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Museum, University of Nebraska State

Insecta Mundi
University of Nebraska - Lincoln Year 

Notes on Polyphylla Harris with a

description of a new species. (Coleoptera:
Scarabaeidae: Melolonthinae)
Delbert A. La Rue
University of California, Riverside, Riverside, CA,

This paper is posted at DigitalCommons@University of Nebraska - Lincoln.
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998 23

Notes on Polyphylla Harris

with a description of a new species.
(Coleoptera: Scarabaeidae:Melolonthinae)

Delbert A. La Rue
Research Associate, Entomology Research Museum
University of California, Riverside, Riverside, CA 92521
Abstract: Polyphylla aeolus La Rue, new species, is described from the Kelso Sand Dunes, Mojave Desert, San Ber-
nardino County, California, U.S.A. Illustrations of dorsal habitus, significant morphological details, and adult genitalic
forms are provided. A description of the type locality including geographical and ecological parameters is presented. Taxo-
nomic problems within the genus and limitations of recently proposed methods of species identification are discussed. A
modified key is provided to distinguish the new species.
The heretofore undescribed females of Polyphylla anteronivea Hardy, P. rnescalerensis Young, P. nubila Van Dyke, and
P. pottsorurn Hardy, are described. The larval host of P. erratica Hardy is reported, and the adult female is redescribed from
pristine specimens. A dorsalhabitus illustration of each female is provided.
Bionomic and distributional data are presented for Polyphylla avittata Hardy, P. cavifrorzs LeConte, P. hirsuta Van
Dyke, P. r~aoraaha~zserasisHardy,P. petiti GuBrin, P. stellata Young, andP. squanziveratris Cazier

Introduction the Mojave River as it emerges from the east end

of Afton Canyon; however, studies conducted by
Continued investigations of sand dune areas in Meek (1989) indicate that a large perennial lalre
western North America by the author and others occupied the Soda Basin and Mojave River wash
have yielded a n abundance of field data regarding prior to the incision of Afton Canyon. Such a lalre
the behavior, ecology, and distribution of the ge- basin may have been a n important source of sand
nus Polyphylla Harris. The most significant is the for the Kelso Dunes.
discovery of a new species from the Kelso Sand
Dunes. California. U.S.A.. described herein. Manv
of these wind-generated dune systems act as natu-
ral reservoirs for moisture from infrequent desert
rains and, consequently, are a habitat for many
rare and endemic Coleoptera (Andrews and Gil-
bert 1992). Hardy (1981), Hardy and Andrews
(1978), and Young (1966, 1986, 1988) have previ-
ously described Polyphylla species from this dis-
tinctive ecological niche.

Origin and ecology of Kelso Sand Dunes

The Kelso Sand Dunes (Fig. 1) lie in the east-

ern Mojave Desert of southeastern California, a t
a n average elevation of 610m. - (2,000
ft.). The val-
ley in which they occur is bordered on the south,
east, and north by the Granite, Providence, and 1. P a r t i a l n o r t h w a r d view of Kelso D u n e s n e a r t y p e
Kelso Mountain ranges respectively. The Bristol locality of P. aeolus n e w species.
Mountains partly block the western margin of the
valley, leaving a gap t h a t allows wind-blown sand The ability of sand dunes to conserve water
to enter. The dunes cover approximately 1,178 km2 has been well documented, (Chadwick and Dalke
(Bowers 1984) and are located a t the southwestern 1965, Hardy and Andrews 1987, Norris and Norris
end of a n elongated sand mass called "Devils Play- 1961, Prill 1968, Sharp 1966, and Shreve 1938,
ground" (Fig. 2). Sharp (1966) suggested that the among others). Consequently, even i n late sum-
source of sand is a broad alluvial apron formed by mer, months after any rainfall, wet sand exists a t
24 Vol. 12, Nos. 1 & 2, March-June, 1998, INSECTA MUNDI

depths of only 15-20 cm below sun-baked dune (Ottawa, Ontario, Canada), H. F. Howden
slopes (Sharp 1966, Tinkham 1962). (Ottawa, Ontario, Canada), M. L. Jameson
The mean annual precipitation, which includes (Lincoln, NE), P. K. Lago (University, MS), R. H.
occasional snowfall, is 140mm (Bowers 1984). McPeak (Battle Ground, WA), S. McCleve
Temperature mean in summer is 26.6"-32.2"C. (Douglas, AZ), G. H. Nelson (Blue Spring, MO), B.
(80"-90°F.), in winter 4.5"-10.O°C. (40"-50°F), with C. Ratcliffe (Lincoln, NE), A. J. Relfschneider
frequent periods of frost of several days duration (Sierra Madre, CA), E. G. Riley (College Station,
(Axelrod 1950:262). TX), D. E. Russell (Oxford, OH), J . Saulnier
The Kelso Dunes, together with other isolated (Yucaipa, CA), P. E. Skelley (Gainesville, FL), B.
Mojave Desert dune fields, form a well defined D. Streit (Tucson, AZ), D. W. Sundberg (San Anto-
floristic community. The flora is characterized by nio, TX), W. B. Warner (Chandler, AZ), G. Walters,
species with Mohavean (15%), Sonoran-Mohavean (La Puente, CA), C. S. Wolfe (Fort Worth, TX), and
(39%), Southwestern (23%), and endemic (9%) af- the author.
finities (Bowers 1982). The floral composition fluc- Description: Holotype: Male (Fig.3). Length:
tuates annually in response to winter precipita- 30.5mm. Greatest width: 13.2mm. (12.5mm. a t
tion. I n years of abundant rain, the dunes are cov- elytral humeri). Body elongate, parallel-sided;
ered with a blanket of annuals, and the perennials head with integumentary color of vertex, front,
support a profusion of vegetative growth. Vegeta- anterior and lateral clypeal margins and angles
tional surveys have been compiled by Bowers black; thoracic integument, clypeal disk, pygidium,
(1984), Dean (1978), Thorne, et al. (1981), and and abdominal segments rufopiceous; elytra,
Tinkham (1973). scutellum, remainder of venter and appendages
Preliminary surveys of the entomofauna in- rufotestaceous; antennal club testaceous. Head:
habiting these dunes were compiled by Tinkham Clypeus 2X wider than long, anterior margins
(1973, 1975) and Andrews, et al. (1979). sharply reflexed, slightly emarginate; disk sunken
behind strongly reflexed anterior margin, lateral
Polyphylla aeolus L a R u e , n e w species margins narrowing to base, not as strongly re-
(Figs. 3-17) flexed, more strongly narrowed i n posterior 113 to
accommodate antennal scape; interior edge of
T y p e m a t e r i a l : Holotype male (California clypeal margins provided with dense contiguous
Academy of Sciences #17132): U.S.A., California: recumbent elliptical white scales intermixed with
San Bernardino County, Kelso Sand Dunes, 2.6 pale yellow acuminate scales; scale configuration
mi. west of junction Kelbaker Rd. and sand dunes, imbricate; clypeal suture slightly emarginate;
31 May 1986, R. A. Cunningham and R. W. Duff clypeal disk with large punctures; each puncture
collectors, a t blacklight. Allotype female (CAS): provided with a suberect long pale yellow seta,
same locality a s holotype, 24 May 1986, D. A. La occasionally interspersed with white scales. Ocular
Rue collector; 721 additional paratypes, same lo- canthi slightly spatulate, approximately 2X longer
cality a s holotype: 24 and 27 May 1986, D. A. La than wide, extending 113 distance across exposed
Rue (159 males, 19 females); 31 May 1986, R. A. portion of eye, with white scales and long testa-
Cunningham and R. W. Duff (238 males, 22 fe- ceous setae concentrated a t apex; frons with small
males); 1-2 J u n e 1990, B. D. Streit, T. H. Weiser, round punctures, each with long pale yellow setae
R. H. Weiser and W. S. Weiser (143 males, 28 fe- or elliptical white scales, scales becoming dense
males); 24 May 1993, D. E. Russell (111 males, 1 and imbricate in a weak flexuous pattern a t lateral
female). Paratypes deposited in these institutional margins above eyes; vertex smooth, glabrous,
collections: California Academy of Sciences (San shining. Pronotum: Evenly convex; approximately
Francisco, CA), United States National Museum of 2X wider than long, widest a t posterior 113; ante-
Natural History (Washington, D .C.), Los Angeles rior angles obtuse; posterior angles rounded and
County Museum of Natural History (Los Angeles, slightly explanate; marginal bead weakly reflexed,
CA), University of Nebraska State Museum with sparse erect testaceous setae anteriorly and
(Lincoln, NE), University of California, Berkeley laterally; lateral marginal bead distinctly serrate;
(Berkeley, CA), University of California, Riverside pronotum anteriorly and posteriorly with dense,
(Riverside, CA), and the private collections of the continuous mat of short, testaceous setae, espe-
following: R. Alten (Rancho Cucamonga, CA) D. C. cially a t articulation; disk sunken a t midline a t
Carlson (Orangevale, CA), R. A. Cunningham anterior margin; with small, glabrous, oblong,
(Chino, CA), R. W. Duff Qowney, CA), B. D. Gill shallow depression mediolaterally; disk clothed
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998 25

