THE ALF.YRODIDS, OR MEALY-WINGED FLIES, OF CALI-FORNIA, WITH REFERENCES TO OTHER AMERICANSPECIES. B}^ Florence E. Be:mis,Of Stanford University, California. INTRODrCTIOX.This paper includes tlie descriptions, iisuall}' with account of eggand larval stage, of nineteen new species of North American Aleyro-didiB, or meah-winged flies, all found in California; a catalogue withreferences, food-plants, and distriV)ution of all the other Americanspecies so far described, and an analytical ke}^ of all the Americanspecies now known. This ke}' is practically that of Quaintancc (1900),expanded and modified to include the author's nineteen new species.The addition of these new forms brings the total number of speciesof American Aleyrodes, so far known, to sixt3^-six.In beginning the study of the Californian Aleyrodid}^, it was foundthat but four species in this little known family of insects had beenrecorded from this State. The accounts of these species consist solel}'of the technical specific descriptions, and are mostly included in thepapers of jVIaskell " and Quaintance.* Quaintance's Monograph enablesone to become acquainted readily with our knowledge of AmericanAleyrodid{\3 up to 1900.The insects themselves have been found to be very i)l(Mitiful, theauthor having collected them from 30 native plants (see p. -174) andfrom various cultivated plants in gardens and conservatories. The\'are so plentiful in some cases as to vie with their near relatives, theCoccids, in economic importance; the author has found the leaves ofthe native live oak {Quercus agrlfoUa), the madrofio {Arhiifiis //len-ziesli), and the sow thistle {Sonchns oleraceus)^ curled, a))normallysmall, and incrusted on the under side with the immature stages; acultivated fern kept in the laboratory and left to its own resources ?Maskell, Trans. New Zeal. Inst., 1895, p. 415. '>Quaintance, Contributions toward a Monograph of the American Alenrodidse(U. S. Agri. Dept., Division of Entomology. Technical Ser., No. 8, 1900).Proceedings U. S. National Museum, Vol. XXVII?No. 1362. 4 ) 1 472 PROCEEDINGS OF THE NATIONAL MUSEUM. vol. xxvii.was almost killed b}- the multiplication of a few meml^ers of onespecies within a year. Already these insects are being fought as apest in the conservatories. On the other hand, the author hasfrequently found colonies of Aleyrodes parasitized to an extentwhich indicates that nature herself will check the threatened danger.The chief reason why these insects might become trouljlesome econom-ically is that man}" of them are omnivorous in food habit, and, notbeing capa])le of strong flight, would, under stress, readily becomehabituated to the nearest food plants.But it is not from the point of view of the economic entomologistthat the author has l)egun the study of the Aleyrodida?. It is ratherwith the hope of throwing some light on their somewhat ambiguouszoological position through a detailed study of their structure andlife history that the work has been undertaken. The curious meta-morphosis of the Ale3n-odids is not definitely understood; whether itshould be called "complete" or "incomplete" is certainly still amooted question. In the present paper the author uses the termi-nology which is already in vogue in the literature of this group, thoughthe terms "pupa" and "pupa-case" are arbitraril}" employed whenapplied to a family in which the metamorphosis ma}" be incomplete.With this structural and developmental study as an idtimate aim theauthor has made a l)eginning in the study of the Aleyrodidse. Anacquaintance with species has necessarily 'been the first step in thework, and this paper is therefore primarily systematic in character.The geographical range covered in collecting the species hereindescribed will be Imt briefly indicated here, as the exact localities aregiven in the text for each species described. It may here be said,however, that all the collecting has been confined to California, speci-mens having been taken in the Santa Clara Valley (Santa Clara County),on the slopes of the Santa Cruz and Sierra Morcna ranges (Santa Claraand San Mateo counties), in Golden Gate Park, San Francisco, in theSan Ramon Valley (Alameda County), at the l)ase of Mount Diablo(Contra Costa County), and to a small extent in Alameda, Napa, andMendocino counties, in southern California, and in the Yosemite Val-ley. King"'s Mountain, often referred to as a collecting ground, is inSan Mateo County and Black Mountain in Santa Clara County.The immature stages may be looked for upon either surface of theleaves, appearing upon plants as dissimilar in hal>it as the plantain andthe oak. Most of the species are omnivorous, while a few seem to beconfined to a single host. It would appear from material collected atpoints scattered from the base of the coast range to its summit that inthe distribution of the Aleyrodidte in this range there are no zonesdefined by altitude, the author having found the greater number ofspecies collected, characteristic of the entire region from valley tosummit. NO. 1362. ALF.YRODIDS OF CALIFORNIA?BEMIS. 