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EN TOMOGRAPHY

Volume 4 1986 pp. 137-183

The Whitefly Genus Aleurocerus Bondar


(Hemiptera, Homoptera, Aleyrodidae)

Louise M. Russell
CooperatingScientist, Systematic Entomology Laboratory, BBII, Agricultural Research
Service, USDA,Beltsville, Maryland 20705
ABSTRACT. — Aleurocerus Bondaris known from Mexico southward through Brazil.
The genusand four species, A.cerifer (Sampson and Drews), A. flavomarginatus Bondar,
A. luxuriosus Bondar, and A. tumidosus Bondar are reviewed and redescribed. Six new
species, A, chiclensis, A. coccolobae, A. colombiae, A. musae, A. palmae, andA. petiolicola
are described. All species areillustrated. Keys are providedfor the separation of immature
stages andfor the identification of species.
This revisionary study of the genus Aleurocerus Bondar was under-
taken in order to provide names for species collected from plants of
commercial value atports of entry into the United States. Thousands of
examples of the genus have been collected by plant quarantine inspec-
tors of the United States Department of Agriculture since 1936 when
specimenswerefirst intercepted. Because previous work on Aleurocerus
is limited to incomplete descriptions andinclusion inlists, it has been
necessary to study the genuscritically. The investigation hasresulted in
aredefinition ofthe group, aredescriptionof the genus andfourspecies,
andthedescription of six new species.
Species of Aleuroceruslive on plants of economic value. Decorative
palms, bromeliads, orchids, and philodendrons,all imported for orna-
mental purposes, andotherplants oftrade, including banana, coconut
palm, and achicle producingtree,are hosts of the whiteflies. Frequently,
leaves are virtually covered by the aleyrodids, plants are rendered
unsightly, and their aesthetic value is lessened by the presence of the
black pupae,their copious, dirty white secretions, and the black fungus
that grows in the honeydew excreted by the insects. The genus is
recorded here from plants belonging to 12 families and from undeter-
mined hosts, and it is known from the southern part of the Nearctic
Region in Mexico through the Neotropical Region to southern Brazil.
Aleurocerus is the only known genusofthe Aleyrodidaein which the
anal area of the pupa terminatesin a bifid horn. The genusalso exhib-
its notable diversity in its wax producing organs.
Unidentified parasites frequently are present in pupae andthird-
stage larvae, but I have not observed them in otherstages.
138

Theclassification of Aleurocerus is based primarily on the morpho-


logical characteristics of the fourth stage larva, commonly known asthe
pupacaseor pupa.A differencein the location of immatures on the host
is the only other observed feature.If the copious secretions of the pupae
were in perfect condition, they doubtless would show differences
between some species, but in the specimens examinedthey are too
imperfect to be helpful. Thebiology of species has notbeen investigated.
Third-stage and second-stage specimens can sometimes be separated
specifically, but first-stage larvae do not exhibit specific differences.
Although adults havesatisfactory diagnostic characters, the specimens
examinedare too fragmentary to permit adequate species descriptions.
Taxonomic study of the pupae of Aleurcerus is an arduous task
becauseit is difficult to obtain mounted specimensin which structures
can be seen clearly, and structures appear differently under different
degrees of bleaching. Mature pupaeare black, brittle, unevenly curved
over the dorsal surface and tend to break and curl during the mounting
process. Since recently emerged pupae have not developed the intense
blackness andrigidity of older specimens, they can be mounted and
studied more successfully than older ones. Removal of somecolor is
necessary, but this must be donecarefully for structures are obliterated
by overbleaching. Immersion in hydrogen peroxideor six parts of hyd-
rogen peroxide combinedwith onepart of ammonia,incorporated in the
mounting procedure, mayresult in a sufficient loss of color. Portions of
the margin and submargin must be flattened if structures and their
relationship to one anotherare to be determinedaccurately; they usually
can be seen moreclearly in detached fragments than in whole bodies.
Whenheavily sclerotized, the venter needs to be mounted separately
from the dorsum ifits characteristics are to be determined.

TERMINOLOGY
Structures used in this study of Aleurocerusare depicted in Fig. 1, a
diagramatic drawing of one-half of the dorsum of a pupa, and are
labeled according to my usage. Although several terms have been
defined by Russell (1943, 1948) or are readily interpretable, it may be
desirable to explain someof them briefly.
The dorsum is the entire dorsal surface and includes areas desig-
nated margin, submargin, subdorsalfurrow, and dorsal disk. The mar-
ginis the narrow distal portion of the dorsum. Marginalteeth are the
distal part of the margin; marginal ridges and furrowsextend from
the teeth to the submargin; ridges may have pale areas of various
shapes. The submarginis a ridge that nearly parallels the curveof the
body margin and extends from the margin to the subdorsal furrow.
Glands(submarginalteeth of Bondar, 1923) are somewhatconical or
rectangularstructures in the submargin. Transverse ridges and fur-
rowsrun transversely to the submargin; the distal ends of ridges,if
pointed, are said to terminate in a tooth. The subdorsal furrow is a
139

QU TTT tt = anterior margi


nal sera
Se marginal teeth
5 _-— marginalridge
7 -ae
- == margginal
inal furr
fur ow
ot ae eve spot
=> submarginal gla
nds
-

—~ dorsal disk
== subdorsal area

¢ submarginal seta
+ transverse molting suture

'- metathoracic seta


+ submarginal ridge
~ subdorsal furrow
~ spinules
—* median and submedian areas

= vasiform orifice
-« homed process
=
_- caudal seta
"== posterior marginal seta

Fig.1. Diagrammatic drawingrej ‘pres


enting one-halfof the dorsum of an aleur
Pupa, ocerine
140

depressed area that separates the submargin from the dorsal disk. The
dorsaldisk is the entire area within the subdorsal furrow. The rachis
is a median ridge on the abdomen. Onthe dorsal disk, a subdorsal
locationis the area between imaginary curved linesprojected cephalad
and caudad to the subdorsal furrow from the outermost points of the
rachis; a submedianlocationis the area just within the imaginary
curved lines andit extends to the median area, the longitudinal center
section of the dorsal disk. The horned processis a modification of
abdominal segment VIII that terminates in a bifid horn; it may have
sclerotized arms extending from its basal angles. The vasiform
orifice, the anal orifice, contains the operculum and lingula. The
venteris the entire ventral surface.
Setae are named from the area or body segment where they occur;
anterior and posterior marginalsetaeare located on the body mar-
gin; submarginalsetaeare on the submargin; metathoracic setae
are on the metathorax; eighth abdominal setae are on abdominal
segmentVIII; caudalsetaeare at the posterior end of the body. spin-
ules vary from minute, needlelike points to relatively large, blunt or
pointed structures. Poresofvarioussize, shape, and structure are pres-
ent. Minute poresoccurin the margin and submargin; some are min-
uscule and obscure while othersare readily visible; some appearas clear
areas with indistinct margin. Disk poresare minute,circular, dark-
rimmed andnonloculate. Locular poresare larger thandisk pores, are
irregular in outline, and have 4-12 loculi; minute columnsprojecting
from the loculi are visible under optimum conditions and are most
distinct when seenin profile; they may be analagousto the porettes of
Russell (1943, 1948) and the pseudopores of Mound (1967). Disk and
locular pores probably are associated with each other althoughin the
submargin their numbers sometimes appearto differ.

CHARACTERS OF VALUE
Differentiating characters are present throughout the dorsum in
pupae andoftenin third-stage larvae; the following are useful. Margin:
Color, numberof teeth between marginalsetae, shapeandsizeofmargi-
nalteeth, length of marginal ridges, absence or presence and shape of
pale areasin ridges, size and location of pores. Submargin: Shape and
size of glands, characteristics of the transverse ridges and furrows,
numberandlocation of disk and locularpores, length of setae. Dorsal
disk: Numberandlocationof disk and locular pores, numberanddistri-
bution of spinules, location of ends of transverse molting suture, loca-
tion of bases of eighth abdominal and caudalsetae, size of the horned
process and location of the vasiform orifice. The basal arms of the
horned process vary considerablyin size and direction and are of limited
usefulness.
Theventeris fairly uniform in pupae andlarvae of Aleurocerus and
doesnot appearto have speciesdifferentiating characteristics. Although
141

the porousor sieve-like appearance ofthe surface mayvarygreatly, this


appears to be a matterof intensity of sclerotization. The length of the
abdominalsetae is not a good separation character.
In adults, characteristics of antennae, sensoria, eyes, setae, and male
genitalia are significant.
Descriptions of the pupae and third-stage and second-stage larvae
are arranged by the margin, submargin,and dorsal disk. Comparative
length and width ofmarginalteeth are for those at the wider parts of the
body; the widthis at the base and the length is from base to apex. The
horned processis described as extending caudad. Species descriptions
are arranged alphabetically.
Holotypes of new species are mounted on slides and are deposited in
the collection of the National Museum of Natural History, Beltsville,
Maryland (USNM). Paratypes andother specimensarein this institu-
tion and elsewhere as indicatedin the text. Repositories are as follows:
California State Departmentof Agriculture, Sacramento (CDA); Florida
State Collection of Arthropods, Gainesville (FSCA); British Museum
(Natural History), London (BMNH); Istituto di Entomologia Agraria
dell’ Universita di Napoli, Portici (IEUN); Zoological Academy of
Sciences of USSR, Leningrad (ZAS). Syntypes of previously described
species presumably are.in the Museu de Zoologia da Universidade de
Sao Paulo, Brazil.

Genus Aleurocerus Bondar


Aleurocerus Bondar,1923:102, 156; Drews and Sampson, 1940.90; Sampson, 1943:199, 218,
1947:49; Sampson and Drews, 1957:692; Mound and Halsey, 1978:27. Type species,
Aleurocerus luxuriosus Bondar,by original designation.
Uraleyrodes Sampson and Drews, 1941:180; Sampson, 1943:178, 220; Mound and Halsey,
1978:27. Typespecies, Uraleyrodesceriferus Sampson and Drews, byoriginal desig-
nation and monotypy.
Mound andHalsey synonymized Uraleyrodes with Aleurocerus on the basis of original
descriptions, I have examined syntypesof luxuriosus and paratypesof cerifer! and verify
the synonymy.
Mound and Halsey (1978) believed that Sampson and Drews were
unaware of Aleurocerus. The latter authors must have known of Aleu-
rocerus, however, because they (1940)cited Bondar 1923, and they (1941)
included a Bondar1923 genusintheir publication. Although the reason
for the Sampson and Drewsoversightis unclear, it may have been in
part because they (1941) placed Uraleyrodes in a new subfamily, the
Uraleyrodinae. Sampson (1943) knew and characterized each genus,
placing Uraleyrodesin the Uraleyrodinae and Aleurocerusin the Aley-
rodinae. Later, in keys to the Aleyrodinae, Sampson (1947) and Samp-
son and Drews (1957) separated Aleurocerus from other genera by the
presenceofits bifid horn. It seemslikely that the authors simply over-
looked the similarity between Uraleyrodes and Aleurocerus.
‘Steyskal (1979, pers. comm.) indicated that words ending in -fer do not take us in the masculine.
Accordingly, the specific nameis changed from ceriferus to cerifer.
142

In describing Uraleyrodes, Sampson and Drews(1941) stated “. . .


vasiform orifice absent; anal opening overhung by bifid caudal horn
bearing long hairs...” This statement doubtless lead to that of Mound
and Halsey (1978), “. .. The abdominalhorns overlaying the vasiform
orifice...’’ Later, Sampson(1943) alleged “... Anal opening overhung by
a bifid caudal horn bearing twolong hairs, the opening being a narrow
transverseslit.”
In his description of Aleurocerus, Bondar (1923) wrote (translation)
“... Vasiform orifice obscured, andthe plate that supports it terminates
posteriorly in two stout horns, turned backward, diverging somewhat
laterally ...” In his characterization of Aleurocerus, Sampson (1943)
asserted “... Vasiform orifice obscured; posterior edge prolonged intoa
bifid horn.”
Uraleyrodes was based on a misconception of the anal area. The
black color of the pupae could easily have concealed features of the
horned processsufficiently to have lead Sampson and Drews (1941) and

|
Sampson(1943)to err in their understandingof it. Actually, the vasi-
form orifice, whichis the anal opening, is present andis almost semicir-
cular rather than a narrowslit. An extension of the derm (“‘plate” of
Bondar (1923)), not the vasiform orifice, terminates in a bifid horn. In
nature the horned processis erect, and the vasiform orifice is located on
its cephalic surface. Whentheinsects are mountedon aslide, the horned
process, if very long, may turn cephalad and obscure the vasiform
orifice, but it usually turns caudad and the vasiform orifice then is
visible in bleached specimens. The eighth abdominalsetae arise near
the base of the horned process. The caudal setae arise on the body
surface, not on the hornedprocess, but their bases may be covered by the
horned process whenit is turned caudad. Althoughcerifer is markedly
distinct from luxuriosus, differences between the two do not appearto be
adequate for considering Uraleyrodes distinct from Aleurocerus.
In Aleurocerus the larval skins tend to remain attached to each other.
Thethird larval skin maybe ontop of the waxy secretion covering the
pupa,the second larval skin remainsonthethird, andthefirst larval
skin rests on the second.
Aleurocerus is closely related to Aleurothrixus Quaintance and
Baker(1914). Pupae of these genera have the submargin separated from
the dorsal disk by a furrow; they havea rachis, at least four pairs of
submarginalsetae on the anterior half of the body, and a pair of sub-
median metathoracic setae; they do not have submedian setae on the
first abdominal segmentor setae on the lingula.
Pupaeof Aleurocerus can be distinguished from those of Aleuro-
thrixus and other described genera by the extraordinary development of
the portion of the posterior segment that terminates in a bifid horn.
Third-state and second-stage larvae also can be distinguished by the
presence of a bifid horn, but a hornis absent from first-stage larvae and
adults.
143

Description of Aleurocerus
Usually living on the lower surface of leaves, occasionally on the
leafstalk, bark or on aerial roots.

