lAg84Te '^ ^-^

A REVISION OF THE NEW WORLD OF THE GENUS RHIZOECUS

(HOMOPTERA: PSEUDOCOCCmAE)

Techmcal Bulletin No. 1522 j, '? f'

-"1 ^4 > •H

0 , ■> J en :.fj

Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE

M/:-. ACKNOWLEDGMENTS

The author is greatly indebted to the following individuals and insti- tutions for the loan of type and other specimens : George W. Dekle, Bureau of Entomology, Division of Plant Industry, Florida Department of Agri- culture, Gainesville ; Roberto H. Gonzalez, Plant Protection Service, Plant Production and Protection Division, FAO, Rome; Michael Kosztarab, Department of Entomology, Virginia Polytechnic Institute and State University, Blacksburg; Raúl MacGregor, Instituto de Biología, Ciudad Universitaria, Mexico, D.F.; Sueo Nakahara, and Plant Health Inspection Service, U.S. Department of Agriculture, Beltsville, Md.; Robert O. Schuster, University of California, Davis; Hamhn H. Tippins, Division of Entomology, Experiment Station, Experiment, Ga. ; Richard F. Wilkey, -Slide Mounts, Bluff ton, Ind. ; and Douglas J. Williams, Commonwealth Institute of Entomology, British Museum (Natural His- tory), London. Special thanks are extended to Douglass R. Miller and Louise M. Russell, Systematic Entomology Laboratory, Agricultural Research Serv- ice, U.S. Department of Agriculture, Beltsville, Md., for their cooperation and assistance during these studies.

CONTENTS Page Economic importance 1 History 2 Host plant relationships 2 Terminology and distinguishing characters 3 Genus Rhizoecus Künckel d'Herculais 6 Key to the New World species of Rhizoecus 6 Descriptions of species 10 Summary 54 Literature cited 54 Index to host plants 55 Index to mealybugs 58 A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS (HOMOPTERA: PSEUDOCOCCIDAE)

By EDSON J. HAMBLETON, cooperating scientist, Systematic Entomology Laboratory, Northeastern Region, Agricultural Research Service

Rhizoecus Kiinckel d'Herculais is a cosmo- In the illustrations certain body structures politan genus occurring most commonly in are portrayed in detail. The dorsal and ventral tropical, subtropical, and warm temperate halves of the body are not shown, because areas. Distribution records for the Americas large outline illustrations of Rhizoecus lack indicate the presence of the Rhizoecus species sufficient detail to be practical in identifying from southern Canada southward into the species. West Indies and South America. By far the All illustrations were prepared with the use largest number of species is reported from of an ocular grid and squared paper. The an- Mexico, California, and Florida, where exten- tennal segments, anal ring, and claws are of sive collecting has been done in recent years. approximately the same magnification for all Elsewhere in the United States no more than species, each about X 1,200; the cephalic plate 5 species are known to occur in any of 20 ad- and rostrum, each about X 600; the circuli, ditional States. cerores, eyes, tubular ducts, and other struc- This study began when I was attempting tures are not always drawn to scale. to make a key to the Rhizoectts species of Florida. As a result of this study it was appar- Types are located in collections of the fol- ent that a reexamination of all North Ameri- lowing institutions hereafter abbreviated as can species would be desirable. Because of a shown: British Museum (Natural History), considerable amount of undescribed material, London (BM) ; California Academy of it was thought that a revision of the genus Sciences, Stanford (CAS) ; California State for the Western Hemisphere would be useful. Department of Agriculture, Sacramento Descriptions and illustrations have been (CDA) ; Florida State Collection of Arthro- prepared for 52 species, of which 12 are de- pods, Gainesville (FSCA) ; Instituto de Bio- scribed as new. Certain morphological char- logía, Ciudad Universitaria, Mexico (IBM) acters generally omitted by workers greatly University of California at Davis (UCD) facilitate the separation of species and are in- University of Georgia, Experiment (UG) cluded here. This bulletin gives a key to the National Collection of Coccoidea, U.S. Na- American species of Rhizoecus and records tional Museum, Washington, D.C. (USNM) ; their host plants and distribution. The study and Virginia Polytechnic Institute, Blacks- is based entirely on adult females. burg (VPI).

ECONOMIC IMPORTANCE

Mealybugs have long been important as fornia, has been well summarized by McKen- pests of farm crops and greenhouse, nursery, zie {1967),^ and ornamental plants. Their destructive role ^ The year in italic after an author's name indicates in American agriculture, particularly in Cali- the reference in Literature Cited, p. 54. TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

The soil-inhabiting mealybugs that feed on strawberries, and prunes in California, the the roots of plants, although less well known only State in which it has been recorded. than those feeding above ground, damage a The three most important species of Rhi- wide variety of commercially grown, subtropi- zoecns in Florida are americanus, floridanus, cal, ornamental plants. Consequently, much and simplex, all widely distributed and eco- effort has been made in California and Flor- nomically important on a variety of nursery ida to determine the presence of subterranean and greenhouse ornamentals. R. pritchardi, mealybugs, their host plants, and the extent originally known only from California, is of injury they cause. Several species belong to present in Florida, in five additional States, the genus Rhizoecus. In the United States, and in Canada. It is chiefly a greenhouse pest four of the six most injurious species of Rhi- causing serious injury to African violets. zoecus are of foreign origin. They are falcifer One of the important factors affecting coffee Künckel d'Herculais, kondonis Kuwana, ameri- production in several countries of Central canus (Hambleton), and símpfeíc (Hambleton). America and South America is directly re- The remaining two, floridanus Hambleton and lated to the feeding of Rhizoecus and related pritchardi McKenzie, are considered endemic mealybugs. Several species also have been re- to Florida and California, respectively. ported attacking other important crops, such as cacao, citrus, cotton, and bananas. Of the 15 species now recognized in Cali- Problems on the biology and control of fornia, falcifer and kondonis are the best mealybugs are complex and warrant further known. The former is a widely distributed investigation. Although some control of Rhi- pest capable of inflicting serious damage to zoecus species has resulted from applying in- many kinds of garden, nursery, and green- secticides in sprays or drenching, the nature house plants. Its distribution in the United of the habitat and lack of perti- States is probably more widespread than rec- nent data on their bionomics undoubtedly ords indicate. R, kondonis also has demon- have limited progress in developing effective strated its potential as a pest of alfalfa. control procedures.

HISTORY Cockerell {189i) described the first United Hambleton, Ripersiella, and Rhizoecus. Ferris States hypogeic mealybug of the tribe Rhizoe- (1953) synonymized Coccidella with Rhizoe- cini and named it Ripersia maritima. J. D. cus, transferred 12 North American species Tinsley (in Cockerell, 1899) transferred this from Coccidella and Ripersiella to Rhizoecus, species to Ripersiella Tinsley, and Cockerell and described 2 new species from California. (1901) described a second species, Ripersiella McKenzie (1960-67) in a series of papers leucosoma. For the next 45 years no addi- synonymized one of Ferris' species, and he tional Rhizoecini taxa were reported in North described 13 new species from Arizona, Cali- America except two introduced species, Rhi- fornia, and Mexico, 3 of which are here con- zoecus falcifer Künckel d'Herculais from sidered synonyms. Miller and McKenzie Europe and R. kondonis Kuwana from Japan. (1971) added a single new species from Mex- I (19i6a) published the first comprehensive ico. Hambleton (1973) described a new spe- paper on mealybugs of the Rhizoecus complex cies from Florida and Georgia, transferred of genera. This article treated 22 New World two species from Ripersiella to Rhizoecus, species and 12 from Africa, Asia, and Europe and (197i) synonymized Ripersiella with that were included in 3 genera, Coccidella Rhizoecus.

HOST PLANT RELATIONSHIPS The American species of Rhizoecus display lies. Records indicate a single host for a few remarkable diversity in host plant selection. species, but most injurious species, such as They are reported here from 62 plant fami- americanus, floridanus, and simplex, are A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS

polyphagous. Food plant and distribution in- ceptable families of economic importance in- formation on pest species has been greatly clude the Gesneriaceae, Leguminoseae, Rosa- expanded owing to their frequent collection ceae, Rubiaceae, Rutaceae, and Sterculiaceae. in nurseries and greenhouses. One can assume Our limited knowledge of this secluded that nursery and greenhouse conditions favor group of subterranean mealybugs is ample the survival of Rhizoecus species on many un- proof of their ability to adapt to varied ecolo- related plants. A similar pattern is indicated gical conditions. Species showing no particu- in open country in Virginia, where I have col- lar host preference would seem to indicate that lected distinctus (Hambleton) on the roots of the most critical limiting factors in their es- 10 different plants representing 6 families within a radius of 50 yards. Host acceptance tablishment would be environmental and not by falcifer and kondonis in California is simi- host selectivity. Successful dissemination re- lar in diversity to that exhibited by the spe- sulting from the movement of living plant cies just mentioned. The family Gramineae, material, soil, and plant debris doubtless will with 19 Rhizoecus species, is the most com- lead to a more widespread distribution of monly infested host family. A few other ac- these .

TERMINOLOGY AND DISTINGUISHING CHARACTERS Most of the terminology in this bulletin is The antennae are clothed with numerous the same as in my (19i6a) previous publica- elongate setae. Also present are enlarged sen- tion and some conforms to common usage. For sory setae whose size and shape are excellent clarity and uniformity, it is hoped that other characters for differentiating species. All five- workers will adopt the terminology used here. segmented antennae have five sensory setae Improper use of the terms "pore'' and **duct'' on segment V, one elongate, spinelike sensory should be corrected. Dimensions of the body seta, and four falcate sensory setae; the are given in millimeters. All other measure- smallest one is borne proximally. In the six- ments are in microns. segmented antennae the terminal segment Body Shape and Size.—Most adult females of has one spinelike sensory seta and three larger Rhizoecus are elongate oval, though some are falcate sensory setae; a smaller fifth sensory broad and others are narrow. Older specimens seta is borne distally near the margin on seg- tend to become more rotund, whereas newly ment V. The larger falcate setae may be long mature adults have not become distended and and slender, short and stout, tapered or clá- appear small when mounted. In the descrip- vate, and the other setae display similar dif- tions here the maximum and minimum lengths ferences in shape. and widths of a species represent the ranges The interantennal space varies consider- for the specimens at hand. Wide variation in ably and generally is not a satisfactory char- dimensions may exist within a species be- acter for differentiating species. tween young adult females and fully mature Eyes.—The presence or absence of eyes is an ones with distended bodies. important specific character. When present, Antennae.—Considerable variation exists in these simple structures may be obscured by the size and shape of the five- or six-segmented the antennae or hidden between folds of the geniculate antennae and in the individual seg- integument, or they may be difficult to observe ments. Differences in the antennae can often because of their small size and weak pigmen- tation. The eyes vary greatly in size and be used to characterize species. In the shorter shape. They are weakly protuberant to com- type most segments are wider than long in paratively prominent and hemispherical to contrast to the reverse condition in the longer globular. Most eyes are round or slightly oval, type. Antennal measurements were made but some are much longer than wide. along the longer margin of each segment and Rostrum.—Only the two distal segments of the divisions number I to VI. this conical structure are referred to in the TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE text. The location of its 20 setae occupy about have two circuli consistently. In some species, the same position in each species. Measure- however, the number of these structures is ments of the length and width of the rostrum variable. In R. campestris Hambleton three and the length of its loop are useful diagnostic to six circuli have been observed on the second characters. Its size and shape are fairly con- to seventh abdominal segments. The circuli stant within a species. The extension of the display much variation in size, shape, and rostral loop may be short, or it may reach to or structure. They may be conical, truncate, or slightly beyond the attachment of the second depressed with narrow or widely sclerotized pair of legs. However, in newly matured fe- rims, and the distal surface may be reticulate, males the rostrum is more robust and rounded, faveolate, or undifferentiated. and the loop appears to be longer than in older Anal Lobes.—The anal lobes may be well de- adult females. veloped, slightly protuberant, or absent. A Cephalic Plate.—This is an irregularly group of three or more elongate setae, several shaped, sclerotized plate situated on the me- auxiliary setae, and trilocular pores laterad dioventral surface of the head anterior to the of the anal ring near the body margin define mouth parts. It is taxonomically useful. The each lobe area. This area, or the lobe itself, plate sometimes has small vacuoles near its may be sclerotized. Individuals may appear to center and normally possesses several body lack sclerotization, even when present, owing setae on its periphery. It is present in most to unsatisfactory mounting procedures. The species. Intraspecific and interspecific varia- size and length of the anal-lobe setae com- tion occurs in the structure and the degree of pared to those of the anal-ring setae are good sclerotization of this plate. Differences in characters for assisting in species separation. staining techniques or condition of specimens The number of elongate lobe setae is also may render the plate difficult to observe. useful. Dorsal Ostioles.—^^These structures are not Anal Ring.—The cellular structure of the too helpful in separating species, and their anal ring is invaluable in separating species, presence sometimes is questionable. They may though heretofore little emphasis has been be heavily sclerotized, conspicuous, and sur- placed on this structure. The nature and dis- rounded by numerous pores and body setae or position of the cells—their size, form, num- weakly sclerotized and with very thin rims. ber, and arrangement—vary little within a Legs.—The size, shape, and nature of the species. In some species the cells of the outer claws and claw digitules are the most useful part of the ring possess spicules, either one to leg characters for separating the Rhizoecns a cell as in falcifer, or three to five as in dis- species. The claws vary greatly in size and tinctus. The cells of the inner part of the ring shape. The claw digitules are either long or are usually larger and more irregular in out- short. The long ones extend to or slightly be- line than the outer ones. Also characteristic yond the claws and usually are dilated api- of the genus is the innermost circle of hemi- cally ; the short ones vary in length and thick- spherical cells that make up part of the ness but seldom reach beyond the midlength darkened area. The comparative differences in of the claws. Leg spines have proven of slight length and thickness of the anal-ring setae value in separating species, and they are omit- with those of the anal-lobe setae are often ted from descriptions. helpful in species diagnosis. Circulus.—The size, shape, and structure Cerores.—The cerores are of maximum value variation of the circuli are reliable differenti- in distinguishing species. They are the most ating characters in over 50 percent of the spe- conspicuous wax-secreting structures in the cies. Normally only a single circulus is present Rhizoecini and differ from the multilocular midventrally on the fourth abdominal seg- disk pores, the triangular and the medioven- ment, but an additional smaller circulus occa- tral pores. Morphologically the ceroris is not sionally is found on the third abdominal a pore because it is composed of either two segment; specimens of a few species seem to or three ducts. The true pore in the Rhizoecini A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS

generally consists of three or more loculi and separating species. These structures when does not possess visible ducts. viewed from above appear as circular objects Bitubular or tritubular cerores are among on the derm. In profile they are readily dis- the more important structures that charac- cernible and should not be mistaken for pores. terize Rhizoecus, Their size, shape, and distri- Tubular ducts vary in size. They are narrow bution are excellent characters for distin- and elongate or short and stout, and they may guishing species. The bitubular ceroris occurs be heavily sclerotized. No distinction is made in only 10 of the 52 species in the Western between the oral-rim and oral-collar tubular Hemisphere. De Lotto (1957) and McKenzie ducts because the thickness of their rims varies (1967) used the term "unitubular pore'' for considerably. Most species, however, appear to what is actually a bitubular ceroris in genimi- have the oral-rim type of duct. In some species latus James from Kenya and disjimctus Mc- the tubular ducts are common and widely dis- Kenzie from California. The elongate ducts in tributed, in others they are scarce or absent. the bitubular cerores of these species appear Mushroom Bodies.—Under high magnifica- to be fused. The bitubular cerores may ap- tion these mushroom-shaped structures re- pear as elongate objects resembling spicules semble tubular ducts, but their exact nature with about one-half of their individual ducts has not been determined. In slide mounted projecting above the derm as in disjîinctiis, specimens they assume various positions, some or elongate and stout with wide ducts weakly of which are shown in figure 32. They meas- protruding above the derm as in mexicamis ure less than one-half the diameter of a triloc- (Hambleton) and solani (Hambleton), and ular pore. with small, elongate, strongly protruding Trilocular Pores.—These pores are present in ducts as in kondonis and menkei McKenzie. all species of Rhizoecus and are among the In the tritubular cerores the three ducts smallest and most common pores in the genus. are placed together in a whorl with their dis- Their relative abundance is of some taxo- tal extremities divaricated. These cerores may nomic significance. The trilocular pore is sub- vary in size and shape interspecifically and triangular to circular and varies in size, each intraspecifically. The walls of their individual having three loculi. These pores occur over ducts may be nearly parallel or taper gradu- the entire derm. ally toward their apices. Several species have Medioventral Pores.—These pores are larger two or three sizes of tritubular cerores— and more elongate than the trilocular pores. large, medium, and small. Where more than They appear conical and vary considerably in one size is present, the smaller one usually is size. They occur in clusters of 4 to 23, each confined to the venter and the larger to the near the medioventral line of the sixth and dorsum. In ornatus (Hambleton) the larger seventh abdominal segments. Three species cerores have short, stout ducts, which are are known to possess these pores. barely longer than wide. Circular Pores.—Appearing as small circles The cerores occur dorsally and ventrally on without depth, these pores are approximately all areas of the body, but are more numerous one-half the diameter of a trilocular pore. on the abdomen. They occur dorsally on the midabdominal seg- Multilocular Disk Pores.—The disk pores are ments in only three species—pritchardi, siib- circular and possess 5 to 12 loculi. They are cyperaUs Hambleton, and pauciportis Ham- important taxonomic characters. These pores bleton. are present in most species and normally Body Setae.—In most species of Rhizoecus occur on the venter in the area of the vulva. the body setae are rather uniformly distrib- When they are abundant, they may be re- uted, but they may be most abundant dor- stricted to the venter, or they may be widely sally on the head and along the body margins. distributed over the derm. They are usually short and slender but may Tubular Ducts.—The size, number, and dis- vary in lenprth and thickness. In a few species tribution of these ducts are important aids in the setae are good differentiating characters. TECHNICAL BULLETIN 1522, U.S. DEPT. OP AGRICULTURE

GENUS RHIZOECUS KÜNCKEL D'HERCULAIS Rhizoecus Künckel d'Herculais, 1878: 163; Hambleton, cus. The remaining 7 genera, of which 4 are 1946a: 50; Ferris, 1953: 426; Williams, 1962: 41; monotypic, include 17 species. Although our De Lotto, 1964: 381; McKenzie, 1967: 370; Ham- present knowledge of the tribe is inadequate bleton, 1973: 63; 1974: 147. Morrisonella Hambleton, 1946a: 16. and further change in its status is inevitable, Coccidella Hambleton, 1946b; 177. it is hoped that a better understanding of its Ripersiella Tinsley, in Cockerell, 1899: 278; Cockerell, known genera will be forthcoming. 1901: 165; Morrison and Morrison, 1922: 54; Ham- The genus may be characterized as follows: bleton, 1946a: 59; De Lotto, 1964: 382; Williams Antennae five- or six-segmented, geniculate, and de Boer, 1973: 241; Hambleton, 1974: 147. Type-species : Rhizoecus falcifer Künckel d'Hercu- sensory setae of terminal segments well devel- lais. oped. Anal ring usually distinct, with six setae, its cellular structure diverse. Anal-lobe I share Ferris' view on the generic struc- area usually undeveloped, with or without ture of Rhizoecus, but I am of the opinion that sclerotization and with three or more elongate the status of the genus can be strengthened setae. Derm with bitubular or tritubular by increased knowledge of characters that cerores of one or more sizes, with trilocular have not been adequately utilized in the past. pores, with or without multilocular disk pores Of major importance are the bitubular and and tubular ducts. Circuli numbering from tritubular cerores, the cellular structure of none to six, of variable size and shape. Head the anal ring, and the sensory setae of the often with an irregularly shaped ventral ce- antennae. Also, the circuli, the claws, and tu- phalic plate anterior to the mouth parts. Eyes bular ducts will provide additional means of present or absent, varying in size and shape. characterizing the genus and will assist in Legs generally spinose, claws of varying the recognition of its species. The composition of Rhizoecus is here con- lengths, digitules short and setose, or elon- sidered identical to my (1973) characteriza- gate, extending beyond tip of claws and di- tion. An effort was made to weigh the value lated apically. Body setae of variable lengths, of morphological details of all taxa in arriv- mostly short. Dorsal ostioles usually distinct, ing at a practical and useful classification. strongly sclerotized, or inconspicuous and pos- Approximately 85 percent of the described sibly absent. Body shape oval elongate to world Rhizoecini species are placed in Rhizoe- broadly ovate.

KEY TO THE NEW WORLD SPECIES OF RHIZOECUS 1. Multilocular disk pores present 2 Multilocular disk pores absent 34 2. Circuli present 3 Circuli absent 13 3. Tritubular cerores present 4 Bitubular cerores present 5 4. Antennae five-segmented californiens Ferris Antennae six-segmented sonomae McKenzie 5. Antennae five-segmented 6 Antennae six-segmented 7 6. With two circuli; anal lobes rounded, protruding, each with 8 to 10 elongate setae; tubular ducts of normal size; body setae conspicuous kondonis Kuwana With three to six circuli; anal lobes not as above, each with three to four elongate setae and several shorter body setae; tubular ducts stout, larger than trilocular pores; body setae inconspicuous campestris Hambleton 7. Digitules short, setose, not extending beyond middle of claws mcnkei McKenzie Digitules long, slender, extending to tip of claws 8 8. Eyes absent; normally with two circuli, but occasionally with one hitiihercidatiis McKenzie Eyes present; normally with one circulus - 9 A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS

9. Bitubular cerores with their ducts appearing to be fused, projecting at least 5,u-6.Li above derm disjunctus McKenzie Bitubular cerores with their ducts separated, projecting less than 5ii above derm 10 10. Circulus large, about same width as anal ring; multilocular disk pores occa- sionally present on head; rostral loop reaching to or slightly beyond insertion of second pair of legs mcvicanus (Hambleton) Circulus small, about one-half width of anal ring; multilocular disk pores confined to abdomen; rostral loop not reaching insertion of second pair of legs 11 11. Anal lobes unsclerotized; legs rather large, stout, about 290|i long; cephalic plate weakly indicated solani (Hambleton) Anal lobes sclerotized; legs moderately small, short, less than 250it long; cephalic plate strongly indicated 12 12. Falcate sensory setae rather stout, weakly clávate, claw digitules appar- ently not dilated apically totonicapanus (Hambleton) Falcate sensory setae more slender, tapering; claw digitules dilated apically gracilis McKenzie 13. Antennae five-segmented; eyes absent 14 Antennae six-segmented; eyes present 15 14. Anal lobes unsclerotized, each with five to seven elongate setae; cerores predominately large; body setae mostly long; multilocular disk pores numerous in vulvar area falcifcr Kiinckel d'Herculais Anal lobes sclerotized, each with three elongate setae and several shorter body setae; cerores predominately small; body setae short, rather sparse; multilocular disk pores scattered in vulvar area nitidalis Hambleton 15. Bitubular cerores present boharti McKenzie Tritubular cerores present 16 16. Tritubular cerores of large size only 17 Tritubular cerores of two or three sizes 20 17. Multilocular disk pores scarce, restricted to vulvar area; tubular ducts present siibcyperalis, new species Multilocular disk pores relatively abundant, widely distributed dorsally and ventrally; tubular ducts absent 18 18. With 21-25 tritubular cerores; anal-ring setae, stout, acute, averaging about 35ILI long; length of longest anal-lobe seta about equal to width of anal ring; multilocular disk pores mostly with 7 loculi spinipes (Hambleton) With 46-74 tritubular cerores; anal-ring setae slender, averaging about 60^i long; length of longest lobe seta considerably greater than width of anal ring; multilocular disk pores mostly with 10 loculi 19 19. Anal lobes protruding, each with four or five elongate setae and three or four shorter auxiliary setae; multilocular disk pores absent from head associatus (Hambleton) Anal lobes seldom showing any protrusion, each with three elongate setae and three smaller auxiliary setae; multilocular disk pores occasionally present on head distincUis (Hambleton) 20. Tritubular cerores of three sizes 21 Tritubular cerores of two sizes 22 21. Tubular ducts present; anal lobes usually distinctly sclerotized; 10-12 small tritubular cerores present ventrally on abdominal segments V-VIII; legs about 366111 long, stout americanus (Hambleton) Tubular ducts absent; anal lobes weakly sclerotized; 11-36 small tritubular cerores present ventrally on abdominal segments IV-VIII; legs about 315M' long, of medium size mayanus (Hambleton) 22. Venter of abdominal segments VI and VII with medioventral pores _ . 23 Venter of abdominal segments without medioventral pores 25 23. Anal lobes unsclerotized; 21-31 small tritubular cerores present ventrally poensis (Hambleton) Anal lobes sclerotized; four small tritubular cerores present ventrally 24 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

24. Multilocular disk pores with 6 or 7 loculi occurring ventrally, 18-25 present on abdominal segments VII-IX; eyes viewed in profile distinctly globular, constricted at their bases; falcate sensory setae long, moderately stout, tapering globocidus (Hambleton) Multilocular disk pores with 10-12 loculi occurring dorsally and ventrally, 35-40 present on abdominal segments VII-IX; eyes viewed in profile hemispherical, not constricted at their bases; falcate sensory setae comparatively short, stout theohromae (Hambleton) 25. Anal-lobe area sclerotized 26 Anal-lobe area unsclerotized 30 26. Tubular ducts absent 27 Tubular ducts present 28 27. Ventrally 10 medium-sized tritubular cerores present on abdominal segments V-VIII; approximately 60 multilocular disk pores occurring ventrally on abdominal segments VII-IX neostangei Miller and McKenzie Ventrally 44 medium-sized tritubular cerores present on abdominal segments IV-VIII; approximately 100 multilocular disk pores occurring ventrally and some dorsally on abdominal segments III-IX stangei McKenzie 28. Antennae normally about 165u, long, apical segment not twice as long as wide; 11 large tritubular cerores present dorsally latiis (Hambleton) Antennae normally at least 200f.i long, apical segment about tv^ice as long as wide; 20 or more large tritubular cerores present dorsally 29 29. Multilocular disk pores scarce, a few present near vulva; circular pores present on segments V and VI paucijjorus, new species Multilocular disk pores abundant, present dorsally and ventrally; circular pores absent graminis (Hambleton) 30. Multilocular disk pores present on both surfaces 31 Multilocular disk pores present on venter only 32 31. Body length less than 1.00 mm; rostral loop extending beyond insertion of second pair of legs; derm with less than 20 tritubular cerores neomexicanus McKenzie Body length at least 1.36 mm; rostral loop not reaching insertion of second pair of legs; derm with more than 30 tritubular cerores caladii Green 32. Tubular ducts absent; circular pores present dorsally across abdominal segments IV-VI; 11-21 multilocular disk pores present ventrally pritchardi McKenzie Tubular ducts present in small numbers; circular pores absent; 5-10 multi- locular disk pores present ventrally 33 33. Falcate sensory setae moderately stout, not tapering; rostrum about 98^1 long; cells in outer part of anal ring ovate elongate; larger cerores not surrounded by body setae nemoralis (Hambleton) Falcate sensory setae elongate, tapering; rostrum about 75¡LI long; cells in outer part of anal ring mostly sinuate; larger cerores surrounded by one to four body setae cyperalis (Hambleton) 34. Normally with two circuli 35 Normally with one circulus, rarely with an additional small one 36 35. Tritubular cerores of one size; apical segment of antennae blunt, with slender, weakly clávate falcate setae; circulus not reticulose at orifice; claws stout bicirculus McKenzie Tritubular cerores of two sizes; apical segment of antennae not blunt, with stout falcate setae; circulus with orifice reticulose; claws elongate ornatus (Hambleton) 36. Eyes absent 37 Eyes present, or sometimes absent from nakaharai 40 A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS

37. Body small, maximum body length 1.53 mm; anal lobes sclerotized; rostrum 51JX-57M- long 38 Body large, minimum body length 1.58 mm; anal lobes unsclerotized; ros- trum 72!Lt-95M' long 39 38. Anal ring about 60LI wide, its setae about 87|Lí long, outer part with 25-30 large, angular, irregularly quadrate, mostly isolated cells; orifice of circulus narrow, less than one-half its basal width floridanus Hambleton Anal ring about 45!LI wide, its setae about 55!LI long, outer part with 19-20 small, elongate, oval cells almost touching end to end; orifice of circulus wide, more than one-half its basal width tropicalis, new species 39. Tritubular cerores abundant, 125-130 present; apical segment of antennae twice as long as wide; sensory seta of segment V short, stout; tubular ducts about same size, of simple design ovatus, new species Tritubular cerores less abundant, 35-50 present; apical segment of antennae less than twice as long as wide; sensory seta of segment V narrow, elongate; tubular ducts varying in size, more complex in design relativus, new species 40. Digitules short, setose, not reaching tip of claws 41 Digitules long, usually dilated at their apices and exceeding tip of claws 42 41. Digitules less than one-half as long as claws; hind claw about 24M. long; rostrum about 69|LI long and 46^1 wide; with 50-55 tritubular cerores; lobe setae shorter and more slender than ring setae arabicus, new species Digitules at least one-half as long as claws; hind claw about 40|LI long; rostrum about 80|Lt long and ISii wide; with 35-40 tritubular cerores; lobe setae about same size as ring setae rtiaritimus (Cockerell) 42. Anal-lobe area sclerotized 43 Anal-lobe area unsclerotized 45 43. Adult female not more than 1.50 mm long; falcate setae stout, weakly clávate; rostrum at least 50^1 long favacirculus, new species Adult female more than 1.50 mm long; falcate setae not clávate; rostrum at least 75^1 long 44 44. Anal ring about 80.u in diameter, its outer part with about 44 cells in double row in places; antennal segment VI twice as long as wide, its larger fal- cate setae tapering apically; rostral loop reaching to insertion of second pair of legs inacgregori, new species Anal ring about 70,ii in diameter, its outer part with 24-28 cells in single row; antennal segment VI not twice as long as wide, its larger falcate setae not tapering apically; rostral loop not reaching insertion of second pair of legs nakaharai, new species 45. Adult female 0.87-1.40 mm long; rostrum not more than 55M. long; anal ring less than 60[i in diameter 46 Adult female 1.40-2.57 mm long; rostrum at least 65M long; anal ring more than 65M^ in diameter 48 46. Cells of outer part of anal ring subquadrate to triangulate; longest anal lobe seta not more than 60,u long insidaris, new species Cells of outer part of anal ring oval to elongate oval; longest anal lobe seta not more than 48|LI long 47 47. Cells of outer part of anal ring elongate, three to four times as long as wide; 8-13 small tritubular cerores present; falcate sensory setae narrow, elongate; rostral loop reaching about one-half distance to insertion of second pair of legs apizacos, new species Cells of outer part of anal ring oval elongate, about twice as long as wide; 40-60 small tritubular cerores present; falcate sensory setae moderately stout; rostral loop reaching insertion of second pair of legs siîïiplex (Hambleton) 10 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

48. With more than 75 tritubular cerores; rostrum at least IOOM' long; cells of outer part of anal ring mostly isolated 49 With less than 75 tritubular cerores; rostrum less than 90^1 long; cells of outer part of anal ring not isolated 50 49. Tubular ducts conspicuous, abundant, short, stout, strongly sclerotized, vari- able in size, many as large as cerores; outer part of anal ring with 48-50 cells; eyes small, rather depressed; with more than 100 tritubular cerores; rostrum about 77|u, wide polyporus, new species Tubular ducts inconspicuous, moderately abundant, elongate, neither vari- able in size nor as large as cerores; outer part of anal ring with 24-31 cells; eyes fairly prominent; with less than 100 tritubular cerores; rostrum about 98^ wide atlanticus (Hambleton) 50. Cells of outer part of anal ring mostly elongate or sinuate; with 20-23 tritubular cerores; rostral loop extending to insertion of second pair of legs chilensis, new species Cells of outer part of anal ring mostly subtriangulate or irregularly ovate; with 30-55 tritubular cerores; rostral loop not reaching insertion of second pair of legs ^1 51. Apical antennal segment usually elongate, about twice as long as wide; sensory seta of segment V weakly lanceolate; rostrum moderately elon- gate, averaging 88^ long, rostral loop extending more than one-half distance to insertion of second pair of legs; claws stout, wide at base; outer part of anal ring with 32-40 subtriangulate to quadrate cells arranged indiscriminately cacticans (Hambleton) Apical antennal segment more robust, seldom twice as long as v/ide ; sensory seta of segment V long, narrow, weakly curved; rostrum stout, broad, averaging 74M' long, rostral loop extending less than one-half distance to insertion of second pair of legs; claws elongate, narrow at base; outer part of anal ring with 22-30 elongate to irregularly ovate cells arranged more uniformly leucosomus (Cockerell)

