- dently nidifying in the old mud nests of Sceliphron cae- mentarium. I have previously recorded* the fact that they use the old cells of this mud-dauber. The spider-wasp, Episyron biguttatus Fab. [S. A. Rohwer]. This Pompilid wasp was seen kicking dirt into a small burrow at the base of the clay bank. She would quickly Sweep in the earth with her front legs and then with rapid beats of the tip of the abdomen she would pound it down. The wasp was taken and the burrow opened; one-half inch below the surface we found a perfectly round cell of the size of a pea, which contained a spider with an egg cemented to the dorsal side of the abdomen. The pipe-organ wasp, Trypoxylon politum Say.** This wasp should probably be included among the . inhabitants, since three large pipe-organ nests were Were built on boards or joists of the porch over the bank, but a more logical and interesting reason for including it was that several of these mothers were seen foraging for spiders which made their homes in the old bee burrows. Thus the species which seemed almost an outsider could affect the life of the community even though in a small way, by reducing the number of spider *Ent. News 24: 396, 1913. **Formerly called T. albitarse. 188 Trans. Acad. Sci. of St. Louis occupants which in turn would have reduced the number of bees and parasites. One nest gave forth its adults between June 26 and 29, 1917. The mud wasp, Sceliphron caementarium Drury. This mud-dauber stands in the same relation to the life of the unit as does the pipe-organ wasp, just men- tioned. These yellow-legged wasps, too, plastered their mud huts on the wooden porch overhead, and in one in- stance even attached one direct to the clay bank, in a depression which I had made in digging out a nest; a crooked, lop-sided, distorted piece of work it was, and it could not have been different in the cramped corner, but with yards of flat surface overhead, why should a sensible wasp choose a spot of this kind! Like the Trypoxylon politum, these wasps were occa- sionally seen entering the burrows in quest of spiders. Their discarded nests about this unit were utilized as homes by Trypozxylon clavatum, Ancistrocerus wnifas- ciatus, and Pseudagenia mellipes and the burglar wasp Chalybion caeruleum made the best of the opportunity to break into them in a most desperate way. The Larrid wasp, Tachysphex terminatus Smith. These little wasps appeared about July 16, 1917. They were digging their burrows in the loose dirt on the top of the clay bank and provisioning their nests with short- horned grasshopper nymphs. Toward the end of the month their numbers became much reduced; on July 30, only three females were in evidence. These wasps are supposed to be essentially sand- loving, but here we found them digging in the loose dirt The Ecology of a Sheltered Clay Bank 189 on top of the hard clay bank, and as a consequence the nests were very shallow, so shallow, in fact, that the prey could be exposed to view by merely blowing the dust away, whereas those burrows found in sandy areas had a much more substantial depth. On August 10, I found that the number nesting had increased to six, which would suggest, though not prove, a second generation. Also it became evident at that time that this wasp sometimes falls prey to the other inhabitants; one dead specimen was taken from the web of a spider occupant of the bank. Only four days later I found these Larrids had suddenly increased to about thirty, with a goodly proportion of males in the lot. Excited courtship and mating were the program of the day. The males outnumbered the females, and the usual fight for partners occurred, three or four males some- times struggling together for the possession of a fe- male. Even during this commotion, however, about ten females managed to attend to home duties, between inter- ruptions, bringing in locust nymphs mostly of the spe- cies Chortophaga viridifasciata De G. (A. N. Caudell) (Fig. 14). Toward the end of August the number of mothers began to diminish; on September 3, one was found dead in its doorway;-this probably indicated the approach of the season of natural death. And so it was, for only one or two were seen after that, and on my farewell visit for the year, Oct. 3, none of these sun- loving creatures were found to have survived the cold, gloomy days just preceding. The wasps have never appeared in early spring, and a few cocoons taken from the bank on April 28, 1921, gave forth adults on June 2; this indicates that this is a summer and fall creature. In 1918 these wasps appeared in slightly greater num- 190 Trans. Acad. Sci. of St. Louis bers than in 1917; by 1920, the population had trebled. Just why this habitually sand-digging Larrid should choose to adopt itself to the yellow clay of the bank, and why after this change, it should flourish and increase in numbers, is still an unanswered question. We see only slight relation between this wasp and the other inhabitants of the clay bank, since these mothers must go abroad for the Orthopterous prey; hence prob- ably it had nothing in common with its neighbors other than occasionally falling prey to the spiders there. The devil’s horse, Stagmomantis carolina Linn. The egg-cases of the devil’s horse were often found plastered to the boards above the bank, and each June the little nymphs could be seen walking about the bank. They are carnivorous, and probably fed upon some of the very small insects about the bank, until they in turn fell prey to spiders that were occupying the old burrows. The paper wasp, Polistes pallipes Lepeletier. During the four years there were probably a dozen nests of P. pallipes on the under side of the porch and attached to the rose bushes in front of the bank; but in so far as I could see, the wasps had no relation to the unit as a whole, since they. made their own nests, and did not get prey from the clay-bank; neither did they fall prey to the inhabitants. Sometimes, they built their nest in narrow spaces between boards, which resulted in nests strangely shaped, but the point which remained most puzzling was that every year some should persist in building in this cramped space between two joists, and making these unusual nests, when there were hundreds of square feet of clear ceiling and wall space ready to accommodate them. The Ecology of a Sheltered Clay Bank 191 The stick-bug, Emesa brevipennis Say [W. L. Me- Atee]. Another insect which should be classified as a pio- neer in the unit, even though it makes no nest, is the stick-bug, Emesa brevipennis. It spent its whole life among the rubbish, boards, ete. piled on top of the bank. The nymphs were seen as early as July 16, and the adults as late as October 12. During their im- mature stages they were inconspicuous, hence not fre- quently noticed, but when full grown and more easily noticeable, they were observed in great numbers. Since they are carnivorous in habit, they were probably very directly responsible for some changes in the popula- tion of the unit. While I have never chanced to see them actually prey upon the inhabitants of the bank, I have seen them feeding upon insects which had been visiting the bank for shelter. One was imbibing the juices of an adult moth, Pyralis cuprealis Hubn. (H. G. Dyar); another was seen to hold an immature (sec- ond stage) short-horned locust in the fore-legs, mantis- fashion, and, with its beak inserted, imbibe its juices for a whole day. Whether they actually captured sleepy bees and wasps, I cannot say. ‘Ants. Crematogaster lineolata Say [W. M. Wheeler]. A crack in the porch post was the home of a large colony of these ants. Its part in the game of give and take being played on this scene was to supply food to the ant-lion larvae, whose pits were in the dust at the base of the bank. The spider, Habrocestum pulex, and also an unidentified ant-mimicking spider were observed feeding upon them. These ants often covered up the cracks in their posts with a gummy secretion. On Au- 192 Trans. Acad. Sci. of St. Louis gust 14, 1917, many winged specimens were in the lot. This colony remained a permanent part of the poput- lation during the five years observation. On my spring trip afield, April 2, 1921, I found on the ground about 400 of these ants, dead. They were still soft; hence they could not have been dead long. More- over, they had not been there on February 26, when I examined the bank. I had previously noted these ants hiding in the cracks in these walls and plugging the crevices with a gum-like substance; hence they could not have merely fallen out. The question remains whether these had come out to meet the spring and had been caught by a severe frost, or whether they were the accumulated dead of the winter, which had been dumped out by the survivors. In another instance, when a Polistes pallipes queen was taken from the nest above the bank for use in homing experiments, a colony of these ants discovered the nest soon after it was left unguarded; they bit into the walls of the cells and completely removed the larvae bit by bit. This task consumed two days, during which time the nest was black with ants. Anti-lion larvae, Myrmeleon immaculatus De Geer {A. N. Candell]. Myrmeleon mobilis Hagen [A. N. Cat- dell]. The numerous insects mining in the bank kicked out quantities of finely pulverized clay, which accumulated at the foot of the bank (Fig. 1,X). This made an excel- lent abode for the ant-lion larvae, which dug their pits of various dimensions (Fig. 15) and fell heir to occa- sional bits of provender brought in for the other nests but dropped, as well as sometimes the spider or ant The Ecology of a Sheltered Clay Bank 193 inhabitants themselves or occasional visitors to the set- tlement. That this was a favorable place for them to flourish and multiply was evidenced by the fact that in 1917, 20 pits were counted at the foot of the bank; in 1918, 150 were there, but most of these were gathered to send to a friend for study. Hence in the following years they appeared again in about the same numbers as in 1917, at the base and also on top of the bank. Of course we should bear in mind that the ant-lions, by virtue of their peculiar mode of development, are well prepared to survive and flourish, so that, like the poor, we shall have them always with us. They neither perish nor migrate if food becomes scarce for a time, but they simply lie dormant, and put off growing and transform- ing for a year or so until food again becomes plentiful. Thus, an examination on October 13 revealed many ant-lions in all stages of development; some were re- constructing their pits in my presence, others were larvae in all sizes from small to large, with pits in pro- Portionately varying sizes. Precisely these conditions were found at all other seasons, from the opening of spring to the close of autumn. This shows that they have no set time of the year for their metamorphosis, but change whenever they have reached a certain stage, regardless of the time of the year. Some that were brought into the house and fed abundantly on flies 8rew rapidly, pupated in a few days and emerged as adults from three to four weeks later. One must not forget to consider the life habits in rela- tion to the environment, when contrasting the develop- ment of the ant-lions with that of the bees. The larvae of the bees are supplied with sufficient food to carry them through their development without their exertion, while 194 Trans. Acad. Sci. of St. Louis the ant-lion larvae must get what they can. With the ant-lions, it is either a feast or a famine—when food is plentiful, they develop to maturity quickly; when it is scarce, they do not die, but combat famine with pa- tience. Herein Mother Nature anticipated Victor Hugo, who said: ‘‘A clock does not stop short at the precise moment when the key is lost!’ An interesting detail in their method of feeding was noticed when a Grapta caterpillar was dropped into one of the pits. The ant-lion captured it and promptly sucked out the juices; five minutes later it was discov- ered that the ant-lion had also snipped the body-wall and was dragging the viscera out through the aperture. The only relation these Jarvae had to the other inhab- itants of the unit, was to eat any insects, such as weevils and caterpillars, which, through accident or misfortune, fell into their pits. I noticed also that occasionally their pits served as good starting points for the burrows of the Larrid wasp, Tachysphex terminatus. The Antho- phora bees were probably too big game for them, but 1 did find one struggling with a dead bee in the pit. Whether the dead bee had fallen into the pit, or the ant- lion had actually attacked and killed a live one which eame within its reach, I do not know. One ant-lion passed five weeks in the pupal stage; hav- ing spun the cocoon on ‘April 16, 1921, it emerged as adult on May 24, A second one spent six weeks in that stage, or from April 6 to May 25, 1921. House-Spider, Theridion tepidariorum Koch [J. H. Emerton]. These house-spiders should be reckoned as among the original inhabitants of the unit, since their webs were The Ecology of a Sheltered Clay Bank 195 occasionally found about the porch above the bank. Their actual activity was observed only once, on October 12, when one was intermittently feeding upon a large, hairy caterpillar caught in its web. I feel confident, however, that, since they are night prowlers, they participated more fully in the life of the community. Elsewhere I have found this species feeding upon a wasp (Odynerus anormis Say). Résumé of Pioneers. To summarize, then, the five years’ work at the clay bank gave us ag pioneers, those which actually opened the way for others, the eighteen species listed below. It is interesting to note that six of this number were soli- tary bees, seven were wasps, and the remaining five be- longed to as many different orders. Carpenter bee. Xylocopa virginica. Mining bee. Anthophora abrupta. Mining bee. Entechnia taurea. Mining bee. Anthophora raut. Halictus bee. Halictus zephyrus. Small carpenter bee. Ceratina calcarata. Mining wasp. Ancistrocerus fulvipes. Eumenid wasp. Ancistrocerus unifasciatus. Spider wasp. Episyron biguttatus. Pipe-organ dauber. Trypoxylon politum. Mud-dauber. Sceliphron caementarium. Larrid-wasp. Tachysphex terminatus. Paper wasp. Polistes pallipes. Devils horse, Stagmomantis carolina. Stick-bug. Emesa brevipennis. Ant. Crematogaster lineolata. 196 Trans. Acad. Sct. of St. Louis Ant-lion larvae. Myrmeleon mobilis. & M. ammaculatus. House spider. Theridion tepidariorum. (c) Renters. In this chapter are enumerated the secondary inhab- itants of the old clay bank, those which came as ten- ants of the burrows made by some of the creatures listed in the foregoing chapter. These, for the most part, were less hardy than the pioneers; they came into the community, evidently quite willing to utilize its ad- vantages and share its comforts, Often they dropped in merely for shelter or food, and remained to the end of their days. These renters, besides using the old dwell- ings, influenced the population of the unit in two ways —by using the other inhabitants for food, and often by being eaten by them. The Pompilid wasp, Pseudagenia mellipes Say. This wasp was occasionally seen about the clay bank. There were four in 1917 and perhaps a dozen in 1920. In ‘‘ Wasp Studies Afield’’ we have told how this crea- ture nests in both new and secondhand domiciles; some- times she makes small mud nests under loose bark (one such nest was found inside an oak-apple lying on the ground), and at other times she modifies the nests of the mud-daubers to suit her needs. The interest of P. mellipes in the bank was the prey which it sought there. The wasps were often seen going into the old bee-burrows, now occupied by spiders, prob- ably in search of food for their young. On a few occa- sions, they were seen going elsewhere for prey and The Ecology of a Sheltered Clay Bank 197 bringing home their booty. The legless spider brought in by one of these P. mellipes was identified as Pisaurina undata Htz. [C. L. Shoemaker] and on another occa- sion a mother was seen at the base of the bank carrying a half-grown Phidippus spider. This versatile little wasp found in the old bee-holes both home and hunting- ground; she was actually seen to drag a spider up out of one burrow—its lair—and down again into another hole near by—her own nest. Thus we see this citizen finding in the clay bank ‘‘all the comforts of home,’’ shelter or a domicile already provided, and food sup- plies available near her door, and she has just as little work to acquire the one as to get the other, whereas in the usual condition, the home-makers get the benefits of one or the other, but seldom of both. These wasps con- tinue their activity throughout a fairly long season, from the last of May to the first of September. Since they made their own mud cells in the old bee burrows, they are classified with the renters, although they had just as much claim to be among the pioneers. Trypoxylon plesium Roh.* [S. ‘A. Rohwer]. Only one specimen was seen here, and it was taken at the clay bank on July 7, 1917. I saw this wasp actually walk over two distinct spider webs without becoming in the slightest way entangled; evidently it was on a spider- hunt when captured. es 198 ' Trans. Acad. Sct. of St. Louis Trypoxylon clavatum Say. [S. A. Rohwer]. This species of wasp in which the male parent guards _ the nest while the female hunts, has been recorded as making use of old beetle burrows in logs, and cleaning out and using the old cells of the mud-dauber wasps. In the clay blank, they were quite abundant, here using the empty bee burrows, partitioning them with mud and filling each compartment with spiders. When this species first made its appearance in 1917, about June 28, there were only three nests, each with its male guarding the doorway, and four days later there were seven. By the time of the next visit, on July 16, these had vanished, and for two weeks not a single T. clavatum was to be found. On July 31, the first one of the second generation appeared. Time dragged on and this one continued to be the sole representative of the spe- cies. Not until the first of September did others appear, and during that week a half-dozen or more were at work. During the latter half of that month, they succumbed to the inevitable. While 7. clavatwm appeared in the fol- lowing years, their numbers became fewer; the species was not holding its own. No doubt one very consider- able factor in the reduction of their numbers was the destruction I wrought in digging up their nests when their colony was not yet strong, in order to study their prey. The relation of these insects to the unit was pri- marily in using the old burrows; with the male on guard in each nest it is not likely that the young of this wasp serves as host to any parasites. A comparison of the contents of their nests with the list of spider inhabitants of the clay bank shows that they must have gone afield The Ecology of a Sheltered Clay Bank 199 for their prey. These details of the life history will be presented in a later publication. Trypoxylon albopilosum Fox [S. A. Rohwer]. This wasp, with habits similar to those of T. clavatum, likewise oceupied the old burrows of the mining bees. It was first observed on June 28, 1917, when several mothers were at work. They always appeared only in such small numbers, however, that I could not accurately define any seasons or generations for them, only that they were to be seen occasionally until early Septem- ber. Since I opened part of the nests to study the con- tents, it was only natural that their population dwindled away, until in 1920 one lone survivor was observed, The relation of this species to their neighbors was the same as that of T. clavatum. The Monobia mud-wasp, Monobia quadridens. Lin- naeus. This wasp, commonly called the carpenter mud-wasp, but, according to our account in ‘‘Wasp Studies Afield”’ not a carpenter at all, can be regarded as only a sec- ondary inhabitant of the clay bank, since it made no nest of its own, but occupied that of the carpenter-bee, Xylocopa virginica. These wasps were seen each year bringing in mud for partitions and caterpillars for food for their young. On one occasion, as previously re- corded, one Monobia mother actually utilized the tunnel ofa mining-bee, carrying in caterpillars and sealing up the opening with mud. This wasp appeared from year to year in very limited numbers, not more than 6 or 8 being present at work at 200 Trans. Acad. Sci. of St. Louis any one time, and most of the time not even that many. Since the shedding-skins of the silver-winged parasite, A. tigrina, were often to be seen adhering to the wood near the holes, it seemed probable that Monobia suf- fered its share of the ravages of this pest. Their season was long; the first appeared about June 26th, 1917, and on October 3, two were seen, still cling- ing to the woodwork over the bank, stunned and half dead with the cold, while a third one was seen to enter its tunnel. In 1918, they appeared in decreased num- bers, and by 1920, only three could be found. I sus- pected that A, tigrina was more responsible than any other factor for their elimination. The grass-carrying wasp, Chlorion (Isodontia) au- ripes Fernald. [S. A. Rohwer]. This wasp, a very conspicuous if not numerous mem- ber of the unit, occupied the old tunnels of the carpenter bee, and occasionally she used the old burrows which had been made in the clay bank by the mining bees. It 1s very easy to tell just where this wasp mother has her nest, for instead of plugging the opening with mud or otherwise concealing it, she stuffs the aperture with grass, drawing each strand in by its middle, so that a broom-like tuft protrudes prominently, thus revealing at once to an experienced eye the location of her nest. This insect occurred in very modest numbers; no more than three or four individuals were at work at any one time. In 1920, neither the wasps nor their nests were to be found there. There was evidence to lead us to think that the silver-winged parasite had played havoe with these wasps, the same as with the other occupants © of the carpenter bee galleries. The Ecology of a Sheltered Clay Bank 201 While these wasps, as well as the Monobia mud wasp, were to be regarded as secondary occupants of the car- penter-bees’ burrows, the relationship was more com- plex, since by occupying these burrows they took just that much space from the carpenter-bee, because each generation of Xylocopa used the old domiciles, in so far as they were adequate, and if the tunnels had been usurped by other species, the mother carpenter-bees were obliged to spend their time and energy at the slow process of hewing wood instead of being fruitful and re- plenishing the earth. Therefore, the only direct bearing that these two wasp occupants had on the inter-relations of life in the unit as a whole, was to deprive the car- penter-bee of some of her rightful nesting-places and to serve as a host for the parasite, Argyromoeba tigrina, for they both gathered their food elsewhere. The blue mud wasp, Chalybion caeruleum Linnaeus. This wasp was heretofore regarded as the mud- dauber wasp making nests very similar to those of the “tyellow-legs.’? On the contrary, I have shown else- where that this wasp does not make nests of her own, but occupies the nests of Sceliphron caementarium, either by breaking into a ‘‘live’’ cell and destroying the Prey or by using the abandoned cells. In so far as the bank was concerned, however, this species was occa- Sionally seen foraging for spiders among the burrows. On several occasions when one was foraging among the Spider webs, she broke through or became entangled, whereupon, in a very skillful manner, she quickly dis- entangled the web from her person. Elsewhere, how- ever, I have often found dead specimens that had been caught in spiders’ webs. 202 Trans. Acad. Sci. of St. Louis These wasps were found to be present at the bank in great numbers on May 27th and 28th. This was before their nesting activities had begun. The significance of their presence has not yet been determined, so I could only state the circumstances. They were on an elm sprout six feet tall, which grew between the rambler rose and the front of the bank. Other insects were there too, so I thought possibly the aphids, which were plen- tiful there, constituted the attraction, but I could not see that they got anything from the aphids; in fact, the plant-lice were on the under side of the leaves while the wasps always walked about on the tops of them. The wasps were constantly active on the shrub, flying and hopping about, running nervously over the foliage and jumping or dropping from leaf to leaf. The per- formance began at about 7:30 a. m. when the first few arrived and became active; the number rapidly increased as time passed. Standing a short distance away, one could see others coming in from various directions to join the dance—for apparently this gathering was some sort of social function—a stag affair, however, for @ dozen or so taken in the net proved to be all males. I suspect that they were awaiting the emergence of their mates, and that if I had been on hand at the proper time, a little later, I could have witnessed some pretty courtship festivities. Leaf-cutter bee, Megachile campanulae Rob. [J. ©. Crawford]. One specimen of this bee was seen at the clay bank; it was at once taken for identification, and therein the prospective founder of a colony was evidently destroyed, since in the following years this bee was not seen again. ' Lhe Ecology of a Sheltered Clay Bank 203 It was on July 30 that this individual entered one of the burrows. When this hole was opened, a quantity of the resinous material was discovered, but I had no way of knowing whether this or another insect had done the storing. The Megachile bees are leaf-cutters, and make nests in hollow stems, but I have elsewhere* recorded that they have been known to make their leafy cups in sheltered places as under clods of earth. Some bees gather resinous substances, but the Megachiles are not known to do so. Another Megachile bee, M. generosa was taken while at rest on the bank June 3, 1921, but it is unlikely that its presence there was anything but accidental. The Osmia bee, Osmia lignaria Say. [S. A. Rohwer]. This little bee did not appear in the community until 1920. In earlier researches I have found that this spe- cies builds its nests in mud nests of Sceliphron caemen- tarium. Since these bees could not offer the excuse of foraging as an explanation for their presence here, I at once suspected that they were replastering the old bur- rows of the other bees here for their own nests in the same way that they utilize the old mud-daubers’ cells. This I soon found to be true. ‘About a half-dozen of them were seen thus occupied about the bank from May 28 to June 19, 1920, and a month earlier in 1921. They Were also observed gathering pollen from the black- berry blossoms near by. Spider, Ariadna bicolor Htz. [C. L. Shoemaker]. Of spiders, this was the most abundant species among the inhabitants of the clay bank, and from the first of — ey *Trans, Acad. St. Louis, 24:1-71, 1922. 204 Trans. Acad. Sci. of St. Louis September until frost their webs became more and more conspicuous. They made funnel-shaped webs, with the points of the funnels extending deep into the old bee burrows, and got a good living from the small fry that became entangled in the webs, such as visiting snout- beetles, ants, chalcids, flies, ete. I was surprised to find this spider actually feeding upon the bug, Reduvius per- sonatus Linn. [W. L. McAtee]. Occasionally, however, the spiders themselves fell prey to some predatory wasp. Spider, Steatoda borealis {[J. H. Emerton]. Among the most important renters of the old burrows in the bank were the spiders, Steatoda borealis. They made light, irregular snares about the bank, more often about the old burrows, where many of the smaller in- sects became their prey. I have often seen Chalcid para- sites and ants, Crematogaster lineolata, entrapped in their snares. The species may rightly be regarded as an inhabitant of the bank, either a renter or a pioneer. They were very numerous in 1917, and only moder- ately so in the following years. They were as abun- dant as ever even so late as October 3, when one spider was making several brave attempts to subdue a large lo- cust, Tettigidae lateralis var. polymorpha Burm. (A. N. Caudell), which was entrapped in its web. Two un- identified beetles were also their victims that day. Résumé of Renters. The following table summarizes the renters in this social group. They number eleven species, of which seven are wasps, two are bees, and two are spiders. All The Ecology of a Sheltered Clay Bank 205 of these utilized to good advantage the burrows or the nests left by the pioneers, and they in turn left many of their burrows for a third tenant. Among these renters one might also include the bees Anthophora abrupta, be- cause they sometimes used their old burrows for a sec- ond season, after enlarging and renovating them. Renters of Old Burrows. Pompilid wasp, Spider wasp, Spider wasp, Spider wasp, Monobia mud-wasp, Grass-carrier wasp, Cow-bird wasp, Pseudagenia mellipes. Try poxylon plesium, Try poxylon clavatum. Try poxylon albopilosum. Monobia quadridens. Chlorion auripes. Chalybion caeruleum. Leaf-cutter bee, Megachile campanulae. Osmia bee, Osmia lignaria. Spider, Ariadna bicolor. Spider, Steatoda borealis. (d) Visitors. This is a list of insects and larger animals which came into the community quite by accident, or in quest of food or shelter. In some cases they remained as permanent additions to the group, but in most cases, their stay was temporary. Many of them influenced the lives of the pio- neers and the renters in one of two ways: either by using them as food, or by giving themselves as food. Whether the word visitor or transients would be the better term to apply to this group is uncertain. Many insects alighted on the bank quite accidentally; others came there for the definite purpose of securing prey; *% 206 Trans. Acad. Sci. of St. Louis some which came as visitors remained at the bank to nest. It is this latter group which prevents one from definitely drawing the line between visitors and resi- dents; the parents of all the occupants must at one time have been visitors, but not all visitors were occupants. Reptiles. On October 3 I noted a snake (Fig. 16) at:the base of the bank, with a small portion of its body still within the rodent burrow, which I suspected was its home. It lay all day long in one position, as figured, but escaped when I attempted to capture it. At that season of the year, with food scarce, it probably had to wait a long time for a morsel, but if this creature had its home at the base of the clay bank all summer, it was no doubt a considerable factor in the control of the insect popula- tion during the season. Common lizards or swifts. During the entire summer of 1917 several common gray lizards made their home among the piled wood on top of the bank, and were undoubtedly a factor in reduc- ing the life of the group. They, too, had enemies, prob- ably larger ones, which I was unable to discover, and these all attacked the lizards in the same way, in the cases observed. During the summer, three dying lizards were found there at different times, all with torn ab- domens and entrails protruding, as shown in Fig. 17. They were always found in the early morning hours, which indicated that the attack had been made during the dark hours of the night. It was strange that these gray lizards did not appear The Ecology of a Sheltered Clay Banh 207 the following season, although they were abundant along the railroad tracks less than two hundred yards away, but instead the blue-tailed lizard or skink appeared in their place. The blue-tailed lizard had evidently sup- planted its predecessor, but since its habits are the same, very probably its function among and relation to the inhabitants and visitors at the bank were practically the same. This idea was strengthened when on three mornings in early June, 1922, a third species Cnemido- phorus sexlineatus [D. M. Cockran], was seen coming out of a burrow and calmly making the rounds of the clay bank, often entering the bee burrows. It would enter head first, with the tail protruding, and often it would perform the quaint feat of, twisting its tail about the turret while its body was inside the tunnel. It seemed to find the burrows adequately roomy, for it would enter head first and also emerge head first. What it got out of the burrows I do not know, but since this lizard was seen to snap at Anthophora bees in the open, I suppose many of these bees were eaten within the burrows. The Chalcid parasites were also in abundance, and wild roaches were often seen in the burrows. The red-eyed flies, Ganperdea apivora, were numerous at just that time, and the skinks were seen successfully capturing these, and a few undetermined insects, in the open. Toads. In 1920 a big toad was a familiar figure about the bank. Before the end of the season, he had the portly figure of a war-time profiteer, and we felt justified in Suspecting that the rest of the population had suffered accordingly. 208 Trans. Acad, Sci. of St. Louis Phoebe bird. Among the vertebrates which were at home at the clay bank was a phoebe bird and her annual family.’ The nest, on one of the posts, was removed each winter, and each spring a new nest appeared in the same place. One year I noted two broods of young in the nest. Whenever I was at work at the clay bank, the mother bird went afar for her food, but often when I came suddenly upon the scene, I saw her quickly fly away from among the old lumber on top of the bank. From this I suspected that she lost no opportunity to get her insect food near at hand when she was not disturbed. Red bird. A red bird nested in the rambler bushes in front of the bank, during the summer of 1920, but in so far as I could see had no influence on the insect colony. Ground mole. There were a few rodent tunnels under the ground at the foot of the bank, and one year there were two distinct openings, but whether the moles or field mice affected the insect life of the bank I cannot say. Man. The human inhabitants of the dwelling whose porch covered the bank were not in the least a factor modifying this unit; further than the first construction of the site, no relationship existed. The house was almost always vacant except at week-ends, and even then none of this biped fauna came near enough to the bank to influence the life, since they mistook the inhabitants for bumble- The Ecology of a Sheltered Clay Bank 209 bees, and hence kept at a discreet and respectful distance. However, one man was a factor to be reckoned with in considering the enemies of the unit—that was the ob- server. That he should rightly be considered as a non- resident part of this ecological unit is evidenced by the fact that, in taking specimens for identification, he dis- rupted the fine balance of nature, but actually no more so than if he had been a lizard and captured these insects for food. He took only those species which were unknown to him, and then took them in minimum numbers. However, in some cases he was guilty of swinging the balance to some point where, without his interference, it would not have stood. Had he not taken the only Halictus, Mega- chile and others, these insects probably would have be- come established in the bank and their progeny would have done their part to modify the life of the community. The reader can easily estimate the extent of these depre- dations by glancing over the list of the inhabitants and noting all the species that have been submitted to ex- perts for identification. I do not want to exaggerate this point absurdly, but I think it is well that we should realize that man is not a thing apart from the other fac- tors in the balance of nature, but that he is only one species among many in the great game of give and take which makes up ecology. Lepidoptera. Pyralis cuprealis Hubn. (H. G. Dyar]. This visitor was found in the beak of a stick-bug, Emesa brevipennis, on July 16, 1917. Two caterpillars belonging to the Pyralididae. [S. B. Fracker] were found promenading on the bank on Sep- tember 12, 1917, and later in the day a dead one was 210 Trans. Acad. Sci. of St. Louis taken from an ant-lion’s pit. Thus, even though they came to the place quite casually, they served the com- munity. Pseudaglossa lubricalis G. [H. G. Dyar]. One speci- men was picked up dead at the bank on September 3, 1917 A caterpillar belonging to the Noctuidae [S. B. Fracker] was feeding on a small plant growing at the base of the bank. Catocala innubens Guen. [Ernst Schwarz]. During the latter part of July, 1918, several of these moths sought daytime shelter here. Herculia olinalis Guen. [H. G. Dyar]. The caterpil- lar of this moth was found walking on the clay bank on April 28. On the next day it spun its silken cocoon and emerged as an adult on May 22. Hymenoptera. Pompiloides sp. [S. A. Rohwer]. This wasp was forag- ing for spiders in the old bee tunnels during August and early September, 1917. Arachnophoctonus ferrugineus Say. One specimen foraging for spiders on July 30, 1917. Pompiloides americanus Beauv. [S. A. Rohwer]. This had come to the bank as a visitor on a foraging eX- pedition, but had become a victim to the prey which she was pursuing and her lifeless remains were removed from a spider web on July 30, 1917. Tachytes peptictus Say. [S. A. Rohwer]. On July 21, 1920, two or three of these green-eyed wasps were at rest on the vegetation at the edge of the bank. The Ecology of a Sheltered Clay Bank 211 Stlaon sp. (S. A. Rohwer) one was nervously walking about the clay bank on July 30, Sphea (Ammophila) nigricans Dahl. [S. A. Rohwer]. This was seen walking about the clay bank, August 14, 1917, evidently seeking a place for a nest. Since her prey is Lepidopterous larvae, we cannot accuse her of having come to the bank for prey. Sphea (Ammophila) procera Klug. [S. A. Rohwer]. On August 16 this large wasp was seen burying a large caterpillar of the Noctuidae [S. B. Fracker]. Since she left her young ‘‘on the door step’’ of the bank, she should be regarded as an inhabitant or a pioneer; but since I took possession of both, she was thus reduced to a visitor. Odynerus (Stenancistrocerus) unifasciatus Say. [S. A. Rohwer]. This Eumenid was seen moistening a spot on the bank with water and biting out and carrying away the mud, June 28, 1920. Polistes variatus was foraging on the rose bush in front of the bank on May 28, 1920. There was hardly any connection or rather only a far fetched one, in the relations to the inhabitants of the bank, in that the P. variatus would probably carry off caterpillars which, in the natural course of events would have become food for the inhabitants. Hoplisus (Pseudoplisus) phaleratus Say. [S. A. Roh- wer]. This species, seen entering a crevice on July 21, 1920, was distinctly new to the bank. Trypoxylon nigrellum Roh. [S. ‘A. Rohwer]. One was seen foraging for spiders in the old bee tunnels of the clay bank on September 6, 1917. Miscophus americanus [S. A. Rohwer]. Two females taken from the bank on August 31, 1918, were probably seeking a nesting site. 212 Trans. Acad. Sci. of St. Louis Vespa germanica Fabricius. In August, 1920, one worker of this hornet visited the bank at short intervals for three whole days, and as described elsewhere, car- ried off each time a specimen of the Chalcid parasite, Monodontomerus. Pseudagenia architecta Say. The first and only speci- men of this wasp was observed on July 7, 1917, to enter many old burrows of the mining-bees. Its be- havior was very much like that of its cousin, P. mellipes when hunting; hence we suppose that it was foraging. This wasp makes tiny twin cells of mud that are some- what barrel-shaped. Megachile generosa Cress. [S. A. Rohwer]. This bee always returned and rested on the bank after { missed capturing her on June 3, 1922, which indicates that her interests were probably other than being merely a visitor. Halictus pectinatus Rob. [S. A. Rohwer]. ‘A male bee of this species was found dead on the bank, September 6, 1917. Photopsis sp. One individual was making itself at home on the bank. This wasp is supposed to be para- sitic; it had evidently found a host among the population. Camponotus herculeanus L. subsp. pennsylvanicus De G. var. ferrugineus Fab. [W. M. Wheeler]. These ants, many of them workers and some with large heads, were going about the bank, often entering burrows. They first appeared in 1921. Beetles. Corymbites inflatus Say. [E. A. Schwarz]. This click- beetle was at rest at the foot of the bank on July 16. This beetle, or others of its kind, arriving there would The Ecology of a Sheltered Clay Bank 213 probably have no effect upon the inter-relations of life except as they might fall prey to some of the spiders or vertebrates. Photinus pyralis Linn. [E. A. Schwarz]. Many of these ‘‘lightning-bugs’’ spent their days near the clay bank about the first of August. Since they are nocturnal insects and are in a sleepy condition during the day, they should have been easy prey ‘to alert prospectors. No actual observations were made, however, on the effect of their presence upon the others. Hypera (Phytonomus) punctata Fabr. [E. A. Schwarz]. Two specimens were taken at two distinct places on the clay bank on July 30, 1917. There may have been more, but their coloration is such as to make them almost indistinguishable from their surroundings. They evidently fell prey to the ant-lion and other car- nivorous creatures. Rhadopterus picipes Oliv. [E. A. Schwarz]. One speci- men of this beetle was taken July 30 from the jaws of an ant-lion. Coleopterous larvae. Some which could not be identi- fied were often taken from the ant-lion pits. These had evidently fallen from the vegetation above, and were a considerable factor in maintaining so large a number of thriving ant-lions. Epicauta marginata Fab. [E. A. Schwarz}. About August 20, several of these blister-beetles were found dead on the bank. On September 4 and 6 there were again large lots of these beetles, all dead. I cannot ex- Plain the phenomenon, unless it be that some creature had captured them and then spat them out, dead, be- cause of their bad taste. : Hyperodes humilis Gyll. [E. A. Schwarz]. One speci- 214 Trans. Acad. Sci. of St. Louis men of this beetle was taken from the jaws of an ant- lion larva on August 14, 1917. Opatrinus notus Say. [E. A. Schwarz]. A dead beetle taken from a spider’s web, Harpalus dichrous Dejean. [E. A. Schwarz]. One dead beetle found on top of the bank, September 4, 1917. Anisotarsus sp. [E. A. Schwarz]. One individual of this species visited the bank on September 6, 1917. Sphenophorus parvulus. Gyll. [E. A. Schwarz]. This snout-beetle was rescued alive from an ant-lion pit, where the enemy was actively endeavoring to make a meal of him on September 7, 1917. Arhopalus fulminans Fab. [E. A. Schwarz]. Several of these were found on the bank on May 28, 1917. Languria mozardi Lat. [E.:A. Schwarz]. Three speci- mens found on the bank on May 28, 1918. Staphylinus cinnamopterus Grav. [E. A. Schwarz]. Of this rove-beetle, the only visitor was found on July 31, 1918. Tanymecus confertus Gyll. [E. A. Schwarz]. This snout-beetle visited the clay bank on May 20, 1920. Orthoptera. Orocharis saltator Uhl. [A. N. Caudell]. Six of these were on a small plant growing at the base of the bank on July 30, They probably contributed to the food sup- ply of certain predacious inhabitants. Gryllus pennsylvanicus Burm. [A. N. Caudell]. 4 small cricket was seen struggling in the jaws of an ant- lion, into whose pit it had fallen on August 14, 1917, and on August 30, one of this species spent a chilly night in one of the bee burrows, and was the first to poke its friendly head out into the light of day next morning. The Ecology of a Sheltered Clay Bank 215 Ischnoptera pennsylvanica De Geer. An egg-case of this roach was found in one of the old bee tunnels. Just what would have been the chances of survival of the young if they had hatched, we can only surmise. An adult was also found hiding in a crevice on September 6. Locust nymph. This visitor was found as prey of the stick-bug, mesa brevipennis. Nemobius carolinus Scud. [A. N. Caudell]. One dead individual on the bank, September 7, 1917. Melanoplus femur-rubrum. One resting here Septem- ber 7, 1917. Tettigidea lateralis var. polymorpha Burm. [A. N. Caudell]. On October 3, 1917, several locusts of this Species were on the clay bank, probably attracted to the spot for shelter from the cold. One was in the web of Steatida borealis Htz., where a battle occurred between the plucky little spider and the big locust. Hapithus agitator Uhler. [A. N. Caudell]. A male visitor of this species fell prey to a spider, Phidippus tripunctatus [J. H. Emerton] on September 6, 1919. Hemiptera. Pselliopus cinctus Fab. [H. E. Gibson]. This bug was found in a spider’s web on September 12. Anasa tristis De Geer. While this bug was seldom seen at the bank during the summer, on October 3, a dozen adults were found quietly at rest among the rub- bish. Each year they appeared there late in the autumn. These bugs hibernate as adults, and had come to the bank seeking crevices in which to spend the winter. They probably appeared upon the scene too late to in- fluence the lives of others. On one occasion (May 20) a young spider, Phidippus sp. [J. H. Emerton], was seen feeding on one of these bugs. 216 Trans. Acad. Sci. of St. Louis Reduvius personatus L. [A. T. McAtee]. This pro- tectively colored bug, concealed in a crevice was sucking the life-blood from a pioneer inhabitant, Ancistrocerus fulvipes, on June 28, 1920, and earlier in these pages I state that this bug was oS upon by the spider, Ariadna bicolor. Arachnida. Phidippus. clarus Keyserling. [C. L. Shoemaker]. One specimen was found on the bank on July 16, 1917. It was probably in quest of food. This is one of the ‘‘wandering’’ spiders, and its wanderings into the unit were evidently not without purpose. Phidippus sp. [J. H. Emerton]. A female actually made a permanent home in one of the burrows, where her face could often be seen peering from the doorway, and occasionally she would catch and devour a Chalcid. Phidippus tripunctatus [J. H. Emerton]. One young one feeding upon a visiting male cricket Hopithus agt- tator Uhler. [A. N. Caudell], on September 6, 1919. Prosthesima sp. [J. H. Emerton]. A female was walk- ing on the bank, June 28, 1920. Thanatus lycosoides Emerton. [C. L. Shoemaker]. One specimen found at the base of the bank on September 23, 1917. Habrocestum pulex Htz. [J. H. Emerton]. One speci- men on the bank September 23, 1917, feeding upon an ant, Crematogaster lineolata. Lycosa scutulata Htz. On July 30, 1917, at twilight, a male of this spider was quietly resting at the base of the bank. The Lycosidae are popularly know? as wolf spiders; they are vagabond, hunting spiders, spinning no snare but chasing their prey upon the The Ecology of a Sheltered Clay Bank 217 ground, This sly watcher undoubtedly had business to transact at the clay bank. Lycosa sp. During the first part of August, several were seen repeatedly at the same place. It seemed that twilight brought out these night prowlers. Xysticus ferox Htz, [C. L. Shoemaker]. One speci- men at the clay bank at dusk, July 30. This is one of the crab spiders, and like the others, probably came in quest of food. Spider. A small unidentified specimen taken from the jaws of the ant-lion on September 3, 1917. Pardosa sp. [J. H. Emerton]. This was another of the wandering hunters which came to the bank October 3, when food elsewhere became scarce, and one specimen was walking about on the bank one sunshiny morning of February 26, 1921. : Drassus sp. [J. H. Emerton]. One at bank October 3, 1917. Gonyleptes sp. Several of these daddy-long-legs were on the bank, and about a dozen had congregated on a small plant on July 30, 1917. Warburton* says the members of this group are ‘without doubt essentially carnivorous.’? Hence the presence of these was prob- ably of serious significance, especially to the smaller Species of residents. Ixodes sp. This tick was abundant each year at the end of July and in the early part of August among the old lumber on top of the bank. Myriapoda. On several occasions during 1917 and 1918, dead speci- mens of Myriapods were found on the bank. Since none *Cambridge Nat. History 4:44, 1909. 218 Trans. Acad. Sci. of St. Louis were found alive and often the dead ones were partly disintegrated, and since these species are known to limit their habitation to rotten woods, I could think of no explanation for their presence here in such condi- tion except the far-fetched theory that they had been brought here by other animals, possibly lizards, and abandoned or spit up. ‘Among those thus found were Euryeurus erythopygus Brandt. [R. V. Chamberlain], Cleidogona caesioannulata Wood. [R. V. Chamberlain], and Julius sp. [R. V. Chamberlain]. Résumé of Visitors. The following table recounts species found among the visitors to the clay-bank community: Snake, Lizards, Toads, Phoebe bird, Red bird, Ground mole, Man, Lepidoptera, Pyralis cuprealis. Caterpillars, Pyralididae and Noctuidae. Lepidoptera, Pseudaglossa lubricalus. Catocala moth, Catocala innubens. Moth, Herculia olinalis. Wasp, Pompiloides sp. Spider wasp, Arachnophoctonus ferrug’ neus. Wasp, Pompiloides americanus. Wasp, Tachytes peptictus. Wasp, = Silaon sp. The Ecology of a Sheltered Clay Bank 219 Pompilid wasp, Leaf-cutter bee, Bee, Parasitic bee, Beetle, Lightning beetle, Beetle, Beetle, Beetle larva, Blister beetle, Beetle, Beetle, Beetle, Beetle, Beetle, Long-horned beetle, Beetle, Rove beetle, Snout beetle, Orthopteron, Cricket, Wild roach, Cricket, Locust, Sphex nigricans. Sphex procera. Odynerus unifasciatus. Polistes variatus. Hoplisus phaleratus. Trypoxylon nigrellum, Miscophus americanus. Vespa germanica. Pseudagenia architecta. Megachile generosa. Halictus pectinatus. Photopsis sp. Camponotus herculeanus. Corymbites inflatus. Photinus pyralis. ° Hypera punctata, Rhabdopterus picipes. _ Unidentified. Epicauta marginata. Hyperodes humilis. Opatrinus notus. Harpalus dichrous. Anisotarsus sp. Sphenophorus parvulus. Arhopalus fulminans. Languria mozardi. Staphylinus cinnamopterus. Tanymecus confertus. Orocharis saltator. Gryllus pennsylvanicus. Ischnoptera pennsylvanica. Nemobius carolinus. Melanoplus femur-rubrum. 220 Trans. Acad. Sci. of St. Louts Orthopteron, Tettigidea laterals var. polymor pha. Orthopteron, Hapithus agitator. ug, Pselliopus cinctus. Squash bug, Anasa tristis. Bug, Reduvius personatus. Spider, Phidippus clarus. Spider, Phidippus tripunctatus. Spider, Phidippus sp. Spider, Prosthesima sp. Spider, Thanatus lycosoides. Spider, Habrocestum pulex. Spider, Lycosa scutulata. Spider, Lycosa sp. Spider, Xysticus ferox. Spider, Pardosa sp. Spider, Drassus sp. Daddy-long-legs, Gonyleptes sp. Tick, Ixodes sp. Myriapod, Euryeurus erythopygus. Myriapod, Cleidogona caesioannulata. Myriapod, Julus sp. The list totals 71 species, and covers a ionhe of forms, from man down to Myriapoda. No attempt was made to list the microscopic organisms, although with proper equipment an interesting list of such specimens could doubtless have been found. It is interesting to note that of these 71 species which were found visiting the clay bank coincident with my intermittent visits 7 were vetebrates, 5 were species of Lepidoptera, 18 Hymenoptera, almost all of which were wasps, 15 were beetles, 7 Orthoptera, 3 species of bugs, 10 Arachnida and 3 Myriapoda. The Ecology of a Sheltered Clay Bank 221 It will be seen that some of the creatures were of little or no importance in the balance of life on the clay bank; for instance the squash-bug, Anasa tristis, which is a vegetable feeder and which few creatures would eat, would have little influence on the life of the social group. On the other hand, some of the visitors wielded a significant influence, as in the case of Vespa germanica which carried off the Chalcid parasites which preyed upon the bees. Many of the visitors might be placed alternatively in still another group, those which might properly be considered as either renters or pioneers, as for instance, the spider-hunter, Pseudagenia archi- tecta, which might be said to carry on its daily busi- ness at the bank, although it maintained its home else- where. Of course the sharp line of demarcation between the groups is difficult to define while the possibility is always open for the visitors to become residents there and participate in all the joys and sorrows incidental to influencing the balance of nature already present, by preying, by being preyed upon or by parasitizing other creatures or by becoming hosts for other para- Sites, (e) Parasites. The following account presents the details of the activities of the last group of the inhabitants of the bank, the parasites, which affected the pioneers and the renters. These exerted their influence upon the balance of population in a variety of ways, and in turn were 1n- fluenced in interesting ways by some creatures from all three groups. 222 Trans. Acad. Sci. of St. Louts Parasitic Hymenoptera. Chalcid parasites, Monodontomerus sp. [J. C. Crawford]. When the observations began on June 25, 1917, and the Anthophora bees, etc., first appeared that spring, hundreds of these parasites were at once on hand. Their emergence was almost simultaneous. The number on this first day of my visit was so great as to make me fear the appalling mortality that must be in store for their hosts, the bees, even if only a small portion of them were to be permitted to beget offspring. They were actively searching over the bank, entering burrows and dancing before the bank in the bright sunshine. I could then get direct evidence only that they had come from the cocoons of Anthophora abrupta, but they were undoubtedly parasitic upon other species as well, since they were found at the bank also in the fall, and the bees, A. abrupta, ended their life cycle in July. One burrow of the mining-bee, when opened, exposed to view fifty Monodontomerus adults; I could not be ab- solutely sure, however, that all of these were young parasites just emerging from this cell; it is possible that some of the sun-dancers had left their companions and crowded into this cell of mixed sexes. In 1920, on July 24, 13 cells of this Anthophora bee were taken into the laboratory, only four of which proved to be infected by this parasite. These four cells gave forth 139 adult parasites, 109 of which were females and 30 males, or an average of about 35 for each bee cell. At such an alarming rate of increase, it is surprising that any bees at all should have escaped them. These parasites throve and appeared in large num bers in each of the seasons of observation. There 1# The Ecology of a Sheltered Clay Bank 223 more than one generation each year. This made a par- ticularly difficult situation for the species of hosts which produced but one generation each year. The first adults that emerged in the spring, lived and disappeared simultaneously with the Anthophora that closed their life cycle about the middle of July. The eggs deposited by the first lot of adult parasites produced adults later in the summer. A careful and frequent examination showed that the Chalcids were not in evidence from the time of their waning and first disappearance, July 16, up until September 3, when they re-appeared after an interval of forty-eight days. This second generation also comprised an enormous population. Many of these were seen also to enter the bee burrows, and some of the bee cells dug up later also contained these Chalcids in the pupal stage. Not all of them succeeded in their parasitic plan of existence; some fell prey to other inhabitants early in life. On the first day of the appearance of the second brood, seven were taken dead from the spiders’ webs nearby, and elsewhere in these pages is recorded the fact that Vespa germanica was making regular trips, carrying them off day after day to her nest. On September 6, while the numbers flying about seemed just as great, still many were found dead about the bank; [ cannot account for this, since this early date seems hardly time for a natural death. These may have been the dead of the previous generation, or those of the Present generation that had perished before emergence, Which some industrious Trypoxylon wasps had swept out of the old burrows that they were renovating for their own use. Mortality was really upon them, how- ever, because by September 12 a marked reduction in 224 Trans. Acad. Sci. of St. Louis their numbers was easily noticeable. On October 3, none at all were abroad, but of course at this date prac- tically all of the life on the bank was still. In 1918, when the carpenter-bees, the Anthophora mining-bees, and the caterpillar wasps all appeared a month earlier, about May 25 (because of temperature conditions), these Chalcid parasites too appeared a month earlier. All of the life in the bank responded to the same laws. In 1918 these parasites were just as numerous but I was surprised to note no increase over the previous year. It would have seemed reasonable to expect that the myriads of the last year would have multiplied as greatly, at the expense of everything else, that this year there would be little or nothing but Chal- cids. But to our surprise the 4. abrupta bees, even under such persecution, were more abundant the second year than the first. This is a good example of the delicate balance in nature’s game of cat and mouse. By June 28, 1918, as many chalcids were dead and strewn about the bank, in burrows and in crevices, a8 were alive and on the wing; thus that date marked the decline of the first generation, which in the preceding year occurred on July 15. SomeAnthophora cells taken a little latter contained live pupae. So, comparing the dates of this species for 1917 and 1918, we see that when the first generation got a thirty days’ earlier start, 80 likewise did the second generation. In 1920 this species appeared in about the same num- bers, and the interval dividing the two generations again came in July. To be exact, on my visit on July 21, not one Chaleid was to be scen. But on the next inspection, September 2, they were out again in great numbers. These of the second generation were, in turn, all gone The Ecology of a Sheltered Clay Bank 225 the first of October. The dates of these two generations were practically the same as in the first year; the first generation ran from late June to mid-July, and the see- ond appeared about the first of September but disap- peared before the end of the month, The animal most abundant in an ecological unit should be regarded as the dominant one. If this rule includes parasites, then we must admit that this Chaleid is the most abundant and dominant insect in the unit. They were already plentiful at the time of my first ob- servation; their hosts were well established and numer- ous, and since each host is capable of bringing to matur- ity, on an average, thirty or so parasites, we see how fa- vorable were all their prospects. And yet we come face to face with the fact that other parasites, Diptera, etc., even though they appear in far smaller numbers, di fully as much destruction to the permanent residents of the bank as did these many little Chalcids. What does it matter to an Anthophora larva, whether it yields its life to produce thirty-nine chalcids or one cuckoo-bee? Thus, we see that the most important aspect of the life of the Chalcids is to keep the bees from overrunning everything by their enormous increase in population, which would doubtless take place without this deterring influence. As a secondary function they serve as food for some species that in no other wise influence the life of the bank, such as non-resident Vespa germanica, and also to feed spiders which, by being thus encouraged to come to or stay on the bank, might also prey upon the other residents. 226 Trans. Acad. Sci. of St. Louis Cuckoo-bees and parasitic wasps. These beautiful green bees, parasitic upon many forms of Hymenoptera, appeared about the bank dur- ing every season of the term of observation. They were directly parasitic upon many of the inhabitants, and were frequently observed following a returning mother to her nest, and were often seen prowling about and entering the nests of others. Only one species was found, Chrysis (Tetrachrysis) coeruleus Fab. [S. A. Rohwer]. These cuckoo-bees ap- peared with the first life on June 25, 1917, and were in almost constant attendance at the bank in summer from that date until near the middle of October. They appeared in great abundance in early summer, flying before the bank in the sunshine, searching over the sur- face and entering numerous crevices and bees’ tunnels, evidently taxing the legitimate life in the bank as much as did the Chalcids. Their numbers seemed to increase as summer advanced; but numerical estimates of this population are difficult to make at the early season. One may count the turrets of the bees, the pits of the ant- lions, or the webs of the spiders, but these transients sneaking about the dwellings of others, could not be definitely enumerated. The next year by August 10 their number seemed much reduced; on August 14 only one specimen Was seen, and none thereafter, excepting dead ones at the foot of the bank. One carcass of this bee was removed from the web of a spider. This was probably the natural time for the death of these bees, and we thought that a new generation would appear soon after. It was interesting to discover that with the disappearance of The Ecology of a Sheltered Clay Bank 227 the cuckoo-bees, the silver-winged parasitic Dipteron made its appearance. ' On August 30, only one cuckoo-bee was present; this seemed a forerunner of the second generation, rather than a straggler from the first, for the next day three were out, and thereafter they increased briskly until a thriving second generation was out and in action. From September 4 to 6 this population was at its height; by September 12 they, along with the other species then about the bank, began to disappear as the chill nights came on, but whether they perished with the cold, or made their natural demise, or merely were in hiding, I cannot say with certainty. However, on one late Sep- tember day, when the bank became warm in the sun- shine, two appeared; one was especially active in annoy- ing a turret-bee. No doubt many of these parasitic bees died in their burrows. Their latest date recorded was October 13. Few of these superchitinous insects fall prey to the hungry neighbors. The rule ‘‘to eat and be eaten’’ does not apply to the cuckoo-bee, which appears to be the armadillo of the insect world. Pseudomelecta interrupta Cress, [S. A. Rohwer]. This bee was seen only once at the clay bank, on July 21, 1920. It entered one of the nests of Anthophora, and was taken for identification as it emerged several minutes later. hice The species must have a highly interesting life his- tory, for Wheeler (Proc. Am. Phil. Soc. 58:14. 1919) classifies it as one of the parasitic bees, derived from the ancestral genus Anthophora and probably parasitic on Anthophora. He questions this later statement, but since it entered the burrow of Anthophora after the 228 Trans. Acad. Sci. of St. Louis manner of a parasite, I think much doubt as to its habits is removed. This of course would mean that P. inter- rupta, having at some time been derived from the genus Anthophora has found a way to escape the Anthophora responsibilities of mining and food gathering by turn- ing parasitic on its own kin, just as seems to be the case with Psithyrus in the nest of Bombus. Mutillids. The cow-killers or velvet ants, with wing- less females and winged males, are well known to be parasitic, and their way of prowling about the clay bank indicates that, like many of our own species, seeking hosts is the sole ambition of their lives. They undoubt- edly found hosts in the clay bank. Dasymutilla ferrugata Fab. [S. A. Rohwer]. On June 28, 1917, one female was seen to enter a dozen burrows of the Anthophora bee, and in some it remained a sufficiently long time to have done mischief. Other mutillid parasites, which remained unidentified, were seen doing precisely the same on July 31, and August 10. Sphaerophthalma scaeva. Two females of this para- site were darting about as if on mischief bent, on Au- gust 30, and on the next day a half-dozen or more were seen. We have previously reared these parasites from the old nests of the mud-dauber, Sceliphron caementar- qum., On the clay bank, besides D. ferrugata and S. scaeva, one would occasionally pick up an unknown specimen of the Mutillids; I suspect that they were seeking hosts, but their numbers were so insignificant in comparison to the other parasites that I do not think they were to be regarded as a ‘‘real force in the community.”’ Sphaerophthalma pennsylvanica Lep. [S. A. Rob- wer]. A small elm shrub, between the clay bank and The Ecology of a Sheltered Clay Bank 229 the rose bushes, seemed especially attractive to a large variety of insects. Flies and parasitic Hymenoptera were particularly abundant in variety, but for large numbers of a single species only two were notable; they were Chalybion caerulewm males, discussed elsewhere in these pages, and males of this species. There was another similar shrub near by, but no such insect popu- lation was apparent upon it, so I concluded that it was the aphids present here which had attracted them to this one, although I could see no actual relationship ex- pressed in their activities. Perhaps in both species, the males were merely marking time until the appear- ance of the females. It is interesting to note, how- ever, that while the males were so near to the clay bank on this occasion, they probably flew here from else- where, for the females of this parasite have never been seen at the bank. Sapyga sp. [S. A. Rohwer]. One was walking about on the bank on June 29, 1917. Members of this family are, according to Rohwer (Conn. Nat. Hist. Survey 22:620. 1916), parasitic on bees and wasps, while Packard (Guide to Study of Insects, p. 134. 