Pupa
Description. Mature pupae with a fringe of white, somewhat fibrous,
crinkly filaments extending outward from the body margin; ribbons of
waxarising from the dorsum;a thin layer of white, paraffin-like wax on
dorsum usually concealed by white, fibrous filaments; white, translu-
cent wax covering the venter, thickest near the body margin.
Ovoid, oval, or broadly oval in outline. Dorsum unevenly convex,
heavily sclerotized, entirely brown or black, or margin yellowish or
reddish. Venter flat, moderately to heavily sclerotized, entirely pale,
brown,or black, or margin yellowish or reddish.
Margin. Reddish, yellowish, dark brownor black. Anterior and poster-
ior setae present. Margin dentate, smallest teeth at anterior and poster-
ior ends of body;thoracic tracheal pore area teeth narrowerthan, or as
wide as otherteeth; abdominal tracheal pore area teeth narrower than
other teeth. A minusculeporein notch betweenteeth or on ridges. Ridges
proximad of teeth, without or with indistinct or distinct pale areas.
Furrowsproximadof notches betweenteeth, narrower than ridges, with
a minutepore.
Submargin. Submarginalridge nearly paralleling body margin, sep-
arated from dorsal disk by the subdorsal furrow. Glands present or
absent; when present smallest at ends of body. Transverse ridges and
furrows weakto strong,their spinules varying from needle-like to blunt
and stout. Disk pores present. Locular pores present. Setae slender, 3
pairs on cephalic segmentsmaller and nearerdistal margin than others;
1 on each thoracic segment and on abdominal segmentI.
Dorsal disk. Eye spotsfaintor distinct, located in subdorsal furrow.
Disk and locular pores present. Rachis present on abdominal segments
I-VIII. Spinuleslocated on rachis. Transverse molting suture slightly
anterior to thoraco-abdominalsuture, curved posteriorly from its mid-
point, recurved and terminating opposite its midpoint or somewhat
anteriorto that at or near the subdorsal furrow. Meso-metathoracic and
abdominalsutures well defined in median area; 8 segmentsvisible in
medianarea of abdomen. Medianlength of cephalic segmentas great as
combined length of subequal thoracic ones; abdominal segment I
slightly longer thanII, II slightly longer than subequal sements III-VII,
each shorter than VIII. Submedian depressions weak,visible on thorax
and abdomen. Pockets small. Submedian metathoracic, eighth abdomi-
nal, and caudalsetae slender, elongate. Modified portion of abdominal
144

segment VIII terminating in an outwardly curved bifid horn; with or


without arms extending cephalad or laterad from base of horned pro-
cess. Vasiform orifice located in horned process, with or without a weak
rim around posterior end,its sides straight near top, curved below,its
bottom extending to anterior marginoforifice; operculum semicircular
or subtriangular,2-3 times wider thanlong; lingula spatula- or toungue-
shaped, shorter than operculum,spinulose, without setae. Caudal fur-
row and caudal ridges absent.
Venter. Appearing minutely porousorsieve-like if densely sclerotized,
smooth if membranous. Marginal wax tubes usually visible. Antennae
lying betweenforelegs and body, 1-segmented,reaching nearly to anter-
ior spiracles; membranous, spinulose,tapered distally; apex apparently
with minute setae and a small spine. Median setae absent anterior to
mouthparts. Segmentationof rostrum obscure, possibly 2-segmented, 3
pairs of minute setae sometimesvisible at apex. Tracheal folds well
defined, thoracic ones usually without, abdominal one with spinules.
Thoracic spiracles smaller than anterior abdominal ones, the latter
smaller than posterior abdominalones. Legsstout, distal margins in a
furrow in derm, withoutdistinct segmentation; often with 2 or 3 minute |
setae near disk on eachleg and | or 2 slightly larger setae on basal area
of middle and posterior legs. Adhesive sacs prominent. Minute circular
pores sometimes visible in submedian area of some abdominal seg-
ments. Abdominalsetae located mesocephaladof posterior abdominal
spiracles. Male organ a bifid sac,

Third-stage larva
Description. Marginal waxy filaments elongate, nearly uniform in
length; dorsal filaments not coalesced.
Elliptical, oval, broadly oval, or ovoid in outline. Dorsum slightly
convex, weakly to moderatelysclerotized,entirely pale or dark brown, or
margin pale. Venter colorless, membranous,flat.
Margin.Anterior andposteriorsetae present. Margin dentate, thoracic
tracheal pore area teeth narrowerthanor as wide asother teeth. Ridges
proximadofteeth; ridges with pale areas of various shapes; each pale
area with 1-5 circular or elliptical minuscule pore-like sections at its
proximalend. Furrows proximadof notches between teeth, each with a
minute pore.
Submargin. Submargin with weak transverse ridges and furrows.
Subdorsal furrow weak. Glands absent. Disk andlocular pores present.
Spinules absent. Setae slender, 3 pairs on cephalic, 1 pair on each
thoracic segment and on abdominal segmentI.
Dorsaldisk. Eye spots present, weak. Disk andlocular pores present.
Rachis weak. Spinules present or absent. Abdomen with 8 segments
visible in median area. Metathoracic, eighth abdominal, and caudal
setae slender, elongate. Modified portion of abdominal segment VIII
145

terminating in an outwardly curvedbifid horn; with or without arms at


basal angles of horned process. Vasiform orifice located in horned pro-
cess, semicircular; operculum semicircular, 2-3 times wider than long;
lingula tongue-shaped, spinulose, without setae. Caudal furrow and
caudalridges absent.
Venter. Marginal wax tubes weak or not visible. Antennae reaching
anterior legs, proximal half stouter than finger-like distal half, spinu-
lose. Median setae absentanterior to mouthparts. Rostrum indistinctly
2-segmented; distal segment apparently with 1 pairof minute setae near
base, 1 at midlength, and 1 at apex. Tracheal folds usually visible.
Thoracic spiracles larger than anterior abdominal ones, as large as
posterior abdominal ones. Legs short, tapered, somewhat bulb-shaped,
curved on distal and proximal margins, usually with 1-3 minute setae
near disk. Adhesive sacs not visible. Abdominalsetae located meso-
cephaladofposterior abdominalspiracles. Male organ absent.

Second-stage larva
Description. Similar to third stage except as follows: Waxy secretions
shorter. Submargin with or without disk and locular pores. Dorsal disk
with or without disk and locular pores and spinules. Abdomen with 7
segments visible in median area, 8 visible in subdorsum. Submedian
cephalic, mesothoracic, and metathoracic setae dagger-shaped; eighth
abdominaland caudal setae slender, elongate. Antennae not reaching
anterior legs. Legs usually with 2 minute setae near disk and apparently
2 farther from disk on middle andposteriorlegs.

First-stage larva
Description. Waxy secretion absent.
Somewhatelliptical in outline, narrowed at anterior end. Dorsum
brown, sclerotized, flat or slightly convex. Venter colorless, membran-
ous,flat.
Anterior and posterior marginal setae present. Margin minutely
dentate, tracheal pore areas not visible. Submarginal] setae of later
immature stages slender, located on body margininstead of on submar-
gin, 3 pairs on cephalic, 1 on each thoracic segment and on abdominal
segment I. Eye spots circular. Submarginal ridge, glands, subdorsal
furrow,rachis, disk and locular pores, and spinules absent. Segmental
sutures poorly defined, 7 segments visible in median area of abdomen, 8
visible in subdorsum. Submediancephalic, mesothoracic, metathoracic,
and eighth abdominalsetae stout, nail-shaped; caudal setae slender,
elongate. Area around vasiform orifice slightly projected, with or with-
out basal arms, without a bifid horn. Vasiform orifice subcircular,its
inner walls smooth; operculum subcircular, nearly filling orifice, its
caudal margin spinulose; lingula tongueshaped, spinulose, without
setae.
146

Venter. Marginal wax tubes andtrachealfolds not visible. Antennae


apparently 2-segmented, reaching to middle legs; basal segment
slender, distal segment slenderer and about 3 times length of basal
segment; 1 minute seta at apex, possibly | minute seta neardistal 4th.
Mediansetae present anterior to mouth parts. Rostrum apparently 2-
segmented, apparently 1 pair of minute setae near base and | pair near
distal end. Tracheal folds not visible. Thoracic spiracles larger than
abdominal ones. Legs jointed, functional; coxa globose, with 1 elongate
seta on proximal margin; trochanterindistinct; femur about as long as
wide, apparently withoutsetae;tibio-tarsus tapering from baseto apex,
1 elongate seta on outer surface of each anterior leg, 1 near midlength
and 1 minute seta just distad of elongate | on distal margin of each
middle and posterior leg; tibio-tarsus with 1 minute seta on proximal
margin and | elongate digitule on distal margin at end of segment;
terminating in a fleshy appendagelocated between 2 stubs. Adhesive
sacs absent. Abdominalsetae located mesocephaladof posterior abdom-
inal spiracles. Male organ absent.

Adults
Description. Males smaller than females.
Head. Vertex broadly curved; with dorsal and ventral slender setae and
minute spinules; without a median ocellus. Compoundeye in2 sections,
1 somewhatcephalad anddorsadof the other, the 2 narrowly separated
or joined by 1-3 facets; facets of the dorsal section smaller than those of
the ventral section; somefacets of ventral section dark, others pale. An
ocellus located at anterior margin of dorsal section of each compound
eye. Antennaspinulose; segment! stout and apparently with or without
1 slenderseta; II stout, longer and with several slender setae; III-VII
slender; III as long as or longer than combined length of IV-VII; IV
shorter than subequal V-VII; distal portion of VII slenderer than
proximal portion; sensoria fringed and with a stub-like sensory seta, or
unfringed and with an elongate sensory seta. Rostrum 3-segmented,
joint between proximal and middle segments sometimes indistinct;
setose.
Thorax. Membranous areas spinulose. Forewing larger than hind-
wing, each wing with a vein (Rs) that approaches the distal margin;
forewing with a diagonal vein (Cu) suggested. Leg 6-segmented; hind
coxa muchlarger than anterior and middle coxae, strongly extended
cephalad on distal side, with a few fine setae; trochanter with 1 strong
seta and sensoria; femur with slender andstoutsetae; tibia with stout,
spine-like setae, some arrangedin a close set row, and with larger, spine-
like apical setae; 1st tarsus longer than 2ndtarsus, with spine-like setae
in 2 rows; 2ndtarsus spinulose, with a few spine-like setae in 2rows and
with 1 dorsal, elongate apical seta; base ofclaw spinulose, with 1 slender
seta on ventral surface; blade-like paronychium longer than claws.
147

Abdomen. Membranous,spinulose, with few to many setae. Disk and


locular pores absent. Area around vasiform orifice slightly enlarged.
Vasiform orifice semicircular to circular; operculum subrectangular, not
filling orifice; lingula spatulate, spinulose, without setae, extending
beyond orifice. Anterior and posterior spiracles conspicuous. Female
with 2 pairs of wax plates. Ovipositor with 3 setae on lateral margins.
Male with 4 pairs of wax plates. Claspers terminatingin a sharptooth,
eachtooth with small, subapicalteeth; aedeagus simple, tubular, curved
dorsad.
DISCUSSION
The pupae of Aleurocerus fall into two major groups based on the
absence or presence of submarginal glands. Glands are absent from
cerifer, coccolobae, colombiae, and petiolicola. A. cerifer represents a
group in which many marginal teeth appearto be double instead of
single, and the submargin is smooth as compared with thatof the other
three species. A. coccolobae lacks visible pale areas in the marginal
ridges and is entirely distinct from colombiae and petiolicola, both of
which have conspicuous pale areasin the marginalridges. The remain-
ing species, chiclensis, flavomarginatus, luxuriosus, musae, palmae,
and tumidosus have submarginal glands. In flavomarginatus and pal-
mae the glands are somewhatconical, and in the other species they are
square or rectangular. In chiclensis and musae submarginaltransverse
ridges with a tooth at the distal end alternate with ridges without a
tooth. In luxuriosus and tumidosusall submarginal transverse ridges
havea tooth at the distal end.