DESCRIPTIONS OF SPECIES Rhizoecus antericanus (Hambleton) legs. Cephalic plate broader than long, its lat- eral projections usually meeting a body seta (Figs. 1-9) near each laterocephalic margin. Dorsal osti- Morrisonella americana Hambleton, 1946a: 18. oles conspicuous, their rims thickly sclero- Rhizoecus americanus: Ferris, 1953: 428; Hambleton, tized, surrounded by 8 or 9 body setae and 15- 1973: 64. 17 trilocular pores. Adult Female.—Ovate elongate. Length, 1.50- Legs moderately long, stout, average length 1.63 mm; width, 0.80-0.89 mm. Antennae six- of segments of hind pair: Trochanter, 55; fe- segmented, average length of segments : I, 41 ; mur, 121 ; tibia, 84 ; tarsus, 76 ; claw, 30 ; claw II, 20; III, 30; IV, 23; V, 21; VI, 45; apical digitules short, setose, reaching half the segment tapered, less than twice as long as length of long, slender claws. wide, with three moderately stout, weakly Circulus absent. Anal lobes slightly protrud- tapered, falcate sensory setae and one nearly ing, usually somewhat sclerotized, each lobe straight, slender, acute sensory seta; segment with one seta 70[i-90pi long, two shorter V with one smaller, narrow, elongate sensory setae, and two or three. auxiliary body setae. seta. Interantennal space equal to combined Anal ring small, approximately 40[i-45[x in length of three terminal segments. Eyes diameter, each of its setae about 56ji-58ji prominent, protuberant, occasionally longer long, slightly more slender than longest lobe than wide, lightly pigmented. Rostrum mod- seta; cellular structure fairly clear; outer erately stout, average about 75^x long, 56^x part of anal ring with 12 irregularly elongate wide; rostral loop extending beyond halfway or sinuate cells, each with 1 and sometimes 2 between insertion of first and second pair of spicules near its center; cells of inner part of A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 11

ring more obscure, seldom more than 6 ob- Florida: Altamonte Springs, 5-VIII-59, C. served, elongate oval. Tritubular cerores of 3 0. Youtsey. Bradenton, 3-IX-70, S. L. Poe. sizes, large size occurring dorsally, 5 or 6 ex- Clarcona, 9, 16-XI-70, 13, 15-IV-71, F. L. tending from head along dorsal line to ab- Ware. Delray Beach, 16-11-71, W. E. Wyles dominal segment VI, 6 submarginally along and R. A. Long. Fern Park, 27-VIII-62, C. 0. thorax and abdomen; medium-sized cerores Youtsey; 22-IV-65, J. W. McLeod; 6-18-XI- about 8 in number, occurring submarginally 70, 16-XII-70, H. M. Van Pelt; 19, 27-1-71, on venter of abdominal segments V-VIII ; 6- G. W. Dekle. Flamingo, 18-11-70, E. J. Ham- 16 small cerores confined to venter of abdomi- bleton. Gainesville, 4-VIII-71, J. Perry. nal segments V-VIII. Multilocular disk pores Grassy Key, 21-X-72, W. H. Pierce. High with 7-10 loculi, most abundant on venter of Point, 15-11-71, E. W. Miller. Largo, 4, posterior segments, elsewhere scattered ven- 16-11-71, K. C. Hickman, E. W. Miller, and trally and dorsally except on head there occur- G. T. Williams. Middle Torch Key, 20-X-72, ring only ventrally when present. Tubular W. H. Pierce. Naranja, 30-IX-72, W. H. ducts about same diameter or slightly larger Pierce. No Name Key, 3-II, 21-X-72, W. H. than trilocular pores, short, strongly sclero- Pierce. N. Fort Meyers, III-71, W. T. Walsh. tized at their bases, scattered dorsally and ven- N. Miami Beach, 18-XII-72, E. J. Hambleton. trally in small numbers over entire derm. Tri- Orlando, 9-II-71, F. L. Ware. Osprey, 5-II- locular pores fairly evenly distributed, more 71, J. R. McFarlin and C. J. Bickner. Palma abundant dorsally. Body setae moderately Sola, 19-IX-72, C. J. Bickner. Palmetto, 1- sparse, short except for few longer ones on VII, 14-X, 15-IX-70, J. R. McFarlin and C. J. head and along body margins. Bickner; 3-VIII-70, C. J. Bickner. St. Peters- Holotype.—Villavicencio, Colombia, 4-XI-44, burg, 12-11-71, K. C. Hickman. Sarasota, 22- E. J. Hambleton. Paratypes.—Twenty-three 11-71, J. R. McFarlin. Seminóle, 12-11-71, from Colombia, Cuba, Ecuador, Jamaica, K. C. Hickman. Snead Island, 11, 16-XI, 24- Trinidad. USNM. XII-70, 2-IV-71, J. R. McFarlin. Sugarloaf Host Plants.—Andropogon virgmiciis, Aralia Key, 3-II-72, W. H. Pierce. Tallevast, 17-11- sp.. Araucaria excelsa^ Areca sp., Arecasirum 71, 9-1-73, J. R. McFarlin. Upper Key Largo, romanzoffianum, Asparagus sprengeri, Cala- 22-VIII-68, R. E. Woodruff. Winter Garden, diîim sp., Calliandra haematocephala, Callian- 21-VIII-67, R. J. Griffith; 9-XII-70, G. W. dra sp., Callistemon sp., C. viminalis, Chamae- Dekle, D. Short, and S. L. Poe; 25-X-72, F. cyparis sp., Chamaedorea elegans, Chlorophy- L. Ware. turn sp., Chry salido carpus liitescens, Chrysan- British West Indies: Trinidad, 23-XI, 7, themum sp., Coccothrinax argéntala, Coffea 31-XII-43, A. H. Strickland. arabica, Collinia sp., Conocarptis erecta, Dief- Colombia: Chinchina, l-IV-56, S. E. Flan- fenbachia mactdata, D. picta, Dizygotheca ders. Fusagasugá (Cund.), 17-VIII-71, F. ele g antis sima, Eragrostis maypurensis, Erno- Mosquero and H. Martin. Villavicencio, 4- dea augusta, Euphorbia milii, Ficus nitida, XI-44, E. J. Hambleton. Gardenia sp., Gnaphalium sp., Guettarda sp., Costa Rica: Las Cruzes, 6 mi E. of San Hemigraphis reptans, Hibiscus rosa-sinensis, Vito, 18-11-70, M. Kosztarab. Hibiscus sp., Isoloma sp., Kentia sp., Kohleria Cuba: Santiago de las Vegas, l-VIII-45, sp., Lantana sp., Liriope sp., Malphighia coc- S. C. Bruner and E. J. Hambleton. cigera, Melaleuca leucadendron, Nephthytis Ecuador: Pichilingue, l-X-44, E. J. Ham- sp., Paspalum conjugatum, P. fimbriatum, bleton. Peperomia pellucida, Phoenix loureirii, Phy- salis pubescens, Pothos sp., Pyracantha coc- Honduras: La Lima, IX, X-67, C. Evers. cinea, Quercus sp., Rhaphiolepis sp., Saint- Jamaica: Interception at Hoboken, N.J., paulia ionantha, undetermined species of ll-IV-27, W. T. Ehiger. Gramineae, Orchidaceae, Urticaceae, fern. Mexico: Vera Cruz, 10 mi SW. of Jalapa, Distribution.—Florida, British West Indies, 13, 16-VII-67, D. R. Miller and J. Villanueva Mexico, Central America, South America. B. 10 mi S. of Xiutetelco, 28-11-72, D. R. 12 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE Miller and F. D. Parker. Interception at one short, slender, weakly clávate sensory Brownsville, Tex., 23-VI-49. seta. Interantennal space less than length of Panama: Canal Zone, Summit Gardens, segment I. Eyes small, protuberant, weakly 14-VII-48, E. J. Hambleton. pigmented. Rostrum short, stout, about 49j.i Puerto Rico: El Yunque, 7-II-72, E. J. long, 45[,i wide; rostral loop reaching about Hambleton. halfway between insertion of first and second Virgin Islands: St. Croix, 21-III-73, G. W. pair of legs. Cephalic plate longer than wide, Dekle and M. Murphey. length about 37ü, narrowed anteriorly, ex- Discussion.—The types and more than 200 tending to or beyond base of antennae. Dorsal additional specimens of americanus have been ostioles inconspicuous, their rims narrow, studied. This species usually has sclerotized weakly sclerotized. anal lobes and tubular ducts, structures that Legs short, moderately stout, average were reported missing in the original descrip- length of segments of hind pair: Trochanter, tion. Multilocular disk pores with 6 loculi were 25; femur, 65; tibia, 58; tarsus, 46; claw, 18; observed with the normal 10 loculi in some claw digitules slender, their tips dilated, ex- Florida specimens collected on Araucaria tending beyond weakly curved claws. excelsa.- With one conical circulus about llf^i wide, In recent years americanus has become a diameter at orifice about 5\i. Anal lobes un- pest of considerable importance in Florida developed, unsclerotized, each lobe area with nurseries. It was first collected in the State three long, slender setae, longest about 48ji at Altamonte Springs in August 1969. Al- long, and four or five shorter body setae. Anal though widely distributed in the State, it is ring clearly defined, averaging approximately unknown elsewhere in the United States ex- 45[i in diameter, its setae 50^ long, about cept an isolated infestation in New Jersey. same length as longest lobe setae but much The diverse host list and wide distribution of stouter; outer part of anal ring with 16-18 this species are indicative of its potential as irregularly sinuate cells; cells on inner part a possible greenhouse pest in other parts of of ring larger, more irregular in shape, num- the country. Costa Rica, Honduras, Panama, bering 8-10, bordered internally by clouded Puerto Rico, Mexico, and the Virgin Islands semicircular cells. Tritubular cerores small, represent new distribution records for this their ducts about 5[i-6|i long, few in num- species. ber, occurring dorsally, three on head, four Among the more important distinguishing on thorax, six or seven on abdomen. Multi- features of americanus are its three sizes of locular disk pores absent. Tubular ducts mi- tritubular cerores, small anal ring, and scle- nute, about one-half diameter of trilocular rotized anal lobes. pore, few in number, at least two each on ab- dominal segments V-VIII. Trilocular pores evenly distributed. Body setae mostly short, Rhizoecus apizacos^ new species longest on posterior abdominal segments. (Figs. 10-15) Holotype.—Tlaxcala State, 26 km N. of Api- zacos, Mexico, 12-VII-67, D. R. Miller and J. Adult Female.—Oval elongate, tapering to- Villanueva B. ; UCD. Paratypes.—Three, same w^ard extremities. Length, 0.97-1.38 mm; data as holotype; one UCD, two USNM. width, 0.41-0.74 mm. Antennae six-segmented, Host Plant.—Unknown. moderately short, stout, closely placed near Distributicn.—Known only from type locality. tip of head, segments in following propor- Discussion.—R. apizacos superficially resem- tions: I, 25; II, 11; III, 14; IV, 11; V, 13; bles simplex, but new species has larger legs, VI, 39 ; apical segment less than twice as long its antennae are more closely placed, and the as wide, with three very slender, rather falcate sensory setae are longer and more strongly curved, falcate sensory setae, sub- slender than those of simplex. Other major equal in length, and one shorter, spinelike differences separating the two species are in sensory seta near its apex; segment V with the key. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 13

Rhizoecus arabicus^ new species short, some longer setae dorsally on head and posterior abdominal segments. (Figs. 16-23) Holotype.—Chinchina, Colombia, April 1956, S. E. Flanders, with one paratype and one un- Adult Female.—Elongate oval. Length, 0.84- identified Neorhizoeciis on same slide; USNM. 1.73 mm; width, 0.46-0.87 mm. Antennae six- Paratypes.—Three, same data as holotype, two segmented, segments in following propor- partially mutilated; two Cota, Costa Rica, tions: I, 35; II, 21; III, 33; IV, 16; V, 17; VI, 20-V-57, E. B. Dixon, on two slides, one slide 45; apical segment rather stout, less than also bearing three immatures of R. campes- twice as long as wide, with three falcate sen- tris; USNM. sory setae and one more slender, shorter, Host Plants.—Coffea arabica, undetermined sharply tapered sensory seta; segment V with Gramineae, in soil. one slender, weakly lanceolate sensory seta. Distribution.—Colombia, Costa Rica. Interantennal space about equal to combined Discussion.—In the key to species, arahicus length of three terminal segments. Eyes mod- occupies the same couplet with maritimiis. It erately prominent, weakly pigmented. Ros- may readily be distinguished from the latter trum rather elongate, about 68fx long, 46fx by its shorter digitules, smaller rostrum, wide; rostral loop extending to or slightly be- larger number of tritubular cerores, and yond insertion of second pair of legs. Cephalic smaller anal ring. The cells of the outer part plate of medium size, irregularly subquadrate, of the anal ring of arahicus are mostly quad- with two vacuoles near its posterior extremity. rate, whereas those of maritimiis are mostly Dorsal ostioles indistinct, appearing as if un- oval elongate. sclerotized. Legs well developed, of moderate size, Rhizoecus associatus (Hambleton) length of segments of hind pair: Trochanter, 43 ; femur, 100 ; tibia, 95 ; tarsus, 57 ; claw, 24 ; (Figs. 24-29) claw digitules short, setose, reaching mid- Morrisonella associata Hambleton, 1946a: 20. length of claws, inner margin of claws almost RhizoecuH associatus: Ferris, 1953: 430; McKenzie, straight. 1967: 372. With one conical circulus, about as long as Adult Female.—Rather stout, elliptical, much wide, its orifice faveolate. Anal lobes undevel- narrower toward head. Length, 1.26-1.94 mm; oped, unsclerotized, each lobe area with one width, 0.58-1.21 mm. Antennae six-segmented, seta about 62|i long, two shorter elongate placed near apex of head, segments in follow- setae, one or two auxiliary setae, and usual ing proportions: I, 39; II, 25; III, 20; IV, 18; trilocular pores. Anal ring well defined, about V, 17; VI, 43; apical segment less than twice 57|.i wide, its setae about 90¡i long, stouter as long as wide, with three stout, apically ta- and longer than lobe setae; outer part of anal pered, clávate falcate sensory setae and one ring with approximately 24-30 fairly large, elongate, nearly straight sensory seta near its irregularly shaped, rather angulate, quadrate apex; segment V with one elongate, narrow to ovate cells, ends of most not touching; inner sensory seta; all segments with extremely part of anal ring with 13-14 larger, irregu- long setae of usual type. Interantennal space larly elongate cells bordered inside by clouded equal to length of segment I. Eyes small, pro- area of semicircular cells. Tritubular cerores tuberant, pigmented. Rostrum long, 75¡i-78^ of medium to small size, their ducts weakly in length, 50u wide; rostral loop extending to tapered, projecting well above derm, 50-55, or slightly beyond insertion of second pair of widely distributed dorsally and ventrally. legs. Cephalic plate large, irregularly trian- Multilocular disk pores absent. Tubular ducts gulate, 54u long, 58u wide, with four or five narrow, elongate, rather uniformly distrib- setae bordering its periphery. Dorsal ostioles uted but less common on head and thorax. weakly developed, inconspicuous. Trilocular pores most abundant on head and Legs moderately short, average length of posterior abdominal segments. Body setae segments of hind pair: Trochanter, 41; femur. 14 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE 80; tibia, 62; tarsus, 52; claw, 30; claw digi- sensory setae are clávate. The larger number tales short, setose, almost reaching midlength of anal-lobe setae and the more protruding of claws. anal lobes readily distinguish associatus from Circulas absent. Anal lobes roundly devel- distinctus, oped, heavily sclerotized, each lobe with 4 or 5 elongate setae of varying lengths, longest Rhizoecus atlanticus (Hambleton), about 80|i, 3 or 4 smaller body setae, and 8-10 new combination trilocular pores. Anal ring about 65[i in diam- (Figs. 30-39) eter, its setae about 63^i in length, stouter than lobe setae; outer part of anal ring with Ripersiella atlántica Hambleton, 1946a: 62. 12 moderately large, elongate cells, each with Adult Female.—Oval elongate, moderately several minute spicules; inner part of ring stout. Length, 1.76-2.20 mm; v^idth, 0.94-1.88 with 6-8 cells larger than those of outer part mm. Antennae six-segmented, elongate, widely and more irregular in shape, lying adjacent spaced, average length of segments : I, 41 ; II, to large darkened semicircular cells. Tritubu- 24 ; III, 29 ; IV, 19 ; V, 20 ; VI, 59 ; apical seg- lar cerores large, varying somewhat in size, ment about twice as long as wide, with three 60-74 occurring dorsally and ventrally over long, rather slender, tapering, falcate sensory entire derm. Multilocular disk pores present and one shorter, sharply tapered sensory seta ; dorsally and ventrally, occurring in irregular, segment V with 1 short, narrow, tapered sen- transverse rows, absent from head. Tubular sory seta. Interantennal space equal to com- ducts absent. Trilocular pores evenly distrib- bined length of segments II-V. Eyes fairly uted, sparse in areas on thorax and between prominent, weakly pigmented. Rostrum very segments. Body setae short, sparse, a few stout, 104|i long, slightly longer than wide; longer setae on head, posterior abdominal seg- rostral loop extending halfway to insertion of ments, and along body margin. second pair of legs. Cephalic plate small, ir- Hololype.—Guatemala, along highway be- regular in shape, with two vacuoles. Dorsal tween Quetzaltenango and Totonicapan, 10,000 ostioles almost indistinguishable. ft, 20-V-45, E. J. Hambleton. Paratypes.—Three Legs fairly large, length of segments of taken with holotype. USNM. hind pair: Trochanter, 52; femur, 118; tibia, Host Plants.—Alchemilla orbictdata, uniden- 107; tarsus, 65; claw, 27; claw digitules long, tified Gramineae. dilated at tips, extending slightly beyond tip Distribution.—California, Guatemala, Mexico. of rather stout, moderately curved claws. California: Santa Cruz, 9-II-64, D. R. Mil- With one elongate, conical circulus. Anal ler and J. A. Froebe. lobes simple, undeveloped, each lobe area with Guatemala: From the type locality. three elongate setae, two about 87fi long, Mexico: 10 mi S. of Texcoco, State of Mex- longer and stouter than third, and four or five ico, 12-VIII-67, D. R. Miller and J. Villanueva smaller auxiliary setae. Anal ring large, 78!i- B. 8511 in diameter, each of its setae shorter and Discussion.—Ferris (1953) doubted that somewhat narrower than largest lobe seta; associatus differed from distinctus. His key outer part of anal ring with 24-31 mostly characters for separating the two species are isolated, irregularly oval to almost circular unreliable because the inner row of "pores" of cells, varying in size, each with tiny spicule; the anal ring appears clouded in all species of cells of inner part of ring approximately 22 Rhizoecus. Staining techniques possibly alter in number, larger, many touching; clouded the degree of cloudiness in some specimens. area indistinct except for large oval cell struc- McKenzie (1967) y using the identical char- ture. Tritubular cerores small, their ducts acters in his key, likewise overlooked the im- long, slender, tapering, and divaricating, pro- portant differences between these mealybugs. jecting well above derm, about 95-100 present The basic differences separating the two spe- dorsally and ventrally in transverse series en- cies are in the key. Also, associatus has longer circling segments. Multilocular disk pores appendages than distinctus and its falcate absent. Tubular ducts fairly abundant, occur- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 15 ring over most of derm. Mushroomlike bodies, each lobe area with three elongate setae, similar to those found in polyporiis, widely subequal in length, longest about 34¡.i long, distributed over derm. Trilocular pores mod- and several shorter body setae. Anal ring erately abundant, evenly distributed. Body about 401-1 wide, its setae about 34ii long, setae longer on head, elsewhere varying in stouter than lobe setae; outer part of anal length, uniformly distributed. ring with 18 elongate oval cells, varying in Halotype.—Guarujá, State of Sao Paulo, Bra- size, mostly with spicules; inner anal ring zil, 22-VIII-35, B. L. Ribeiro and E. J. Hamble- with 8-10 larger, more irregularly elongate ton ; on slide with four paratypes. Paratypes.— cells, lying adjacent to rather indistinct Ten on two slides. USNM. clouded area. Tritubular cerores small, their Host Plant.—Axonopus sp. ducts short, 20-24 present, most occurring Distribution.—Known only from the type lo- dorsally. Multilocular disk pores absent. Tu- cality. bular ducts smaller in diameter than trilocu- Discussion.—No other known species of lar pore, present dorsally and ventrally, scat- Rhizoecus resembles atlanticus. Its large, tered on abdomen, sparse on thorax, absent broad rostrum, finely tapered ceroris ducts, from head. Trilocular pores rather sparsely and unusual cellular structure of the anal ring distributed over entire derm. Body setae set it apart from all other species. This mealy- mostly short, slender. bug was originally collected along the Atlantic Holotype.—Trinity County, 6 mi NE. of seacoast with R. nitidalis and Brevicoccus Weaverville, Stony Creek on Lake Trinity, clavisetosus Hambleton. Calif., 22-V-63, R. F. Wilkey; UCD. Para- types.—Two, one USNM. Rhizoecus bicirciilus McKenzie Host Plants.—Ceanothiis piimilus, Eriogo- mim sp., Lotus scopariiis, unidentified Grami- (Figs. 40-45) neae. Rhizoecus bicirculus McKenzie, 1967: 374. Distribution.—Type locality and the follow- ing records: California: Merced County, 18 Adult Female.—Elongate oval. Length, 1.10- mi W. Los Banos, 17-11-67, T. R. Haig. Santa 2.55 mm; width, 0.40-1.26 mm. Antennae six- Barbara County, South Ridge Station, Santa segmented, short, stout, weakly clávate, length Cruz Island, 6-V-68, D. R. Miller and R. W. of segments : I, 21 ; II, 12 ; III, 17 ; IV, 10 ; V, Rust. South Rim Station, Santa Cruz Island, 11; VI, 28; apical segment broad, with three 9-V-68, D. R. Miller. fairly stout, clávate, falcate sensory setae and Discussion.—This unusual species is easily one narrow, elongate sensory seta; segment recognized by its two large circuli, its short, V with very short, clávate sensory seta. Inter- stout antennae, and small rostrum. Contrary antennal space less than width of segment I. to McKenzie's belief, bicirculus is not closely Eyes small, inconspicuous. Rostrum small, related to cacticans. The two species are not about 42(1 long, 28[i wide; rostral loop not similar morphologically. reaching halfway to insertion of second pair of legs. Cephalic plate longer than wide, about Rhizoecus hituherculatus McKenzie 28^ long. Dorsal ostioles small, weakly scle- rotized. (Figs. 46-54) Legs short, stout, length of segments of Rhizoecus hituherculatus McKenzie, 1960b • 743- 1967- hind pair: Trochanter, 24; femur, 54; tibia, 376. 47; tarsus, 30; claw, 12; claw digitules long, Adult Female.—Oval elongate. Length, 0.82- dilated at their tips, exceeding basally stout, 1.20 mm; width, 0.34-0.40 mm. Antennae six- weakly curved claws. segmented, moderately stout, segments in fol- With two depressed circuli situated ven- lowing proportions: I, 33; II, 19; III, 31; IV, trally on segments III and IV, measuring 14; V, 17; VI, 44; apical segment tapered, lOfi-13^ in diameter, their circular rims twice as long as wide, with three fairly stout, strongly sclerotized. Anal lobes undeveloped. weakly tapered, falcate sensory setae and one 16 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE elongate, spinelike sensory seta; segment V and R. 0. Schuster. Three mi N. of Placer- with one shorter, weakly lanceolate sensory ville, 18-V-61, R. 0. Schuster. seta. Interantennal space equal to length of Indiana: Lawrence County, Spring Mill segment I. Eyes absent. Rostrum about 65u State Park, 7-XI-71, J. A. McBride. (In rot- long, 45LI wide; rostral loop extending to or ting wood and soil, via Berlese trap.) beyond insertion of second pair of legs. Ce- North Carolina: Duke University Forest, phalic plate about 38u long, narrower anteri- 24-11-45, A. S. Pearse. orly. Dorsal ostioles well developed, strongly Discussion.—This small native species is one sclerotized, posterior pair surrounded by con- of the few with bitubular cerores. The pres- centration of trilocular pores and body setae. ence of a circulus, the bitubular cerores, and Legs relatively short, stout, length of seg- lack of eyes distinguish bituberadatKS from ments of hind pair: Trochanter, 35; femur, its nearest relatives. The multilocular disk 77; tibia, 66; tarsus, 50; claw, 17; claw digi- pores of this species display a considerable tules weakly dilated apically, extending be- difference in size and their loculi vary from 8 yond claws. to 12 instead of from 10 to 12, the number in With one faveolate, depressed, conical cir- most species. Tubular ducts, reported to be culus, its orifice about llj^i in diameter. Anal absent in the original description, are present lobes weakly developed, each with small scle- in bituberculatus. rotized patch, three elongate setae, subequal in length, longest approximately 52u long. Rhizoecus boharti McKenzie Anal ring well defined, diameter 45[.i, its setae about 6S[i long, slightly weaker than (Figs. 55-61) strongest lobe setae; outer part of anal ring Rhizoecus holiarti McKenzie, 1960a: 139. with approximately 14-16 oval, elongate cells of varying lengths, each with spicule; inner Adult Female.—Elongate oval, narrower to- part of ring with smaller number of larger, ward apex. Length, 1.05 mm; width, 1.00 mm. more irregular cells lying adjacent to cellular Antennae six-segmented, relatively small, darkened area. Bitubular cerores small, elon- length of segments: I, 33; II, 22; III, 26; IV, gate, relatively stout, their ducts projecting 17; V, 19; VI, 41; apical segment about twice well above derm, 45-55 occurring dorsally and as long as wide, with three elongate, falcate ventrally. Multilocular disk pores more abun- sensory setae and one elongate, spinelike sen- dant ventrally in abdominal and thoracic sory seta near apex; segment V with one small, areas, scattered and in small groups, dorsally short, rather narrow sensory seta. Inter- limited to posterior abdominal area. Tubular antennal space equal to length of segment I. ducts scattered dorsally and ventrally. Triloc- Eyes small, protruding. Rostrum elongate, ular pores uniformly distributed dorsally, on 66j^i long, 36|.i wide; rostral loop reaching to venter in patchlike areas. Body setae varying insertion of second pair of legs. Cephalic plate in length, rather sparse, usually occurring broadly rounded, weakly pigmented. Dorsal with trilocular pores. ostioles well developed, their rims strongly Holotype.—Placer County, 4 mi W. of New- sclerotized. castle, Calif., 21-III-58, W. H. Lange, L. M. Legs comparatively small, length of hind Smith, and R. 0. Schuster; UCD. Paratypes.— leg segments : Trochanter, 29 ; femur, 72 ; Many, same location as holotype, 15-IV-58, tibia, 64; tarsus, 44; claw, 17; claw digitules 10-III-59, L. M. Smith and R. 0. Schuster; elongate, their tips weakly dilated, exceeding UCD, two USNM. moderately stout claws. Host Plant.—Unknown, taken in soil under Circulus absent. Anal lobes undeveloped, QuercMS wislizenii. each lobe area with sclerotized patch, three Distribution.—California, Indiana, North Car- elongate setae, longest about 42j^i long, and olina. several short body setae. Anal ring 45fi wide, California: Type locality and near Nash- its setae about 44|.i long, slightly longer and ville, El Dorado County, 25-1-58, L. M. Smith stouter than lobe setae ; outer part of ring with A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 17

16 oval, elongate cells with spicules; inner rather long, averaging about 88|i long, 65\i part of ring with 10-12 larger, more irregular wide; rostral loop extending to or slightly be- cells lying adjacent to diffused shaded area. yond halfway to insertion of second pair of Bitubular cerores small, their ducts elongate, legs. Cephalic plate irregularly triangulate, weakly tapered, about 55 present, occurring 35[i wide at base, with two large vacuoles. dorsally and ventrally. Multilocular disk pores Dorsal ostioles inconspicuous, their rims present, confined almost entirely to posterior lightly pigmented. abdominal segments, dorsally only two pores Legs of normal size, moderately stout, elon- present on segment VIII. Tubular ducts short, gate, length of segments of hind pair: Troch- smaller in diameter than trilocular pore, dis- anter, 51; femur, 124; tibia, 104; tarsus, 62; tributed over entire surface. Trilocular pores claw, 20; claw digitules elongate, weakly di- distributed dorsally and ventrally, fewer pres- lated at tips, extending beyond stout, curved ent between segments. Body setae mostly claws. short, slender. With one strongly sclerotized, conical cir- Holotype.—Cochise County, 5 mi SE. of culus, about as long as wide. Anal lobes un- Apache, Ariz., ll-VIII-58, R. M. Bohart; developed, unsclerotized, each lobe area with UCD. three elongate setae, one measuring 75^-95u Host Plant.—Unknown, collected in soil via long, remaining two shorter, more slender, Berlese trap. and not as long as anal-ring setae. Anal ring Distribution..—Known only from type local- approximately 75ji in diameter, its cellular ity. ^ structure clear, distinct, ring setae ranging Discussion.—This rather unique species with between 92|i and 105¡i long; outer part of anal six-segmented antennae and bitubular cerores ring with 32-40 subtriangulate to quadrate, has no circulus. These features alone separate indiscriminately arranged cells of large size; it at once from all known species of Rhizoeciis cells of inner part 20-22, somewhat larger in the Western Hemisphere. than those of outer part, irregular in shape, lying end to end, bordering clouded area of Rhizoecus cacticans (Hambleton) semicircular cells. Tritubular cerores small, (Figs. 62-69) between 50 and 55 present, occurring dorsally and ventrally, usually 4 or 5 to segment, their Ripersiella cacticans Hambleton, 1946a: 64. individual ducts not reaching far above derm. Rhizoecus cacticans: Ferris, 1953: 432; McKenzie, 1960b: 745; McKenzie, 1967: 379; Hambleton, Multilocular disk pores absent. Tubular ducts 1973: 65. minute, of smaller diameter than trilocular Rhizoecus epiphylli Ferris, 1953: 442; McKenzie, pore, distributed over entire derm but more 1960b: 745. prominent on abdomen. Trilocular pores nu- Rhizoecus leucosomus (Cockerell) : McKenzie, 1967: merous, evenly distributed. Body setae 20!.i- 398 (misidentification in part) ; Gonzalez and 55n long, rather uniformly arranged. Charlin, 1968: 112 (misidentification). Holotype.—Cayambe, Ecuador, ll-X-44, E. J. Adult Female.—Oval elongate, stout. Length, Hambleton. Paratypes.—Twenty-three from 2.25-2.57 mm; width, 0.98-1.18 mm. Anten- Argentina, California, Ecuador, and Germany. nae six-segmented, rather elongate, widely USNM. spaced, average length of segments: I, 39; II, Host Plants —Bromiis unioloides, Cyperiis 25; III, 45; IV, 18; V, 20; VI, 55; apical seg- sp., Ditdleya farinosa, Echeveria sp., Eleusine ment usually twice as long as wide, with three indica, Epiphylkmi sp., Holcus lanatus, Kal- rather narrow, elongate, falcate sensory setae anchce SD., K. tomentosa, Kleinia sp., Lobivia and one more slender, strongly tapered sen- shaferi, Lobivia sp., Lolium perenne, Mammil- sory seta; segment V with one shorter, nar- laria sp., Mesembryantliemuni sp.. Opuntia row, elongate sensory seta. Interantennal sp., Sempervivuni sp., S. tectoriim. Senecio space equal to approximately length of seg- mikanioides, undetermined Cactaceae. ments IV-VI taken together. Eyes moder- Distribution.—California, Florida, Argentina, ately large, protuberant, pigmented. Rostrum Chile, Ecuador, Honduras. 18 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