473The adults may roadily be found rcstino- on the under sides of theleaves. In colU'eting-, if the adults take to Hl.g'ht, it is only necessaryto remain (|uietly iti wait, for they usuall}' return from this upwardflight, alighting in almost the same places from which thej^ arose; theauthor has frequently tluis disturl)ed a femal(> in the act of egg-laying,and has seen her return to the same leaf when the disturbance ceased.The sure method of securing adults is, of course, to breed them frompupa' which have been carefully isolated.All the species described have been placed in the genus Aleyrodes.Where the author has described species from immature forms only (ausage adopted by systematic students of this family from the begin-ning), there was no means of definite generic identitication, as thegeneric characters lie in the wnng venation of the adult. But as allthe adults found belonged to Aleyrodes it was deemed best to place allforms examined provisionally under the one genus.The author has had to depend on the pupa-cases for the identiticationof species, there being very little specific dift'erence in th(^ adults.True, some have such distinct characters as immaculate wings and y^X-low body, but there are others with similarly unmarked wings in whichthe abdomen is yellow and the head and thorax brownish, while stillothers have wings bearing dusky spots and the l)odies with regulardark-brown markings. But as apparently identical adults issue fromwidely difl'erent pupa-cases, even the constant adult characters have nosystematic value unless correlated with the pupal characters.In the determination of the adults here described the specimens inquestion have in every case been bred b}' the author from their pupa?in the laboratory, or taken out of doors in the act of issuing from thepupa-case. In the determination of larval stages the following precau-tions have been taken for securing accurac}': In determining the begin-ning of the tirst stage the author has used onh' specimens obtained b}^capturing the larvie immediateh' upon their issuance from the eggs.In determining later stages advantage has l)een taken of the insects'ha])it of preserving the larval moults, which may be found on the dor-sum in regular succession from the first to the latest, which rests imme-diately upon the dorsum of the pupa-case. These moults have ))eenremoved, mounted in glycerin jelly or in Canada balsam, their charac-ters studied and their dimensions taken from comparison with identicalstages found on the same leaf.The following new^ species included in this paper, all from Califor-nia, are described under the following names: Alej/rodes nxtdron!, A.sjjlendois, A. q^iaintancei, A. sta?fo7'di,, A. et^rans, A. ititen'ogatioius,A. QnaKl'elli^ A. dlasemus^ A. extra/n'ens, A. merlin i, A. tn'/h/iatue, A.am?iic()h(^ ^[. jmiinostis, A. ///y/v//;.v, ^1. /'r/'dcscefix^ A. tentdoddta.s^ A.hell<)x.The Alevrodidie taken in California have ])een found t>n the follow- 474 PROCEEDINGS OF THE NATIONAL MUSEUM. vol. xxvii.ing native food plants: Rhamniis californica^ Rhainnus crocea^ Umbel-I'ularia californica^ Heteroineles arbutifolia, Quercus agrfolia^ Querciisdensi-fiora^ Clematis Ugustlcifolia^ Opulaster capitatiis^ Lonicera invo-Imcrata, Rhus diversiloha^ Primus ilicifolia^ Arbutus menziesii^ SalixIcevigata., Wasldngtonia nuda, SympJioricarpos racemosus, Ribes gluti-nosum, Ceanothus californicus, Ruhiis vItifalius, Troximmi sp., Sojichusolepaceus. Convolvulus sepinni. Convolvulus arvens is, Aesculus califar-nica, Plantago major, Solanum dauglasii, Arctostaphylas manzanita,Arcta8ta])liylas sp. (unnamed species from the Yosemite), Quercuschrysolejns, and Eriadictyon califarnicuin.The author's thanks are due to Prof. V. L. Kellogg, under whosedirection the work has been done in the entomological laboratory ofStanford University; also to Superintendent MacLaren, of GoldenGate Park of San Francisco, and to Mr. Adolph Holme, in charge ofthe conservatory of the park; also to Mr. Edward M. Ehrhorn, hor-ticultural officer of Santa Clara County, and to Mr. George A. Cole-man for specimens; also to Prof. T. D. A. Cockerell for material,helpful suggestion, and valuable criticism, and to Prof. A. L. Quaint-ance, the well-known special student of Aleyrodidse, who has mostkindly examined the entire manuscript of this paper. The authordesires to make an especial acknowledgment to Miss Mary H. Well-man, of Stanford University, wdio has made all the drawings fromnature.The