KEY TO STAGES OF IMMATURE ALEUROCERUS


1. Without submedian cephalic and mesothoracic setae; submedian
metathoracic setae elongate and slender; 8 segments visible in
median area of abdomen. .......... errr er eet riTe 2
— With submedian cephalic and mesothoracic setae, these and subme-
dian metathoracic setae short and dagger- or nail-shaped; 7 seg-
ments visible in median area of abdomen. 3
2. Legs with distal margin curved and longer than proximal! margin;
molting sutures present. ...............6.08 fourth stage (pupa)
— Legswith distal and proximal margins evenly curved and subequal
in length; molting sutures absent. .................- third stage
3. Legs somewhatbulb-shaped, notdistinctly jointed, terminating in a
disk; anal area with a bifid horn; eighth abdominalsetae elongate,
dissimilar in shape andsize to dagger-shaped cephalothoracicsetae;
submarginal ridge usually visible; median setae absent anterior to
mouthparts, ........ 06006 c cece eee eee eee eee e es secondstage
— Legs normalleg-shaped,distinctly jointed, terminatingin a fleshy
appendage;anal area withouta bifid horn; eighth abdominalsetae
short, similar in shape and size to nail-shaped cephalothoracic
148

setae; submarginalridge not visible; median setae present anterior


to mouth parts. 2.2.0.0... 0c cee eee c cee eee eee ee eee first stage

Key To PUPAE OF ALEUROCERUS

. Marginalteeth short, broadly curved, some notched and appearing


double; submarginaltransverse ridges and furrowsindistinct, with
needle-like spinules near marginal ridges; posterior 4 pairs of sub-
marginal setae each 70-80, long; horned process unusually large,
strongly projected; vasiform orifice located at or near midlength of
horned process. .......... 000 e cece cece eee ee eee teen eens cerifer
Marginalteeth longer, narrowly to broadly curved, not notched and
appearing double; submarginal transverse ridges and furrowsdis-
tinct, without needle-like spinules near marginal ridges; posterior 4
pairs of submarginalsetae each less than 614 long; horned process
smaller, less strongly projected; vasiform orifice located in or nearer
base of horned process except in tumidosus....
. Submarginalglands absent.
Submarginal glandspresent. .
. Margin reddish in unbleached, paleiin bleachedinsects, contrasting
wo

sharply with the color ofthe body; marginal teeth nearly contiguous,
thoracic tracheal pore area teeth similar to other teeth; marginal
ridges without pale areas; submarginal transverse ridges strong,
each usually with 2 rowsof stout, blunt spinules and with a strong
tooth at distal end; rachis with comparatively few spinules, 3-4 rows
located anteriorly at endsof each of abdominal segments II-VII but
absent posteriorly from these segments. ............ coccolobae
Margin black in unbleached,dark in lightly bleached insects, sim-
ilar in color to the body; marginal teeth '4 to the width of a tooth
apart, thoracic tracheal pore area teeth narrower than otherteeth;
marginal ridges with conspicuous pale areas; submarginal trans-
verse ridges less strong, ridges without spinules or with a few small
ones, with a weaktooth at distal ends; rachis with numerousspin-
ules, located around ends and in submedianareaof each of abdomi-
nal segments II-VI or TI-VIL. ow... cece eee eee ee 4
. AbdominalsegmentVIIIwith a furrow posteriorto the 7th abdomi-
nal suture and with an inverted, crescent-shaped pocket anterior to
the vasiform orifice; marginalridges each with a flask-shaped pale
area; transverse ridges extending across % width of submargin;
mesothorax without submedian spinules on dorsal disk ..........
cee eee eee e eect eee en ees colombiae
Abdominal segment VIII without a furrow posterior to the 7th
abdominalsuture and withoutan inverted, crescent-shaped pocket
anterior to the vasiform orifice; marginal ridges each with a pear-
shaped pale area; transverse ridges extending across ¥ width of
submargin; mesothorax with submedian spinules on dorsal disk.
nn petiolicola
149

5. Margin yellowish or reddishin unbleached, pale in bleached insects,


contrasting sharply with the color of the body; submarginalglands
narrowlyconical to thimble-shaped,their apices narrowlyor broadly
curved andtheir lateral margins weakly to stongly convergent ...
teen cease enna een mee ea tesa sana teeta tence deen ee erat eenaeseeees 6
— Margin black or dark brown in unbleached, darkin lightly bleached
insects, similar in color to the body; submarginal glands nearly
squareor rectangular, their apices truncate andtheir lateral mar-
gins parallel ... .7
6. Submarginal glands narrowly conical, their margins strongly con-
vergent, the largest ones smaller than closest marginalteeth around
entire body; ends of transverse molting suture opposite meso-
metathoracic suture ........ 0.0. c cece eee ee ene flavomarginatus
— Submarginal glands varying from broadly conical with margins
moderately convergent and apices moderately curved to thimble-
shaped with lateral margins nearly parallel and apices broadly
curved, the largest ones larger than closest marginalteeth except at
ends of body; ends of transverse molting suture opposite its mid-
point .. palmae
7. Submarginal transverse ridges each with rows of spinules; alter-
nateridges each witha strongtoothatdistal end; intervenient ridges
without teeth and centers of distal ends opposite spaces between
glands 22... eee eeeet eee eee n een neces 8
— Submarginaltransverse ridges each with 2-3 rows of spinules and a
strongtoothatdistal end; intervenient ridges absent andcenters of
distal ends of furrows opposite spaces between glands... .9
8. Submarginal glands with proximalends of adjacent glands joined
and appearingscalloped, their lateral margins straight; spinules
absent from rachis on abdominal segmentI; basesof caudalsetaein
subdorsal furrow ..........- 0000.20 cece cece eee e eens chiclensis
— Submarginal glands with proximalendsnotjoined and not appear-:
ing scalloped, appearing instead to blend into transverse ridges,
their lateral margins notched; 5-7 spinules at ends of rachis on
abdominal segmentI; bases of caudal setae cephalad of subdorsal
furrow underneath horned process ............0..000seeee musae
9. Tracheal pore area teeth as wide as other marginal teeth; submar-
ginal transverse furrows narrow at proximal ends but as wide as
transverse ridges at distal ends; disk and locular pores compara-
tively sparse on dorsal disk, no more than5 pairs on any abdominal
SESMENE ©... eee eee eee ee cent eee eee ee luxuriosus
— Tracheal pore area teeth narrower than other marginalteeth; sub-
marginal transverse furrows of uniform width and much narrower
thantransverse ridges; disk and locular pores comparatively numer-
ous on dorsal disk, 8-16 pairs on each of abdominal segments
TIAVI o.oo e eee eee eee e eects tumidosus
150

KEY TO THIRD-STAGE LARVAE OF ALEUROCERUS


Third-stage larvae of coccolobae and tumidosus are unknown.
. Marginalteeth broadly curved, many notched and appearing dou-
ble; submarginaltransverse ridges weak or notvisible; posterior 4
pairs of submarginal setae each 48-56y long ............- cerifer
Marginalteeth broadly curved or not, but not notched and appear-
ing double; submarginal transverse ridges weak to moderately
strong; posterior 4 pairs of submarginal setae each less than 25u
JONG oo ccc cece ee cee cnet een been eeeeeeeeuee 2
. Posterior 4 pairs of submarginal setae each 124 or more long ... 3
Posterior 4 pairs of submarginal setae each not more than 12y
TONG cece teen ee cee eee ete e eee been ence 4
. Posterior 4 pairs of submarginal setae each 12y long; abdominal
segmentVIII with a weak furrow posterior to 7th abdominalsuture;
spinules absent posteriorly from median area of rachis on each of
abdominal segments ITI-VI ..............2-..00200255 coiombiae
Posterior 4 pairs of submarginal setae each 20 long; abdominal
segment VIII without a furrow posterior to 7th abdominal suture;
spinules present posteriorly in median area of rachis on each of
abdominal segments IJI-VI ................0ee eee petiolicola
. Spinules present anteriorly but absentposteriorly at endsof rachis
on each of abdominal segments II-VI .............. luxuriosus
Spinules present anteriorly and posteriorly at ends of rachis on each
of abdominal segments II-VI ..............022.0 0.0 e eee eee 5
. Posterior 4 pairs of submarginal setae each 8-10, long; marginal
teeth wider than long ......... 00.0. e cee ce ence eee chiclensis
Posterior 4 pairs of submarginal setae each less than 8 or 1 pair
more than 10z long; marginal teeth wider than long or longer than
Wid 21. cece cence ete eee eet e ee teeeee 6
. Marginal teeth longer than wide, their lateral marginsnearly paral-
lel ++. musae
Marginal eir lateral margins
CLT16 7
. Marginal teeth nearly contiguous; submarginal transverse ridges
and furrowsnotvisible or faintly suggested near margin; 4th-7th
pairs of submarginalsetae each 2-4, long; bases of caudalsetae at
distal edge of submargin ..................... flavomarginatus
Marginalteeth ‘4-4 width of a tooth apart; submarginal transverse
ridges and furrowsclearly visible across most of submargin;4th and
5th pairs of submarginalsetae each 10-12, long, 6th and 7th pairs
each 6-8u; bases of caudal setae at midwidth of submargin ..
webbed e cece eee eee eee eee e eee e tee ee a eee eet e nett eee ce ee pai
151

Aleurocerus cerifer (Sampson and Drews)


(Fig. 2)
Uraleyrodes ceriferus Sampson and Drews, 1941:180-181; Sampson, 1943:220.
Aleurocerus ceriferus (Sampson and Drews), Mound and Halsey, 1978:27
Living on the lower surface of leaves.

Pupa
Description. Nearly oval in outline, narrowed at anterior end, 1-1.50
mm long, 0.75-1 mm wide.
Margin. Yellowishin unbleached, pale in bleached specimens. Anterior
setae 40-48, long, bases 18-24 teeth apart; posterior 48-60, bases 32-36
teeth apart. Teeth 1-3 times wider than long, apices broadly curved,
somenotched andappearing double, appearing as globules in the body
margin when proximal margins arevisible (focus on ventral surface),
contiguousto 2 width of a tooth apart; thoracic trachealpore area teeth
similar to other teeth. Ridges 3-4 times lengthofteeth, an oval pale area
sometimes visible, each ridge with a minute pore near midlength. Fur-
rows each with a minute pore neardistal end.
Submargin. Glands absent. Transverse ridges and furrowsvisible on
distal 2 of submargin; proximal 6/7 smooth andwithout spinules, distal
1/7 withfaint, etch-like lines and needle-like spinules. Disk pores located
irregularly fromdistal 3rd to midline of submargin, 5-7 on cephalic and
2 or 3 pairs on each other body segment. Locularpores in an irregular
row proximadof and as numerousasdisk pores. Anterior pair of setae
6-8 long, 2nd and 3rd pairs each 8-12y, posterior 4 pairs each
70-80p.
Dorsaldisk. Pairs of both disk and locular pores located about as
follows: Cephalic segment | submedian, 2-4 subdorsal; prothorax 1
submedian; mesothorax 2 or 3 submedian; metathorax 2 submedian, 2
subdorsal; abdominal segment I 0 or 1 submedian; II 3 or 4 submedian,
7-10 subdorsal; III 4-6 submedian, 4-6 subdorsal; IV 4 or 5 submedian,
4-6 subdorsal; V 5 submedian, 2-4 subdorsal; VI 3-6 submedian; VII 2
submedian,2 or 3 subdorsal; VIII | or 2 submedian near basal arms of
horned process, 1 or 2 submedian near bases of caudalsetae, 0 or | in
subdorsal furrow. Spinules located anteriorly just proximad of ends of
rachis as follows: Abdominal segmentII 0 or 2; III 2 rows; [V-VI each 2
rows; VII 2 rows. Transverse molting suture terminating nearly oppo-
site its midpoint. Metathoracic setae 120-140u long; eighth abdominal
300-350x, bases on baseof horned process caudad ofbasal arms; caudal
300-350, bases cephalad of subdorsal furrow underneath horned pro-
cess. Horned processvery strongly projected,its basal armslarge. Vasi-
form orifice located at or near midlength of horned process, about 504
wide; operculum about 24, long, 48u wide.
152

Fig. 2. Aleurocerus cerifer.A, Parasitized pupa, paratype. B, Third stage. C, Second


stage. D,First stage. E, Section of margin and submargin ofpupa.F,Section ofdorsum of
pupa showing hornedprocess turned cephalad.
153

Third-stage larva
Description. Broadly oval in outline.
Margin. Teeth 1-2 times wider than long, apices broadly curved or
truncate, many notched and appearing double; thoracic tracheal pore
area teeth similarto other teeth. Ridges 3 times length ofteeth, each with
an oval pale area.
Submargin.Transverse ridges and furrows weakornotvisible. Both
disk and locular pores in 3 pairs on cephalic and in | pair on each other
body segment. Anterior 3 pairs of setae each 2u long, posterior 4 each
48-56p.
Dorsaldisk. Both disk and locular poresin 1 submedian and1 subdor-
sal pair on each body segment. Spinules absent. Metathoracic setae
120-140, long; eighth abdominal250-300,, bases at ends of basal arms
of horned process; caudal 240-260y, bases at midwidth of submargin.
Horned process strongly projected, its basal arms large.