California: Berkeley, 7-VI-48, C. A. Han- Rhizoecus caladii Green sen. Dale City, 18-1-57, D. J. Bingham and E. L. Labadie. Davis, 11-11-65, W. Russell; (Figs. 70-71) 24-11-65, Mrs. W. A. Williams. Fallbrook, Rhizoecus caladii Green, 1933: 53. 11-III-57, K. F. Sims. Fontana, l-V-34, E. P. Bradbury. Leucadia, 8-XII-55, D. Sprague. Adult Female.—Oval elongate, moderately Niles, 25-11-48, A. E. Pritchard. Ontario, stout. Length, 1.36-1.65 mm; width, 0.74-0.91 26-XII-48, C. R. Tower. Pacifica, 8-II-64, mm. Antennae six-segmented, average length D. R. Miller and J. A. Froebe. Santa Cruz, of segments: I, 38; II, 21; III, 33; IV, 28; V, 26-III-64, R. W. Nutter. Watsonville, 29-XI- 22 ; VI, 48 ; apical segment longest, with three 57, D. H. Shaw. rather long, slender, gradually tapering, fal- Florida: St. Petersburg, 9-II-73, C. K. cate sensory setae and one slender, spinelike Hickman. Sanford, 15-11-63, C. 0. Youtsey. sensory seta near its apex; segment V with Argentina : Locality unknown, intercepted one short, slender falcate seta. Interantennal at Washington, D.C., 23-XI-36, D. P. Limber space about equal to combined length of seg- and W. B. Wood. ments V and VL Eyes protuberant, lightly Chile: Cautín Province, 1 km N. Villarrica, pigmented. Rostrum elongate, about 78|i long, 29-IV-65, R. H. Gonzalez. Cuesta de Puca- 5l!i wide ; rostral loop not reaching insertion lan. La Calera, 1961, L. M. Smith. La Cruz, of second pair of legs. Cephalic plate not ob- 4, 15-IV-61, L. M. Smith. One km E. Maiten- served. Dorsal ostioles clearly defined, their cillo, 16-III-61, L. M. Smith. Olmue, 21-VI- rims sclerotized, surrounded by numerous tri- 61, L. M. Smith. Valparaiso Province, Que- locular pores and body setae. brado El Soldado, El Cobre, 15-VII-61, L. M. Legs fairly large, average length of seg- Smith. Jardim Botánica Nacional, Vina del ments of hind pair: Trochanter, 55; femur, Mar, 16-VI-61, 4-VII-61, L. M. Smith; 15- 123; tibia, 88; tarsus, 76; claw, 33; claw digi- VI-65, R. H. Gonzalez. tules short, setose, not reaching one-half Ecuador: Alausi, 7-X-44, Cayambe; 11-X- length of narrov/, elongate, finely pointed 44, Salcedo; 4-X-44, E. J. Hambleton. claws. Honduras: La Lima, X-67, C. Evers. Circulus absent. Anal lobes very weakly Discussion.—This species has been confused protruding, each lobe area with one seta 78ii with leiLcosomiis, floridamis, and the new spe- long and two shorter setae. Anal ring about cies chilensis and nakaharai described herein. 45u in diameter, its structure poorly defined, The misidentification of cacticans is believed cells appearing elongate; its setae 50ji-60}i to have been due primarily to overlooking de- long. Tritubular cerores of 2 sizes, their ducts tailed morphological structures. Characters short, stout, about 20 of larger size occurring of major importance in separating cacticans dorsally, 6 extending along median line from from its nearest relatives are its unsclerotized head to abdominal segment VI, 4 submargin- anal lobes, elongate rostrum, length of apical ally on abdomen; small cerores occurring ven- antennal segment, and the number, shape, trally, total of 14 present submarginally and and size of the cells in the outer part of the near median area of abdominal segments V- anal ring. In floridamis and nakaharai the VIII. Multilccular disk pores present ven- anal lobes have rather small, elongate sclero- trally, moderately crowded near vulva, scat- tized areas. Several differences separate leit- tered elsewhere on abdomen and thorax, absent cosomiis and cacticans. The former possesses from head. Tubular ducts small, scarce, few a shorter and stouter rostrum, less elongate near abdominal margins. Trilocular pores apical antennal segment, and fewer elongate, rather evenly distributed. Body setae short, irregularly ovate cells in the outer part of the sparse, inconspicuous. anal ring. These cells are also smaller and Topotypes.—Three, Surinam, Dutch Guiana, more uniformly arranged in lencosomns. lO-X-31, G. Bunzli; BM. Other characters may be referred to in the Host Plants.— Caladiiim bicolor, Coffea arab- key. ica. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 19

Distribution.—Known only from tvpe local- lobe seta and usually about as long as latter; ity. ^ outer part of anal ring with 14-16 small, uni- Discussion.—Redescribed from three topo- form, elongate, oval cells, each with spicule; types determined by F. Laing. It is not known inner ring with 10 or 12 larger, irregularly if the three topotype specimens were seen by elongate cells bordered inside by darkened Green. I am convinced nevertheless that they area of semicircular cell structure. Tritubular are conspecific with Green's type specimen of cerores of 2 sizes present, their ducts short caladii, and stout, smaller size less numerous, more R. caladii resembles jooensis from Colombia common ventrally, total number of cerores but differs from it in lacking clusters of medio- ranging between 130 and 200, situated mostly ventral pores and by possessing shorter and in rows across segments and scattered on less conspicuous body setae and narrower head. Multilocular disk pores present, confined claws. to posterior abdominal segments ventrally, from 27 to 45 distributed along margins of Rhizoecus californicus Ferris segment VII, around vulva, and across seg- ment IX. Tubular ducts rather elongate, with (Figs. 72-80) broadly rounded sclerotized bases, distributed Rhizoecus californicus Ferris, 1953: 434; McKenzie, on both surfaces, as many as four on each 1967: 381. abdominal segment. Trilocular pores fairly evenly distributed, less common in thoracic Adult Female.—Elongate oval, moderately area. Body setae sparse, mostly short, slender. stout. Length, 1.20-1.50 mm; width, 0.75-1.00 Lectotype.—Berkeley, Calif., 23-IV-46, H. M. mm. Antennae five-segmented, weakly clá- Butterfield, with five paralectotypes on one vate, average length of segments: I, 22; II, slide, E. 0. Essig collection, UCD. 24; III, 19; IV, 17; V, 60; apical segment Host Plants.—Artemisia donglasiana, Dipsa- more than twice as long as wide, with four cus sp.. Fragaria chiloensis, Heliaiitlms sp., falcate sensory setae of medium size, longest Polygomim sp., in soil under chaparral, in about equal to width of segment V, and one grassland. narrow, spinelike sensory seta. Interantennal Distribution.—California. space less than length of segment V. Eyes California: Berkeley, 2-III-58, L. M. Smith. usually inconspicuous, weakly pigmented, One mi W. Bolinas, 19-1-60, W. G. Iltis and often difficult to observe. Rostrum short, aver- C. Judson. Two mi NE. Bolinas, 27-VIII-63, age length 55fi, width 42|í; rostral loop D. R. Miller. Davis, 23-IX-61, M. E. Irwin. rather short, seldom extending to insertion of Goat Rock, lO-X-67, H. L. McKenzie. Guer- second pair of legs. Cephalic plate broader neyville, 30-V-60, C. Judson. Millbrae, San than long, about 50|i wide. Dorsal ostioles Andres Lake, XI-57, D. W. Price. Palermo, prominent, their rims heavily sclerotized. 23-X-70, E. Remmurs. Six mi E. Point Legs stout, exhibiting some variation in Reyes, 1-III-60, L. M. Smith and R. 0. size, hind pair measurements of largest speci- Schuster. men: Trochanter, 44; femur, 92; tibia, 83; Discussion.—The original description of cali- tarsus, 55; claw, 22; claw digitules long, di- fornicus by Ferris {1953) was based on a col- lated at tips, extending beyond stout, curved, lection of six female specimens mounted on a acute claws. single slide. Since no type was designated, I With one rather small, truncate to dome- have designated the specimen on the right shaped circulus with reticulated oral surface. side of the slide, nearest the label to the right, Anal lobes showing some development, each as the lectotype. with strongly sclerotized area equal in size to In addition to the types, more than 28 spec- width of anal ring, with three elongate setae, imens of this interesting California species subequal in length and size, and several body have been studied. It is readily distinguished setae. Anal ring about 40|i in diameter, each bv having five-segmented antennae, an aver- of its setae slightly more slender than largest age of over 150 tritubular cerores of two 20 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE sizes, multilocular disk pores, and a circulus. dium size, with elongate, weakly pigmented The heavily sclerotized anal lobes are also a ducts whose external bases are surrounded by good diagnostic character. Although con- oval-shaped sclerotized collar, occurring dor- sidered absent in the original description, tu- sally, 1 on head, 5 on thorax, 14 on abdomen bular ducts are present in calif omictis. It may along middorsal line and submarginally. also be noted that the eyes are so weakly pig- Multilocular disk pores confined to venter, mented that they often are not observed. some 40-48 present on abdominal segments VII-IX. Tubular ducts unusually large, about Rhizoecus campestris Hambleton 6|i long, rather flask-shaped, their diameter exceeding that of trilocular pore, scattered (Figs. 81-88) over entire derm. Trilocular pores fairly evenly distributed except on thorax and between ab- Rhizoecus cam'pestris Hambleton, 1946a: 51; Ferris, 1953: 436. dominal segments. Body setae slender, short to medium in length, distributed in same Adult Female.—Elongate oval. Length, 1.30- manner as pores. 1.53 mm; w^idth, 0.46-0.60 mm. Antennae Holotype.—Retalhuleu, Guatemala, 16-V-45, strongly geniculate, clávate, five-segmented, E. J. Hambleton. Paratypes.—Three, taken segments measuring as follows: I, 43; II, 22; with holotype. USNM. III, 19; IV, 13; V, 69; apical segment more Host Plants.—Coj^^ea arabica, unidentified than tv^ice as long as wide, with three stout, Compositae. weakly clávate sensory setae, one shorter clá- Distributioiio—Costa Rica, El Salvador, vate seta near its base, and one elongate spine- Guatemala. like sensory seta near apex. Interantennal Costa Rica: Coto, 13-IV, 20-V-57, E. B. space less than length of apical segment. Eyes Dixon. absent. Rostrum small, average length about El Salvador: Two mi S. Quetzaltepeque, 17- 58}!, average width 40^x; rostral loop extend- VII-61, M. E. Irwin. ing to insertion of second pair of legs. Ce- Guatemala: Retalhuleu, 16-V-45, E. J. phalic plate irregular in shape, about 32[.i Hambleton. San Rafael Pie de la Cuesta, De- long, wider posteriorly, sometimes with sev- partamento de San Marcos, December 1963, eral vacuoles. Dorsal ostioles conspicuous, N. Jesus Escobar. their rims well sclerotized. Discussion.—R. campestris is one of the few Legs of medium size, length of segments of species with multiple circuli and five-segmented hind pair: Trochanter, 42; femur, 103; tibia, antennae. Perhaps it is more closely allied to 77; tarsus, 56; claw, 20; claw digitules slen- kondonis than to any other species. It is dis- der, acute, reaching about half the length of tinguished from the latter by its smaller size, elongate, moderately stout claws. shorter and less abundant body setae, fewer With two to six truncate circuli, varying in multilocular disk pores, and particularly by size and occurring along venter from abdomi- its poorly developed anal lobes, each with 3-4 nal segment II to VII. Anal lobes weakly de- elongate setae instead of 8-10 as in kondonis, veloped, unsclerotized, each lobe area with three to four elongate setae, longest sometimes Rhizoecus chilensis^ new species 82[i in length, four to five shorter auxiliary setae, number of trilocular pores with some (Figs. 89-93) adjacent to bases of larger setae. Anal ring about 40[i in diameter, its setae 62pi long, more Rhizoecus leucosomus (Cockerell) : Gonzalez and Char- lin, 1968: 112 (misidentification). slender than largest lobe seta; outer part of anal ring with 12-14 narrow, elongate cells, Adult Female.—Elongate ovate, narrower to- each with elongate spicule; inner part of ring ward head. Length, 1.07-1.43 mm; width, 0.29- with 6-10 larger, more elongate, and irregu- 0.58 mm. Antennae six-segmented, closely lar cells lying adjacent to darkened area of spaced, average length of segments: I, 34; II, semicircular cells. Bitubular cerores of me- 18; III, 28; IV, 15; V, 18; VI, 44; apical seg- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 21

ment not twice as long as wide, with three sembles leucosomus but may be separated slender, falcate sensory setae and one shorter, from it by its smaller size, narrower body, spinelike sensory seta; segment V with one shape of its rostrum, its longer rostral loop, short, slender sensory seta. Interantennal and more elongate cells of the outer anal ring. space less than width of segment I. Eyes rath- er inconspicuous, weakly pigmented, slightly protuberant. Rostrum of medium size, 68|-i Rhizoecus cyperalis (Hambleton) long, 53fi wide; rostral loop extending to in- (Figs. 94-97) sertion of second pair of legs. Cephalic plate irregularly elongate, reaching base of anten- Morrisonella cyperalis Hambleton, 1946a: 22. nae, about 50|i long, 33|i wide. Dorsal ostioles Rhizoecus cyperalis: Ferris, 1953: 438; McKenzie, 1967: 383 (misidentification). inconspicuous, their rims thinly sclerotized. Legs of medium size. Average length of seg- Adult Female.—Broadly ovate. Length, 1.42- ments of hind pair: Trochanter, 38; femur, 1.58 mm; width, 0.84-1.02 mm. Antennae six- 91 ; tibia, 79 ; tarsus, 50 ; claw, 23 ; claw digi- segmented, widely spaced, average length of tules long, slender, dilated at their tips, extend- segments: I, 37; II, 21; III, 34; IV, 20; V, 17; ing to or slightly beyond moderately elongate, VI, 41 ; apical segment less than twice as long curved, acute claws. as wide, with three elongate, moderately stout, With one conical circulus about 22ji wide at weakly tapered, falcate sensory setae and one base. Anal lobes undeveloped, without scleroti- narrower, spinelike sensory seta; segment V zation, each lobe area with three elongate with single, short, stout, almost lanceolate sen- setae, one about Glji long, slightly longer and sory seta. Interantennal space about equal to stouter than remaining two. Anal ring well combined length of segments III-VI. Eyes developed, structurally distinct, about 681.1 rather prominent, appearing little longer than wide, its setae approximately 84|i long, much wide. Rostrum elongate, about 75^i long, 42¡.i stouter and longer than lobe setae; outer part wide; rostral loop not reaching insertion of of anal ring with 26-28 mostly small, narrow, second pair of legs. Cephalic plate apparently elongate to oval elongate cells; inner part of lacking. Dorsal ostioles well developed, their ring with 12-16 much larger, irregular cells rims thickly sclerotized. lying adjacent to clouded area of large hemi- Legs moderately stout, average length of spherical cells. Tritubular cerores medium to segments of hind pair : Trochanter, 51 ; femur, small, their ducts narrow, few in number, 122; tibia, 85; tarsus, 75; claw, 24; claw digi- about 20-23 present, more common dorsally. tules very short, acute, not reaching middle of Multilocular disk pores absent. Tubular ducts weakly curved claws. short, smaller than trilocular pore, occurring Circulus absent. Anal lobes weakly pro- in small numbers on both surfaces. Trilocular truding, unsclerotized, each lobe area with one pores rather evenly distributed, with clear seta approximately 50u long and two shorter areas more common ventrally. Body setae slender setae. Anal ring about 40LI in diam- mostly short, longer on head, and posteriorly eter, its setae 60u long, as stout as lobe setae; on abdominal segments. outer part of anal ring with sinuate to oval, Holotype.—El Granizo, Olume, Valparaiso elongate cells, their number impossible to Province, Chile, 5-IV-61, L. Campos; USNM. determine because of condition of type. Tri- Paratypes.—Three, same data as holotype, one tubular cerores of 2 sizes, small and medium, each, IBM, UCD, and USNM. their ducts short, stout, at least 20 of medium- Host Plant.—Unknown, taken from soil. sized cerores occurring dorsally and submar- Distribution.—Known only from type local- ginally over body segments, majority sur- ity. Three immature females believed to be rounded by 1-4 body setae; 4-6 small cerores this species were taken 4 km W. of Cherquenco, present ventrally near body margins of poste- Cautín Province, Chile, 26-IV-65, by R. H. rior abdominal segments. Multilocular disk Gonzalez. pores scarce, only five to seven occurring ven- Discussion.—This new species somewhat re- trally near vulva. Tubular ducts small, few in 22 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE number, observed near body margins on pos- tral loop not reaching insertion of second pair terior abdominal segments. Trilocular pores of legs. Cephalic plate longer than wide, nar- moderately abundant, evenly distributed. row toward apex, about 33fx long. Dorsal osti- Body setae short, slender, rather sparse. oles not conspicuous, weakly sclerotized. Holotype.—Santiago de Maria, El Salvador, Legs moderately short, stout, average length 20-VIII-44, E. J. Hambleton. Paratypes.— of segments of hind pair: Trochanter, 25; fe- Three taken with holotype. USNM. mur, 58; tibia, 53; tarsus, 36; claw, 14; claw Host Plant.—Cyperus tenerrimus. digitules long, slender, dilated at their tips, Distribution.—Known only from the type lo- extending beyond sharp, apically curved claws. cality. With one conical circulus, about 12[x wide Discussion.—McKenzie (1967) recorded cyp- at base. Anal lobes undeveloped, with small, eralis from Adiantum sp., San Mateo County, elongate sclerotized patch between setae, each Calif., 1955. After close examination of the lobe area with three slender setae, longest two rather poor specimens, I am convinced about 43[i long, several body setae, usual derm that they were misidentified and confused pores and cerores. Anal ring 40ji wide, its with pritchardi, which they closely resemble. setae slender, slightly stouter and longer than Furthermore, pritchardi has been taken on lobe setae; outer part of anal ring with 12 the same host, on the same date, and in the sinuate cells, many with spicules; inner part same vicinity, and was recorded as such by of ring with 10 larger, more irregular cells McKenzie (1967). lying adjacent to circle of dark, rounded cells. This Central American species is closely Bitubular cerores present, their two ducts allied to nemoralis, pritchardi, and suhcypera- often lying parallel or appearing to be parti- Us, resembling all three in body shape, anal ally fused, elongate, conical, about 8|i-9j.i ring, and antennae. The presence of one to long, projecting well above derm, occurring four body setae in close proximity to most of over entire surface but more abundant dor- the larger tritubular cerores, fewer cerores, sally across segments and scattered on head. and more elongate, tapering, falcate sensory Multilocular disk pores present on both sur- setae separate cyperalis from nemoralis. The faces, 17-28 on venter of posterior abdominal latter is larger with fewer anal-lobe setae and segments, fewer on dorsum in same general more conspicuous body setae. The characters area. Tubular ducts about same size as trilocu- that separate cyperalis from pritchardi are its lar pores, their rims strongly sclerotized, tubular ducts, more cerores, and absence of rather evenly distributed over entire derm. small circular pores. Trilocular pores sparse. Body setae variable in length, not abundant. Rhizoecus disjunctus McKenzie Holotype.—Corona (Prado Dam), Calif., 27- XII-64, D. R. and J. F. Miller; UCD. Para- (Figs. 98-103) types.—Two taken with holotype, one UCD, Rhizoecus disjunctus McKenzie, 1967: 385. one USNM. Host Plants.—Encelia sp. ?, Eriogomim fasci- Adult Female.—Elongate oval. Length, 0.75- cidatiim, Eriogomim sp. 1.43 mm; width, 0.40-0.56 mm. Antennae six- Distribution.—California, Mexico. segmented, rather short, stout, clávate, closely California: Beaumont, 24-XII-67, D. R. spaced, average length of segments: I, 22; II, Miller. Corona, 27-XII-64, D. R. and J. F. 13 ; III, 21 ; IV, 13 ; V, 12 ; VI, 33 ; apical seg- Miller. ment stout, not twice as long as wide, with Mexico: Three mi S. Zapotitlan, Puebla, 2- three long, slender, falcate sensory setae and III-72, D. R. Miller and F. D. Parker. one spinelike sensory seta near apex; segment Discussion.—The elongate, conical bitubular V with one much shorter, narrow sensory seta. cerores of disjunctus separate this unique spe- Interantennal space less than width of seg- cies from any other Rhizoecus in the Western ment I. Eyes hemispherical, pigmented. Ros- Hemisphere. This type of ceroris is similar to trum of medium size, 53|i long, 40^ wide ; ros- the one found in genicidatus from East Afri- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS

ca, first discussed by me (19i6a). These inter- Tritubular cerores large, their ducts stout, 46- esting cerores, referred to by DeLotto (1957) 54 occurring rather uniformly on both sur- and McKenzie (1967) as ''unitubular pores," faces, 3 on head, 3-7 on each body segment. are actually bitubular in nature ; the two elon- Multilocular disk pores occurring dorsally and gate ducts lie almost parallel. The cerores of ventrally in irregular rows across segments, disjiinctiis differ from those of geniculatus by but more abundant on venter, scarce on head. being larger, wider at the surface, and more Tubular ducts absent. Trilocular pores fairly strongly tapered at their apices. numerous and evenly distributed. Body setae variable in size and length, evenly distributed, Rhizoecus distinctus (Hambleton) some longer setae on head and across abdomi- nal segments. (Figs. 104-110) Holotype.—Falls Church, Va., 14-XI-45, Morrisonella distincta Hambleton, 1946a: 24. Floyd Andre. Paratypes.—Two, one taken with Rhizoecus distinctus: Ferris, 1953: 440. holotype, one Cabin John Bridge, Md., 27-III- 00, T. Pergande. USNM. Adult Female.—Oval elliptical, broad across Host Plants.—Avdropogons-p., Artemisia vul- middle. Length, 1.28-2.22 mm; width, 0.74- garis, Aster sp., DactijUs glomerata, Lespedeza 0.81 mm. Antennae six-segmented, strongly euneata, Malus sp., Qnercus coccinea, Rnhus geniculate, closely spaced near apex of head, argutus, Solidago sp., Tridens flavus, Verbas- average length of segments : I, 33 ; II, 22 ; III, cum, thapsus, unidentified moss. 21; IV, 20; V, 17; VI, 43; segments clothed Distribution.—Maryland, Virginia. with long, slender setae, apical segment with Maryland : Cabin John Bridge, 27-III-00, T. three stout, weakly clávate, falcate sensory Pergande. Libertytown, 25-VIII-44, Floyd setae and one narrow, elongate, spinelike sen- Andre. sory seta; segment V with one narrow, elon- Virginia: Falls Church, 20-VIII, 29-X, gate, curved sensory seta. Interantennal space 19-XI-44, Floyd Andre. Mt. Vernon, 3-XII- less than width of antennal segment I. Eyes 44, Floyd Andre. Purcellville, lO-VI-66 21- small, protuberant, pigmented. Rostrum elon- V-68, 21, 23-IV-69, 29-X, 5-XII-70, 20-11-71, gate, 66(1 long, 40{i wide ; rostral loop reaching E. J. Hambleton. near insertion of second pair of legs. Cephalic Discussion.—I have collected this species on plate large, more or less triangulate, about SSi^i 10 difiTerent host plants near Purcellville, Va. long, with seven to eight slender body setae This additional material enabled me to verify bordering its periphery. Dorsal ostioles incon- the diflferences between associates and distinc- spicuous, their rims with little sclerotization. tus. (See discussion on p. 14.) Legs rather short, stout, average length of segments of hind pair : Trochanter, 36 ; femur, 76; tibia, 55; tarsus, 44; claw, 25; claw digi- Rhizoecus falcifer Kiiiickel d'HercuIais tules short, setose, less than half the length of claw, latter long, slender, moderately curved (Figs. 111-117) beyond middle. Rhizoecus falcifer Künckel d'Herculais, 1878: 163. Circulus absent. Anal lobes weakly protrud- Ripersia terrestris Newstead, 1895: 213. ing with heavily sclerotized patch surround- Rhizoecus terrestris (Newstead): Fernald, 1903: 113. ing lobe setae, each lobe area with three elon- Rhizoecus africanus Brain, 1915: 65. gate setae, longest about 74(i long, and several Rhizoecus decoratus Green, 1926: 177. Rhizoecus moruliferus Green, 1933: 52; Williams 1961- smaller body setae. Anal ring about 55ji wide, 93. its setae rather short, averaging about 58ii Rhizoecus falcifer: Rau, 1937: 267; Hambleton, 1946a: long, shorter and stouter than lobe setae ; outer 53; Ferris, 1953: 444; Williams, 1962: 47; Mc- part of anal ring with 10-12 elongate cells, each Kenzie, 1967: 389; Hambleton, 1973: 65. with 2-5 spicules; inner part of ring with 8 Adult Female.—Elongate oval, rather large. larger, irregular elongate cells lying next to Length, 1.91-4.28 mm; width, 0.59-1.93 mm. darkened area of 10-12 large subcircular cells. Antennae five-segmented, average length of 24 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE segments : I, 66 ; II, 28 ; III, 51 ; IV, 41 ; V, 112 ; Host Plants.—Agapanthtis sp., Ajuga sp., segment I stout, apical segment longest, more Anthémis tinctoria, Aquilegia sp., Biixiis sem- than twice as long as wide, with four moder- jjervirens, Cestritm sp., Chrijsanthemiim frii- ately stout, elongate, tapering, falcate sensory tescens, Chrijsanthemiim sp., Codiaeum sp., setae and one slender, spinelike sensory seta. Coffea arabica, C. liberica, Cttpressiis sp., Cyn- Interantennal space about equal to twice odon dactylon, Fragaria sp.. Gardenia sp., length of antenna! segment I. Eyes absent. Heiichera sp., Howeia helmoreana, H. forster- Rostrum IO811 long, 6811 wide; rostral loop iana, Iris sp., Jasminum sp., Kalmia sp., Lotus reaching about halfway to insertion of second sp., Lycopersicon esculentiim, Ophiopogon sp,, pair of legs. Cephalic plate irregularly quad- Pelargonium sp., Phoenix roehelenii, Picea rate, about 4811 long, usually with two or three abies, Tropaeohim majus, Watsonia sp., Zan- vacuoles. Dorsal ostioles prominent, heavily tedeschia aethiopica, various Gramineae, mis- sclerotized, bordered by numerous body setae cellaneous flowering plants, in soil and leaf and trilocular pores. mold. This species has also been reported from Legs normal, size in proportion to body the following hosts in California, but I have length, average length of segments of hind not seen specimens: Acacia sp.. Anemone hu- pair: Trochanter, 73; femur, 184; tibia, 141; pehensis var. japónica, Aralia sp., Carex sp.. tarsus, 98; claw, 40; claw digitules short, se- Citrus sinensiSy Delphinium sp., Erodium mos- tose, reaching about half the length of moder- chatum, Escallonia rubra, Hibiscus sp., Ligus- ately long, stout, curved claws. trum ovalifolium, Matthiola sp.. Petunia sp.. Circulus absent. Anal lobes roundly protrud- Phoenix canariensis. Piper sp.. Prunus sp., ing, unsclerotized, each lobe area with six Ribes sp., Sijringa vulgaris, Thymus vulgaris, elongate, stout setae, longest about 188^ long, Veronica sp., Vitis sp. and four to six shorter auxiliary setae. Anal Distribution.—California, Florida, Missouri, ring relatively small, about 62\i in diameter, New Jersey, New York, Dutch Guiana, Mexico. its setae long, slender, averaging about 102f.i California: Berkeley, 8-VII-44, E. 0. Essig; long; outer part of ring with 10-12 irregularly 15-11-59, A. E. Pritchard; 25-IX-59, L. A. Foi- sinuate cells, most with spicule ; inner part of son; 17-IV-60, Bill Paul. Davis, 24-V-62, F. ring containing 8 larger, irregular, elongate E. Strong; 6-VIII-63, H. Lange. El Cerrito, cells lying adjacent to circle of darkened semi- 23-11-60, Lora Wiegmann. Fairfax, 5-V-58, circular cells. Tritubular cerores present in 2 W. Erickson. Gardena, 26-V-58, Hener. Healds- sizes, 175-200 present, larger size occurring burg, 1-II-62, W. R. Michie ; 25-III-65, J. Oga- dorsally on head, thorax, and across abdominal wa. La Jolla, 18-V-63, R. A. Lewis. Los segments; medium-sized cerores present ven- Angeles, 9-VI-17, R. S. Woglum; 17-VII-25, trally, few on head and thorax, more numer- L. E. Myers. Modesto, 5-X-56, G. E. Wilhite. ous across abdominal segments. Multilocular Montebello, 16-IV-31, L. S. Jones. Oakland, disk pores present on both surfaces, more 19-VII-39, E. 0. Essig; 30-IV-52, G. B. Laing; abundant ventrally, especially above and below 29-11-59, A. E. Pritchard. Palo Alto, 25-III-41, vulva and on posterior abdominal segments, Duncan. Piedmont, lO-V-59, K. S. Hagen. Rio elsewhere scattered ; dorsally more common on Vista, 14-IX-64, W. H. Lange. Sacramento, head and sides of thorax. Tubular ducts fairly 27-IV, 8-V-63 ; 30-IX-67, H. L. McKenzie. Sa- numerous, present on all segments dorsally linas, 17-IV, 19-V-62, W. H. Lange and L. and ventrally, but more common ventrally on Lanini. San Angelmo, 20-VII-59, J. L. Joos. posterior abdominal segments. Trilocular pores San Jose, 11-1-21, L. R. Cody and E. 0. Essig; numerous and evenly distributed. Body setae 15-V-61, G. S. Myers; 13-IV-21, R. D. Hart- variable in length, most shorter than 30^i, man; 25-IX-59, W. Allen; 18-X-59, W. Allen. some about 55fi, some along body margins San Rafael, Aug. 1959, R. Hunsinger. Santa lOOn long, more abundant and evenly distrib- Barbara, 31-III-42, J. B. Steinweden and S. uted over dorsal surface. Smith. Santa Paula, 14-X-33, E. L. Smith. Types.—Location not determined. Probably Santa Rosa, 15-V-61, M. E. Imsdale. Vallejo, in Muséum d'Histoire Naturelle, Paris. 13-VII-44, E. 0. Essig. Walnut Grove, 10-XI- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 25