throughout with broad, white and pale yellow glabrous and shining; distal segment of maxillary
scales rising from scattered to contiguous, round palpus conical, subequal in length to basal seg-
punctures; with slender, long, pale yellow, erect to ments combined; clypeopleuron with nearly con-
suberect setae rising medially, becoming sparse tiguous white scales that form feeble flexuous pat-
mediolaterally, concentrated a t center of midline tern; anterior tibiae strongly bidentate; femora,
to form small tuft or feeble scopula (Fig. 9); prono- tibiae and tarsomeres intermittently covered with
t u m trivittate, with a mixture of dense, contiguous heavy to sparse, white scales and pale yellow se-
to imbricate, elliptical, white and pale yellow tae; antennal scape with dense pile of long pale
scales forming narrow vitta a t midline; lateral vit- yellow setae, antennal club 2.7 (linear measure-
tae formed a s irregular, crescentic mass of con- ment) or 4 (along curve) times length of basal
tiguous to imbricate white and pale yellow scales; segments combined. Genitalia: (Figs. 11-14).
a small area of contiguous white scales medially Allotype: Female (Fig. 4). Length: 29.5 mm.
between lateral vittae and pronotal marginal bead, Greatest width: 14mm. (12.5mm. a t elytral hu-
appearing a s white, subreniform, elliptical spot to meri). Differs from holotype in these respects:
unaided eye. Scutellum: Posterolateral margins more robust; thoracic integument, clypeal disk,
glabrous, slightly sunken medially, covered a t cen- and pygidium rufotestaceous; antennal club and
ter with dense patch of contiguous to imbricate appendages rufopiceous. Head: Anterior clypeal
white, acuminate scales; with sparse, decumbent, margin not a s strongly reflexed, especially medi-
pale yellow setae present anteromedially. Elytra: ally, disk more shallowly sunken; vestiture re-
Elytra approximately 1.5X longer t h a n wide; nar- duced on frons and clypeus. Pronotum: Discal se-
rowing posteriorly and broadly rounded, margin tae more pronounced (Fig. lo), lateral marginal
explanate a t elytral humeri, slightly so postero- bead not a s coarsely serrated; mat of short, testa-
laterally; marginal bead slender, provided with 1-3 ceous setae a t articulation approximately 2X
rows of minute pale yellow spiculae posteriorly; longer than i n male. Scutellum: Pale yellow setae
elytral suture with short pale yellow setae, be- medially forming feeble scopula. Elytra: Submar-
coming dense and longer a t elytral declivity; ely- ginal vittae continuous and unbroken. Venter: Ba-
tral surface with deep rugose punctures and no- sal segment of maxillary palpus slightly longer,
ticeable vittae; elytral vittae appearing distinct apical segment slightly flattened apically; anten-
and nearly continuous to unaided eye, microscopi- nal club with 5 antennomeres, with 5 antenno-
cally discontinuous and reduced to irregularly meres composing basal stem; anterior tibiae
shaped clumps of imbricate white lanceolate strongly tridentate; metafemora enlarged, flat-
scales; sutural vittae continuous, formed of white tened, subequal i n length to those of male. Genital
contiguous scales; vestiture of white scales nearly plates: (Fig. 15).
contiguous below elytral declivity near external
margins; intervals uniformly covered with pale Variation i n paratype series
yellow lanceolate scales, scales rarely contiguous
or imbricate, appearing slightly larger than those Males (651) (Figs. 5-8). Length: 23.0-32.5mm.
forming the elytral vittae, frequently meeting ely- Greatist width: 9.5-13.7mm. Differ from holotype
tral vittae; elytral surface with short, suberect, in following respects: color of clypeus rufotesta-
scattered pale yellow setae that are more evident ceous to piceous; pronotum and abdominal seg-
a t humeral declivity and along lateral elytral mar- ments rufotestaceous; antennal club a n d minor
gins, more evident microscopically (30X). Pygid- appendages rufotestaceous; elytral humeri
ium: Length subequal to width; margins reflexed, piceous; Head: Anterior clypeal margin sinuous,
basal portion, including strongly reflexed margin, not a s strongly reflexed; scale vestiture of disk
with numerous, erect, pale yellow setae, setae con- pronounced, obscuring integument beneath, or
tinuing towards base along margin to postero- reduced to clypeal margins; vertex with scattered
lateral 113; disk nearly contiguously covered with large punctures. Pronotum: Serration of lateral
white scales, scales becoming sparse a t midline marginal beads reduced; erect to suberect, pale
and mediolaterally mesad of outer margin. Venter: yellow setae of disk more pronounced or reduced
Head and thorax densely pilose, setae pale, testa- (abrasion?). Scutellum: Scale vestiture reduced;
ceous, obscuring ventral surface; abdominal seg- pale yellow, decumbent setae reduced or absent.
ments connate, each segment with basal glabrous Elytra: Rarely, pale yellow interstitial scales so
band of scattered to contiguous white scales and abundant a s to give specimen olivaceous-colored
pale yellow short erect setae; surface of clypeus cast; short, pale yellow setae absent (abrasion?);
26 Vol. 12, Nos. 1 & 2, March-June, 1998, ZNSECTA MUNDI

