DEATH VALLEY JUNE BEETLE

PROJECT 1995

for

NATIONAL BIOLOGICAL SURVEY and

DEATH VALLEY NATIONAL PARK

by

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MIAMI UNIVERSITY

OXFORD, OHIO TABLE OF CONTENTS

SECTION PAGE

ABSTRACT ...... 1

INTRODUCTION ...... 1

METHODS

sampling strategy ...... 5

genetic analysis ...... 7

RESULTS

genetic data...... 8

sampling results ...... 9

DISCUSSION

taxonomic status ...... 13

aistii5ution...... ; ...... 1'5~=~------

biology ...... ;...... 17

sexual attraction ...... 21

predators ...... 22

threats ...... 22

CONCLUSION ...... 25

ACKNOWLEDGEMENTS ...... ~ ...... 25

LITERATURE CITED ...... ·... 26 INDEX TO TABLES, FIGURES, AND APPENDICES

TABLE # PAGE

1. Collection sites for june beetle survey ...... 38

2. Enzyme and buffer systems used for electrophoretic analysis of Polyphylla ...... 44

3. Allele frequencies for P. anteronivea ...... 45

4. Allele frequencies for P. erratica ...... ~ ...... 45

5. Genetic variability at 14 loci for all populations of P. anteronivea and P. erratica (standard errors in parentheses) ...... 46

6. Matrix of genetic similarity and/or distance coefficients between P. anteronivea and P. erratica ...... 4 7

7. Matrix of genetic similarity and/or distance coefficients for selected Polyphylla taxa...... ,' ...... 48

8. Estimated acreage for Polyphylla erratica ...... 49

1. Location of Death Valley National Park...... 8

2. June beetle census locations along the Amargosa River drainage and the Saline and Eureka Valleys in Death Valley National Park...... 29

3. Polyphylla erratica survey sites along the Amargosa River drainage ...... 30

4. Polyphylla anteronivea survey sites in Saline and Eureka Valleys ...... 31

5. Polyphylla anteronivea male ...... 32

6. Polyphylla erratica male and female ...... 33

7. Saline Valley Dunes. (A) north side of lake with mesquite covered dunes; (B) west side of lake with creosote and open sand ...... 34 FIGURE # PAGE

8. Alkali flats with Distichlis saltgrass (A) Ash Meadows; (B) Saratoga Springs ...... 35

9. Female Polyphylla erratica in her burrow about eight inches below the soil surface ...... 36

10. Polyphylla erratica impaled on mesquite thorn by loggerhead shrike at Saratoga Springs ...... 37

APPENDIX

A Hardy, A.R., and F.G. Andrews. 1978. Studies in the Coleoptera of western sand dunes. 1. Five new Polyphylla Harris. Pan.-Pac. Ent. 54:1-8.

B Young, R.M. 1988. A monograph of the genus Polyphylla Harris in America north of Mexico (Coleoptera: Scarabaeidae: Melolonthinae). BulL of the Nebraska State Mus. 11: 1-115. 1995 DEATH VALLEY JUNE BEETLE REPORT

David Russell Department of Zoology, Miami University, Oxford, OH 45056

ABSTRACT

The beetle genus Polyphylla (Coleoptera: Scarabaeidae: Melolonthini) is a taxonomically difficult group. This genus has had over 73 names proposed for the 28 currently recognized species north of Mexico. Five of these species are currently listed as Category 2 organisms-­ candidates for federal listing as threatened or endangered but for which insufficient information is known. The purposes of this study were: 1) to ascertain the taxonomic status of the Category 2 species Polyphylla anteronivea and P. erratica using data from allozyme electrophoresis; 2) to survey Death Valley National Park and adjacent areas to determine the present distribution of both taxa; 3) to study present populations in light of potential threats to their existence. A total of 144 sites were surveyed over three collecting periods between April 28 and June 29, 1995. Fourteen loci from 11 enzyme systems were examined using starch gel electrophoresis. Analysis of 38 P. anteronivea and 44 P. erratica revealed significantly different =~~=a;Hel~f=rebeetles to be restricted in range, although common in appropriate habitats. Habitat alterations, primarily by water removal, represent the greatest potential threat to these species.