Second-stage larva
Description. Oval in outline.
Margin. Teeth barely visible, some notched and appearing double.
Submargin. Transverseridges and furrows notvisible. Disk and locu-
lar pores absent. Anterior 3 pairs of setae each 2u long, posterior 4 each
45u.
Dorsaldisk. Disk and locularpores absent. Spinules absent. Cephalic,
mesothoracic, and metathoracic setae dagger-shaped, each about 40u
long; eighth abdominalsetae slender, about 200, bases near ends of
basal armsof horned process;caudal slender, about 200u,bases on distal
edge of submargin. Horned process strongly projected, its basal arms
small.

First-stage larva
Description. Elongate ovoid in outline, narrowed anteriorly.
Submarginalsetaeof later immaturestages located on margin, each
pair 8 long. Submedian setae nail-shaped; cephalic 36-40u long; meso-
thoracic and metathoracic each 40-44u; eighth abdominal 32,, bases
just cephalad of basal arms ofprojected area nearly opposite anterior
margin of vasiform orifice; caudal slender, broken in specimens ex-
amined, bases on body margin.

Specimens examined
Paratypes, on woodyvine, State of Nayarit, Mexico, XI-1925, Ferris
No. 10, lent by W.W. Sampson, 3 mounted pupae. Several unmounted,
154

and mounted specimensas follows: 5 pupae (+fragments), 5 third stage


(+ fragment), 2 secondstage,| first stage (+ fragments), | adult male and
1 adult female (removed from pupae) on Cissus sp. (Vitaceae), Lancetilla
Valley, Dept. Atlantida, Honduras, 12-I-1928, P.C. Standley.

Notes
Pupae of Aleurocerus cerifer are distinctive in having broadly
curved, notched, marginalteeth, an unusually large horned process, and
in lacking prominent submarginal transverse ridges and furrows. A.
ceriferis strikingly distinct from other described species of Aleurocerus.

Aleurocerus chiclensis, new species


(Fig. 3)
Living on the lower surfaceof leaves.

Pupa :|
|
Description. Ovoid in outline, 0.90-1.20 mm long, 0.60-0.75 mm wide. {
|
Margin. Black in unbleached, dark in bleached specimens. Anterior a
setae 24, long,posterior 32u, basesof each pair 16-20 teeth apart. Teeth
slightly longer than wide, apices moderately curved, 4 width of a tooth
apart; thoracic tracheal pore area teeth narrower than other teeth
Ridges about % length of teeth, apparently without pale areas. Furrows
each with a minute pore at proximalend.
Submargin. Glandsaslong as wideto slightly longer than wide; trun-
cate at distal ends, proximalendsof adjacent glands joined and appear-
ing scalloped; lateral margins straight, parallel, nearly contiguous; cen-
ters of distal ends opposite marginal furrows. Transverse ridges
extending across '2-% width of space between glands and subdorsal
furrow;each ridge with 2 rowsof stout, blunt spinules; alternate ridges
with a strongtooth at distal ends pointed at centers of glands; interveni-
ent ridges without a tooth, distal ends opposite spaces between glands.
Transverse furrows narrowerthan ridges, without or with a few slender
spinules. Disk pores located irregularly along proximal endsof trans-
verse ridges, 6-8 pairs on cephalic and 2 or 3 pairs on each other body
segment. Locular pores proximad of and apparently slightly less
numerous than disk pores. Anterior 3 pairs of setae each 4-6u long,
posterior 4 each 20-24y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows. Cephalic segment 1 submedian, | or 2 subdorsal; prothorax 1
submedian,1 subdorsal; mesothorax 1 submedian, 1 or 2 subdorsal;
metathorax 0 or 1 submedian, 2 or 3 subdorsal; abdominal segment I 1
submedian; II 0 or 1 submedian,1 or 2 subdorsal; III 2 submedian,1 or 2
subdorsal; IV 2 submedian,1 subdorsal; V 1 submedian, 1 subdorsal; VI
155

c
Fig. 3. Aleurocerus chiclensis. A, Pupa, holot
ype.B, , Section of margin, submargin,
broken dorsal disk, and venter. C,Third stage.
ir
156

1 submedian,1 subdorsal; VII 1 submedian, 1 subdorsal; VIII 1 subdor-


sal. Spinules located just proximadofendsof rachis as follows: Abdom-
inal segments II-VII each 2-3 rows anteriorly; II-V each also 1-7 spin-
ules posteriorly. Transverse molting suture terminating opposite meso-
metathoracic suture. Metathoracic setae 140, long; eighth abdominal
broken in specimens examined, bases adjacent to basal arms of horned
process; caudal 120y, bases in subdorsal furrow. Horned process weakly
projected, its basal arms small. Vasiform orifice located in base of
horned process, about 36u wide; operculum 12y long, 32 wide.

Third-stage larva
Description. Broadly oval in outline.
Margin. Teeth slightly wider than long, apices curved broadly; thoracic
trachealpore area teeth narrower thanotherteeth. Ridges 4 longer than
teeth, each with a subcircularpale area.
Submargin. Transverseridges and furrows extending across submar-
gin; someridges witha tooth atdistal end. Both disk and locular pores in
2 pairs on each of cephalic and prothoracic segments andin 1 pair on
each other body segment. Anterior3 pairsof setae each 1-2, long, poster-
ior 4 each 8-10.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment 1 submedian, 1 subdorsal; mesothorax and
metathorax each 1 submedian, 1 subdorsal; abdominal segment I 1
submedian; III-VIII each 1 subdorsal. Spinules located on ends of
rachis as follows: Abdominal segmentI 0-4 anteriorly; I1 3-7 anteriorly,
1 or 2 posteriorly; III 9-13 in 2 rows anteriorly, 2-4 spinules posteriorly;
IV-VI each 10-12 in 2 rows anteriorly, 5-10 spinules around ends and
posteriorly; VII 5-8 anteriorly, 0 or 1 posteriorly. Metathoracic setae
broken in specimens examined but more than 160, long; eighth abdomi-
nal broken but more than 80u, bases near ends of basal armsof horned
process; caudal 145y, bases ondistal submargin. Horned process moder-
ately projected,its basal armslarge.

Second-stage larva
Description. Ovoid in outline.
Margin.Teeth slightly wider than long, curved from base.
Submargin. Transverse ridges and furrowsbarelyor not visible. Disk
andlocular pores absent. Anterior6 pairs of setae each 2y long, posterior
pair 4u.
Dorsaldisk. Both disk and locularpores in 1 subdorsalpair on each of
mesothorax, metathorax, and abdominal segments ITI-VIII. Spinules
absent. Metathoracic setae dagger-shaped, 40u long; other setae and
other structures absent from specimen examined.
157

Specimens examined
Holotype, on a chicle producingtree [perhaps Manilkara Zapota (L.)
Van Royen (Sapotaceae)], Mexico, intercepted at quarantine at Browns-
ville, Texas, 30-VIII-1935, G.A. Pfaffman. Paratypes, a few unmounted,
and 5 mounted pupae(+ fragments), 3 third stage, and 1 fragmentary
second stage, taken with the holotype.

Notes
Pupae of Aleurocerus chiclensis differ from those of the closely
related A. musae in having the lateral margins of the submarginal
glandsstraight and the proximalendsof adjacent glands joined, thus
giving a scalloped appearance. In musae the lateral margins of the
glands are notched and their proximal ends appearto blend into the
submarginalridges. In chiclensis spinules on the rachis are much less
numerous than in musae, and the basesofthe caudal setae are located in
the subdorsalfurrow insteadof farther cephalad underneath the horned
processasthey are in musae.

Aleurocerus coccolobae, new species


(Fig. 4)
Habit not observed.
Pupa
Description. Elongate oval in outline, 1.25-1.45 mm long, 0.65-0.75
mm wide.
Margin. Reddish in unbleached,pale in bleached specimens. Anterior
setae 28u long, bases 14-16 teeth apart; posterior broken in specimens
examined, bases 26-34 teeth apart. Teeth as wide as longto slightly
wider than long, apices moderately curved, nearly contiguous to 1/5
widthof a tooth apart; thoracic tracheal pore area teeth similarto other
teeth. Ridges 1% times length of teeth, each with an oblong pale area.
Furrowseach with a minute pore near midlength.
Submargin. Glands absent. Transverse ridges strong, irregular, ex-
tending across submargin at anterior end of body, across 2% width of
submargin elsewhere;each ridge with 1 or 2 rowsofstout, blunt spinules
and a strongtoothat distal end pointed at marginalfurrows. Transverse
furrows irregular, narrower than ridges, without spinules. Disk pores
located irregularly near midline of transverse ridges, 8-10 pairs on
cephalic and 3-5 pairs on each other body segment. Locular pores prox-
imadof and apparently less numerousthan disk pores. Anterior 3 pairs
of setae each 4u long, posterior 4 each about 12y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment 3 submedian,1 or 2 subdorsal; prothorax 1
submedian, | or 2 subdorsal; mesothorax 2 submedian, 2 subdorsal;
158

Fig. 4. Aleurocerus coccolobae. A, Pupa, holotype. B, Section of margin, submargin,


anddorsaldisk.

metathorax 1 or 2 submedian, 4 subdorsal; abdominal segmentI 1


submedian, 2 subdorsal; II 2 submedian, 2 subdorsal; III 1 or 2 sub-
median, 5 subdorsal; IV 1-3 submedian, 3-5 subdorsal; V 1-3 submedian,
3 or 4 subdorsal; VI 1-3 submedian,1 subdorsal; VII 1 or 2submedian,1
subdorsal; VIII 1 or 2 submedian, 1 or 2 subdorsal. Spinules located in 3

Alsaei haaes’
or 4 rowsanteriorly just proximadof endsof rachis on each of abdomi-
nal segments II-VII. Transverse molting suture terminating opposite a
point slightly posterior to its midpoint. Metathoracic setae broken in
specimens examined; eighth abdominalbroken in specimens examined
but more than 100z long, bases on or adjacent to basal arms of horned
process; caudal 200, bases cephalad of subdorsal furrow just laterad of
horned process. Horned process moderately projected, its basal arms
small to moderately large. Vasiform orifice located in base of horned
process, about 44, wide; operculum about 16long, 40z wide.

Specimens examined
Holotype, on Coccoloba padiformis Meisn. (Polygonaceae), Colom-
bia, 18-VII-1935, Perez Arbelaez. Paratypes, a few unmounted and 6
mounted pupaetaken with the holotype.
159

Notes
Aleurocerus coccolobae resembles A. colombiae andA.petiolicola in
lacking submarginalglands.It differs from these species conspicuously,
however, in the color of the body margin, the closeness of the marginal
teeth, the prominenceof the submarginalridges and submarginal spin-
ules, and in the comparatively small numberof spinules on therachis.