vittae not a s broken, discontinuous, forming iden- Bentham (Big Galleta grass), Crotolt mojavensis
tifiable lines, or broken and fragmented, forming a Ferguson (Croton), Petalolzyx thurberi Gray
scattered pattern of irregular clumps. Pygidium: (Sandpaper Plant), and scattered Larrea triden-
Scale vestiture uniformly reduced; pale yellow se- tata (Mocifino and Sess6) Coville (Creosote Bush).
tae reduced to basal tip. Venter: Heavy, dense pile Range cattle heavily utilize the Hilaria for grazing
of testaceous setae slightly obscuring surface. purposes; to what extent their activity has im-
F e m a l e s (70). Length: 27.5-33.0mm. Greatest pacted the ecology of this area has not been ascer-
width: 13.0-16.5mm. Differ from allotype in fol- tained.
lowing respects: Clypeal and thoracic integument Taxonomy. Polyphylla aeolus will r u n to cou-
rufopiceous; antenna1 club rufotestaceous. Prono- plet 20 of Young's (1988) key to the species of
tum: Serration of lateral marginal beads reduced; Polyphylla ("Aedeagal Characters Primarily Ex-
pale yellow setae on disk forming dense scopula. cluded"). The following modification will serve to
Scutellum: Pale yellow setae reduced or absent. separate P. aeolus:
Elytra: White interstitial scales; vitta reduced to
scattered, irregular clumps . 20 (19'). Elytra with long erect setae, excluding
R e m a r k s . Polyphylla aeolus is externally suture; yellow interstitial scales meeting vitta
similar to P. arguta Casey, which has a primarily without glabrous area between .........................
Great Basin distribution. However, P. aeolus may ............................................ rugosipenlzis Casey
be distinguished from P. arguta, and all other 20' Elytra without long erect setae, excluding su-
known species in the genus, by the combination of ture; white (or rarely yellow) interstitial scales
pronotal and elytral setae, pale yellow interstitial not meeting vitta, area between glabrous .........
squamae, consistent reddish-brown elytral color, .......................................................... diffracta Casey
and imbricate squamae. All these character states 20" Elytra with sparsely scattered short sub-erect
are lacking in P. arguta (Young 1988:39-40). pale yellow setae, excluding suture; yellow in-
Polyphylla aeolus is the 3rd species to have the terstitial scales frequently meeting vitta, areas
imbricate scale configuration specifically noted in between discontinuously glabrous to unaided
the literature, the others being P. erratica Hardy, eye .................................... aeolus La Rue, n. sp.
and P. anteronivea Hardy (Hardy and Andrews
1978:4-5; Young 1988:78). Furthermore, ecological The large size, dorsal vestiture, and distribu-
parameters of both species differ significantly: P. tion of this species indicates that it is a member of
arguta is a montane species that primarily occu- the P. decelnli~teataspecies complex (sensu Young
pies pifion-juniper woodlands, generally in, but not 1988) which includes P. arguta, P. dece~nliizeata
restricted to, the upper latitudes of the Great Ba- (Say), and P. rnortahaltselzsis Hardy. However, ex-
sin; P. aeolus inhabits a n isolated sand mass sur- amination of the male aedeagus appears to con-
rounded by xeric desert i n the lower latitudes of tradict these conclusions because it displays simi-
the Mojave Desert. larities to those of P. anteroaivea and P. erratica of
Etymology: This species is named after Aeo- the P. halnrnolzdi species group which are distin-
lus, keeper of the winds in Greek mythology; in guished by "the lack of setae over the entire prono-
reference to the natural forces which have created tal surface, lack of clumped squamae, unique red-
and formed Kelso Sand Dunes, the type locality. dish-brown elytral color, and the aedeagus which
Distribution: Despite intensive entomological does not broaden a t the cleft, with the apex paral-
exploration in the Mojave Desert of California, this lel-sided to broadly spatulate, and appearing trun-
large, unique, psammophilous species has been cated when viewed dorsally" (Young 1988).
collected only a t the southern edge of the Kelso Polyphylla aeolus exhibits only the reddish-brown
Sand Dunes. This may indicate that it is endemic elytral color, but has pronotal setae, clumped
to these particular dunes. More than half of the squamae, and a gradually rounded aedeagal apex,
known North American Polyphylla are restricted (Figs. 11-14). These latter characters would clearly
to a unique, isolated habitat (Young 1988). exclude it from this complex and raise a question
The type locality is located approximately 4.2 to the taxonomic validity of species groups (sensu
km. west of Kelbaker Road, and slightly north of a Young 1988) based upon the male aedeagus.
graded dirt road that provides ingress to this por- Adult behavior: Most sand dune inhabitants
tion of the dunes (Fig. 2). The area consists of possess physiological, morphological, and behav-
semistabilized dune hummocks that support ioral adaptations allowing them to exist in such
abundant growths of Hilaria rigida (Thurber) environments (Andrews and Gilbert 1992). Many
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998 27