69, Rogers. Watsonville, 19-IV-67, L. R. Gil- mented. Rostrum of medium size, elongate, 53¡i logly. Whittier, 29-VII-33, Stickney. long, 39ji wide; rostral loop extending almost Florida: Location unknown, specimens col- to insertion of second pair of legs. Cephalic lected from a palm originating in Florida, 20- plate triangulate, its base about 38^-44{i wide, XI-53, at station in Iowa, W. S. Craig, collector. with three vacuoles and several body setae. Dor- Missouri : St. Louis, 13-VI-41, T. E. Birkett. sal ostioles inconspicuous, lightly sclerotized. New Jersey: E. Rutherford, 1936, G. Rau. Legs small, average length of segments of Summit, 3-1-21, A. I. Bourne and J. G. Mac- hind pair : Trochanter, 34 ; femur, 73 ; tibia, 68 ; donald. tarsus, 40; claw, 19; claw digitules long, slen- New York: Ithaca, October 1940, 21-11-41, der, their tips dilated, reaching to or slightly W. G. Bodenstein; 13-IX-38, W. E. Heming; beyond moderately stout, curved claws. 14-X-50, C. Johansen; April 1955, J. Naegle. With one circulus, about 20|i wide at base, Dutch Guiana: Surinam, 1932?, G. Bunzli. its orifice faveolate. Anal lobes undeveloped, Mexico: Location unknown, taken at quar- each lobe area with small sclerotized patch be- antine. Nogales, Ariz., 4-III-68, G. Ehni. tween elongate setae, latter broken at their Discussion.—Known by the common name bases. Anal ring about 55|i in diameter, its setae ''ground mealybug,'' this species is one of the averaging about 70[i long; cells of outer part of better known cosmopolitan members of the ring varying in size and shape, irregularly genus. Although it was described from France ovate, about one-half of 28 cells with spicules ; and is widely distributed in Europe, it was not inner ring cells fewer in number and varying discovered in the United States until 1917. greatly in shape and arrangement, lying next Since then it has at times been a troublesome to contiguous ring of large darkened cells. Tri- pest. tubular cerores of medium size, 48-55 evenly R. falcifer is the largest species of the genus. distributed dorsally and ventrally, not more It is easily recognized by its five-segmented an- than 4 or 5 to an abdominal segment. Multi- tennae, the clusters of anal lobe setae, the many locular disk pores absent. Tubular ducts smaller tritubular cerores, and lack of circulus. Al- in diameter than trilocular pore, generally dis- though considered by some workers to lack tu- tributed over entire derm. Trilocular pores bular ducts, these structures are present in most numerous dorsally, in some areas sparse considerable numbers over the entire body sur- on venter. Body setae mostly short, moderately face in mature females. In general appearance, sparse. falcifer resembles kondonis. The presence of Holotype.—Coto, Costa Rica, 20-V-57, E. B. small bitubular cerores and circuli in the latter Dixon. Paratype.—One, taken with holotype. separates the two species immediately. USNM. Host Plant.—Unknown, specimens taken in soil under Coffea arabica. Rhizoecus favacirculus^ new species Distribution.—Known only from type locality. (Figs. 118-125) Discussion.—This species closely resembles arahicus, which is also found on coffee roots, Adult Female.—Elongate ovate, of medium but favacirctihis differs by possessing long claw size. Length, 1.28 mm; width, 0.60 mm. Anten- digitules, shorter antennae, and stouter falcate nae six-segmented, rather short, widely spaced, setae. The smaller legs and shape of the anal average length of segments: I, 33; II, 15; III, ring cells of favacircuhis also are distinctive. 22; IV, 11; V, 11; VI, 33; apical segment cone- shaped, longer than wide, with three short, stout, weakly clávate, falcate sensory setae with Rhizoecus floridanus Hambleton tapered apices and one slender, spinelike sen- (Figs. 126-132) sory seta; segment V with one fairly elongate, clávate sensory seta. Interantennal space equal RhÍ2:occns floridanus Hambleton, 1973: 67. to combined length of segments IV, V, and VI. Adult Female.—Broadly ovate, narrower in Eyes prominent, globose, not strongly pig- area of head and thorax. Length, 1.20-1.63 mm; 26 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE width, 0.65-0.80 mm. Antennae six-segmented, Aralia sp.. Araucaria excelsa, Arecastrum rom- of medium size, average length of segments: anzo ffianum, Bambnsa sp., Billhergia sp., Buxus I, 31 ; II, 19 ; III, 25 ; IV, 17 ; V, 16 ; VI, 37 ; api- sp., Calliandra sp., Callistemon rigidus, Carissa cal segment slightly less than twice as long as grandiflora, Carissa sp., Chry salido carpus lu- wide, with three moderately long, stout, falcate tescens, Citrus mitis, Citrus sp., Conocarpus sensory setae and one slender, acute sensory erecta, Cortaderia selloana, Cuphea sp., Diejfen- seta; penultimate segment with one smaller, bachia sp., Distichlis spicata, Dizygotheca ele- rather elongate, weakly clávate sensory seta. gantissima, Dracaena marginata, Eremochloa Interantennal space equal to about twice length ophiuroides, Eugenia sp.. Gardenia thunbergia. of segment I. Eyes absent. Rostrum relatively Hoya sp.. Ilex cornuta cv. Burfordii, /. opaca, short, measuring about 57|i long, 40^ wide; L rotunda, L vomitoria, Ixora sp., Jasminum rostral loop extending to or near insertion of sp., Lachnanthes tinctoria, Leucophyllum fru- second pair of legs. Cephalic plate irregularly tescens, Panicum sp.. Philodendron selloum, triangulate to quadrate, wider posteriorly, Philoxerus vermicularis. Phoenix canariensis, about 38^ across its base, with indistinct vac- Pluchea sp.. Prunus angustifolia, Pyracantha uoles. Dorsal ostioles weakly sclerotized, incon- sp., Quercus sp.. Rhododendron sp.. Sida sp.. spicuous. Viburnum suspensum, undetermined Grami- Legs of medium size, average length of seg- neae, Palmaceae, from rotting wood of dead ments of hind pair : Trochanter, 37 ; femur, 81 ; maple. tibia, 69; tarsus, 51; claw, 22; claw digitules Distribution.—Florida, Georgia, Indiana, long, their tips dilated, surpassing narrow, elon- Maryland. gate, weakly curved claws. Florida : Alva, 17-X-72, J. C. Denmark, Apop- With one conical circulus, variable in size, ka, 11, 15-11-71, C. L. Speaker; 14, 19-VI-72, width at base varying from 21ji to 35^x, its apex F. L. Ware; lO-XI-72, P. Gibson and W. H. finely faveolate. Anal lobes undeveloped, each Pierce; 3-VIII-72, J. F. Knauss. Arcadia, lobe area with small, narrow sclerotized patch 26-11-71, G. P. Lamb. Boynton Beach, 4-1-73, between three elongate setae, longest measur- W. H. Pierce. Bradenton, 5-1-71, J. R. McFar- ing about 60|i long. Anal ring about 60ji in lin; 11-1-71, S. L. Poe. Casselberry, 26-XII- diameter, its setae considerably longer and 63, C. 0. Youtsey. Clearwater, 12-11-71, E. W. stouter than lobe setae, averaging about 87|i Miller. Cudjoe Key, 2-1-73, E. J. Hambleton. long; outer anal-ring cells large, diversiform, Dade City, 30-IX-72, 15-XI-72, W. H. Pierce. 24-33 present, mostly isolated; inner part of Delray Beach, 11, 15-11-71, W. E. Wyles and anal ring with 14-20 cells of similar size and R. A. Long. Eau Gallie, 8-II-71, H. C. Levan. shape lying adjacent to inner darkened area of Englewood, 29-VI-71, C. J. Bickner. Fairvilla, semicircular cells. Tritubular cerores small, 2, 22-11-71, 21-XII-71, F. L. Ware. Flamingo, v/alls of their individual ducts almost parallel, 18-11-70, E. J. Hambleton. Ft. Meyers, 31-1- 36-40 present dorsally and ventrally, most abun- 73, W. A. Padgett. Gainesville, 19-IV-67, dant on abdomen and rather evenly distributed G. W. Dekle; 29-III-67, G. W. Dekle and C. elsewhere. Multilocular disk pores absent. Tu- Lyons. Gillette, 23-IV, 23-V-71, C. J. Bickner. bular ducts small, widely scattered on both sur- Grant, 19-11-71, H. C. Levan. Key West, l-I- faces but absent from dorsal surface of head. 73, E. J. Hambleton. Lakeland, 6~VI-69, J. W. Trilocular pores fairly evenly distributed. Body McLeod. Largo, 10, 11-11-71, C. K. Hickman setae mostly short. et al. Lockhart, 30-XII-63, 6-1-64, R. J. Grif- Holotype.—Pembroke, Fla., 2-VIII-67, H. G. fith. Mango, 19-11-71, E. R. Simmons. Naples, Schmidt. Paratypes.—Many from numerous 4-III-71, W. T. Walsh and W. A Padgett. No localities in Florida and from Spaulding Coun- Name Key, 21-X-72, W. H. Pierce. N. Miami ty, Ga. Holotype and five paratypes USNM ; two Beach, 18-XII-72, E. J. Hambleton. Oneca, paratypes each BM, UCD, UG, VPI; remaining 27-V-71, J. R. McFarlin. Orlando, 20-11-69, paratypes FSCA. F. L. Ware; 10-11-71, W. W. Smith and E. R. Host Plants.—Aechmea chantinii, A. orlandi- Fatic; 21-V-71, F. L. Ware; 26-11-71, D. A. ana, Andropogon virginicus, Anthémis sp., Grady and E. W. Ensign; 31-VII-72, F. L. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 27 Ware. Osprey, 5, 10, 15-11-71, J. R. McFarlin mented, and constricted at their bases. Rostrum and C. J. Bickner. Palma Sola, 19-X-72, C. J. of medium size, about 73u long, 48¡-i wide; Bickner. Palmetto, 25-11-71, J. R. McFarlin. rostral loop extending to or slightly beyond in- Pembroke, 14-IV, 2-VIII-67, H. G. Schmidt. sertion of second pair of legs. Cephalic plate Pinellas Park, 18-11-71, C. K. Hickman. Plant irregular in shape, 30u-40^i wide, with two or City, 19-11-71, D. A. Vaughn. Plymouth, 29-1- three vacuoles near its center. Dorsal ostioles 71, W. W. Smith and E. R. Fatic; 19-V-71, rather prominent, their rims rather heavily H. M. Van Pelt; 12-X-71, H. M. Van Pelt; sclerotized. 19-X-72, P. Gibson. Punta Gorda, 16-V-71, Legs well developed, average length of seg- G. P. Lamb. Sarasota, 22-11-71, J. R. McFarlin ments of hind pair: Trochanter, 50; femur, and C. J. Bickner. Sebastian Inlet, 12-11-71, 115; tibia, 88; tarsus, 64; claw, 28; claw digi- H. C. Levan. Seffner, 12-11, 17-VIII-71, D. A. tules short, setose, not reaching middle of long, Vaughn. Tall Timbers, 12-XII-69, H. H. Tip- narrow, weakly curved claws. pins. Tampa, 3, 15-11-71, S. A. Fuller and Circulus absent. Anal lobes slightly pro- C. W. Hale; 25-VIII-72, C. W. Hale. Terra truded and weakly sclerotized, each with three Ceia, 15, 18-IX-72, J. R. McFarlin. Vineland, slender setae averaging about 50^i long. Anal 29-XI-72, F. L. Ware. W. Melbourne, 12-n-71, ring about 47^i in diameter, its structure not H. C. Levan. W. Palm Beach, 11, 15-n-71, clearly defined, ring setae slightly stouter than 4-ni-71, M. L. Messec. Windermere, 28-XI-73, anal-lobe setae, longer setae at least 581^1 long; F. L. Ware. Winter Haven, ll-Vni-68, H. C. cellular structure of outer part of ring appear- Burnett. ing narrow and elongate, possibly with less Georgia: Spaulding County, 16-V-68, H. H. than 13 cells; inner part of ring with larger, Tippins. undiiferentiated cells. Tritubular cerores of 2 Indiana : Bluff ton. Wells County, 19-XII-72, sizes, 16-18 of large size on dorsum, 6 along R. F. Wilkey. each lateral margin, 4-6 along middorsal Hne; Maryland: Grasonville (Kent Narrows), 24- small to medium-sized cerores occurring on ven- IX-42, H. S. McConnell. ter of abdominal segments VI-VIII. Multilocu- Discussion.—R. floridanus most closely re- lar disk pores of 6 or 7 loculi confined to venter sembles tropicalis from Guatemala but is easily of posterior abdominal segments in area of recognized by its more elongate, apical anten- vulva, 18-25 present. Medioventral pores on na! segment, more slender falcate setae, and venter, usually occurring in two groups, one on larger and more numerous triangulate to abdominal segment VI with none to five pores, quadrate anal-ring cells. other posterior to it on abdominal segment VII with four to nine pores. Trilocular pores uni- Rhizoecus globoculus (Hambleton), formly distributed but not abundant. Body new combination setae rather fine, sparsely distributed. Holotype.—Non Pareil Estate, Trinidad, Brit- (Figs. 133-140) ish West Indies, 3-III-44, A. H. Strickland. Morrisonella globocula Hambleton, 1946a: 25. Paratypes.—Two taken with holotype; five, Marper Estate, Trinidad, 5-1-44, A. H. Strick- Adult Female.—Oval elongate, moderately land ; one, Trinidad, 6-VI-35, E. J. H. Berwick stout. Length, 1.05-1.36 mm; width, 0.60-0.76 USNM. mm. Antennae six-segmented, average length Host Plants—Cojffea arabica, Theobroma of segments: I, 37; II, 22; III, 26; IV, 21; V, cacao. 19; VI, 45; apical segment longer than vv^ide, Distribution.—Known only from the type lo- v^ith three medium-sized, gently tapered, fal- cality. cate sensory setae and one narrow, elongate, Discussion.-This species runs parallel with spinelike sensory seta ; segment V with one nar- theobromae in the key to the New World spe- row, elongate sensory seta. Interantennal space cies of Rhizoecus, Although both species attack equal to combined length of segments III, IV, cacao and are rather unique in having medio- and V. Eyes prominent, globular, weakly pig- ventral pores, globoculus is readily distin- 28 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE guished from the former by its large globular abdominal segment, occasionally observed ven- eyes, its multilocular disk pores of six or seven trally near body margins. Multilocular disk loculi, and the more elongate, tapering falcate pores variable in number, 11-64 located pos- setae. teriorly on venter from apex to abdominal seg- ment VI, few occurring dorsally on abdomen. Rhizoecus gracilis McKenzie Tubular ducts short, stout, readily observed, scattered dorsally and ventrally over most of (Figs. 141-149) derm. Trilocular pores evenly distributed except Rhizoecus gracilis McKenzie, 1961: 45; McKenzie, 1967: between segments where they are sparse or 392. missing. Body setae short, slender, uniformly Adult Female.—Elongate oval. Length, 0.95- distributed. 2.07 mm; width, 0.30-0.83 mm. Antennae six- Holotype.—Cahill Ridge, Calif., 7-V-60, D. W. segmented, moderately short, stout, average Price. Paratypes.—Several taken with holo- length of segments : I, 32 ; II, 18 ; III, 25 ; IV, type, also from 2 mi W. of Independence, 6-V- 17; V, 15; VI, 38; apical segment slightly less 60, A. S. Menke and F. D. Parker, and at Susan- than twice as long as wide, with three medium- ville, Calif., 17-V-58, R. W. Gerhardt. Holo- sized, rather slender, falcate sensory setae and types and paratypes UCD ; paratypes CDA, one shorter, tapering, spinelike sensory seta; USNM. segment V with one much shorter, narrow sen- Host Plants.— Achillea millefolmm, Artemi- sory seta. Interantennal space equal to about sia californica, Artemisia sp., A. tridentata, width of basal antennal segment. Eyes hemi- Atriplex sp., Brickellia sp., Chrysothammis spherical, not strongly pigmented. Rostrum viscidiflorus, Eriogonum heracleoides, Eriophyl- relatively small, 66^i long, 42fi wide; rostral him confertifloriim, Franseria chamissonis, loop extending to about halfway to insertion Grindelia camporum, Gutierrezia sp., Haplo- of second pair of legs. Cephalic plate slightly pappus canus, H. spinulosus ?, undetermined longer than wide, with two vacuoles and four Cactaceae, Chenapodiaceae, Compositae, Grami- body setae. Dorsal ostioles not strongly devel- neae, Leguminoseae, Loranthaceae, in soil, oped, their rims thinly sclerotized and weakly under rocks, beneath chaparral. pigmented. Distribution.—Arizona, California, Colorado, Legs rather small, stout, average length of Idaho, Montana, New Mexico, Oklahoma, Ore- segments of hind pair : Trochanter, 35 ; femur, gon, Texas, Virginia, Mexico. 81 ; tibia, 72 ; tarsus, 48 ; claw, 15 ; claw digi- Arizona: Bisbee, 2-VIII-66, D. R. Miller. tules long, dilated at their tips, extending be- Five mi NE. Douglas, 2-VIII-66, D. R. Miller. yond short, stout claws. California: Barrego Springs, 27-1-65, D. R. With one broadly truncated circulus, width Miller. Cahill Ridge, 7-V-60, D. W. Price. Co- across its base about 20}i-28fx, its orifice ches Prietos, Santa Cruz Island, 18-VI-67, 10- reticulated. Anal lobes undeveloped, each lobe V-68, D. R. Miller. Two mi W. of Independ- area with small, elongate sclerotized patch ence, 6-V-60, A. S. Menke and F. D. Parker. and three elongate setae subequal in length, Fifteen mi N. of Kramer Junction, 28-XII-64, longest about 51|i long. Anal ring approxi- D. R. and J. F. Miller. Two mi N. of Lompoc, mately 48|i wide, its setae averaging 5S[i long, 27-VI-66, D. R. Miller. Merced, 18-1-67, T. R. stouter and somewhat longer than lobe setae; Haig. Monitor Pass, 9-VIII-64, D. R. Miller. outer part of anal ring with 14-16 sinuate Ten mi N. of Pescadero, 8-II-64, D. R. Miller to oval, elongate cells, each with long spic- and J. A. Froebe. Susanville, 17-V-58, W. ule; inner part of ring with at least 10 large, Gerhardt. Trabuco Canyon, 28-III-64, D. R. irregular, elongate cells lying adjacent to well- Miller and J. A. Froebe. Valley Center, 27-III- defined clouded area of semicircular cells. Bi- 64, D. R. Miller and J. A. Froebe. Warner tubular cerores small, moderately stout, their Springs, 26-1-65, D. R. Miller. W. Patterson, ducts about 7\i long, shghtly tapered and divari- 27-IV-66, D. R. Miller. cated, widely distributed dorsally, 5-8 on head, Colorado: Holly, 29-VI-70, D. R. Miller. La- 25-28 on thorax, usually 4-7 spread across each mar, 30-VI-70, D. R. Miller. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 29

Idaho: Craters of the Moon, 8-VIII-67, Rhizoecus graminis (Hambleton) D. R. Miller and D. S. Horning. Five mi E. of Montpelier, 3-VIII-67, D. R. Miller and D. S. (Figs. 150-154) Horning. Twenty mi N. of Spencer, 5-VIII-67, Morrisonella graminis Hambleton, 1946a: 28. D. R. Miller and D. S. Horning. Rhizoecus graminis: Ferris, 1953: 446; McKenzie, 1967: 394. Montana: Boyes, ll-VIII-70, D. R. Miller. Nine mi SE. Livingston, 28-VIII-64, D. R. and Adult Female.—Oval elongate. Length, 1.26- J. F. Miller. 1.85 mm; width, 0.54-0.76 mm. Antennae six- New Mexico: Carrizozo, 6-VIII-66, D. R. segmented, elongate, segments I and VI rather Miller. Five mi NW. Cedarvalle, 6-VIII-66, stout, length of segments as follows : I, 57 ; II, D. R. Miller. Three mi S. Oscuro, 5-VIII-66, 26; III, 37; IV, 27; V, 28; VI, 70; apical seg- D. R. Miller. Seven mi W. of Silver City, ment robust, about twice as long as wide, with 6-IX-68, D. R. Miller and J. E. Lauck. three moderately stout, falcate sensory setae, Oklahoma : N. of Boise City, 30-VI-70, D. R. tapering gradually apically, and one much nar- Miller. rower, spinelike sensory seta ; segment V with Oregon: Bandon, 6-VIII-68, D. R. Miller one much shorter, stout, clávate sensory seta and R. F. Denno. Twelve mi SW. of Plush, narrow at its base. Eyes rather prominent, pro- 3-VIII-68, D. R. Miller and R. F. Denno. Ten tuberant, lightly pigmented. Rostrum elongate, mi W. of Vale, 4-VIII-70, D. R. Miller. 96}-i long, 61^1 wide; rostral loop reaching in- sertion of second pair of legs. Cephalic plate Texas: Dumas, l-VII-70, D. R. Miller. relatively small, with two vacuoles and two Virginia: Mountain Lake, 24-X-70, D. R. body setae. Dorsal ostioles conspicuous, their Miller. rims heavily sclerotized and surrounded by Mexico: Apizaco, Tlaxcala, 16-VII-67, D. R. numerous body setae and pores. Miller and J. Villanueva B. Jalapa, Vera Cruz, Legs well developed, elongate, average length 16-VII-67, D. R. Miller and J. Villanueva B. of segments of hind pair: Trochanter, 63; fe- Fifteen km SW. of Miraflores, Vera Cruz, 13- mur, 135; tibia, 122; tarsus, 73; claw, 28; claw VII-67, D. R. Miller and J. Villanueva B. Seven digitules very short, setose, less than half the km N. of Perote, Vera Cruz, 28-11-72, and 7 km length of stout, elongate, curved claws. SW. of Perote, 29-11-72, D. R. Miller and F. D. Without circulus. Anal lobes weakly pro- Parker. Texcoco, 12-VII-67, D. R. Miller and truded, partially sclerotized, each lobe area J. Villanueva B. with three elongate setae, longest about 88u Discussion.—Since its discovery in California long, and several shorter body setae; trilocular in 1960, gracilis has been collected in nine ad- pores crowded at base of larger setae. Anal ring ditional States and Mexico owing largely to the about 51LI in diameter, its structure not clearly intensive and productive collecting by D. R. visible, its setae averaging about 67fi long, Miller and his associates. Except for seven lo- slightly narrower and much shorter than longer cality records previously reported by McKenzie lobe setae ; outer part of anal ring apparently (1967) for California, the distribution records with 14-16 small, elongate cells; inner part of listed here are new. With its recent discovery ring with cells larger and less numerous. Tri- in Virginia, gracilis appears to be the most tubular cerores of 2 sizes present, 20 of larger widely distributed indigenous species in the size occurring dorsally, 6 along middorsal line United States. from head to abdominal segment VI, 7 along The species most closely resembling gracilis each body margin or submarginally ; small tri- is totonicapamis from Guatemala. The falcate tubular cerores with about 25 present ventrally, sensory setae of gracilis are slender and taper 4 on thorax, 21 distributed across abdominal apically, whereas those of totonicapamis are segments IV-IX. Multilocular disk pores nu- somewhat stouter and weakly clávate distally. merous, present dorsally and ventrally, more In gracilis the rostrum is slightly longer and profuse ventrally in bands across abdominal there appear to be fewer bitubular cerores than segments. Tubular ducts small, their collars in totonicapamis. sclerotized, slightly larger in diameter than tri- 30 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE locular pore, commonly observed ventrally much stouter than lobe setae ; outer part of anal on posterior abdominal segments. Mushroom ring with 28 large, subquadrate to triangulate bodies present dorsally and ventrally. Trilocu- cells, mostly touching end to end; inner part lar pores apparently with tiny prongs, rather of ring with 10-12 irregular, more elongate uniformly distributed except scarce on thorax. cells lying adjacent to darkened area of cellular Body setae variable in size, distributed in same structure. Tritubular cerores small, their ducts manner as trilocular pores. narrow, elongate, projecting more than one- Holotype.—Oakland, Calif., 29-1-38, E. 0. half their length above body surface, 55-60 of Essig. Paratypes.—Twelve taken with holo- them occurring on both surfaces, widely dis- type, 7 on one slide with holotype, 5 on 2 addi- tributed, as many as 6 per abdominal segment. tional shdes. USNM. Multilocular disk pores absent. Tubular ducts Host Plant.—Agrostis sp. small, few in number, observed more readily Distribution.—California. The type locality, along body margins. Trilocular pores moder- and Colma, 12-XII-63, C. S. Koehler. ately abundant, scattered. Body setae short, Discussion.—Although R. graminis keys to inconspicuous. the same couplet as latiis from Ecuador, the Holotype.—Darwin Research Station, Gala- two species are easily separated. The antennae pagos Islands, 30-1-64, R. 0. Schuster; UCD. and leg segments of graminis are much larger, Host Plant.—Hippomane mancinella. the eyes less protuberant, and the number of Distribution.—Known only from the type lo- multilocular disk pores is greater than in latus. cality. Discussion.—At least three species of Rhizoecus are now known from the Galapagos Rhizoecus insularis^ new species Islands : lattis, insularis, and a third apparently (Figs. 155-158) undescribed species of which only an immature female has been seen. The large cellular struc- Adult Female.—Elongate oval. Length, 1.24 ture of the anal ring, the small rostrum, and mm ; width, 0.65 mm. Antennae six-segmented, number and size of the tritubular cerores are of average size, segments measuring as follows : distinguishing features of insularis. I, 22 ; II, 17 ; III, 25 ; IV, 13 ; V, 14 ; VI, 33 ; api- cal segment less than twice as long as wide, with three moderately stout, elongate, falcate Rhizoecus kondonis Kuwana sensory setae and one slender, spinelike sensory (Figs. 159-166) seta ; segment V with one shorter, narrow sen- sory seta. Interantennal space equal to com- Rhizoecus kondonis Kuwana, 1923: 55; Hambleton, bined length of segments I and II. Eyes semi- 1946a: 56; Ferris, 1953: 448; McKenzie, 1960b: circular, rather prominent. Rostrum small, 749; McKenzie, 1967: 394; Kuwai and Takagi, 1971: 177. 47^1 long, 44^ wide; rostral loop extending to Rhizoecus spinosus McKenzie, 1960b: 753; McKenzie, about insertion of second pair of legs. Cephalic 1967: 404. NEW SYNONYMY. plate not distinguishable. Dorsal ostioles incon- spicuous, their rims very thinly sclerotized. Adult Female.—Elongate oval. Length, 1.58- Legs short, segments of hind legs measuring 2.24 mm; width, 0.62-0.92 mm. Antenna five- as follows: Trochanter, 35; femur, 72; tibia, segmented, moderately stout, placed near apex 68; tarsus, 46; hind claw missing; claw digi- of head, segments with following measure- tules long, slender, their tips dilated, reaching ments : I, 55 ; II, 34 ; III, 38 ; IV, 25 ; V, 95 ; api- slightly beyond claws. cal segment almost three times as long as wide, With one conical, weakly faveolate circulus segment I very stout, with four relatively stout, measuring about 16ji at its base. Anal lobes short, weakly tapered, falcate sensory setae and slightly or not developed, each lobe area with one narrow, elongate, spinelike seta gently three small, delicate setae, longest about 60!.i curved. Interantennal space less than width of long. Anal ring clearly defined, about 55j_i wide, segment I. Eyes absent. Rostrum averaging its setae averaging approximately 66[i long, about 75[i long, 53u wide ; rostral loop reaching A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 31 almost to insertion of second pair of legs. Ce- largonium inquinans, Portulaca grandiflora, phalic plate occasionally obscure, wider than Primus sp., Rosa sp., Rumex sp., Scabiosa sp., long, triangulate, with 2 small vacuoles near Stellaria media, Watsonia sp., leaf mold, decay- its center and 8-10 short setae along or near its ing log. border. Dorsal ostioles well developed, strongly Distribution.—California, Guatemala. sclerotized, and surrounded by numerous setae California: Berkeley, 27-X-28, E. 0. Essig. and pores. Butte County, 15-X-62, M. Morse. Clarksburg, Legs large, well developed, average length of 23-XI-60, V. E. Berton. Davis, 12-XI-58, 'segments of hind pair: Trochanter, 59; femur, L. M. Smith ; 9-XI-60, D. W. Price and R. 0. 137; tibia, 109; tarsus, 81; claw, 34; claw digi- Schuster; 15-X-65, Mrs. Rose. Two mi N. of tules very short, acute, less than half the length Dixon, ll-V-66, A. G. Gentile. Healdsburg, of slender, curved claws. 12-1-62, P. F. Wright. Hopland, 2-VIII-67, Usually with two truncate circuli, smaller T. Erickson. Knight's Landing, 24-IV-59, one on abdominal segment III, one on segment F. C. Raney. Menlo Park, 11-III-21, G. F. IV averaging about 29[i in diameter, both cir- Ferris. Mix Canyon, 25-VIII-60, R. 0. Schus- culi with thinly sclerotized orifices and some ter. Placer County between Wheatland and reticulation. Anal lobes well developed, roundly Newcastle, 26-IX-62, V. Marble. Placerville, protruding, apparently unsclerotized, each with 11-11-44, M. Austin. Sacramento, 8-V-63, cluster of 8-10 elongate setae, varying in length, H. L. McKenzie. St. Helena, 12 mi N. of city, longest about 98!i long. Anal ring about 54ji in 8-IV-61, F. C. Raney. San Jose, lO-XII-58, diameter, well differentiated, its setae averag- A. E. Pritchard. San Leandro, 9-VI-59, A. E. ing about 60^ long, more slender than largest Pritchard. Santa Monica, 18-IV-40, in quar- lobe setae; outer part of anal ring with 18-20 antine from St. Helena. Santa Rosa, 7-XI-58, fairly uniform, oval, elongate cells, each with B. Houston. Stockton, 3-XI-70, J. Gianelli. spicule ; inner part of ring made up of about 10 Yuba City, 26-V-59, K. Uriu. Two mi S. of larger, more elongate, and irregular cells, Yuba City, 23-ni-62, L. M. Smith. clouded area with little or no visible design. Bi- Guatemala: San Marcos, San Marcos Prov- tubular cerores small, each with 2 ducts arising ince, 4-XI-63, N. Escobar. from sclerotized cone with oval-shaped base, Discussion.—The distribution of kondonis in 25-30 uniformly distributed on dorsum. Multi- the United States is restricted to California, locular disk pores concentrated ventrally in where it has become a pest of considerable area of vulva and from segment VII posteriorly importance (McKenzie, 1960b). Its presence to apex, absent elsewhere, 65-106 present. Tu- in Guatemala may indicate a wider distribution bular ducts present on both surfaces but most in the hemisphere than records signify. common on abdomen, their diameter about size ^ Kuwai and Takagi (1971) have called atten- of trilocular pore. Trilocular pores uniformly tion to some discrepancies between Japanese distributed over most of derm. Body setae nu- specimens of kondonis and the descriptions and merous, variable in length, some measuring figures given by Ferris (1953) and McKenzie 100|i long. (1967). Ferris' indequate illustration and di- Types.—According to S. Takagi (personal agnosis of kondonis apparently mislead Mc- communication), Kuwana's types of kondonis Kenzie when he (1960b) prepared the descrip- were probably lost in a fire following an earth- tion of spinosus. No mention was made of its quake in Tokyo, Sept. 1, 1923. Two topotypes similarity to the Japanese species until 1967. taken on roots of orange, Wakayama, Japan, ^ I have carefully checked the types and nine ad- in November 1926 and identified by Kuwana, ditional specimens labeled spinosus and find no formerly a part of the Stickney collection, are difference between them and those labeled kon- in the USNM. The types of R. spinosus are in donis. The type specimen of spinosus has what UCD. appears to be a poorly defined and weakly scle- Host Plants.—Caréna; sp. ?, Celtis occidentalism rotized circulus on segment III in addition to Citrus sp., Coffea arabica, Fragaria sp., Ligust- the normal larger one on segment IV. Occasion- rum sp., Medicago sativa, Nerium sp.. Pe- ally only a single circulus may be present in 32 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE species normally having multiple circuli. Of about 60^ long, slightly stouter and longer than the 95 specimens of kondonis studied, 35 were shorter lobe setae ; outer part of anal ring with immature females, all with a single circulus. 10-12 elongate, oval cells, each with spicule; One specimen identified by McKenzie as spino- inner part of ring with irregularly elongate, sus has two circuli, the normal number for somewhat larger cells lying adjacent to concen- kondonis. Since there appear to be no consistent tric ring of large, ovate, darkened cells. Tritu- differences between the two, I consider them bular cerores of 2 sizes, larger size confined to the same, and spinosus thus becomes a junior dorsum, 1 on head, 5 on thorax, 5 on abdomen ; synonym of kondonis. 10-12 medium-sized cerores, most occurring on The roundly protruding anal lobes beset with venter of posterior abdominal segments. Multi- numerous elongate setae, the normal presence locular disk pores present on both surfaces, at of two circuli in the adult female, the five- least 50 observed ventrally from apex of abdo- segmented antennae, and bitubular cerores men to abdominal segment VII, elsewhere very are important distinguishing characters of few scattered widely on head, thorax, and ab- kondonis. domen. Tubular ducts present, their collars slightly larger in diameter than a trilocular pore, most common ventrally but also present Rhizoecus latus (Hambleton), dorsally. Trilocular pores sparse, uniformly new combination distributed. Body setae sparse, mostly short (Figs. 167-174) but few longer ones on head and along body margins. Morrisonella lata Hambleton, 1946a: 30. Holotype.—Pichilingue, Ecuador, 2-X-44, E. Adult Female.—Broadly ovate. Length, 1.35- J. Hambleton. Paratypes.—Five taken with 1.61 mm; width, 0.82-1.07 mm. Antennae six- holotype. USNM. segmented, widely spaced near apex of head, Host Plants.—Hippomane mancinella, un- segments measuring as follows: I, 38; II, 20; determined Gramineae. III, 25 ; IV, 20 ; V, 17 ; VI, 45 ; apical segment Distribution.—Ecuador, Galapagos Islands. tapered, with three moderately stout, falcate Ecuador: From the type locality. sensory setae and one elongate, slender, acute Galapagos Islands : Darwin Research Station, sensory seta; segment V with one elongate, Santa Cruz Island, 25-1, 30-1-64, R. 0. Schuster. slender sensory seta. Interantennal space equal Discussion»—Sixteen specimens from an un- to combined length of four distal antennal seg- determined host taken in the Galapagos Islands ments. Eyes prominent, protuberant, pig- answer the description of latus but differ in the mented, slightly longer than wide. Rostrum of following respects: The largest adult female medium size, 75\i long, 48pi wide; rostral loop measures 1.43 mm, whereas most specimens extending to beyond halfway to insertion of consist of newly emerged females with nondis- second pair of legs. Cephalic plate small, tri- tended bodies that measure less than 1.00 mm angulate to quadrate, with indistinct vacuoles in length ; the rostrum is shorter, more rounded and body seta on each laterocephalic margin. at its apex; the sensory seta on antennal seg- Dorsal ostioles conspicuous, their rims thickly ment V is weakly clávate apically. In another sclerotized. collection from the same locality, taken on Hip- Legs of normal size, average size of segments pomane mancinella, two specimens of latus of hind pair : Trochanter, 50 ; femur, 107 ; tibia, were taken with the species insularis. 80; tarsus, 64; claw, 26; claw digitules short, setose, reaching almost to middle of fairly stout, weakly curved claws. Rhizoecus leucosomus (Cockerell) Circulus absent. Anal lobes slightly promi- (Figs. 175-181) nent, each sclerotized and with one long and two shorter elongate setae, longest about 80^i- Ripersiella leucosoma Cockerell, 1901: 165. 85|i long, and several auxihary setae. Anal ring Ripersiella leucosoma: Fernald, 1903: 115; Bueker, rather small, about 41^x in diameter, its setae 1931: 151; Hambleton, 1946a: 66. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 33