Aleurocerus colombiae, new species


(Fig. 5)
Coccids Brown in Kempf 1968:401.
Living on stemsandleaf petioles (Brown, 1978, pers. comm.).

Pupa
Description. Broadly oval in outline, 1.40-1.50 mm long, 1.00-1.05
wide.
Margin. Black in unbleached, dark in lightly bleached specimens.
Anterior setae 25-40, long, bases 10-12 teeth apart; posterior 55-60u
bases 20-22 teeth apart. Teeth about 2 times wider than long, apices
moderately to broadly curved, 4 to nearly the width of a tooth apart;
thoracic treacheal pore area teeth narrowerthanotherteeth. Ridges 3-4
times length of teeth, each with a flask-shaped pale area. Furrows each
with a minute pore near proximal end.
Submargin. Glands absent. Transverse ridges irregular, extending
across % width of submargin, rough but without, or with a few spinules;
most ridges with a weak tooth at distal ends pointed at marginal fur-
rows. Transverse furrows irregular, narrower than ridges, without
spinules. Disk pores located in an irregular row at proximal 3rd of
transverse ridges, 7 or 8 pairs on cephalic and 2-4 pairs on each other
body segment. Locular pores proximadofdisk pores and proximal ends
of transverse ridges and apparently as numerous as disk pores. Anterior
3 pairs of setae each 8u long, posterior 4 each 30z.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment 5 or 6 submedian, 3 subdorsal; prothorax 1
submedian, 0 or 1 subdorsal; mesothorax and metathorax each 3 or 4
submedian,3 or 4 subdorsal; abdominal segment 1 or 2submedian, 3 or
4 subdorsal; II 1 submedian, 2-4 subdorsal; III 1 submedian, 1 or 2
subdorsal; IV 2 or 3 submedian, 2 subdorsal; V 1 or 2 submedian, 2
subdorsal; VI 3 or 4 submedian, 1 subdorsal; VII 3 or 4 submedian, 1
subdorsal; VIII 2 or 3 submedian distad of horned process, 1 or 2 sub-
median posterior to horned process proximadof bases of caudal setae, 2
or 3 subdorsal. Spinules unusually numerous,located as follows: Meta-
thorax 6-10 submedian on each half of segment; on rachis, abdominal
segmentI 0-4 anteriorly, 4-8 rows posteriorly; II-VII each 4-6 rows
160

Fig.5. Aleurocerus colombiae. A, Pupa, holotyp


dorsaldisk. C, Third stage. D, Second stage. e. B, Section of margin, submargin, and
161

around endsand 1-3 rows extending across submedian to medianarea;


VII 0-6 spinules anteriorly. Transverse molting suture terminating
opposite meso-metathoracic suture. Metathoracic setae 200u long; eighth
abdominal 220u, baseson or adjacentto basal armsof horned process;
caudal 130, bases in subdorsal furrow. Abdominal segmentVIII witha
subcircular furrow nearly paralleling posterior abdominal suture and
an inverted, crescent-shaped pocket anterior to vasiform orifice. Horned
process weakly projected, its basal arms small. Vasiform orifice located
in base of horned process, about 604 wide; operculum about 20u long, 454
wide.

Third-stage larva
Description. Broadly oval in outline.
Margin. Teeth wider than long, apices broadly curved; thoracic tra-
chealpore area teeth narrowerthanotherteeth. Ridges 4 times length of
teeth, each with an elongate pale area.
Submargin.Transverse ridges and furrows extending across submar-
gin; ridges occasionally with a weak tooth pointed at distal ends. Both
disk and locular pores in 3 pairs on cephalic andin | or occasionally 2
pairs on each other body segment. Anterior3 pairs of setae each 4 long,
posterior 4 each 12y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment 2 or 3 submedian; prothorax 1 or 2 submed-
ian, 1 subdorsal; mesothorax 0-2 submedian, 1 subdorsal; metathorax
1-3 submedian,1 subdorsal; abdominal segmentI 1 submedian; II-VII
each 1 submedian, 1 subdorsal; VIII 2 subdorsal. Spinules unusually
numerous, located on rachis as follows: Abdominal segmentI 0-4 anter-
iorly, 2-5 posteriorly; II-VI each with 12-20 in 2 or 3 irregular rows
anteriorly and across submedian to medianarea and aroundends, 4-8
spinulesposteriorly; VII 10-15 anteriorly, 2-5 posteriorly. Metathoracic
setae 130x long; eighth abdominal 160u, bases at ends of basal armsof
horned process; caudal 120u, bases on distal submargin. Abdominal
segment VIII with a weak furrow posterior to 7th abdominal suture.
Horned process moderately projected, its basal arms rather small.

Second-stage larva
Description. Broadly ovalin outline.
Margin. Teeth wider than long,apices broadly curved.
Submargin.Transverseridges andfurrowsfaintly visible on distal %
of submargin.Both disk and locularporesin 1 pair on cephalic segment.
Anterior3 pairs of setae each 2u long, posterior 4 each 4p.
Dorsal disk. Both disk and locular pores in 1 subdorsal pair on each
body segment except 2 pairs on abdominal segment VIII. Spinules
162

absent. Cephalic, mesothoracic, and metathoracic setae dagger-shaped,


each 32-48long;eighth abdominal slender, 1204 long, bases near ends of
basal armsof hornedprocess; caudal slender, 120u, bases on distal sub-
margin. Horned process weakly projected, its basal arms small.

Specimens examined
Holotype, on the upper stem of an aroid plant, possibly Dieffen-
bachia sp. (Araceae) (Brown, 1978, pers. comm.), near Anchicaya, 3.2
km E. Rio Aguaclara on old Cali Road, Colombia, 19-III-1967, R.B.
Root and W.L. Brown. Paratypes on stems andleaf petioles, a few
unmounted and 6 mountedpupae, 11 third stage, 5 secondstage, taken
with the holotype. Paratypes in BMNH and FSCA.

Notes
In discussionof the ant Octostruma inca Brown and Kempf, Kempf
(1968) quoted Brown asstating, “One group of workers was found tend-
ing coccids on an aroid stem.” Brown (1978, pers. comm.) wrote that he
had not examined theinsects carefully.
Aleurocerus colombiae and A. petiolicola are unusual in living on
stemsandleafpetioles instead of on leaves. The two species are closely
related. Pupae of A. colombiae are distinctive in having a well defined
furrow and an inverted, crescent-shaped pocket on the eighth abdomi-
nal segment.A.petiolicola lacks these characteristics.

Aleurocerus flavomarginatus Bondar


(Fig. 6)
Aleurocerus flavomarginatus Bondar, 1923: 161-162; Mound and Halsey, 1978:27.
Aleurocerus spp. Mound and Halsey, 1978:27, in part.
Living on the lower surfaceof leaves.

Pupa
Description. Ovoid in outline, 1-1.25 mm long,0.75-0.95 mm wide.
Margin.Reddish or yellowish in unbleached, pale in bleached speci-
mens. Anterior setae 24u long, bases 14-16 teeth apart; posterior 32u,
bases 16-18 teeth apart. Teeth slightly wider than long, apices brodly
curved, 4-4 width of a tooth apart; thoracic tracheal pore area teeth
narrower thanotherteeth. Ridges extending to proximal ends of spaces
between submarginalglands,2 times length ofteeth, each with an elon-
gate pale area extending into space between submarginal glands.
Furrowseach with a minute pore near midlength.
Submargin. Glands smaller than nearest marginal teeth; narrowly
conical, tapered from broad base to narrowly curved apices, as long as
wide; proximalendsof adjacent glands joined and appearingscalloped;
163

Fig. 6, Aleurocerus flavomarginatus. A, Pupa, syntype. B, C,Sections of margin,


submargin, anddorsal disk. D, Third stage.

apices of glands opposite marginal furrows. Transverse ridges extend-


ing across % width of space between glands and subdorsalfurrow; each
ridge with 2 rowsof stout, blunt spinules; alternate ridges with a strong
tooth at distal ends pointed at centers of glands; intervenient ridges
without a tooth, distal ends opposite scallops between glands. Trans-
verse furrows narrowerthan ridges, with minute spinules. Disk pores
located irregularly just proximad of transverse ridges, 6-8 pairs on
cephalic, 2 or 3 pairs on each thoracic, and1 or 2 pairs on each abdomi-
nal segment. Locular pores located proximadof and as numerousas disk
pores. Anterior 3 pairs of setae each 4long, posterior 4 each 12u.
164

Dorsal]disk. Pairs of both disk and locular pores located about as


follows: Cephalic segment 2 submedian, 1 or 2 subdorsal; prothorax 1
submedian, | or 2 subdorsal; mesothorax 1 submedian, 1-3 subdorsal;
metathorax | submedian,1 subdorsal; abdominal segmentI 1 subdor-
sal; II 1 submedian,1 subdorsal; [II 1 submedian,| or 2 subdorsal; IV 1
or 2 submedian, 1 subdorsal; V 1 or 2 submedian, 1 subdorsal; VI 2
submedian, 1 or 2 subdorsal; VII 1 submedian, 1 subdorsal; VIII 1
subdorsal. Spinules located on ends of rachis as follows: Abdominal
segment I 0-7; II-VII each 3-5 rows anteriorly; II-VI each also 3-7
spinules posteriorly. Transverse molting suture terminating opposite
meso-metathoracic suture. Metathoracic setae broken in specimens
examinedbut more than 120, long; eighth abdominal 200x,baseslater-
ocephaladof vasiform orifice; caudal 200u, bases cephalad of subdorsal
furrow just laterad of horned process. Horned process weakly projected,
basal arms very small or absent. Vasiform orifice located in base of
horned process, 44-50, wide; operculum 18-20, long, 32-40wide.

Third-stage larva
Description. Broadly ovoid in outline.
Margin.Teeth as wide as long,apices broadly curved; thoracic tracheal
pore area teeth narrowerthan other teeth. Ridges ‘4 longer than teeth,
each with an elongate pale area.
Submargin. Transverse ridges and furrowsnotorfaintly visible across
submargin; ridges occasionally with a tooth at distal ends. Both disk
and locular pores apparently in 3 pairs on cephalic andin 1 pair on each
other body segment. Anterior 3 pairs of setae each 1-2y long,posterior 4
each 2-4y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment1 submedian, 1 subdorsal; mesothoracic and
metathorax each 1 submedian, 1 subdorsal; abdominal segment I 1
submedian; II-VIII each 1 subdorsal. Spinuleslocated on ends ofrachis
as follows: Abdominanl segmentI or 5 anteriorly; II-VI each 10-13 in 2
or 3 rows anteriorly, 3-7 spinules posteriorly; VII 5-8 anteriorly, 3-5
posteriorly. Metathoracic setae broken in specimens examined but more
than 120y long; eighth abdominal about 200,, bases near endsof basal
armsof hornedprocess; caudal about 240u,bases on distal submargin.
Horned process weakly projected, its basal arms small.

Second-stage larva
Description. Broadly ovoid in outline.
Margin.Teeth as long as wide, apices broadly curved.
Submargin. Transverse ridges and furrowsvisible on distal /-% of
submargin. Both disk and locular pores in 1 pair on cephalic segment.
165

Spinules absent. Anterior 6 pairs of setae each 1-2, long,posterior pair


4u. .
Dorsaldisk. Both disk and locular pores in 1 subdorsal pair on each
body segment. Spinules, 1 submedian located anteriorly or posteriorly
on each of abdominal segments IV-VI. Cephalic, mesothoracic, and
metathoracic setae dagger-shaped, each 32u long; eighth abdominal
slender, 200u, bases distad of basal arms of horned process; caudal
slender, 100u, bases on distal edge of submargin. Horned process very
weakly projected, its basal arms very small.

Specimens examined.
Syntypes, on unidentified plants of the “matta,” Belmonte [Bahia],
Brazil, XII-1922, G. Bondar, a few unmounted and 3 mounted pupae (+
fragments), | third stage. Several unmounted, and mounted specimens
as follows: 21 pupae, 5 third stage on coconut (Cocossp.) (Palmae), Erin,
Trinidad, 22-I-1919, C.B. Williams (BMNH); 10 pupae(+ fragments), 7
third stage, 3 second stage on coconut (Cocos sp.), St. Augustine, Trin-
idad, 1958-59, L.A. Mound; 6 pupae on Musasp. (Musaceae), St. Augus-
tine, Trinidad, 1958-1959, L.A. Mound.
Notes
The specimenscollected by C.B. Williams were recorded as Aleuroce-
rus sp. by Mound and Halsey (1978).
Bondar (1923) stated (translation) “. .. When on theleaf, the insect
produces a white, abundant, curly wax forminga pile on the top of the
insect and coveringit, but it does not make a tube asin the preceeding
species” /tumidosus).
Aleurocerus flavomarginatus is closely related to A. palmae; their
relationship is discussed underthe latter species.