dune-inhabiting species of Scarabaeidae seem to down and outward, and the distal tip are depicted
be otherwise unable to survive in desert areas as guttiform, suggesting the parameres are cylin-
(Hardy and Andrews 1987). Polyphylla aeolus is drical for their outer length. My observations indi-
apparently a n obligate to this pattern. cate, in dorsal aspect, the parameres of P. a n -
The following notes of adult behavior are teroltivea and P. erratica are clearly separated and
summarized from observations by several collec- scarcely touch a t the anteroventral recurved por-
tors over several years: tion of the distal tip, which terminates to a gradu-
I n late afternoon, male activity began as rapid, ally rounded point and is clearly not truncated. I n
irregular flights several meters above the sand caudal view, the dorsal ridge is elevated into a
surface. Females were evident in open areas away sharp carina, and the dorsal portion of the
from vegetation, partially emerged from the sand parameres is slightly obliquely flattened giving the
with only the head and pronotum exposed (Fig. distal tip a n elongated, trapezoidal appearance.
16). The apparent variability of the male aedeagal
As dusk approached, male flights became less structure may give some justification and compre-
erratic and more purposeful, and females were hension to Hardy's statement (1981), with which I
found fully emerged and resting on the sand sur- agree, that a species complex based upon this
face. Males flew rapidly upwind, returning in a character "is not entirely accurate."
slow zig-zag flight (indicative of osmoclinotaxic Additional collecting in sand dune areas and
orientation). Males ultimately alighted within a similar habitats in the southwestern United States
few centimeters of a female, and several matings has yielded several heretofore unknown female
were observed (Fig. 17). Copulation ranged from Polyphylla. Their descriptions follows:
75 to 120 seconds, after which the females imme-
diately began to burrow into the sand. I n a few Polyphylla anteronivea H a r d y
instances, females began to re-enter the sand with (Fig. 18)
the male still i7t copula (R. A. Cunningham, pers.
comm.). Evidently, females attract males via a sex Description of female: Length: 27.5mm
pheromone as several males were drawn to a sin- Greatest width: 14.5mm (12.5mm a t elytral
gle female elytron (B. D. Streit, pers. comm.). humeri). Differs from male in the following re-
After twilight, males were readily attracted in spects: robust, parallel-sided, somewhat wider a t
large numbers to black-light, white-light, and the posterior third; integumentary color including ap-
glow of a Coleman lantern. I n these cases, visible pendages deep rufotestaceous; outer margins of
adult activity ceased several hours after sunset, clypeus, protibia, and pygidium, rufopiceous.
because no additional specimens were found a t the Head: Clypeus subquadrate, anterior margin not
lights nor could be located anywhere on the sand as reflexed, disk shallowly depressed, provided
surface. On other dates, male flight activity ceased with deep large punctures more pronounced;
shortly after sunset (B.D. Streit, pers. comm.). clypeal suture slightly emarginate; scale vestiture
Females were not observed to fly, and most limited to outer marginal areas of clypeus and
were encountered on the sand surface. However, a frons; long, pale yellow setae abundant throughout
number of females were excavated a t depths of disk; ocular canthi conical, slightly spatulate. Pro-
10-20 cm by locating the distinctive burial points notum: Strongly convex, elevated and appearing
on the sand similar to those illustrated by Hardy swollen, widest a t anterior 113; uniformly covered
and Andrews (1986:135. Fig. 7.) (R. W. Duff, pers. with white oval scales except weakly reduced a t
comm.). These measurements coincide with the anterior 112, scale vestiture of disk not obscuring
varied depth of the damp sand interface, where pronotal integument as in male, forming single
females presumably spend the daylight hours. medial vitta with symmetrical areas of heavy im-
Discussion: Further aedeagal analysis of P. bricate white scales; with dense area of long, re-
a7ttero7tivea (38 male topotypes) and P. erratica (50 cumbent, pale yellow setae posterolaterally be-
male topotypes) displays significant differences tween central vitta and lateral pronotal margins,
from those illustrated by Young (1988:11, Figs. 8a additional long, recumbent, white and pale yellow
and 8d). Young's illustrations show that, in dorsal setae located anteromedially within central vitta.
aspect, the parameres meet from the basal 112 (P. Scutellum: Transverse; covered with contiguous
alttero~tivea)or basal 113 (P. erratica) to the distal white, lanceolate scales, heavily so in some areas;
tip, which is truncated; in caudal aspect, the dorsal pale yellow setae present medially, posterolateral
ridge of the parameres, which gradually curve margins glabrous. Elytra: 1.5X longer t h a n wide;
28 Vol. 12, Nos. 1 & 2, March-June, 1998, INSECTA MUNDI

margins glabrous. Elytra: 1.5X longer than wide; individual would regain flight posture and repeat
scale vestiture reduced to scattered individual the activity. (D. E. Russell, pers. comm.).
white, lanceolate scales, except in sunken area
above humeri, along lateral margins and length of Polyphylla mescalerensis Y o u n g
sutural area, where they are close to contiguous; (Fig. 19)
setae absent except along suture. Pygidium:
Slightly transverse; uniformly covered with white Description of female: Length: 23.5mm.
scales, except for faint glabrous midline; yellow Greatest width: 12.0mm (9.5mm a t elytral
setae sparsely scattered throughout disk, becom- humeri). Differs from male in the following re-
ing more dense a t posterior marginal bead. Venter: spects: robust, widest a t posterior third; pronotal
Head and thorax with dense pile of testaceous se- integument posterolaterally piceous, ventral pu-
tae which obscures ventral surface, except for area bescence pale testaceous, pygidium and abdominal
of glabrous metasternal midline; abdominal seg- segments rufopiceous. Head: Clypeus distinctly
ments lacking heavy scalation as in male; apical transverse, 2X wider than long, anterior margin
maxillary palpomere conical, subequal in length to not strongly reflexed, anterior angles acute and
basal 2 palpomeres combined; antennae reduced to elevated, disk weakly depressed, suture slightly
5 antennomere club, with 5 antennomeres in basal emarginate; vertex and frons convex; ocular canthi
stem. Profemora and protibiae shorter in length not as elongate; pale testaceous setae of surfaces
than male; metafemora enlarged, flattened, sube- not as coarse and shorter in length. Pronotum: An-
qua1 to length in male; combined length of tar- teromedially with shiny glabrous area; scale vesti-
someres shorter; meso-, metafemora and meso-, ture of vittae and lateral margins reduced; shallow
metatibiae densely covered with long testaceous punctures of disk larger in diameter, nearly conti-
setae. guous, coarser posteromedially. Scutellum: Slight-
Description based on one example with the ly longer, longitudinal midline with pale testa-
following data: U.S.A., California: Inyo County, ceous setae. Elytra: Sutural costa scarcely indi-
Saline Valley Dunes, 26-27 May 1990, R. A. Cun- cated; scales on lateral marginal vittae and poste-
ningham, D. E. Russell, B. D. Streit collectors, rior elytral declivity larger. Pygidium: Laterally
"taken on surface of sand dune in copulo." (B. D. with scales and pale setae reduced, revealing in-
Streit collection). tegument. Venter: Head and thorax covered with a
R e m a r k s : The presence of pronotal setae dense pile of pale testaceous setae which obscures
would exclude this species from the "P.lzarn~nol~di ventral surface; terminal segment of maxillary
complex," as defined by Young (1988:5). Since palpus short, flattened dorsally, subequal in
many females in the genus were unknown a t the length to 2 basal palpomeres combined; antennae
time of his monograph, Young's species groups are reduced to 5 antennomere club, with 5 antenno-
based solely on male characters. meres in basal stem; protibial dentition distinctly
Until recently, P. al~terol~iueamales were more developed, closer together in distribution;
known to exhibit only nocturnal flight behavior. At metafemora enlarged, flattened, shorter t h a n i n
the time the female was collected, in mid after- male; tarsi shorter; 2 distal spines of meso-, meta-
noon, males had been flying for some time. The tibiae noticeably developed; metatibial spines
mating pair was located on the leeward side of the somewhat flattened, rectangular, projecting obli-
dune mass, approximately 1.5 meters above the quely; meso-, metafemora, meso-, metatibiae cov-
sand-hardpan interface. Additional males were ered with long, pale testaceous setae.
collected out on the hardpan substrate, far re- Description based on one example with the
moved from the dune influence (B. D. Streit, pers. following data: U.S.A., New Mexico, Chaves
comm.). County, 9.5 miles W. of Caprock, 27 July 1989, E.
Presence of strong winds and airborne sand G. Riley and C. S. Wolfe collectors (E. G. Riley
does not curtail flight activity. Males were ob- collection).
served flying a t midday i n a sand storm with 65-80 Remarks. The female was discovered on the
kph (40-50 mph) winds. During these conditions, side of a dune swale, approximately one hour after
flight was slow and laborious several centimeters dark. The area consisted of active dune hummocks
above the wind protected leeward dune face. Bee- with Quercus harvardii Rydberg (Shinnery Oak)
tles reachng the dune crest, directly in the pre- distributed on the dune crests (E. G. Riley, pers.
vailing wind's path, tumbled several centimeters comm.).
down the leeward slope. After several seconds, a n
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998 29