cotypes of all the species described in this paper are depositedin the collections of the U. S. National Museum.LIFE HISTORY AND HABITS.In a number of species thei"e have been six stages verified, namely:Egg, three larval, a pupal, and an adult. In all the species whichhave been under oljservation, the eggs are laid in a circle or an arc ofone, one or more rows deep, and three to twenty-eight eggs have beencounted in a place; occasionalh^ they are found in an irregular group,but always close together. Each is laid singly, the female standingwith her wings somewhat outstretched and her head at the center ofthe future circle, her body forming the radius. As the eggs leave theabdomen, she raises the tip of her body above the usual level; aftereach IS deposited, she swings the posterior part of her bod}^ laterallyfor a short distance and lays another. This is kept up until oviposi-tion is completed or she is disturbed. Often several females will beseen standing near each other upon a leaf where there are no eggs;they keep moving in a restless manner, and gradually the leaf l>ecomescoated with minute, white granules of wax similar to that which isupon their bodies and wings; where there is but one insect at workthe wax is regularly circular in shape, l)ut where there are more it isirregular. Usually the eggs are found upon these places, and are N0.13G2. AtEYRODIDS OF CALIFORNIA?BEMIS. 475more or less covered with wax; they are elliptical in shape and cuincdto a greater or less degree. At first they are white or pale yellow^,but as the em])ryo develops the color becomes darker; the younghatch ill about ten to thirteen days, the egg opening along the innercurve from the apex toward the base. The pedicel, which is such anoticeable feature of the eggs of the Aleyrodidie, is a prolongation ofthe chorion, and can be seen within the body of the gravid female,frecjuently attracting attention by its dark brown color.The embryo lies with its hi^ad toward the apex of the shell, andabout the sixth day after ])eing laid the reddish eyespots and orange-colored visceral mass ])egin to show plainly. In the to the pupa-case. Specimens were seenactive for eight or more hours. One lived for fifty-three hours anddied without attaching itself. This may have been because the leaf waswithering, although when removed to another it did not atfix itself.At first the lar^';e are very convex and entirely free from secretionof any kind, but within an hour after emerging from the Qgg thebeginning of the marginal ))aiid of wax is seen. When the food wasallowed to dry somewhat so that it wilted, the young larvte secreted acoating of wax, which was not present under other circumstances.The author has not l)een able to determine the time Avhich elapsedbefore the first molt or between the successive molts. From thefact that the larval exuviiv are alwa3's on the dorsum of the succeed-ing stage, and that they are folded back with the cephalic portion ofthe ventral surface uppermost, it is almost certain that the skin breakson the ventral surface or along lateral margins. This is the reverseof the mode in the pupal stage, where the imago issues through a rentmade along the longi-dorso-medial and thoi-aco-abdominal sutures, theflaps being folded back. When a parasite emerges from a pupa-casethere is left an irregular round hole in the dorsum ot the thorax andcephalic region; this characteristic opening makes it possible to deter-mine the extent of parasitization among the empty cases. The mmi-ber of larval stages has been determined from the molts, which areuniforndy found upon the dorsum of the pupa-cases.In the first stage the larva? are always semitransparent, withfunctional legs and anteniuw and in the majority of species have fromseven to nine latero-margiiial hairs. After the lirst molt the cuticle maybegin to grow darker in color ainl thicker in texture, until the culmi- 476 PROCEEDINOS OF THE NATIONAL MVSEUM. vol. xsvii.nation is readied in the thick black pupa-case, or the color and texturema}" remain approximately^ the same as in the tirst stage; when the latterobtains, some of the structural stages can be followed by superficialexamination. The degeneration, or loss of legs and antenntv, and thepermanent disappearance of the latero-marginal hairs take place withthe molt of the first skin; in the second stage there is no externaltrace of either legs or antennae, but in the third and beginning of thefourth stage, the reduced legs and occasional!