H. Morrison.
On Some Trophobiotic Coccidæ From British Guiana.
Psyche 29:132-152, 1922.

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132 Psyche [August
ON SOME TROPHOBIOTIC COCCIDB FROM BRITISH GUIANA.~ '
Bureau of Entomology, Washington, D. C.
Representatives of all of the species of Coccidae discussed in the following pages were collected by Dr. W. M. Wheeler of Bussey Institution, Harvard University, in the course of his investigatoins at the Tropical Research Sta-tion of the New York Zoological Society in British Guiana and were recently submitted to the writer for determination. Critical study of the material available for examination in the National Collection of Coccidae has shown that two of these species have also been collected in certain of the West Indian islands by the writer and others, and this opportunity has been taken to add these records to those from British Guiana.
SUBFAMILY MARGARODINX
GENUS STIGMACOCCUS HEMPEL.
Stigmacoccus asper (Hempel)
A number of specimens of this species were received with the following note by Dr. Wheeler :
"No. 757.
Kartabo, B. G. Sept 5, 1920. Taken from a huge colony of Crematoqaster sp. (near o.cuta Fabr.) nesting under bark of a large standing tree. The nest covered an area of more than 12 square feet and contained several hundred coccids enveloped in black carton. The young coccids were golden yellow, the older darker."
IPublished with the permission of r.he Secretary, U. S. Department of Agriculture. -The htimate relations describ'ed-as existing between ants on the one hand and the various Homoptera-on the other hand-have a common peculiarity. In all of these cases the ants are supplied with food in the form of an excretion or secretion elaborated from the juices of the plant^. Wasmann has therefore designated these relationships as trophoblosis to distinguish them from the cases of myrmecophily proper-." (Wheeler, Ants, etc. New York, Columbia Univ. Press, 1013, p. 360).




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19221 On some Trophobiotic Coccidce from British Guiana 133 Only preadult specimens and one partially molted adult could be located in the material submitted, and mounts of such specimens have been carefully compared with the corresponding stages from cotype material received by the Bureau of Entomo- mology from Prof. Adolph Hempel, the describer of the species. Some slight differences in the length and stoutness of the heavy spines crowding the derm of the preadult have been observed, but no other morphological characters that even suggest that the specimens from British Guiana represent another species than S. asper. In the absence of other stages, as larvae and well developed females and males, in which specific differences might appear, it seems best to regard the specimens from British Guiana as Hempel's species.
SUBFAMILY DACTYLOPIINB
GENUS Ph'E UDOCOCCUS WESTWOOD
Pseudococcus bromeliae (Bouch6)
References.-Ferris, Journ. Econ. Ent. vol. 12. no. 4, Aug.
1919, p. 196-Morrison, Phil. Journ. Sci. vol. 17, no. 2, Aug. 1920, p. 173.-Cockerell, Science n.s. vol. 55, no. 1492, 1922, p. 351. After extended crit,ical study the writer has placed as this species the materia,l from Dr. Wheeler's collections listed below together with his notes on the same.
"No. 116. Kartabo, B. G. July 17, 1920. In the cavities of the stems of a seedling
Cecropia (angulata I. W. Bailey).
The ants belonged to a black species of Azteca not yet identified." "No. 247. Kartabo, B. G. July 23, 1920. The coccids were in the hollow bases of the leaf-petioles of a Tachigalia paniculata Aublet with a black species of Azteca (Azteca foveiceps Wheeler) ."
"No. 367. Kartabo, B. G. Aug. 3, 1920. In node-like stem swellings of Cordia nodosa Lani. var. hispidissima Freser, with the ant Brachymyrmex sp. (probably heeri Forel)." No number. "Kartabo, July-Aug. 1920. In the swellings