INTRODUCTION The beetle genus Polyphylla (Coleoptera: Scarabaeidae: Melolonthini) is found discontinuously throughout the Northern Hemisphere. Of the 65 species known worldwide (Dalla Torre 1912, DeWailly 1948, 1993), 32 are found in the New World. Since the genus was first described by Harris (1841) over 73 names have been proposed for the 28 currently recognized species in America north of Mexico (Young, 1988). Numerous attempts have been made in the last 150 years to identify morphological characters that provide accurate species identifications. Indeed Polyphylla as a whole appears replete with apparent homoplasies and convergent evolution as its members have independently evolved similar characteristics to survive in extremely hostile environments. The present systematics of Polyphylla makes identification tentative at best. Fifteen of the 28 currently described species of Polyphylla north of Mexico are found in a single sand dune system or on a single mountain top (Young, 1988). Frequently, collection location is used to differentiate specimens because of a lack of substantial morphological characters. Five species of Polyphylla are currently Category 2 species-- candidates for federal listing as threatened or endangered but for______which insufficient information is known to make a formal decision.

The purpose of this study was threefold: 1) to ascertain the taxonomic status of the Category 2 species Polyphylla anteronivea and Polyphylla erratica using data from allozyme electrophoresis; 2) to survey Death Valley National Park and adjacent areas to determine the present distribution of both taxa; 3) to study the present populations in light of potential threats to their existence. The present distribution of both species was thought to be (Hardy and Andrews, 1978) primarily within the borders of· Death Valley National Park (Fig 1). Though described as separate species by Hardy and Andrews (1978) their current taxonomic status is in

2 question. As Young (1988) stated in his monograph "... there are certainly gradations between the two [Po anteronivea and P. erratica] which can only be clearly identified by the collecting locality". Upon extracting the aedeagi (not done by Hardy and Andrews (1978) at the time of the original description) Young (1988) found no significant differences and stated there "was insufficient data to decide whether these two names represent valid, somewhat cryptic species, or a single, perhaps clinal, taxon. This species [Po anteronivea] is very closely related to P. erratica, so closely III fact that I doubt that both names represent biological species."

The Saline Valley Snow-front June Beetle (P. anteronivea) IS known only from the sand dune complex in the Saline Valley of Inyo Co., CA (Fig 2B). With the signing of the Desert Bill in early 1995, Saline Valley was incorporated into Death Valley National Park. Typical individuals are easily distinguished from closely related specIes (especially P. erratica) by the thick covering of white scales on the pronotum, scutellum, and anterior elytra with the distal half of the elytra reddish brown without vittae (Fig 5). However, worn specimens frequently show little or no scalation making them very difficult to distinguish from similarly worn P. erratica. Hardy and Andrews (1978) distinguish P. anteronivea from P. erratica as follows: "P. anteronivea has slightly less well developed clypeal angles, an apically broadened clypeus (quadrate in P. erratica), less elongate and apically less pointed scales, and an anterior elytral impression which results in a transverse carina from the scutellum towards the humeral area. II Because of the extremely remote location of the sand dunes (about 70 km from the nearest paved

3 road) and scarcity of collections, nothing is known about their biology. Males are most frequently collected at blacklights within an hour or two of dark. The female is currently undescribed in the literature, but a single individual was recently collected. She appears flightless, similar to many other dune endemic Polyphylla (pers. observ.). The Death Valley June Beetle (P. erratica), another desert species (Fig 6), is found primarily along the Amargosa River in southern Death Valley (Fig 2A). The type locality and largest known concentration of P. erratica is Saratoga Springs along the Amargosa River drainage in the southern edge of Death Valley National Park. The Amargosa River is approximately 120 km in length, and only three short reaches contain surface water on a year around basis. In years when precipitation is· abundant, a continuous stretch of water can cover one-half to two-thirds of the river channel. Polyphylla erratica is found in perennially damp, salt encrusted alkali flats, particularly those covered with Distichlis grass [salt grass]. Similar to P. anteronivea, females are flightless and are collected by excavating them from their burrows. Unpredictably triggered by currently unknown stimuli, males periodically fly in the late morning and afternoon as opposed to the normally post-sunset flight period. In agreement with Hardy and Andrews' (1978) observations, males that fly during the day do not appear to be attracted to blacklights after dark. However, for unknown reasons, and particularly when no daytime activity is noted, males are readily attracted to blacklights at dusk. This differs from' P. anteronivea where males are frequently collected at blacklights after dark even if males were observed flying

4 earlier m the day.