}" the minute antennajmay be made out; in the fourth stage the wings are present and thelegs and antenna? are more like those of the adult, but are still unseg-mented; in the later part of this stage they are approximately as inthe adult, and sex can be readily distinguished. The mouthparts seemto be smaller in the second and third stages; often in the latter theycan not be made out, but in the late pupal stage they occupy relativelythe same space as in the first larval.All of the pupa? secrete "honeydew,'' sometimes in such quantitiesthat the leaf around tile case and the dorsum of the pupa is coveredwith it; in some species there are seen minute, blunt tubes on theapex of the lingula, through which the fluid may be excreted (fig. 47,Plate XXXIII). When the "honeydew" is emitted the operculum islifted, the lingula is protruded, dorsally recurved, and the dropthrown with considerable force (fig. 46, Plate XXXIII). The liquidis sweet, and when exposed to the air it becomes thick and finallyhardens. The frequent appearance of fungus in and about the casesis probably induced by the presence of this medium, as it is in theCoccidge. On Chammdoreci sp., an introduced plant from Mexico whichwas kept in the Golden Gate Park Conservatory, San Francisco, theauthor saw many large, black ants busily engaged in gathering ' ' honey-dew," acting as ants do with Aphids.Leaving the pupa-case is a slow and toilsome process; the imagooften struggles for hours before it is free and ready for flight. Afterleaving the case, it usually remains quiet for a few minutes. At thistime it is paler in color than it will be later, and its wings are dampand crumpled; soon it begins to walk very slowly, and after going ashort distance crouches upon the leaf as if exhausted; gradually itswings unfold and straighten into definite shape, the color becomesvivid and the granular secretion of wax from which the family hasderived its name, begins to appear.The adults have a peculiar manner of flight; when disturbed theyrise in an almost vertical direction, and, if not further molested, alightnearly where they were in the first place. This habit may account forthe females returning to the same leaf where they were laying eggswhen disturbed, for on several occasions they have been observed toresume ^g^ laying within a short distance from the place where theirother eggs had been deposited.Although the adults possess mouthparts and an alimentary canal, No.lse^ ALEYRODIDS OF CALIFORNIA?SEMIS. 477 close observation on the part of the author has failed to reveal themfeeding; Prof. A. L. Quiiintanco, however, in a letter, writes that hehas frequentl}' observed adults of ^1. ahutllo)icii.s llaldenian feeding.Experiments for determining the average duration of adult life werenot satisfactory, as the .specimens had to be confined in an artificialenvironment that would not give results identical with natural condi-tions. From data gathered, it is probable that there are at least twobroods each 3'ear, the adults in this \icinity emerging fi'om the pupa-cases and laying in April and May, and again in September, October,and November of the same year.Family ALEYRODIDiE.Small to minute insects infesting plants; oviparous; metamorphosisincomplete (?); immature stages quiescent, attached by sucking mouthparts to the leaves; adults free and active, covered with granules ofwhite wax.Adults of both sexes with four wings, which are held horizontallyand extend beyond the abdomen when the insect is at rest. Wingsrounded, pure white or w^itli dusky spots, margins golden-yellow andserrulate or "beaded'' all around, each serrulation with three to fiveminute, delicate hairs; color and serrulations of the costal marginmore pronounced; forewings with a single, median vein; costal mar-gin bearing nine spines at base. Color of bod}^ 3^ellow; head andthorax usualh' darker; the entire body occasional!}' with conspicuousbrown markings. Head small, convex above, rounded anteriorly.Eyes two; red, brown, or l)lack; either more or less constricted nearthe middle, reniforni, or divided into two lobes, the lesser of which isanterior, In-ighter in color, and W' ith smaller facets. Anterior to eacheye is a single small ocellus. When mounted the divided eyes appearsingle, and only a careful inspection under a high power of micro-scope will show the diti'erence in structure. Antenme of seven seg-ments, the first of which is shortest and the third longest; the firsttwo segments are simple and stout; segment one. cup-shaped; segmenttwo, subpyriform; segments three to seven, inclusive, cylindrical andclosely ringed with minute hairs. Legs long and slender, feet withdimerous tsirsi, terminating in three claws, of which the middle one issmaller than the other two, and bears a number of spines. Rostrumprojecting from the under side of head, composed of a single segment,at th(? apex of which are the seta^ that form the suctorial tul)e; fromthe l)as(^ arises the long, 8-segmented mcntum, sul)cylindrical in shape,free for its entire length, and inclosing the rostral setie. Thorax withabout e