Polyphylla nubila V a n D y k e male and a pair ilt copulo were found crawling on
(Fig. 20) the sand surface with no evident burrow or vegeta-
tion i n the immediate vicinity. Males emerged a
D e s c r i p t i o n of female: Length: 25.5mm. few hours after sunset and were readily attracted
Greatest width: 12.5mm (1l.Omm a t elytral to a 175 watt mercury vapor light. A number of
humeri). Differs from male i n the following re- males were found hidden among dried grass up to
spects: robust, subparallel, widest a t posterior 113; ten meters away from the light source. Males of
color of elytra, pygidium, ventral surfaces, lateral Polyphylla decemlineata (Say) were collected a t
margins of prothorax, clypeal disk, and antenna1 the same light station (D. E. Russell, pers. comm.).
scape deep rufotestaceous; frons, anterior clypeal Additional specimens examined, add distribu-
margin, central thoracic disk, elytral humeri, and tional, and phenological data: California, S a n Luis
outer margin of protibia rufopiceous; tarsi, and Obispo County: Atascadero, 25 July 1967, S. Allen
minor appendages testaceous. Head: Clypeus dis- collector; San Luis Obispo, 26 April 1956, W.A.
tinctly transverse, angles quadrate, anterior mar- Wallace collector; Poly Grove, 18 May 1956, R. Goo
gin not a s reflexed, disk weakly depressed, with collector; Paso Robles, 10 April 1956, C. Blundell
scattered large punctures; clypeal suture slightly collector; (all i n D. A. La Rue collection).
emarginate; vertex with long, pale testaceous se-
tae and large coarse, nearly contiguous punctures Polyphylla pottsorum H a r d y
and 1 or 2 rows of white, lanceolate scales a t outer (Fig. 21)
margin above eyes; Pronotum: Distinctly convex,
somewhat impressed a t midline and anterolater- D e s c r i p t i o n of female: Length: 21.5mm.
ally; long testaceous setae of pronotal disk concen- Greatest width: 9.0mm (8.5mm a t elytral humeri).
trated anteromedially to form scopula; appearing Differs from male in the following respects:
univittate to the unaided eye, distinctly trivittate slightly more robust; color of elytra and append-
microscopically. Scutellum: Transverse, disk with ages rufotestaceous, pygidium rufopiceous. Head:
contiguous oval white scales and sparse pale testa- Clypeus quadrate, anterior margin not a s strongly
ceous setae medially. Elytra: Approximately 1.5X reflexed, slightly bisinuate, disk weakly depressed;
longer t h a n wide; vittae reduced; pale yellow vestiture reduced, vertex slightly wider; setae of
lanceolate scales and sparse, suberect, pale yellow, ocular canthi reduced in number. Pronotum: Wid-
setae more evident a t lateral declivity. Pygidium: est a t anterior third; appearing univittate, reduced
Covered with short recumbent sparse pale testa- testaceous setae a t articulation shorter i n length.
ceous setae and white oval scales. Venter: Head Scutellum: Subglabrous except for small area of
and thorax with dense, pale testaceous setae that white lanceolate scales medially. Elytra: White
obscure ventral surface; terminal palpomere short, scales scattered sparsely throughout disk, disk
truncate, slightly flattened dorsally, shorter in appearing glabrous to unaided eye. Pygidium:
length t h a n 2 basal palpomeres combined; anten- Covered with scattered, sparse, white, lanceolate
nae reduced to 5 antennomere club, with 5 anten- scales and short, recumbent, pale setae. Venter:
nomeres i n basal stem; profemora and protibiae Surface of head and thorax not a s densely setose;
shorter i n length; metafemora enlarged, flattened; abdominal segments sparsely covered with short
combined length of tarsomeres shorter; meso- white scales and long pale setae; terminal maxil-
rnetafemora and meso-, metatibiae with long, tes- lary palpomere short, indistinctly truncate, equal
taceous setae. in length to basal 2 palpomeres combined; anten-
Description based on one example with the nae reduced to 5 antennomere club, with 5 anten-
following data: U.S.A., California: San Luis Obispo nomeres i n basal stem; combined length of tar-
County, Atascadero, 15 J u n e 1991, D. E. Russell someres shorter, metafemora enlarged, flattened.
collector (D. E. Russell collection). Description based on one example with the
R e m a r k s : Adults of this species normally be- following data: U.S.A., Texas: Ward County,
come active following the first warm weather in Monahans Sand Dunes, 19 July 1976, R. W. Duff
June, occasionally a s early a s April, with some collector (R. W. Duff collection).
populations active until late July. The type series R e m a r k s : The female was located crawling a t
was collected i n late May (Van Dyke 1947). night on the sand surface i n a n area of active
Specimens were collected i n a n area of sandy, parabolic dunes with scattered Calamovilfa gigan-
loose soil with Adeltostorna (chamise), Quercus tea (Nuttall) Scribner and Merrill (Giant Sand
(oak), and scattered grasses and annuals. The fe- Reed) and Quercus harvardii Rydberg (Shinnery
30 Vol. 12, Nos. 1 & 2, March-June, 1998, INSECTA MUNDI