METHODS A) SAMPLING STRATEGY Because adult emergence is heavily dependent upon environmental conditions, with initial beetle emergence varymg from year to year by as much as three weeks, three separate collecting periods were used during this 1995 study: 27 April - 3 May

(primarily for P. erratica), 18-28 May (for both species), and 20-30 June (primarily for P. anteronivea). Previous collection records for P. anteronivea show this species usually emerges several weeks later than P. erratica from Saratoga Springs. Two different methods were used to collect specImens. Males flying during the day were collected with aerial butterfly nets. Other specImens were collected with mercury vapor (white) and/or ultraviolet (black) lights which were operated within a few hours """""----"--.~~=~~~~~~~~~~=~~~~=~~==~=~=~= after sunset. Ten to 15 unattended battery-operated 22-watt blacklights equipped with a baffle, funnel, and collecting bucket were used on any given night of trapping. The bucket traps were dug into the ground flush with the soil line. These traps allowed simultaneous sampling of several different localities and were very important for collecting individuals from low density populations or locating new populations. A solar switch activated the lights at dusk and beetles attracted to the lights were collected, live, in a bucket below the light. A portable mercury vapor light set-up consisted of a 1000 w Honda generator with a 77 m cord along which four 22-w blacklights

5 were mounted every 15.5 m, with two 160 w mercury vapor lights usually mounted at the generator and at the opposite end of the cord. The mercury vapor lights are considerably brighter than the blacklights and contain the entire light spectrum, not exclusively limited to the ultraviolet wavelengths as are blacklights. Some specIes of beetles appear to be more attracted to some wavelengths of light than others; hence, in some situations mercury vapor lights (which contain all wavelengths including those found in blacklights) appear to attract a broader representative of beetles than blacklights. Also, and probably more importantly, the mercury vapor lights are much more powerful and can be seen from a greater distance in this largely flat unobstructed terrain, thus potentially drawing beetles from a greater distance. Our intentions were to conduct a mark and recapture experiment to determine the distance that Polyphylla were attracted to the lights. Unfortunately, because of difficult weather conditions (wind and rain) and relatively small numbers of beetles observed this year, this project was not conducted. However, our use of these collecting methods allowed sampling of relatively large areas, increasing our ability to attract and collect beetles even when present in low numbers. Females, were located by looking for emergence holes in the ground or by following males. Females were excavated from their burrows with a shovel. At each site sampled, a global positioning system (GPS) location was taken fixing the position to within ± 100m. General notes on each site were recorded, including presence of habitat disturbances, types of vegetation present, and description of soil· conditions (wet

6 vs. dry).

B) GENETIC ANALYSIS All Polyphylla were collected live and either frozen in liquid

nitrogen III the field or express mailed live to Miami University for freezing at -80C. Specimens remained frozen until processed for genetic analysis. For sample preparation, a beetle was thawed on Ice, the abdomen detached, and if a male, the genitalia removed and mounted for subsequent comparisons. The tissue within the thorax was scraped out and divided among five micro-centrifuge tubes-four for refreezing for later tests or comparisons. The tissue from the fifth tube was homogenized in a 1: 1 volume of tissue to grinding solution (2% Triton X-100). After centrifugation, the supernatant was either absorbed ontoWhatman #3 filter paper wicks and inserted directly into 12.5% or 15% starch gels or placed in wells for cellulose acetate plate inoculation. Electrophoresis and histochemical staining followed standard procedures (Hebert, 1989; Acquaah, 1992). Analysis of 11 enzyme complexes on 6 buffer systems revealed a total of 14 well-differentiated loci (Table 2). The patterns for each enzyme locus were photographed and alleles were interpreted and scored on the basis of relative mobility.. An internal standard was included so reference values could be used for initial comparisons of alleles from different runs. These comparisons were later confirmed using line-up gels or plates. The computer program BIOSYS-1 (Swofford and Selander, 1981) was used to calculate allelic frequencies, distance coefficients, and genetic variability estimates.

7 RESULTS A) GENETIC DATA Of the 14 loci examined in 38 P. anteronivea (mean sample size 34 individuals [not all individuals were scorable at each locus]) and 44 P. erratica (mean sample size 43 individuals), both were fixed for the same allele at three loci. Polyphylla anteronivea was fixed at eight loci (57%), with three additional loci with an allele present at greater than 95% frequency; a total of 11 (79%) fixed or nearly fixed loci (Table 3). In comparison, P. erratica had three fixed loci and two other loci with an allele present at greater than 95% frequency; this only represents 36% of the total examined (Table 4). The same three loci were fixed in the two species and the two alleles found at greater than 95% frequency in P.erratica were also found in high frequency in P. anteronivea. Significantly different allele frequencies exist for four loci (DIA, MPl1, MPI2, PEP). Polyphylla anteronivea appears to possess three unique alleles (DIA-A, MDH2-C, and MPH-A) at the 14 loci compared. It is interesting to note a possible gene duplication at the PEP locus. All P. anteronivea were fixed for both the Band E alleles; while P. erratica had both Band E yet they were not the most frequent alleles. The apparent gene duplication occurred after the separation of the two populations. All 38 P. anteronivea individuals possess the BE alleles while the majority of P. erratica (58%) share the C allele, only 13% have the B allele and 18% the E allele (A, D, and

F alleles accounted for the remaining 11 % combined). Two heterozygous PEP BE individuals were expected under Hardy- Weinberg conditions; however, none were observed.