Aleurocerus luxuriosus Bondar


(Fig. 7)
Aleurocerus luxuriosus Bondar, 1923:157-159; Costa Lima, 1928 (1927):100, 1936:151;
Drews and Sampson, 1940:90; Sampson, 1943:213; Silva etal., 1968:112; Mound and
Halsey, 1978:27.
Living on the lower surfaceof leaves.

Pupa
Description. Oval in outline, 1.10-1.50 mm long, 0.75-0.95 mm wide.
Margin.Black in unbleached, dark in bleached specimens. Anterior
setae 36u long, bases 18-20 teeth apart; posterior broken in specimens
examined,bases 28-30 teeth apart. Teeth as long as wide, apices broadly
curved, 1/6 width of a tooth apart;thoracic treacheal pore area teeth
similarto other teeth. Ridges % length of teeth, apparently without pale
areas. Furrows each with a minute pore near proximalend.
166

Fig. 7. Aleurocerus luxuriosus. A, Pupa, syntype. B, Section of dorsal disk. C, Second


stage. D, Third stage. E, F, Sections of margin, submargin, and dorsaldisk of pupa
167

Submargin. Glands as long as wide;truncate at distal ends, proximal


endsof adjacent glands joined and appearingscalloped;lateral margins
notched,parallel, nearly contiguous; centerof distal ends opposite mar-
ginal furrows. Transverse ridges irregular, extending across % width of
space between glands and subdorsalfurrow; each ridge with 2-3 rows of
stout spinules and a strong tooth at distal end pointed at centers of
glands. Transverse furrowsirregular, as wide as ridges at distal ends
but narrower at proximal ends, with stout spinules, their distal ends
opposite spaces between glands. Disk pores located irregularly along
proximal ends of transverse ridges and a few proximadofridges,
number on cephalic segment not determined, 2-4 pairs on each other
body segment. Locular pores proximad of and as numerous as disk
pores, some near subdorsal furrow. Anterior 3 pairs of setae each 5-6
jong, 4th and 5th each 32-36y,6th and 7th each 12u.
Dorsal disk. Pairs of both disk and locular pores located as follows:
Cephalic segment3 or 4 subdorsal; prothorax 3 submedian; mesothorax
and metathorax each 1 or 2 submedian, 2 or 3 subdorsal; abdominal
segmentI 1 submedian, 2 or 3 subdorsal; II 1 submedian, 2-4 subdorsal;
III 1 submedian, 2 or 3 subdorsal; IV-VII each 1 submedian, 1 or 2
subdorsal; VIII 2 or 3 subdorsal caudad of horned process. Spinules
located anteriorly on endsof rachis as follows: Abdominal segmentII
3-4 rows;III-VII each 5-6 rows. Transverse molting suture terminating
nearly opposite its midpoint. Metathoracic setae broken in specimens
examined; eighth abdominal 240 long, bases on or adjacent to basal
arms of horned process; caudal broken in specimens examined, bases
cephalad of subdorsal furrow underneathor just laterad of horned pro-
cess. Horned process rather strongly projected, its basal arms small.
Vasiform orifice located in base ofhornedprocess, about 60wide;oper-
culum 20g long, 32u wide.
Third-stage larva
Description. Broadly ovalin outline.
Margin. Teeth wider than long, apices broadly curved to subtruncate,
marginsparallel; thoracic tracheal pore area teeth similar to other teeth.
Ridges 3 times length of teeth, each with a poorly defined pale area.
Submargin.Transverse ridges and furrows extending across submar-
gin; ridges rarely with a weak tooth at distal ends. Both disk and locular
poresin 3 pairs on cephalic, in 2 pairs on eachthoracic, and in 1 pair on
each abdominal! segment. Anterior 3 pairs of setae each 1-2, long, pos-
terior 4 each 4-5y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment1 submedian, 1 or 2 subdorsal; prothorax 0 or
1 subdorsal; mesothorax and metathorax each 1 submedian, 1 or 2
subdorsal; abdominal segment I 1 submedian;II-VII each | or 2 subdor-
sal; VIII 1 submedian. Spinules located anteriorly on endsor rachis as
168

follows: Abdominal segmentII 0-4; III-VII each 16-26in 3 or 4 rows.


Metathoracic setae about 155y long; eighth abdominalbroken in speci-
mens examined but more than 220u, bases proximadof ends of basal
armsin 1 specimen, laterocephalad of vasiform orifice in 1 specimen;
caudal broken in specimens examined, bases on proximal submargin.
Hornedprocess strongly projected, its basal armslarge in 1 specimen,
absentin 1 specimen.
Second-stage larva
Description. Ovalin outline.
Margin. Teeth wider than long, apices subtruncate.
Submargin. Transverse ridges and furrows visible across % of sub-
margin. Both disk and locular pores in 1 pair on each body segment.
Anterior 3 pairs of setae each 2 long, posterior 4 each 4.
Dorsal disk. Pairs of both disk and locular pores located as follows:
Cephalic segment 1 submedian; prothorax 0 or 1 submedian; meso-
thorax | submedian; metathorax 1 submedian,1 subdorsal; abdominal
segments III-VIII each 1 subdorsal. Spinules absent. Cephalic, meso-
thoracic, and metathoracic setae dagger-shaped, each 40y long; eighth
abdominal slender, broken in specimens examined, bases near ends of
basal arms of horned process; caudalslender, about 90y, bases on distal
edge of submargin. Horned process strongly projected, its basal arms
fairly large.
First-stage larva
Description. Elliptical in outline.
Submarginalsetae of later immaturestages located on margin, each
pair 2u long. Submediansetae nail-shaped; cephalic, mesothoracic, and
metathoracic each 44, long; eighth abdominal nail-shaped, broken in
specimens examined, bases just cephalad of basal arms of projected
area; caudal slender, broken in specimens examined, bases on body
margin.
Specimens examined
Syntypes, on unidentified Myrtaceae, Bahia, Brazil, Jacy Bondar; a
few unmounted and 12 mounted pupae(+ fragments), 2 third stage, 2
secondstage, 2 first stage, 2 adult females.

Notes
Bondar (1923) wrote (translation) “. .. When on theleaf it is sur-
rounded by white rather soft wax issuing from the margin androlled
dorsad, but leaving it somewhat exposed. The cast skins of previous
molts are frequently stuck in the wax.” He described the adult male and
femaleof A. luxuriosus.
Pupaeof Aleurocerus luxuriosus have an unusually large number of
spinules on the rachis. The speciesis closely related to A. tumidosus.

é
169

Aleurocerus musae, new species _


(Fig. 8)
Living on the lower surfaceof leaves.
Pupa
Description. Ovoid in outline, 1-1.50 mm long, 0.60-0.90 mm wide.
Margin. Black in unbleached, dark in bleached specimens. Anterior
setac 28-32. long, bases 22-24teeth apart; posterior 38-40y, bases 26-28
teeth apart. Teeth as long as wide, apices moderately curved, 4-4 width
of a tooth apart; thoracic tracheal pore area teeth narrower than other
teeth. Ridges % length of teeth, apparently without a pale area. Furrows
each with a minute pore at proximal end.
Submargin.Glandsas longas wideto slightly longer than wide; trun-
cate at distal ends, proximal ends appearingtoblend into submarginal
ridges; lateral margins notched,parallel, nearly continguous, center of
distal ends opposite marginal furrows. Transverse ridges extending
across 2 width of space between glands and subdorsal furrow; each
ridge with 2 rowsofblunt spinules; alternate ridges with a strong tooth
at distal endspointed at centersof glands;intervenientridges without a
tooth, distal ends opposite spaces between glands. Transverse furrows
much narrower than ridges, with a few slender spinules. Disk pores
located irregularly along proximalendsoftransverse ridges, 6 or 7 pairs
on cephalic, 2 or 3 on each thoracic, and 1 or 2 on each abdominal
segment. Locular pores proximad of and apparently as numerous as
disk pores. Anterior 3 pairs of setae each 4-6y long, posterior 4 each
16-20y.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment1-3 submedian, 1 or 2 subdorsal; prothorax 1
or 2 submedian, 1 subdorsal; mesothorax 1 submedian, 1 subdorsal;
metathorax 1 submedian, 1 or 2 subdorsal; abdominal segment I’ 1
submedian; II 1 or 2 submedian, 2 subdorsal; IIT 1 or 2 submedian, 1
subdorsal; IV 2 or 3submedian, 1 or 2 subdorsal; V 2 or 3 submedian, 1 or
2 subdorsal; VI 2 submedian, 0 or 1 subdorsal; VII 1 submedian, 0 or 2
subdorsal; VIII 0 or 1 submedianlaterocephalad of eighth abdominal
setae. Spinules located on endsofrachis as follows: Abdominal segment
15-7 posteriorly; II 3-4 rows anteriorly, 5-7 spinules posteriorly; IIJ-V
each 2-4 rowsanteriorly, 6-10 spinules posteriorly; VI 3 rows anteriorly,
0-4 spinulesposteriorly; VII 3 rows located anteriorly. Transverse molt-
ing suture terminating opposite meso-metathoracic suture. Metathor-
acic setae broken in specimens examined but more than 188, long;
eighth abdominal 210-250, bases on basal armsof horned process;
caudal 200-2504, bases cephalad of subdorsal furrow underneath
horned. Horned process moderately projected, its basal arms small to
large. Vasiform orifice located in base of horned process, 40-48 wide;
operculum 16-20u long, 32-444 wide.
170

Fig. 8. Aleurocerus musae. A, Pupa, paratype. B, Section of margin,


submargin, and
dorsaldisk. C, Third stage. D, Second stage.
171

Third-stage larva
Description. Broadly ovoid in outline.
Margin. Teeth slightly longer than wide, apices moderately curved,
lateral margins nearly parallel; thoracic tracheal pore area teeth nar-
rower than other teeth. Ridges about 2 times length of teeth, each with
anelliptical pale area.
Submargin. Transverse ridges and furrows extending across submar-
gin; ridges with a weaktooth at distal ends. Both disk and locular pores
in 2 or 3 pairs on cephalic segment and in 1 pair on each other body
segment. Anterior3 pairs of setae each 3-4, long,posterior 4 each 4-6.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment 1 submedian, 1 subdorsal; mesothorax 2
submedian, 2 subdorsal; metathorax 1 submedian, 1 subdorsal; abdom-
inal segment I | submedian; II 1 median (not paired), 1 subdorsal;
III-VIII each 0 or 1 median, 1 subdorsal. Spinules located on ends of
rachis as follows: Abdominal segmentII 6-8 in 2 rows anteriorly, 1-3
spinules posteriorly; III 7-11 in 2 rows anteriorly, 2-4 spinules poster-
iorly; TV 8-14 in 2 rows anteriorly, 2-6 spinules posteriorly; V 8-14 in 2
rowsanteriorly, 3-5 posteriorly; VI 9-11 in 2 rowsanteriorly, 4-7 spin-
ules posteriorly; VII 5-7 in 2 rows anteriorly. Metathoracicsetae 125p
long; eight abdominal 120y, bases at or proximadof ends ofbasal arms
of horned process; caudal 140, bases at midwidth of submargin.
Horned process moderately projected, its basal arms small to large.

Second-stage larva
Description. Broadly oval in outline.
Margin.Teeth slightly longer than wide, apices moderately curved,
lateral margins nearly parallel.
Submargin.Transverse ridges and furrowsbarelyor notvisible. Disk
and locularpores absent. Anterior3 pairs of setae each 2u long, posterior
4 each4p.
Dorsal disk. Both disk and locularpores in 1 submedianor subdorsal
pair on each body segment. Spinules absent. Cephalic, mesothoracic,
and metathoracic setae dagger-shaped, each 32-36long; eighth ab-
dominalslender, 140, bases near endsof basal armsof hornedprocess;
caudal slender, 75y, bases near distal edge of submargin. Horned
process strongly projected, its basal arms small.

Specimens examined
Holotype, on Musa paradisiaca L. (= M. sapientum L.) (Musaceae),
Guatemala City, Guatemala,intercepted at quarantine at San Antonio,
172

Texas, 25-VII-1965, D. Johnston. Paratypes, a few unmounted and 8


mounted pupae, 4 (+ fragments) third stage, 3 second stage, 2 adult
males, 2 adult females taken with the holotype.