Rhizoecus leucosomus: Ferris, 1953: 450; McKenzie, Host Plants.—Bahia dissecta, Cypertis esctt- 1967: 398. lentus, Escoboria tuberculosa, Mammillaria sp., Rhizoecus cacticans (Hambleton) : McKenzie, 1967: 400 (misidentification in part). Nerium oleander, Phleum pratense, Poa sp., Psoralea tenuiflora, Sempervivum sp.. Sorghum Adult Female.—Oval elongate, moderately halepense, unidentified Gramineae, under stones stout. Length, 1.60-2.88 mm; width, 0.70-1.55 with ants. mm. Antennae six-segmented, rather widely Distribution.—Arizona, California, Colorado, spaced; segments in following proportions: I, New Mexico, Ohio, Texas, Virginia, Washing- 34 ; II, 24 ; III, 34 ; IV, 18 ; V, 20 ; VI, 53 ; apical ton, D.C., Mexico. segment seldom twice as long as wide, with Arizona: 19 mi E. McNary, 8,500 ft, 18-VII- three elongate, medium-sized, falcate sensory 40, L. P. Wehrle. Williams, l-IX-68, D. R. setae and one narrow, spinelike sensory seta; Miller and J. E. Lauck. segment V with one shorter but elongate sen- California: Calexico, 30-III-72, R. A. Flock sory seta. Interantennal space equal to com- and D. Conn. Norco, 4-X-65, N. W. Getz and bined length of two apical segments. Eyes com- J. H. Preston. Sacramento, 22-IX-65, S. Scrib- paratively small, not strongly pigmented, ner. San Clemente, 22-X-65, A. Johnson. slightly protuberant. Rostrum short, stout, Colorado: Boulder, 13-VIII-23, R. Shotwell. averaging about 74|i long, 64[i wide; rostral New Mexico: Las Vegas, 11-IV-Ol, W. P. loop seldom reaching halfway to insertion of Cockerell. second pair of legs. Cephalic plate fairly large, Ohio: In quarantine at Washington, D.C., longer than wide, about 45ji long, with two 4-XII-22, W. B. Wood. large vacuoles, and usually bordered by three or Texas: Corpus Christi, 4-VIII-71, S. Naka- four setae. Dorsal ostioles plainly evident, hara. weakly sclerotized. Virginia: Chincoteague, 17-VIII-71, D. R. Legs of average size, segments of hind pair Miller. Narrows Marina, Seashore State Park, measuring as follows: Trochanter, 48; femur, 9-V-71, D. R. Miller and S. Nakahara. 122; tibia, 104; tarsus, 63; claw, 25; claw digi- Washington, D.C., 1-III-27, H. Y. Gouldman. tules elongate, dilated apically, extending be- Mexico: In quarantine at Nogales, Ariz., yond moderately stout, curved claws. 9-IX-67, J. M. Kaiser. Seven mi N. Perote, With one truncated, cone-shaped circulus. Vera Cruz, 28-11-72, D. R. Miller and F. D. Anal lobes undeveloped, each lobe area with one Parker. elongate seta about 76|i long and two shorter Discussion.—The type material of leucoso- elongate setae. Anal ring large, about 64fi in mus, consisting of six specimens mounted on diameter, its structure distinct, its setae stouter one slide, was remounted in February 1931 than lobe setae, about 89!i long; outer part of and again in May 1944. Since a type was not anal ring with 23-32 mostly oval, elongate designated by Cockerell, I hereby designate cells and some distinctly shorter, irregularly specimen No. 1 at the extreme left on the type rounded cells, many lying end to end but not slide as the lectotype. The midlegs and hind always touching; inner part of ring with 16-20 legs are missing, but otherwise the lectotype is larger, irregularly elongate cells lying next to in good condition. The five paralectotypes are clouded area of semicircular cells. Tritubular also in fair condition except some missing ap- cerores small, 5 on head, 10-13 on thorax, 18-20 pendages and lobe setae. on abdomen, more prevalent on dorsum, few Until recently leucosomus has been known submarginally on venter. Multilocular disk only in the Western United States. It has been pores absent. Tubular ducts smaller than triloc- confused with cacticans, floridanus, and naka- ular pores, present on both surfaces. Trilocu- harai. The San Clemente record by McKenzie lar pores uniformly distributed. Body setae (1967) for California is correct, but the re- short, inconspicuous. maining ones refer to cacticans and nakaharai. Lectotype„—Las Vegas, N. Mex., 11-IV-Ol, The similarity of leucosomus to cacticans W. P. Cockerel]. Paralectotypes.—Five with and what may be intermediate forms of these lectotj^pe on one slide. USNM. species sometimes makes their identification 34 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE difficult. In most specimens they may be readily bodies less than one-half diameter of trilocular separated by comparing details of the antennae, pore, scattered widely over body surface. Tri- rostrum, and anal ring. locular pores fairly abundant, more common dorsally. Few minute, simple circular pores observed ventrally across midabdominal seg- Rhizoecus macgregori^ new species ments. Body setae short, rather sparse. (Figs. 182-191) Holotype.—Apatzingan, Michoacan, Mexico, 19-XI-65, Raúl MacGregor; USNM. Para- Adult Female.—Oval elongate, moderately types.—Two, taken with holotype; IBM. stout. Length, 1.44-1.95 mm; width, 0.65-1.06 Host Plant.—Gossypium hirsutum. mm. Antennae six-segmented, average length Distribution.—Known only from type locality. of segments: I, 44; II, 26; III, 40; IV, 23; V, 26 ; VI, 56 ; apical segment almost twice as long Discussion.—The sclerotized anal lobes, size as wide, with three rather stout, tapering, fal- and abundance of the conspicuous tubular ducts, cate sensory setae and one elongate, spinelike and the presence of minute mushroomlike tub- sensory seta; segment V with short, much ular ducts and circular pores distinguish this smaller sensory seta. Interantennal space equal cotton-inhabiting species from all other mem- to com.bined length of segments II and III. bers of the genus. Eyes small, pigmented, not strongly protuber- It is with pleasure that I dedicate this species ant. Rostrum fairly stout, elongate, about 85|i to Dr. Raúl MacGregor L. of the Instituto de long, 56fi wide ; rostral loop reaching more than Biología, Universidad Nacional, Mexico, D.F. halfway to insertion of second pair of legs. Cephalic plate irregularly triangulate, approxi- mately 45|i-50|i wide, with three or four vacu- Rhizoecus maritimus (Cockerell) oles and bordered by several body setae. Dorsal (Figs. 192-196) ostioles well developed, their rims narrow, sclerotized. Ripersia inaritima Cockerell, 1894: 42. Legs rather elongate, average length of seg- Ripersiella marítima: Cockerell, 1899: 278; Fernald, ments of hind pair : Trochanter, 48 ; femur, 129 ; 1903: 115. tibia, 115; tarsus, 69; claw, 24; claw digitules Morrisonella maritima: Hambleton, 1946a: 31. Rhizoecus maritimus: Ferris, 1953: 452; Hambleton, elongate, dilated apically, extending beyond 1973: 68. basally stout, curved claws. With one prominent, strongly sclerotized Adult Female.—Elongate oval. Length, 1.17- subconical circulus, measuring about 39|i at its 2.51 mm; width, 0.34-1.33 mm. Antennae six- base, its orifice 20{i wide, finely reticulated. segmented, average length of segments: I, 40; Anal lobes little or not protruding; each lobe II, 25; III, 36; IV, 17; V, 21; VI, 55; apical area with small sclerotized patch between three segment almost twice as long as wide, with elongate setae, longest about 88ji long, and few three medium-sized, tapering, falcate sensory short body setae and trilocular pores. Anal ring setae and one acute, spinelike sensory seta ; seg- large, about 81|i in diameter, its setae some- ment V with one much smaller, weakly curved what stouter and longer than lobe setae ; outer sensory seta. Interantennal space about equal part of anal ring containing 42-45 small oval to combined length of segments V and VI. Eyes cells, in places forming double row ; inner part prominent, rather globose, pigmented. Rostrum of ring with about 20 larger, more irregular stout, length about 84j.i, width, 73ji; rostral cells. Tritubular cerores small, 40-60 submar- loop usually short but sometimes reaching in- ginally and across body segments dorsally and sertion of second pair of legs. Cephalic plate ventrally. Multilocular disk pores absent. Tu- approximately 50|.i long, triangulate or quad- bular ducts conspicuous, varying in diameter rate, with two or three vacuoles and six or more from size of trilocular pore to that of tritubular small body setae. Dorsal ostioles fairly well ceroris, stout, strongly sclerotized, widely dis- developed, their rims weakly pigmented, lightly tributed over entire derm. Minute mushroom sclerotized. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 35

Legs moderately stout, average length of I hereby designate the second specimen from segments of hind pair : Trochanter, 55 ; femur, the right on slide No. 1 as the lectotype. 120 ; tibia, 109 ; tarsus, 73 ; claw, 39 ; claw digi- R. maritim us may be readily distinguished tules slender, setose, varying in length but usu- from its closest allies by its long, slender claws, ally about half the length of narrow, elongate, stout rostrum, and claw digitules that are about weakly curved claws. half as long as the claws. With one large truncate circulus, its broad, sclerotized base about 45LI in width. Anal lobes undeveloped, unsclerotized, each lobe area with Rhizoecus m ay anus (Hainbletoii) three elongate setae, longest about 90|.i long, (Figs. 197-204) remaining two slightly shorter. Anal ring large, about 87¡.i in diameter, its structure MoD-isoiicIIa maya}ia Hambleton, 1946a: 32. clearly defined, ring setae each about 85u-90u Rhizoecus maijayins: Ferris, 1953: 454. long, about same thickness as longer lobe setae ; Adult Female.—Oval elongate, moderately outer anal ring composed of 36-40 rather small, stout. Length, 0.89-2.00 mm; width, 0.41-1.04 oval, or irregularly polygonal cells, some not mm. Antennae six-segmented, widely spaced, touching, some with short spicules; cells of average length of segments : I, 38 ; II, 20 ; III, inner part of ring much larger, 26-28 present, 28; IV, 22; V, 18; VI, 46; apical segment cone- more elongate, rectangular, mostly touching shaped, with three moderately stout, weakly end to end, sometimes forming double row and tapering, falcate sensory setae and one slender, lying adjacent to clouded area of large semi- spinelike sensory seta ; segment V with one nar- circular cells. Tritubular cerores small, their row, elongate sensory seta. Interantennal space ducts slightly tapered, occurring on both sur- about equal to combined length of antennal seg- faces, but more common dorsally, 4-5 on head, ments I-IV. Eyes small, protuberant, lightly 10-12 on thorax, 18-20 on abdomen. Multilocu- pigmented. Rostrum elongate, 73¡.i long, 48^ lar disk pores absent. Tubular ducts small, nu- wide; rostral loop extending halfway or more merous, distributed over entire derm. Trilocular to insertion of second pair of legs. Cephalic pores well distributed but sometimes sparse in plate broader than long, about SOj^i long, 48^i areas across segments. Body setae variable in wide, with body seta on each laterocephalic size and length, longer setae on head, thorax, margin. Dorsal ostioles moderately conspicuous, and along body margins. their rims heavily sclerotized, surrounded with Lectotype.—Sea Cliff, N.Y., 7-V-94, Nathan numerous pores and body setae. Banks, mounted on one slide with five paralecto- Legs of medium size, average length of seg- types. Paralectotypes.—Five additional speci- ments of hind pair : Trochanter, 44 ; femur, 99 ; mens mounted on two slides. USNM. tibia, 79; tarsus, 66; claw, 27; claw digitules Host Plants,—Faucaria tigrina, Iva frutes- short, setose, reaching about half the length of cens var. oraría, Rhizophora mangle, Spartina fairly stout, weakly curved claws. alterniflora, S, patens. Circulus absent. Anal lobes weakly protrud- Distribution.—Florida, Massachusetts, New ing, with small, irregular sclerotized area at Jersey, New York. base of larger setae, each lobe with one seta Florida: Bailey's Bluff, 7-III, 3-VIII-71, about 72¡.i long and two shorter elongate setae, G. T. Williams. Cedar Key, 19-VII-70, G. W. several auxiliary setae and body pores. Anal Dekle. Tampa, 17-11-71, E. R. Simmons. ring small, about 43|_i in diameter, its structure Massachusetts: Cape Cod, July 1968, W. J often indistinct, setae averaging about 58j.i Wall. long, shorter and more slender than largest lobe New Jersey: Greenwich, l-XI-44, G. Rau. setae ; outer part of anal ring with 10-12 oval New York : Hampton Bays, Long Island, 14- cells, twice as long as wide, most with spicules V-57 ; North Sea, Long Island, 19-V-57, John and touching end to end; cells of inner part Vack. usually larger and more irregular ; shaded area Discussion.—Since Cockerell (1894) did not not clearly distinguishable. Tritubular cerores designate a holotype in his original description, of 3 sizes, all with short stout ducts, distributed 36 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE as follows: Large size, 12[x-13[x in diameter, Rhizoecus menkei McKenzie confined to dorsum, 5 extending from head to (Figs. 205-211) abdominal segment VI along or near middorsal line ; medium-sized cerores, 9pi-ll|i in diameter, Rhizoecus menkei McKenzie, 1962: 673. occurring ventrally, 1 on head, 3 or 4 along submarginal borders of abdominal segments Adult Female.—Oval elongate, small. Length, V, VII, and VIII ; small cerores, 5\i-7\i in diam- 1.19 mm; width, 0.65 mm. Antennae six- eter, present ventrally from segments IV to segmented, average length of segments : I, 35 ; VIII, 0-3 on IV, 1-5 on V, 2-10 on VI, 5-13 on II, 18; III, 27; IV, 17; V, 17; VI, 45; apical VII, and 3-8 on VIII. Multilocular disk pores segment about twice as long as wide, with three occurring in some numbers on venter of poste- moderately stout, elongate, weakly tapering, rior abdominal segments and in transverse falcate sensory setae and one slender, acute, rows across posterior borders of segments spinelike sensory seta ; segment V with one nar- V-VII, elsewhere, except on head, usually row, shorter sensory seta. Interantennal space widely isolated on both surfaces. Tubular ducts equal to combined length of segments I and usually absent. Trilocular pores rather evenly II. Eyes rather prominent, slightly longer than distributed, not abundant. Body setae short, wide, weakly pigmented. Rostrum elongate, moderately sparse. about 67^ long, 44|i wide ; rostral loop extend- Holotype.—Puerto Barrios, Guatemala, 9-V- ing to or slightly beyond insertion of second 45, E. J. Hambleton. Paratypes.—Four taken pair of legs. Cephalic plate fairly prominent, with holotype, 13 from same location and date measuring 33|i long, 47ii wide. Dorsal ostioles on different host and from Retalhulew, Guate- well developed but weakly sclerotized. mala, 17-V-45, E. J. Hambleton. USNM. Legs with average length of segments of hind Host Plants.—Cymbopogon citratus, C. nar- pair : Trochanter, 44 ; femur, 88 ; tibia, 84 ; tar- dus, Echinochloa colonum, Eleusine indica, sus, 53; claw, 24; claw digitules short, setose, Musa paradisiaca var. sapientum, Panicum not half as long as rather stout, elongate, maximum, Paspalum conjugatum, Setaria geni- curved claws. culata, Xanthosoma robustum, undetermined With one thin-rimmed truncate circulus, its Labiatae. apex faveolate. Anal lobes slightly protruding, Distribution.—Guatemala, Honduras. each lobe area with prominent sclerotized patch, Guatemala: Puerto Barrios and Retalhulew, one seta about 66^i long, two shorter setae, and usual pores and body setae. Anal ring small, as recorded previously. approximately 50[i wide, its structure clearly Honduras: La Lima, l-IX-60, L. Roth; IX, defined, its setae about as long as longest lobe X-67, Carlos Evers. seta, anterior pair somewhat stouter than other Discussion.—This species was originally de- two pairs, latter about as stout as lobe setae; scribed from two imported oil-producing outer part of anal ring with 12-14 elongate grasses, Cymbopogon citratus and C nardus. cells, wide at middle with tapered ends, each These plants are grown commercially in Guate- with spicule ; cells of inner part of ring larger, mala. R, mayanus is here reported on roots of more irregular in shape; clouded area contain- banana in Honduras. ing some large globular cells anteriorly. Bitubu- The larger specimens of mayanus resemble lar cerores with ducts about 8|x long, not americanus; the two species each possess three strongly divaricated, occurring dorsally and sizes of tritubular cerores, but the rostrum of ventrally, 7 on head, 22-25 on thorax, 48 on mayanus is more elongate and its anal ring cells are larger and less sinuate than those of ameri- abdomen, most addominal segments with 5-7. canus. Type specimens of mayanus lack tubular Multilocular disk pores more abundant ven- ducts, yet occasionally an isolated duct is pres- trally from posterior apex to posterior border ent in the Honduran specimens. On the average, of abdominal segment VII, elsewhere usually mayanus has twice as many small tritubular absent except few occurring dorsally along sub- cerores as americanus. marginal borders of segment VIII. Tubular A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 37

ducts very small, about one-half diameter of With one large, truncate, faveolate circulus, trilocular pore, present in small numbers over about 4S[i at its widest diameter, its orifice entire derm. Trilocular pores and short body about 27|i wide, comparable in width to anal setae uniformly distributed over body surface. ring, strongly sclerotized. Anal lobes unde- Holotype.—One mile N. of Tepic, Nayarit, veloped, unsclerotized, each lobe area with one Mexico, 27-VIII-59, A. S. Menke and L. A. seta about 50\x long and two shorter setae. Anal Stange. Paratype.—One taken with holotype. ring 43[i wide, its structure clearly defined, UCD. anterior pair of ring setae about 56^ long, Host Plant.—Unknown, taken in soil. fairly stout, other 2 pairs shorter and about Distribution.—Known only from type locality. half as stout as anterior pair; outer part of Discussion.—R. menkei is more closely re- ring consisting of 11 or 12 rather uniform sinu- lated to hituherculatus than to any other spe- ate cells, each with prominent spicule near its cies. However, the prominent eyes, shorter legs, center; inner part of ring with 10-12 larger, more elongate cells of the outer part of the anal more irregularly elongate cells lying adjacent ring, somewhat longer and more numerous to circular clouded area composed of semicircu- bitubular cerores, and absence of multilocular lar or triangulate cells. Bitubular cerores pres- disk pores from the head and thorax distin- ent, their individual ducts stout, about 9\x long, guish menkei from bitiiherculattis. Contrary to weakly projecting above derm, occurring dor- McKenzie's belief, both of these species have sally and ventrally in about following propor- tubular ducts. tions : 8-9 on head, 20 on thorax, 40 on abdo- men, most abundant dorsally. Multilocular disk pores present on both surfaces but more nu- Rhizoecus mexicanus (Hambleton), merous ventrally on three caudal abdominal new combination segments, occasionally an isolated one occurring (Figs. 212-217) elsewhere. Tubular ducts common, more than half as long as ceroris ducts and smaller in diam- Ripersiella mexicana Hambleton, 1946a: 67. eter than trilocular pore, widely scattered over Adult Female.—Elongate oval. Average entire derm. Trilocular pores sparse ventrally, measurements of two specimens, length, 1.60 especially in thoracic area, elsewhere more or mm; width, 0.87 mm. Antennae six-segmented, less evenly distributed. Body setae variable in comparatively small, moderately widely spaced, length, most short and sparsely distributed over derm. average length of segments : I, 28 ; II, 16 ; III, 30 ; IV, 15 ; V, 14 ; VI, 39 ; apical segment about Holotype.—Mexico, locality unknown, inter- twice as long as wide, with three long, moder- cepted at quarantine in San Francisco, Calif., 3- ately slender, falcate sensory setae and one VI-35. Paratype.—One taken with holotype spinelike sensory seta; segment V with one and mounted on same shde. USNM. much narrower and shorter sensory seta. Inter- Host Plant.—Undetermined succulent and antennal space equal to combined length of seg- cactus. ments I-III. Eyes small, moderately protuber- Distribution.—Texas, Mexico. ant, lightly pigmented. Rostrum elongate, about 73u long, 471.1 wide ; rostral loop reaching to or Texas: Locality unknown, taken at St. slightly beyond insertion of second pair of legs. Petersburg, Fla., via Texas, 7-III-73, C. K. Cephalic plate very small, insignificant, with Hickman. two visible vacuoles. Dorsal ostioles not ob- Mexico : Locality unknown. served. Discussion.—This species was inadvertently Legs of medium size, length of segments of omitted by Ferris (1953) and by McKenzie hind pair : Trochanter, 33 ; femur, 79 ; tibia, 67 ; {1960h, 1967). R. mexicanus closely resembles tarsus, 44; claw, 14; claw digitules elongate, solani but may be distinguished by its larger dilated, reaching beyond rather short, stout circulus, smaller bitubular cerores, and more claws. slender falcate setae. 38 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

Rhizoecus nakaharai, new species fined to dorsum of abdomen. Trilocular pores rather evenly distributed on both surfaces but (Figs. 218-227) less abundant on thorax. Body setae short, Rhizoecus cacticans (Hambleton) : McKenzie, 1967: sparse, longer and more numerous across pos- 381 (misidentification in part). terior abdominal segments. Rhizoecus leucosomus (Cockerell) : McKenzie, 1967: 398 (misidentification in part). Holotype.— Discovery Bay (Jefferson County), Wash., 5-IX-66, Sueo Nakahara; USNM. Para- Adult Female.—Elongate ovate. Length, types.—Five taken with holotype; USNM. 1.25-3.22 mm; width, 0.56-1.73 mm. Antennae Forty additional paratypes from following lo- six-segmented, moderately stout, average cations: California: Coches Prietos, Santa length of segments: I, 44; II, 24; III, 36; IV, Cruz Island, Santa Barbara County, lO-V-68, 22 ; V, 23 ; VI, 51 ; apical segment slightly less D. R. Miller; El Centro, Imperial County, 25- than twice as long as wide, with three rather 11-63, G. 0. Poinar and D. W. Ricker ; one-half narrow, elongate, falcate sensory setae and one hii S. Gold Lake, Sierra County, 3-XI-66, T. R. shorter, narrow, spinelike sensory seta; seg- Haig; N. Grizzley Island, Solano County, 20- ment V with one short, narrow, occasionally X-69, R. 0. Schuster; Los Banos, Merced weakly clávate sensory seta. Interantennal County, 23-IV-68, T. R. Haig; Rohnersville, space equal to combined length of antennal seg- Humboldt County, 29-III-56, B. F. Kemp. ments V and VI. Eyes weakly protruding, often Idaho: Three mi N. Ashton, Fremont County, difficult to observe, and may be absent. Ros- 21-VIII-64, D. R. and J. F. Miller; Craters of trum stout, slightly longer than wide, about 77^i the Moon National Monument, Butte County, long, 68LI wide; rostral loop short, seldom 4-VII-67, D. S. Horning; 8-VIII-67, D. R. reaching halfway to insertion of second pair of Miller and D. S. Horning; 20 mi N. Spencer, legs. Cephalic plate usually longer than wide, Clark County, 5-VIII-67, D. R. Miller and D. S. quadrate or triangulate, with two or three Horning. Montana: II/2 nii SSW. Winett, vacuoles and bordered with several body setae. Petroleum County, via W. H. Burleson, 1971. Dorsal ostioles well developed. Oregon: Mapleton, Larie County, 5-VIII-68, Legs robust, average length of segments of D. R. Miller and R. F. Denno. Washington: hind pair: Trochanter, 45; femur, 108; tibia, Blewett, Chelan County, 4-VIII-66, Sueo Naka- 96; tarsus, 60; claw, 24; claw digitules elon- hara. Thirty-six UCD, four USNM. gate, their apices dilated, extending beyond Host Plants.—Agropyron sp., Carex sp., Cy- basally stout, elongate, curved claws. perns rotnndiis, Distichlis spicata, Dudleya With one conical circulus, its base measuring greenei, Eriogonum heradeoides, Jnncus balti- between 20^ and 30fx. Anal lobes undeveloped, ens, Lithospernmm riiderale. each lobe area with small sclerotized patch between three elongate setae, longest about 74|.i Distribution.—California, Idaho, Montana, long, and two much shorter setae. Anal ring Oregon, Washington. well defined, its periphery often hexagonal- Discussion.—This species has been confused shaped rather than circular, about 69|i in diam- in the literature with cacticans and ¡éticosO'ïmis. eter; its ring setae approximately 85^ long, R. nakaharai resembles lencosomits, particu- much stouter and longer than lobe setae ; outer larly in the size and shape of the rostrum and part of anal ring with 24-48 rather small, oval, the nature of the cellular structure of the anal elongate, or triangulate cells, many almost ring. Hov/ever, the presence of an elongate touching; inner part of ring with 18 large sclerotized area on the anal lobes is character- irregular cells bordered inwardly by circle of istic of nakaharai and will separate it at once large, ovate dark cells. Tritubular cerores small, from leucosomus and cacticans. 36-46 present, 7-10 dorsally along median line It is a pleasure to name this species in honor from head to abdominal segment VIII, and of my friend, Sueo Nakahara, whose dedication 21-36 dorsally and ventrally, most along sub- to the collection and identification of the Coc- marginal borders. Multilocular disk pores coidea has helped increase our knowledge of absent. Tubular ducts small, few, usually con- these insects. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 39