8 P. erratica had a direct count mean heterozygosity per individual of 0.18 (S.E. ± 0.05) with P. anteronivea at 0.13 (S.E. ± 0.07). The mean number of alleles per locus for P. anteronivea was 1.6 compared to 2.6 for P. erratica (Table 5). If the presence of more than one allele at a locus is considered a polymorphism then the percentage of loci polymorphic for P. anteronivea was 0.43, much less than the 0.75 for P. erratica.

For P. anteronivea, the direct count mean heterozygosity of 0.13 . was slightly greater than the expected value of 0.11 under

Hardy-Weinberg equilibrium condition. In contrast, for P. erratica the expected allele frequency was 0.32, while the direct count was only 0.18, 56% of the expected value. Chi square tests for deviation from Hardy-Weinberg equilibrium proportions were significant (p< 0.05) in 8 of 11 (3 were fixed) calculations (1 expected by chance) for P. erratica and 1 of 6 (8 were fixed) calculations for P. anteronivea. The majority of deviations were a deficiency of heterozygotes; the notable exception was the greater number of

observed heterozygotes at the PEP locus in P. anteronivea with 38 of 38 individuals sharing the BIE allele pattern. A Nei (1978) unbiased minimum distance value of 0.17 and a Modified Rogers' distance (Wright, 1978) of 0.42 was calculated between the two species (Table 6).

B) SAMPLING RESULTS A total of 144 sites were surveyed over the course of three collecting periods (Table 1). Ideally, distributional studies should be done over several consecutive years; however, those areas in which

9 beetles were detected this summer can be assumed to represent the minimum beetle distribution. Polyphylla anteronivea is certainly restricted to the Saline Valley Dunes (Fi g 4). Sixteen trap sites in and directly adjacent to the dunes contained P. anteronivea. A total of 38 P. anteronivea were collected from 24 sites in a total of 36 trap nights. Traps within 100 m of the dunes failed to attract a single Polyphylla. The beetles are found primarily in both the dunes along the northeast and north sides of the lake playa in the bottom of Saline Valley. One collection was recorded in a previous insect survey from a small set of sand dunes on the southeast side of the lake. Polyphylla erratica is found in a much larger geographical area. Individuals were collected or have been recorded from Fairbanks Spring in Ash Meadows National Wildlife Refuge in the north, south to Death Valley Junction, Shoshone, and Tecopa. A large population is found at Saratoga Springs in southern Death Valley. No beetles have been observed north of Saratoga Springs toward Badwater in Death Valley, north of Ash Meadows towards Beatty, nor south of Dumont Dunes in Salt Creek along Highway 127 (Fig 3). A total of 69 P. erratica were collected; 47 netted while day flying and 22 collected

III 5 sites from 120 trap nights. Although not seen during this study, P. erratica males have been observed emerging in large numbers, with several hundred individuals seen flying about at a given time (Evans and Cunningham, pers. observation.). In an attempt to determine generation times a 1.5 m by 1.5 m patch of alkali with Distichlis grass in the Amargosa River channel at Saratoga Springs was excavated to a depth of about 0.5 m on April

] 0 28, 1995. Although no adult beetles were observed nor emergence holes noted, in previous years numerous males and females had been observed in this vicinity. In 1995, no adult beetles (or pupae) were excavated, but eight larvae were collected. All larvae were large, with head capsule widths ranging between 7.2 and 7.6 mm. In one of the only studies on the biology of Polyphylla, Van Steenwyk and Rough (1989) showed for P. decemlineata (although P. erratica adults are slightly smaller overall) that 1st ins tar larvae head capsule widths range between 2.4-3.2 mm, 2nd instar larvae 4.0-5.5 mm, and 3rd instar larvae between 6.0-8.7 mm, with approximately three weeks being spent in the pupal stage prior to adult emergence. No adults were seen during the initial 27 April through 1 May, 1995 survey in any area. On May 18, 1995, during early stages of the second survey in Saratoga Springs, only three males were collected in blacklight traps and no day flying males were observed. This is much later in the .~~======~====~~.-----~-~~------year than previously when males had been observed as early as April 25 in this area, with peak emergence usually occurring by the second week of May. On May 19, 1995, a second 1.5 by 1.5 m patch, immediately adjacent to the previous site, was excavated resulting in five females, four males, and four (3-3rd instar, - I-2nd instar) larvae. The females were 15-23 cm below the soil surface (Fig 9) with 3 of the 5 having tunnels with emergence holes at the surface. The remaining two did not appear to have dug their way to the surface yet. Two of the males were less than 5 cm below the soil surface with no tunneling beneath them; they apparently dug back in the soil after the previous night's flight. If the 3rd instar larvae were still 2