Oak) (R. W. Duff, pers. comm.). Polyphylla erratica Hardy

This species was previously known from the (Figs. 22-23)
Monahans Sand Dunes in Ward and Winkler
Counties, Texas, and the Mescalero Dunes, Chaves Young's description and illustration of the fe-
County, New Mexico. Collecting in intervening male of Polyphylla erratica Hardy (1988:78 and
and adjacent sand habitats indicates the distribu- Fig. 72) was based on a n abraded, damaged speci-
tion of P. pottsorutn is far more extensive: Texas: men and does not accurately depict the true form.
Andrews County, 1.5 miles S. Junction Highways I t is here redescribed and illustrated from pristine
115 and 181, 6 July 1991, C. S. Wolfe and E. G. specimens.
Riley collectors; New Mexico: Lea County, 23.5 Length: 27.0-28.5mm. Greatest width: 12.0-
miles and 26.5 miles W. of Hobbs, 4 July 1991, C. 14.5mm (11.5-12.0mm a t elytral humeri). Differs
S. Wolfe and E. G. Riley collectors; Eddy County, 5 from male in the following respects: robust, widest
miles E. of Loco Hills, 5 July 1991, C. S. Wolfe a t posterior 112; color of head, clypeal disk, anten-
collector. nal club, and protibial dentition piceous. Prono-
The Andrews County, Texas, locality is a con- turn: With nearly contiguous to imbricate, oval,
tinuation of the Monahans Sand Dune system, white scales, becoming sparse anteromedially,
with similar sand composition and plant commu- leaving small, crescentic, glabrous area on either
nity (C. S. Wolfe, pers. comm.; see remarks under side of central vitta. Venter: Surface of head and
P. monahansensis below). thorax covered with dense, pale, testaceous setae
The Lea and Eddy County, New Mexico, locali- and lanceolate, white scales, weakly obscuring sur-
ties are composed of extensive deep reddish dunes face; terminal maxillary palpomere short, conical,
that are distinct from those found in the Mona- shorter in length than basal 2 palpomeres com-
hans system, suggesting they are of a n older, finer bined; metafemora enlarged, flattened, tarsi
grained composition (sensu Norris and Norris shorter.
1961:612). However, the floral community does not Description based on 5 examples with the fol-
differ significantly (C. S. Wolfe, pers. comm.). lowing data: U.S.A., California, San Bernardino
Polyphylla pottsorum is herein defined as a County: Amargosa River near Hwy. 127, 4 May
complex of geographical phenotypes differing prin- 1986 (I), R. A. Cunningham collector (R. A. Cun-
ciply by male dorsal coloration. Although some ningham collection); 30 April 1986 (3), 6 May 1986
specimens appear to be quite distinctive, they in- (I), D. A. La Rue collector (D. A. La Rue collec-
variably represent intrapopulational phenotype tion).
extremes. Remarks: Aside from sexually dimorphic
Populations from New Mexico differ signifi- characters, females are remarkably similar to
cantly from the nominate phenotype of west Texas males in this species.
and are briefly characterized as follows: Females were located by observing small ag-
Chaves County, Mescalero Sand Dunes ("9.5 gregations of 3 to 8 males hovering above or
miles W. Caprock, 1.7 mi. S.E. of Highway 380"). crawling on the soil surface. Examination of these
The coloration of the pronotal integument is a pale locations yielded a female still within or partially
reddish brown rather than the typical piceous, and emerged from a burrow. I n one instance, a male
the elytra are more of a golden yellow. was noted to physically displace a partially
Lea County, "23.5 miles and 26.5 miles W. emerged female from the burrow to attain a copu-
Hobbs." The elytral coloration ranges from deep latory position (personal observation).
reddish brown to black (appearing dark chestnut Polyphylla erratica is a halophytic species,
brown microscopically) with a n even gradient be- with larvae, pupae, and adults occurring i n moist,
tween the two. A female from "26.5 miles west of salt encrusted soil. Larvae were taken a t the roots
Hobbs" differs from the nominate phenotype de- of Distichlis divaricata Beetle (Salt Grass), which
scribed above by a rufopiceous pronotal integu- is abundant throughout the Amargosa River bot-
ment and deep chestnut brown elytra. tom. Polyphylla erratica is undoubtedly restricted
Eddy County, "5 miles E. Loco Hills." The col- to this unique ecotone as the known distribution
oration of the pronotal integument ranges from follows the moist environs of the Amargosa River
deep chestnut brown to black; the elytra from yel- system.
lowish brown, a s in the Mescalero Dunes popula- Males of this species exhibit a variety of flight
tion, to a deep chestnut brown, with a n even gra- behaviors. Active flight has been observed from
dient between the 2. late morning to early afternoon (R. A. Cunning-
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998

ham, pers. comm.), and from late afternoon to well July 1984, R. W. Duff collector; Highway 78, vi-
into dusk (pers. observation). Specimens have also cinity of Green Road, 19 July 1982, R. W. Duff
been taken i n a blacklight trap (D. E. Russell, collector; 19 July 1987, R. A. Cunningham collec-
pers. comm.) indicating that males are not re- tor; 2.1 miles N. of Laguna Dam, 24 July 1984,
stricted to diurnal or crepuscular flights as was R.W. Duff collector; San Bernardino County, 13
previously assumed (Hardy and Andrews 1978). road miles NE of Earp, Highway 62, 3 August
As with P. altteronivea, strong prevailing winds do 1973, R.W. Duff collector; Riverside County, Dos
not seem to curtail flights but may reduce the Palmas Spring, Coachella Valley, 30 May 1992,
number of active adults (pers. observation). R.W. Duff collector.
Ravens, shrikes, and coyotes were observed to Remarks: Young's (1988:78) statement "...
opportunistically feed on the adults of this species. this is a species of the true xeric desert" is mis-
Specimens were found impaled on vegetation by leading and conveys the impression that P. cavi-
shrikes, and coyote scats examined were composed frolts inhabits the open, arid desert. Ecologically, it
almost entirely of undigested adult P. erratica is nearly entirely restricted to desert riparian or
fragments (R. A. Cunningham and R. W. Duff, similar habitats. The recorded distribution follows,
pers. comm.). or is adjacent to, natural or manmade water-
D i s t r i b u t i o n r e c o r d s : The following data courses and drainage ways that include rivers,
hopefully will provide additional knowledge to- canals, and irrigation ditches. A unique record is
ward understanding the ecology and distributions from Dos Palmas Spring, a n isolated area of
of those species considered. Unless otherwise indi- spring-fed ponds and extensive Washinggtonia filif-
cated, specimens are in the collections of the re- era (Linden) Wendland (California Fan Palm).
spective collectors.
Polyphylla hirsuta V a n Dyke
Polyphylla avittata H a r d y U.S.A., Arizona: Santa Cruz County, Patago-
nia Mountains, Mt. Washington, vic. Forest Serv-
U.S.A., UTAH: Washington County, Snow ice Road #I28 and Duquesne Road, 5300' elev.,l2
Canyon State Park, 14 and 15 July 1978, 30 June August 1991, R.A. Cunningham collector,l5 watt
1986, G. H. Nelson collector, a t blacklight. BL; Duquesne Road (Forest Service Road #61) 7.8
R e m a r k s : This area is composed of white and miles E. Junction Highway 82, 4800' 'elev., 13
red Navajo sandstone with numerous black lava August 1991, L. G. Bezark, R. A. Cunningham, D.
beds. The sand dunes are lighter in color and not E. Russell collectors (D. E. Russell collection).
a s extensive a s those comprising the type locality, R e m a r k s : Both localities are in the Patagonia
Hurricane Dunes, approximately 30 km. to the Mountains near the type locality, Mt. Washington
east. The plant community is essentially Artelnisia (Van Dyke 1933). However, the localities differ
(sagebrush), with scattered scrub oaks (G. H. Nel- ecologically. Males were taken a t 1,590 m. (5,300
son, pers. comm.). Hardy and Andrews (1978:2) ft.) elevation, in predominately oak woodland with
noted that the predominant vegetation a t the type some scattered pines and junipers. Specimens
locality was Artelnisia filifolia Torrey (cited as were taken a t 1,440 m. (4,800 ft.) elevation, in a
"filiformis"). sandy area best described as oak-mesquite transi-
tion with several large Juglans major (Torrey)
Polyphylla cavifrons LeConte Heller (Arizona Walnut). Polyphylla halnmondi
LeConte was sympatric a t the lower elevation. The
U.S.A., Arizona: Maricopa County, Coon's first significant summer monsoons had recently
Bluff, near Gila River, 20 July 1987, 22 July 1990, drenched the area comprising both localities, re-
16 August 1990, R. W. Duff collector; Yuma sulting in a phenomenal number of insects a t mer-
County, Ehrenberg, rest area, 22 August 1989, R. cury vapor and blacklight stations during subse-
W. Duff collector; quent nights (R. A. Cunningham, D. E. Russell,
U.S.A., California: Imperial County, 6 miles E. pers. comm.).
of Holtville, East Highline Canal, 10 June 1989, J . Young's (1988:63) record of this species from
Beierl and E. Barchet collectors (R. A. Cunning- "Patagonia Mountain" (singular) is probably a ty-
ham collection); Titsworth Road and East Highline pographical error. This name comprehensively ap-
Canal, 24 July 1992, R.W. Duff collector; Junction plies to the mountain range; no singular mountain
of Highway 111 and Alamo River, 19 July 1987, R. peak by this name exists in Arizona or adjacent
A. Cunningham collector; Two Rivers rest area, 20 Mexico.
Vol. 12, Nos. 1 & 2, March-June, 1998, INSECTA MUNDI