1876) says that in Southern Europe Sapyga repanda is parasitic in the nests of Xylocopa violacea; Sharp, however, (In- sects, Pt. 2, p. 100) says that Sapyga 5— punctata was seen carrying caterpillars. : Sphecodes sp. Two specimens of this red-bellied parasitic bee were seen going in and out of the burrows of the resident bees on June 28, 1920. Diptera. Of all the Diptera taken at the bank, not one speci- men was other than parasitic; the group as a whole may 230 Trans. Acad. Sci. of St. Louis be justly regarded as doing as much damage to the law- ful inhabitants as either the Chrysits or the Chalcids. The Diptera, however, came later in the season, and possibly sought different victims. Dipterous parasites were first seen on July 30, 1917—a half-dozen represent- ing two or three species. As days went by, they became more numerous in species and numbers. The details be- low relate their activities. Copecrypta ruficauda Coq. [C. H. T. Townsend]. This fly was seen to rest for several minutes on the clay bank on August 10. While I was wondering how I could capture it, it flew to my wrist and became so in- tent on drinking the perspiration that I picked it up easily in my hand. It was the only specimen seen there. Archytas aterrima R. D. [C. H. T. Townsend]. This fly belonging to the family Tachinidae, and commonly called the tachina fly, is parasitic on other insects. One specimen was taken at the bank on July 31. Like Cope- crypta ruficauda, this fly was taken when it flew to my hand to drink the perspiration. Sarcomacronychia trivittata T. [C. H. T. Townsend]. The first time this fly was seen at the bank was September 6, when two specimens were taken. They spent their time flying about the bank, often resting upon it and seldom going far away from it; this con- duct indicated that there must be some attraction there. A third individual was observed on October 3, 1917. Parametopia sp. [C. H. T. Townsend]. One such pest was seen to follow the wasp, Ancistrocerus fulvipes, as she entered her nest laden with a caterpillar on Sep- tember 12. A second individual was scen to follow the same species of wasp on October 3, 1917, as she was 8% ing to the nest. This does not mean that this fly has The Ecology of a Sheltered Clay Bank 231 become so specialized as to require a specific host, for we have often seen it behave so with other wasps else- - where, Argyromoeba anale Say. One specimen of this para- sitic fly was on the bank September 7, 1917. Argyromoeba oedipus Fab. [F. Knab]. Six specimens of this parasite were seen about the clay bank on July 31, one of which actually entered a burrow of the white- banded bee, Entechnia taurea. Argyromocba fur O. S. [F. Knab]. One was at rest on the clay bank on September 3, 1917. In 1918, this species, along with all the other life in the unit, appeared about a month earlier, or on May 28. (The 1918 material was identified by C. T. Greene.) Argyromoeba tigrina De G. [F. Knab]. This para- Sitic fly (Fig. 18) first appeared on August 8, 1917. At first there were only three individuals; two days later they were present in very great numbers. They frisked and flitted about in the sunlight, stopping to rest for long periods on the wooden part of the porch and in the direct sunlight. When at rest thus they could easily be Picked up with the fingers. The peculiar life history of this fly enables us to know absolutely that they had not flown to the bank from afar, but had actually emerged on the scene. The tell-tale evidence is this: in trans- forming to adulthood, this species leaves the home of its host while it is still in the pupal stage and, upon gaining its freedom transforms into the adult and leaves the old shedding-skin (Fig. 19) on the surface near the point where it emerged. In many places on the face of the clay bank, and at its base, up among the Wooden partitions very near to the tunnels of the car- Penter-bees, Monobia wasps or grass-carrier wasps, and 232 Trans. Acad. Sci. of St. Louis even protruding from the bee burrows, were these shed- ding skins adhering conspicuously. This indicated just which species were probably their hosts. That they were quite numerous I was also certain, since I picked up more than sixty of these shedding-skins, and of course these were not all. While there is sufficient evidence that these para- sites prey upon the inhabitants, and while they spend most of the sunshiny hours each day in flying to and fro before the openings of the burrows or resting near by, and although I have watched for hours at a time, I have never yet seen one of these creatures attempting to enter a burrow. I especially watched for this, since the great Fabre has discovered for another species of this genus some complicated aspects of life history. True I have often seen the females hovering over a bur- row, poising on the wing above the aperture and dipping down again and again, much in the manner of a dragon- fly depositing her eggs under the water, and I suspected that with each dip she deposited an egg but, like Fabre, I have never discovered the egg. Fabre seems to think, for his species, that the parent fly oviposits by merely dropping a minute egg while flying over the surface of the mud walls which contain the grubs of the host, and that the larvae hatching from the eggs are wonderfully adapted for breaking into the masonry to reach the host by being provided with a very horny, deflexed head, armed in front with stiff bristles and under the body with several pairs of elongate setae serving as organs of locomotion. In the species under present consideration, the eggs are probably deposited in this manner, although some other species of this genus, as A. oedipus, ate known to enter the bee burrows. Several females of A. The Ecology of a Sheltered Clay Bank 233 tigrina were seen repeatedly to fly violently against the wood beside the Xylocopa or other burrows; then often I could see the tip of the abdomen enlarge, as it was bent downward and tightly pressed against the rough surface, and moved in a circular motion as it rubbed against the wood. In only one instance did I actually See something fall, but I was not near enough to see whether it was an egg or excrement. So the problem of the life history of this parasite remains unsolved. It would at first appear that A. tigrina came too late in the season to affect the bees, Anthophora abrupta, since their lives had terminated before the advent of the parasites. This might be true, and A. tigrina might not affect this species but might be a direct cause of the gradual lessening of the numbers of the white- banded bees, Entechnia taurea, which were contempo- raneous. However, if Fabre’s theory holds true for this species and these parasitic young possess those won- derful adaptations for reaching their host even through prison walls, then the parasites would affect the dormant generation of Anthophora, and other species as well. There is additional evidence, however, in the fact that I found the shedding-skins on the face of the bank, but not on the top, and A. abrupta builds only in the face of the bank. This would lead us to think that A. abrupta is a host despite the fact that the adults are not con- temporaneous. I want to repeat that in constant watching during these dipping maneuvers, I failed to see an egg drop, but from the insect’s behavior I felt sure that something was happening. It is reasonable to believe that the egg is minute, and with the observer four feet away (one can- 234 Trans. Acad. Sci. of St. Louis not successfully come closer) something might happen beyond one’s sense of sight. These adults are flower-loving insects. On August 13, two specimens were seen on flowers in a field some dis- tance away. It may well be that this would account for the reduced numbers during the morning hours, while at noon they were present in full force; perhaps they were out in the blossoms satisfying their morning hunger, or again perhaps they had gone out into the vegetation to spend the night (their sleeping habits have not yet been ascertained) and were slow in returning to the day’s activity. With the cold nights of early September they began noticeably to wane, and none were seen alive after September 12. These flies were seen throughout the season in vary- ing abundance. It seemed at first that with their play- ing and dancing about the bank they were merely mark- ing time, but, since they continued it all summer, it ap- pears that they are the kind of beings that can seemingly dance and frolic through life and yet make a success of living. Some were seen in copulo on July 26, 1922. One pair remained in copulo, back to back, for a half hour, resting on a shady portion of the bank. In 1918, this fly was no exception in the phenomenot of all life appearing a month earlier. A few individuals were seen out on July 17, 1918, as against ‘August 1, 1917. Until the end of July they were very abundant, and like their ancestors of yesteryear, they kept busy most of the time, doing nothing in particular, unless they oviposited on the sly. The shedding-skins as well as the flies gave evidence of an increase in population. This increase was probably at the expense of the dwellers in the wood-tunnels above, for as I have already show? The Ecology of a Sheltered Clay Bank 235 these and the white banded bees declined simultaneously with the increase in this parasite. I do not know what happened to the species in 1919, but in 1920 they appeared in limited numbers, and in 1921, they were again on the increase. This fly evidently has a sensational criminal life his- tory, which I hope some day will be brought to light. Ganperdia apivora Aldrich [J. M. Aldrich]. In June, 1920, a dozen or more specimens of this red-eyed diptron were flying about the burrows in the clay bank, in a manner strongly indicating parasitic habits, although I did not at any time actually see them enter the burrows, and the same observation was made on May 27 to May 28, 1921. These bees are undoubtedly parasitic on the mining bees in the clay bank, since I actually observed them hatching in the laboratory from cells of Antho- phora abrupta on May 20 to 22, 1921, from material taken about nine miles north of the clay bank. This species is described by J. M. Aldrich* from spec- imens taken near St. Louis in 1877. It is very interest- ing, too, that forty-five years ago this fly was found in a clay bank about the nests of the mining bee, Antho- phora abrupta. That it is parasitic upon these bees is evident from the statement in the paper cited, that one ‘of the bee cells contained cocoons which gave forth adult flies of this species in the following Spring. In the first week of June, 1922, hundreds of these red- eyed flies were out at the bank and were then being freely preyed upon and devoured by the lizard (Cnemt- dophorus sealineatus) which made itself at home about the place. *Proc. Ent. Soc. Wash. 21:106. 1919. 236 Trans. Acad. Sct. of St. Louis Lepidophora lepidocera Weid. [C. T. Greene]. This parasite was present at the clay bank on July 1, 1922. It was often seen bobbing up and down on the wing before the burrows, but if eggs were dropped in the process, none were ever discovered. Parasitic Beetles. Hornia minutipennis Riley [E. ‘A. Schwarz]. This beetle, which is parasitic upon the mining bees, was found only twice at the clay bank. One was taken at the base of the bank on May 27, 1920; it deposited many eggs in captivity the next day, and died soon after. An- other was taken from the cell of A. abrupta on October 2, 1920; it emerged in the warm laboratory on March 19, 1921, promptly ate of a piece of apple and consumed part of an Anthophora larva which was given it; the re- mainder of the larva it allowed to dry up. It lived five weeks. 7 C. V. Riley* describes this species from specimens reared from the cells of Anthophora sponsa taken from elay banks near St. Louis. Piercet tells us that the genus Hornia is known to be parasitic on Hymenoptera; the larvae are conveyed to the host by some other insect —that is, they are passively conveyed. Their feet are specialized for clinging, not digging. Their food is generally honey; hence the mouth parts are reduced. Hornia shows degeneracy in the adult stage with com- plete loss of the wings, and almost entire reduction of the elytra. Fabret has found a parasitic beetle, Sitaris humeralis *Trans. Ent. Soc. St. Louis, 3:563-565. 1877. Univ. Nebraska Studies, 4:153. 1904. arp, Insects. Pt. II, p. 272. The Ecology of a Sheltered Clay Bank 237 by name, living at the expense of bees of the genus Anthophora. ‘‘The eggs of Sitaris are deposited in the earth in close proximity to the entrances of the bees’ a ar a single female producing upwards of two thousand eggs. ..... They hatch in about a month, producing a tiny triungulin of black color; the larvae, however, do not move away, but, without taking any food, hibernate in a heap until spring, when they be- come active. Although they are close to the abodes of the bees, they do not enter them, but attach themselves to any hairy object that may come near them, and thus get onto the bodies of Anthophora and are carried to the nest.’’? At first it seemed strange that at the clay bank at Wickes they were present only in such small num- bers, but this is explained by the statement from the Same work that the triungulins distribute themselves on all sorts of unsuitable insects, so that it is possible that not more than one in a thousand succeeds in gain- ing access to the Anthophora nests. Résumé of Parasites. To summarize this group, I have found in the clay bank parasites to the number of 19 species. This is highly interesting in the light of the relation of this number to the number of pioneers and renters. Of the first there were 18 species, many of which could not possibly be host of these parasites; of the latter there were 11 species, including spiders and others which likewise could not serve as host. Leaving out of consid- eration all the non-host species of both the pioneers and the renters, we find the parasites greatly outnumbering legitimate residents of either class. The following table recounts the parasites that visited the bank. 238 Trans. Acad. Sci. of St. Lowis 3 Parasites. Chalcid parasite, Monodontomerus sp. Cuckoo-bee, Chrysis (Tetrachrysis) coeruleus. Parasitic bee, Pseudomelecta interrwpta. Cow-killer or velvet ant, Dasymutilla ferrugata. Cow-killer or velvet ant, Sphaerophthalma scaeva. Cow-killer or velvet ant, Sphaerophthalma pennsyl- vanica. Parasitic Wasp, Sapyga sp. Parasitic Bee, Sphecodes sp. Parasitic Fly, Copecrypta ruficauda. Parasitic Fly, Archytas aterrima. Parasitic Fly, Sarcomacronychia trivtt- tata. Parasitic Fly, Parametopia sp. Parasitic Fly, Argyromoeba anale, Parasitic Fly, Argyromoeba oedipus. Parasitic Fly, Argyromoeba fur. Parasitic Fly. Argyromoeba tigrina. Parasitic Fly. Ganperdia apivora. Parasitic Fly. Lepidophora lemdocera. Parasitic beetle. Hornia minutipennis. It will be seen from the table that, with the exception of one beetle, all of the parasites belonged to the orders Diptera and Hymenoptera. Of the former there were 10; of the latter, there were 8, comprising one chalcid, three species of cow-killers or velvet ants, one wasp and two species of bee which only recently phylogenetically speaking, have acquired parasitic habits, and one spe- cies of cuckoo-bee. With this appalling array it is @ wonder that any host can continue to exist; there is 20 The Ecology of a Sheltered Clay Bank 239 telling how many species have already become extinct through this agency during the existence of the clay bank. (C) RELATION OF POPULATION TO ENVIRONMENT, (a) Relation to Temperature. Insects know not dates; May or December, June or January mean nothing to them other than heat or cold. The insects’ calendar is the thermometer. Instead of saying that all the spring Hymenoptera, regardless of species, emerged during the latter part of June, I should say that they emerged when the average mean tempera- ture for fifteen days reached 73°. Let us not forget that June has nothing in its makeup to mean anything to the Hymenoptera, but 73° for fifteen days at the springtime of the year means to them life, sunshine and food. My attention was first attracted to this phenomenon in noting that in 1918 the life appeared from the clay bank about a month earlier than in 1917, and in 1920 © they appeared at almost the same time as in 1918. In other words, the hymenopterous life about the place was in its spring fullness on June 25, 1917, May 28, 1918, and May 30, 1920. The average mean temperature for the fifteen days preceding and including June 25, 1917, was 73 1/25°; the average mean temperature for fif- teen days preceding and including May 28, 1918, was 73 4/25°; and for the same period preceding May 30, 1920, was 731/2°. These temperature figures of course are only relative; the clay bank was facing the eastern sun, whereas, the temperature records were made in the shade; let us remember, too, that the life potential buried in the clay bank was likewise in the shade.* This corre- *The temperature records are from the U. S. Meteorological Re ports taken at St. Louis, twenty miles north of this site. 240 Trans. Acad. Sct. of St. Louis lation works out so beautifully that I shall want to place this material experimentally at a temperature of 73° in November or February, to see if then the develop- ment and maturity occurs likewise. Had the 1917 pop- ulation (which emerged in abundance about June 25th) emerged on May 28, as did the 1918 lot, it would have spent the 15 days before its emergence at a temperature of 66 2/15°. This temperature phenomenon seemed to hold in gen- eral for most of the spring Hymenoptera, and I found them all appearing at one time, after the manner of Anthophora abrupta for example. The bank yielded its contents of this species all at one time each year, within a space of two or at most three days. Simulta- neously appeared also their Chalcid parasites, Mono- dontomerus. The green cuckoo-bees burst forth at their appointed time, and so too the caterpillar wasp, Ancis- trocerus fulvipes; the Monobia mud wasp, the Trypoxy- lon, ete., ete. The only exception was the carpenter- bee, Xylocopa virginica which appeared some time earlier.* This species is a wood dweller—not a cave dweller— and would, therefore, be influenced by temperature somewhat differently from those which live in the com- pact earth. Not alone in 1917 did I see this simultane- ous emergence of the various species and of all individ- uals of each of these species, but in 1918 and 1920 as well. It was as though some one touched an electric button and that button was labeled 73° F. The white-banded bees E. taurea and the dipterous parasites, demanding a higher temperature and hence *We do not know how this bee hibernates, but observations given elsewhere indicate a period of hibernation in the adult stage. The Ecology of a Sheltered Clay Bank 241 emerging later, conformed nicely to the precedent set by the spring Hymenoptera in the year 1918, when the meteorological conditions were very different from 1917. This bee population emerged in 1917 on July 16 and in 1918 with a more favorable temperature 18 days earlier on June 28. In this species I found, in so far as the records were kept, that they responded, by emerging from the pupal condition, to a temperature of 77°. Using the same weather records as a basis, we find that for the fifteen days preceding their emergence on July 16, 1917, we had an average mean temperature of 77 2/15°. In 1918, their appearance was earlier, June 28, and the average mean temperature for the period of fifteen days preceding this was 77 2/3° F. Thus, we can see in the emergence of this hymenop- terous population a very definite response to tempera- ture, and not an emergence correlated with the length of time the organisms had spent in the ground as larvae or pupae. This is quite reasonable, as is already known, the time spent in the various stages is lengthened by cold, and the development is accelerated by warmth. ‘All this seems, in so far as these Hymenoptera are con- cerned, not to apply merely to a single species, but to indicate a law influencing all life under these conditions. (b) Relation of population to light and sunshine. The Hymenoptera are generally regarded as lovers of the light and sunshine, and especially is this true of the flower-frequenting bees and wasps; they are gen- erally to be seen foraging among the blossoms, and then indeed may we call them children of the sun. Here we are interested in the three phases: (a) the relation of the position of the nests to the sunlight, (b) the rela- 242, Trans. Acad. Sci. of St. Louis tion of the sun-loving parasites to the position of the nests of their hosts, and (c) the action of parasite to- ward host as the nest of the host is in the sunny or the. shady portion of the bank. Position of the clay bank in relation to the sunlight. We have seen in Fig. 2, how the clay bank was protected above by the porch and faced the eastern sun. Three clumps of rambler rose-bushes stood in front of the bank, one at the north end and two spread over the south half. These were densely covered with flowers and foliage during the summer.* Since two rose-bushes were in the southern half, the portion of the clay bank just behind them was much more shaded than the north- ern half. In this brilliantly illuminated area between the bushes the nests of 4. abrupta abounded; here they left their old burrows, an unmistakable record of the life which had been there. It was in this portion of the bank also that in 1917, the most of the contemporaries of the Anthophora congregated to build their nests. It would be better to say that here they emerged, and then re- — mained in almost the precise spot to nidify, but this in turn only means that their ancestors at some time ac tually chose this spot or congregated here, so the signi- ficance is the same. In order to realize the sharp con- trast in the nesting activities, between the sunny poT- tion of the bank and the shaded areas, Figs. 3 and 4 may be compared. Fig. 4 shows the northern half (the photograph was taken after a dry year, when few bees made turrets, and these few had fallen or had been *This picture was taken in October, when most of the foliage had fallen. The Ecology of a Sheltered Clay Bank 243 harvested for study), or that portion which got the max- imum amount of sunlight. This is in striking contrast to Fig. 3, the southern half, which enjoyed only a lit- tle filtered sunlight. One can readily see that the de- gree of illumination made a great difference in choice of site by the A. abrupta. The white-banded bees, En- techma taurea, the caterpillar-wasps, Ancistrocerus ful- vipes, and the Trypoxylon wasps did not give evidence of so direct a dependence upon this factor. I have often wondered just what is the correct explanation of this correlation of warmth to longevity. I have discovered in other studies that in organisms that take no food as adults (Saturniid moths) increased warmth tends to accelerate the activity and thus exhaust the vital energy so that the life of the organism is cut to a much shorter duration than it would have been at a lower temperature where activity is retarded. So shall we say that be- cause A. abrupta chooses the warmest place in the sun, her activity is intensified and her life soon spent? Or, on the contrary, shall we say that her habit of choosing the sunny situations is a fortunate adaptation in that it enables her to get through with the necessary duties of life in the short period of time allotted to her, and that probably only those individuals which do exercise this choice will be able to finish their life-work and leave progeny? It should be noted that all the other insects mentioned in this study have a much longer time for their duties than A. abrupta, since they either lived the entire season, or had more than one generation each season. In 1917, the white-banded bees greatly outnumbered the Anthophora. While a few of the nests of the former were in the face of the bank, and often took a somewhat 244 Trans. Acad. Sci. of St. Louis vertical position (Fig. 20E) most of them were hort- zontally resting on top of the bank, near the edge (Fig. 21E). All of the nests of this species enjoyed an abun- dance of the sunlight; those on top of the bank were made very near to the front edge. However, during the years that followed this species decreased in num- bers so that by 1920 it was barely holding its own, whereas A. abrupta increased amazingly, as described elsewhere. Another interesting factor enters into the story here, however; during the years under observa- tion the white-banded bees gradually extended their nest- ing area. In 1917 most of them were in among the An- thophora nests near the sunlight, either in the face of the bank or near the edge of the top (Figs. 20E and 21K) with merely the beginning of a settlement at the extreme south end and another at the north end. In 1918, they were almost all in these two end groups, and none among the Anthophora nests. In 1919 a few were at the south end, but most of them were on the top, farther back than previously, and in 1920 to 1922, all of the burrows found were far back on the bank and entirely out of the direct sunlight. In the meantime Anthophora abrupta was in- creasing in numbers, always building on the sunny face of the bank. Moreover they were gradually spreading over the southern portion of the bank, until by the end of the period the north half of the dark barren portion seen in Fig. 3 was abundantly inhabited; Fig. 22 is the same area as in Fig. 3. Not without cause, however; was this significant change; in the spring of 1918 the tenants of the house had cut the ramblers almost to the ground, so for the seasons immediately succeeding, that portion of the bank was also fully exposed to the sunlight. The Ecoloyy of a Sheltered Clay Bank 245 The parasites also love the sunlight. This point leads us at once into a maze of interrelations between sun- light, parasites and hosts. The very short life cycle of Anthophora abrupta eliminates from consideration prac- tically all of the Dipterous parasites, because by their early emergence they escape the parasites and their suc- ceeding generation is practically provided for before the most of the parasites are born. Hence Anthophora has to struggle against only the Chalcids, the cuckoo-bees and possibly one species of Diptera in order to main- tain her species. While these three parasites did evi- dently prey upon this Anthophora population to some extent, their success depended to a far greater extent upon the white-banded bees as hosts since the duration of the life of the latter was more than twice that of the Anthophora bees, and since the white-banded bees emerg- ing later in the summer were the possible hosts of more than three times as many species of parasites. It is little wonder then, that the species which was being rapidly exterminated left the sunshine and the enemies which swarmed in it, and sought the ill-lighted recesses of the bank. Of course, it is really a problem whether these bees actually left the sunshine and crept away, or whether the parasites had merely exterminated all those which persistently remained in the sunny areas. With nations of bees—as with nations of men—there is not place in the sun for all. The numerous and the mighty occupy the favored places, and any others that wish to live may occupy the remaining corners of the earth until such a time as the powerful wish to usurp the places in the shade also. Usually the only hope of the ultimate survival of such oppressed species depends upon their powers of adaptation. Thus run the histories of the sur- 246 Trans. Acad. Sci. of St. Louis vival of the fittest, the one proving its fitness by re- maining in the sunshine thereby accelerating its devel- opment, but curtailing its life, and the other proving its strength by continuing its existence in spite of tre- mendous opposition. One man’s meat is another man’s poison; one bee’s sunshine is another bee’s death. We eagerly await further developments in this com- petition, but at present we cannot forgo the temptation to speculate upon it. Will the white-banded bees, thus hard pressed, be finally exterminated? If so, what will be the effect upon Anthophora in return? Perhaps by that time they will have become so strong that, like the oak and the ivy, they will be able to carry on prosper- ously the work of the mining-bees and also maintain the various parasites without seriously feeling the strain. But again, if this happens, what will become of those later parasites which depended upon the white-banded bees, etc., those which, in fact, have killed the goose that laid the golden eggs? They will be compelled either to change their mode of living in some way, to migrate and perhaps perish in the attempt, or to quit living. If, how- ever, the parasitic Diptera adhere to their tendency to confine themselves to sunny areas, and if at the same time the white-banded bees can adapt themselves to their dim corners where they have taken refuge, there is a pos sibility that the white-banded bees may again increase and at some time regain prestige. Of course, these last Sentences are merely speculative, and yet it is by pre cisely such cat and mouse methods as these that nature lets her many species run on in constant and deadly competition. The Ecology of a Sheltered Clay Bank 247 (c) Relation to cold, cloudiness, darkness. Up with the early morning sun, at 6:30 in fine weather the mining-bees of all three species, the carpenter-bees and the Monobia mud wasps were all heartily at work. They began early and worked diligently while the sun shone and then relaxed soon after midday as soon as the bank was in the full shadow. I have said elsewhere that these insects know no calendar but the thermome- ter; again we might well say they know no clock but the sunlight. I hope some day to find out just what is these bees’ program of daily work when their nests are in the normal light, i. e. not shaded by artificial struc- tures during any part of the normal hours of sunlight. While the action described above was their usual pro- gram, they sometimes made exceptions to this habit. These digressions I have been unable satisfactorily to explain. Often during the early part of June the An- thophora bees were seen at work at 5:30 a. m., and con- tinued busy until sunset. Despite the fact that May 28 was cloudy, some 20 mothers were actively engaged in carrying in orange-colored pollen. On the other hand, When on June 16 the temperature dropped twenty de- grees, their activities were greatly lessened, and they did not venture out to resume their work until 9 o’clock on the 17th. Also after cold nights or rainy mornings and toward the end of the season, even when the sun did shine, often they did not come out early, but waited until the bank was thoroughly warmed up. Sometimes, when a run of cold days came in early autumn (in this detail I am speaking of the white-banded bees), they did not get out to work for two or three days; suddenly they Seemed to be seized by the impulse to work, and then 248 Trans. Acad. Sci. of St. Louis they would come out even on cold and gloomy days and work hard, striving to complete their task almost as if they felt in some way that time and life were ebbing fast. It will perhaps lead to a clearer understanding if we pass from these generalizations to the record of what actually happened after the rainy afternoon of August 29 and other cloudy and cold days. I merely give ex- cerpts from the original notes. ‘August 30. The rain fell until 2 p. m. yesterday, after that hour no insects were abroad. The night was rather cold and today it was 10 a. m. before the first insect appeared about the bank. A black cricket poked his head out of one of the bee-holes where he had spent the night. Soon several wasps, Tachysphex terminatus, were scraping away the soil about their nests. A few specimens each of Ancistrocerus fulvipes, Monobia qua- dridens and Xylocopa virginica, and one each of Try- poxylon clavatum, Chlorion auripes, and three white- banded bees, Entechnia taurea were at work at their va- rious duties of nidification: these were all observed ab 11:30 a. m. and all seemed unusually industrious, as if to make up for the time lost in yesterday’s rain. As it neared noon, the white-banded bees appeared in greater abundance, and the parasitic bee-fly, Argyromoeba tig- rina, made its appearance. That the sunshine was a fac- tor in bringing out the bees is evidenced by the fact that only from the nests that had been warmed by the di- rect sunlight had the bees crept out to work. The mothers from the nests in sunny sites were the first to be active in bringing loads of pollen, while those nests not so favored showed no external signs of life. I have previously stated that this species of bee, Entechnia The Ecology of a Sheltered Clay Bank 249 taurea, will burrow without regard to the light, that it chooses dark areas as well as well-lighted ones without any apparent detriment to the work, but here it is ap- parent how, especially after a cold night, the sun indi- rectly influenced the amount of work done by warming up the nest and routing the sleepers. After all we must admit, that despite the fact that these bees often built their nests back in the shadow of the porch, they seldom did work when the sun had passed beyond the zenith, leaving the bank more shaded. It is interesting to note that in the afternoon, after the deeper shadows fell upon the bank at about 2 p. m. the activities of several spe- cies were at their minimum. An analogous condition existed on dark or cloudy days. The insects behaved much as the population of a normal site would at twi- light. As the shadows lengthened, some spiders would become active, thus adding the suggestion of night- prowlers to the twilight setting. “‘September 4. While I have elsewhere recorded that as the sun warms up the bank, more and more of the Entechina bees come out of their holes and begin work, it is now my pleasant duty to record that this rule is not absolute, for today the behavior is reversed. Yes- terday they were working intensely at their digging; now it is dull and dreary after the heavy rain in the night, yet the bees have been busily at work carrying in pollen. In so far as I can see, this reversal of behavior can be due only to the fact that much rainy weather lately has caused a set-back in their work, and they be- &in to ‘realize’ that if ever they are going to get their domestic duties done before Gabriel sounds his trumpet, they will have to be up and doing despite weather con- ditions. i 250 Trans. Acad. Sci. of St. Louis ‘‘September 12. The last two or three nights have been very cold and today at 9:45 the only life about the bank is a lone Tachysphex terminatus, an unfortunate caterpillar in the jaws of an ant-lion, and about ten gray lizards. None of the white banded females are in evi- dence, but in the garden back of the house, eight males were this morning found huddled in the cups of as many wild bindweed blossoms where they had spent the night. The flowers had not closed, evidently because of the low temperature, and these male bees found lodging there all night, and even at ten o’clock in the forenoon, they were in a torpid and lethargic condition, heavily laden with dew. This discovery so late in the season also shows that they are not shorter-lived than the females as we have found for other wasps and bees. By noon the bank has warmed up perceptibly, and a few of the sun-loving insects are out; one Entechnia taurea hovers over her nest; one cuckoo-bee, Chrysis coeruleus lurks suspiciously about. At two o’clock the first carpenter- bee, Xylocopa virginica, and an Ancistrocerus fulvipes are diligently caring for their nests. It is 3 p. m. be- fore the first carpenter mud-wasp, Monobia quadridens, becomes active about the wood tunnels overhead, and even the Chalcid parasites, Monodontomerus sp. lin ger to this late date. These meager numbers of each species are no doubt indicative of the entire population at this time of the year. It is interesting, too, to note that despite the fact that most of the activities have heretofore ceased somewhere about two in the after- noon, we now find these remaining mothers working far into the late hours of the afternoon. The day is too short for them; they have their tasks and are intent upoD finishing them, and in such stress are not guided abso- The Ecology of a Sheltered Clay Bank 251 lutely and blindly by instinct, yet they do modify their behavior according to their needs. Whether the sun- shine is likewise warming the lizards, or whether the appearance of possible food excites them, I do not know, but at any event these vertebrates, heretofore sluggish, become animated and active at the same time as the re- mainder of the inhabitants. “‘September 23. Only a few stragglers representing some of the more important groups of the inhabitants, remain at this late date, and they are indeed few in number. It is evident that the cold of autumn is a fac tor in reducing their numbers, but we should like to determine precisely to what extent this is the deter- mining factor, or in how far the weather conditions directly influence their longevity. If the optimum con- ditions for their existence prevailed for several months longer, it might retard but would not abolish their mor- tality. In other words, to what extent is longevity in- herent in the organism, and how far is death due to some external calamity? “October 3. Today the most of the few individuals to be seen certainly give one the impression that they have outlived their allotted time—‘yet is their strength labor— and sorrow.’ Three survivors of EZ. taurea linger help- lessly about, as if they had been overlooked by death and did not know how to conduct themselves in the land of the living, where they no longer belong. Indeed no activity is now possible for them, because the flowers from which they have gathered pollen are long since withered. Two Ancistrocerus fulvipes mothers are feebly finishing the closure of their nests. Two or three Monobia mud-wasps are seen on the rafters above the bank, stunned and half dead with the cold. The only 252. Trans. Acad. Sci. of St. Louis inhabitant showing any semblance to normal activity is the grass-carrying wasp, Chlorion auripes. Four moth- ers have been busy all this dull, gray day carrying in blades of grass to plug up their nests in the old bur- rows of the carpenter-bees. All the creatures of the unit are now practically ready for their long winter sleep. Soon the bank, frozen or snow-covered, will look utterly barren and lifeless, yet within that mass of earth is life —potential life that in the warm days of spring will burst forth with renewed vigor, ready to battle again in sharp competition for the possession of a small portion of the earth which they may call home.”’ With the approach of cold weather the carpenter-bees crowd into their tunnels and quietly await death. This I suspected one spring when I opened a nest in a piece of wood which contained six adults, all dead. During eold days they often remain in the cells, crowded close together, and come out again when the day grows warmer. I have seen them come out for a little while as late as October 13. On another occasion in early September, I watched in vain for two days, for the oc cupants of a nest. Concluding at length that they had joined the heavenly choir, I proceeded to investigate with an ax. At the first blow, a loud chorus of buzzing greeted me, and then one by one twelve bees, each an- grier than the last, flew out of the tunnel. This startling experience made me wonder if it is not possible that part if not all of them, hibernate in this way instead of dying with the first cold of autumn. It seems to me that in mild winters or in the southern states they might easily hibernate thus. There is some evidence that the clearness of the light may be a factor in the homing behavior of the mining The Ecology of a Sheltered Clay Bank 253 bees. When they returned from the field on their reg- ular day trips, each tumbled precipitously into its own hole without exploring the region or seldom trying out any of the burrows before finding its own. (Figs. 1 and 4 show how similar and how close together the burrows were). But one evening between 7 and 7:30, when it was almost dark and bees were returning home, many hesitated and hovered about before the colony and even entered several holes before finding their own. Two factors may enter into this inability to find their hole, viz., the condition of weariness and the failing light. This seems to us a logical foregone conclusion, and it would be quite superfluous to mention it except for the fact that a certain school of investigators argue for a sixth sense to serve these Hymenoptera in their homing flight or the ability to return home as if by magic. (d) Relation to rain. All of the mining bees carried water for their excavat- ing. Rain helped them substantially by filling the wagon- ruts in the road and giving them a supply of water nearby. When these puddles dried up, they were obliged to go a long way for water, and in some cases they did without water or stretched a gulletful over so much earth in the burrow that the result was not plastie mud that could be shaped into turrets, but crumbly, slightly moist pellets that could only be kicked out of the hole. The bees did not leave their burrows during a rainy spell, and I have never yet seen a mining-bee return to the nest during a storm. I did, however, watch five Monobia mud-wasps come home during a heavy down- pour. They came in at intervals, all dripping wet and slow of flight, and crept into their holes. The 254 Trans. Acad. Sci. of St. Louts carpenter-bees nesting in the same place did not come home during the storm. They saved themselves a drench- ing, and after an hour or so, when the storm had ceased, they came flying blithely in. We have recorded else- where in these pages that the rain drops on the leaves near by were eagerly lapped up by 4. abrupta and used in their work of excavating, and I should not be sur- prised if drops of dew in the early morning were gath- ered for the same purpose. While we find evidence of a certain amount of de- struction of life by rainstorms, insects of a great variety beaten down by the rain, and their nests in the earth cut open or buried deeply by the erosion of the soil, yet the rain is not without its benefits also. It has been noticed that immediately after a rain following a long dry period, there was a surprising increase in the pop- ulation of burrowing Hymenoptera. For a time I east this aside as mere coincidence, until the evidence seemed too strong to be thus discarded. Why this sudden ap- pearance after the rain, and what is the direct action of the rain upon their emergence? There is no physiolog- ical connection, I am sure, but merely a physical relation. During a period of drought the earth had become ex- tremely hard; hence the emergence of these young ground-dwellers was rendered impossible. Perhaps they could wait for a time, but surely not for long. We have no way of knowing how many or what proportion of them perished in their futile attempts to break through the brick-like earth. The rain came and softened the earth, and lo! they appeared as if by magic. No doubt there was heavy elimination here, but in this case one cannot call it the survival of the fittest; it was only the survival of the lucky, and how do we know but the The Ecology of a Sheltered Clay Bank 255 lucky ones in this case may have been the least fit to per- petuate the race? In the activities of the turret-build- ing bees also, we find wet or dry weather an important factor in their nest building. In dry years, the turrets are almost entirely omitted and in the case of A. abrupta, when for the scarcity of water, much labor is put into hard digging, the number of young is diminished. (e) Relation of inhabitants to plant life. At first thought, one sees but little relation between the plant and the animal life of the bank. The vegeta- tion about the base of the bank, scanty as it was, har- bored numerous insects, such as small beetles, beetle larvae and caterpillars, which sometimes dropped into the ant lion pits. The grass-carrier wasp gathered grass for her nest; the mining bee, the carpenter bee and the Halictus went to the plants for nectar and pollen. The caterpillar wasps A. fulvipes visited the plants for their nectar and their caterpillars; the Trypoxylon and the Tachysphex wasps went to the vegetation for their spl- ders and grasshoppers respectively. As we have re- lated before, the flowers of the wild morning glory served so well as bachelor quarters for the males of the white- banded bee that the females were obliged to fight for even a chance to harvest the pollen from the flowers. The bees came in from the surrounding vegetation bring- ing pollen of various colors—yellow, orange, white, or light green. One often found against the side of the bank splashes of this pollen which had been lost in their homeward flight. For some time I suspected an intimate correlation between the dates of the persimmon blossoms and the emergence of the Anthophora bees. In the spring of 256 Trans. Acad. Scr. of St. Louis 1920, I watched to compare the dates. A large number of Anthophora abrupta pupae which had been brought into the laboratory early in April gave forth their young in May;* the males emerged from May 10 to 25 and the females only from May 22 to 25. On May 27, I visited Wickes and found both sexes abundant about the clay bank; this indicates that the time of emergence for the controls in the laboratory was about the same as those out-of-doors. Previously I had recorded the coincident dying off of these bees with the dropping off of the per- simmon blossoms, from which they gathered their pollen and food. On May 27, I visited the permisson tree near their colony, and found it laden with large buds, but no insects were about the tree. The next morning, sev- eral branches bore flowers; and lo! dozens of Antho- phora bees were already present, joyously busy about the new blossoms. While this may at first seem to be only coincidence, or to show that the bees had simply turned to any flowers that they could find, we should not be hasty in dubbing it mere chance until we have given due consideration to the fact that the benefits of the association of blossoms and bees may be mutual, and may be of as great impor- tance to the one as to the other, or until we have more data on these interrelations. (f) Death by violence, and natural death. To eat and to be eaten seems to be the law of the wild. In the interrelation of life in the clay bank, we have re- peatedly noticed how one insect falls prey to another. We seldom think of natural death among the insects, *Trans. Acad. Science St. Louis, 24 735, 1922. The Ecology of a Sheltered Clay Bank 257 but this, too, occurs, and often to a much greater extent than we usually suspect. During this study at the bank, I have often picked up dead and enfeebled insects. Not all the Chalcids fell prey to the spiders and ant lions; I have removed hundreds from the crevices where they had crept in to die. Many mining-bees were picked up dead or limp at the end of their season and many more were pulled out of their burrows. The cuckoo-bees were picked up dead, too—and what enemy could wound a cuckoo-bee in its thick armor? The blister-beetles, Hpi- cauta marginata, dropped dead from the plants near the base of the bank. Others found there whose death was not accounted for, were Harpalus dichrous Dejean (E. A. Schwarz), Tachysphex terminatus, Trypoaylon sp., Pseudagenia mellipes, Halictus pectinatus, Anthophora, and Xylocopa virginica. Thousands of dead Chalcid parasites were found in the burrows when they were examined during the winter. One often found newly dead female bees at the foot of the bank. This was very puzzling, until one day I actually saw where lay the trouble. A returning female A. abrupta found that during her absence another bee of the same species had appropriated her nest. A fight ensued, in which the usurper was thrown bodily to the ground and after a few twitches of the legs was dead. Other fights of a less serious nature were often observed. On one occasion I saw a female with part of her body protruding from the burrow. I pulled her out with the forceps, and found her tenaciously clinging with her jaws to a second female that had evidently intruded in the burrow. The next day also I saw the yellow, pollen- laden legs protruding from a bee which seemed to be in agony. I pulled it out with the forceps, and it, too, pug- 258 Trans. Acad. Sci. of St. Louis naciously clung to another bee, which had evidently taken advantage of the owner’s absence to usurp her home. (g) Relation to Soil conditions. Not all of the soil that came out of the burrows went into the making of the turrets. Much of it was kicked out and fell on the ground below. This was neither dust nor pellets, but characteristic minute bits or granules. A very considerable heap of these granules had accu- mulated in front of the center of the bank, where the greatest activity occurred. This strip of drift meas- ured 70 inches in length, 6 inches wide and from ¥% to 21% inches in depth. This of course gave the Tachysphes. terminatus wasps an excellent medium in which to make their nests, and afforded a material which made pit- making for the ant-lions a pleasure. It seems superfluous to mention the fact that, had the nature of the soil been very different from what it was, the colony of insects, especially the principal characters, the mining-bees, would never have been there, since 4 firm, clayey soil of this nature is quite essential to mas- onry of their kind; and too, if these pioneers had not thrown out a large mass of granular soil to the bottom of the bank, there would have been no ant lions, for these never dig pits in firm soil. (D)conc.upinG REMARKS, After following the great things and the little things in the lives of these tiny creatures, we may actually com- pare the whole to a play upon the stage. The environ ment corresponds to the stage; the dominant members correspond to the leading characters, the secondary spe The Ecology of a Sheltered Clay Bank 259 cies to the essential but subordinate characters. The individual animals adjust themselves to one another, and to the environment, as the personalities in the play adjust themselves to one another and to the conditions. ‘‘In both groups some individuals are dominant, some used and useful, some are tolerated, some pick up the crumbs, still others are predatory or parasitic, and all must be mutually adjusted to one another and to the environment.’’ On this little stage of the clay bank we have seen en- acted the tragedies and joys of life and love, birth and death of many creatures; their tolerance of, or indiffer- ence to one another, their craftiness in preying upon one another and their alertness in escaping persecution, the artisans at work, the visitors and the varying outcome of their visits—all these things and many others per- taining to the lives of a hundred insect species for a day, a season, or a succession of years. We have seen the pioneers blazing the trail, the rent- ers following hard by, and the parasites ever on hand keeping in check the otherwise rapidly increasing races ; we have seen some that are transitory and some that visit, prefer to remain, and may prove either a benefit or a burden to the group. And finally we have seen their behavior toward the elements of nature, to sunlight and cold, to darkness and to rain; all these little creatures would come and go, act and react, conscious or unconscious of one an- other and of the environment. All of the relations and interrelations which have controlled them, and all of the influence which they have wielded upon each other, to bring this social unit to the degree of development or balance in which we find it in the present years of the 260 Trans. Acad. Sci. of St. Louis study, must have occurred within a short period, for the existence of the clay bank covered less than a score of years. How many bee kingdoms or families have grown, lived and crumbled to oblivion here in that time, we have no way of knowing. Under our very eyes we have seen one grow to an unwieldy size at the expense of another that has almost crumbled to decay. It may seem naive for us to inquire into the mean- ing of all the little commonplace phenomena which we have observed here, and to search for many more not yet observed. A casual reader might say ‘‘Of course the organisms adjust themselves to each other and to their environment; why all this fuss about it?’’ Per- haps the same casual observer would have said to Isaac Newton ‘‘Of course the apple will fall; why ask questions about it??? And yet while we glibly call the phenomena commonplace, we must admit that we really know so little about the vital points in the lives of these organisms in this one microcosm that if we were to at- tempt to modify the balance of the population there we should bungle the delicate adjustments so that our ex- periments would not end anywhere near the point where we aimed; and until we can thus direct the controlling factors, we cannot boast an understanding of the sub- ject. ‘‘Every reflective biologist must know that no living being is self-sufficient, or would be what it is, or be at all, if it were not part of the natural world, although no truth is easier to lose sight of.’’ W. D. Brooks. Sie ss SeSnecease F< hee ies oe Fig. 1. Fig. 2. Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE Puiate XIV The work of An nthophora ey ag Not all hi bis excavated soil went ie turret building, but much of i s kicked out, and formed at the base (x) - medium for he habitel of ant- lion larve. , The clay bank under the porch. Trans. Acap. Sci. oF St. Lours, VoL. XXV PLATE XIV 264 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PLaTeE XV Fig. 3. a ey ‘ewer of a ba bank. Note the absence of burrows in contrast with figt Fig, 4, ce EN. of the clay bank. EUs. “VOU. AN “AD. Sct. OF S AC TRANS. 266 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PLaTE XVI Fig. 5. The mining-bee, Anthophora abrupta, (2 x natural size). Fig. 6. The mining-bee, Anthophora raui, (2 x natural size). Fig. 7. The white banded bee, Entechnia taurea, (2 x natural size). Trans. Acap. Sci. oF St. Lours, Vor. XXV PiatTe XVI 268 Trans. Acad. Sci.‘of St. Louis EXPLANATION OF PLATE PLate XVII 2 the Pig. 8: The “Ham and Bud” club. Note the clay bank under south-half of the porch. Trans. Acap. Sci, or St. Lours, Vou. XXV PuaTe XVII 9 -~ 70 Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE Puate XVIII Fig. 9. /The‘ turrets ia : AE eee E, Entechnia taurea; arrow, Anthophora ra Fig. 10. The u was place position.) turned turrets of Anthophora raui. (A wad of cotton d in the opening of each turret to show its up-turned Trans, Acap:. Sci. or Str, Louts, Vou. AXV PLATE XVIII 272 Trans. Acad. Sci. of St. Lowis EXPLANATION OF PLATE Puate XIX The Cells of Anthophora raui. The pup of Anthophora raui, A portion of the tunnel, and the orifice of the nest of Halictus zephyrus. Nymphs of Chortophaga_ viridfasciata. Ant-lion pits in the dust at the foot of the bank. Trans, Acap. Sci. or. Sr. Lours, Vou. XXV Pirate XIX Trans. Acad. Sci. of St. Louis EXPLANATION OF PLATE PLaTE XX A conspicuous inhabitant of the bank. Gray lizards inhabited the bank. Silver winged fly, Argyromoeba tigrina, (2 x natural size). The shedding skins of Argyromoeba tigrina. Trans. Acap. Sci. of Sr: Louis, Vout. XXV PLATE XX Trans. Acad. Sci. of St. Lowis EXPLANATION OF PLATE Pirate XXI Fig. 20, 4 portion of the face of the clay bank, showing turrets (E) Entechnia faurea, and (U) Anthophora abrupta. (Thi shows the relative size of the turrets of the two species. tu is horizontally made CE)' of penne taurea. (S) Pig. 21; he Pe upturned turrets of Anthophora ra Fig. 22. Three years later; the same area as Fi ig 3; the nests of “st abrupta are gradua ally spreading southward as the sunlig shaded _ portions. reaches the previously Louris, VoL. XXV 1. OF Sr TrANS. Acapb, Sc FLORIDA FLOWERS AND INSECTS. CHARLES RoBERTSON. Except when Orlando is specified, the observations recorded here were made at Inverness. Observations on blooming seasons did not extend beyond April. Flowers indicated as blooming the last of April probably bloomed longer. SIGNS, ABBREVIATIONS, ETC. ab = abundant; ¢ = collecting pollen; esp = collecting stray pollen; f = feeding on pollen; fq = frequent; fsp = feeding on stray pollen; gn= gnawing; in cop = in copula; lp = labial palps; Ma = non-social long-tongued bee flower; Mas = social long-tongued bee flower; Mi = non-social short-tongued bee flower; Mis = social short- tongued bee flower; n= non-pollinating; 0 = ornitho- philous; Pol = polytropic; R = red and all dark colors; S = sphingophilous; se = sucking nectar and collecting pollen; sf = sucking nectar and feeding on pollen; W = white; Y = yellow, greenish to orange; ¢ = male; ? = female; % — worker. The family ending ‘¢idae’’ is omit- ted, as Halict for Halictidae. The months are Ja, F, Mr, Ap, My, Jn, Jl, Au, S, O, N, D. When no other indications are used after a name, the insect is counted as sucking nectar legitimately and ef- fecting pollination. Dates after ‘‘visitors observed’’ in- dicate the period within which the observations were made. Signs in parentheses, like ‘¢Halictidae (2, ¢)”’ mean that all of the following Halictidae were females and all collecting pollen. Numbers in. parentheses, for example (6,65), refer to the titles and pages in the bibliography. In tables they are for species and visits or (277) 278 Trans. Acad. Sct. of St. Louts individuals. The plant names used here were taken from Small, ‘‘ Flora of the Southern United States.’’ GENERAL SYNONYMICAL LIST. The following is a list of all of the insects taken on flowers, with authors’ names which are not repeated in the special lists. Names which occur often enough to justify it are abbreviated as follows: Bs. = Boisduval, Cm. = Cramer, Cq. = Coquillett, Cr. = Cresson, Ed. = Edwards, F. = Fabricius, L. = Linnaeus, Lp. = Lepele- tier, Lw. = Loew, Mc. = Macquart, Mg. = Meigen, Rb. = Robertson, Rh. — Rohwer, Ss. = Saussure, Sm. = Smith, Tn.—Townsend, Wd.= Wiedemann, Wk. = Walker, Wl. = Williston. Names in parentheses are the . correct ones. BIRDS. Trochilidae: Trochilus colubris L. LONG-TONGUED BEES (30). Anthophoridae: Centris sp., Emphoropsis floridana Sm.; Ap.:'Apis mellifera L.; Bomb.: Bombias scutellaris Cr., B. separatus Cr., Siahais americanorum F., B. im- patiens Har.: : Clergine- Ceratina dupla Say; Emphor.: Melitoma tenes Say; Epeol.: Epeolus zonatus Sm.; Eucer.: Florilegus condignus Cr., Melissodes sp., M. var- iabilis Rb.; Megachil.: Ashmeadiella floridana Rb., type, Coelioxqs 8. dentata Say, C. sayi Rb., Diceratosmia con- junctoides Rb., type, Lithurgus ibboaie Sm., Megachile floridana Rb. ie, M. generosa Cr., M. lanuginosa Sm., M. mendica Cr. , Sarogaster georgicus Cr.; Nomad: Cen- trias rubicundus Oliv., Cephen fervidus Sm.; Stelid: Dianthidium curvatum Sen D. (Anthidiellum) notatum Latr., D. (A) perplexum fae Xylocop.: Xylocopa mi- cans Los .. X. virginica Drury. Florida Flowers and Insects 279 SHORT-TONGUED BEES (27). Andrenidae: Opandrena scutellaris Rb., type; Collet: Colletes sp., C. sp., C. americanus Cr., C. distinctus Sm., C. latitarsis Rb., C. thoracicus Sm.; Halic: ‘Agapostemon splendens Lp., Augochlora fulgida Sm., A. sumptuosa Sm., Chloralictus apopkensis Rb., type, C. ashmeadii Rb., type, C. floridanus Rb., type, C. longiceps Rb., type, C. nymphalis Sm., C. reticulatus Rb., type, C. tegularis Rb., Evylaeus nelumbonis Rb., E. pectoralis Sm., Odon- talictus capitosus Sm., Oxystoglossa sp., O. austrina Rb., type, O. matilda Rb., type, Panurg: Perdita obscurata Cr.; Prosopid: Prosopis flammipes Rb., type, P. floridana Rb., type, P. schwartzii Ckll. OTHER HYMENOPTERA (100) Bembicidae: Bembix spinolae Lp., Bicyrtes insidiatrix Handlirsch, B. ventralis Say, Microbembex monodonta Say; Cercer: Cerceris bicornuta Guerin, C. insolita Cr., C. austrina Fox, type, C. rufopicta Sm., C. verticalis Sm.; Chalcid: Leucospis affinig Say, L. robertsoni Crawford, type, L. slossonae Weld, Orasema sp. Spilochalcis flam- meola Cr.; Chrysid: Chrysis intricata Brulle, Hedychrum violaceum Brulle, Tetrachrysis venusta Cr.; Crabron: Anacrabro robertsoni Rh., type, Hypocrabro decemma- culatus Say, Solenius scaber Lp.; Ewmen: Eumenes fra- ternus Say, E. smithii Ss. Leionotus apopkensis Rb., type, L. arvensis Ss., L. australis Rb., type, L. bicornis Rb., type, L. bifurcus Rb., type, L. boscii Lp., L. floridanus Rb., type, L. foraminatus Ss., L. fulvipes Ss., L. fundati formis Rb., type, L. histrio Lp., L. histrionalis Rb., L. megaera Lp., L. molestus Ss., L. oculeus Rb., type, L. Saecularis Ss., L. turpis Ss., Monobia quadridens L., Odynerus erinnys Lp.; Larr.: Notogonidea argentata Bv., Tachysphex apicalis Fox, type, T. laevifrons Sm., Tachy- tes aurulentus F., T. breviventris Cr., T. duplicatus Rh., 280 Trans. Acad. Sci. of St. Louis type, T. pepticus Say var. floridanus Rh., type, T. rob- ertsoni Rh., type; Nysson: Nysson aequalis Patton; Oxybel: Oxybelus floridanus Rb., type, O. fulvipes Rb., type; Pemphredon: Psen maculipes Fox, type; Philanth: Philanthus carolinensis Banks, P. eurynome Fox, P. ven- tilabris F.; Pompil: Allocyphonyx maurus Cr., Anoplius illinoensis Rb., Aporinellus fasciatus Sm., Arachnoproc- tonus ferrugineus Say, Ceropales robinsonii Cr., Episy- ron biguttatus F., E. posterus Fox, type, Lophopompilus philadelphicus Lp., Planiceps calcaratus Fox, type, P. dubius Fox, type, P. minor Fox, type, Poecilopompilus navus Cr., Pompiloides americanus Bv., P. argenteus Cr., P. marginatus Say, P. subviolaceus Cr., P. tropicus L., Psammochares tenebrosus Cr., Sericopompilus plutonis Banks; Sphec.: Chalybion caeruleum L., Isodontia cin- erea Fern., I. exornata Fern., Priononyx thomae F., Psammophila violaceipennis Lp., Sphex gracilis Lp., 8. nigricans Dahlbom, S. pictipennis Walsh, S. procera Klug, 8. vulgaris Cr.; Scoli: Campsomeris plumipes Dru., C. quadrinotata F., Discolia nobilitata F., Elis floridanus Rh., type, E. propodialis Rh., type, E. robertsoni Rh., type; Tenthredin: Pseudosiobla robusta Kby.; Tpht: Tiphia floridana Rb., type, T. vulgaris Rb.; Vesp: Polis- tes americanus F., P. pallipes Lp., P. rubiginosus Lp, Vespula maculata L.; Vipion: Cardiocheiles nigriceps Viereck, Microbracon vernoniae Ash. DIPTERA (104). Agromyzidae: Milichia indecora Lw., M. robertsonii Cq., type, Milichiella arcuata Lw.; Anthomy: Bithoraco- chaeta leucoprocta Wd., Coenosia ovata Stein., C. sexno- tata Mg., Homalomyia prostrata Rossi, Limnophora narona Wk., Phorbia platura Mg.; Bibion: Dilophus orbatus Say; Bombyli: Anthrax lateralis Say, ‘A. lucifer F., Bombylius sp., Systoechus vugaris Lw., Systropus Florida Flowers and Insects 281 macer Lw., Toxophora amphitea Wk., T. virgata O. S.; Conop: Conops excisus Wd., Dalmannia vitiosa Cq., Physocephala sagittaria Say, Zodion nanellum Lw.; Ephydr: Notiphila bicolor Cr., type, N. carinata Lw., Ochthera exculpta Lw.; Musc: Chrysomyia macellaria F., Lucilia caesar L., L. sericata Mg., L. sylvarum Mg,, Musca domestica L., Synthesiomyia brasiliana B. & B.; Ortal: Euxesta notata Wd., Tephronota humilis Lw., Oscin: Chlorops unicolor Lw., Siphonella cinerea Lw.; Sapromyz: Pachycerina verticalis Lw.; Sarcophag: Heli- cobia helicis Tn., Metoposarcophaga pachyprocta Parker, Ravinia floridensis Ald., R. quadrisetosa Cq., Sarcophaga assidua Wk., S. bullata Parker, S. incerta Wk., S. utilis Ald., Sarcophagula occidua F.; Seps: Sepsis violacea Mg.; Stratiomy; Nemotelus glaber Lw., Odontomyia cincta Oliv., O. trivittata Say; Syrph: Allograpta obliqua Say, Baccha clavata F., B. tarchetius Wk., Ceria signi- fera Lw., Eristalis albiceps Me., E. dimidiatus Wd., HE. transversus Wd., Helophilus divisus Lw., H. similis Mc., Mallota cimbiciformis Flln., Mesogramma boscii Me., M. marginata Say, Microdon viridis Tn., type, Milesia vir- giniensis Drury, Orthoneura nitida Wd., Paragus tibialis F'illn., Psilota buceata Me., Spilomyia hamifera Lw., Syrphus americanus Wd., Tropidia albistylum Me., Volu- cella fasciata Mc., V. sexpunctata Lw., V. vesiculosa F., Xylota pigra F.; Taban: Tabanus sparus Whitney; Tachin: Archytas lateralis Mc., Atrophopalpus angusti- cornis Tn., type, Atrophopoda singularis Tn., Chaeto- glossa picticornis Tn., type, C. violae Tn., type, Cylin- dromyia nana Tn., Ennyomma globosa Tn., type, Gonia capitata De Geer, G. senilis Wl., Gymnoprosopa polita Tn., type, Hypostena floridensis Tn., type, H. vander- wulpi Tn., Masiphya brasiliana B. & B., Pachyophthal- mus floridensis Tn., Phasioclista metallica Tn., Phasiop- sis floridensis Tn., type, Phorocera edwardsii W1., Plagi- Zaz Trans. Acad. Sct. of St. Louts prospherysa parvipalpis Wulp, Polistomyia histrio Wk., Senotainia rubriventris Mc., S. trilineata Wulp, Siphocly- tia robertsonii Tn., type, Siphona geniculata De Geer, Siphophyto floridensis Tn., type, Siphosturmia rostrata Cq., Spallanzania hesperidarum W1., Sturmia distincta W4., Trichopoda pennipes F., Xanthomelana atripennis Say; Trypet: Neaspilota vernoniae Lw., Urellia solaris Lw. LEPIDOPTERA (53). Danaidae: Danaus archippus F., D. berenice Cm.; Hes- peri: Achalarus lycidas 8. & A., Amblyscirtes vialis Ed., Ancyloxypha numitor F., Atalopedes cempestris Bs., Atrytonopsis loammi Whitney, Cocceius pylades Scudder, Epargyreus tityrus F., Goniurus proteus L., Hylephila phylaeus Drury, Lerodea eufala Ed., Megistias fusca G. & R., Pamphila attalus Ed., Paratrytone aaroni Skinner, Polites baracoa Lucas, P. brettus Bs., P. cernes B. & L., Prenes ocola Ed., Thanaos brizo B. & L., T. juvenalis F., T. martialis Scudder, T. terentius S. & B., Thorybes daunus Cm.; Lycaen: ‘Atlides halesus Cm., Hemiargus hanno Stoll, Strymon cecrops F., S. melinus Hbn.; Nymphal: Dione vanillae L., Junonia coenia Hbn., Phy- ciodes tharos Drury, Vanessa atalanta L., V. virginiensis Drury ; Papilion: Papilio eresphontes Cm., P, glaucus L., P. marcellus Cm., P. philenor L., P. polydamas L., P- polyxenes F., P. troilus L.; Pier: Catopsilia eubule L., Eurema delia Cm., E. euterpe Menetries, E. nicippe Cm. Zerene caesonia Stoll; Rhiodin: Calephelis virginiensis Gray; Satyr: Cissia sosybius F.; Sesi: Synanthedon rubristigma Kellicott, S. geliformis Wk.; Arcti: Lerina incarnata Bs., Utetheisa bella L.; Noctu: Psychomorpha epimenis Drury; Syntom: Scepsis fulvicollis Hbn. COLEOPTERA (15). Buprestidae: Aemaeodera tubulus F.; Carab: Lebia Florida Flowers and Insects 283 pumila De jean; Cerambyc: Typocerus zebratus F.; Chry- somel: Donacia piscatrix Lac.; Cler: Trichodes apivo- rus Germar; Dermest: Attagenus piceus Oliv., Orphilus glabratus F.; Hlater: Cardophorus gagates Erichson; Lampyr: Chauliognathus marginatus F., Polemius limba- tus Lec.; Mordell: Mordella melaena Germar; Scarabae: Euphoria sepulchralis F., Trichius (Trichiotinus) affinis Gory, T. delta F., T. piger F. HEMIPTERA (8). Lygaeidae: Melanocoryphus facetus Say; Pentatom: Euschistus crassus Dallas, E. servus Say; Phymat: Phy- mata erosa L. var. guerini Lp. & S.; Pyrrhocer: Arhaphe carolina H. Schaeffer, Largus longulus Stal; Reduvi: Zelus cervicalis Stal, Z. bilobus Say. When a genus contains only one species, the specific name is omitted in the tables. Thus Apis = Apis melli- fera &. Insects were identified for me by J. M. ‘Aldrich, W. H. Ashmead, Nathan Banks, D. W. Coquillett, J. C. Craw- ford, E. T. Cresson, R. A. Cushman, W. J. Fox, G. H. French, C. A. Frost, A. B. Gahan, C. A. Hart, Samuel Henshaw, L. O. Howard, C. W. Johnson, Charles Liebeck, J. R. Malloch, H. M. Parshley, R. R. Parker, Theo. Per- gande, S. A. Rohwer, C. H. T. Townsend, P. R. Uhler, and S. W. Williston. Considerable difficulty has been found in getting the insects identified. I have been compelled to meddle with the business of the systematists and describe some bees and Eumenidae, or go without names. This has put me in the class of the jack-of-all-trades, giving the impres- sion that I was collecting and describing bees from any- where and everywhere, and I am offered specimens in exchange for local ones. 284 Trans. Acad. Sct. of St. Louis Sometimes insects sent by me for identification have been retained and identified for the persons who retained them, and it is often an accident that I find out the names. A chalcid was identified for me as Leucospis distinguen- dus and afterwards described as L. robertsont. A syrphid identified as Microdon coarctatus was afterwards de- scribed as M. viridis. The same specimen of Notiphila was identified for me by three different dipterologists as Dichaeta brevicauda, Notiphila carinata and N. bicolor. One of these defended this practice because the correct name had not been assigned. But that would not be fol- lowed by any one who distinguished what he knew from what he was guessing at. The best way to get species identified is to describe them yourself. One who will give you no aid in determin- ing them will go 1000 miles to suppress them as synonyms. Some author will describe species as if they differed only in size and color and deposit the types in a distant museum. You are not expected to read his descriptions but must examine the types to see what he described. His descriptions are ignored for many years. The spe- cies are described under other names, which become as- sociated with an extensive literature. Finally the types are examined and familiar names are upset. The only competent thing is for the systematists to clear up the old names at first. My experience is that the best time to get species iden- tified is when an author is working on the group, when he is competent to determine species or compare types. It is hard to strike authors at that time. I have had proba ble new species of Cerceridae lying in my collection while three authors, one after the other, were working on them. About the time you desire to send cercerids to one of these, he is working on may-flies or spiders, like a flit- Florida Flowers and Insects 285 ting schmetterling. I happened to strike Townsend when he was working on Tachinidae and got many of them identified, nine being types of new species. Descriptions of bees in which the sexes are separated as distinct species are worthless for identification. Nomenclature is synonymical, not binomial. No one has any authority to ignore priority*, but the historically correct name often has no relation to the literature. ACANTHACEAE. CaLopHanes Ostonerroia, Ma., R.