I I or 3 weeks from eclosion, as data from P. decemlineata suggest, this would project the continued adult emergence well into mid-June, a late emergence date not normally recorded for this area. On May 27, Doug Threloff, the Resource Specialist for Death Valley National Park, captured 17 males flying between 7:00-7:45 PM in an area approximately the size of a football field. On June 21, 1995, during the mid to late June survey, another 1.5 by 1.5 m patch was excavated immediately adjacent to the previous sites. Seven 1st instar larvae, with head capsule widths from 2.2-2.5 mm, were collected. The larvae were in two clumps, one cluster surrounding a chamber, about 18 cm below the soil surface, containing a dead, largely decomposed female. It appeared the eggs were laid in close proximity to the area in which she died. In addition to the 1st instar larvae, a solitary 2nd instar larva (head capsule size was 4.7 mm) was noted. No adults were seen during the day or collected at blacklights at night suggesting that the adult flight season was probably over. Only one adult P. erratica has been recorded in July (Hardy and Andrews, 1978) and this was probably from Ash Meadows, 90 km to the north, where emergence usually occurs several weeks later than the more southern, lower elevation Saratoga Springs site. On June 21, 1995 Doug Threloff .collected 13 male and one female P. erratica 4.6 km northeast of Death Valley Junction. The males were flying over the Distichlis grass between 7:00-7:40 PM. The female was walking on the ground. This area is about 75 km north of Saratoga Springs and within about 15 km of Ash Meadows National Wildlife Refuge. On June 28, 1995, 48 males were collected in Ash Meadows in an area roughly 20 m x 60 m

1 2 flying between approximately 5:30-8:30 PM. An estimated 15-20 km!h wind was blowing. Some males would work their way up­ wind, zig-zagging until they would suddenly veer off, flying down wind and gradually work their way back up over the same ground. Other males would fly in a direct line up-wind when they would suddenly make a quick turn, flying down-wind 50-60 m and then would zIg-zag back up-wind.

DISCUSSION A) TAXONOMIC STATUS The biological and genetic results of this study indicate that contrary to Young's (1988) assertions, although closely related, P. anteronivea and P. erratica represent valid species. A Nei (1978) genetic distance coefficient of 0.17 (and a Modified Rogers' Distance of 0.42), combined with the presence of unique alleles, indicate that these two speCIes represent distinct taxa. In addition, biologically,

" " these two speCIes occupy different habitats with P. anteronivea occupying sand dunes (Fig 7) and P. erratica occupying moist alkali! Distichlis grass mudflats (Fig 8). These two species should continue to genetically differentiate since there is a lack of gene flow and differing environmental pressures. Genetically, these species appear to be closely related sister taxa. A Nei D value of 0.17, is on the lower end of values considered sufficient to differentiate species, but is nearly indistinguishable from the 0.18 value determined between Polyphylla decemlineata from Oroville, CA (Butte Co.) and P. nubila, another Category 2 species from coastal CA (San Luis Obispo Co.). In a previous study (Russell

13 and Guttman, unpub.), these two specIes had the lowest Nei D value of the Polyphylla surveyed (Table 7); however, they are morphologically quite distinct and have been placed in different species complexes under Young's (1988) current classification scheme. Similarly, the lowest Modified Rogers' distance coefficient determined was 0.40 between P. decemlineata (combination of three populations from CA, NV, and CO) and P. nubila restricted to San Luis Obispo Co., CA. Another closely related pair of species, P. devestiva from ID (Owohee Co.) and P. barbata, another Category 2 species, from Santa Cruz Co., CA, had a Modified Rogers' distance value of 0.43. The 0.42 value between P. anteronivea and P. erratica certainly falls within species level values in this genus. The genetic data for these sister species suggest that P. anteronivea arose from P .. erratica. Through a founder event, which seems unlikely with flightless females, or more probably through a range constriction, the Saline Valley population became genetically ~~=~~~~=~~==:~=~~~~~==~~.------­ isolated from the main population. During the Pleistocene, Death Valley was an overflow basin filling with glacial meltwaters and run- off from the Sierra Nevada mountains as well as from pluvial precipitation that w:as higher than today's levels. Death Valley until

10,000 years ago was the 185 m deep Lake Manly. Th~ playa of the Death Valley basin had only ephemeral lakes since the Pleistocene; these were usually only centimeters deep and lasted a few weeks after major storms; however, 2,000 years ago one lake may have lasted longer since it was over 10 m deep (Collier, 1990). The repeated expansions and contractions could easily have isolated small populations in pockets of suitable habitat. Cut off from gene