Notes
The pupaeof Aleurocerus musaeare the only ones studied in which
the proximalendsofthe submarginalglandsareseparate from adjacent
glands andblendinto the transverse ridges instead of being joined and
having a scalloped appearance. The species is closely related to A.
chiclensis.

Aleurocerus palmae, new species


(Fig. 9)
Living on the lower surfaceof leaves.

Pupa
Description. Ovoid in outline, 0.95-1.50 mm long, 0.55-1 mm wide.
Margin.Reddish in unbleached, pale in bleached specimens. Anterior
and posterior setae each 24-28, long, bases of each pair 16-24 teeth
apart. Teeth as wideas long,apices broadly curved, 4-4 widthof a tooth
apart; thoracic tracheal pore area teeth narrower than other teeth.
Ridges extending to proximal ends of spaces between submarginal
glands, 2 times length of teeth, each with a flask-shaped clear area
extendinginto space between submarginalglands. Furrows each with a
minute pore near midlength.
Submargin. Glandslarger than nearest marginal teeth except some-
|
times nolarger at anterior, and smaller than teeth at posterior end of
body; varying in shape from broadly conical with margins irregular and
slightly convergent and apices moderately curved to thimble-shaped
with lateral margins nearly parallel; proximal endsof adjacent glands
joined and appearing scalloped; apices of glands opposite marginal
furrows. Transverse ridges extending across 2 width of space between
glands and subdorsal furrow; each ridge with 2 rowsof stout, blunt
spinules; alternate ridges with a strong tooth at distal ends pointed
at centers of glands; intervenient ridges without a tooth, distal ends
opposite scallops between glands. Transverse furrows narrower than
ridges, with minute spinules. Disk pores located just proximadoftrans-
verse ridges, 5-9 pairs on cephalic, 2-4 pairs on each thoracic, and 2 or 3
pairs on each abdominal segment. Locular pores proximadof and as
numerousas disk pores. Anteriorpairof setae 4u long, 2nd 8y,3rd 16y,
4th and 5th each 20-28, 6th and 7th each 12-16.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment | or 2submedian, | or 2 subdorsal; prothorax
1 submedian, 1 subdorsal; mesothorax 1 or 2 submedian,1 or 2 subdor-
Fig. 9. Aleurocerus paimae. A, Pu pa, holotype. B, Third stage. C,
Section of margin,
submargin, and dorsaldisk of pupa. D, First
stage. E, Second stage. F, Third stage with
white, fibrous, waxy covering,
rT
174

sal; metathorax 1 or 2 submedian, 2 subdorsal; abdominal segmentI 1


submedian, 1 or 2 subdorsal; II 1 or 2 submedian, 1 subdorsal; III 2 or 3
submedian, | subdorsal; [V2-4 submedian, 1 subdorsal; V 2-5 submed-
ian, 1 subdorsal; VI 1-4 submedian, | subdorsal; VII 1 or 2submedian,1
subdorsal; VIII 1 subdorsal. Spinules located on ends of rachis as
follows: Abdominal segmentI 0-5 anteriorly or laterally, or 1-3 poster-
iorly; II 2-3 rowsanteriorly, 1-8 spinules posteriorly; [II-VI each 3 or 4
rows anteriorly, 4-10 spinules posteriorly; VII 2-4 rows anteriorly, 0-3
spinulesposteriorly. Transverse molting suture terminating oppositeits
midpoint. Mctathoracic setae 140-200, long; eighth abdominal 200-
240u, bases on or adjacent to basal arms of horned process; caudal
160-220, bases cephalad of subdorsalfurrow underneathorjust laterad
of horned process. Horned process weakly to moderately projected, its
basal arms small to large. Vasiform orifice located in base of horned
process, 40-48wide; operculum 16-20, long, 36-40, wide.

Third-stage larva
Description. Broadly ovalin outline.
Margin.Teeth slightly wider than long, apices broadly curved; thoracic
tracheal pore area teeth narrower than other teeth. Ridges 2 times
length of teeth, each with an oblong pale area.
Submargin. Transverse ridges and furrows extending across submar-
gin; ridges with a tooth at distal ends. Both disk and locular pores

ai
usually in 2 pairs on cephalic andin 1 pair on each other body segment.
Anterior 3 pairs of setae each 4, long, 4th and 5th each 10-12y, 6th and
7th each 6-8y.
Dorsal disk. Pairs of both disk and locular pores located as follows:
Cephalic segment 1 submedian; prothorax 1 submedian; mesothorax 1
submedian,1 subdorsal; metathorax 1 submedian,1 subdorsal; abdom-

ti ica
inal segment I 1 submedian; III-VII] each 1 subdorsal. Spinules located
on rachis asfollows: Abdominal segment I 0-2 anteriorly and/orposter-
iorly; If 11-16 around ends andalong anterior and posterior margins;
III-VI each 11-26 in 3-4 rowsanteriorly, 4-12 spinules posteriorly and
extending into median area; VII 6-12 in 2 rowsanteriorly to posteriorly.
Metathoracic setae 160-200y long; eighth abdominal200-220u,bases at
ends of basal arms of horned process or distad of anterior margin of
vasiform orifice; caudal 180-200u, bases at midwidth of submargin.
Horned process moderately projected, its basal arms small to large or
rarely absent.

Second-stage larva
Description. Elliptical in outline.
Margin.Teeth as long as wide, apices moderately curved.
Submargin. Transverse ridges and furrowsvisible on distal % of sub-
175

margin. Both disk andlocular pores in 1 pair on each body segment.


Anterior 3 pairs of setae each 2, long, posterior 4 each apparently 4.
Dorsal disk. Pairs of both disk and locularpores located as follows:
Cephalic segment 2 submedian; mesothoracic and metathorax each 1
submedian, | subdorsal; abdominal segments III-VI and VIII each 1
subdorsal. Spinules sometimes absent; usually 1 submedian located
anteriorly on each of abdominal segments II-VI or IV-VI. Cephalic,
mesothoracic, and metathoracic setae dagger-shaped,each 32-40. long;
eighth abdominalslender, 160-180, bases near ends of basal arms of
horned process; caudal slender, 100-110z, bases on distal submargin.
Horned process moderately projected, its basal armslarge.
First-stage larva
Description.Elliptical in outline.
Submarginal setae of later immature stages located on margin,
anterior 3 pairs each 4, long, posterior 4 each 6-84. Submedian setae
nail-shaped; cephalic 36 long; mesothoracic and metathoracic each
es

48-52u; eighth abdominal 28, bases opposite anterior magin of vasi-


form orifice; caudal slender, 12 long, bases on body margin.

Specimens examined
Holotype, on Chamaedoreaelegans Mart. (Palmae), Mexico, inter-
cepted at quarantine at Dallas, Texas, 28-VI-1966, G.M. Hess. Para-
types, unmounted and 1 mounted pupa taken with the holotype.
Additional paratypes, intercepted at quarantine by plant quarantine
inspectorsofthe U.S. Department of Agriculture between 1936 and 1983,
consist of numerous unmounted andthe following mounted specimens:
Approximately 2000 pupae, 300 third stage, 250 secondstage, 100 first
stage, 54 adult females, 2 adult males. Hosts ofthese collections are as
follows:
Palmae: Arecastrum Romanzoffianum (Cham.) Bece.(= Cocos plum-
osa Hook.); Chamaedoreaelegans Mart., C. glaucifolia H. Wendl., C.
oblongata Mart., C. Tepejilote Liebm., Chamaedorea sp.; Cocos nucifera
L.; Howea (= Kentia) sp.; Phoenix Roebelenii O’Brien; undetermined
palms. The paratypes were intercepted from the following states and
locations in Mexico: Chiapas (Arriaga), Chihuahua, Judrez), Coahuila
(Piedras Negras), México (México City, México D.F., Tenancingo),
Michoacan, Morelos (Cuernavaca), Nuevo Leon (Linares), San Luis
Potosi (Tamazunchale), Tabasco, Tamaulipas (Matamoros, Nuevo Lar-
edo, Reynosa, Tampico), Veracruz (Paplanta, Tuxpan, Tuxtla, Vera-
cruz), Yucatan (Mérida), unstated localities; Guatemala (Petén), un-
stated localities; Honduras, unstated locality.
Paratypesof A. palmaeare deposited in CDA, FSCA, BMNH, IEUN,
and ZAS,
Additional specimensthatI identify as A. palmaebutdonotinclude
in the type series are as follows:
176

Araceae: Philodendron sp. (12 pupae, 1 second stage, 1 first stage),


Mexico: San Luis Potosi (Tamazunchale), Vera Cruz (Fortin).
Bromeliaceae: Tillandsia sp.; unidentified bromeliads.(43 pupae, 13
third stage, 5 second stage). Unstatedlocalities in Costa Rica, Panama
Canal Zone, Colombia, Ecuador.
Moraceae: Artocarpussp. (5 pupae, | adult male), locality unknown.
Orchidaceae: Epidendrum alatum Batem., E. cochleatum L., Epi-
dendrum sp.; Oncidium ornithorhynchum HBK., Oncidium sp.; uniden-
tified orchids. (153 pupae,9 third stage, 2 second stage). Mexico: Chiapas
(Arriaga), México (México D.F., unstated localities), Veracruz (Fortin,
Veracruz); Belize; Guatemala (San José, unstated localities); Honduras
(San Pedro Sula, unstatedlocalities); Nicaragua; Panama CanalZone;
Peru.
Unidentified forest tree (37 pupae, 16 third stage, 16 secondstage),
Panama.

Notes
Aleurocerus palmae andA.flavomarginatus are similar morpholog-
ically andtheir recognitionas two distinct species may be questionable.
In the hundreds of specimens examined, however, two segregates are
clearly evident. Pupae of two forms can be separated by differences in
the shape andsize of the submarginalglands as compared with the size
of the marginalteeth. Investigations of a biological or genetic nature
shouldbe initiated in order to determinethe correct status of the two.

Aleurocerus petiolicola, new species


(Fig. 10)
Living on stemsandleafpetioles.

Pupa
Description. Broadly ovalin outline, 1.25-1.95 mm long,0.85-1.25 mm
wide.
Margin. Black in unbleached, dark in lightly bleached specimens.
Anterior setae 30u long, bases 16-18 teeth apart; posterior 45u, bases
16-20 teeth apart. Teeth 2 times wider than long, apices moderately
curved, 4 width of a tooth apart; thoracic tracheal pore area teeth
narrowerthanotherteeth. Ridges 3 times length of teeth, each with a
pear-shaped pale area. Furrows each with a minute pore near mid-
length.
Submargin.Glands absent. Transverse ridges weak,irregular, extend-
ing across % width of submargin; distal half without, proximal half
with small spinules; most ridges without, rarely with a weak tooth at
distal ends pointed at marginal furrows. Transverse furrows weak,
ere
177

Fig. 10. Aleurocerus petiolicola. A, » Pupa, holotype. B, D, Sections


of margin, submar-
gin and dorsal disk. C, Third stage.
178

irregular, narrower than ridges, without or with a few spinules. Disk


pores located in | or 2 irregular rows at proximal 3rd of transverse
ridges, 5 or 6 pairs on cephalic, 3 or 4 pairs on each other body segment.
Locular pores proximadoftransverse ridges, as numerousas disk pores.
Anterior 3 pairs of setae each 20-24u long, posterior 4 each 36-48u.
Dorsal disk. Pairs of both disk and locular pores located about as
follows: Cephalic segment3 or 4 submedian, 4 subdorsal; prothorax 3
submedian, 3 subdorsal; mesothorax 2 submedian, 2 or 3 subdorsal;
metathorax 1 or 2submedian, | or 2subdorsal; abdominal segment I 0 or
2 submedian,0 or | subdorsal; I] 2 submedian, 2 subdorsal; III 3 or 4
submedian, 1 subdorsal; IV 3-5 submedian, 1-3 subdorsal; V 3 or 4
submedian, 1 subdorsal; VI 2-4 submedian, 1 subdorsal; VII 1 or 2
submedian, 1 subdorsal; VIII 0-2 submedianlaterad of horned process,
1 submedian proximadof base of caudal setae, 2 subdorsal. Spinules
unusually numerous,located as follows: Mesothorax 10-30 submedian
on each half of segment; metathorax 15-30 submedian on eachhalf of
segment; on ends of rachis, abdominal segmentI 4-6 anteriorly, 8-18
posteriorly; II-VI each 3-4 rows extending around ends and approach-
ing median line of body anteriorly and posteriorly; VII 3-4 rowsanteri-
orly, 1 or 2 rows posteriorly; VIII 3 or 4 spinules anteriorly. Transverse
molting suture terminating opposite meso-metathoracic suture. Meta-
thoracic setae 175-200. long; eighth abdominal180u, bases adjacentto
endsof basal arms of horned process; caudal 180-200n, bases in subdor-
sal furrow. Horned process weakly projected, its basal arms small. Vasi-
form orifice located in base of horned process, 40-52 wide; operculum
20-244 long 32-42y wide.