—The stems rise 1 dm. or less, and bear three or four flowers open at a time. The corolla is 2 to 3 cm. long, expands 2m. The throat is 1 em. long and 7 to 8 mm. wide, narrowing rather strongly to the tube which is 1 em. long. The corolla is blue, but the throat below is marked with purple. The flowers are nototribe and proteran- drous. The throat is so wide that a bee which presses its thorax against the anthers readily effects pollina- tion. Butterflies can suck the nectar without much prob- ability of effecting pollination. Mr. 15-Ap. 29, 17 spe- cies, 100 individuals observed, Ap. 4-9. Lone-roneurp Bess (4:14)—Anthophor.: Centris ¢ 1; Bomb.: Bombus americanorum ? 1; Eucer. (2): Melis- sodes sp. 1, M. variabilis 11. Swort-roncuep Bezxs (3:5, ? )—Halict.: Agapostemon 1, Odontalictus 1, Oxy- Stoglossa austrina 3, n. Orner Hymenoptera (1:1)— “There are people who ignore history just as there are people who legislate to make more than one-half of one per cent of alcohol intoxicating instead of being appro- priated as food, to limit a patient’s needs to one pint in 10 days, and to propagate the fake biology which 1 satis ev: olution. ea fd 286 Trans. Acad. Sci. of St. Lowis Sphec.: Sphex procera 1. Drprera (1:12)—Bombyl.: Systoechus 12. Lepmoprera (8:68, n.)—Hespert.: Lero- dea 1, Pamphila 4, Polites baracoa 2, P. brettus 47; Nym- phal.: Junonia 1; Papilion.: Papilio philenor 3, P. poly- damas 3; Pier.: Eurema nicippe 7. Ruetuia Hummus, Ma., R.—Mr. 12-Ap. 28, 3 visitors observed, Mr. 14-26. LeprpopTera (3)—Hesperi.: Pamphila; Papilion.: Pap- ilio philenor, P. polydamas. ANONACEAE. Asmina Opovara, Mi, W.—Mr. 15-Ap. 26, 4 species and 9 individuals observed, Ap. 8-14. Conzorrera (3:8)—Cerambyc.: Typocerus 1; Scar- abae.: Trichius affinis, 1, T. piger 6. Hemrprera (1:1)— Pentatom.: Euschistus crassus 1. APOCYNACEAE, Amsonia Cruata, Ma., R.— The stem is terminated by a conspicuous panicle of pale bluish flowers. The corolla is tubular, with a border of 5 lobes. It meas ures 14 mm. across. The tube is slender and measures 7 mm. in length. In the upper part, occupied by the anthers and stigma, it is broader. The throat is densely bearded, shielding the pollen from intruders. The anthers surpass the stigma, so that self-pollina- tion is not likely. The pollen is discharged in a mass- The stigma is lateral and appears quite viscid. When es insect inserts its tongue the viscid matter may readily adhere to it and catch the pollen. Long-tongued bees commonly show a mass of pollen 0? their maxillary laminae, but butterflies never do. It is doubtful whether butterflies ever earry the pollen. Mr. 8- Florida Flowers and Insects 287 Ap. 26, visitors observed Mr, 18-Ap. 25. The following 5D species and 49 individuals carried pollen: Lone-toneuep Bzzs (5:49)—Anthophor.: Centris ¢ 2, 2; Bomb.: Bombus americanorum °?, 6; Eucer. (82): Florilegus 7, Melissodes sp. 18, M. variabilis 16. The following 28 species and 112 individuals carried no pollen: Lone-roncuep Brzs (7:19)—Anthophor.: Centris 3; Bomb.: Bombus americanorum 4; Eucer.: Florilegus 9, Melissodes sp. 1, M. variabilis 3; Megachil. ( ¢ ): Lith- urgus 1, Megachile generosa 2. Oraer HyMENoPTERA (3:5)—Sphec.: Isodontia cinerea 1; Scoli.: Camp- someris quadrinotata 1, Discolia 3. Drerera (2:12)— Bombyli.: Bombylius 1, Systoechus 11. Lepmoprera (15:75)—Dana.: Danaus berenice 1; Hesperi.: Hylephila 3, Megistias 1, Pamphila 6, Polites baracoa 41, P. brettus 1, P. cernes 9, Thorybes 3; Nymphal.: Dione 1, Junonia 1; Papilion.: Papilio cresphontes 1, P. marcellus 1, P. phile- nor 4, P. troilus 1; Pier.: Catopsilia 1. Cotnoprera (1:1) —Lampyr.: Chauliognathus 1. ASCLEPIADACEAE, Asciepias AmpLextcauuis, Mas, R.—Mr. 8-Ap. 26, 33 Species and 237 individuals observed, Mr. 19-Ap. 23. The following 17 species and 66 individuals carried Pollinia; h—on tarsal hairs, lp=on labial palps: Lone-roncurp Bess (1:3)—Megachil.: Megachile gene- rosa é 2, Ip, 3. SHort-roncuep Brees (2:2? )—Halict.: Augochlora sumtuosa h, 1, Oxystoglossa austrina Ip, 1. OrHer Hymenoprera (5:15)—Bembic.: Bembix h, 2, Microbembex h, 1; Ewmen.: Leionotus foraminatus, on claws, 1; Sphec.: Priononyx h, 10; Scoli.: Discolia h, 1. (1:1)—Bombyli.: Systoechus, on pulvilli 1, Lept- 288 Trans. Acad. Sct. of St. Louis poprera (8:45, h)—Hesperi.: Achalarus 1, Cocceius 1, Pamphila 11, Polites baracoa 23, P. brettus 2, P. cernes 2, Thorybes 4; Nymphal.: Junonia 1. The following 172 individuals, belonging to 27 species, carried no pollinia: Lone-toncuep Bess (2:32)—Megachil.: Coelioxys 8- dentata ¢, 1; Megachile generosa 31. SHonrtT-ToNGUED Brzs (3:8? )—Halict.: Chloralictus floridanus 2, C. nym- phalis 1, Oxystoglossa austrina 5. Orner HyMENOPTEBA (7:26)—Bembic,: Bembix 3, Microbembex 2; Eumen.: Leionotus arvensis 2; Larr.: Tachytes pepticus var. florl- danus 2, type; Sphec.: Priononyx 15; Scoli.: Campsome- ris plumipes 1, C. quadrinotata 1. Drerera (2:3)— Conop.: Zodion 2; Taban.: Tabanus 1, Leprooprera (11:99) —Dana,.: Danaus archippus 1, D. berenice 3; Hespert.: Cocceius 2, Pamphila 9, Polites baracoa 11, P. cernes 1, Thorybes 1; Nymphal.: Junonia 2; Papilion.: Papilio marcellus 1, P. philenor 1; Pier.: Zerene 1. CoLEoPTERA (1:2)—Lampyr.: Chauliognathus 2. Hemrprera (1:2)— Reduvi.: Zelus cervicalis 2. The Lepidoptera show the highest percentages “ gor cies and individuals both with and without pollinia. OF the species, 42.19%’ but of the individuals 31.2% show pol- linia. Of Other Hymenoptera individuals, 36.5% show pollinia. Only 11.1% of bee individuals show them. hs most common bee, Megachile generosa, showed 34 in duals, of which only 3 carried pollinia, and then only on the labial palps. Ascuepias Vertiomiata, Pol, W.—Mr. 4-17, one visitor observed at Orlando, Mr. 4. Orner Hymenoprera (1)—Tenthredin.: Pseudosiobla, pollinia on legs, tongue and face. Florida Flowers and Insects 289 AURANTIACEAE, Cirrus Aurantium, Ma, W.—Visitors observed Mr. 31. Lone-toneureD Bers (3)—Bomb.: Bombus americano- rum ¢, sc, ab; B. impatiens ? 3; Xylocop.: Xylocopa virginica 2. Orner Hymenoptera (3)—Larr.: Tachytes aurulentus ; Scoli.: Campsomeris quadrinotata fq; Vesp.: Vespula. Lepmoprera (3)—Nymphal.: Junonia; Pap- thon.: Papilio philenor; Pier.: Catopsilia. CACTACEAE, Opuntia Austnina, Ma., Y. Mr. 30-Ap. 29, 15 species, 51 individuals observed, Ap. 7-25. Lone-roneurp Bzzs (7:31)—Bomb.: Bombias scutella- ris %, 1; Eucer.: Melissodes variabilis 3, se, 13; Megachil.: ‘Ashmeadiella ¢, c, 1; Lithurgus ¢ 2, se, 13; megachile generosa ¢?, 1, M. lanuginosa ¢, 1, mendica ¢,1. SHort-roncuep Bzzs (2:4, 2, ¢)—Halict.: Agapostemon 3, Chloralictus numphalis 1. Lzpmop- Tera (4:5)—Hesperi.: Pamphila 1, Polites brettus 2; Papilion.: Papilio philenor 1; Pier.: Eurema nicippe 1. Couzoprera (2:11, f)—Mordell.: Mordella 1; Scarabae.: Trichius piger 10, in cop. CAPRIFOLIACEAE. Lowicrra Sempsrvinens, Ma. (0) R. Bros (1)—Trochil.: Trochilus ab. Lepiwoprera (1)—Papilion.: Papilio cresphontes. CISTACEAE, Hewianroemum Carournianum, Ma., Y. F. 2-Ap. 29. The stems rise a few inches and bear solitary, yel- low flowers an inch or more in width. The petals are obovate and are expanded horizontally. The numerous 290 | Trans. Acad. Sci. of St. Lows stamens are also horizontal, their dehiscent anthers fac- ing upwards. In this position there is no chance of the pollen touching the stigma. The large stigma is sessile. If a large bee loaded with pollen lands upon the flower it must dust the stigma before touching the anthers. Cross-pollination in this case is between different plants. If the insect does not bear pollen, it may readily effect self-pollination. The flower is visited exclusively for pollen, the petals, as in most pollen-flowers, soon falling. Females of small Halictidae, Chloralictus, Evylaeus and Oxystoglossa, were observed collecting pollen. COMPOSITAE. Brruanpirera Susacauuis, Mas, Y.—The heads are Single on the scapes and measure about one inch across. There are 8 to 10 broad, yellow ray-flowers which are pistillate. The lobes of the style are long, nearly erect, with stigmatic edges. The disc florets have nectar-bearing tubes 4 mm. deep. They are staminate, the pollen being carried out on a long style brush, as in Silphium. Ja. 10-Ap, 29, 14 visitors observed F. 27-Ap. 4: Lone-tonevep Bzzs (2)—Megachil.: Coelioxys sayi °5 Megachile lanuginosa ¢. SHort-roncuep Buss (5, ?)— Halict.: Augochlora sumptuosa, Chloralictus floridanus sc, Evylaeus nelumbonis, E. pectoralis, Odontalictus s¢. (2)—Bombyli.: Anthrax lucifer; Tachin.: Chae- toglossa picticornis. Leproprera (5)— Hesperi.: Polites baracoa, Thorybes; Nymphal.: Phyciodes; Pier.: Eurema nicippe; Rhiodin. : Calephelis. On ten days, Mr. 7-21, the following 300 individuals were taken on flowers: Lone-roneusp Bzzs (8)—Ap. 1, Epeol. 3; Megachil. 4. SHort-ronevep Brrs (216)—Halict, 216. Drermra (9)— Florida Flowers and Insects 291 Bombyli. 5, Syrph. 4. Lepmoprzra (67)—Arcti. 1, Hes- peri. 43, Nymphal. 12, Papilion. 7, Pier. 4. Odontalictus capitosus 2 , belonging to Halictidae, 8.3% of the families represented, showed 65.3% of the indivi- duals. Carpuvus Sp., Mas, R.—One visitor observed at Orlando, Mr. 21 Lone-roneuep Brzs (1)—Emphor.: Melitoma ¢ °. _Lycopzsm1a ApHytia, Mas, R.—Mr. 21-Ap. 29, 5 spe- cles and 15 individuals observed, Ap. 6-12. Lone-ronevep Bzzs (1:1)—Megachil.: Lithurgus ¢, 1. SHorT-toneurep Brzs (1:1, 2, ¢, n)—Halict.: Chloralictus floridanus. Lepmoprera (2:12)—Hesperi.: Pamphila 6, Polites brettus 6. Conzoprera (1:1, f, n)—Scarabae.: Trichius affinis 1. ERICACEAE, Batopenpron Arsoreum, Ma, W.—The trees rise sev- eral metres and are fairly white with flowers, which are arranged in small racemes. The corolla is 4 mm. wide at the thoat, so that the bodies of small insects are admitted. The lobes are slightly reflexed, enabling bees to cling to them more readily. The corolla is 7 mm. long, its tube 5mm. The filaments have a pair of awns, which when struck, cause a downpour of dry pollen. Nectar is secreted by an epigynous disc. To reach it a tongue must be thrust between the filaments. The latter are hairy, which helps to protect the nectar and to cause a pollen discharge. The flowers are homogamous. Cross-pollination is se- cured by the stigma being far in advance of the anthers. 292 Trans. Acad. Sci. of St. Louts In their first visits bees may effect self-pollination. In the absence of insects self-pollination may occur by the pollen falling upon the slightly projecting rim of the stigma. F’. 22-Ap. 24, 26 species and 68 individuals ob- served, Mr. 16, 23, Ap. 5. Lone-roneveD Bres (5:25)—Anthophor.: Empho- ropsis ¢, sc, 7; Bomb.: Bombias separatus %, se, 1, Bombus impatiens ? %,sc, 9; Xylocop.: Xylocopa micans @, 1, X. virginica ¢ ¢°, 7. SHort-roncueD Berss (3:7, 2 )—Halict. (sc): Agapostemon 5, Evylaeus pecto- ralis 1; Prosopid.: Prosopis flammipes 1, type. OTHER Hymenoptera (8:17)—Eumen.: Leionotus apopken- sis 1, L. megaera 2, Monobia 5; Scoli.: Campsomeris plumipes 2, C. quadrinotata 4; Vesp.: Polistes ameri- eanus 1, P. pallipes 1, P. rubiginosus 1. Drprera (2:3)—Musc.: Chrysomyia 1; Syrph.: Milesia 2. Leri- DoPTERA (8:16)—Hesperi.: Cocceius 1, Epargyreus 1, Goniurus 1; Thanaos juvenalis 1; Papilion.: Papilio cres- phontes 2, P. marcellus 3, P. philenor 6, P. troilus 1. Gayussacia Hrrrenza, Ma., W.—F. 17, 18, 9 species and 28 individuals were observed at Orlando. Lone-ronevep Bess (1:1 2 )—Bomb.: Bombias separa- tus 1. Swort-roncurp Brxs (1:5 ¢)—Halict.: Augo- chlora fulgida 5. Oraer HyMEnoprera (7 :22)—Eumen. (at holes, n): Eumenes smithii 1, Leionotus histrio 3, Odynerus 7, perforating; Scoli.: Campsomeris plumipes LG. ae ear ce! Q iat (at holes, n): Polistes ameri- canus 4, P. pallipe Preris Nirma, Ma., R.—The stems rise one OF two metres and the wie of the branches are crowded with pendulous flowers. The corolla is cylindraceous Florida Flowers and Insects 293 with a gibbous base. It is from 7 to 9 mm. long and is contracted in the throat to 1 or 2mm. The stamens and stigma are included. The filaments are crumpled and at the apex are two-awned, so that an entering tongue is certain to touch them and shake out the loose pollen. Blooms February 2-April 26, 1 visitor observed, February 19. Lepwoprera (1)—H espert.:; Thanaos juvenalis. Viccintum Nitipum, Ma., W.—The corolla is white, often with a reddish tinge, urceolate, 5 mm. long. The mouth is so constructed that all except the smaller insects are excluded, and for ordinary ones a tongue 5 mm. long is needed to obtain the nectar. The bee touches the exserted stigma before the included anthers. The anthers have scarious tips with terminal pores through which the light pollen is sifted down upon the visitor. Nectar is secreted by the thickened epigynous disc. Ja. 24- Ap. 23, 18 visitors observed, F. 16-M. 7. Lone-ronevep Brzs (2)—Anthophor.: Emphoropsis ¢ %, fq; Megachil.: Megachile lonuginosa ¢. SHorr- TONGUED Brxs (1, 2 )—Halict.: Chloralictus longiceps. Orner Hymenoprera (3)—Eumen.: Odynerus, perforat- ing, n; Scoli.: Campsomeris plumipes fq, C. quadrinotata fq. Lepworrmra (12)—Hesperi.: Cocceius, Polites bara- coa, Thanaos juvenalis, T. martialis, T. terentius, Thorybes; Nymphal.: Vanessa virginiensis; Papilion.: Papilio marcellus, P. philenor, P. troilus; Pieri.: Eurema nicippe, Zerene. Xouisma Ferrucmvza, Ma., W.—The ends of the branches are crowded with white flowers. 294 Trans. Acad. Sci. of St. Louis The corolla is nearly globular, 3 or 4 mm. long by 4 mm. wide at the base and 2 mm. wide at the 5-toothed throat. The 10 stamens and the style are included. The anthers have terminal pores and hold their tips against the style. A bee’s tongue disturbs the anthers and the pollen is sifted out. The pollen does not fall upon the stigma, though its edge may receive some. The pendulous position favors bees, which easily hang upon the flowers, but butterflies also effect cross-pollina- tion. The mouth of the corolla is so narrow that only bees with tongues about 4 mm. can drain the nectar. F. 2-Ap. 24, 6 visitors observed, F. 2-Ap. 5. Lonc-ronevep Bzes (3)—Anthophor.: Emphoropsis 3 2,8c, fq; Ap.: Apis; Bomb.: Bombus impatiens ? ¥- Oruer Hymeyorrera (1)—Scoli.: Campsomeris quadri- notata, Lxpipoprera (2)—Hesperi: Thanaos martialis; Papilion.: Papilio marcellus. ERIOCAULONACEAE. Exrocauton sp., Mis, W—F. 7-Ap. 26, 9 species and 29 individuals observed, F. 29. Lone-ronevep Bess (1)—Ceratin.: Ceratina ¢, 1. SHort-roneurp Bugs (4:23)—Halict. (2): Chloralictus ashmeadii 12, C. nymphalis 5, Evylaeus nelumbonis 1; Prosopid.: Prosopis schwartzii ¢ 9,5. Drerera (3:4)— Syrph.: Masogramma boscii 1, M. marginata 2, Tropidia 1. Lepwoprrera (1:1)—Nymphal.: Phyciodes 1. EUPHORBIACEAE. Cyiposcotus Strmutosus, Ma., W.—F. 8-Ap. 29, 2 visi- tors observed, Mr. 19, Ap. 4. Drerera (1)—Bombyli.: Anthrax lucifer. LepmoPTERA (1)—Hesperi.: Thorbyes. Individuals observed Mr. 16, 17, 19, 21, were 15 Hesperiidae. Florida Flowers and Insects 295 HYPERICACEAE. Ascyrum TrrrapetaLum, Ma., Y.—The four yellow petals are horizontal, the flower measuring an inch across. The numerous stamens dehisce in _ succes- sion. Since the stamens are turned more or less out- ward, the anthers do not touch the stigmas. An insect coming with pollen may readily effect pollination before touching the anthers. Otherwise, it may effect self-pol- lination. The flower seems to be visited only for pollen. F, 2-Ap. 26, 4 visitors observed, F. 18-Mr. 23. Lone-roneuzp Bes (1)—Megachil.: Megachile lanu- ginosa ?, c, and n, cutting petals. SHort-ronauzp BEEs (1)—Halict.: Chloralictus nymphalis °, ¢. Drerera (1) —Syrph.: Mesogramma marginata f. Conzoprera (1)— Buprest.: ‘Aemaeodera f. LEGUMINOSAE. Croratarta Rorunprrouia, Ma. Y. — The flowers are separated on the racemes, so that only a few on different stems are open at the same time. The insect must land on each flower separately, and the flowers must receive pollen from separate stems and commonly from separate plants. The flowers are yellow. The banner is nearly orbicu- lar, measuring 8 mm. across and is erect or reflexed. Its claw projects forward, and with the calyx serves to hide the nectar so that a tongue 3 mm. long is needed to drain it. At base the banner has cushions which serve the bee as a head-rest when forcing down the keel. The wings are short, of an orange red color, forming a path-finder, and are folded close around the keel, so that the latter dispenses entirely with its color. The base of the wings lie upon the base of the keel and their tips so closely envelope it that all are forced down together and, when 296 Trans. Acad. Sci. of St. Louts released, return to their former position. The keel is saccate at base, turns abruptly upward and ends in a slender beak. The stamens are monadelphous, a round opening in the tube at base above allowing access to the nectar. When the flower opens five oblong anthers have discharged their pollen and five rounded anthers are still indehiscent. In the bud the long anthers surpass the others. The five stamens with long anthers rise to the tip of the keel, shed their pollen, and then withdraw. The five stamens with round anthers then lengthen, their indehiscent an- thers serving to push the pollen up into the apex of the keel. Then they shed their own pollen where it will be easily carried out by the style-brush. The style bends abruptly upward, and has a fringe of hairs above and a circle of hairs about the tip. When the keel is depressed, the style pushes out a ribbon of pollen. If the keel be further depressed, the style itself appears carrying out more pollen upon its brush. Ja. 21- Ap. 29, 13 species, 90 individuals observed, F. 20-Ap. 20. Lone-ronevzp Bzxzs (9:71)—Anthophor.: Centris ?, 1; Megachil.: Ashmeadiella floridana ¢ 2, s¢, 13, type, Diceratosmia ¢, 1, Megachile generosa ¢ 2, 9, M. lanu- ginosa ¢ 2, sc, 20, M. mendica 2 , 7, Sarogaster ¢ %, 8¢ 9; Stehd. (8 %, sc): Dianthidium notatum 4, D. per- plexum 7. Dirrera (1:1, n)—Bombyli.: Systoechus 1. Lepmwoprera (3:18, n)—Hesperi.: Polites baracoa 16, Thanos brizo 1, Thorybes 1. Eryrorina Herpacea, Ma. (0), R.—Several stems rise 2 to 4 feet and bear scarlet flowers in racemes 1 to 2 feet long. The flowers are about 2 inches long: The calyx tube is 10 mm. with short lobes. The wings and keel are almost entirely included in the calyx tube, Florida Flowers and Insects 297 serving merely as a protection to the stamens on the lower side. The banner is folded longitudinally. At base it enfolds the stamens, wings and keel and is itself enfolded by the calyx tube. ‘Access to the nectar is on each side of the free filament. The stamens are of unequal length, so that the anthers form a pollen surface 12 mm. in length. They are enclosed in the infolded ban- ner, but their tips are slightly exposed below. Visitors must have a proboscis 14% inches or more to reach the nectar conveniently. The stigma stands among the an- thers and may easily receive their pollen, but the way neglected flowers fall seems to show that self-fertiliza- tion does not occur. It is evidently adapted to humming birds, and dusts their throats with pollen. F. 26-Ap. 29. Brrps (1)—Trochil.: Trochilus ab. Lupinus Dirrusus, Ma., R.—Often the plants grow in clusters a few feet across, which increases the conspicu- ousness of the flowers. The racemes are erect and densely flowered. The flower is 15 mm. long. The banner has its blade entirely exposed, since it has mainly an attractive func- tion. At base it runs forward and is creased so as to give a head-rest to bees depressing the keel. It is blue with a longitudinal white spot, which forms a path-finder. It is nearly orbicular, 12 mm. across. The wings form a colored envelope of the keel. They are coalescent at tip and move with the keel so as to keep it concealed. The keel is faleate, without color, and is bent abruptly upward, ending in a sharp beak, as in Crotalaria. The keel petals are coalescent above and below, leaving a small opening at the tip. 298 Trans. Acad. Sct. of St. Louis The absence of an opening in the stamen tube shows that nectar is wanting, and that the flowers are adapted to female nest-making bees, but they are visited by other insects. In the bud the oblong anthers dehisce, filling the tip of the keel with pollen. When the flower opens, the round anthers are still closed. They no doubt aid in forcing the pollen of the long anthers into the tip of the keel and then dehisce. The tip of the style is provided with a brush of hairs, which, when the keel is depressed, forces the pollen out of the tip of the keel. A long ribbon of pollen first ap- pears and then the tip of the style. The style returns to its former position when the keel is released, and then, when the keel is forced down, again sweeps out the pol- len. F. 7-Ap. 26, 29 species, 93 individuals observed, Mr. 5-Ap. 5. Lone-roneuzp Brzs (10:35)—Anthophor.: Emphorop- sis 2, se, 1; Ap.: Apis sc, 2; Bomb.: Bombus american- orum 2,1; Megachil.: Coelioxys octodentata ¢, 1, C. sayl é ?,5, Megachile floridana 3,1, type, M. generosa ¢ %, 7, M. lanuginosa 3 9, s¢.,7 , M. mendica ¢ 2, sc., 8, Saro- gaster 6,2. Sworr-roneuep Buss (2:9)—Halict.: Augo- chlora sumptuosa 2°, ¢., 2, Chloralictus floridanus 2, fsp-, 7, type. Leproprera (17:49, n.)—Hesperi.: Achalarus 1, Cocceius 2, Hylephila 1, Lerodea 6, Megistias 1, Pamphila 3, Polites baracoa 2, P. brettus 7, Thanaos juvenalis 1, T. martialis 1, Thorybes 13; Lycaen.: Strymon melinus 1; Nymphal.: Junonia 2, Vanessa virginiensis 1; Papilion.: Papilio philenor 5, P. troilus 1; Pier.: Eurema nicippe 1. T1um Inroysum, Ma., Y.—The stems are prostrate, the short, dense-flowered racemes only turning up from the ground. The flower is pale yellow and measures 11 mm. The calyx tube measures 3 or 4 mm. Florida Flowers and Insects 299 The banner runs forward for 5 mm., its claw, as well as the claws of the other petals, being held by the calyx tube. To reach the nectar a tongue 4 or 5 mm. long is needed. The banner then bends upward, being con- cave in front and strongly reflexed on the sides. As it appears in front, it measures 7 mm. high and 4 mm. wide. The folds render it quite rigid, so that it resists upward pressure. The wings are free, lying close to the keel, their tips being bent to the right. This requires the bee to land on the right side and, as in 7. obcordatum, pollen contact is limited mainly to the left cheek. This slight disposi- tion of the wings makes the flower pleurotribe. A process in each wing fits in a pit in the keel, requiring the parts to move together. On the base of each wing is a hook-like process which clasps the stamen tube and pre- vents the wings and keel from being thrust aside. The keel petals are united. When the keel is de- pressed, all of the stamens are exposed. When freed, the keel again covers them. The stigma receives pollen from the bee before the anthers touch it. Self-pollination is hardly probable. Access to the nectar is on each side of the free stamen. F. 19-Ap. 16, 2 visitors observed, F. 25. Lermorrera (2, n.)—Hesperi.: ‘Amblyscirtes, Polites baracoa. Trum Oscorpatum, Ma., R.—It grows flat on the ground, and bears short racemes of pale-bluish flowers, the tip of the keel being of a darker blue. The flower is 10 mm. long, the banner 5 mm. wide. The banner runs forward for 6 mm., when it turns obliquely upward. At base the banner clasps the other petals and is itself 300 Trans. Acad. Sct. of St. Lowis clasped by the calyx, so that it resists the entrance of small insects and limits the visitors to those having a tongue 5 or 6 mm. long. On the base of the banner is a large pale spot crossed by purple lines which run in- wards, forming pathfinders. The keel petals are united. A process in each wing fits in a depression in the keel, so that keel and wings are depressed together. Both wings turn to the right, which requires the bee to land on the right. The bees taken on the flowers had pollen on the left cheek, so that the flower is pleurotribe. When the keel is forced down, the stigma and after it the dehiscent anthers strike the bee, the stigma thus receiv- ing pollen from other flowers. Self-pollination can hardly occur if the flowers remains undisturbed. F. 16- ‘Ap. 23, 5 species and 19 individuals observed, F'.17-29. Lone-toncugp Brzs (2:13, ¢ )—Megachil.: Diceratos- mia 4, Megachile lanuginosa 9. Lxepmorrera (3:6, n.)— Hesperi.: Amblyscirtes 3, Polites baracoa 1, Thorybes 2. Viora AcutiForta, Ma., W.—The flower is 9 mm. long, the abcordate banner 6 mm. wide. The banner is pale bluish, marked in the middle with purplish streaks, which, with the purple tip of the keel, form pathfinders. Elsewhere the flower is white. The banner has a broad base, enclosing the other petals. It runs forward for 6 mm., thus limiting access to the nectar. The wings lie ulonseido the keel, hiding it from view, their tips far surpassing it. They are united with the keel and move with it. The keel is short (6 mm.), its petals united. Access to the nectar is on each side of the free filament. When the keel is de- pressed the anthers are exposed, but most of the pollen is deposited on the style-brush. The style turns up and Florida Flowers and Insects 301 is bearded near the tip. Mr. 9-Ap. 26, 4 visitors ob- served, Mr. 11-Ap. 3. Lone-toneurp Bers (1)—Anthophor.: Emphoropsis 2. Lepworrera (3, n.)—Hesperi.: Polites baracoa; Nym- phal.: Phyciodes; Pier.: Eurema euterpe. LENTIBULARIACEAE, Prinevicuna Exatior, Ma., R.—Mr. 16-Ap. 3, 2 visitors observed at Orlando, Mr. 16. Lone-ronevrp Brzs (2, 2, ¢., reversing)—Megachil.; Megachile lanuginosa, M. mendica. Pincurcuta Lurea, Ma., Y.—F. 19-Mr. 24, 2 visitors ob- served, F. 19. SHort-toncuep Brzs (2, ?)—Halict.: Agapostemon, Odontalictus. Pinevicuta Puma, Ma., W.—Each plant commonly bears a single scape. Only one flower is open at a time, so that cross-pollination between distinct plants is usual. The corolla varies from white to bluish-purple. The white form has the tube yellow within, 4 or 5 reddish Streaks on the upper wall forming pathfinders. The pur- plish form has the tube darker purple within and the Same pathfinders. The corolla is 10 to 16 mm. long and 7 to 10 mm. wide. The tube measures from 5 to 7 mm., the spur 4 to 5 m., so that a tongue 9 to 12 mm. long is needed to drain the spur. The tube of the corolla is only 2 or 3 mm. wide. The stigma is just above the mouth of the spur, where the proboscis of the visitor is certain to strike it. The lower wall of the tube is folded inward longitudinally and is covered with glandular hairs, requiring the in- 302 Trans. Acad. Sci. of St. Louis sect’s tongue to pass nearer the upper wall and so in contact with the stigma. The stigma is a valvular flap, turned backward over the two anthers. When a but- terfly withdraws its tongue the stigma is turned forward and the tongue comes in contact with the anthers. The flower is nearly erect and its border almost hori- zontal. It remains open all day. It may also be visited by night-flying lepidoptera. J. 24-Ap. 5, 10 visitors ob- served, F. 18-23. SHort-roncurp Brzs (1)—Halict.: Evylaeus pectoralis @. Lepmwoprrera (9)—Hesperi.: Amblyscirtes, Lerodea, Megistias, Polites baracoa, Thanaos juvenalis ; Nymphal.: Phyciodes; Papilion.: Papilio philenor; Pier.: Eurema nicippe, Zerene. Urrrcunara Inriara, Ma., Y.—The scape is supported on the water by a whorl of leaves and bears sev- eral flowers. The corolla is 18 mm. long and 9 to 12 mm. wide. The upper lip is 11 mm. long and 9 mm. wide. In its basal concavity is situated the stigma and two stamens. The lower lip has a palate which rises 5 mm. above its base and shuts against the upper lip, conceal- ing stamens and pistil and closing the spur. The flower is yellow, some reddish streaks on the palate forming path-finders. Below, the lower lip is three-lobed and lies in contact with the spur. The spur is saceate, notched at the tip, has a large mouth and is 8 mm. long. The valve-like stigma lies upo? and conceals the anthers. Later it rises so as to expose them. Nectar is secreted by the spur. The flowers Te main open a few days. Cross-pollination results from insect visits and self-pollination can hardly occur. Only one or two flowers on the scape are open at a time. The flower is evidently adapted to bees which know how to Florida Flowers and Insects 803 depress the palate. Butterflies might steal nectar with- out transferring pollen. Plants were found rising from the bottom in 91% feet of water in Tsala Apopka Lake. The flowers sometimes open while they are still two or three feet below the sur- face. After the flowers reach the surface the stems sep- arate from the ground but remain anchored by surround- ing plants. Ja. 7-Ap. 20, 5 visitors observed, F. 19. SHORT-TONGUED Bezs (3, 2 )—Halict.: Chloralictus reti- culatus, Evylaeus nelumbonis; Prosopid.: Prosopis schwartzii. Lxepmoprera (2, n.)—Hesperi.: Thorybes; Pier.: Catopsilia. Urricutarra Susutata, Mi., Y.—The slender scape rises 3 or 4 inches and bears two or three flowers, only one of which is open at a time; so that pollination is between dis- tinct plants. Sometimes the plants form thin patches quite attractive to insects. The flower is 6 to 11 mm. long, the lower lip 5 to 8 mm. wide. In a general way it resembles U. inflata. The flower is entirely yellow. The spur measures 5 to 7 mm. The lower lip forms a convenient landing place, and the weight of the insect opens the spur. When insect visits fail, self-pollination probably oc- curs from the stigma curling up so as to touch the pollen. Ja. 11-Ap. 4, 2 visits observed, F. 28, Mr. 2. Lermoprera (2, n.) — Hesperi.: Polites baracoa, Thanaos terentius. MALVACEAE, Sma Ruomerroua, Mi., W.—Ja. 11-Ap. 28, 1 visitor ob- served, Mr. 22. Lepmorrera (1)—Hesperi.: Polites baracoa. 304 Trans. Acad. Sci. of St. Louis NYMPHAEACEAE. Castauia Renirormis, Mi., W.—F. 17-Ap. 20, 3 visitors observed at Orlando, Mr. 18. Ap. 2. Suort-ronevep Bzzs (2)—Halict.: Agapostemon ?, ¢.; Prosopid.: Prosopis schwartzii ¢ ?, f. Drprera (1)— Ephydr.: Notiphila bicolor fq., type. Little plants, growing in shallow water, with pedun- cles 9-19 dm. long, and petioles 18-25 dm., probably be- long to a different species. See Nymphaea odorata (3,125). One visitor observed at Orlando, F’. 24. Suort-roncuzep Bees (1)—Halict.: Evylaeus nelum- bonis ?,¢. Nympuaza ‘Apvena, Mi., Y. (3,123)—Ja. 11-Ap. 26, 6 visitors observed at Orlando, F. 18-Ap. 11. SHort-roncuep Bees (3)—Halict. (2, se.) : Agaposte- mon, Evylaeus nelumbonis ab.; Prosopid.: Prosopis schwartzii . Drerzra (2)—Ephydr.: Notiphila carinata; Syrph.: Helophilus divisus. Cotzoprera (1)—Chryso- mel.: Donacia f., gn., in cop., ab. OLEACEAE. OsmaytHus Americana, Mis., W.—The ends of the branches are full of flowers. The corolla with its 4 broad, revolute lobes measures 5 mm. across. The tube is 2 or 3 mm. long and 2 mm. wide in the throat, so that the nectar is quite accessible. F. 22-Mr. 31, 5 visitors observed, F’. 22, 26. Lone-roncuEp Brgs (2)—Anthophor.: Emphoropsis ¢ j Xylocop.: Xylocopa virginica ¢. Orner HyMENoPTEBA (1)—Vesp.: Polistes rubiginosus. LepmopTERA (2)—Ly- caen.: Strymon cecrops; Papilion.: Papilio marcellus. Florida Flowers and Insects 305 ORCHIDACEAE, Isiprum Gracriz, Ma., W., Spiranthes gracilis in 7, 52— F’. 17-Mr, 16, 1 visitor observed at Orlando, F. 17. Lone-roncurp Bess (1)—Megachil.: Megachile lanu- ginosa ¢, pollinia on maxillary laminae. Lrmoporum Sp., Ma., R., Calopogon pulchellus in 1—F. 21-Mr. 16, 9 visitors observed at Orlando. Lone-roneurp Bres (2, 2 )—Bomb.: Bombias separa- tus n.; Megachil.: Megachile lanuginosa with pollinia. SHort-roncurp Bzzs (6, ? )—Halict.: Augochlora fulgida with pollinia, A. sumptuosa with pollinia, Chloralictus ashmeadii n., Evylaeus nelumbonis n., Odontalictus n., Oxystoglossa sp., n. Orner Hymenoprers (1)—Eumen.: Leionotus oculeus n., type. POLYGALACEAE, Potyeata Nana, Ma., Y.—The flowers grow in close spikes. The early ones are sessile and their flowers ap- pear to be habitually self-fertilized. The later spikes are borne on stems two or three inches high. The flowers are pale yellow, the tip of the keel more orange-yellow. The keel bears a crest of 8 slender lobes. Pollen is de- posited on a terminal tuft on the style. The stigma is lateral. Nothing seems to prevent pollen from the stamens being carried down to the stigma, though in- Sects will no doubt effect cross-pollination. A probos- cis 4 mm. long can easily exhaust the nectar. Ja. 26- Ap. 24, 1 visitor observed, Mr. 2. Lone-ronevep Bzzs (1)—Megachil.: Megachile lanu- ginosa 3. 306 Trans. Acad. Sci. of St. Louis Ponyeata Ponyeama, Ma., R.