Rhizoeciis netnoralis (Haiiibleton) Holotype.—Along roadside, between Quetzal- tenango and Totonicapan, Guatemala, 10,000 ft (Figs. 228-234) elevation, 20-V-45, E. J. Hambleton. Para- MorrisoncUa nemoralis Hambleton, 1946a: 33. types.—Two taken with holotype. USNM. Rhizoeciis nemoralis: Ferris, 1953: 456. Host Plants.—Alcliemilla orbiculata, Citrus s ¡neu sis. Adult Female.—Broadly ovate. Length, 1.64- Distribution.—At type locality and Antiqua, 2.08 mm; width, 1.00-1.19 mm. Antennae six- Guatemala, 25-V-45, E. J. Hambleton. segmented, moderately stout, widely spaced, Discussion.—Ferris (1953) was not convinced average length of segments: I, 46; II, 24; III, that nemoralis and cypemlis were distinct. 39; IV, 22; V, 21; VI, 50; apical segment less This was probably because of the limited than twice as long as wide, tapered, with three amount of material available for study. Based moderately stout, bluntly tapered, falcate sen- on my studies it is clear that these species are sory setae and one spinelike sensory seta ; seg- closely related. Until more material of cypera- ment V with one short, stout, lanceolate sensory Us is available, it seems advisable to consider seta. Interantennal space equal to combined the two as distinct. See discussion under length of segments II-VI. Eyes small but prom- eyperalis, page 22. inent, subconical, pigmented. Rostrum elongate, 98|.i long, 51ii wide ; rostral loop reaching almost to insertion of second pair of legs. Cephalic Rhizoeciis neomexicanus McKenzie plate absent. Dorsal ostioles prominent, heavily (Figs. 235-240) sclerotized, surrounded by numerous pores and body setae. Rhizoeciis veomexicanns McKenzie, 1962: 674. Legs stout, average length of segments of Adult Female.—Oval elongate. Length of hind pair: Trochanter, 67; femur, 135; tibia, holotype, 0.83 mm; width, 0.37 mm. Antennae 100 ; tarsus, 86 ; claw, 27 ; claw digitules short, six-segmented, length of segments as follows: acute, extending to about one-half the length I, 29 ; II, 19 ; III, 17 ; IV, 16 ; V, 13 ; VI, 34 ; api- of long, stout claws. cal segment tapered, less than twice as long as Circulus absent. Anal lobes weakly protrud- wide, with three rather stout, falcate sensory ing, without sclerotization, each lobe area with setae, their tips weakly tapered, and one elon- three elongate setae, longest about 92[.i long gate, more acute sensory seta ; segment V with and five to eight short body setae and numerous one elongate, narrow sensory seta. Interanten- pores. Anal ring about 49|i wide, its structure nal space equal to less than length of segment fairly distinct, its setae averaging about 72j.i I. Eyes small, with little pigmentation. Ros- long, shorter and more slender than largest lobe trum approximately 63!.i long, 44|.i wide ; rostral seta ; outer part of anal ring with 10-12 irregu- loop extending little beyond insertion of second larly elongate cells, some with elongate spicules ; pair of legs. Cephalic plate triangular, about cells of inner part somewhat larger, 8 lying 35^1 wide. Dorsal ostioles moderately conspicu- adjacent to clouded area of large oval-shaped ous, their rims weakly pigmented. cells. Tritubular cerores of 2 sizes, larger or Legs relatively short, of medium size, seg- medium size occurring dorsally, 5 along dorso- ments of middle pair of legs as follows : Tro- median line from head to abdominal segment chanter, 25; femur, 70; tibia, 47; tarsus, 42; VIII ; 5 or 6 along submarginal borders ; 28-37 claw, 18; claw digitules short, acute, about one- small cerores present ventrally near body mar- half as long as slender, acute claws. gin and scattered elsewhere. Multilocular disk Circulus absent. Anal lobes somewhat pro- pores numbering 5-11, confined to venter near duced and slightly sclerotized, each lobe with vulva. Tubular ducts few, also occurring in three elongate setae of varying lengths, longest area of vulva. Trilocular pores rather numerous about 45LI, slightly longer and stouter than ring and evenly distributed. Body setae conspicuous, setae. Anal ring about 38|.i wide, its setae aver- variable in length, longest setae approximately aging about 44j.i long; outer part of anal ring 50(:x long. not well defined, cells appearing to be longer 40 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

than wide ; inner part of ring indistinct. Tritu- 21 ; V, 22 ; VI, 55 ; apical segment not twice as bular cerores short, stout, of two sizes, larger long as wide, with three elongate, medium cerores present only on dorsum, one on head, thick, rather bluntly tapered, falcate sensory five on thorax, one on abdominal segment V, setae and one spinelike sensory seta; segment and two on segment IX ; smaller cerores occur- V with one elongate, narrow, weakly clávate, ring ventrally, two on abdominal segment V, curved sensory seta. Interantennal space equal four on VII, and two on IX. Multilocular disk to combined length of segments IV-VI. Eyes pores present on both surfaces but more nu- small, rather prominent, lightly pigmented. merous ventrally from posterior border of ab- Rostrum 86¡i long, 62ji wide; rostral loop ex- dominal segment VII to apex, few scattered tending only slightly beyond apex of rostrum. elsewhere on abdomen and thorax, absent from Cephalic plate broader than long, 33ji long, 39u head. Tubular ducts distributed lightly over wide, with body seta on each dorsolateral mar- entire derm. Trilocular pores most numerous gin. Dorsal ostioles well developed, heavily scle- dorsally, sparsely scattered ventrally. Body rotized, with numerous trilocular pores along setae short, some longer setae on head and their rims. along posterior abdominal margins. Legs rather large, segments of hind pair Holotype.—Alazán, Vera Cruz, Mexico, 15- measuring as follows: Trochanter, 60; femur, VIII-59, A. S. Menke and L. A. Stange. Para- 124 ; tibia, 101 ; tarsus, 77 ; claw, 29 ; claw digi- type.—One taken with holotype. UCD. tules short, setose, not reaching middle of Host Plant.—Unknown, taken in jungle soil. rather long, stout, weakly curved claws. Distributian.—Known only from the type lo- Circulus absent. Anal lobes weakly protrud- cality. ing, each lobe area with small sclerotized patch, Discussion.—Three slides labeled Rhizoecus three elongate setae, longest about 66j-i long, neomexicanus McKenzie represent the type and usual pores and setae. Anal ring not clearly material, which forms the basis for the previous defined, about 53ji wide, its setae about same redescription of this species. Unfortunately thickness as lobe setae but slightly shorter than there are two species among the five individuals longest, averaging 63pi long ; outer part of ring mounted on the three slides. One slide labeled probably with no more than 10-12 elongate, "type^^ has the true neomexicanus and an im- oval cells most with small spicules ; cells of inner mature female of tropicalis, new species. A part of ring and clouded area not discerned. second slide labeled "paratype" contains one Tritubular cerores of 2 sizes, 19 large ones oc- female of neomexicanus in the process of molt- curring dorsally, 7 along middorsal line from ing and one apparently mature female of tropi- head to abdominal segment VI, 12 submargin- calis. On the third slide, also labeled *'paratype'' ally along thorax and abdomen; 10 medium- is an adult female of tropicalis but no neomexi- sized cerores present ventrally from abdominal canus. The holotype of neomexicanus probably segments V to VIII. Multilocular disk pores had not attained its full life size and is not in present on venter only, about 60 located from the best condition; therefore, its redescription posterior margin of abdominal segment VII to is incomplete. anal ring and few between or near legs. Tubu- R, neomexicanus is smaller and has fewer lar ducts absent. Trilocular pores distributed tritubular cerores than caladii. dorsally and ventrally. Body setae rather short, slender. Rhizoecus neostangei Miller and McKenzie Holotype.—Jalapa (Rio Cedeno), Vera Cruz, (Figs. 241-245) Mexico, 16-VIII-67, D. R. Miller and J. Vil- lanueva B. ; UCD. Rhizoecus neostangei Miller and McKenzie, 1971: 588. Host Plant.—Rhus sp.. Adult Female.—Oval elongate, stout. Length, Distribution.—Known only from the type lo- 1.86 mm; width, 1.08 mm. Antennae six- cality. segmented, moderately stout, widely spaced, Discussion.—This species is closely related to length of segments : I, 44 ; II, 23 ; III, 26 ; IV, R. stangei but differs from it in lacking dorsal A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 41

multilocular disk pores and in possessing fewer from head to abdominal segment VIII ; smaller medium-sized tritubular cerores on its venter. cerores numbering between 45 and 51 ventrally in more or less transverse rows across abdomi- nal segments IV-IX ; segment VII with as many Rhizoecus nitidalis Hambleton as 15 in some specimens. Multilocular disk pores (Figs. 246-250) of five to seven loculi distributed on both sur- faces, but more numerous on posterior abdomi- Rhizoecus nitidalis Hambleton, 1946a: 57. nal segments, gradually diminishing in num- bers anteriorly and absent from frons. Tubular Adult Female.—Broadly ovate, stout. Length, 1.70-3.06 mm; width, 0.94-1.99 mm. Antennae ducts about same diameter as trilocular pores, five-segmented, large, average length of seg- apparently confined to abdominal area. Trilocu- ments: I, 55; II, 25; III, 45; IV, 31; V, 98; lar pores almost circular, rather uniformly dis- segment I rather stout, apical segment approxi- tributed. Minute circular pores half the size of mately three times as long as wide, with four trilocular pores scattered over most of derm. stout, elongate, strongly tapered, falcate sen- Body setae usually short, variable in size and sory setae, posterior seta smaller, shorter, and length, some longer setae along body margins and on head. one narrow, spinelike sensory seta near apex. Interantennal space equal to slightly more than Holetype.—Guaruja. State of Sao Paulo, Bra- width of segment I at its base. Eyes absent. zil, 22-VIII-35, B. L. Ribeiro and E. J. Hamble- Rostrum elongate, averaging 106¡i long, TOji ton. Paratypes.—Five mounted with holotype wide; rostral loop extending halfway or more on same slide and 30 from other host at same to insertion of second pair of legs. Cephalic location. USNM. plate twice as wide as long, 82^ wide, 39^ Host Plants.—Axonoptis sp., Cenchriis echin- long, with three to six vacuoles near and along atiis, Paspahim vaginatiim. its posterior border and several body setae along Distribution.—Known only from the type lo- anterior margin. Dorsal ostioles conspicuous, cality. their rims narrow but strongly sclerotized. Discussion.—R. nitidalis resembles falcifer Legs well developed, large, with considerable in general appearance. It differs in having scle- variation in size, average lengths of segments rotized anal lobes with fewer elongate setae, of hind pair : Trochanter, 67 ; femur, 160 ; tibia, fewer and different distribution of cerores, its 119; tarsus, 80; claw, 40; claw digitules very multilocular disk pores with fvve to seven loculi, short, finely setose, less than one-half the length and in having shorter and more sparsely dis- of long, narrow, weakly curved claws. tributed body setae. Circulus absent. Anal lobes slightly protrud- ing, each lobe with irregularly rounded scle- rotized patch, three elongate, stout setae, long- Rhizoecus ornatus (Hambleton), est about 8l!i long, five or six short auxiliary new combination setae, and small concentration of pores. Anal (Figs. 251-258) ring about 58J.I in diameter, of simple, concise design, its setae averaging about 60!_i long, Ripersiella ornata Hambleton, 1946a: 70. shorter and weaker than largest lobe seta; Adult Female.—Oval elongate. Length, 1.44 outer part of anal ring with 10 or 12 elongate, mm; width, 0.75 mm. Antennae six-segmented, pointed cells, each with spicule; cells of inner relatively small, length of segments : I, 29 ; II part of ring irregularly elongate and larger, 10 20; III, 24; IV, 16; V, 16; VI, 37; apical seg- usually touching and lying adjacent to undif- ment less than twice as long as wide, with three ferentiated shaded area. Tritubular cerores of stout, bluntly tapered, falcate sensory setae 2 sizes present, larger about 12u wide, smaller and one elongate, spinelike, apical sensory seta; about 5^1 wide, both with short, stout ducts; segment V with one short, stout, lanceolate sen- large cerores occurring dorsally, 16 present on sory seta constricted near its base. Interanten- median line and along or near body margins nal space less than width of segment I. Eyes 42 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

rather prominent, protruding, weakly pig- Because of their unusual morphological char- mented. Rostrum about 60^ long, 38|i wide; acters, they differ from other members of the rostral loop extending to or slightly beyond in- genus. This species may be distinguished by sertion of second pair of legs. Cephalic plate its tritubular cerores with their short, stout present, weakly pigmented, not well defined. ducts. Dorsal ostioles inconspicuous, their rims thin, Borchsenius (1949) described a species under weakly sclerotized. the name ''Rhizoecits ornatus'' from the Cri- Legs of medium size, length of segments of mea. Since ornatiis Borchsenius is a secondary hind pair: Trochanter, 35; femur, 74; tibia, homonym of ornata, the name of the Crimean 65 ; tarsus, 41 ; claw, 25 ; claw digitules elongate, species should be changed. The Trinidad and weakly dilated apically, reaching tip of long, Crimean taxa are both valid species. narrow, curved claws. With two elliptical circuli about 21fx wide, with thinly sclerotized rims, their orifices re- Rhizoecus ovatus^ new species ticulose, with 15-18 polygonal-shaped cells. Anal lobes undeveloped, unsclerotized, each (Figs. 259-263) lobe area with three slender, elongate setae, Adult Female.—Broadly ovate. Length, 1.58 longest about 38|i long and three or four body mm; width, 1.05 mm. Antennae six-segmented, setae. Anal ring unique, 65^ wide, its setae comparatively short and stout, each about 55}.i widely spaced, average length of segments: I, long, setae shorter than diameter of ring ; outer 48 ; II, 26 ; III, 47 ; IV, 21 ; V, 26 ; VI, 65 ; apical segment twice as long as wide, with three nar- part of anal ring with 10 large, isolated, oval, row, elongate, strongly tapered, falcate sen- elongate cells, 2-3 times longer than wide, each with prominent spicule; inner part of ring sory setae and one spinelike sensory seta; seg- with 6-8 more elongate cells lying adjacent to ment V with one shorter, narrow sensory seta. Interantennal space equal to combined length darkened area or concentric ring of large oval cells. Tritubular cerores of 2 distinct types, of segments V and VI. Eyes absent. Rostrum larger ceroris strongly depressed with short rather large, conical, 95j.i long, 73|i wide; ros- ducts, about 7[i in diameter, occurring dor- tral loop extending almost halfway between sally in interrupted rows across segments VII insertion of second and third pair of legs. Ce- and VIII, 9-11 on VII, and 19-22 on VIII; phalic plate inconspicuous, weakly sclerotized, smaller cerores of more normal type with short its shape undiscernible. Dorsal ostioles well ducts, about 4^ in diameter, usually encircled developed, but with weakly sclerotized rims. by trilocular pores, more common on head and Legs long, average length of segments of thorax and along body margins on both sur- hind pair: Trochanter, 55; femur, 134; tibia, faces. Multilocular disk pores absent. Tubular 101 ; tarsus, 55 ; claw, 28 ; claw digitules slen- ducts not observed and considered absent. Tri- der, elongate, setose, reaching to or slightly locular pores distinctly triangulate, rather nu- beyond stout, strongly curved claws. merous, tending to form bands across abdomi- With one truncate circulus, its orifice about nal segments. Body setae small, some longer 14u in diameter. Anal lobes undeveloped, un- setae across abdominal segments, along body sclerotized, each lobe area with two stout, elon- margins, and on head. gate setae, longest about 86j-i long, and one Holotype.—Trinidad, British West Indies, 3- shorter, more slender seta and five or six III-44, A. H. Strickland. Paratypes.—Two smaller auxiliary setae. Anal ring large, well from same location, 6-VI-35, E. J. H. Berwick. defined, about 78^ wide, its setae about same USNM. size as lobe setae but appearing to be slightly Host Plants.—Coffea arabica, Theobroma longer ; outer part of anal ring consisting of cacao. 35-40 rather small, irregularly quadrate or Distribution.—Known only from type locality. rounded cells, most being longer than wide, Discussion.—R. ornatus is one of several in- isolated, and in places forming a double row ; teresting hypogeic mealybugs from Trinidad. inner part of ring with about 20 much larger A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 43

and irregularly oval, elongate cells, their ends rotized, surrounded by numerous pores and touching and lying adjacent to ill-defined body setae. darkened area. Tritubular cerores small to Legs apparently rather large, all broken ex- medium, their ducts weakly tapered, at least cept right hind leg, its dimensions as follows: 45-50 occurring dorsally, and uniformly dis- Trochanter, 66; femur, 138; tibia, 90; tarsus, tributed from head to abdomen. Multilocular 88; claw, 30; claw digitules probably short, disk pores absent. Tubular ducts rather promi- setose, not observed ; claws stout. nent, short, stout, 6u-7u long, slightly smaller Circulus absent. Anal lobes weakly protrud- in diameter than tritubular ceroris, distributed ing, each lightly sclerotized, and with three dorsally and ventrally, anterior abdominal seg- elongate setae, longest about 85u long, (most ments each with 7 to 11 tubular ducts. Triloc- of setae lost in mounting), and with concentra- ular pores more abundant dorsally. Body setae tion of pores. Anal ring about 46u wide, its short, longer, and more abundant dorsally. structure not clearly defined, its setae shorter Holotype.—Mexico, location unknown, taken and somewhat more slender than largest lobe in quarantine at Bellflower, Calif., 2-IV-40, seta ; outer part of anal ring with no more than L. E. Meyers ; UCD. Paratypes.—Three taken 12-14 oval, elongate cells, some with spicules; with holotype, one UCD, two USNM. inner part with fewer similar cells, darkened Host Plant.— Mammülaria sp. area indistinct. Tritubular cerores present in Distribution.—Known only from Mexico. 2 sizes, larger about 8¡.(-9^i in width, their indi- Discussion.—With the exception of falcifer, vidual ducts short, stout, 20 occurring dorsally, this is the only known Mexican species that 6 along median line from head to abdominal lacks eyes. The presence of prominent tubular segment VII, 7 submarginally along both sides ducts, their size, abundance, wide distribution, from head to abdominal segment VIII ; smaller and slender, tapering, falcate sensory setae are cerores, normally 8 occurring ventrally, 4 on distinguishing features of this species. One thorax, 1 pair each on segments VII and VIII specimen has circuli on the venter of each of in submarginal areas. Multilocular disk pores three abdominal segments, a normal-sized cir- inconspicuous, only three, each with six locuh, culus on abdominal segment IV, and a smaller near vulva. Tubular ducts present, strongly circulus on each of segments III and V. sclerotized, equal to or larger in diameter than trilocular pores, more numerous near vulva, few scattered elsewhere. Trilocular pores fairly Rhizoecus pauciporus^ new species large, variable in size, some almost circular, most abundant, and uniformly distributed dor- (Figs. 264-268) sally. Circular pores about one-half the diam- Adult Female.—Broadly ovate. Length, 1.87 eter of trilocular pore present dorsally in small mm; width, 1.05 mm. Antennae six-segmented, numbers, more visible across and near poste- rather stout, tapering toward apex, length of rior borders of segments V and VI. Body setae segments: I, 50; II, 22; III, 40; IV, 24; V, 21; usually short, relatively sparse, some longer setae on head. VI, 50; apical segment about twice as long as wide, with three moderately stout, elongate, Holotype.—Ten mi NW. Comitan, Chiapas, falcate sensory setae and one smaller, elongate, Mexico, 22-VIII-67, J. Reddell, J. Fish, and T. spinelike sensory seta near its apex; segment Evans ; USNM. V with one shorter, fairly stout, lanceolate sen- Host Plant.—Unknown, taken in surface de- sory seta. Interantennal space equal to com- bris. bined length of segments I, II, and III. Eyes Distribution.—Known only from type locality. prominent, longer than wide, strongly pig- Discussion.—The shape of the antennae and mented. Rostrum elongate, 103|_i long, 64|.i eyes, the sparsity of multilocular disk pores, wide; rostral loop nearly reaching insertion of its well-defined tubular ducts, and presence of second pair of legs. Cephalic plate apparently circular pores separate this species from its absent. Dorsal ostioles prominent, heavily scle- relatives. 44 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

Rhizoecus poensis (Hambleton), posterior abdominal area, occurring on both new combination surfaces over entire derm. Tubular ducts ab- sent. Trilocular pores uniformly distributed. (Figs. 269-273) Medioventral pores in 2 clusters ventrally on abdominal segments VI and VII, anterior clus- Morrisonella poensis Hambleton, 1946a: 35. ter with 5-12 pores, posterior cluster with 5-14 Adult Female.—Elongate oval, moderately pores. Body setae moderately dense, variable stout. Length, 1.54-1.96 mm; width, 0.77-1.12 in size, 10ji-55|i long, longest on head and mm. Antennae six-segmented, widely spaced, along body margins. average length of segments: I, 45; II, 28; III, Holotype.—Bogota, Colombia, at 10,000 ft, 30 ; IV, 27 ; V, 22 ; VI, 56 ; apical segment ta- 27-X-44, E. el. Hambleton. Paratypes.—Nine pered, not quite twice as long as wide, with taken with holotype, two mounted on same three moderately stout, weakly tapered, falcate slide with holotype. USNM. sensory setae and one spinelike sensory seta; Host Plant.—Poa annua. segment V with one narrow, extremely elon- Distribution.—Known only from type locality. gate sensory seta. Interantennal space about Discussion.—R. poensis is one of three spe- equal to combined length of segments IV-VI. cies having two clusters of medioventral pores. Eyes prominent, protuberant, weakly pig- Both glohoculus and theobromae possess these mented. Rostrum stout, 77^ long, 65^ wide; pores and are closely related to poensis. How- rostral loop extending approximately halfway ever, poensis may be distinguished by its un- to insertion of second pair of legs. Cephalic sclerotized anal lobes, presence of multilocular plate poorly defined, wider than long, about disk pores on the head, and the more slender 44|i wide, containing numerous indistinct vac- falcate setae. uoles. Dorsal ostioles inconspicuous, thinly rimmed, weakly sclerotized, anterior pair al- most indistinguishable, posterior pair with Rhizoecus polyporus^ new species numerous body setae surrounding them. (Figs. 274-282) Legs robust, average length of segments of hind pair: Trochanter, 61; femur, 140; tibia, Adult Female.—Oval elongate. Length, 1.73 103 ; tarsus, 85 ; claw, 33 ; claw digitules short, mm; width, 1.02 mm. Antennae six-segmented, setose, reaching middle of moderately stout, elongate, rather stout, length of segments: I, curved claws. 44 ; II, 30 ; III, 50 ; IV, 24 ; V, 29 ; VI, 62 ; apical Circulus absent. Anal lobes simple, weakly segment at least twice as long as wide, with developed, unsclerotized, each lobe with three three elongate, fairly slender, falcate sensory elongate setae, longest about 85[i long, and six setae and one shorter, spinelike sensory seta to eight short auxiliary setae. Anal ring about near its apex; segment V with one much 50!i in diameter, its setae long and slender, shorter, lanceolate sensory seta. Interantennal slightly narrower than stoutest lobe seta, aver- space about equal to length of segment VI. Eyes aging about 73fx long; outer part of anal ring small, rather depressed, pigmented. Rostrum with 10-12 narrow, elongate cells, each with stout, 106f>i long, 77fi wide ; rostral loop extend- spicule; inner part of ring with cells similar ing between insertion of second and third but somewhat larger; clouded area with oval- pair of legs. Cephalic plate arrowhead-shaped, shaped cells. Tritubular cerores of 2 distinct longer than wide, about 60|i long, 37ji wide, sizes, their ducts short, stout, larger size about with vacuoles near its center and two body 8!i-9|i in diameter, occurring dorsally, 7-10 setae on its posterior margin. Dorsal ostioles along each body margin, 5-10 along middorsal well developed, strongly sclerotized. line; smaller cerores about 6ii-7|i in diameter, Legs rather large, segments of hind pair occurring ventrally, 5-9 submarginally along measuring as follows: Trochanter, 61; femur, each side and usually 1 pair near midventral 156 ; tibia, 136 ; tarsus, 77 ; claw, 23 ; claw digi- area on each of segments V-VIII. Multilocular tules setose, elongate, extending beyond stout, disk pores relatively abundant, especially in apically curved claws. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 45

With two circuli (probably only one occur- Rhizoecus pritchardi McKenzie ring normally), one large, truncate, about 50[,i in diameter, 34u across its finely faveolate (Figs. 283-289) orifice, located on segment IV, and smaller one Rhizoecii.^ fjritchardi McKenzie, 1960b: 749; McKenzie, of similar shape on segment III. Anal lobes 1967: 400: Hambleton, 1973: 69. undeveloped, without sclerotization, each lobe Rhizoecus cypcralis (Hambleton): McKenzie, 1967: 383 area with three elongate setae, two of them (misidentification). about 80|i long, third much shorter, and sev- Rhizoecus eluminatus McKenzie, 1960b: 747; McKenzie, 1967: 387. NEW SYNONYMY. eral body setae and two cerores. Anal ring con- spicuous, measuring S9\i in diameter, its struc- Adult Female.—Elongate oval, moderately ture clearly defined, its setae stouter and longer stout. Length, 1.62-2.10 mm; width, 0.95-1.33 than lobe setae, anterior pair about 95[.i long; mm. Antennae six-segmented, rather stout and outer part of anal ring consisting of 48-50 widely spaced, average length of segments: I, rather small, oval, diversiform, isolated cells, 40 ; II, 21 ; III, 33 ; IV, 18 ; V, 18 ; VI, 43 ; apical in places forming double row; inner part of segment narrowest, not twice as long as wide, ring with circle of larger, irregularly shaped with three moderately stout, falcate sensory cells, about 20 lying adjacent to shaded un- setae and one spinelike sensory seta near apex ; differentiated area. Tritubular cerores of me- segment V with one much shorter, lanceolate dium size, their individual ducts about lOji sensory seta. Interantennal space equal to long, rather strongly divaricating and weakly twice width of segment I. Eyes small, often tapered, projecting well above derm and occur- weakly pigmented, difficult to observe. Rostrum ring dorsally and ventrally in large numbers, elongate, 88[^i long, 52ji wide; rostral loop not approximately 130 on head and thorax, 108 on reaching insertion of second pair of legs. Ce- abdomen; dorsally cerores are arranged in phalic plate absent. Dorsal ostioles well devel- rather uniform rows across abdominal seg- oped, their rims strongly sclerotized. ments. Multilocular disk pores absent. Tubular Legs normal for size of , moderately ducts short, stout, strongly sclerotized, 5ji- stout, elongate, average length of segments of 6|i in diameter, slightly smaller |:han cerores, hind pair: Trochanter, 49; femur, 109; tibia, present over entire derm, some abdominal seg- 82 ; tarsus, 71 ; claw, 25 ; claw digitules short, ments with as many as 20. Minute tubular acute, reaching about to middle of stout, curved ducts, referred to as ''mushroom bodies,'' scat- claws. tered widely over both surfaces. Trilocular Circulus absent. Anal lobes slightly devel- pores fairly numerous, more abundant dorsally. oped or weakly protruding, unsclerotized, each Body setae variable in length, short, evenly lobe with one long and two shorter elongate distributed. setae, longest about 84|i long, and several body Holotype.—State of Sonora, Mexico, 29-111- setae. Anal ring comparatively small, about 67, intercepted at quarantine at Nogales, 50[i wide, its setae shorter and narrower than Ariz. ; USNM. longest lobe seta ; outer part of anal ring with Host Plant.—Unknown, taken in cacti debris. 12 elongate, oval cells, each with spicule; inner Distribution.—Known only from the type lo- part of ring with cells of similar shape but cality. larger; darkened area indistinct (anal ring of this species often assumes vertical position Discussion.—This species and relativus are upon mounting and thus makes it difficult to alike in having large, unusually stout tubular ascertain the nature of the cellular structure). ducts and an abnormally large number of tri- Tritubular cerores of 2 sizes, 25-28 of larger tubular cerores. Both the size of the tubular size present dorsally, 3 on head, 8-10 on tho- ducts and large number of cerores are uncom- rax, 13-14 on abdomen; slightly smaller cerores, mon for Rhizoecus. R. polyporiis differs from 4-6 occurring submarginally on abdomen. relativus in having longer legs and antennae, Multilocular disk pores, varying in size and larger circulus, slender, nontapered sensory shape, 11-21 present ventrally, usually near setae, and more numerous cerores. vulva on segments VIII and IX, occasionally 46 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE 1 or 2 on VII. Tubular ducts absent. Trilocular of eyes. Close examination of the types indi- pores abundant, well distributed, some areas cates the presence of small, inconspicuous eyes of head and thorax occasionally with few pores. in both species. Examination of a series of Circular pores occurring dorsally, very small, pritchardi specimens reveals that the eyes may about one-half size of trilocular pore, from be present or absent. In all other respects elumi- few to as many as 22 scattered across posterior natus and pritchardi are similar and therefore borders of abdominal segments IV, V, and VI. are here treated as synonyms. The name Body setae variable in size, short and long setae "pritchardi'' has been selected as senior syno- about evenly distributed. nym because of its usage in economic literature. Holotype.—Colma, Calif., 17-1-57, E. L. La- R. pritchardi is readily separated from its badie and D. J. Bingham. Paratypes.—Several closest allies by the absence of tubular ducts taken with holotype, UCD; paratypes also and presence of circular pores. It has more CAS, CDA, USNM. Holotype of eliiminatus, multilocular disk pores than either cyperalis Bayside, Calif., 24-IX-54, W. D. Thomas; para- or nemoralis, two of its nearest relatives. types taken with holotype. UCD, one paratype USNM. Host Plants.—Achillea ageratifolia, AdiaU'- Rhizoecus relativus^ new species him sp., Arctostaphylos sp.. Chrysanthemum (Figs. 290-293) sp., Geiim coccineum, Lantana sp., Polygala crotalariodes, Saintpmdia ionantha, Sinningia Adult Female.—Elongate oval. Length, 1.95 speciosa. mm; width, 1.01 mm. Antennae six-segmented, Distribution.—California, Florida, Illinois, moderately stout, length of segments: I, 36; Maryland, Massachusetts, New York, Pennsyl- II, 25; III, 40; IV, 25; V, 24; VI, 53; apical vania, Canada. segment robust, not twice as long as wide, with California : Bayside, 24-IX-54, W. D. Thomas. three elongate, moderately stout, tapering, fal- Bellflower, 12-1-67, T. and H. Ryner. Berke- cate sensory setae and one shorter, spinelike ley, 25-IX-59, A. E. Pritchard. Colma, 24-11- sensory seta ; segment V with one much smaller, 55, W. E. Davis. Escondido, 21-11-56, A. A. short, lanceolate sensory seta. Interantennal Church. Napa, 30-XI-64, D. R. Hall. San space equal to about combined length of seg- Bruno, 29-IV-58, A. E. Pritchard. San Fran- ments I and II. Eyes absent. Rostrum stout, cisco, 29-IV-60, R. Michelsen. Santa Barbara, approximately 72|.i long, 65^ wide; rostral 18-11-60, M. Suskin. Visalia, 11-III-64, Phil loop reaching midway between first and second Hemphill. pairs of legs. Cephalic plate longer than wide, Florida: Apopka, 1-IV, ll-V-71, R. M. Rem- irregular in outline, narrow anteriorly, about ington. Gainesville, 23-V-68, E. Mercer. 53fi long, 40ji wide, with four to five body Largo, 22-IV-66, J. R. McFarlin. setae on its borders. Dorsal ostioles inconspicu- Illinois: Elizabeth, 5-IV-64, F. F. Smith. ous, hind pair more prominent, all with weakly Maryland: Baltimore, 22-IV-65; Beltsville, sclerotized rims. 15-XII-65, 2-III-70, F. F. Smith. Cheverly, Legs comparatively short, stout, (hind pair 4-XI-71* W. W. Cántelo. and one of second pair broken), length of seg- Massachusetts: Waltham, 26-III-70, A. G. ments of midleg : Trochanter, 41 ; femur, 94 ; Gentile. tibia, 69 ; tarsus, 48 ; claw, 19 ; claw digitules New York: Ithaca, 8-XII-61, D. S. Welsh. elongate, weakly dilated, extending beyond Scotia, 25-III-65, G. V. Johnson. rather stout, acute, curved claws. Pennsylvania : York, 22-VII-65, F. F. Smith. With one medium-sized truncate circulus, Canada: Niagara Falls, Ontario, 22-XI-48, 34fx at its base, 13fi across its orifice. Anal lobes R. Sheppard. undeveloped, unsclerotized, each lobe area with Discussione—McKenzie (1960b), in describ- three elongate setae, longest about 77{x long, ing eluminatus, indicated the close relationship several shorter setae, body pores, and several between it and pritchardi, pointing out that cerores. Anal ring about 68ji in diameter, its eluminatiis differed principally in the absence cellular structure well defined, its ring setae A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 47