Polyphylla monahansensis H a r d y (D. A. La Rue collection).

Remarks: Young (1988) reported that the ge-
U.S.A., TEXAS: Winkler County, Junction of nus Polyphylla occurs worldwide between 15" and
State Highway 115 and Farm Road 874, NE of 53" North latitude, with the southern-most New
Kermit, rest area, 28 July 1978, R. W. Duff collec- World record as "Honduras, 3.2 miles SW of Valle
tor; Andrews County, 1.5 miles south of Junction de Angeles."
Highways 115 and 181, 6 July 1991, C. S. Wolfe The Nicaragua record represents a consider-
and E. G. Riley collectors. able distributional extension as it is south of 14"
U.S.A., NEW MEXICO: Lea County, 4 miles North latitude. I t also represents a new country
south of Jal, Highway 18, 6 July 1991, C. S. Wolfe record for the genus.
and E. G. Riley collectors.
R e m a r k s : The Texas localities are a continua- Polyphylla stellata Young
tion of the Monahans Dune system, with similar
plant community (R. W. Duff, C. S. Wolfe, pers. U.S.A., California: Sacramento County,
comm.). Polyphylla pottsorurn was sympatric a t Sloughhouse, 20 June 1961, Rooney Brothers col-
both sites. lectors, Japanese Beetle Survey (D. A. La Rue col-
The New Mexico locality consists of extensive lection); same county, Carmichael, American
deep reddish dunes a s described above for P. pott- River, near Ancil Hoffman State Park, 24 July
sorurn i n Lea and Eddy Counties. Polyphylla pott- 1989, 25 July 1992, D. E. Russell collector, mer-
sorurn was not collected a t this site (C. S. Wolfe, cury vapor light.
pers. comm.). Aside from this locality, P rnonahan- R e m a r k s : This species inhabits native grass
sensis was known to be sympatric with P. pott- and oak woodlands often adjacent to riparian habi-
sorurn throughout it's distribution. This represents tats in the Sacramento Valley.
a new state record for this species in New Mexico. At Carmichael, males were abundant in areas
Young (1988:52) recorded P. monahar~sensis of "manicured" grass with scattered oaks (i.e., pic-
from Mexico based on a single example with the nic and golf course areas). This suggests P. stellata
following data: "Mexico, Chihuahua, between may be remarkably tolerant ecologically. Speci-
Yepachic and Tomachic (correct spelling: Temosa- mens have also been collected a t Discovery Park,
chic), large canyon bottom, 31 July 1984, Doug Sacramento County, and the Antioch Dunes, Con-
Mullins collector" (S. McCleve collection, on loan to tra Costa County (D. E. Russell, pers. comm.). The
R. M. Young, Cody, WY). According to the collec- undescribed females of this species have been col-
tor, this locality is in the Sierra Madre Occidental lected from the Antioch locality. Unfortunately,
range in west-central Chihuahua a t a n elevation of these specimens were not made available to the
1,500-1,650 m. (5,000-5,500 ft.). The habitat con- author for this study.
sists of oak-pine woodlands surrounding a small
stream. The specimen in question was taken a t Polyphylla squamiventris Cazier
blacklight with Plusiotis beyeri Skinner, Diplotaxis
sp., and Phyllophaga sp., all of which are generally U.S.A., Texas: Presidio County, Redford,
associated with this ecotone (D. Mullins, pers. Highway 170, 6 June 1992, C. S. Wolfe collector,
comm.). below streetlight.
Polyphylla monahansensis is a sand obligate Remarks: A male of this uncommon species
restricted to regions of arid climate in southeast- was taken in a n area surrounded by cultivated
ern New Mexico and adjacent western Texas. fields bordering the Rio Grande River, approxi-
While it would not be surprising for t h s species to mately 24 km. east of Presidio, the type locality (C.
occur in suitable habitats in northern Mexico, it is S. Wolfe, pers. comm.). Possibly, P. squarniventris
extremely unlikely t h a t it would exist in a mon- continues to inhabit undisturbed remnants of na-
tane environment as described above. Taking into tive vegetation remaining along the river.
account the striking disparity of ecological pa- Polyphylla harnmondi was also collected under the
rameters, I consider this record to be erroneous. same streetlight.