14 flow from the mam population and living within an increasingly hostile environment insect in most of these pockets died out. It is interesting from a theoretical perspective to note that P. anteronivea and P. erratica appear to fit the classic peripheral isolate model of speciation, whereby small, peripherally isolated populations have become phenotypically distinct, possibly in response to locally extreme physical conditions. Speciation is thought to be rapid because small populations are more quickly affected by the simultaneous effects of genetic drift and selection than are larger popUlations. Additionally, peripheral or founder popUlations are initially cut off from the main population and the founder individuals lack some genes found in the main or ancestral species. This theory has previously been illustrated by example only and not through systematic study (Ridley, 1993), but P. anteronivea and P. erratica appear to show systematic evidence in support of this model. Results from the genetic analysis in this study revealed P. anteronivea contain only a subset of the alleles present in P. erratica. Tliey· have only a few potentially unique alleles not found in P. erratica (these may have evolved after the split) while, geographically, our distributional survey has shown P. anteronivea isolated in an extremely harsh environment on the periphery of the range of a geographically more widespread species.

B) DISTRIBUTION There is no current overlap in the ranges of P. anteronivea and P. erratica. Polyphylla anteronivea is confined to the Saline Valley Dunes. These beetles are found primarily in the dunes along

1 5 the northeast and north sides of the lake playa although individual beetles have been recorded from small sets of sand dunes on the southeast side of the lake. No individuals have been noted flying even 100 m away from the dune edge. Thus, the total habitat is probably only about 1000 hectares.

Polyphylla erratica, by contrast, IS found in a much larger geographical area. Polyphylla erratica were collected or have been recorded from Fairbanks Spring in Ash Meadows National Wildlife Refuge in the north, south to Death Valley Junction, Shoshone, and Tecopa. A large population is then found at Saratoga Springs in southern Death Valley (Fig 3). Their range, while apparently confined to the alkali/ Distichlis grass communities, is found to be discontinuous along the Amargosa River, over 120 km in length. There are no records of Polyphylla nor were Polyphylla found

III Eureka Valley or in northern Death Valley; historically, areas most likely to have supported populations allowing continuous distribution from present day populations into Saline Valley for P. erratica. It was surprising that no P. erratica were collected· in the extensive alkali/ Distichlis grass community in the north end of Death Valley, north of Stovepipe Wells. These alkali flats extend for several km along Death Valley Wash between the Cottonwood and .the Grapevine Mountains. The absence of beetles in this area is probably the result of the flightless nature of the females, making dispersal or recolonization after localized extinctions nearly impossible. Polyphylla erratica are found on the east side of the Funeral Mountains at Ash Meadows National Wildlife Refuge, only 75 km southeast on a direct line from Stovepipe Wells. However, within

1 6 Death Valley, P. erratica have not been collected north of Saratoga Springs (about 130 km south of Stovepipe Wells) along the Amargosa River or along Salt Creek in the Badwater Basin. A few small patches of Distichlis grass are found throughout the Badwater Basin up through Furnace Creek to nearly the southern edge of Stovepipe Wells, but individually or combined these patches do not appear sufficient to support populations of P. erratica allowing the colonization of the extensive alkalilDistichlis flats north of Stovepipe Wells.

C) BIOLOGY

Very little IS known about the biology of both specIes. Casual observations made over approximately the last 15 years reveal Polyphylla erratica to be possibly bivoltine with observations of large emergences followed by a smaller emergence the next year. However, this initial larval survey suggests that the majority of beetles cycle annually with only few individuals continuing development into a second year. However, the development and emergence of this species is probably closely linked to environmental conditions and to development time, though usually one year, may fluctuate between annual and biannual depending upon local conditions. The larvae appear to be most commonly associated with, and probably feed on, the roots of Distichlis spicta (salt grass) in perennially damp soils, frequently in the actual river bed (these areas rarely contain standing water). Females typically remain in their burrows in the soil, although periodically a female is observed crawling on the soil surface. Males typically fly during daylight