—Mr. 22-Ap. 29, 3 visitors observed, Ap. 20. Lone-roneueD Brrs (1)—Stelid.: Dianthidium per- plexum ?. SHort-ToneveD Begs (1)—Halict.: Oxystoglossa aus- trina ?. Lepipoprera (1)—Hespert.: Pamphila. POLYGONACEAE, THYsANELLA Fiwperata, Mis., W.—Ninety-six species, 5999 individuals observed, N. 11-21. The Lower Hymenoptera are 48.9% of the species and 42.5% of the individuals, while the bees are 17.7% of the species and 40.5% of the individuals. Odontalictus capitosus, 1% of the species, showed 17.8% of the individuals. Lone-roncurp Brzs (6:87)—Ap.: Apis sc., 78; Bomb.: Bombus impatiens ¢, 3; Megachil.: Coelioxys sayi ¢, 1, Megachile lanuginosa ¢ ?,3,M. mendica ¢,1; Xylocop.: Xylocopa virginica ?, 1. SHort-ronevep Brzs (11:156)—Collet. (2): Colletes sp. se, 10, C. americanus 1; Halict.: Agaposte- mon ¢, 12, Augochlora fulgida é, 1, A. sumptuosa $, 1, Chloralictus floridanus 4 ¢, sc., 12, C. nymphalis 2, 5, C. reticulatus ¢ 2,3, Evylaeus pectoralis 2, 1, Odontalictus é 2,8¢., 107, in cop., Oxystoglossa austrina ¢, 3. Orner Hymenorrmra (48:255)—Bembic.: Bembix 1, Bicyrtes insidiatrix 3, B. ventralis 4; Cercer:: Cerceris bicornuta 1, C. insolita 4, C. verticalis 1; Chalcid.: Leucospis affinis 3, L. slossonae 3; Crabron.: Anacrabro robertsoni 1, type, Solenius 1; Eumen.: Kumenes fraternus 5, E. smithii 3, Leionotus australis 1, L. bifureus 1, L. fulvipes 1, L. histrio 3, L. molestus 1, L. oculeus 2, L. saecularis 8, L. turpis 1, Odynerus 21; ‘Florida Flowers and Insects 307 Larr.: Tachysphex apicalis 2, T. laevifrons 1, Tachytes breviventris 1, T. duplicatus 1, type, T. robertsoni 1, type; Nysson.: Nysson 3; Philanth.: Philanthus ecaro- linensis 27, P. eurynome 1, P. ventilabris 8; Pompil.: ‘Allocyphonyx 48, Episyron posterus 1, Pompiloides marginatus 1, P. tropicus 4, Sericopompilus 1; Sphec.: Priononyx 11, Sphex gracilis 3, S. nigricans 1, 8. pic tipennis 1, 8. procera 6, S. vulgaris 1; Scoli.: Camp- someris plumipes 2, Elis floridanus 36, type, E. propo- dealis 6, type, E. robertsoni 3, type; Vesp.: Polistes amer- icanus 14, P. rubiginosus 1; Vigion.: Cardiocheiles 1. Diprera (13:34)—Bombyli.: ‘Anthrax lateralis 1, A. lucifer 1, Systropus 2; Conop.: Conops 1; Sarcophag.: Helicobia 1; Syrph.: Baccha clavata 6, Orthoneura 1, Volucella fasciata 6; Tachin.: Archytas 5, Gonia capi- tata 2, Phasiopsis 1, Siphosturmia 2, Trichopoda 5. Lerioprera (16:57)—Hesperi.: Goniurus 4, Hylephila 3, Lerodea 1, Polites baracoa 11, P. brettus 2; Lycaen.: Hemiargus 21, Strymon cecrops 1, S. melinus 5; Nym- phal.: Junonia 1, Phyciodes 1; Pier.: Catopsilia 1, Eure- ma delia 2, E. nicippe 1, Zerene 1; Satyr.: Cissia 1; Syntom.: Seepsis 1. Cotgorrera (2:9)—Lampyr.: Chauliognathus 1; Scara- bae.: Trichius delta 8. Hemiprera (1:1)—Phymat.: Phymata 1, n. RHAMNACEAE, Czanoruus MicropHyiuvs, Pol., W.—Ja. 24-Ap. 14, 178 Species and 971 individuals observed Mr. 1-31. The Lower Hymenoptera show 34% of the species and 42.2% of the individuals; the flies 33.7 and 29.5; the bees 14 and 17. The flowers are pretty decidedly sphecopoly- tropic. The Diptera and Lepidoptera show a decline in 308 Trans. Acad. Sct. of St. Louts percentage of individuals. The list yields types of 25 new species. Lone-roncurep Brzs (9:32)—Epeol.: Epeolus ¢ 2, 3; Megachil.: Coelioxys 8-dentata ¢, 2, C. sayi # 2, 9, Megachile generosa 3, 3, M. lanuginosa ¢, 8, M. men- dica ¢, 3; Nomad.: Cephen 2, 2; Stelid.: Dianthidium eurvatum ¢,1, D. perplexum ¢, 1. SHort-toncuep Bess (16:134)—Collet. (¢ ¢%): Col- letes distinctus 6, C. thoracicus 40; Halict. (¢): Aga- postemon 1, Augochlora fulgida 2, A. sumptuosa S8¢., 21, Chloralictus apopkensis 1, C. ashmeadii 2, C. flori- danus 27, C. longiceps 1, C. nymphalis se., 20, C. reti- culatus 1, C. tegularis 1, Evylaeus pectoralis 1, Odon- talictus 2, Oxystoglossa austrina 1, type; Prosopid.: Prosopis floridana 7, type. Orner Hymenorrera (60:410) — Cercer.: Cerceris austrina 47, type, O. rufopinta 18, C. verticalis 8; Chalcid.: Leucospis affinis 5, L. robertsoni 9, type, Ora- sema 2, Spilochalcis 1; Chrysid.: Chrysis 2, Hedychrum 6, Tetrachrysis 6; Crabron.: Hypocrabro 5, Solenius 28; Eumen.: Eumenes fraternus 8, E. smithii 7, Leionotus apopkensis 3, type, L. australis 9, type, L. bicornis 9, type, L. bifureus 7, type, L. boscii 9, L. foraminatus 7, L. fulvipes 1, L. fundatiformis 1, type, L. histrio 9, L. histrionalis 9, L. megaera 1, L. molestus 1, L. oculeus 33, L. saecularis 41, L. turpis 11, Monobia 2, Odynerus 6; Larr.: Tachysphex apicalis 2, type, T. laevifrons 6, Tachytes aurulentus 4; Oxybel.: Oxybelus floridanus 2, type; Pemphredon.: Psen maculipes 2, type; Philanth.: Philanthus eurynome 22; Pompil.: Allocyphonyx 2; Aporinellus 1, Arachnoproctonus 1, Ceropales 1, Episy- ron biguttatus 9, E. posterus 5, type, Planiceps caleara- tus 3, type, P. dubius 1, type, P. minor 1, type, Poecilo- pompilus 2, Pompiloides argenteus 1, P. marginatus 4, Florida Flowers and Insects 309 P. subviolaceus 5, Psammochares 1; Scoli.: Campsomeris plumipes 8, C. quadrinotata 1; Sphec.: Isodontia exor- nata 2, Psammophila 1, Sphex vulgaris 8; Tiphi.: Tiphia floridana 4, type; Vesp.: Polistes americanus 2, P. ru- biginosus 1; Vipion.: Microbracon 1. Diptera (60:287)—Agromyz.: Milichia indecora 21, M. robertsonii 2, type, Milichiella 4; Anthomy.: Bithor- acochaeta 1; Bibion.: Dilophus 1; Bombyli.: Anthrax lateralis 21, A. lucifer 1, Bombylius 17, Toxophora am- phitea 11, T. virgata 2; Conop.: Conops 4, Dalmannia 1; Ephydr.: Ochthera 1, pred.; Musc.: Chrysomyia 2, Syn- thesiomyia 1; Sapromyz.: Pachycerina 2; Sarcophag.: Helicobia 9, Metoposarcophaga 1, Ravinia floridensis 1, R. quadrisetosa 15, Sarcophaga assidua 1, S. bullata 1, S. incerta 1, 8. utilis 1; Syrph.: Allograpta 2, Baccha tarchetius 1, Mesogramma boscii 1, M. marginata 5, Orthoneura 2, Paragus 2, Spilomyia 1, Volucella fasciata 6, V. vesiculosa 1, Xylota 2; Taban.: Tabanus 4; Tachin.: Archytas 3, Atrophopalpus angusticornis 2, type, Atro- phopoda 1, Chaetoglossa picticornis 7, type, C. violae 5, Cylindromyia 3, Ennyomma globosa 4, type, Gonia sen- ilis 4, Gymnoprosopa polita 36, type, Hypostena flori- densis 6, type, Masiphya 1, Pachyophthalmus floridensis 6, type, Phasoclista 1, Phorocera 1, Plagiprospherysa 6, Polistomyia 1, Senotainia rubriventris 11, S. trilineata 2, Siphoclytia robertsonii 6, type, Siphona 13, Siphophyto floridensis 10, type, Spallanzania 2, Sturmia 1; Trypet.: Neaspilota 2, Urellia 4. Lerwoprera (21:28)—Arcti.: Utetheisa 1; Hesperi.: Achalarus 1, Amblyscirtes 1, Cocceius 1, Lerodea 2, Megistias 1, Pamphila 2, Polites baracoa 1, Thanaos juvenalis 1, T. martialis 1, Thorybes 2; Lycaen.: Stry- mon cecrops 1, S. melinus 1; Nymphal.: Junonia 2, Phy- ciodes 1, Vanessa virginiensis 1; Papilion.: Papilio 310 Trans. Acad. Sci. of St. Louts troilus 1; Per.: Eurema euterpe 1; Rhiodin.: Calephelis 4; Sest.: Synanthedon rubristigma 1, S. geliformis 1. Coxteoptera (7:39)—Cerambyc.: Typocerus 18; Cler.: Trichodes 1; Dermest.: Attagenus 12, Orphilus 4; Elater.: Cardiophorus 1; Lampyr.: Chauliognathus 1; Scarabae.: Euphoria 2. Hemiprera (5:41)—Lygae.: Melanocoryphus 1; Penta- tom.: Euschistus crassus 7; Pyrrhocor.: Arhaphe 2, Largus 22, in cop.; Redwvi.: Zelus bilobus 9. ROSACEAE, Amyepauus Persica, Ma., R.—Sixteen visitors observed, . 10. Lone-roncurp Brzs (1)—Anthophor.: Emphoropsis ¢ ?, fq. Suorr-roneuep Brzs (3, 2° )—Halict.: Chloralic tus longiceps, C. nymphalis, C. tegularis fq. Oruer Hy- MENOPTERA (4)—Humen.: Leionotus histrio, Odynerus fq.; Scoli.: Campsomeris plumipes ab., C. quadrinotata. Dirrera (4, n.)—Agromyz.: Milichiella; Ortal.: Huxesta; Sarcophag.: Helicobia; Seps.: Sepsis. Leprnoprera (4)— Dana.: Danaus archippus; Papilion.: Papilio philenor, P. polyxenes; Pier.: Catopsilia. _Lavrocerasvs Carouinsana, Mis., W.—First 1000 indi- viduals taken on the flowers, F. 5-Mr. 4, follow: Lone-roncurp Brzs (96)—Anthophor. 1, Ap. 9%. SHort-toncump Bes (573)—Andren. 527, Collet. 21, Halict. 25. Oren Hymenoprera (13)—Chalcid. 4, Ewmen. 7, Larr.1, Vesp,1. Dierera (288)—Acalyptratae 11, An- thomy, 29, Bibion. 1, Conop. 1, Muse. 118, Sarcophag. 32, Syrph. 74, Tachin, 22. Lepmorrera (30)—Rhopalocera 29, Heterocera 1. Florida Flowers and Insects 311 Opandrena scutellaris ¢ 2, sc., showed 52.7% of the in- dividuals and is probably more important than all of the rest together. Papus Sezotina, Mis., W.—F. 3-Ap. 2, 29 species, 102 individuals observed, Mr. 14, 15. Lone-roneuEp Bess (1:15)—Ap.: Apis se. 15. Suorr- TONGUED Buss (4:7, 2? )—Collet.: Colletes sp. 1; Halict. (se.) ::Agapostemon 3, Chloralictus reticulatus 1, C. tegu- laris 2. Orner Hymenoptera (3:14)—Sphec.: Chalybion 1; Scoli.: Campsomeris plumipes 4, C. quadrinotata 9. Diptera (14:40)—Anthomy.: Limnophora 1; Bombyli.: Bombylius 2; Conop.: Zodion 3; Musc.: Chrysomyia 20, Lucilia caesar 1, Musca 1; Stratiomy.: Nemotelus 1, Odon- tomyia cincta 1, O. trivittata 1; Syrph.: Allograpta 1, Kristalis dimidiatus 1, Helophilus similis 1, Psilota 1, Tropidia 5, in cop. Lepmoprera (6:23)—Hesperi.: Achal- arus 2, Polites baracoa 10; Nymphal.: Junonia 2, Papi- lion.: Papilio eresphontes 2, P. marcellus 2, P. philenor 5. Cotzoprera (1:3)—Lampyr.: Chauliognathus 3. Prunus Umpetiata, Mis., W.—Ja. 18-Mr. 16, 89 visi- tors observed F, 8-14. Lone-roncuzp Buss (5)—Anthophor.: Emphoropsis é , ab.; Ap.: Apis. se., ab.; Bomb.: Bombias scutel- laris @ ; Xylocop. (8 2): Xylocopa micans, X. virginica s¢., fq. SHort-ronauep Buss (11)—Andren.: Opandrena scutel- laris 3 9 , SC, in cop., type; Halict. (¢): Agapostemon, Chloralictus apopkensis se., type, C. ashmeadii se., type, C. longiceps, type, C. nymphalis sc., C. reticulatus sc., type, C. tegularis sc., Evylaeus pectoralis sc., Oxysto- 312 Trans. Acad. Sct. of St. Lows glossa matilda, type; Prosopid.: Prosopis schwartzii é ¢, ab., in cop. Orner Hymenoptera (13)—Chrysid.: Chrysis; Crab- ron.: Solenius; Eumen.: Eumenes fraternus, EB. smithii ab, Leionotus bifurcus, L. floridanus type, L. his- trio ab., L. saecularis, Odynerus ab. ; Scoli.: Campsomeris plumipes; Vesp.: Polistes americanus, P. pallipes, P. rubiginosus. Diptera (32) — Agromyz.: Milichiella; Anthomy.: Coenosia ovata, ©. sexnotata, Limnophora, Phorbia; Bombyli.: Anthrax lateralis; Conop.: Physocephala, Zodion; Musc.: Chrysomyia; Oscin.: Siphonella; Sar- cophag.: Helicobia, Ravinia quadrisetosa, Sarcophaga assidua, Sarcophagula; Syrph.: Allograpta, Ceria, Eris- talis albiceps, E. transversus, Helophilus similis, Meso- gramma marginata ab., Orthoneura, Psilota fg., Syrphus, Tropidia, Volucella fasciata, V. sexpunctata, Xylota fq.; Tachin.: Archytas, Hypostena vanderwulpi, Plagipros- pherysa, Siphona, Xanthomelana. Lepmworrera (28)—Dana.: Danaus archippus ab.; Hesperi.: Epargyreus, Goniurus, Lerodea, Paratrytone, Polites baracoa, Thanaos juvenalis ab., T. martialis; Lycaen.: Atlides, Strymon eecrops fq., S. melinus fq.3 Nymphal.: Dione, Junonia, Phyciodes fq., Vanessa ata- lanta, V. virginiensis; Papilion.: Papilio cresphontes, P. glaucus, P. marcellus ab., P. philenor ab., P. polydamus, P. polyxenes, P. troilus fq.; Pier.: Catopsilia, Zerene; Rhiodin.: Calephelis; Arcti.: Lerina; Noctu.: Pachy- morpha Rusvs Vintosus, Mi., W.—F. 13-Ap. 26, 6 visitors ob- served, Mr. 13. The list is fragmentary. Lerworrera (6) — Hesperi.: Pamphila, Thorybes; Nymphal.: Junonia; Papilion.: Papilio cresphontes, P. philenor ab., P. troilus. Florida Flowers and Insects 313 RUBIACEAE. GeLsEMiIum Sempervirens, Ma., Y. — The corolla is 30 to 35 mm. long, its five-lobbed border ex- panding to the same extent. The base is 25 mm. long. From a narrow part below, which is 8 mm. long by 2 mm. wide, it widens regularly to a throat which is 15 mm. wide. The throat within is rich orange. In the short-styled form the 4 lobed stigma rises 10 mm. The 5 filaments are attached to the corolla for 8 mm., their free ends exposing the anthers 6 mm. beyond the mouth of the tube. The stamens form a central col- umn, the anthers being extrorse. The bee touches the stamens exteriorly and inserts its tongue between the filaments. The stigma lobes project between the fila- ments. The tube is narrowed by the filaments, the pistil and the stigma. A tongue at least 10 mm. long is neces- sary to drain the nectar. There is a tendency to sterno- tribe zygomorphism. The flower is horizontal, the bor- der vertical, the upper lobes somewhat reflexed. The stamens are bent a little upward, and Emphoropsis lands upon them and enters on the upper side of the flower. In the long-styled form the stigma is less exserted than the anthers in the short-styled form and the stamens are longer than the pistil of that form. The tube is ob- structed by the anthers. The style is declined toward the lower side. Self-pollination might be effected by insects in the short-styled form. In the other form it can hardly occur. D. 7-Ap. 3, 5 visitors observed, F. 3-26. Long-tonauep Bers (1)—Anthophor.: Emphoropsis é 2, ab. Lepmoprera (4)—Hesperi.: Goniurus; Papilon: Papilio philenor; Pier.: Catopsilia, Eurema nicippe. 314 Trans. Acad. Sct. of St. Louis Hovustonia Rorunpirouia, Ma., W.—The flowers often appear as mats a foot or more in extent, and are quite attractive to insects. The corolla is salver-form, with four ovate white lobes, the tube yellowish. The tube is 6 mm. long and nectar is secreted at its base. | In the short-styled form the stigma stands in the throat, while the stamens rise 3 mm. above. The stamens are separated (3 to 4 mm.), so as to apply pollen to the heads or other parts of the visitors, where it will be read- ily received by the stigma of the long-styled flower. The long-styled form can only deposit its pollen on the tongues of the visitors, where it will touch the stigma of the short-styled form. The stigma rises 3 to 4 mm. above the mouth, the anthers being included. The throat is narrowed by the introrse anthers. Ja. 5 to Ap. 26, 28 visitors observed, Ja. 30-F 24. SHorRT-ToNGUED Brzs (1)—Halict.: Agapostemon ¢. Orner Hymenoptera (1)—Scoli.: Campsomeris plumipes. Lepmortera (26)—Dana.: Danaus archippus; 7 espert.: Goniurus, Hylephila, Lerodea, Polites baracoa, P. bret- tus, Prenes, Thanaos juvenalis, T. martialis, T. terentius, Thorybes; Lycaen.: Strymon cecrops, 8. melinus; Nym- phal.: Dione, Junonia, Phyciodes, Vanessa virginiensis; Papilion.: Papilio marcellus, P. philenor, P. troilus ; Pier.: Catopsilia, Eurema delia, E. euterpe, E. nicippe, Zerene ; Rhiodin.: Calephelis. RUTACEAE. Facara Cuava-Hercuuis, Mis., Y—Mr. 16-Ap. 16; first 1000 individuals taken on staminate flowers, Mr. 17-AP. 2, follow: Lone-toneuep Brzs (171)—Anthophor. 3, Ap. 128, Bomb. 3, Epeol. 1, Megachil. 28, Nomad. 3, Stelid. 3, Florida Flowers and Insects 315 Xylocop. 2. SHort-ronavep Bees (683)—Andren. 32, Col- let. 259, Halict. 192. Orner Hymenoprera (108)—Cercer. 2, Crabron, 8, Eumen. 37, Ichneumon. 1, Larr. 1, Nysson. 1, Philanth, 24, Pompil. 5, Scoli. 7, Sphec. 4, Tiphi. 10, Vesp.8. Drprera (177)—Bombyli. 7, Conop. 37, Muse. 86, Syrph. 15, Tachin. 32. Lepmorrera (51)—Rhopalocera 50, Heterocera 1. Cotroprera (6)—Lampyr. 6. Hemie- TERA (4)—Pentatom. 4. The Colletidae, about 2.7% of the families, showed 25.9% of the individuals. Colletes thoracicus had 256 individuals. The Colletidae, Halictidae and Apidae, about 8.3% of the families, made up 57.9% of the indi- viduals. SCROPH ULARIACEAE. InysantHEs GranprFrtora, Ma., R.—The flower is blue, the lobes more or less white, especially at the tips. The upper lip is small and cleft, the lower spreading, with three large rounded lobes. On the lower wall of the corolla the filaments of the two lower sterile stamens form two ridges which are covered with yellow pollen-like glands. The filaments have about the middle an awn- like process. The stigma lies under the upper lip. The two fertile anthers are coherent and lie on the upper wall about 2 mm. below the stigma. In this position self-pollination is fairly impossible. The insect’s tongue touches the stigma before the anthers. The stigma is sensitive and closes after contact. When the tongue touches the anthers and is withdrawn, it turns back the lower lobe of the stigma and closes it. The tube is narrow, rendered more so by the sterile filaments, perhaps entirely excluding short tongues and requiring long tongues to incline to the opposite side 316 Trans. Acad. Sci. of St. Louis where they will touch the anthers and stigmas. The tube is about 6 mm. The flower is nototribe, but insects may land on any side and insert their tongues. F. 15-Mr. 23, 4 visitors observed, F’, 21-Mr. 7. SHort-toncuep Brzs (1)—Halict.: Chloralictus ash- meadii 2, fq. Leprpoprera (3)—Hesperi.: Polites bara- coa, Thanaos juvenalis; Nymphal.: Phyciodes. Linarra Canapensis, Ma., R. (2, 228; 5, 585)— F. 17-Ap. 26, 31 visitors observed at Orlando and Inverness, F. 17- Mr. 24. Lone-Toneuzp Bzzs (6)—Ap.: Apis; Megachil.: Coeli- oxys sayi 2, Megachile lanuginosa ¢ ?, M. mendica 2; Nomad.: Centrias 2 ; Stelid.: Dianthidium notatum 8. SHort-Toneuep oa (6, 2 )—Halict.: Agapostemon, Augochlora fulgida, A. sumptuosa, Chloralictus longi- ceps type, Evylaeus pectoralis, Odontalictus. OTHER Hymenoptera (1)—Scoli.: Campsomeris quadrinotata. Diptera (3)—Bombyli.: Toxophora amphitea; Syr ph. (f., n.): Baccha clavata, Mesogramma marginata. Ler! DOPTERA (15)— Hesperi.: Ancyloxypha, Atalopedes, Lerodea, Polites brettus, P. cernes, Prenes; N: ymphal. : Junonia,. Phyciodes, Vanessa virginiensis; Papilion.: Papilio marcellus, P. philenor; Pier.: Catopsilia, Eure- ma euterpe, Zerene; Arcti.: Utetheisa. PENTSTEMON Hirsutus, Ma., W.—Mr. 30-Ap. i, 1 visi- tor observed, Ap. 8. Lone-Tonevep Bers (1)—Bomb.: Bombus america- norum 2, SOLANACEAE. Puysauis Arenicona, Mi., Y.—Mr. 28-Apr. 29, 1 visitor observed Ap. 6. Florida Flowers and Insects S17 SHoRT-TONGUED Bzzs (1)—Collet.: Colletes latitaris 3. Sotanum Nicrum, Ma., W.—F. 24-Ap. 29, 1 visitor observed at Orlando, F. 24. Lone-roncuep Brzs (1)—Bomb.: Bombus impatiens, 2,6, UMBELLIFERAE, Hyprocoryie Umpeixiata, Pol., Y. (4, 456, 459)—F. 27- Ap. 26, 33 species and 65 individuals observed at Or- lando, Mr. 15-20. SHort-toneven Brzs (1:1)—Panurg.: Perdita ¢, 1. Orner Hymenoptera (10:18)—Larr.: Notogonidea 4, Tachysphex apicalis 1; Oxybel.: Oxybelus fulvipes 1, type; Pompil.: Anoplius 1, Lophopompilus 1, Pompi- loides americanus 4, P. argenteus 3, P. marginatus 1, P. subviolaceus 1; Tiphi.: Tiphia vulgaris 1. Dr- TERA (19:41) — Agromyz.: Milichiella 1; Anthomy.: Bithoracochaeta 1, Homalomyia 1, Limnophora 4; Musce.: Chrysomyia 2, Lucilia sericata 1, L. sylvarum 4, Musca 4; Ortal.: Tephronota 7; Oscin.: Chlorops 1; Sarcophag.: Ravinia floridensis 1, Sarcophaga bullata 2; Syrph.: Baccha tarchetius 1, Mesogramma boscii 5, M. marginata 2, Microdon viridis 1, type, Orthoneura 1; Tachin.: Phasioclista 1, Senotainia trilineata 1. CoLEOPTERA (3:5)—Carab.: Lebia 1; Lampyr.: Chauliognathus 3, Polemius 1, VERBENACEAE. Lantana Camara, Mas., Y.—Three visitors observed at Orlando, F. 24, Lepworrera (3) — Beek: Thanaos juvenalis; Pa- Pilion.: Papilio marcellus, P. philenor. 318 Trans. Acad. Sci. of St. Louis Lantana Oporata, Mas., R.—Ja. 24-Ap. 11, 17 visitors observed, F. 16-Mr. 19. Drerera (1, f, n) —Syrph.: Mallota. Lepmoprera (16)—Dana.: Danaus archippus; Hesperi.: Atrytonop- sis, Cocceius, Lerodea, Megistias, Polites baracoa, Thanaos juvenalis; Nymphal.: Junonia, Phyciodes, Van- essa virginiensis; Papilion.: Papilio cresphontes, P. phil- enor, P. troilus; Pier.: Eurema euterpe, E. nicippe, Zerene. VIOLACEAE. Viota Lancrouata, Mi.. W.— The flowers are small. The petals are white, the lower one streaked with purple, lateral not bearded. The flower is not adapted to pollen- collecting bees. The spur is short. From the tip of the style a narrow passage leads to the base of the tube, a distance of about 4mm. The stigma is beaked and only surpasses the anthers a little, but enough to prevent self- pollination. Insects land on the upper petals and approach the nec- tary from above. The flower is therefore sternotribe, the place of pollen-contact being the under surface of the proboscis. Ja. 19-Ap. 26, 16 visitors observed, Ja. 19- F. 20. SHort-ToncuEp Bzxzs (3, 2° )—Halict.: Chloralictus apopkensis, C. ashmeadii, Odontalictus. DrprERa (3)— Tachin.: Chaetoglossa picticornis, C. violae fa, ‘ype Siphona fq. Leprmoprera (10)—Hesperi.: Amblyscirtes, Cocceius, Goniurus, Megistias, Polites baracoa, Thanaos juvenalis; Nymphal.: Phyciodes; Papilion.: P apilio philenor; Pier.: Eurema delia, E. euterpe. Florida Flowers and Insects 819 FLOWER CLASSES. Ma.—Non-social long-tongued bee flowers (including 0) are 07.1% of the flowers observed, receive 34.4 and 28.6% of the total and the pollinating visits. The class re- ceives 71% of the pollinating visits of long-tongued bees and 38.2 of those of Lepidoptera. These are respectively 32.4 and 38.9% of the total visits to Ma. Ma shows for colors R 40.6, Y 25.0 and W 34.3%, and for visits to each color 49.1, 15.7 and 35.1%. Of the total visits to R, Y and W, Ma receives 71.0, 44.3 and 18.9%. Ma shows maxima of long-tongued bees, 70.5, short- tongued bees 33.6, and Lepidoptera 48.9. Of total non-pollinating visits, Ma shows 78.4%, of those of Lepidoptera 88.3, flies 75.0, Lower Hymenoptera ~ 71.4 and short-tongued bees 66.6%. Many insects get pollen or nectar without effecting pollination. Ma shows maxima of Trochil, Anthophor, Bomb, Eucer, Megachil, Stelid, Halict, Bombyli, Hesperi, Nymphal, Papilion, Pier, Scoli, long-tongued bees, total Lepidop- tera. Mi.—The non-social short-tongued bee flowers are 17.8% of the flowers observed and receive 6.2% of the Visits. Lepidoptera are 57.4% of the visitors and short- tongued bees 22.2. The Lepidoptera are not better adapted but more common. The colors are R 10, Y 30 and W 60%, and the visits R 7.4, Y 16.6 and W 75.9. But the percentages of total Visits to Mi, R, Y and W are 1.9, 8.4 and 7.4. Mas.—The social long-tongued bee flowers are 10.7 %o of the flowers and show 8.4% of the visits. Of the visits to Mas, the Lepidoptera show 53.4%. 320 Trans. Acad. Sct. of St. Louis Mis.—The social short-tongued bee flowers are 8.9% of the flowers and show 26.4% of the visits. Of the visits to this class, the lower Hymenoptera show a maximum of 28.2%, the flies with 27.0 and the Lepidoptera with 23.1. Of the visits of the flies, lower Hymenoptera and short-tongued bees, 39.9, 36.5 and 28.8 are to this class. Mis shows maxima of Xylocop, Collet, Lycaen, Sphee, Bembie, Crabron, Philanth, Syrph, Conop, Stratiomyidae. Pol.—The polytropic flowers are 5.3% of the flowers and receive 24.4% of the visits. Of the visits to this class, the flies show a maximum of 37.2, the lower Hymenoptera with 33.4. Of the visits of the flies and lower Hymenop- tera 47.0 and 39.8% fall under Pol. Cases referred to Pol on account of the species of vis- itors may be shown to belong to Mis when the individuals are counted. In Fagara clava-herculis the short-tongued bees forming 9.3% of the families, were 48.5% of the individuals. Pol shows maxima of Eumen, Pompil, Oxybel, Tiphi, non-aculeata; Tachin, other Calyptratae, total Muscoidea, total flies, lower Hymenoptera, Coleoptera, Hemiptera. Social and non-social flowers —While the social (Mas, Mis, Pol) are only 25% of the flowers observed by me at Inverness and Orlando, they receive 59.2% of the visits. Of 437 flowers observed at Carlinville, Illinois (8, 158), the social, 45.4% of the total, received 78.9% of 13,971 pollinating visits. The differences are 34.2 and 33.5. Couors. Red.—Red flowers are 32.1% of the flowers observed and receive 23.8% of the visits. Of the visits to red, the Lepidoptera made 49.2%. Red shows maxima of Eucer and Megachilidae. Florida Flowers and Insects Sal Yellow.—Shows 25% of the flowers observed and 12.1% of the visits. Of the visits to yellow, 23.8% are made by flies and 23.8 by Lepidoptera. White—Shows 42.8% of the flowers and 64% of the visits. The lower Hymenoptera show 84.2, flies 76.7, Coleoptera and Hemiptera 64.5, short-tongued bees 56.7, Lepidoptera 54.9, long-tongued bees 41.1% under white, all maxima. Of total visits to white, the Lepidoptera show 27.9%, lower Hymenoptera 27.0, flies 23.2, short- tongued bees 10.6, long-tongued 7.5. General colors—As we pass from red through yellow to white, the long-tongued bees show 38.2, 20.5, 41.1; Lepi- doptera 36.1, 8.8, 54.9; short-tongued bees 25.9, 17.3, 56.7; Coleoptera and Hemiptera 12.9, 22.5, 64.5; lower Hymen- opetera 10.7, 5.0, 84.2; flies 8.3, 14.8, 76.7. Or long- tongued bees and Lepidoptera 36.7, 11.9, 51.3; other in- sects 13,3, 12.2, 74.4. The determining condition is that red generally shows the deepest seated nectar and white the shallowest. At Carlinville, Illinois, were observed 13,971 pollinating visits to 437 flowers, of which 30.2% were red, 30.6 yellow and 39.1 white. Long-tongued bees show 34.0, 31.4, 34.5; Lepidoptera 36.8, 23.7, 39.3; short-tongued bees 11.9, 35.8, 52.1; flies 7.7, 32.9, 59.3; lower Hymenoptera 7.5, 29.4, 62.9; Coleoptera and Hemiptera 6,3, 31.3, 62.2. Long-tongued bees and Lepidoptera show 34.6, 29.2, 36.0; other insects 8.9, 32.9, 58.0. Species and individuals.—In the case of 18 species in which the individual insects were taken as they came, bees shows 24.9% of 477 pollinating visits and 37.4% of 2,679 individuals. In the case of 41 flower species observed at Carlinville, Illinois, bees were 31.6% of 405 species, made 40.2% of 1,452 visits and were 57.4% of 7,391 individuals (9: 70.) 322 Trans. Acad. Sct. of St. Louis INSECTs. Long-tongued bees.—In the case of 338 visitors and 867 visits, the long-tongued bees show 8.8% of the species and 11.7% of the visits; 13% of the pollinating and 1.9% of the non-pollinating.. The maximum is 70.5% under Ma. For colors, they show 38.2% to red and 41.1 to white, or 6.1 more and 1.7 less than the percentages of colored flowers observed. The Megachil, Bomb and Anthophor, 33.3, 13.3 and 6.6% of the long-tongued bees, make 44.1, 14.7 and 11.7% of the long-tongued bee visits. These, with Eucer and Stelid, have maxima under Ma. The Xylocopidae show maxima under Mis. Short-tongued bees——Being 7.9% of the species, these show 11.9% of the visits; 12.4% of pollinating and 8.8 of non-pollinating visits. The maximum is 33.6 under Ma. They show 25.9% to red and 56.7 to white, or 6.27 less and 13.9 more than the percentages of those colors. All of their non-pollinating visits are to red flowers for which they are too small. The Halict, 59.2% of the short-tongued bees, make 85.5% of the short-tongued bee visits. They show maxima under Ma. The Collet show maxima under Mis. Lower Hymenoptera.—Being 29.5% of the visitors they make only 20.5% of the visits; 21.4% of the pollinating and 13.7 of the non-pollinating. A maximum of 39.8% is under Pol. They show 10.7 under red and 84.2 under white, or 21.4 less and 41.4 more than the percentages of those colors. The Eumen., Pompil. and Scoli., 21, 19 and 6% of the lower Hymenoptera, make 27.5, 14.0 and 14.67 of the lower Hymenoptera visits. They show maxima as follows: the Scoli under Ma; the Spec, Bembic, Crabron and Philanth under Mis; the Eumen, Pompil, Oxybel, Tiphi and non-aculeata under Pol. Florida Flowers and Insects aac Fles.—These are 30.7% of the visitors and make 19.3% of the visits; 20.3% of the pollinating and 11.7 of the non-pollinating. The maximum is 47% under Pol. They show 8,3 under red and 76.7 under white, 23.8 less and 33.9 more than the percentages of those colors. They show maxima as follows: the Bombyli under Ma; the Syrph, Conop and Stratiomy under Mis; the Muscoidea under Pol. They show in percentages of fly species and visits: Bombyli 6.7, 10.7; Syrph 23.0, 26.7; Tachin 27.8, 23.2; other Calyptratae 14.4, 16.0; total Calyptratae 42.3, 39.1; total Muscoidea 61.7, 54.7. Lepidoptera.—Are 15.6% of the species and make 82.5% of the visits; 29% of the pollinating and 58.8% of the non-pollinating. The maximum is 48.9% under Ma. They show 36.1 under red and 54.9 under white, or 4 and 12.1 more than the percentages of those colors. Of the non-pollinating visits 80% are to red. These show percentages of Lepidoptera species and visits: Hesperi 41.5, 44.6; Papilion 13.2, 18.7; Nymphal 9.4; 12.7; Pier 9.4, 12.4. These families show maxima under Ma, The Lycaen show maxima under Mis. Coleoptera and Hemiptera.—Are 6.8% of the visitors and make 3.5% of the visits. The maximum of 48.3% is under Pol. They show 12.9% under red and 64.5 under white, or 19.2 less and 21.7 more than the percentages of those colors, Early visits at Inverness and Carlinville—Comparing Percentages of the visits observed at Inverness, Ja.-Ap., with those observed at Carlinville before July shows some remarkable results. The Lepidoptera are the most con- Spicuous with 29 against 6.4. The lower Hymenoptera show 21.4 and 10.1, resembling the late visits at Carlin- ville, 22.7. ‘The lower Hymenoptera and Lepidoptera 324 Trans, Acad. Sci. of St. Louis make 51.7% of the early visits at Inverness and only 16.5% of those at Carlinville. The short-tongued bees show 12.4 and 27.7, while the long-tongued show 13.0 and 20.7. The bees make 48.5% of the early visits at Carlinville and 25.4% of those at Inverness. The short-tongued bees and flies make 56.9% of the early visits at Carlinville and 32. is of those at Inverness. BIBiLioGRAPHY. 1. Robertson, C. 1887 (12). Fert. Calopogon. Bot. G. 12 :288-91. 2. ——,1888. Zygomorphy. Bot. G. 13:146-51, 203-8, 224-30. 3. ——, 1889 (5). Fis. I. Bot. G. 14:120-6: 4. ——, 1890. Fis. Umbelliferae. St L. A. 5:449-50. 5. ——, 1891 (2). Fis. Asclep.Scroph. St. L. A. 5 :569-98. 6. ——, 1891 (3). Fis. VI. Bot. G. 16:65-71. 7. ——, 1893 (2). Fis. X. Bot. G. 18 :47-54. 8. ——, 1923 (10). Visits of Insects. Psyche 30 :158-69. 9. ——, 1924 (9). Fls. XXIII. Bot. G. 78 :68.84. me eee oy ph? a aged Bat] Transactions of the Academy of Science of St. Louis a “ps ieee AA Se rongpe Volume XXV, No. 9 FIELD STUDIES IN THE BEHAVIOR OF THE NON-SOCIAL WASPS PHIL RAU FIELD STUDIES IN THE BEHAVIOR OF THE NON-SOCIAL WASPS CHAPTER CHAPTER CHAPTER CHAPTER CHAPTER CHAPTER Co A ™™ER ~ te, xy. VE. VOT. PHIL RAU CONTENTS The life and habits of two beetle-hunting wasps. Cerceris raui Cerceris bicornuta An amputator of spiders’ legs. Pseudagenia mellipes The grass-carrier wasp. Chlorion auripes An assortment of twig dwellers. Diphlebus biparitor Diphlebus tenax Silaon niger Solenius interruptus Stigmus fraternus raui. Hypocrabro stirpicolus Wasps of the genus Odynerus and their nesting habits. Odynerus (Stenodynerus) conformis... Odynerus (Stenodynerus) pennsy wk POT TT ea eh ee eerie Soy cea Odyneris Leucomme lars cceescecseceveenveeeeneneen ate Odynerus foraminatus ... saith Odynerus capra The patriarchate wasps of the genus Trypoxylon, Try posylon Clavat urn a neenenn mmm Trypoxylon politum Seen Trypoxylon sates Ee ag ee Trypoxylon texense .. Rigo SR The nesting habits of the ae mud-dauber Sceliphron caementari The prey of Sctliphows: ibaa: = PAGE 325 337 INTRODUCTION, This work on the behavior of twenty-three species of solitary wasps was all done in the vicinity of St. Louis, Mo., between the years 1917 and 1923. As in previous studies, all insect material was sub- mitted to experts for identification, and their names ap- pear in brackets throughout the work. The photographs Were made by the author, and the drawings were made by Dr. Gustave Dahms of this eity. A debt of gratitude is due to my wife for valuable assistance on the manu- Script. ‘Observation sets the problem; experiment solves it, always presuming that it can be solved, or at least if powerless to yield the full light of truth, it sheds a cer- tain gleam over the edges of the impenetrable cloud.’’ —J. H. FABRE, “So the first task of comparative psychology is to find out the instinctive equipment of any animal studied. In- stincts are, however, well worth study for their own sake. An instinctive fear of a certain enemy may be as truly useful to an animal as sharp teeth or protective colora- tion. Instincts are the expression of structures and func- tions of the nervous system, and are as real and as im- portant matters for the biologist as are bones and blood vessels.’’ —E, L, THORNDIKE. ‘It may be noted that all the activities of ants, their reflexes and instincts as well as their plastic behavior, gain in precision with repetition. In other words, all their activities may be secondarily mechanized to form habits, in the restricted sense of the word. This is tanta- mount to say that even the reflexes and instincts are not So stereotyped but that they may become more so by ex- ercise during the lifetime of the individual. And not only do ants thus form habits, but . . . these habits when once formed are often hard to break. It is certain that many instincts . . . are at first complete and indefi- nite and are guided to their proper course by stimuli that affects the organism at a later period. .... There is little doubt, moreover, that the more fixed or stereo- typed instincts are phylogenetically the older.’’ —wmM. M. WHEELER. Sa Nats = Field Studies of the Non-Social Wasps 325 CHAPTER I. Tue Lire Hasirs or Two BEEtLe-HUNTING WASPS. THe Jerseypate Cercents, Cerceris raui Roh. [S. A. Rohwer]. Jerseydale (fig. 26), some 30 miles south of St. Louis, consists of a railroad sign-post, a milk-can platform, and a well of excellent cold water; this equipment renders it easily recognizable as the center of a thinly populated but prosperous dairying community. Besides the well, Jerseydale has other attractions, un- noticed perhaps by the men who daily drive up to the platform to exchange their full cans for empty ones. In the hard road under the hoofs and wheels abound the Cerceris wasps, Cerceris raui. This wasp (fig. 27) was described by Rohwer from material submitted from three distinct localities: Lake View, Kan.; Wickes, Mo., and Jerseydale, Mo.