Third-stage larva
Description. Broadly oval in outline.
Margin. Teeth as wide as long, apices moderately curved; thoracic
tracheal pore area teeth narrower than other teeth. Ridges 2 times
length of teeth, each with an elongate pale area.
Submargin.Transverseridges and furrows extending across submar-
gin; ridges withouta tooth at distal ends. Both disk andlocular pores in 3
or 4 pairs on cephalic, in 2 pairs on each thoracic andin or 2 pairs on
each abdominal segment. Anterior 3 pairs of setae each 12y long, pos-
terior 4 each 20p.
Dorsal disk. Pairs of both disk and locular pores located as follows:
Cephalic segment 3 submedian, 1 or 2 subdorsal; prothorax 0 or 1 sub-
median; mesothorax 1 submedian, 1 or 2 subdorsal; metathorax 1 or 2
submedian, 1 subdorsal; abdominal segmentI 1 submedian; II-III each
0 or 1 submedian, 0 or 1 subdorsal; IV-V 0 or 1 submedian,1 subdorsal;
VI-VII 0 or 1 submedian, 0 or 1 subdorsal; VIII 0 or 1 subdorsal; a
subdorsalpore usually presentif a submedianpore is absent. Spinules
located on endsofrachis as follows: Abdominal segmentI 0-3 anteriorly,
179

1-4 posteriorly; II 1-5 anteriorly, 3-8 posteriorly; [II-VI each 8-16 in 2


rowsanteriorly, 4-10 spinules posteriorly and occasionally 2-4 extend-
ing accross submedian area; VII 6-10 anteriorly, 3 or 4 posteriorly.
Metathoracic setae 140, long; eighth abdominal 2004, bases near basal
armsofhornedprocess; caudal 165,, bases on distal submargin. Horned
process weakly projected, its basal arms small.

Specimens examined
Holotype, on stem of Stromanthe lutea (Aubl.) G.F. Mey. (= Calathea
lutea) (Marantaceae), in forest at Las Cruces, east of San Vito, Costa
Rica, 16-II-1970, Michael Kosztarab. Paratypes, a few unmounted and 7
mounted pupae taken with the holotype; a few unmounted and 4 frag-
mentary mounted pupae, and 7 third stage, on stems of Philodendron sp.
(Araceae), Arboretum, Organization of Tropical Studies Research Sta-
tion, La Selva, Porto Viejo area, Costa Rica, 14-I-1970, Michael Kosz-
tarab; unmounted fragments, 2 mounted pupae (+ fragments), and 2
third stage onpetiole ofleaf of Araceae, Panama,intercepted at quaran-
tine at Miami, Florida, 3-VI-1953, Alfred Mills.

Notes
Pupaeof Aleurocerus petiolicola are unusualin lacking teeth at the
distal ends of most submarginalridges. It is the only speciesof the genus
whose pupae are knownto have spinules on both the mesothorax and
metathoraxofthe dorsal disk. Spinules are present on the metathorax of
the closely allied species A. colombiae.

Aleurocerus tumidosus Bondar


(Fig. 11)
Aleurocerus tumidosus Bondar, 1923:159-161; Costa Lima, 1928(1927):100, 1936:151; Silva
et al., 1968:112; Mound and Halsey, 1978:27.
Pupa
Description. Oval in outline, 1.40-1.50 mm long,0.90-1.10 mm wide.
Margin.Colornot observed in unbleached specimens;entirely dark, or
apicesof teeth yellowish in mounted specimens.Anterior setae 24 long,
bases 20-22 teeth apart; posterior broken in specimens examined,bases
24-32 teeth apart. Teeth as long as wide, apices moderately curved, 4-'4
width of a tooth apart; thoracic trachealpore area teeth narrower than
other teeth. Ridges as long as teeth, apparently withoutclear areas.
Furrowseach with a minute pore near midlength.
Submargin. Glandsaslong as wide; truncate at distal ends, proximal
ends of adjacent glandsjoined and appearingscalloped; lateral margins
notched, parallel, nearly contiguous; centers of distal ends opposite
marginal furrows. Transverse ridges irregular, extending across
180

width of space between glands and subdorsal furrow; each ridge with
2-3 rows of stout spinules and a strong tooth at distal end pointed at
centers of glands. Transverse furrows poorly defined, much narrower
than ridges, without or with a few stout spinules, their distal ends
opposite spaces between glands. Disk pores located irregularly along
proximal ends and a few proximad of transverseridges, 6-8 pairs on
cephalic and 3 or 4 pairs on each other body segment. Locular pores
proximad of and as numerous as disk pores. Anterior 3 pairs of setae
each 4, long, posterior 4 each 32,
Dorsaldisk. Pairs of both disk and locular pores unusually numerous,
located as follows: Cephalic segment2 or 3 submedian,2 or 3 subdorsal;
prothorax 2 submedian, 3 or 4 subdorsal; mesothorax 3 or 4 submedian, 3
subdorsal; metathorax 1-3 submedian, 2-5 subdorsal; abdominal seg-
mentI 1-3 submedian, 3-5 subdorsal; II 4-8 submedian,6-8 subdorsal;
IIL 5-8 submedian, 5-8 subdorsal; IV 4-7 submedian, 4 or 5 subdorsal; V
4or5 submedian,3-5 subdorsal; VI 2-4 submedian,2-4 subdorsal; VII1
or 2 submedian, 1-3 subdorsal; VIII 3 subdorsallaterad of hornedpro-
cess, 1 or 2 subdorsal caudadof horned process. Spinules located anter-
iorly on endsof rachisas follows: Abdominal segmentI 0 or 1; II 3 rows;
III-VII each 4-5 rows. Transverse molting suture terminating opposite
its midpoint. Metathoracic setae 150long; eighth abdominal about
250x, bases on or adjacent to basal armsofhornedprocess; caudal 200y,
bases cephaladof subdorsal furrow underneathorjust laterad of horned
process. Horned process ratherstrongly projected, its basal arms small
to fairly large. Vasiform orifice located at midlength of horned process,
about 60wide; operculum 24-28y long, 44-48) wide.

Specimens examined
Syntypes, on ‘cipo preto’ ou ‘cipo caboclo’ [Davilla rugosa Poir.]
(Dilleniaceae), Bahia, Brazil, G. Bondar, 6 mounted pupae.

Notes
Bondar(1923) stated (translation), “... The marginal teeth are of a
dark yellow transparentcolor, black at the base...” Since the marginal
teeth are entirely dark in some mounted specimens, I assume they are
black in unbleached insects and that Bondar’s description referred to
teeth of bleached, mountedinsects. The waxy covering of the pupa was
described by Bondar(1923) as follows(translation) “... When on the
leaf, the insect is completely hidden by wax which surges from the
margin, directing itself to the top and forming abovethe insect a tube 3
or 4mm long, whose extremity supports the black skins coming from the
moltingof the insect and which [skins] coveror close the tube. Laterally,
aroundthe baseof the tube, the rather abundant wax makes a kind of
fringe.”

4
181

Fig. 11. Aleurocerus tumidosus, A, Parasitized pupa, syntype. B, Section


of margin and
submargin. C, Section of margin, submargin, subdorsal furrow,
and dorsaldisk. D, Sec-
tion of dorsaldisk, submargin, and margin,
182

Pupaeof Aleurocerus tumidosus have an unusually large number of


disk and locular pores on the dorsal disk, being distributed across the
rachis and into the subdorsal furrow on some segments. Thespecies is
allied to A. luxuriosus.

Aleurocerus sp.
Aleurocerus sp. Mound and Halsey, 1978:27,in part.
Mound and Halsey (1978) recorded species from Musasp.from Trini-
dad. A slide from the British Museum (Natural History), labeled
banana, Trinidad, Jan. 1959, L.A. Mound, contains only two horned
processes dissected from two pupal cases. They are not identifiable to
species.

ACKNOWLEDGMENTS
1 am grateful to A.B. Hamon, Florida State Collection of Arthropods, Gainesville,
Florida, the late W.W. Sampson,Lafayette, California, and David Hollis, British Museum
(NaturalHistory), London, England, for the loanof specimens. AndI thank W.L.Brown,
Cornell University, Ithaca, New York,for kindly furnishing information on specimens he
collected in Colombia.Thanksare extendedto A.B. Hamon(FSCA), J.B, Lipes, Columbia,
Maryland, and D.R. Smith, Systematic Entomology Laboratory, Agricultural Research
Service, U.S. DepartmentofAgriculture, for their critical reviews of the manuscript. I am
indebted to other members of the Systematic Entomology Laboratory for additionalaid:
HN. Proctor for typing the manuscript, G.C. Steyskal for checking the spelling of specific
names, S. Nakahara for his opinion onspecial details, and M.B.Stoetzel for photographing
specimensandproviding helpful suggestions.I extend mysincere appreciationto all who
haveassisted in any way.
183

LITERATURE CITED
Bondar, G. 1923. Aleyrodideos do Brasil. Sec. Agr. Ind. Obras. 183 pp. (Bahia.)
Browr, W.L. 1968. In W.W. Kemp.Miscellaneousstudies on neotropical ants, [V.(Hymen.
optera, Formicidae). Stud. Entomol. (n.s.). 11 (fase. 1-4):369-416.
Costa Lima, A.M.da. 1928. Segundo cétalogo systematicdos insectos que vivem nas
plantas do Brasil e ensaiode bibliographia entomoldgica brasileira. Arch. Esc. Super.
Agr. Med. Vet. 8(1-2) (1927):69-301. (Aleyrodidae pp. 89-101.) (Nictheroy, Rio de
Janeiro.)
_____.. 1936. Terceiro catdlogo dos insectos que vivem nas plantas do Brasil, Min. Agr.
Esc. Nac. Agron., Dir. Estat. Prod., 460 pp. (Aleyrodidaepp. 144-159.) (Rio de Janeiro.)
Drews, E.A. and W.W. Sampson. 1940. A list of the genera and subgenera of the Aleyrodi-
dae. Bull. Brooklyn Entomol. Soc. 35:90-99.
Mound,L.A. 1967. A new species of whitefly (Homoptera: Aleyrodidae) from ferns in
British glass houses. Proc. R. Entomol. Soc. London (B) 36(1-2):30-82.
Mound, L.A.and §.H.Halsey. 1978. Whitefly of the world. A systematic catalogue of the
Aleyrodidae (Homoptera)with host plant and natural enemy data. Brit. Mus. (Nat.
Hist.) Publ. 787, 340 pp.
Quaintance, A.L. and A.C. Baker. 1914. Classification of the Aleyrodidae. PartIT. U.S.
Dept. Agr. Bur. Entomol. Tech. Ser. 27 (pt. 2):95-109.
Russell, L.M. 1943. A new genusand four new species of whiteflies from the West Indies
(Homoptera, Aleyrodidae). Proc. Entomol. Soc. Wash. 45:131-141.
—_. 1948. The North American species of whiteflies of the genus Trialeurodes. U.S.
Dept. Agr. Misc. Publ. 635:1-85.
Sampson, W.W. 1943. A generic synopsis of the hemipterous superfamily Aleyrodoidea.
Entomol. Amer.(n.s.) 23:173-223,
——____. 1947, Additionsand correctionsto “A generic synopsis of the Aleyrodoidea.” Bull.
Brooklyn Entomol. Soc. 42:45-50.
Sampson, W.W. and E.A. Drews. 1941. Fauna Mexicana.IV. A reviewof the Aleyrodidae of
Mexico. An. Esc. Nac. Cienc. Biol. 2(2-3):143-189, (Mexico, D.F.)
___. 1957. Keysto the generaof the Aleyrodinae andnotes on certain genera. (Homop-
tera: Aleyrodinae). Ann. Mag. Nat. Hist. 9 (ser. 12) (1956):689-697.
Silva, A.G. d’Araujoe. et al. 1968. Quarto catdlogo dos insetos que vivem nas plantas-do
Brasilseus parasitos e predadores. Part IT. 1:1-622. (Aleyrodidaepp.106-120.) (Rio de
Janeiro.)

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