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134 Psyche [August
of the stems of Cordia nodosa Lani. var. hispidissima Freser, with Allomerus octoarticulatus Mayr, the typical ant of this myrmecophyte.''
Nos. 209 and 175 have previously been identified in con- nection with Dr. Wheeler's extended account of the curious social beetles which he found in this region.
An exmination of these specimens in comparison with an extensive series of individuals from many other tropical and subtropical areas has indicated that this species is subject to a certain degree of variation in respect to the structural characters that are at present regarded as of taxonomic value, this variation occuring particularly in the number of spines in the different marginal cerarii, the shape of the ventral chitinized area at the caudal apex of the body, and the number of antenna1 segments. In the specimens collected by Dr. Wheeler the cerarian .
spines average slightly more numerous for corresponding cerarii than in typical forms from pineapple, that is, for example, in the same cerarius in ten specinlens instead of, say, four having four spines and the other six, three spines each, the proportions , may be six with four spines and four with three spines to each. The chitinized ventral thickening in nearly all of the specimens from pineapple is irregularly quadrate, at most only slightly longer than wide, but in the specimens from Dr. Wheeler, while the variation is marked, the average shape of each thickening is distinctly elongate, in this respect more nearly resembling the shape as figured by Ferris (ref. cited) for the species than the usual shape in the specimens from pineapple. The accompanying figure indicates this variation quite clearly. Finally, P. bromelice normally has 8-segmented antennae, while in a majority of the specimens sent by Dr. Wheeler these are 7-segmented; however, since these specimens show a range of from six to eight segments there seems no ground for attaching any taxonomic significance to the presence of one less segment in t,he majority of the antennae examined.
Prof. Cockerel1 (ref. cited) has recently called attention to the fact that the identification by modern coccidologists of the mealybug commonly cccuring on pineapple as the "Coccus



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19221 On some Trophobiotic Coccidce from British Guiana 135 bromeliae" of BoucM rests on a very unstable basis due to the ambiguity of the original description. As a matter of fact, it appears to be entirely impossible to determine positively what species was described by Bouch6 under this name, and our present notion of the species really dates from Signmet's redes- cription in 1875. As Prof. Cockerel1 suggests, his "Dactylopius brevipes" described from Jamaica on pineapple is identical with the species at present recognized as P. brornelice, but as Pseudo- coccus brevipes has never attained any standing among coccid workers, while several at present recognize and ident,ify a certain mealybug as P. bromehce, the writer has preferred to retain the older name regardless of the uncertainty as to what was actually described by Bouch6, at least until his species is clearly shown to be something other than the one at present given the name bromelice.
Pseudococcus rotundus sp. nov.
Occuring in cavities in stems of the host, attended by ants. A.dult Female.-All specimens available preserved in liquid, so nothing regarding normal external appearance can be given; maximum length as mounted 3 mm., maximum width 2.75 mm., specimens in preservative prop~rt~ionately somewhat more elongate, flattened beneath but strongly convex above; derm clearing completely on treating with potassium hydroxide, except for the appendages and the ventral chitinized areas; antennae normally 8-segmented, not unusual, the range of measurements in microns of segments in four individuals as follows: 1, 72-79; . 11, 68-79; 111, 54-64; IV, 36-46; V. 46-54; VI, 43-57; VII, 50-54; VIII, 107-114; legs normal, stout, hind femur and tibia with numerous tiny pores, those on femur frequently clustered, digitules probably normal, but all injured, claw stout, without denticle; beak elongate triangular, with a narrow basal collar and two much larger, distinct segments; with the usual two pairs of dorsal ostioles; normally wit,h 17 pairs of cerarii each composed of spines with accessory set% and a cluster



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136 Psyche [AU~US~
of triangular pores, but some of those on the thorax often greatly reduced or entirely wanting; the numbers of spines in the different cerarii showing the following range in four speci- mens: I (anterior), 3-4; 11, 3-4; 111, 2-3; IV; 0-3; V, 1-3; VI, 0-2; VII, 1-5; VIII, 0-2,; IX, 2-4; X, 2-4; XI, 3-4; XII, 2-4; XIII, 2-4; XlV, 4-5; XV, 4-5; XVI, 6-9; XVII (anal lobe). 11-22; in mounted specimens, wit,h the posterior two or three cerarii on each side dorsal and the remainder ventral; apex of abdomen not produced into lobes; ventral chitinous thickening elongate, bearing several setae; apical setae about 145p long; longest and anal ring seta about 93p long; anal ring of normal form and construction, with inner and outer rows of pores but with the setae more numerous than in other species of the genus, there being three larger primary set% and from five to ten smaller supplementary setae on each half of the ring,t,he smaller setae about half the length of the larger; dorsally with small triangular pores only. these scattered rather uniformly over the surface except along the body margin and at the cerarii where they are more numerous; ventrally with the triangular pores over most of the surface, with a few large, circular, multilocular disk pores around the genita,l opening, with two median trans- verse rows of tiny tubular ducts just anterior to these, and with a very few disk pores at the spiracuiar openings; cerarian spines conical, varying greatly in size; body with numerous, but scattered, slender setae, these averaging larger dorsally and most abundant along the body margin; with a single, large, q~adra~te median ventral cicatrix posterior to the hind legs* Immature Siaqes.~None a,vailable for examination. This species has been described from five specimens with the following da,ta, according to Dr. Wheeler's notes: "No. 87.
Barakara, B. G. July 15, 1920. In the cavities of the stems of Cecropia anqulata 1. W. Bailey. The ants in this Cecropia were a species of Azteca not yet identified." (Holotype). "No. 89.
Barakara, B. G. July 15, 1920. In the cavities of