longer and stouter than lobe setae, longest weakly tapered, falcate sensory setae and one about 75LI long; outer part of anal ring with spinelike sensory seta near apex; segment V 36-38 cells varying considerably in size and with one shorter, clávate sensory seta. Inter- shape, some rounded, elongate, or triangulate, antennal space about equal to length of segment rather indiscriminately arranged; inner part I. Eyes weakly protruding, lightly pigmented. of ring with 14-16 irregularly elongate, larger Rostrum medium size, about 50u long, 35i.i cells lying next to darkened area consisting wide; rostral loop extending almost to insertion of about 10 hemispherically shaped cells. Tri- of second pair of legs. Cephalic plate irregular tubular cerores small to medium, projecting in outline, 37u long, 35ü wide, containing two well above derm, their ducts divaricating, pres- vacuoles and bordered by five or six body ent on both surfaces but more abundant dor- setae. Dorsal ostioles inconspicuous, lightly sally, total of about 85 on head and thorax, 60 sclerotized. on abdomen. Multilocular disk pores absent. Legs small, rather stout, segments of hind Tubular ducts short, stout, strongly sclero- pair measuring as follows: Trochanter, 28; fe- tized, diameter varying from that of triangular mur, 58; tibia, 56; tarsus, 36; claw, 15; claw pore to ceroris, scattered widely over both sur- digitules elongate, dilated at tips, extending to faces. Mushroom bodies widely distributed end of long, rather acute claws. over entire derm. Trilocular pores fairly uni- With one more or less truncate circulus, 13¡.i formly distributed, more numerous on poste- wide at its base, its orifice roughly faveolate. rior abdominal segments but sparsely arranged Anal lobes undeveloped, unsclerotized, each in area along anterior and posterior margins lobe area with three slender, elongate setae, of segments. Body setae short, varying some- longest about 37u long. Anal ring about 46j.i what in length, evenly distributed. wide, its setae averaging about 50u long, Holotype.—Three mi NE. Nogales Station, stouter than lobe setae ; outer part of anal ring Cruz Co., Ariz., l-VIII-66, D. R. Miller; UCD. consisting of 17-23 oval cells with spicules; Host Plant.—Undetermined. inner part of ring with about 14 larger diversi- Distribution.—Known only from type locality. form cells lying adjacent to circular area of Discussion.—The presence of large tubular darkened hemispherical cells. Tritubular cer- ducts separates this species from leucosomiis. ores small, variable in number, 48-68 present, As its name suggests, relativiis is closely re- most numerous dorsally, their ducts elongate, lated to polyporus, but it differs from the latter about 7^1-8^1 long, and projecting well above in having shorter legs and antennae, a smaller derm. Multilocular disk pores absent. Tubular circulus, differently shaped falcate setae, and ducts small, elongate, present on both surfaces fewer cerores. In relativiis the eyes are lacking but more numerous ventrally. Trilocular and the microscopic mushroom bodies are pores evenly distributed, not numerous. Body poorly defined. setae short, slender, rather sparse. Lectotype.—Sao Paulo City, Sao Paulo, Bra- zil, lO-IV-35, B. L. Ribeiro and E. J. Hambleton. Rhizoecus simplex (Hambleton) Paralectotypes,—Four taken with lectotype and (Figs. 294-298) mounted on same slide, seven additional para- lectotypes taken on different hosts in same lo- Ripersiella simplex Hambleton, 1946a: 73. cality mounted on four slides. USNM. Rhizoecus cacticans (Hambleton): McKenzie, 1967: 381 (misidentification in part). Host Plants.—Brassaia sp., Calendula sp., Rhizoecus simplex: Hambleton, 1973: 69. Carissu grandiflora, Coccoloha iivifera, Cordy^ line sp., Cryptanthus sp., Dieffenbachia sp., Adult Female.—Oval elongate. Length, 0.87- Dizygotheca ele g antis sima, Erigeroîi bonarien- 1.39 mm; width, 0.38-0.66 mm. Antennae six- sis, Eriobotrya japónica, Euphorbia niilii, Gar- segmented, short, moderately stout, average dénia jasminoides, Hamatocactus setispinus, length of segments : I, 25 ; II, 14 ; III, 20 ; IV, Hederá helix, Hoya carnosa cv. exotica, Ixora 12; V, 13; VI, 31; apical segment less than coccínea, Neoregelia sp., Nephrolepsis exaltata, twice as long as wide, with three fairly stout, Oxalis martiayia, Panicnm sp., Peperomia sp., 48 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE Pilea microphylla, Plantag o sp., Saintpaulia whereas in cacticans there are 32-40 larger, sp., Strelitzia reginae, Zygocactus sp., Z. trun- subtriangulate to quadrate cells in the outer catiis. part of the ring. Distribution.—California, Florida, New York, Washington, Brazil. California: Berkeley, Alameda Co., 4-XI-58, Rhizoecus solani (Hambleton) H. L. McKenzie and A. E. Pritchard. (Figs. 299-303) Florida: Apopka, 21-1-71, F. L. Ware. Bradenton, 1-1-71, J. R. McFarlin. Eau Gallie, Ripersiella solani Hambleton, 1946a: 75. 8-II-71, H. C. Levan. Englewood, 16-1-73, Rhizoecus solani: Ferris, 1953: 458. C. J. Bickner. Fairvilla, 2-II-71, F. L. Ware. Adult Female.—Elongate oval. Length, 1.35- Fort Lauderdale, 14-11-72, J. R. Halstead. Gainesville, 19-VI-70, John Perry. Gotha, 1.53 mm; width, 0.66-0.71 mm. Antennae six- segmented, rather widely spaced, average l-VIII-61, R. J. Griffith. Homestead, 23-IV- 71, J. H. Knowles. Key West, 1-1-73, E. J. length of segments: I, 32; II, 18; III, 31; IV, Hambleton. Leesburg, 13-V-71, A. L. Bentley. 17; V, 17; VI, 45; apical segment not quite Lockhart, 29-1-71, F. L. Ware. Orlando, twice as long as wide, with three moderately 10-11-71, W. W. Smith and E. R. Fatic; 21- stout, bluntly tapered, falcate sensory setae V-71, F. L. Ware. Osprey, 5-II-71, J. R. and one elongate, spinelike sensory seta; seg- McFarlin. Oxford, 22-1V-71, A. L. Bentley. ment V with shorter, narrow sensory seta. Palma Sola, 7-XII-67, J. R. McFarlin. Pom- Interantennal space equal to combined length pano Beach, lO-VI-63, M. L. Bank. Sebastian of segments V and VI. Eyes fairly prominent, Inlet, 12-11-71, H. C. Levan. Snead Island, weakly pigmented. Rostrum elongate, meas- 2_IV-71 ; Tallavast, 30-III-71, J. R. McFarlin. uring 73fi long, 41(1 wide ; rostral loop reaching Upper Key Largo, 3-II-68, R. E. Woodruff. almost to insertion of second pair of legs. Ce- W. Melbourne, 9, 12-11-71, H. C. Levan. Win- phalic plate weakly indicated, appearing to be ter Garden, 11-1-65, 26-11-65, R. J. Griffith; longer than wide. Dorsal ostioles inconspicuous. 19-11-68, F. L. Ware. Zellwood, 28-IX-73, P. Legs rather stout, average length of seg- Gibson and W. Pierce. ments of hind pair: Trochanter, 37; femur, New York: Syracuse, XI-72, 27-11-73, C. A. 96; tibia, 81; tarsus, 59; claw, 20; claw digi- tules elongate, weakly dilated apically, extend- Cooke. ing to or slightly beyond rather stout claws. Washington: Bellingham, 4-IX-65, C. A. With one medium-sized conical circulus, 25|i Leckie; lO-VI-68, B. J. Landis. across its base, its orifice about 10(i in diameter, Brazil : The type locality. finely faveolate. Anal lobes weakly protruding, Discussioii.—Because a holotype was not in- unsclerotized, each lobe with three elongate dicated in the original description of simplex, setae, two of them longer and stouter than the I hereby designate the first specimen on the third, longest about 74ji long. Anal ring ap- left in the bottom row in a field of five individ- pearing slightly wider than long, measuring uals on slide No. 1 as the lectotype. 49(1 in diameter, its setae about as long and The study of a long series of specimens from stout as lobe setae; outer part of ring with Florida has proved invaluable in confirming 12-15 elongate, sinuate cells practically touch- the validity of simplex. Its present distribu- ing end to end, each with spicule near its cen- tion, including the new record for New York, ter; cells of inner part of ring 10-12, larger represents a considerable extension in the and more irregular in shape, lying adjacent to known range of the species. clouded area of subcircular cells. Bitubular R, simplex is about one-half the size of cacti- cerores present, about 13^i long, rather stout, cans, and both species differ in the size and 45-48 present mostly dorsally and submargin- shape of their antennae and rostra. The cells ally, 21-22 on head and thorax, 24-26 on abdo- of the outer part of the anal ring in simplex men, walls of ducts parallel, two ducts lying are mostly oval elongate and number 17-23, side by side, or with their bases diverging, pro- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 49 jected part slightly above derm. Multilocular slender, dilated at tips, reaching beyond mod- disk pores 45-50, occurring across venter of erately stout, curved claws. posterior three abdominal segments and occa- With one elongate, conical circulus, about as sionally 1 over same area dorsally. Tubular wide at base as long. Anal lobes undeveloped, ducts small, widely distributed over derm. Tri- each lobe area with small sclerotized patch, locular pores moderately abundant and uni- three elongate setae, longest about 75|i long, formly distributed. Body setae variable in small concentration of pores, and several body length, most short, not prominent. setae. Anal ring about 5l!i wide, its structure Holotype.—Guatemala City, Guatemala, 12- distinct, its anterior pair of setae stoutest, re- V-45, E. J. Hambleton. Paratypes.—Three maining pairs about as stout but shorter than taken with holotype. USNM. longest lobe seta; outer part of anal ring with Host Plant.—Solanum nigrum. 17-20 small, elongate, oval cells, most with spic- Distribution.—Guatemala, Mexico. ules; inner part of ring with 14 much larger, more irregular cells lying adjacent to darkened Guatemala : The type locality. area of semicircular cells. Tritubular cerores Mexico: 15 mi S. Llera, Tamaulipas, 24-11- of medium size, about l\i long, walls of indi- 72, D. R. Miller and F. D. Parker. vidual ducts almost parallel, projecting half Discussion.—As indicated under the discus- their length above derm, about 30 dorsally along sion of mexicanus, that species and solani are median line and submargin, on venter, about closely related. See page 37 and the key to spe- 15 occurring on submargin. Multilocular disk cies for their major differences. pores more abundant ventrally in area of vulva, as many as 28 on segment VIII, 26 across base of segment VII, 18 on segment IX, several on Rhizoecus sonomae McKenzie VI and areas opposite base of rostrum, else- (Figs. 304-309) where scattered; on dorsum, 3 present on VI, 3 on VII, 9 on VIII, 6 on IX. Tubular ducts Rhizoecus sonomae McKenzie, 1960b: 751; McKenzie, small, evenly distributed on both surfaces, few 1961: 47; McKenzie, 1964: 269; McKenzie, 1967: 402. on head. Trilocular pores fairly uniformly dis- Rhizoecus hrowni McKenzie, 1961: 43; McKenzie, 1967: tributed, more sparse ventrally, especially in 376. NEW SYNONYMY. thoracic area. Body setae varying somewhat in length but most short and rather inconspicuous. Adult Female.—Elongate oval. Length, 0.91- Holotype.—Two mi W. of Petrified Forest, 1.84 mm; width, 0.46-0.91 mm. Antennae six- Sonoma County, Calif., 23-11-59, W. R. Bauer segmented, average length of segments: I, 33; and J. S. Buckett ; UCD. Paratype.—One from II, 20 ; III, 36 ; IV, 21 ; V, 21 ; VI, 51 ; apical seg- same locality, 25-X-59, J. S. Buckett; CDA. ment about tv^ice as long as wide, with three Holotype of hrowni, 5 mi NW. Spanish Flat, long, narrow, falcate sensory setae and one dis- Napa County, 1-III-61, S. W. Brown; UCD. tal, spinelike sensory seta; segment V with one Host Plant.—Unknown, taken in leaf mold shorter, narrow, weakly clávate sensory seta. and trash from soil under Juniperus sp. and Interantennal space equal to about combined Quercus wislizenii. length of segments III and IV. Eyes weakly Distribution.—California : Type locality. Gra- protuberant, lightly pigmented. Rostrum mod- ton, 20-11-60, C. L. Judson. Lucerne, l-VI-61, erately stout, 71|i long, 53fi wide; rostral loop R. 0. Schuster. Wooden Valley, 29-IV-62, extending to about insertion of second pair of S. F. Bailey. legs. Cephalic plate usually slightly wider than Discussion.—In attempting to key the species long, well sclerotized, bordered anteriorly by sonomae and hrowni on the basis of their origi- three small body setae. Dorsal ostioles well nal descriptions, certain characters appeared developed, inconspicuous. to differentiate the two forms. However, when Legs of average size, length of segments of types and other available material were stud- hind pair: Trochanter, 45; femur, 94; tibia, ied, it became apparent that they could not be 86; tarsus, 63; claw, 22; claw digitules long, separated. In all specimens the sclerotization 50 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

of the anal lobes is very weak, the claw digi- slightly tapered, about ll|i long, present on tales are weakly dilated, and the anal ring both surfaces, 3 on head, 8 on thorax, 12-14 structure, antennae, rostra, and other charac- on abdomen. Multilocular disk pores with six ters show no marked differences. Because of to seven loculi, present dorsally and ventrally, these similarities, browni is here considered a few but uniformly scattered, absent from head. junior synonym of sonomae. Tubular ducts absent. Trilocular pores sparse, widely distributed. Body setae short, sparse, somewhat longer on head and on dorsum of Rhizoecus spinipes (Hambleton) posterior abdominal segments. (Figs. 310-315) Holotype.—Howard County, Ark., 8-VI-36, W.F. Turner; USNM. MorrisoneUa spinipes Hambleton, 1946a: 36. Host Plants.— Andropogon rhizomatiis, Pani- Rhizoecus spinipes: Hambleton, 1973: 70. ctim sp., undetermined Gramineae, in soil under peach tree. Adult Female.—Broadly elliptical. Length, 1.32-1.64 mm; width, 0.91-1.19 mm. Antennae Distribution.—Arkansas, Florida, Georgia, six-segmented, short, stout, closely placed near Mexico. apex of head, segments in following lengths: Arkansas : Type locality. I, 23; II, 18; III, 15; IV, 12; V, 11; VI, 34; Florida: Gainesville, lO-X-67, K. R. Lang- apical segment stout, less than twice as long as don. wide, with three stout, strongly clávate, api- Georgia: Chattooga County, 4-XII-73, R. J. cally acute, falcate sensory setae and one spine- Beshear. like sensory seta near apex; segment V with Mexico: Jalapa (Rio Cedeno), Vera Cruz, one narrow, elongate, curved sensory seta. 16-VII-67, D. R. Miller and J. Villanueva B. Interantennal space about equal to width of Discussion.—This unique species is distin- segment I. Eyes small, rather prominent, guished by its short, stout, acute anal-ring weakly pigmented. Rostrum almost twice as setae, presence of four elongate setae on each long as wide, 57|i long, 31ji wide; rostral loop anal lobe, shape of its falcate sensory setae, extending slightly more than halfway to inser- and multilocular disk pores with six or seven tion of second pair of legs. Cephalic plate spade- loculi. shaped, rather large, about 44^ long, with as R. spinipes was omitted by Ferris (1953) many as 10 small body setae associated with it. and by McKenzie (1960b, 1967). Dorsal ostioles poorly developed, anterior pair apparently absent, posterior pair inconspicu- ous, weakly pigmented. Rhizoecus stangei McKenzie Legs small, robust, rather spinose, length of (Figs. 316-320) segments of hind pair : Trochanter, 30 ; femur, 58; tibia, 47; tarsus, 35; claw, 21; claw digi- Rhizoecus stangei McKenzie, 1962: 676. tules not reaching half the length of long, Adult Female.—Oval elongate. Length of slender claws. type, 1.29 mm; width, 0.66 mm. Antennae six- Circulus absent. Anal lobes undeveloped, segmented, stout, length of segments: I, 58; each lobe area with elongate sclerotized patch, II, 33 ; III, 36 ; IV, 27 ; V, 27 ; VI, 64 ; apical seg- with four slender, elongate setae, longest about ment less than twice as long as wide, with three 48j.i long. Anal ring about 45|i in diameter, its moderately stout, falcate sensory setae tapering setae shorter than its diameter, stout, apically toward their extremities and one spinelike sen- acute, about 35¡i long; outer part of anal ring sory seta near apex ; segment V with one elon- with 12 rather large, elongate cells isolated gate, narrow, slightly clávate sensory seta. from one another and with tiny spicules ; inner Interantennal space about equal to combined part of ring with 8-10 similar cells lying adja- length of segments V and VI. Eyes prominent, cent to undifferentiated shaded area. Tritubu- rather globose, weakly pigmented. Rostrum lar cerores large, their individual ducts stout. comparatively stout, length 83|i, width 6A\x; A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 51

rostral loop reaching insertion of second pair Rhizoecus siibcyperalis^ new species of legs. Cephalic plate approximately S3[i long and 46j.i wide, with two body setae on anterior (Figs. 321-323) border. Dorsal ostioles well developed, heavily Adult Female.—Broadly ovate elongate, sclerotized. stout. Length, 1.60-1.81 mm; width, 0.81-1.00 Legs rather large, length of segments of mm. Antennae six-segmented, short, rather hind pair : Trochanter, 59 ; femur, 123 ; tibia, stout, widely spaced, average length of seg- 101; tarsus, 79; claw, 34; claw digitules very ments : I, 41 ; II, 19 ; III, 35 ; IV, 18 ; V, 18 ; VI, short, setose, hardly reaching half the length 41 ; apical segment not twice as long as wide, of long, stout, curved claws. tapering, with three comparatively narrow, Circulus absent. Anal lobes slightly protrud- elongate, falcate sensory setae and one elon- ing, each lobe with small sclerotized patch, gate, spinelike sensory seta; segment V with three elongate setae, longest about 51¡.i long, one shorter, smaller sensory seta. Interanten- remaining two shorter, smaller, and with six nal space equal to combined length of segments to eight short body setae. Anal ring about 53u I-III. Eyes small, often inconspicuous, weakly wide, its setae of variable lengths and averag- pigmented. Rostrum elongate, 82!.i long, 55fi ing about 60|.i long, all longer and about as wide; rostral loop reaching halfway to inser- stout as stoutest lobe seta; outer part of anal tion of second pair of legs. Cephalic plate ab- ring with 10 rather large, elongate, oval cells, sent. Dorsal ostioles conspicuous, their rims some with spicules and most isolated from each strongly sclerotized. other ; inner part of ring with about same num- Legs of average size, length of segments of ber of cells of similar size; clouded area not hind pair: Trochanter, 46; femur, 105; tibia, clearly defined. Tritubular cerores fairly nu- 75; tarsus, 68; claw, 24; claw digitules short, merous, 2 sizes present, larger about ll^i wide, acute, about reaching middle of elongate, their ducts 8^-9|i long, stout, about 18 occur- curved claws. ring dorsally, 5 along median line from head to Circulus absent. Anal lobes with some pro- abdominal segment VI and 12 or 13 submar- trusion, unsclerotized, each lobe with three or ginally from thorax to abdominal segment four elongate setae, longest approximately VIII; smaller or medium ceroris 5|.i-7^ wide 78^1 long, and body pores concentrated around and of similar shape, 44 or more occurring ven- them. Anal ring about 51j.i wide, usually resting trally in rows across segments, of these, 23-28 at angle or vertically in mounted specimens, its are on segments VII and VIII. Multilocular setae each about 57|.i long, as stout as but disk pores abundant, over 100 observed on ven- shorter than longest lobe seta; outer part of ter of abdominal segments III-IX, more nu- anal ring with 10-12 elongate, oval, or sinuate merous in area of vulva and across posterior cells with spicules; inner part of ring with borders of segments V-VIII, few occurring dor- larger cells of similar shape but more irregular sally or near lateral margins, absent from head in outline. Tritubular cerores large, of one size, and thorax. Tubular ducts absent. Trilocular 18 occurring dorsally, 3 on head, 7 on thorax, pores rather evenly distributed over both sur- 8 on abdomen. Multilocular disk pores 4-10, faces. Body setae most small, inconspicuous. confined to venter in area of vulva. Tubular Holotype.—One mi N. Tepic, Nayarit, Mexico, ducts small, few, located on venter of abdomi- 21-VIII-59, L. A. Stange and A. S. Menke. Par- nal segments VIII and IX. Trilocular pores atype.—One taken with holotype. UCD. evenly distributed over both surfaces. Circular Host Plant.—Unknown, collected from soil. pores minute, sparse, occurring across dorsum Distribution.—Type locality. of midabdominal segments. Body setae rather Discussion.—ß. stangei seems to bear some short, inconspicuous. relation to americanus and mayanus. The Holotype.—Portal, SW. Research Station, major difference between them is that stangei Cochise County, Ariz., 3-VIII-66, D. R. Miller; has only two sizes of tritubular cerores and UCD. Paratypes.—Five taken with holotype, possesses many more larger cerores on its ven- on four slides; six additional on six slides as tral surface. It is also similar to neostangei. follows: Arizona: Bisbee, Cochise County, 2- 52 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE VIII-66, D. R. Miller; Madera Canyon Station, conspicuous, their rims weakly pigmented, sur- Santa Cruz County, 31-VII-66, D. R. Miller. rounded by numerous pores and body setae. Idaho: Craters of the Moon National Monu- Legs of average size, length of segments of ment, Butte County, 4-VII-66, D. S. Horning; hind pair : Trochanter, 41 ; femur, 96 ; tibia, 8 mi NE. Minkcreek, Franklin County, 3-VIII- 74; tarsus, 55; claw, 25; claw digitules short, 67, D. R. Miller and D. S. Horning. Nine UCD, setose, hardly reaching half the length of long, two USNM. rather stout, weakly curved claws. Hast Plants.—Lithospermum ruderale, Ur- Circulus absent. Anal lobes weakly devel- tica hjallii, under rocks, in soil. oped, each lobe with light, irregular sclerotiza- Distribution.—Arizona, Idaho. tion, three slender, elongate setae, longest about Discussion.—Twelve specimens in the UCD 60[i, (others broken off in type), and several collection mislabeled cyperalis, eluminatus, and small body setae. Anal ring about 40fi in diam- pritchardi form an integral group of individ- eter, its setae about same thickness but shorter uals with like characters and are here consid- than longest lobe seta, measuring about 52Lí ered new and described under the name subcyp- long; outer part of ring containing 12 narrow, eralis. The true cyperalis from Central Amer- elongate cells with spicules and often touching ica does not appear to be present in California, end to end; cells of inner part of ring with Arizona, or Idaho ; eluminatus has been synon- about same number, somewhat larger but not ymized with pritchardi. The latter species has well defined and lying next to darkened area 30-35 cerores of 2 sizes that occur both dor- with large, oval cell structure. Tritubular sally and ventrally, whereas subcyperalis has cerores present in two sizes, larger about 8\x only 18 cerores of 1 size and they are restricted wide occurring dorsally, five along median line to the dorsum. from head to abdominal segment VI, eight sub- marginally ; cerores about 5\x wide, present ven- trally, one pair each on abdominal segments V Rhizoecus theobromae (Hambleton), and VII near midventral area. Multilocular new combination disk pores most numerous on venter of poste- (Figs. 324-328) rior abdominal segments, 35-40 present, else- where, except on head, few pores widely scat- Morrisonella theobromae Hambleton, 1946a: 39. tered on both surfaces. Tubular ducts scarce, occurring on venter, opposite anal ring on ante- Adult Female.—Elongate oval, slightly con- rior anal-lobe area. Trilocular pores uniformly stricted behind thorax. Length, 1.39-1.70 mm; distributed, most numerous dorsally. Medio- width, 0.66-0.75 mm. Antennae six-segmented, ventral pores in two clusters across midven- comparatively short, widely spaced near apex tral surface of abdominal segments VI and VII, of head, segments measuring as follows : I, 33 ; 12-31 pores in each cluster. Body setae mostly II, 18 ; III, 19 ; IV, 20 ; V, 17 ; VI, 42 ; apical seg- short, some longer setae on head, prothorax, ment less than twice as long as wide, sharply and posterior abdominal segments. tapered, with three stout, apically obtuse, fal- cate sensory setae and one elongate, spinelike Holotype.—Pichilingue, Ecuador, l-X-44, sensory seta; segment V with one elongate, E. J. Hambleton. Paratypes.—Three (one stout, clávate sensory seta. Interantennal mounted with holotype on same slide) and space equal to combined length of segments V three preadult females. USNM. and VI. Eyes rather prominent, hemispherical, Host Plant.—Theobroma cacao. pigmented. Rostrum of medium size, length Distribution.—Known only from type locality. about 67ji long, 48ji wide; rostral loop extend- Discussion.—This species, like the West In- ing slightly beyond insertion of second pair of dian globoculus, has the peculiar medioventral legs. Cephalic plate irregularly shaped, approxi- pores and is found on the same host. Its stouter mately 28|i long by 48fi wide, with several vac- falcate setae, less globose eyes, and more abun- uoles and usually with body seta at each latero- dant multilocular disk pores distinguish it from cephalic margin. Dorsal ostioles moderately globoculus. Tubular ducts are present in theo- A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 53

hromae, though the original description of the Tubular ducts about 5|.i long, distributed on species stated that they were absent. both surfaces in small numbers over entire derm. Trilocular pores fairly abundant, more numerous on dorsum. Body setae varying in Rhizoecus totonicapanus (Hambleton) length, some longer setae nearly 40(.i long on (Figs. 329-333) head and abdomen. Holotype.—Along roadside between Quetzal- RiperHiella totonicapana Hambleton, 1946a: 76. tenango and Totonicapan, Guatemala, 20-V-45, Rhizoecus totonicapanus: Ferris, 1953: 460. E. J. Hambleton. Paratype.—One taken with Adult Female.—Oval elongate. Length, 1.35- holotype. USNM. 1.43 mm; width, 0.67-0.71 mm. Antennae six- Host Plant.—Alchemilla orbictdata. segmented, moderately widely spaced, segments Distribution.—Known only from type locality. measuring as follows : I, 25 ; II, 17 ; III, 19 ; IV, Discussion.—In addition to the differences 14; V, 13; VI, 39; apical segment less than pointed out between this species and gracilis twice as long as wide, with three elongate, mod- (p. 29), the latter species is larger and the cells erately stout, weakly clávate, falcate sensory in the outer part of its anal ring are much nar- setae and one elongate, spinelike sensory seta rower than those in totonicapanus. located near apex; segment V with one short, clávate sensory seta. Interantennal space about Rhizoecus tropicalis^ new species equal to combined length of segments I-III. Eyes rather small with little pigmentation. (Figs. 334-339) Rostrum of average size, 62\x long, 41[i wide; rostral loop extending more than halfway to Rhizoecus neomexicanus McKenzie, 1962: 674 (mis- identification in part). insertion of second pair of legs. Cephalic plate subquadrate, about 30¡i long, 40^i wide, with Adult Female.—Oval elongate. Length, 1.29 two vacuoles and several body setae on its bor- mm ; width, 0.64 mm. Antennae six-segmented, ders. Dorsal ostioles almost unrecognizable, moderately short, stout, strongly geniculate, without pigmentation. length of segments as follows: I, 21; II, 17; Legs short, length of segments of hind pair : III, 19; IV, 15; V, 14; VI, 33; apical segment Trochanter, 33; femur, 68; tibia, 55; tarsus, less than twice as long as wide, with three 45; claw, 18; claw digitules long, slender, and slender, bluntly tapered, weakly clávate, fal- appearing setose, extending to tip of relatively cate sensory setae and one narrow, more acute stout claws. sensory seta ; segment V with one shorter, clá- With one conical circulus, about 16^1 wide vate sensory seta. Interantennal space about across its base. Anal lobes not protruding, each equal to combined length of segments V and lobe area with small, irregular sclerotized VI. Eyes absent. Rostrum of medium size, 51u patch, three slender setae, longest about 60|i long, 45|i wide; rostral loop almost reaching long, several body setae and pores. Anal ring insertion of second pair of legs. Cephalic plate about 50[.i wide, its setae slender, anterior pair longer than wide, with four or five body setae stoutest and about 62[i long; outer part of anal along its periphery. Dorsal ostioles weakly scle- ring with 12-14 elongate, oval cells with spic- rotized, inconspicuous. ules; inner part of ring with 10-12 larger, Legs short, stout, length of segments of hind more irregular cells lying next to shaded area pair: Trochanter, 33; femur, 58; tibia, 55; of rounded cell structure. Derm with bitubular tarsus, 35; claw, 15; claw digitules elongate, cerores small, elongate, with ducts about 9[.i their tips dilated, extending beyond rather long, weakly tapered, divaricating, present dor- stout, weakly curved claws. sally and ventrally, 19-22 on head and thorax, With one small, roundly truncate circulus, 23-27 on abdomen. Multilocular disk pores about 13ji wide at its base, its apex widely 31-43, present largely on venter of posterior faveolate. Anal lobes undeveloped, each lobe abdominal segments, few occurring on dorsum. area with small, irregularly elongate, sclero- 54 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE tized patch between one long and two shorter Holotype.—San Rafael, Departamento San setae (mostly broken in types). Anal ring about Marcos, Guatemala, 28-XII-63, J. G. Rodriguez. 45ii in diameter, its setae approximately 55u Paratypes.—Two taken in Guatemala in 1964 long, much stouter than lobe setae; outer part by Jesus Escobar, USNM; two. Alazán, State of anal ring with 19-21 elongate to ovate, of Vera Cruz, Mexico, 15-VIII-59, A. S. Menke loosely arranged, spiculate cells; inner part of and L. A. Stage, UCD. ring with 10-12 larger, more elongate, and Host Plant.— Coffea arabica. somewhat stouter cells, dark area poorly de- Distribution.—Guatemala, Mexico. fined. Tritubular cerores small, their ducts about 6!.i-7!.i long, present dorsally and ven- Guatemala : Type locality. trally, 17-18 on head and thorax, 21-24 on ab- Mexico: Alazán, Vera Cruz, 15-VIII-59, domen, as many as 5 on abdominal segment A. S. Menke and L. A. Stange. This collecting VIII. Multilocular disk pores absent. Tubular record includes three specimens mistaken for ducts smaller in diameter than trilocular pores, neomexicamis. (See discussion under R. neo- widely distributed but most common ventrally, mexicanus, p. 40.) five or six present on most abdominal seg- ments. Trilocular pores rather sparsely distrib- Discussion.—The structure of the anal ring, uted, very few in some areas of abdominal shape of the rostrum, and faveolate circulus segments. Body setae short, rather sparse. are distinguishing characters of this species.

SUMMARY

In this bulletin 52 species of Rhizoecus bleton and Ripersiella Tinsley are assigned to Kiinckel d'Herculais are recognized, of which Rhizoecus. A key is presented to adult females 12 are described as new ; 33 of the 36 previously of New World species. All taxa are described described New World species are retained as or redescribed and pertinent structures are valid, and 3 names are synonymized; 7 species illustrated. Host plant and distribution records originally described under Morrisonella Ham- are listed for each species.

LITERATURE CITED

BORCHSENIUS, N. iS. DE LOTTO, G. 1949. FAUNA OF THE USSR. HOMOPTERA, PSEUDO- 1957. THE PSEUDOCOCCIDAE DESCRIBED BY H. C. cocciDAE. Akad. Nauk Zool. Inst. (n.s. 38) JAMES FROM E. AFRICA. Brit. Mus. (Nat. 7, 383 pp. Hist.) Ent. Bui. 5(5): 185-232. BRAIN, C. K. 1915. THE COCCIDAE OF SOUTH AFRICA. Roy. SoC. 1964. OBSERVATIONS ON AFRICAN MEALYBUGS. Brit. Africa Trans. 5 (2) : 65-194. Mus. (Nat. Hist.) Ent. Bui. 14 (8) : 343- BUEKER, E. D. 397. 1931. MEALY-BUGS (COCCIDAE) OF NESTS OF ANTS. FERNALD, M. E. Colo. Univ. Studies 18 (3) : 151-162. 1903. CATALOGUE OF THE COCCIDAE OF THE WORLD. COCKERELL, T. D. A. Mass. Agr. Expt. Sta. Bui. 88, 360 pp. 1894. A MARITIME SPECIES OF COCCIDAE. Insect Life 7(1): 42-44. FERRIS, G. F. 1953. ATLAS OF THE SCALE INSECTS OF NORTH 1899. TABLES FOR THE DETERMINATION OF THE AMERICA, VI, THE PSEUDOCOCCIDAE (PT. Il). GENERA OF COCCIDAE. Canad. Ent. 31 (10) : Pp. 279-506. Stanford Univ. Press, Stan- 273-279. ford, Calif. GONZALEZ, R. H., and CHARLIN, R. 1901. NEW AND LITTLE-KNOWN COCCIDAE I, RIPER- 1968. NOTA PRELIMINAR SOBRE LOS INSECTOS COC- SIELLA AND CEROPUTO. Biol. Soc. Wash. COIDEOS DE CHILE. Rev. Chilena de Ent. Proc. 14: 165-167. 6: 109-113. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 55

GREEN, E. E. 1926. OBSERVATIONS ON BRITISH COCCIDAE X. ENT. 1962. THIRD TAXONOMIC STUDY OF CALIFORNIA Monthly Mag. 62: 172-183. MEALYBUGS, INCLUDING ADDITIONAL SPECIES FROM NORTH AND SOUTH AMERICA (HOMOP- 1933. NOTES ON SOME COCCIDAE FROM SURINAM, TERA: COCCOIDEA: PSEUDOCOCCIDAE). Hilgar- DUTCH GUIANA WITH DESCRIPTIONS OF NEW dia 32 (14) : 637-688. SPECIES. Stylops 2 (3) : 49-58. HAMBLETON, E. J. 1964. FOURTH TAXONOMIC STUDY OF CALIFORNIA 1946a. STUDIES OF HYPOGEIC MEALYBUGS. Rev. de MEALYBUGS, WITH ADDITIONAL SPECIES FROM Ent. (Rio de Janeiro) 17 (1-2) : 1-77. NORTH AMERICA, SOUTH AMERICA AND JAPAN (HOMOPTERA: COCCOIDEA: PSEUDOCOCCIDAE). 1946b. A NEW NAME FOR A MEALYBUG. Biol. Soc. Hilgardia 35 (10) : 211-272. Wash. Proc. 59: 177.