1 7 hours, usually in the late afternoon, or for about three hours

commencing at dusk. Interestingly, if males are observed flying III the late afternoon no males are collected in that area in light traps after dark. However, if no daylight activity is noted, males in the same area are attracted to lights after dusk. Males are extremely strong fliers and if disturbed can cover great distances quickly, often flying until out of sight. Polyphylla erratica collected from Ash Meadows along the northern edge of their range appeared morphologically similar to southern representatives from Saratoga Springs, with the exception that most were only one-half to two-thirds the size. Whether the SIze difference was more a factor of larval food availability or actual genetic differences between populations cannot be determined with information currently available. An analysis of gene flow between representative populations along the corridor from Ash Meadows to Saratoga Springs was unsuccessful due to a deterioration of specimens from Shoshone and - Saratoga Springs, rendering tliefr------­ electrophoretic patterns unreadable. Ash Meadow populations of P. erratica appear to peak in mid-June, later than the typical mid-May peak in Saratoga Springs, although similar to the mid-June flight period of P. anteronivea.· The differences III emergence times between P. erratica populations are probably due to a combination of environmental factors associated with an almost 600 m difference III elevation between the higher Ash Meadows and Saratoga Springs, and also possibly the more northerly location of Ash Meadows. Little has been determined about the biology of P. anteronivea. Larvae are unknown and the only female collected was found in the

1 8 late afternoon crawling on the sand beneath a saltbush (Atriplex sp.) along the edge of a dune. The vegetation present in areas were P. anteronivea were collected were much more variable than for P. erratica. The northern most dunes are covered with Mesquite (Prosopis spp.) have large quantinies of sediment mixed into the dune sand (Fig 7 A). A considerable amount of dead wood, in some cases whole hillocks, IS evident in these patches. Saltbrush (Atriplex ssp.) and Iodine Bush (Allenrolfia occidentalis) are also common on or at the base of the dunes. This contrasts with the vegetation along the northeast-side where the dunes are open sand with patches of Creosote Bush (Larrea tridentata) around the margins (Fig 7B). Saltbrush (Atriplex ssp.) is common in the flats bordering the dunes. Males are most frequently collected at blacklights starting about 20 minutes after sunset and continuing until about 11 :00 P.M. Males have been observed flying in late afternoon (app. 4:00 P.M.) although their daytime flights appear to coincide with barometric pressure fluctuations or increased humidity. They are usually seen flying only during unstable, cloudy days frequently with scattered rain drops. Being extremely strong fliers males were once seen flying right above the sand surface III 50-60 km/h wind. Attempts to conduct life cycle studies similar to P. erratica were unsuccessful as no larvae, females, or pre-emergent males were found. Because of the tremendous variation in environmental conditions, results of a single year's distributional/ biological study can be misleading, largely underrepresenting areas in which Polyphylla occur and giving misleading biological information associated with unusual climatic conditions. Environmental

1 9 conditions for collecting were difficult this year in no way appearing to be typical for an average year. In addition to the regular gale force winds, the combination of below normal maximum daytime temperatures at Park Headquarters at Furnace Creek for April and May (mean maximum temperature was 31 0C in April, 1995 [80 year mean was 32.1 oCl and 34.7 °C in May, 1995 [80 year mean was 37.3 oCl) and usually heavy precipitation in January made 1995 a highly unusual year, This apparently contributed to the late emergence of P. erratica. All weather measurements were recorded from the nearest weather station at Death Valley National Park Headquarters in Furnace Creek approximately 100 km north of Saratoga Springs. These reading describe general conditions at Saratoga Springs but do not reflect the localized differences frequently encountered in this desert environment. However, by June, 1995, daytime temperatures frequently exceed 43 0C at Furnace Creek and during our study, several days exceeded 46 0C (one was 50.6 OC). Average maXImum temperatures for June, 1995 was 41.1 °C still under the 80 year average of 42.8 °C. The environment throughout the area is extremely harsh. Weather data from Furnace Creek since 1912 show a mean daytime temperature of 38.5 °C in May, reaching mean temperatures of .46.4 0C by July. This is the hottest site in the Western Hemisphere and a record of 56.70 C in the shade at Furnace Creek (probably hotter in Badwater, 100 ft farther below sea level) is only 1.00 C less than the hottest record in the world (Collier, 1990). Daytime ground temperatures can reach over 900 C. The 80 year average for total annual rainfall is 4.6 cm; 1995 had a total of 8.3 cm of which 6.6 cm

20 fell in Furnace creek in January. Additional ram fell to the south of Furnace Creek in April and May, but was unrecorded at Furnace Creek. Some years there is no rainfall at a given point in the valley. The little rain that does reach the ground is subject to an evaporation rate of 375 cm per year (Collier, 1990). The ability for larvae to hold over an additional year, or to have the plasticity to delay emergence until environmental conditions are most favorable, would be a great advantage in this extreme environment. The possibility certainly exists for the generation time to exceed three, or possibly four years as has been recorded for P. decemlineata (Richter, 1966).