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19221 On some Trophobiotic Coccidce from British Guiana 137 the stems of Cecropia angulata I. W. Bailey. The ants belonged .to a species of Azteca not yet identified.'" (Paratypes).
No number. "Kartabo, B. G. August 1920. In stems of Cecropia with Azetca sp. (Paratypes).
The types are in the U. S. National Collection of Coccidse, The proliferation of the anal ring set% in this species is a marked digression from the normal condition in the genus Pseudococcus and might be regarded as of sufficient importance to justify the separation of the species from that genus. How- ever, most, if not all, of the other structural characters of the species a,ppear to be characteristic of Pseudococcus and it has therefore been placed in that genus.
Genus Farinococcus, gen. nov.
A member of the group of which Pseudococcus is the typical genus: body oval, antennae %segmented, legs normal, without tiny pores, claw without denticle, beak short triangular, 2- segmented, spiracles stout with large opening and numerous disk pores near mouth, two pairs of dorsal ostioles, cerarii very large, composed of many lanceolate spines, set= and pores and joined into a continuous band anteriorly; anal lobes not de- veloped, wit)h ventral chitinized thickening and apical seta, anal ring normal, with pores and six setae, derm with triangular pores dorsally and ventrally and many multilocular disk pores ven- trally in genital region, no other pore types, derm with slender set= dorsally and ventrally, with a single median ventral ci- catrix. Genotype.-F'arinoccusus multispinosus, sp. nov. This genus diverges from the normal Pseudococcine type most conspicuously in the marked development of the cerarii and the great increase in the number of ventral disk pores, and, to some extent, in the enlargement of the spiracles. Farinococcus multispinosus sp. nov.
Occuring in cavities in the stems of the h~st~attended by ants. Adult female.-Preserved in liquid, so nothing regarding the normal external appearance can be given; color of alcoholic



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138 Psyche [August
specimens mostly a light yellow brown; body elongate oval, tapering somewhat behind the middle; average length 3.5 mm., width 2.25 mm; uniformly elongate oval when flattened on a slide; clearing completely, except for appendages and ventral chitinized thickenings, on treating with potassium hydroxide; antennae normal1 y &segmented, not unusual, the range of measurements in microns of the different segments as follows: I, 68-78; 11, 64-71; 111, 46-53; IV, 28; V, 43-57; VI, 43; VII, 46; VIII, 110-118; legs fairly large, stout, the claw wit,hout denticle, the digitules normal, those of claw slightly knobbed at tip, all the legs without tiny pores, but all the cox= with large distinct ba,sal aerola.tions, and the femora and tibise with similar, but quite indistinct, very large aerolat'ions on the upper halves of each; beak short amd stout triangular, two segmented; spiracles large and stout, with a number of disk pores near mouth; with two pairs of heavy, thick-lipped dorsal ostioles, cerarii very strongly developed, those on the posterior abdominal segments narrowly but distinctly isolated from each other, those on the anterior portion of the body united to form a narrow continuous submarginal band made up of hundreds of closely crowded lanceolate spines and triangular pores flanked within by dozens of slender setse aggregated to form more or less distinct groups; posterior cerarii made up of numerous lanceolate spines (apical about 70, preapical about 85, ante- penultimate about 80, fourth from last about 70, next about 65, next about 55) and within each of these ventrally a triangular cluster of setse and in and around both spine and set% clusters numerous triangular pores; anal lobes not developed, ventral chitinous thickening heavily chitinized, irregular in shape, en- closing three or four setse; apical setz set off just outside the ventral thickening, large and stout as compared with the anal ring setse, perhaps three times the length of these (broken); anal ring placed dorsally well anterior to the body apex, small, of normal structure, with double pore row and six setse, the derm on each side of the ring protruding and overlapping to form t-wo longitu- dinal folds covering the ring, except for a linear or oval median slit; derm dorsally with numerous, uniformly distributed,