Polyphylla petiti G u k r i n Acknowledgements

NICARAGUA, Nueva Segovia: 15 km. N. of I am indebted to Robert W. Duff (Downey, CA)

Jalapa, (no date), J . S. Maes collector, U.V. light for informing me of the existence of the new
INSECTA MUNDI, Vol. 12. Nos. 1 & 2. March-June. 1998 33

Polyphylla described herein and for providing a Five new Polyphylla Harris. Pan. Pac. Ent.
female of P. pottsorum. Special gratitude is ex- 54(1):1-8.
tended to Brett C. Ratcliffe (Lincoln, NE), and Hardy, A. R., and F. G. Andrews. 1986. Studies
William B. Warner (Chandler, AZ) for advice, con- in the Coleoptera of western sand dunes. 2.
structive suggestions, and reviewing various drafts Notes on four Scarabaeidae from the Algodo-
of the manuscript. Thanks are also due to Max E. nes Dune system. Coleop. Bull. 40(2):127-139.
Badgley (Biological Photography, Moreno Valley, Hardy, A. R., and F. G. Andrews. 1987. Studies
CA) for photographing the types and undescribed in the Coleoptera of western sand dunes. 3.
females; Richard A. Cunningham (Chino, CA) for Remarks upon Serica from some Nevada sand
a n additional female of P. erratica; Doug Mullins dunes with descriptions of two new taxa. Co-
(Tucson, AZ) for field data and habitat description leop. Bull. 41(2):173-179.
in Chihuahua, Mexico; Gayle H. Nelson (Blue Meek, N. 1989. Geomorphic and hydrologic impli-
Spring, MO) for field data and habitat description cations of the rapid incision of Afton Canyon,
of P. avittata from Snow Canyon, Utah;'Edward G. Mojave Desert, California. Geology 17:7-10.
Riley (College Station, TX) for females of P. mes- Norris, R. M., and K. S. Norris. 1961. Algodo-
calerelzsis and P. pottsorum; David E. Russell nes dunes of southeast California. Geol. Soc.
(Oxford, OH), for providing the female of P. nu- Amer. Bull. 72:605-620.
bila; Barney D. Streit (Tucson, AZ) for computer Prill, R. C. 1968. Movement of moisture in the
expertise, field photographs, and the female of P. unsaturated zone in a dune area, southwest-
a7zterolzivea; and Charles S. Wolfe (Fort Worth, ern Kansas. U.S. Geol. Survey Prof. Paper
Texas) for specimens and field data. 600D:l-12.
Sharp, R. P. 1966. Kelso Dunes, Mojave Desert,
References CA. Geol. Soc. Amer. Bull. 77:1045-1074.
Shreve, F. 1938. The sandy areas of the North
Andrews, F. G., A. R. Hardy, and D. Giuliani. American desert. Association of Pacific Geog-
1979. The coleopterus fauna of selected Cali- raphers 4:ll-14.
fornia sand dunes. Report to the BLM on con- Thorne, B. L., B. A. Prigge, and J. Henrick-
tract CA-960-1285-1288-DEOO. California De- son. 1981. A flora of the higher ranges and the
partment of Food and Agriculture, Sacra- Kelso Dunes of eastern Mojave Desert in Cali-
mento. 142 pp. fornia. Aliso 10:71-186.
Andrews, F. G., and A. J. Gilbert. 1992. Cadiz Tinkham, E. R. 1962. Studies in Nearctic Desert
hardyi, a new genus and species of leaf beetle Sand Dune Orthoptera. 6. A new genus and
from western North American sand dunes. three new species of large sand-treader camel
Coleop. Bull. 46(1):4-14. crickets from the Colorado Desert with keys
Axelrod, D. I. 1950. Evolution of desert vegeta- and notes. Bull. So. Calif. Acad. Sci. 61(2):89-
tion in western North America. Carnegie Inst. 111.
Wash. Pub. No.590:215-306. Tinkham, E. R. 1973. Flora of the Kelso Dunes.
Bowers, J. E. 1982. Plant ecology of inland dunes Unpublished report prepared for the U.S. De-
in western North America. Jour. Arid Envi- partment of Interior, Bureau of Land Man-
ronments 5:199-220. agement, California Desert District Office,
Bowers, J. E. 1984. Plant geography of south- Riverside, California.
western sand dunes. Desert Plants 6(1):31-54. Tinkham, E. R. 1975. Endemicity in the fauna
Chadwick, H. W., and P. D. Dalke. 1965. Plant and flora of the Kelso Dunes and the threat of
succession on sand dunes in Fremont County, the dune buggies. Unpublished report pre-
Idaho. Ecology 46:766-785. pared for the U.S. Department of Interior, Bu-
Dean, L. E. 1978. The California desert sand reau of Land Management, California Desert
dunes. iv + 71 pp. Unpublished report pre- District Office, Riverside, California.
pared for the U.S. Department of Interior, Bu- Van Dyke, E. C. 1933. Two new species of Scara-
reau of Land Management, California Desert baeidae. Pan. Pac. Ent. 9(3):115-116.
District, Riverside,. CA. Van Dyke, E. C. 1947. Western Coleoptera. Pan.
Hardy, A. R. 1981. Polyphylla Harris in Baja Pac. Ent. 23(4):160-161.
California. Coleop. Bull. 35(3):299-302. Young, R. M. 1966. A new species of Polyphylla
Hardy, A. R., and F. G. Andrews. 1978. Studies and a designation of two lectotypes. Jour.
in the Coleoptera of western sand dunes. 1. Kans. Ent. Soc. 39(2):233-236.
34 Vol. 12, Nos. 1 & 2, March-June, 1998, WSECTA MUNDI

Young, R M. 1986. Three new species of North Young, R. M. 1988. A monograph of the genus
American Polyplzylla Harris. Trans. Nebr. Polyphylla Harris in America North of Mexico.
Acad. Sci. 14:47-50. Bull. Univ. Nebr. St. Mus. 11(2):1-115.

Fig. 2. Map of Kelso Dunes and vicinity.

INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June. 1998 35

Fig. 3-10. Fig.3, Polyphylla aeolus, holotype male, dorsal habitus (0.75X); scale equals 5mm. Fig. 4,
Polyphylla aeolus, allotype female, dorsal habitus (0.75X); scale equals 5mm. Fig. 5.-8, Polyphylla aeolus,
male paratype variation (0.75X); scale equals 5mm. Fig. 9. Polyphylla aeolus, holotype male, left lateral
view of pronotum (5X); scale equals 2.5mm. Fig. 10. Polyphylla aeolus, allotype female, left lateral view of
pronotum (5X); scale equals 2.5mm.
36 Vol. 12, Nos. 1 & 2, March-June, 1998, INSECTA MUNDI

Fig. 11.-15. Polyphylla aeolus, holotype male, parameres; scales equal lmm. Fig. 11, dorsal view; fig.
12, lateral view; fig. 13. ventral view; fig. 14. caudal view; fig. 15, allotype female, ventral view of genital
plates; scale equals lmm.
INSECTA MUNDI, Vol. 12, Nos. 1 & 2, March-June, 1998 37

Fig. 16-21. Fig. 16. Polyphylla aeolus, female emerging from sand substrate; fig. 17. Polyphylla aeo-
lus, male and female iit copulo; fig. 18. Polyphylla ai~teroitiveaHardy, dorsal habitus of female (0.75X);
scale equals 5mm; fig. 19. Polyphylla ~nescalerelzsisYoung, dorsal habitus of female (0.75X); scale equals
5mm; fig. 20. Polyphylla itubila Van Dyke, dorsal habitus of female (0.75X); scale equals 5mm; fig. 21.
Polyphylla pottsoruin Hardy, dorsal habitus of female (0.75X); scale equals 5mm; fig. 22.-23. Polyphylla
erratica Hardy, dorsal habitus of females showing variation (0.75X); scales equal 5mm.

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