1973. FLORIDA MEALYBUGS OF THE GENUS RHIZOE- 1967. MEALYBUGS OF CALIFORNIA WITH , CUS WITH DESCRIPTION OF A NEW SPECIES. BIOLOGY AND CONTROL OF NORTH AMERICAN Ent. Soc. Wash. Proc. 75 (1) : 62-71. SPECIES. 525 pp. Univ. Calif. Press, Berkeley. 1974. THREE NEW SPECIES OF RHIZOECUS FROM NEW MILLER, D. R., and MCKENZIE, H. L. ZEALAND WITH NOTES AND REDESCRIPTION 1971. SIXTH TAXONOMIC STUDY OF NORTH AMERI- OF OTHERS. New Zeal. Jour. Zool. 1 (2) : CAN MEALYBUGS, WITH ADDITIONAL SPECIES 147-158. FROM SOUTH AMERICA (HOMOPTERA: COCCOI- KUNCKEL D'HERCULAIS, J. DEA: PSEUDOCOCCIDAE). Hilgardia 40 (17): 1878. HISTOIRE DE LA COCHENILLE VIVANT SUR LES 565-602. RACINES DES PALMIERS DE LA SECTION DES MORRISON, H., and MORRISON, E. SEAFORTHIA. Soc. Ent. de France Ann. 1922. A REDESCRIPTION OF THE TYPE SPECIES OF 8 (5) : 161-164. COCCIDAE BASED ON SPECIES DESCRIBED BY KuwAi, S., and TAKAGI, K. MASKELL. U.S. Nati. Mus. Proc. 60 (12) : 1971. DESCRIPTIONS OF THREE ECONOMICALLY IM- 1-120. PORTANT SPECIES OF ROOT-FEEDING MEALY- BUGS IN JAPAN (HOMOPTERA: PSEUDOCOCCI- NEWSTEAD, R. DAE). Appl. Ent. and Zool. 6 (4) : 175-182. 1895. OBSERVATIONS ON COCCIDAE. Ent. Monthly KUWANA, I. Mag. 31: 213-214. 1923. DESCRIPTIONS AND BIOLOGY OF NEW OR LITTLE- RAU, G. T. KNOWN cocciDS FROM JAPAN. Japan Dept. 1937. EXTEMPORANEOUS REPORT ON SCALE INSECTS Agr. and Com. Imp. Plant Quar. Sta. Bui. IN A GREENHOUSE AT BRONX PARK, N.Y. 3: 1-67. N.Y. Ent. Soc. Jour. 45 (2) : 267. MCKENZIE, H. L. WILLIAMS, D. J. 1960a. A NEW SUBTERRANEAN RHIZOECUS MEALY- 1961. CHANGES IN NOMENCLATURE AFFECTING BUG FROM ARIZONA. Pan-Pacific Ent. 36 SOME COCCOIDEA (HOMOPTERA). Ent. (3) : 139-141. Monthly Mag. 97: 92-93.

1960b. TAXONOMIC STUDY OF CALIFORNIA MEALY- BUGS, WITH DESCRIPTIONS OF NEW SPECIES 1962. THE BRITISH PSEUDOCOCCIDAE (HOMOPTERA: (HOMOPTERA: COCCOIDEA : PSEUDOCOCCIDAE) . COCCOIDEA). Brit. Mus. (Nat. Hist.) Ent. Hilgardia 29 (15) : 681-770. Bui. 12 (1) : 1-79. — and DE BOER, J. A. 1961. SECOND TAXONOMIC STUDY OF CALIFORNIA 1973. THE TAXONOMY OF SOME NEW ZEALAND MEALYBUGS, WITH DESCRIPTIONS OF NEW PSEUDOCOCCIDAE (HOMOPTERA: COCCOIDEA). SPECIES (HOMOPTERA: COCCOIDEA: PSEUDO- Roy. Ent. Soc. London, Trans. 125 (2) : COCCIDAE). Hilgardia 31 (2): 15-52. 227-252.

INDEX TO HOST PLANTS

Page Page Acacia sp. 24 Aechmea orlandiana L. B. Smith 26 Achillea ageratifolia (Sibth. and Sm.) Boiss. 46 Agapanthus sp. 24 Achillea millefolium L. 28 Agropyron sp. 38 Adiantum sp. 22,46 Agrostis sp. 30 Aechmea chantinii (Carrière) Baker 26 Ajuga sp. 24 56 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

Page Page AlcheTnilla orbiculata R. and P. 14, 39, 53 Coffea arabica L 11,13,18, 20, 24-25, 27, 31, 42, 54 Andropogon rhizomatus Swallen 50 Coffea liberica Bull ex Hiern 24 Andropogon sp. 23 Collinia sp. 11 Andropogon virginicus L. 11,26 Conocarpus erecta L. 11,26 Anernone hupehensis var. japónica Cordyline sp. 47 (Thunb.) Bowles and Stearn 24 Cortaderia selloana (Schult.) A. and G. 26 AntheTnis sp. 26 Cryptanthus sp. 47 Anthémis tinctoria L. 24 Cuphea sp. 26 Aquilegia sp. 24 Cupressus sp. 24 Aralia sp. 11,24,26 Cymbopogon citratus (DC.) Stapf 36 Araucaria excelsa R. Br. 11-12. 26 Cymbopogon nardus (L.) Rendle 36 Arctostaphylos sp. 46 Cynodon dactylon (L.) Pers. 24 Areca sp. 11 Cyperus esculentus L. 33 Arecastrum romanzoffianum (Cham.) Becc. 11,26 Cyperus rotundus L. 38 Artemisia californica Less. 28 Cyperus sp. 17 Artemisia douglasiana Bess. 19 Cyperus tenerrimus Liebm. 22 Artemisia sp. 28 Dactylis glomerata L. 23 Artemisia tridentata Nutt. 28 Delphinium sp. 24 Artemisia vulgaris L. 23 Dieffenbachia maculata (Lodd.) Asparagus sprengeri Regel 11 G. Don cv. Amoene 11 Aster sp. 23 Dieffenbachia picta Schott 11 Atriplex sp. 28 Dieffenbachia sp. 26,47 Axonopus sp. 15,41 Dipsacus sp. 19 Bahia dissecta (A. Gray) Britt. 33 Distichlis spicata (L.) Greene 26,38 Bamhusa sp. 26 Dizygotheca elegantissima (Veitch) Billbergia sp. 26 Vig. and Guill. 11,26,47 Brassaia sp. 47 Dracaena marginata Lam. 26 Brickellia sp. 28 Dudley a farinosa (Lindl.) Britt. and Rose 17 Bromus unioloides Kunth 17 Dudleya greenei Rose 38 Buxus sempervirens L. 24 Echeveria sp. 17 Buxus sp. 26 Echinochloa colonum (L.) Link 36 Caladium bicolor (Ait.) Vent. 18 Eleusine indica (L.) Gaertn. 17,36 Caladium sp. 11 Encelia sp. 22 Calendula sp. 47 Epiphyllum sp. 17 Calliandra haematocephala Hassk. 11 Eragrostis maypurensis (H.B.K.) Steud. 11 Calliandra sp. 11,26 Eremochloa ophiuroides (Munro) Hack. 26 Callistemon rigidus R. Br. 26 Eriger on bonariensis L. 47 Callistemon sp. 11 Eriobotrya japónica (Thunb.) Lindl. 47 Callistemon viminalis Cheel 11 Eriogonum fasciculatum Benth. 22 Carex sp. 24, 31, 38 Eriogonum heracleoides Nutt. 28, 38 Carissa grandiflora (E. Mey.) A. DC. 26,47 Eriogonum sp. 15,22 Carissa sp. 26 Eriophyllum confertiflorum (DC.) Gray 28 Ceanothus pumilis Greene 15 Ernodea angusta Small 11 Celtis occidentalis L. 31 Erodium moschatum (L.) L'Her. ex Ait. 24 Cenchrus echinatus L. 41 Escallonia rubra (Ruiz and Pav.) Pers. 24 Cestrum sp. 24 Escoboria tuberculosa (Engelm.) Chamaecyparis sp. 11 Britt. and Rose 33 Chamaedorea elegans Mart. 11 Eugenia sp. 26 Chlorophytum sp. 11 Euphorbia milii Desmoulins 11, 47 Chry salido carpus lutescens (Bory) Wendl. 11,26 Faucaria tigrina (Haw.) Schwant. 35 Chrysanthemum frutescens L. 24 Ficus nitida Thunb. 11 Chrysanthemum sp. 11, 24, 46 Fragaria chiloensis (L.) Duchesne 19 Chrysothamnus viscidiflorus (Hook.) Nutt. 28 Fragaria sp. 24, 31 Citrus mitis Blanco 26 Franseria chamissonis Less. 28 Citrus sinensis (L.) Osbeck 24, 39 Gardenia jasminoides Ellis 47 Citrus sp. 26,31 Gardenia sp. 11,24 Coccoloba uvifera (L.) L. 47 Gardenia thunbergia L. f. 26 Coccothrinax argentata Bailey 11 Geum coccineum Sibth. and Sm. 46 Codiaeum sp. 24 Gnaphalium sp. 11 A REVISION OF THE NEW WORLD MEALYBUGS OP THE GENUS RHIZOECUS 57

Page Page Gossypium hirsiituTn LI. 34 Melaleuca leucadendron (L.) L. 11 Grindelia caTnporum Greene 28 Mesembryanthemum sp. 17 Guettarda sp. ., 11 Musa paradisiaca var. sapientum Kuntze 36 Gutierrezia sp. 28 Neoregelia sp. i_„ 47 Hamatocactus setispinus Britt. and Rose _ . 47 Nephrolepsis exaltata (L.) Schott 47 Haplopappiis canus (Gray) Blake 28 Nephthytis sp. 11 Haplopappus spiniilosiis (Pursh) DC. _ 28 Nerium oleander L. 33 Hederá helix L. _ 47 Neriiim sp. 31 Helianthus sp. _ 19 Ophiopogon sp. 24 Hemigraphis reptans T. Anders, ex Hemsl. _„ 11 Opuntia sp. 17 Heuchera sp. _ 24 Oxalis martiana Zuce. __ 47 Hibiscus rosa-sinensis L. _ 11 Panicum maximum Jacq. 36 Hibiscus sp. _ 11,24 Panicum sp. J 26,47,50 Hippomane Tnancinella L. 30,32 Paspalum conjugatum Berg. __ 11,36 HolcuH lanatus L. __ 17 Paspahim fimbriatum H.B.K. 11 Howeia belmoreana (C. Moore and Paspalum vaginatum Swartz 41 F. V. Muell.) Becc. . . . 24 Pelargonium inquinans (L.) L'Herit et Ait. 31 Howeia forsteriana (C. Moore and Pelargonium sp. 24 F. V. Muell.) Becc. __ __ 24 Peperomia pellucida H.B.K. 11 Hoya carnosa (L.) R. Br. cv. Exotica . 47 Peperomia sp. 47 Hoya sj). _ 26 Petunia sp. 24 Ilex cornuta cv. Burfordii 26 Philodendron selloum C. Koch 26 Ilex opaca Ait. „_ _ 26 Philoxerus vermicularis (L.) Beauv. 26 Ilex rotunda Thunb. 26 Phieum pratense IJ. 33 Ilex vomitoria Ait. 26 Phoenix canariensis Chaub. 24,26 Iris sp. 24 Phoenix loureirii Kunth 11 Isoloma sp. _ 11 Phoenix roebelenii O'Brien 24 Iva frutescens var. oraria (Bart.) Physalis pubescens L. 11 Fern, and Grisc. 35 Picea abies (L.) Karst. 24 Ixora coccinea L. 47 Pilea microphylla (L.) Liebm. 48 Ixora sp. 26 Piper sp. 24 Jasminum sp. 24,26 Plantago sp. 48 Juncus balticus Willd. 38 Pluchea sp. 26 Juniperus sp. 49 Poa annua L. 44 Kalanchoe sp. 17 Poa sp. 33 Kalanchoe tomentosa Baker 17 Polygala crotalariodes Ham. ex DC. 46 Kalmia sp. 24 Polygonum sp. 19 Kentia sp. 11 Portulaca grandiflora Hook. 31 Kleinia sp. 17 Pothos sp. 11 Kohleria sp. ^ . 11 Prumis angustifolia Marsh. 26 Lachnanthes tinctoria (Walt.) Ell. 26 Prunus sp. 24,31 Lantana sp. „^ 11,46 Psoralea tenuiflora Pursh 33 Lespedeza cuneata (Dumont) G. Don 23 Pyracantha coccinea Roem. H Leucophyllum frutescens Johnst. . 26 Pyracantha sp. 26 Ligustrum ovalifolium Hassk. 24 Quercus coccinea Muenchh. 23 Ligustrum sp. 31 Quercus sp. 11,26 Liriope sp. 11 Quercus wislizenii A. DC. 16,49 Lithospermum ruderale Dougl. ex Lehm. 38,52 Rhaphiolepis sp. H Lobivia shaferi Britt. and Rose 17 Rhizophora Tnangle L. 35 Lobivia sp. 17 Rhododendron sp. 26 Lolium perenne L. 17 Rhus sp. 40 Lotus scoparius (Nutt. in T. and G.) Ottley 15 Ribes sp. 24 Lotus sp. 24 Rosa sp. 31 Lycopersicon csculentum Mill. 24 Rubus argutus Link 23 Malpighia coccigera L. 11 Rumex sp. 31 Mahis sp. 23 Saintpaulia ionantha Wendl. 11, 46 Mammillaria sp. 17,33,43 Saintpaulia sp. 42 Matthiola sp. 24 Scabiosa sp. 31 Medicago sativa L. 31 Sempervivum sp. 17,33 58 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

Page Page Sempervivum tectorum L. 17 Thynuis vulgaris L. 24 Senecio viikanioides Otto ex Walp. 17 Tridens flaims (L.) Hitchc. -_ 23 Setaria genimdata (Lam.) Beauv. 36 Tropaeolum majns L. 24 Sida sp. 26 Urtica lyallii Wats. 52 Sinningia speciosa (Lodd.) Hiern 46 Verhascum thapsus L. 23 Solanum nigrum L. 49 Veronica sp. 24 Solidago sp. 23 Viburnum suspensuTïi Lindl. „_ 26 Sorghum halepense (L.) Pers. 33 Vitis sp. 24 Spartina alterniflora Loisel. 35 Watsonia sp. 24,31 Spartina patens (Ait.) Muhl. 35 Stellaria media (L.) Cyrill. 31 Xanthosoma robustum Schott 36 Strelitzia reginae Ait. 48 Zantedeschia aethiopica (L.) Spreng. 24 Syringa vulgaris L. 24 Zygocactus sp. 48 Theobroma cacao L. 27,42,52 Zygocactiis trnncatus (Haw.) K. Schum. 48

INDEX TO MEALYBUGS

(Valid names are in roman type, synonyms in italic) Page Page africanus Brain 23 maritimus (Cockerell) 9,13,34-35 americanus (Hambleton) 2,7,10,12,36,51 may ana Hambleton 35 apizacos Hambleton 9,12 mayanus (Hambleton) 7,35-36,51 arabicus Hambleton 9, 13, 25 menkei McKenzie 5-7,36-51 associatus (Hambleton) 7,13-14,23 Tuexicana Hambleton 37 atlántica Hambleton 14 mexicanus (Hambleton) 5,37,49 atlanticus (Hambleton) 10,14-15 Morrisonella Hambleton 6, 10, 13, 21, 23, 27, 29, bicirculus McKenzie 8,15 32, 34-35, 39, 44, 50, 52 bituberculatus McKenzie 6,15-16, 37 moruliferus Green 23 boharti McKenzie 7,16 nakaharai Hambleton 8-9,18, 33, 38 browni McKenzie 49-50 nemoralis (Hambleton) 8,22,39,46 cacticans (Hambleton) 10,15,17-19,33,38,47-48 neomexicanus McKenzie 8,39-40,53-54 caladii Green 8,18, 40 neostangei Miller and McKenzie 8, 40, 51 californicus Ferris 6,19-20 nitidalis Hambleton 7,15,41 campestris Hambleton 4, 6, 20 ornata Hambleton 41-42 chilensis Hambleton 10,18, 20 ornatus (Hambleton) 5,8,41-42 Coccidella Hambleton 2, 6 ovatus Hambleton 9,42 cyperalis (Hambleton) 8, 21-22, 39, 45-46, 52 pauciporus Hambleton 5, 8, 43 decoratus Green 23 poensis (Hambleton) 7,44 disjunctus McKenzie 5, 7, 22-23 polyporus Hambleton 10,15, 44-45, 47 distinctus (Hambleton) 3-4,7,14,23 pritchardi McKenzie 2,5,8,22,45-46,52 eluminatus McKenzie 45-46, 52 relativus Hambleton 9, 45-47 epiphylli Ferris 17 Rhizoecus Künckel d'Herculais 1, 6 falcifer Künckel d'Herculais ^— 2-4, 6-7, 23, 25, 41, 43 Ripersia Signoret 2, 34 favacirculus Hambleton 9, 25 Ripersiella Tinsley __._ 2, 6, 14, 32, 34, 37, 47-48, 53-54 floridanus Hambleton 2, 9,18, 25, 27, 33 simplex (Hambleton) 2,9,12,47-48 geniculatus James 5,22-23 solani (Hambleton) 5,7,37,48-49 globoculus (Hambleton) 8,27,44,52 sonomae McKenzie 6, 49-50 gracilis McKenzie 7, 28-29 spinipes (Hambleton) 7,50 graminis (Hambleton) 8,29-30 spinosus McKenzie 30-32 insularis Hambleton 9, 30, 32 stangei McKenzie 8, 40, 50-51 kondonis Kuwana 2-3, 5-6, 20, 25, 30-32 subcyperalis Hambleton 5, 7, 22, 51-52 lata Hambleton 32 terrestris Newstead 23 latus (Hambleton) 8,30,32 lencosoma Cockerell 32 theobromae (Hambleton) 8,44,52 leucosomus (Cockerell) 2,10,17-18,20-21,32-33,38,47 totonicapana Hambleton 53 macgregori Hambleton 9, 34 totonicapanus (Hambleton) 7,29,53 marítima Cockerell 2,34 tropicalis Hambleton 9,27,40,53 A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 59

\?^p^^^

Syjm^.

f

A/ .:/ll:: ®/

/^ji/- 11 10

FIGURES 1-13.—Rhizoecus americanus: 1, Terminal segments of antenna; 2, cephalic plate; 3, rostrum; 4, tritu- bular ceroris, lateral; 5, tubular duct, lateral and ventral; 6, multilocular disk pore; 7, anal ring, right half; 8, hind claw; 9, trilocular pore. R. apizacos, new species: 10, Rostrum; 11, cephalic plate; 12, hind claw; 13, tritubular ceroris, lateral and ventral. 60 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 14-23.—Rhizoecus apizacos, new species: 14, Terminal segments of antenna; 15, anal ring, right half. R, arabicus, new species: 16, Trilocular pore; 17, anal ring, right half; 18, terminal segments of antenna; 19, tubular duct, lateral and ventral; 20, hind claw; 21, cephalic plate; 22, circulus, lateral; 23, rostrum. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 61

::•;;:.<2^;

^'''^P 25

FIGURES 24-29.—Rhizoecus associatus: 24, Terminal segments of antenna; 25, cephalic plate; 26, rostrum; 27, anal ring-, right half; 28, tritubular ceroris, lateral and ventral; 29, hind claw. 62 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 30-39.—Rhizoecus atlanticus: 30, Cephalic plate; 31, terminal segments of antenna; 32, mushroom bodies, various aspects; 33, tritubular ceroris, lateral; 34, rostrum; 35, tubular duct, lateral; 36, anal ring, right half; 37, eye, lateral; 38, circulus, lateral; 39, hind claw. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 63

\ /

/ A 41

54

FIGURES 40-54.—Rhizoecus hicirculus: 40, Anal ring, right half; 41, cephalic plate; 42, terminal segments of antenna; 43, circulus, dorsal; 44, hind claw; 45, rostrum. R. bituberculatus: 46, Bitubular ceroris, lateral; 47, anal ring, right half; 48, terminal segments of antenna; 49, rostrum; 50, cephalic plate; 51, tubular duct, lateral; 52, multilocular disk pore; 53, circulus, dorsolateral; 54, hind claw. 64 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 55-65.—Rhizoecus boharti: 55, Terminal segments of antenna; 56, cephalic plate; 57, anal ring, right half; 58, hind claw; 59, tubular duct, lateral; 60, rostrum; 61, bitubular ceroris, lateral. R. cacticans: 62, Terminal segments of antenna; 63, hind claw; 64, cephalic plate; 65, anal ring, right half. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 65

FIGURES 66-80.—Rhizoecus cacticans: 66, Rostrum; 67, circulus, lateral; 68, tubular duct, lateral; 69, tritubular ceroris, lateral. R. caladii: 70, Hind claw; 71, rostrum. R. californicus: 72, Trilocular pore; 73, termi- nal segments of antenna; 74, tubular duct, lateral; 75, anal ring, right half; 76, cephalic plate; 77, tritu- bular ceroris, lateral; 78, hind claw; 79, rostrum; 80, circulus, lateral. 66 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES Sl-SS.—Rhizoecus campestris: 81, Terminal segments of antenna; 82, tubular duct, lateral and ventral; 83, cephalic plate; 84, rostrum; 85, anal ring, right half; 86, anal lobe, right; 87, bitubular ceroris, lateral; 88, hind claw. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 67

FIGURES 89-97.—Rhizoecus chilensis, new species: 89, Terminal segments of antenna; 90, cephalic plate; 91, hind claw; 92, rostrum; 93, anal ring, right half. R. cyperalis: 94, Terminal segments of antenna; 95, rostrum; 96, tritubular ceroris and body setae, lateral; 97, hind claw. 68 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 98-110.—Rhizoecus disjunctus: 98, Terminal segments of antenna; 99, bitubular cerores, lateral; 100, anal ring, right half; 101, cephalic plate; 102, rostrum; 103, hind claw. R. distinctus: 104, Terminal segments of antenna; 105, tubular duct, lateral; 106, hind claw; 107, tritubular ceroris, lateral; 108, cephalic plate; 109, rostrum; 110, anal ring, right half. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 69

123 124

FIGURES 111-125.—Rhizoecus falcifer: 111, Anal ring, right half; 112, tritubular ceroris, lateral and ventral; 113, rostrum; 114, hind claw; 115, multilocular disk pore; 116, cephalic plate; 117, tubular duct, lateral and ventral. R. favacirculus, new species: 118, Trilocular pore; 119, cephalic plate; 120, anal ring, right half; 121, terminal segments of antenna; 122, hind claw; 123, tubular duct, lateral; 124, circulus, dorso- lateral; 125, rostrum. 70 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 126-139.—Rhizoecus floridanus: 126, Terminal segments of antenna; 127, tubular duct, lateral; 128, circulus, lateral; 129, cephalic plate; 130, hind claw; 131, rostrum; 132, anal ring, right half. R. glohocu- lus: 133, Medioventral pores, ventral and dorsal; 134, eye, lateral; 135, rostrum; 136, multilocular disk pore; 137, cephalic plate; 138, terminal segments of antenna; 139, hind claw. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 71

144 143 142

146 148

152 153

FIGURES U0-153.~Rhizoecus globoculus: 140, Cluster of medioventral pores. R. gracilis: 141, Terminal seg- ments of antenna; 142, cephalic plate; 143, tubular duct, lateral; 144, eye, lateral; 145, anal ring, right half; 146, hind claw; 147, circulus, dorsal; 148, bitubular ceroris, lateral; 149, rostrum. R. graminis: 150, Rostrum; 151, cephalic plate; 152, hind claw; 153, tubular duct, lateral and ventral. 72 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

^^c

162 164 163

FIGURES 154-165.—Rhizoecus graminis: 154, Terminal segments of antenna. R. insularis, new species: 155, Circulus, dorsolateral; 156, hind claw; 157, rostrum; 158, anal ring, right half. R. kondonis: 159, Anal ring, right half; 160, bitubular ceroris, lateral; 161, cephalic plate; 162, circulus, lateral; 163, hind claw; 164, rostrum; 165, tubular duct, lateral. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 73

173

FIGURES 166-174.—Rhizoecus kondonis: 166, Terminal segment of antenna. R. latus: 167, Eye, left, and trilocu- lar pore, lateral; 168, tubular duct, lateral; 169, cephalic plate; 170, anal ring, right half; 171, terminal segments of antenna; 172, rostrum; 173, multilocular disk pore; 174, hind claw. 74 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 175-188.—Rhizoecus leucosomus: 175, Tritubular ceroris, ventral; 176, terminal segments of antenna; 177, tubular duct, lateral; 178, anal ring, right half; 179, cephalic plate; 180, hind claw; 181, rostrum. R, macgregoriy new species : 182, Circulus, dorsal ; 183, tritubular ceroris, lateral ; 184, tritubular ceroris, ventral; 185, rostrum; 186, cephalic plate; 187, hind claw; 188; tubular duct, lateral and ventral. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 75

FiGVRES 1S9-195.—Rhizoecus macgregori, new species: 189, Terminal segments of antenna; 190, anal ring, right half; 191, trilocular pore. R. maritimus: 192, Rostrum; 193, cephalic plate; 194, terminal segments of antenna; 195, hind claw. TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 196-20S.—Rhizoecus maritimus: 196, Anal ring, right half. R. may anus: 197, Terminal segments of antenna; 198, rostrum; 199, cephalic plate; 200, hind claw; 201, anal ring, right half; 202, tritubular ceroris, large, ventral; 203, tritubular ceroris, medium, ventral; 204, tritubular ceroris, small, ventral. R. menkei: 205, Cephalic plate; 206, hind claw; 207, circulus, dorsolateral; 208, anal ring, right half. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 77

220

FIGURES 209-223.—Rhizoecus menkei: 209, Terminal segments of antenna; 210, rostrum; 211, bitubular ceroris, lateral. R. mexicanus: 212, Terminal segments of antenna; 213, anal ring, right half; 214, bitubular ceroris, lateral; 215, hind claw; 216, rostrum; 217, circulus, dorsal. R. nakaharai, new species: 218, Cephalic plate; 219, hind claw; 220, trilocular pore; 221, eye, lateral; 222, tubular duct, ventral; 223, circulus, dorsal. 78 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

232

FIGURES 224.-2Z2.-BMzoecus nakaharai, new species: 224, Terminal segments of antenna; 225 tntubularc^^^^^^^^^ ventral; 226, anal ring, right half; 227, rostrum. R. nemoralis: 228, Anal ring, right half; 229, terminal segments of antenna; 230, trilocular pore; 231, hind claw; 232, tubular duct, ventral. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 79

FIGURES 2SS-245.—R hizo ecus nemoralis: 233, Rostrum ; 234, tritubular ceroris, ventral. R. neomexicanus: 2Z^, Tubular duct, lateral; 236, tritubular ceroris, lateral; 237, cephalic plate; 238, terminal segments of antenna; 239, middle claw; 240, rostrum. R. neostangei: 241, Rostrum; 242, cephalic plate; 243, trilocular pore; 244, terminal segments of antenna; 245, hind claw. 80 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

'M^ ^ 247

246 s

"^■•/^i '\..„..;r-^ 248

252

251 253

255 254 256

FIGURES 246-256.—Rhizoecus nitidalis: 246, Terminal segment of antenna; 247, rostrum; 248, cephalic plate; 249, hind claw; 250, anal ring, right half. R, ornatus: 251, Tritubular ceroris and trilocular pores, ventral; 252, hind claw; 253, tritubular ceroris, lateral; 254, small tritubular ceroris and trilocular pores, ventral; 255, circulus, dorsal; 256, rostrum. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 81

FIGURES 2bl-2QZ.—Rhizoecus ornatus: 257, Anal ring, right half; 258, terminal segments of antenna. R, ovatus, new species: 259, Rostrum; 260, tubular duct, ventral; 261, anal ring, right half; 262, terminal segments of antenna; 263, hind claw. 82 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 264-273. Rhizoecus pauciporusy new species: 264, Terminal segments of antenna; 265, multilocular disk pore; 266, rostrum; 267, eye, lateral; 268, hind claw. R. poensis: 269, Cephalic plate; 270, anal ring, right half; 271, terminal segments of antenna; 272, hind claw; 273, rostrum. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 83

284 285 286

FIGURES 274-286.—R hizo ecus polyporus, new species: 274, Rostrum; 275, cephalic plate; 276, circulus, dorsal; 277, eye, lateral; 278, small circulus, dorsolateral; 279, tubular duct, lateral and ventral; 280, anal ring, right half; 281, hind claw; 282, terminal segments of antenna. Ä. pHic/^arcii: 283, Hind claw; 284, trilocu- lar pore; 285, tubular duct, ventral; 286, rostrum. 84 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

7,r ^^^^„

^>^\--^.'

288

287

293 294 295 292

FIGURES 287-295.—Rhizoecus pritchardi: 287, Terminal segments of antenna; 288, large tritubular ceroris, ventral; 289, small tritubular ceroris, ventral. R. relativus, new species: 290, Terminal segments of antenna; 291, anal ring, right half; 292, cephalic plate; 293, middle claw. R. simplex: 294, Rostrum; 295, cephalic plate. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 85

307 306

FIGURES 296-308.—Rhizoecus simplex: 296, Terminal segments of antenna; 297, hind claw; 298, anal ring, right half. R, solani: 299, Hind claw; 300, terminal segments of antenna; 301, anal ring, right half; 302, ros- trum; 303, bitubular ceroris, lateral. R, sonomae : 304, Anal ring, right half; 305, hind claw; 306, cephalic plate; 307, circulus, lateral; 308, rostrum. 86 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

310 312

O ......

315 '■'v!M'F" 314

W^^^"^^ 318

316 317

FIGURES 309-319.—Rhizoecus sonomae: 309, Terminal segments of antenna. R. spinipes: 310, Rostrum; 311, anal ring, right half; 312, hind claw; 313, terminal segments of antenna; 314, cephalic plate; 315, multi- locular disk pore. R. stangei: 316, Rostrum; 317, anal ring, right half; 318, cephalic plate; 319, hind claw. A REVISION OF THE NEW WORLD MEALYBUGS OF THE GENUS RHIZOECUS 87

327

FIGURES 320-327.—Rhizoecus stangei: 320, Terminal segments of antenna. R. svbcyperalis, new species: 321, Hind claw; 322, rostrum; 323, terminal segments of antenna. R. theobrovme: 324, Cephalic plate; 325, hind claw; 326, rostrum; 327, anal ring, right half. 88 TECHNICAL BULLETIN 1522, U.S. DEPT. OF AGRICULTURE

FIGURES 328-339.—Rhizoecus theobromae: 328, Terminal segments of antenna. R. totonicapanus: 329, Rostrum; 330, hind claw; 331, anal ring, right half; 332, cephalic plate; 333, terminal segments of antenna. R. tropi- calis, new species: 334, Circulus, dorsolateral ; 335, anal ring, right half ; 336, terminal segments of antenna; 337, rostrum; 338, cephalic plate; 339, hind claw. i^ U.S. GOVERNMENT PRINTING OFFICE: 1976 O 589-531