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192.21 On some Trophobiotic Coccidce from British Guiana 139 triangular pores, ventrally with these pores less numerous, but also uniformly distributed, except at and near the body margin, where they are close1 y crowded, with numerous large circular multilocular disk pores in a heavy collar around the genital opening and in transverse bands on three segments anterior to genital collar, and quinquelocular disk pores of approximately the same size as the genital pores in numbers at the spiracles; no other pore sorts located; dorsally and ventrally with numbers of scattered, slender, faintly lanceolate setse; with a single large quadrate median ventral cicatrix posterior to the hind legs. Preadult female.-In general resembling the adult quite closely, differing in the smaller size, the 7-segmented antennae, t'he smaller and less developed spiracles, the reduced numbers of spines, seta? and pores in the cerarii and the very great reduction in the number of ventral disk pores.
No other stages have been available for examination. This species has been described from four mounted and a few unmounted specimens received from Dr. Wheeler with the following note :
"In the ca,vities of t,he stems of Triplaris surinamensis Chanc. with Pseudomyrma sp. nov. (allied to Ps. triplaridis Forel.) "
The types are in t'he U. S. National -Collection of Coqcidz. This genus is at present merely a convenient dumping ground for those species of mealybugs having the number of antenna1 segments reduced and tlhe cerarii usually much reduced in number or even wanting. Until the genera of the true mealy- bugs are based on an adequate morphological foundation, little else can be done with apparently new species than to place them, in some such convenient genus as this one, and it is on this basis that the two species described below have been assigned to a position here.




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Psyche
Ripersia petiolicola sp. vov.
Occuring in the hollow bases of the leaf petlioles of the host, attended by ants.
Adult Female.-Preserved in liquid leaving none of the se- cretionary covering available for examination, average length as mounted, 1.3 mm., width 0.875 mm; clearing completely on treating with potassium hydroxide, except for the appen- dages; antennse small, placed fairly close together at anterior apex of head, not unusual, normally 6-segment,ed, but frequently 5 and rarely 4-segmented, the normal measurements in microns ranging as follows: I, 33-39, 11, 29-36; 111, 36-43; IV, 18-25) V, 18-25; VI, 50-61; legs normal, fairly stout, claw without denticle, hind femur and tibia each with a number of pores; beak triangular, with an incomplete basal collar and two distinct segments; with the usual t,wo pairs of dorsal ostioles; only the two anterior and the two posterior cerarii developed to a definitely recognizable state, the anal pairs with relatively large conical spines, several triangular pores and four to six accessory setae, the spines in the other recognizable cerarii slender, lanceolate, setalike, the pores less numerous and the accessory setae fewer or wanting; probably with a more or less complete series of cerarii around the body margin, but these not definitely re- cognizable, due to the separation of the component parts; anterior cerarii with three spines in each, all others with two; anal lobes slightly indicated, with the apical cerarii underlaid by a faint suggestion of chitinization, with a small, irregular, elongate ventral thickening and an apical sets about 100p long; largest anal ring seta about 78~; anal ring normal with the usual pores and six setae; body, as far as noted, with only two sorts of derm pores, a few multilocular disk pores around the genital opening and smaller triangular pores scattered rather uniformly over the surface, but more numerous along the margin and in the cerarii; body dorsally with some scattered, slender, lanceo- late setae and ventrally with even fewer, distinctly longer and more slender, hair-like setse; with a single, indistinct, transverse oval ventral cicatrix posterior to the hind legs.