* While only fragmentary notes were gathered from the first two localities, at Jerseydale the life secrets of this species were more fully discovered. One salient feature of their habits is that in all three places their nests were dug chiefly in the hard-packed roads, where traffic was heavy, and only a very few nests on the outskirts of a colony occurred at the roadside. The hole which this species makes is an excellent piece of workmanship. There is no attempt at concealment of the burrow, such as we find practiced, sometimes very skillfully, by some other species of wasps. Her nest is So inaccessible in its situation that she can well afford to work boldly and openly before all men, without fear of her home being molested, unless by parasites which are cunning enough to elude her and gain access by her own doorway. The burrows in each of the three habitats *Also taken later at Wesco, Mo., 108 miles west of St. Louis. 326 Trans. Acad. Sci. of St. Louts were found in the middle of a well-beaten road, where the earth was so hard packed as to be almost impenetra- ble with ordinary tools. ‘A spading-fork, a digging-trowel and a pruning-knife each in turn proved ineffectual in the conquest, and only by the aid of a hatchet was I finally able to follow a very few burrows to their termini. The work was rendered increasingly difficult by the fact that the nests were in the beaten track, and at the ap- proach of each automobile or wagon it was necessary for the perspiring investigator to take flight to a safe dis- tance to escape the wrath of the irate drivers who seemed not to appreciate the beauty of the quest of truth when it led to the digging of chuck-holes in their roadway. At Lake View the road leading to a country club was beaten so hard by the automobile traffic that I could not dig up a single nest. At Jerseydale the yellow clay was very sticky when wet and flinty when dry, and at Wicks (fig. 28) the admixture of clay and rocks rendered digging most difficult. At the two latter places, however, a few nests were found at the side of the road which could be dug out to sufficient depths to learn some details of their interiors. The mouth of the hole is always beautiful in its sym- metry; it is 14 inch in diameter, and around the aperture is a mound of chips or granules of earth which have been brought up from below (see fig. 29; a ten-cent coin nearby shows the relative size). The burrow is amply large; the diameter is sufficient to permit the occupant easily to turn around or even to make her toilet while within. The tunnel is long and tortuous. In the places where rocks are present in the soil this is, of course, the logical re- sult, but the same condition exists to an extreme degree in clear soils also. The illustration (fig. 30, %4 nat. size) gives an idea of the rambling course of one of these chan- Field Studies of the Non-Social Wasps 327 nels, but, of course, an incomplete idea, since a diagram can show the curves in only one plane, and not the curves toward and away from the observer. Another was de- scribed in detail in my notes as follows: This hole went straight down for 2 inches, then curved toward the south- east for 3 inches, then southwest for 4 inches, made a sharp turn toward the northwest and downward for 4 inches, westward and still down for 2 inches, upward and west 2 inches, then curving downward and south about 4 inches. Along this section of the tunnel were three cells containing fat Cerceris larvae and beetle remains. These cells were oval and about 114 inches long and % inch in diameter. Beyond this the tunnel continued slightly up- ward and westward for 3 inches more, and in the end, in a sort of pocket, were 3 more fat beetles. This indi- cates that the wasp digs her tunnel and makes several pockets at the terminus for brood cells. The tunnel varied slightly in diameter, the minimum being about % inch. The total length of this one was 24 inches, and the cells were situated 17 inches below the surface of the ground. One can readily see what an enormous task this mother wasp had accomplished, to have made so large an excavation in soil which was so nearly impenetrable that I myself could hardly dig it. The excavation of another burrow was begun in the absence of the mother wasp. The hole was likewise very tortuous, and at a depth of 15 inches it turned directly under the hardest part of the road, where the tools broke and the digging had to be abandoned. Meanwhile the mother returned, and so persistent was she in trying to enter the wreck of her home that twice I picked her up in the forceps with perfect ease. She fairly fought to get back to the place, and dug for more than an hour trying to follow her old burrow, which she recognized so 328 Trans. Acad, Sci. of St. Louis surely despite its mutilation. I left her still working feverishly at her task of reconstruction. Since in the three places where these wasps have been found they occurred in settlements or colonies, one won- ders if they do not dig their way out of their subter- ranean winter dormitory in the spring and then turn around and enlarge their exit-passage for their new bur- rows. It seems incredible that one wasp is able to dig so large a tunnel in so short a time. I earnestly hope that the reader will be satisfied with the details of these few burrows, so I may mercifully be spared the task of digging out another one. So far as I could learn from others partly excavated, the tortuous course and the dimensions of these are fairly typical. I have not been so fortunate as to see the beginning of the digging of the burrow. Later on in the process, however, when the tunnel was only being enlarged or ex- tended, I have observed the outward evidence of work going on within. There is then little display to attract the attention of the passer-by; the wasp in her dark tun- nel down under the earth digs up the soil—I know not how—and packing a considerable mass behind her body and above her in the tunnel, backs up and pushes it out above the surface with her abdomen. This soil comes out, not in loose, powdery form, but in granules packed together to form plugs or sausage-like masses, which protrude for a time above the surface and later, when thoroughly dry, collapse in a heap at a touch or breeze. This digging was observed several times early in the day, before the wasps had come out of their holes to brush back the dirt from around the mouth of the burrows, and depart for refreshments or prey. Hence I cannot say with fairness that these Cerceris are not early risers, for I do not know how early they begin their Field Studies of the Non-Social Wasps 329 work in the dark gallery, but they do not open up their doors, which are closed each night, until the sun is high and hot in the summer sky. This statement implies that they sleep in their burrows, and so they do; at least the females of the population which possess burrows sleep there; I have not yet been able to find, by careful scrutiny of the vicinity, where the males find their night’s lodging. I have never yet seen the males enter the burrows, although they are frequently to be seen pursuing the fe- males as they come and go. They are small and agile, and difficult to apprehend. The sexes are so different in appearance that they look like two different species, the female a large, reddish-colored insect, and the other much smaller and yellow. Mating occurs on the wing or while the home-maker goes to and fro about her work; no special frolic or festival has been observed to accom- pany this function, such as occurs in several species of wasps. The owners of the burrows come home and enter the nest early, about 3:30 to 4:30 o’clock, and push up a plug of loose earth to close their door snugly for the night. This roof remains in position remarkably well, in spite of the fact that it is so soft that it will collapse if tickled with a straw. On one afternoon when a shower occurred early, I visited the colony and was surprised to find that even at that time of day, which is usually the wasps’ busy hour, these good housewives had all hurried home to shut up the house before the storm broke; every door of the fifty or sixty nests was closed from within! In covering the hole the wasp backs up with the ab- domen extended straight and the load of earth under the abdomen and behind the legs. Thus she pushes the load up to a point 14 inch from the top, where she presses the dirt against one side of the hole. Thus she brings up a 330 Trans. Acad. Sci. of St. Louts few more loads, condensing and packing the earth against one wall; then she brings up more dirt to close the re- maining aperture. Even then she does not stop, but adds many layers to the plug from the inside; we are made aware of this by the movement of the plug as it heaves up with the vigorous pressure from underneath as each load is packed in place. In many cases the wasps seem to deem their day’s work done at 4 o’clock, and by 6 most of the holes are partly or completely closed for the night. It is a pretty sight in the morning to see the wasp first venturing out. The granules of clay covering the hole quiver and part, the head appears, and the opening is silently made wider. Then the wasp pauses to peer cau- tiously about (fig. 31); this habit of cautious recon- naissance is a very characteristic gesture of this species. She creeps so stealthily to the edge of the hole that one is hardly aware of the motion, and pauses with her face at the aperture and peers about with a gaze so intent that one can almost feel it; if she is not satisfied that all is safe, she drops stealthily back; if all is well, she darts away like a flash, leaving the burrow wide open. She has retreated thus when I moved noiselessly at a distance of 4 feet; this would indicate that her range of vision en- compasses at least that distance. Upon returning home, she seems to have no trouble in locating her burrow, al- though it is one among so many all alike, in the roadway or in the sparse grasses nearby. She does not loiter, but plunges headlong into the burrow, almost before one has time to get a good view of her. Sometimes she circles about on the wing a few times before she seems to get her bearings to make an accurate descent directly upon her burrow. Every observation upon the Cerceris raui population at Jerseydale indicated that this wasp uses only two Field Studies of the Non-Social Wasps 331 species of Coleoptera as food for her young, viz., a short, fat weevil, Thecesternus humeralis Say [E. A. Schwarz], and the long, slim weevils, Liaus concavus Say [B, A. Schwarz] (fig. 32). Less extensive observations at Lake View, Kan., and Wickes, Mo., gave the same results, but the data there were not sufficient to be conclusive. Of 22 females which were caught as they returned to the nest, 16 carried Thecesternus humeralis, and 6 had the long, _ Slender beetles, Lixus concavus. In the several brood- cells which I had the good fortune to explore, the former beetles were present in far greater numbers. In some cells all the provision was of that species. About 15 to 22 beetles seems to be the ration allowed each young wasp. In all of the cells wherein the larva had already worked over the food supply provided for it, we found that it had separated each beetle into the three com- ponent parts, the head, thorax and abdomen. The head separates easily from the thorax in these beetles, and the larva undoubtedly takes its first meal from the soft tis- Sues at the point where the thorax joins the abdomen. It is wonderful to see how clean these beetle shells have been licked when the larva is done with them. (Fig. 33 Shows the beetle remains after the feast). One wonders how it is possible for the larva to clean out the contents So completely without breaking through the body wall at various places, but when one sees the remarkable adaptation of its anatomy for just this purpose, one can readily see how it succeeds in reaching all parts of the body cavity. While it in general looks like all wasp larva, the oral end tapers into a nice point so that it is possible for this part to enter the abdomen through one opening and reach clear to the other end. The head part sways 332 Trans. Acad, Sci. of St. Louts from side to side, and the jaws move about seeking the food when it is taken from them. The beetles, when they are removed from the cells, are motionless, but if they are taken from the mother wasp as she brings them in, and placed in a vial, they soon re- vive so that they climb over one another very actively. Hence one doubts if they have actually been stung. The 22 taken from the wasp mothers, as previously stated, were very active when examined the same day; when in- spected two days later there was much excrement in the vials, indicating that life had continued for some time, but the beetles were dead; whether death had been due to the confinement or the sting, I do not know. Close observation has taught me that these wasps do sometimes sting their prey, but how general this habit is has not been ascertained. One of them stung a live weevil which { substituted for her own, while another did not. When the wasp was out foraging I placed a small ball of cotton in the mouth of her burrow; when she returned and at- tempted to enter, she dropped her prey in alarm and flew away, and when she returned she found I had re- moved the plug. Meanwhile I had also exchanged her quiet weevil for a live one, which was so active that I could hardly keep it near the hole. The mother returned, flew directly into the hole as usual, and after a few sec- onds crept to the top, poked her head out cautiously, grasped the beetle in her jaws and dragged it in; if a sting was administered it must have been done under- ground. Many wasps, in foraging or bringing in building mate- rial, take a direct flight to and from the place of interest, either with or without circles of orientation. This species, so far as I have observed, follows a rather strange course; she does not fly ‘‘as a bird flies,’’ or ‘‘in a bee Field Studies of the Non-Social Wasps 333 line,’’ but she literally makes a round trip. Several which I watched going to the cornfield on the other side of the railroad track, left the nest flying northward; after fifty feet or so they circled around toward the east, and after an absence of from 3 to 8 minutes they came back with their prey, by way of the south. Just how general this habit of circular flight is I do not know. They work with surprising rapidity in bringing in their victims. I timed a number of them on different occasions and they usually took only 3 to 8 minutes to capture a beetle and return. They lost no time in finding the open nest, plunged in, and stored the booty speedily. It was noticed that practically all of the weevils brought in by these Cerceris were covered with a crust of earth. That raises the question: do these beetles oceupy subterranean habitations where these wasps are obliged to dig for them? The wasps seem to find them so easily and in such large numbers that there must be an abun- dant supply of them near, perhaps in the cornfield. To ascertain the hunting habits of these wasps one ought to know something of the two species of beetles that they hunt, Livus concavus, properly known as the rhubarb cureulio, and Thecesternus humeralis. Of the latter spe- cies, Mr. E, A. Schwarz writes: ‘‘The life history of this weevil still remains unknown, but the imago is quite common under dried cow dung in our prairie states, or under stones in dry situations in Indiana, western Mis- souri, Nebraska, Colorado, New Mexico and Texas. Most of the specimens in our collection are covered with a thick crust of dirt, which shows that the earlier stages are passed underground and that the larvae will be found In the roots of some plants. Oceasionally specimens of the imago have been found above ground on the stems of various plants. * * * A somewhat allied species, 334 Trans. Acad, Sci. of St. Lows Rhigopridius tucumanus Heller, from Argentina, has been bred from the tuber of a potato.’’ The weevil makes so small a burden that the Cerceris can carry it very easily and swiftly. In all of the cases which I could observe closely, the wasp carried the prey, clasped in her middle legs, with the ventral side of the weevil against the under side of her own body, and in some cases it appeared that she grasped its proboscis in her jaws. She drops her burden quickly if disturbed. One wasp returned with her weevil under her body, and as she found difficulty in finding her hole, which had ac- cidentally been covered, I pressed close to see her method of search. She braced herself with her hind feet while she dug in the mound of dust with her free fore-feet. Not gaining much headway in this manner, she dropped her weevil and then worked faster with four feet. Thus she eventually found her hole on the other side of the mound. Meanwhile, I exchanged the weevil which she had laid down, for one borrowed from another wasp. Several times she went into the hole, as if to make sure that everything was right, and came out again. At last she approached the weevil, took it in her mouth, rolled over and stung it. In my eagerness I came too close and frightened her away. After ten minutes more of nervous coming and going, she again took the weevil in her mouth, curled her body almost into a circle in order to reach the prey with her sting, propping herself up meanwhile on her two protruding wings, while she inserted her sting on the ventral side of the victim and kept it there for almost a minute, her abdomen pulsating all the time. All this happened on the mound of loose soil, only a half inch from her hole, which was now open; nevertheless she took up her weevil, flew into the air, circled about a Field Studies of the Non-Socral Wasps 335 few times, alighted again in the normal manner and plunged in. While another nest was being observed, a troublesome weed that grew near it was pulled up; this changed the topography of this tiny area. Upon returning several hours later we found four weevils lying out on top of the ground, and the returning wasp even then circled and buzzed about the hole for fully half an hour before she would enter. It was evident that she had discarded the weevils lying there in the sun; either she had dropped them as she had come back from foraging and had lost them in her confusion at finding the familiar landmarks altered, or she had suspected that, since the region had been changed, her nest had probably been meddled with and so, like other wasps of our acquaintance, she had thrown out the stores she already had as though they were contaminated. One nest was especially conspicuous because at that point in the road the surface of the ground was covered with black cinders, while the circular mound of subsoil around the hole was gray. This contrast made the nest itself a conspicuous landmark for the returning wasp. I watched until the wasp left home; then with the point of the trowel IT carefully removed this gray earth and replaced it with a mound of black cinders, just like the surrounding medium. I was quite proud of my new ac- complishment of building wasps’ nests, for the imitation was, to my eye, very accurate except for the color of the material. It was, of necessity, a hasty job, for the rightful proprietor returned from her trip in only two minutes. She flew direct to the location of her hole as usual, but just above it she brought herself up with a Jerk and flew to and fro over it in an agitated manner for several minutes. Her confusion was evident; twice 336 Trans. Acad. Sci. of St. Louis she dipped down on the wing as if ready to enter, then withdrew in alarm; again she tried to scrape away the cinders a little to one side, as if thinking that her burrow had been covered by accident, as frequently happens dur- ing the mother’s absence. Thus she continued to search in all the crevices and possible places. For the first five minutes of this hunt, she carried her prey with her; then she dropped it until she could find her burrow. After that she searched on foot thoroughly over all the area within a dozen inches of her hole, only occasionally wandering in circular flights over a wider area. She seemed sure of the general region—the cinder area—but the ring of gray dust around her hole was not to be found; therefore she did not recognize it as her own even after poking her head in three times, and went back to scratch among the cinders in various points for another quarter of an hour. After she had spent a full half hour in this faithful search, I tried to make reparation by bringing a trowelful of gray dust of just the right shade and fineness and putting it nicely around the hole. As soon as I withdrew she re- turned on the Wing to continue the search, and as she came within sight of the place, she dashed straight to it and tumbled in headlong, in the old familiar manner! She remained in longer than usual this time, then passed out and in several times, making short flights of orientation about the place each time. Her faith had been severely shaken and she had to reassure herself many times. Fur- thermore, things were not yet just as they should be; my clumsy hand had spilled some cinders in her burrow, and she had to make the necessary corrections and get ac quainted with the place anew. At last, with one more careful flight of orientation, she departed for the fields to resume her foraging. These wasps keep so diligently to their nesting business Field Studies of the Non-Social Wasps 337 that I have found little else to record concerning them. They neither loiter nor dance, fraternize nor fight. Only a few times during the summer were they seen feeding quietly on the flowers of buckbrush and goldenrod near their nests. The wasps continued to work the old burrows until late in the summer, but no new ones appeared during the latter part of the season. They continued to push up dirt and carry in prey; hence I surmised that they were ex- tending the burrows and putting in additional pockets. During the week of September 12-18 they disappeared ; the earth over their holes was soon beaten down by rain and traffic. The place and manner of their death was not learned. A number of other species were seen prowling about the premises of Cerceris’ home. Their purposes or meth- ods of approach were not actually ascertained, but we know the character of some of them well enough to feel justified in holding our suspicions. A Parametopia sp. entered one hole and was captured as it emerged. In early August many Hedychrum violaceum Brulle [S. A. Rohwer] (fig. 34) were seen entering the burrows, and in two instances Lyroda sublita Say [S. A. Rohwer] in- truded. A Megachile was also seen hovering about the burrows of Cerceris, but I could not catch her in any mis- demeanor. Several specimens of the Dipteron, Exopro- Sopa fasciata Macq. [C. T. Greene] were taken as they persistently hovered above the Cerceris burrow at Jer- Seydale, Tue Bu-sue Huntress, Cerceris bicornuta Gurrty [S. A. Rohwer]. Cerceris bicornuta, like C. raui, digs holes in the earth and uses bill-bug beetles for food. The various species 338 Trans. Acad. Sci. of St. Louis of Cerceris do not all have the same habits; there is much diversity within the genus, in both nesting sites and prey eaptured. While most members of the genus nest in the ground, we have a few that make nests in twigs, e. g., C. finitima Cress., and in getting prey for the young, certain species catch weevils, Buprestid and other beetles, while others seek bees. If there are such things as plant societies, then surely we may say insect societies exist. When one compares the wasp population of our baseball diamond at St. Louis with that of the school yard at Pevely, thirty miles dis- tant, then one must surely say that certain types of areas attract certain insect societies. The two places gave us Bembia nubilipennis, Odynerus dorsale, Cerceris bicor- nuta, and one single specimen at each place of Cerceris fwmipennis. The proportion of each species in the popu- lation as a whole was likewise similar. One frequently finds the hole of this wasp closed when the digging is still in progress, but it is easily located and partially identified by a pile of loose dirt over it (fig. 35), at the center of which is usually a group of a half dozen rolls of dust, tightly packed, just as the wasp has pushed it out of the burrow. This detail indicates that her method of excavating is similar to that of C. raw. She digs and pushes the dirt up to the upper part of the channel where it forms a plug; when at intervals she pushes this whole stopper out above the ground the masses of dirt retain their sausage-like form for a time after they have been forced out, until they become thor- oughly dry and crumble to dust in the wind. Certain beetles excavate in the same way. It seems that excavat- ing and provisioning must be carried on at the same time, for often when she pushes out a pack of dirt, she throws out some of the bill-bugs as well. Four beetles were Field Studies of the Non-Social Wasps 339 found in the dust on the top of one of these nests. These weevils ‘‘play possum,’’ and one cannot always tell whether they are dead from the sting or merely in a feint, but these showed no indication of life, even when they were pinned without the usual formalities of a cyanide bottle. : These wasps dig with surprising rapidity. In one nest where the loose dust on the surface was brushed away at 7 o’clock one evening, the next morning at 9 there was a mound of earth 2 inches in diameter and 34 inch high which had been newly thrown out of the burrow. It Seemed impossible that this mound could have been thrown up since daylight; hence I suspect that the mother Verceris had been at work during the night. After all, how can they tell day from night when they are at work in the inky darkness of their underground tunnels? Rain works real hardship to these little creatures. One of my records tells of a nest in which the work was pro- gressing nicely, and each morning showed a mound of fresh dirt on the surface. A steady rain soon flattened the mound, but the plug held for several hours; a down- pour eventually washed it down, however, and the hole was flooded. The next day I watched the nest and con- cluded that the owner had been drowned, for during sev- eral hours of sunshine she did not appear. In the middle of the afternoon, however, I saw her again carrying in her eatch of bill-bugs! How she could have escaped death by drowning I cannot see. In other nests which we watched, however, the plugs were soon washed down; the occupants of most of these nests never reappeared. Three survivors tried to rebuild their homes, but were defeated by heavy rains. Thus we see that the rain here was a factor in exterminating the adult population; how the young fared in these circumstances we do not know. 340 Trans. Acad, Scr. of St. Louts ‘After the excavating is once done, the workers seem to have no objection to leaving the mouth of the burrow open. They are frequently to be seen on sunny days, with the face just within the aperture of the hole, their bright eyes shining as they gaze into the outer world. But if one comes too near, they, like C. raui, creep down, cautiously, almost imperceptibly; one does not realize that they are moving until they have vanished from sight. While Cerceris bicornuta persistently chooses for her nesting site a hard-packed soil, yet a number of times I have found evidence that she gladly avails herself of assistance in her digging by utilizing some hole already begun. In one case, one extended a neat hole left by a peg which had been driven into the ground and pulled out. In other instances they used and lengthened the old holes from which Bembiz nubilipennis had emerged (fig. 36). While they do not seem actually to follow up the Bembix or depend upon them for their assistance, they frequently have the opportunity of availing themselves of it, since their choice of the same environment often brings them together. When one finds insects established in a city lot, one is inclined to think of them as relies from a preciviliza- tion period, clinging to their former habitat despite the tightening about them of urban conditions; one is sur- prised to learn that species new to the neighborhood are coming in and becoming established under our very eyes- Intensive Hymenoptera collecting in this vicinity for eleven years had not revealed one specimen of this wasP- The small area where they are now established was dur- ing the four years, 1913 to 1916, subjected to my very intensive study; observations were made almost every¥ day as I crossed the field, yet I never found a single speci- Field Studies of the Non-Social Wasps 341 men of C. bicornuta. In 1917 I was out of the city, and the next summer, on June 30, the first specimen was cap- tured; during the following week six others were found nesting there. This shows how quickly and firmly they become established when they find a desirable environ- ment. C. bicornuta preys upon three species of bill-bugs, Sphenophorus placidus, S. zeae and S. parvulus. Bill- bugs are very destructive to cereal and forage crops, and the Cerceris wasps are one of the important natural checks. The blue-green bill-bug, 8. parvulus, is a widely distributed upland species usually infesting timothy and bluegrass, but often injuring wheat, oats, barley and rye. Pupation occurs either in the corms or in the soil. A large blue cuckoo-bee, Hedychrum violacewm, en- tered the burrow of this wasp and remained within for twenty minutes; what her errand was we can only sur- mise, Adults of this species have been found feeding on the flowers of the buckbrush. 342 Trans. Acad, Sci. of St. Louis CHAPTER II. Aw Amputator oF Sprpers’ Leas. Pseudagenia mellipes Say [S. A. Rohwer]. The genus Pseudagenia belongs to the family Psam- mocharidae, whose habits of nesting are in rather un- stable conditions, since each species shows peculiarities of its own. Natural selection has not, up to the present date, created fixed habits for the genus. There is, how- ever, one department in the work of this genus that has become rigidly established, that is the cutting off of the legs of the spiders stored as food for the young. All species whose habits have been studied show this trait, although in P. mellipes (fig. 37) it is possibly not so rigid as in other species, for in numerous cases I have found that only a part of a spider’s legs have been removed. As in the genus Trypoxylon, we find great versatility of behavior within certain species and within the genus. To Francis X. Williams* falls the credit of having brought to light some highly interesting behavior infor- mation on seven species of Pseudagenia which were stud- ied in the Philippine Islands, with the aid of which one can point out the evolution of the nesting behavior, from the simple, single mud cells of certain species to that masterpiece of economy, the work of P. mellipes. Versatility in behavior, or adaptability, seem to be the qualities which have led some individuals, at least, of P. mellipes to a method of nesting that is most economi- cal of labor, materials and time. In connection with the evolution of the nesting habits of Pseudagenia, the fol- lowing statement by Williams is indeed significant: ‘* The “Philippine Wasp Studies. Rept. Exp. Sta. Hawaiian Sugar Plant. Assoc., Bull. 14, pp. 79-108, 1919. Field Studies of the Non-Social Wasps 343 genera Macromeris, Paragenia, and Pseudagenia and others [of the Psammocharidae] are mason wasps, hav- ing advanced beyond the digging stage still adhered to by the majority of the family. They build cells of clay or other earth-like material; they may construct these in sheltered or unsheltered places above the ground, more rarely in burrows.’’ As Williams says, ‘‘having advanced beyond the digging stage,’’ to a mud-daubing cell-maker, So certain individuals of the species P. mellipes, in their cell-making, have gone a step forward in eliminating the carrying of the building materials. We find that in P. mellipes this labor-saving method of using materials at hand, or rather of finding the building materials in the Shape of a lump of mud plastered on the wall, and then fashioning it into cells, entails the habit of carrying water. This method, when first I discovered it, greatly surprised me, since I thought only the Odynerus wasps, and Anthophora bees were capable of so ingenious a method. Since then I have found this same feat done by Chalybion caeruleum, which shows at least that this habit can penetrate families regardless of structure (tax- onomy). That this water-carrying habit is more deep- Seated in the genus than the newly acquired habit of using mud nests already at hand is evidenced by the fact that Williams tells us of a species in the Philippine Islands which follows this method, showing that habit persists in spite of remoteness. Another habit that dis- tance has not obliterated is the method of using the dorsal tip of the abdomen, flexed under the head, as a rounded smoothing tool, in fashioning the nest. This habit we observed also in Pompiliodes tropicus,* and in Williams’ report he records this habit for three species of Pseudagenia, “Wasp Studies Afield, p. 54, 1918. 