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19221 On some Trophobiotic Coccidce from British Guiana 141 Immature stages.--None available for examination. This species has been described from three specimens having the following collection data according to Dr. Wheeler's not,es: 'No. 247. Kartabo, B. G. July 23, 1920. The coccids were in the hollow bases of the leaf-petioles of a Tachigalia paniculata Aublet with a black species of Azteca (Azteca foveiceps Wheeler .) " . .
The types are in the U. S. National Collection of Coccidse. Ripersia subcorticis sp nov.
Occuring under the bark of the host, attended by ants. Adult Female.-Preserved in fluid, so nothing regarding the normal external appearance can be given; color of alcoholic specimens ranging from yellow brown to dull purple, but usually with a purple tinge; stout oval, flattened beneath, fairly convex above, average length of mountled specimens 2 mm., width 1.5 mm; clearing completely, except for appendages, on t>reating with potassium hydroxide; antenna? not unusual, normally 7- segmented, but frequently with six, the range of measurements in microns of those available for examination as follows: I, 47-54; 11, 39-50; 111, 25-3.5; IV, 22-32; (111 and IV where combined, 57-61), V, 22-29; VI, 28-32; VIIJ, 71-82; legs normal, the claw without denticle, the hind coxa, but not the femur and tibia, with a number of pores; beak elongate triangular, with an obscure basal collar and two dist,inct segments; with the usual two pairs of dorsal ostioles; cerarii, as such, not distinctly developed, but with loose marginal or ventral submarginal clusters of slender, conical to slightly lanceolate spines, a,c- companied by a closer grouping of the small triangular pores, these clusters distinctly isolated on the posterior a,bdominal segments, but forming an almost continuous but inconspicuous band around the anterior margin of the body; anal lobes barely indicated, the apical seta immediately adjacent to the posterior cluster of spines, length not certain, probably about one and a half times that of the anal ring setae, without any trace of ventral chitinous thickenings; derm dorsally with only the small tri-



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142 Psyche [August
angular disk pores, these scattered rather uniformly over the body surf ace ; ventrally with these pores distributed uniformly except at margin close to and in the "cerarii," with some large circular, i~ultilocular disk pores around the genital opening and in narrow transverse rows on the two segments anterior to this, with a very few, tiny tubular ducts near the genital opening and with much larger, short tubular ducts in clusters just within the "cerarii" on the last five or six abdominal segments, these ranging from four to eight in number in the three clusters anterior to the apical but with only one or rarely two in the one or two anterior groups, the apical group median, usually with about twelve pores in it; derm dorsally with fairly uniformly scattered, very slender, lanceolate setae, ventrally with longer, slender hair-like setae; anal ring of normal form, with inner and outer rows of pores and with three primary and usually four smaller, secondary setae on each half, longest anal ring seta about llOp; with a single median quadrate ventral cicatrix posterior to the hind legs.
Imma,ture stages.-None available for examination. This species has been described froin four mounted and a few unmounted specimens collected by Dr. Wheeler with the following note :
"No. 79. Bmakara, B. G. July 15, 1920. Taken in a large colony of the ant Tranopelta gilva Mayr, under the bark of a living tree. The white coccids were present in great numbers over the whole surface of the wood. The ant is usually sub- terranean, its nest under bark being very exceptional." The types are in the U. S. National Collection of Coccidse. In t,he multiplication of the anal ring set% is found a con- dition exactly ~ompara~ble to that in Pseudococcus rotundus, just described, and it is also similar to the structural modifications of the anal ring in certain other relat'ed forms known tJo be pro- tected and attended by ants, such as Lachnodiella cecropice, suggesting the possibility that the relationship between the ant and mealybug may have had some influence on the modification in structure.




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19221 On some Trophobiotic Coccidoe from British Guiana 143 SUBFAMILY COCCING,.
GENUS AKERMES COCKERELL
Following Prof. Newstead's lead, the two species discussed below are continued in the genus Akermes, although a com- parison of the two with the genot>ype raises a very serious question as to the correctness of such generic association. Akermes quinquepori (Newstead)
Reference,-Newstead Bull. Ent. Res. vol. 7, 1917, p. 349. This species is represented in the material submitted by Dr. Wheeler by three lots of specimens. Dr. Wheeler's notes in regard to these are as follows:
"No. 76. Barakara, B. G. July 15, 1920. Found in a large colony of ants (Camponotus novogrenadensis Mayr) which had their nest within that of Hamitermes excellens Emerson (MS.) in the bark of a large standing t'ree. The galleries of the ants interdigitated but did not communicate with those of the ter- mites. The coccids were in the ant galleries and attached in great numbers and in all st,ages to the surface of the tree itself. It was difficult t30 remove them without breaking them. A large piece of the inner bark with the coccids was removed with large coccids attached to it. The insects were seen to exude copious drops of clear honeydew from the brown anus on the dorsal surface,"
"No. 87.
Barakara, B. G. July 15, 1920. In tlhe cavities of a stem of a Cecropia angulata I. W. Bailey. The ants in this