344 Trans. Acad. Sci. of St. Louis The stability of these two habits, in groups of or- ganisms separated from each other by half the girth of the earth, lends a tinge of romance to the subject. Shall we say that these habits are instinctive and of so strong a type that they persist regardless of time and space, or shall we rather say that these groups of insects, m the two remote regions, have independently acquired the same habits, through the similarity of their problems or through the similarity of their environment? I said that in the evolution of nest-building activities, P. mellipes is in advance of any other member of the Pseudagenia clan, despite the fact that the Philippine Pseudagenia are likewise water-carriers and likewise use the abdomen as a tool wherewith to fashion the nest. The facts are that the oriental wasps still carry the building material to the site, while some individuals of our P. mellipes have advanced to the point where the carrying proclivities are eliminated. Not all of our P. mellipes have abandoned their habit of carrying clay, for we have formerly recorded three and four-celled nests taken from under loose pieces of bark (fig. 40), and a four-celled nest that was found in an oak-apple still hanging on the tree. All of these nests in due season gave forth P. mellipes, and both situations certainly entailed the carrying of mud from a distance. Other nests which must have involved the new method of building, have been found in old nests of the pipe-organ wasp, Trypowylon politum (fig. 63), and in the abandoned nests of Sceliphron caementarium (fig. 38), where these wasps had built their little cells within the larger ones of both species. Thus we see the most recent accomplishment of the species, that of carTy- ing water, to be so new as not to have permeated the race, and the pioneers in this habit are the wasps observed at an obscure corner of the earth called Wickes. Field Studies of the Non-Nocial Wasps 345 It is fascinating indeed to think of our own little hand- ful of P. mellipes as displaying perhaps a more advanced point in their evolution than other members of even their own species, and exhibiting a type of behavior more ad- vanced than that of any other member of the genus whose habits have been recorded. In this connection we must not overlook one Philippine species, P. nyemitawa, whose unique habits are recorded by Williams. This wasp pastes its nest, consisting of two or three cells, on tree trunks, in rather exposed situations, but the structure seems proof against wind and weather, being varnished over with a tree gum. The cells are made of an earth- like substance such as is used by the termites for their Coverways. ‘‘The wasps first sip up some water from Some convenient hollow or edge of stream, and the ball of earth subsequently gathered is worked around in her mouth until it assumes the right consistency, when it is plastered on the building site with the dorsal tip of the abdomen. * * * Before the cell group of two or three cells is completed a partial coat of varnish may be put on. The cells finished are more closely united with mud. x When Pseudagenia is through with the mud she turns her attention to a tree gum which she works over in her mouth-parts and spreads on precisely as she did the mud. Then without intermission she brings in an- other final coating or rather patching material—a pale 8tay lichen—which she works up in her mouth into a Sort of viscid paste also applied like mud and varnish, but the nest is blotched rather than completely covered.” There is a small outbuilding at Wickes which harbored a number of old mud nests of Sceliphron caementarium in addition to a few nests of Polistes pallipes. For two years, 1917 and 1918 the inhabitants therein were ob- jects of study. The ecological succession of life, even in 346 Trans. Acad. Sci. of St. Louis so small a room, was as clear as it was interesting. To recapitulate the story of the occupants, the original builders, Sceliphron, had made their nests and left them to the insect community; then the succeeding inhabitants followed in peaceful succession. First after the Sceli- phron came renters by the name of Trypoxylon clava- tum. These were abundant, and used the old mud nests with only slight modification. After T. clavatwm had been watched and studied here for two years, it was suddenly discovered, when the season of 1919 opened, that clavatum was no more; that a species never here- tofore seen in this building was present in fair numbers, and had taken possession of the situation for its abode. This new citizen in the community, that had the art of making dainty little nests out of old tumble-down, twice- used mud huts, was Pseudagenia mellipes. In previous descriptions and in the paragraphs preced- ing I have already shown that P. mellipes is a fairly versatile little creature. In a former work, when we re- corded that this insect was found in the old cells of Scelt- phron (fig. 38 shows the old nests broken to expose to view the mud cells of P. mellipes), and discovered heavy mandible marks on the outside of the nest, we registered the suspicion that this was the work of P. mellipes. This suspicion was confirmed by the observation of this lot of little creatures in action. The following extracts from my notebook will give the details. At 12:50, on September 6, I noticed a one-celled nest of Sceliphron with a P. mellipes aboard. The opening was half sealed in a manner which was new to me; @ wall was being built across the aperture from the two opposite sides, leaving a vertical crack across the center about 5 mm. wide. After five minutes the opening Was smaller. My chief attention was at that time concel- Field Studies of the Non-Social Wasps 347 trated upon other insects; when again at liberty, twenty minutes later, I found the cell completely sealed, and then I ruefully realized that I had lost a rare opportunity to observe this species in action; to find whether she got the mud from elsewhere to seal the aperture, or in some way worked out the mud from various parts of the nests. For the next hour I watched the wasp busy at her work on various parts of the cell, principally at the newly made plug, in a very interesting manner. With her jaws she would scrape very energetically about the rough wall of the shanty, trying apparently to remove some of the sur- face of the wood, but in reality she was scraping together whatever she could get (often it was mostly dust and spi- der web) which she worked in her jaws on her way back to the nest a few inches away. When she reached the nest the mouthful was usually applied to the plug which had just been placed. She would press the small ball of dusty web against the plug, flex her abdomen directly under her mouth, emit a tiny globule of water apparently from the anus, and, with the flat-iron movement that we have elsewhere described for Pompiloides tropicus, Smooth and rub the material into the plug. This she did repeatedly. The rubbish she garnered was a motley mass; it was chiefly dust and spider web, with occa- Sional small splinters, bits of disintegrated insects and vegetation, which, combined with the added moisture, was worked into the nest. Not all of these reinforce- ments were directed solely to the plug, but many appli- cations were placed promiscuously about the nest. To see something entirely new in waspdom, and to See it done repeatedly, was indeed enough to hold one alert to catch every detail. To be sure, we had previ- ously discovered that a near relative, Pompiloides ‘ropicus, likewise bends the abdomen under the body 348 Trans. Acad, Sci. of St. Louis for the same purpose, and that distant relatives (Ody- nerus) carry water in their throats. These are near approaches to the method now discovered, but are not its counterpart. When I saw that this P. mellipes re- peatedly left the building by a certain crack, and that when she returned she was able to gather three pellets of reinforcing material, I formed the deduction, based largely on experience with Odynerus, that this was also a water-carrying species. Yet the puzzling ques- tion remained unanswered: If she fills her crop with water, how does it go through the body so quickly to be emitted from the opposite extremity a few minutes later? The drop of water was each time handled with such lightning speed that, I must confess, I might have been deluded; perhaps the drop came from the mouth, and the abdomen was so bent as to whip it instantly from the mouth. I craned my neck all the more; I even used my magnifying glass, but the work was done too quickly for the eye to catch the movements. For an hour while the process of reinforeing contin- ued, the little mother always went out for water through the crack nearest the nest and regularly came back through the same crack. Only once did she enter on her return through another crack a few inches toward the north. Did she go from there to her nest? No, she left mmediately by the way she had éome; presently her face appeared at the right opening and she proceeded direct to her nest and resumed her work. Place mem- ory was again demonstrated. When at last she seemed to consider her work done and failed to return, my attention was directed to the behavior of some half dozen other P. mellipes in the puild- ing, but of this nest and its contents we shall have more to say later. Field Studies of the Non-Social Wasps 349 A second P. mellipes was watched for fifteen minutes as she nervously walked on the walls or flew from place to place within the narrow confines of the shack. She was seen to enter a dozen empty cells of mud wasps, and one in particular claimed her attention. An hour’s wait, however, revealed no further activity on the part of this prospector. A third individual was seen to return often to a new and partly wet cell of S. caementarium which was in course of construction, and I am sure would have started something had not the owner, returning with her mouth- ful of mud, routed her with such hearty vigor that P. mellipes was discreet and never returned. ‘A fourth P. mellipes on an exploring expedition likewise entered several old nests but did not remain in any of them. They generally seem hard to please or for some unex- Plained reason they do a large amount of prospecting before they finally select a mud hut to remodel, although from my standpoint the offerings of available second- hand domiciles are all about equally good or bad. A fifth P. mellipes had a sixth one close at its heels as they both entered through a crack in the wall. The first one entered an unsealed cell and the other one still followed; after a few seconds, a wrangling was heard within and soon after they appeared fighting at the orifice, the victor threw the rival out, and as she fell toward the ground she expanded her wings and flew out at the door. The other remained within. Before I left an hour later, I inserted a blade of grass in the cell to ascertain if she was still present, whereupon she fled in alarm. Late that afternoon I had opportunity to return, and found her not in, but I did notice that the opening of the old cell had in the meantime been enlarged, and the jaw-marks gave evidence that some 350 Trans. Acad. Sct. of St. Louts of the clay had been bitten away from the periphery. By the next morning, more of the clay around the open- ing had been eut away, and inserting a stem of grass I sensed that some structural work had been done with- in. Late that afternoon I paused a moment just before train-time and removed the nest, incomplete as it was, to take home, and was surprised to see that more clay had been taken from about the aperture. Looking with- in, I could see at the far end a completed and open cell. Now to return to an examination of the first cell, whose closure I witnessed as recorded above. This mother had taken the unused cell and modified it in accordance with the plan illustrated in the diagram (fig. 39). In cell ‘‘A’’ the inside walls were smooth and slick, and a black spider of the species Phidippus tri- punctatus [J. H. Emerton] was within. Like many other individuals of this species observed, this wasp had failed to remove all of the legs of its prey, but those which she had taken off were amputated in a workman- like manner — that is, with a nice straight cut a few millimeters from the base. Again, she differed from others in her work in that the spider’s remaining legs were not completely paralyzed. How often we read in the text-books that the wasp stings the prey so that the body remains fresh food for the young, while the legs are paralyzed to protect the young from injury. In this spider the body seemed dead, but the remaining legs were active and clung to my pencil or finger with great tenacity. After the lapse of five days it was in about the same condition of activity as when the nest was first opened and 15 days later this spider shed its skin. This little incident may have a deeper meaning, too. It is possible that this species directs her stings to affect other parts besides the legs, for what would Field Studies of the Non-Social Wasps 351 be the utility of stinging so as to paralyze the legs, in a species which has the habit of cutting off the legs of its prey? An egg, now discolored, was sealed to the left lateroventral surface of the abdomen. Cell ‘‘B”’ was likewise smooth on the inside; all of its roughness faced into the empty cell ‘‘C’’ where it would do no harm since it was empty. In ‘‘B”’ one spider, a young Philodromus sp. [J. H. Emerton] was found. This was dead, bore a dried, shriveled egg upon its dorsum, and had only three legs. The five that had been removed were the first, third and fourth on the left side and the third and fourth on the right. All these seemed to have been cut at the joint between the coxa and the femur, leaving clean-cut stumps. ‘‘D’’ was a very large cell between the last brood cell and the plug, and con- tained sufficient space for at least three P. mellipes cells, but without taking advantage of this space she had pro- ceeded to plug up the orifice. We ought to pause here to ask how the habit was acquired of sealing up the open- ing of the tunnel or old cell when her own cells within were sufficiently sealed, since in many situations of her nest, as under bark of trees, ete., she cannot seal up the cavity °r Space containing her cells. Fig. 39 again shows the great versatility of this little creature. In this case she did not actually build the usual thimble-shaped cells, but modified her work to resemble that of Trypoxylon clavatwm (fig. 54), and actually built walls to separate the larve instead of following the more elaborate plan of building separate cups for each. In contrast with the precaution of leay- ing an air space, as seen at ‘‘D”’ in fig. 39, and sealing the opening, we see (fig. 38) where the old mud-dauber’s cells were completely filled with thimble-like cells of P. mellipes and in other cases some of them were partly 352 Trans. Acad. Sci. of St. Louis exposed to view, not having the plug at the opening. All of this shows that ‘‘habit’’ has not yet bound this creature to her wheel, and it suggests that this species’ instinct is today in a very unstable condition and will bear watching for a solution of some of the most per- plexing problems of psychic evolution. The promising prospect of wresting from P. mellipes some of the secrets of her nest-building caused me to return to Wickes. But on this second visit, in Septem- ber, the outhouse that had previously been of interest proved void of P. mellipes. At 5:35, however, almost twilight, I found in a dark corner of another shed nearby a P. mellipes atop the nest of a mud-dauber. The nest was more or less dilapidated, with whole sides torn out of certain cells. At first this condition aroused little attention, since I thought the nest had merely met with some calamity such as an attack by birds, or disintegra- tion by the elements. Later investigations, however, showed that this destruction was the work of P. mellipes; hence I give briefly the condition of the nest when found. One cell that had been sealed empty had the entire top removed; a second upper cell had likewise had the top torn away, but this had been a sealed cell, and the pupal case of the Sceliphron was hanging over one wall, where evidently it had been pushed by P. mellipes as she was demolishing the cell and carrying the mud inside for her new cells. The two cells underneath were of special interest. One had the wall about the opening removed, forming a rather large circular aperture, twice the size of the original opening. This cell will hereafter be re- ferred to as ‘‘X.’? The mandible-marks were vividly etched about the periphery. The other under cell was the main item of interest, because the P. mellipes was at the time building her own cells therein of the mud Field Studies of the Non-Social Wasps 353 which she obtained by breaking down and carrying away other parts of the old nest. When she had first begun to make her cells herein, she had evidently gotten her mud from the very same cell, for enough had been re- moved about the opening to enlarge it to two or three times its normal size and the edge was all scarred with mandible-marks. This cell is known as ‘‘H.’’ P. mellipes was on the nest at the time of its discov- ery, but left as I approached. After a lapse of nine minutes she returned, flew into cell ‘‘X,’’ presently came out and flew into her own cell ‘‘H.’’? She went in head first; the abdomen partly protruded and showed every indication that she was at work within the cell. Evidently when she entered cell ‘‘X’’ she removed a mouthful of mud from one of the walls, and within the cell which she had appropriated she was busily fashion- ing it into her own cells. The next two mouthfuls proved the correctness of this assertion, for she was next ac- tually seen biting out mouthfuls of mud from the broken cell on top and carrying this material into the cell she was reconstructing. Since these pellets were not car- ried out, it is evident that they went into her masonry. Thus she removed six loads from the wall and took them down into her cell; then she went out and returned in 12 minutes. Throughout the period of observation She often left. the nest, undoubtedly for water, and re- turned each time after an interval of about two min- utes. The light was dim in the corner of the shed under the roof where she was at work, and I could not see whether she moistened the dirt first before removing it or wet it as she was actually at work biting it out with her jaws. Of only this much I was certain: that the soil as she carried it in her mouth was wet and glistening, and very close scrutiny revealed that the spot where she got 354 Trans. Acad, Sci. of St. Louis her mouthful of dirt was always dry. Odynerus geminus was lavish in the use of water, when mouthful by mouth- ful she bit away the chimney over her nest, but this eco- nomical little creature either bit away every particle that contained water or bit out the dry dirt and disgorged upon it just enough water to make it of the right con- sistency. She came repeatedly to the wall of the nest ap- parently empty-handed, made actual mud in her jaws for short periods and then went away. Working thus, she continued to remove the foreign portions of the nest, carry the mud to a certain cell and there to fashion her own dainty little thimble-shaped cells. Often, as she worked, her hind legs extended out to view, and fre- quently I could see her curl the abdomen under the body, as previously described. Again I tried hard to deter- mine the question which had puzzled me on previous 0¢c- casions, whether the water oozes out of the tip of the abdomen when it is so inflexed or whether the drops come from the mouth and are whipped into place by the tip of the abdomen as it begins to rub down the mud, but again I was unsuccessful in the dim light. The speed of her work may be judged from the fol- lowing sample of her comings and goings, and the num- ber of mouthfuls of mud removed after each trip. She went out at 5:40 p. m., returned at 5:42; out two min- utes, then removed 8 mouthfuls of mud and worked it into her own architecture, consuming 6 minutes at the task; out again at 5:48, in at 5:49, absent one minute, removed 7 loads of mud in the same fashion in the fol- lowing 6 minutes; out for 2 minutes, then transferred 6 loads of mud to her own structure in three minutes; out again for three minutes, and in the following 5 min- utes took out 6 loads of dirt. Field Studies of the Non-Social Wasps 355 The wall-structure of this Sceliphron nest had not been reinforced in the usual way; consequently it was thin and quantities of it were required to make the com- pact cell of P. mellipes. Hence the tops of three large cells had been demolished and much additional material taken from the other two cells, to put into the making of the three stocky little cells which had been built with- in the one chosen mud-dauber’s cell. The P. mellipes builder did not each time return direct to one spot to get her mud, but walked all over the nest in a searching attitude, sometimes more than once, be- fore stopping to get her mouthful. Sometimes she went to the same place on two consecutive trips, but more often she landed at a new spot each time. ‘At about 6 0’clock the cup-shaped cell was completed very near to the opening, and the next few loads brought in were used to fill in the niches about the new cell. When she left at 6:08 and did not return at the usual time, I suspected, since her nest seemed completed, that she had gone in search of a spider. I waited until 7 p. m., when it was quite dark in the shed, but she did not return. Occa- sional visits during the two days following showed me no evidence of the return of the mother. Perhaps she had met with a tragedy; perhaps I had frightened her away by trying to observe the progress of her work by the aid of lighted matehes. The cell which she had completed before her disappearance was found to be empty ; the middle one contained a spider, Phidippus sp. [J. H. Emerton], and had a discolored egg fastened across the dorsal surface of the abdomen. The remain- ing cell, the oldest of the three, contained an almost full 8rown P. mellipes larva and a few fragments of spiders. It was replaced and on September 24 it spun its cocoon. 356 Trans. Acad, Sci. of St. Lows In the living-room of the club-house this yellow-legged Pompilid was once seen crossing the floor with her spider. When I came too near she flew to the window, but presently she returned slowly and cautiously. Dur- ing her absence I examined the spider, a young Phidip- pus [J. H. Emerton] and found that it had only one leg, a front one; the other seven had been bitten off close to the body. Thus it seems that each individual P. mel- lipes trims the legs of her spiders exactly to suit her own fancy, and not in accordance with any law or fixed instinct. She left the spider lying on its back; when she returned she mounted it without changing its posi- tion, which made the ventral side of the spider rest against the ventral side of the wasp. Her body, and that of the spider, are about the same size, so she nicely _ covered it as she trundled it to a point some three feet away. Despite the length of her legs she was unable to lift it from the floor, so it rubbed against the rough boards until I thought surely it would get a splinter in its back. She stopped alongside a loose board and with- out leaving her spider or changing her position in the slightest degree, she paused and rested for a full half hour. Sometimes, while still in the same position, she would lift her hind legs and stroke her body with them = in toilet-making. The second half hour was drag- ging wearily on; her capacity for waiting seemed to exceed mine. I blew my breath gently but directly upon her; this immediately aroused the antenne to very wild movements, but soon her agitation cooled and she quieted them. This, however, did not induce her to move, but she became quite tame, so that I could approach her near enough to see that with her mandibles she held the abdominal tip of the spider as it lay beneath her. Suc- cessive blowings at length roused her to move on with Field Studies of the Non-Social Wasps 357 it for a distance of two feet, where she lapsed again into the same loitering, sitting astride her spider. I wished to let her alone and observe her natural capacity for loafing, but I could not remain longer so took both wasp and spider. In the foregoing pages we have remarked upon the unusual versatility of this little wasp. This character- istic is generally regarded as an advantageous faculty, but we must not forget that versatility may lead to stu- pid, labor-wasting ways as well as ingenious, economical methods. Witness the following story of wasted time and energy. This P, mellipes was seen carrying a spider under her body in the usual way, i. e., venter to venter and the anal tip grasped in the wasp’s jaws, while the first pair of legs held it close to the wasp’s body. Thus the wasp walked actively about on the face of an almost perpen- dicular bank, and entered about a dozen abandoned Anthophora burrows, coming out very soon from all ex- cepting the last; there she remained for some time. I placed a test-tube over the hole to take her when she Should emerge, but when she saw the tube she darted back into the depths of the burrow, but soon again ven- tured to the orifice. Seeing that something was wrong, she went back into the hole again, got her spider and attempted to escape with it, which I permitted. After that she carried this spider in and out of more than thirty holes, always holding it in the same manner, going into one hole twice without seeming to know it, becom- ing entangled once in a spider’s web, from which she cleared herself with a few well-placed kicks, once tum- bling down with her burden from the top of the bank, but spreading her wings when half way down and re- gaining her position. Eventually, she came out of one 308 Trans. Acad, Sci. of St. Louis of the many holes empty-handed. With a probe I found the hole so shallow that the spider was easily poked out. It was legless; the legs had all been bitten off very near to the body. This is my first record of a totally de- legged spider. The victim, a Pisaurina wndata Htz. [C. H. Shoemaker], was evidently dead. For the next half hour she continued to entertain herself by going into other abandoned holes. It is possible that this par- ticular wasp had a mud cell hidden in one of the bee- burrows and, becoming confused at not finding the right cell, abandoned the prey. P. mellipes, too, has her enemies. In September, 1919, a cuckoo-bee, Tetrachrysis pattoni Aaron [S. A. Rohwer], was reared from one of its cocoons. A brief summary of the nesting habits of the mem- bers of this genus, as far as they have been reported, might be of interest here in a comparative way, although finer details of the methods of manipulation are needed before we can get much from a study of their relation- ships through this behavior. The pots of Agenia punctum* are made of mud and are shaped like oval jars, each smaller than a cherry stone. Those of A. hyalipennis affect a conoid form, narrow at the base and wider at the mouth like a primi- tive drinking cup. The nests of both species are glazed on the inner surface; this makes them waterproof. (Fabre, The Mason Wasp, p. 84, and Sharp, Insects, Pt. 2, pp. 105-106.) Agenia carbonaria contrives a nest much like a wide- mouth bottle. It appears that this insect has not learned *Many of these insects were formerly known under the generic name of Agenia, but since the name was preoccupied in the Hemiptera, he genus is now named Pseudagenia (Banks, Journ. N. Y. Ent. Soe 19:221, 1911). No attempt is made in these reviews to standardize the nomenclature, but each name is the one used by the author cited. Field Studies of the Non-Social Wasps 359 the secret of kneading its material with saliva, so its pots have not the proper permanency, and for this reason they are not placed in exposed situations. (Step, Mar- vels of Insect Life, p. 426; Sharp, Insects, Pt. 2, pp. 105-106.) Pseudagenia adjuncta makes tiny mud cells in the old cocoons left in the nests of Trypoxylon politum, and it also plasters larger mud cells to the same nests. (Rau, Journ. Anim. Behay., 6; 42, fig. 21, 23.) Agenia subcorticalis builds nests in old Pelopoeus cells, taking the dirt from the walls of the nests in which She is building, moistening the dirt with water as she works. (Hartman, Bull. Univ. Texas, 6 : 51.) Pseudagenia blanda builds stout cells of mud, cylin- drical and thick walls, ranged alongside each other, Sometimes as many as four cells in a nest; they are placed in the fissure of a vine or at the base of a tree. (Williams, loc. cit., p. 97.) Pseudagenia makilingi makes a two-celled mud nest which is hidden in a curled-up leaf. (Williams, loc. cit., p. 97.) Agenia bombycina makes clusters of little mud cells % crevices. (Peckham, Wasps Social and Solitary, p. 4.) Pseudagenia architecta makes twin cells of mud. (Rau, Wasp Studies Afield, p. 84.) These nests are probably smilar to those described by the Peckhams (Wise. Geol. & Nat. Hist. Surv., 2; 165-166), but they do not men- tion any attachment of the cells in pairs. Ashmead (Psyche 7; 66. 1896) records that the nests are thimble- Shaped, and are found under bark, logs and rocks. This brief description by Ashmead is applied also to three other species, A. bombycina, A. corticalis and A. mellipes. 360 Trans. Acad, Sci. of St. Louis Agenia variegata. This species is recorded as bur- rowing in the ground. (Riley, Am. Nat. 8; 8. 1874.) This, however, needs verification, since no other record shows similar behavior for members of this genus; Bouvier says (Psychic Life of Insects, p. 190) that A. variegata closes the entrance of her nest with balls of spider web which she compresses into a tight wad, al- though he does not state what kind of nest this species makes. Pseudagenia nyemitawa. This wasp plasters a two- or three-celled nest on trunks or logs. (Williams, loc. cit., p. 98.) Pseudagenia caerulescens makes a little mud nest within the silken retreat of a jumping spider, the web of which is usually in a crack in a bamboo stump. (Wil- liams, loc. cit., p. 100.) Pseudagenia sp. had enlarged a burrow in a honey- combed log. Within, a short tunnel was found which contained three cells, each supplied with a spider. (Wil- liams, loc. cit., p. 101.) Pseudagenia macromeroides shows indications of a semi-social habit; the mud cells are placed in groups in sheltered places. (Williams, loc. cit., p. 102.) Pseudagenia okowa is a twig-nesting wasp. She par- titions the hollows of slender twigs into 2, 3 or 4 cells, and stores them with the delegged spiders. The parti- tions are of mud, and in addition the outer one is smeared with a gummy substance (Williams, loc. cit., p. 103). Thus we see that the genus Pseudagenia, more often called Agenia, although it belongs to a family of wasps that dig burrows, comprises for the most part potter wasps that build neat pots of various shapes a¢ _ cording to the species, in various localities, with or with- Field Studies of the Non-Social Wasps 361 out garnishments, according to whether the nest is in the open or in protected situations. All of these species carry mud from afar to build their nests. Sometimes, however, some individuals of P. mellipes eliminate this work by building their nests in a daub of dry mud and carrying only the water to this point of operations, thus Saving a great amount of the work. Not all individ- uals of P. mellipes behave in this manner; this variation shows at least that this habit is new to the species and probably in an incipient stage of higher development. One must not overlook, too, that Hartman has discov- ered that Agenia subcorticalis uses the cells of Pelopoeus merely as cavities in which to build her own small cells of the ancestral type. Thus she may have as many as five of her own cells inside a single chamber of a mud- dauber’s nest. Indeed, A. subcorticalis too goes a step farther and not only closes each of her individual cells, but builds a plug over the opening to the large chamber, thus offering to her enemies an additional rampart. The dirt is taken from the very nest in which she is building her own; she gnaws off pellets, after having moistened the dirt with water from her gullet. When one sees the variations in nesting habits within each of these two species, A. Mellipes and A. subcorti- calis, and compares the behavior of the various species within the genus, one cannot help but look toward the day when naturalists will be able to show the graduated Series from the nest of most primitive structure to one of great complexity and the relationship between the nesting habits and anatomical characteristics. 362 Trans. Acad, Sci. of St. Louis CHAPTER III. Tue Grass-Carrier Wasp, Chlorion (Isodontia) auripes Fern. The behavior of this very interesting creature was incompletely told in ‘‘Wasp Studies Afield,’’ pp. 203- 206, and Trans. Acad. Sci. 25; 199-201, 1926. Since that time I have been able to add, piecemeal, several details of its life history. In 1917 we occasionally encountered the striking spectacle of a wasp flying high with a long blade of grass or strand of excelsior in its jaws. We finally traced these wasps to carpenter-bee burrows in wooden beams, where they were carrying in this mate- rial, and so ended our information on the habits of the species. ‘‘Thus the season ended without our having ascertained whether they used this material for bedding, for food, or as a plug to close the orifice.’’ With our curiosity thus aroused, we were ever on the watch for their burrows in the wood. We have previously quoted Packard, who tells how these wasps use analagous material for plugging their nests. More recently we were pleasantly surprised to find a note by F. X. Williams* which has some bearing on the phylogeny of this habit. He records that a slen- der wasp from Philippine Islands (species not recorded) “‘of the Zsodontia group, provided with slim mandibles and legs unfitted for digging, was observed gathering tomentum or wool-like material from the under side of a green leaf, and she evidently makes use of some pre- existing hollow as a nesting place, dividing the cells therein with the material gathered.’’ My first interesting discovery was a pine board in horizontal position, above a doorway in an abandoned *Philippine Wasp Studies, p. 119, 1919. Field Studies of the Non-Social Wasps 363 cabin near Kirkwood. A typical carpenter-bees’s old hole had been filled in with grass, tightly packed. The diameter of the hole at the opening was 7/16 inch (see 1 in fig. 41). This opening led to a gallery going east for a few inches (see 2 in fig. 41), then turned back sharply and went toward the west (just below 3) for 8 inches.