D) SEXUAL ATTRACTION The erratic zig-zagging, up-wind flight of the males indicated the probable use of a sex pheromone by the females to attract the male. The use of sex pheromones by female Polyphylla has been observed and recorded by others (Young, 1988; VanSteenwyk and Rough, 1989). Interestingly, some males would suddenly alight and begin digging beneath the alkali crust; sometimes two or three males would congregate in the same spot. In three instances, females were dug up from beneath where males alighted-two from burrows adjacent to where the males landed, but in one case, a female was excavated although no exit hole was noted. This female was more than 15 cm below the surface, still in her pupal chamber with her cast pupal skin with no evidence of tunneling. Males must have the ability to detect minute quantities of pheromone, considering it was released by a female beneath the surface of the ground, with unequal diffusion through the soil and still detectable by the males

2 1 under windy conditions. The pheromone appears concentrated in the abdomen. A female was accidentally cut in half while attempting to excavate her from a burrow; a number of males were attracted to the abdomen placed on the soil surface but neglected the head and thorax placed about a meter away (D. Russell, pers. observ.).

E) PREDATORS Natural predators include loggerhead shrikes that will capture active males and skewer them on mesquite thorns for later consumption (Fig 10) and coyotes, whose scats, at times of large emergences, are composed entirely of beetle parts (Evans, pers. comm.) Nothing has been recorded regarding other predators or parasites. However, wasps in the insect family Tiphiidae were abundant in the blacklight samples. Tiphiids are commonly parasites of beetle larvae, particularly "white grubs", a designation frequently given to many scarab larvae, such as Polyphylla. ~=~======~=====~ ------

F) THREATS These beetles are adapted to extremely harsh environmental conditions that are not conducive to human cohabitation. With 17 positive survey sites for the presence of this species, the total range of P. anteronivea is confined to the Saline Valley Dunes, now entirely contained within the borders of Death Valley National Park. Since it is adapted to the harsh conditions in Saline Valley, and protected from man-made disturbances, it appears secure. Burro numbers could become a threat and should be monitored to ensure that their populations do not adversely affect the vegetation through over-

22 grazmg and trampling. Because of the highly restricted distribution of P. anteronivea, this species is certainly susceptible to natural, catastrophic events, but this possibility has existed since the divergence of this species and no evidence was noted of conditions that might compound the problems of survival. Under the current park service guidelines there appears to be little man-made threat to this specIes. Similarly, P. erratica, although it occupIes a larger geographic range than P. anteronivea, is still extremely localized in its distribution. However, these beetles appear to be present in all of their historical localities (with the exception of a P. erratica recorded from Pahrump, NV). During this survey, no alkali/Distichlis grass areas could be located in or around Pahrump. The human population of this area has expanded. rapidly over the past five years and the habitat may have been lost or the beetle label is in error; the beetle may have been collected in the general geographic area with Pahrump being the nearest city [beetles caught near the eastern edge of Ash Meadows might easily qualify for the latter possibility]. Polyphylla erratica appears to be confined to the alkali/ Distichlis grass community found discontinuously along about a 150 km stretch of the Amargosa River channel. A rough estimate of acreage that is perennially damp with a cover of alkali and salt grass that would be potentially occupied by P. erratica is approximately 4750 acres (Table 8). A mosaic of jurisdictional responsibilities oversee these areas. Sites with positive Polyphylla identifications included 6 National Park Service locations, 11 Bureau Land Management locations, and 5 U. S. Fish and Wildlife Service locations (Fig. 9).

23 Additional beetles are undoubtedly located on private lands around Shoshone and Tecopa. Despite the fact that the Amargosa River is largely underground and open water is nonexistent, periodic flooding of the Distichlis grass community is certainly normal in high ram years. Consequently, this area would be under little developmental pressure. Despite the fact that some off-road vehicular activity does

occur III all of these study areas the overall impact to the moist alkali areas appears negligible. Environmental conditions are extremely harsh. Temperatures are high and considering the saline content, lack of shade, and sparse vegetation, this area is unsuitable for grazing of range animals. Some wild horses are found near Death Valley Junction which cause minor trampling problems, but as with the off-road vehicles, their impact appears minimal. With the exception of development for geothermal purposes or a significant increase in mining activity from the largely dormant claims scattered throughout the area, there appears to be little threat to P. erratica throughout its limited range. The greatest potential threat for P. erratica wouJd include changes in groundwater hydrology in event of groundwater pumping. Some pumping is currently occurring in the area. The groundwater that comes to the surface at Ash Meadows is derived from a carbonate rock aquifer which covers the southern one-third of Nevada. In the event that pumping up-gradient from Ash Meadows affects spring discharge rates, the hydrology of the river (and the persistence of damp soils) could be jeopardized.

24 CONCLUSION There appears to be no imminent threat to either P.

anteronivea or P. erratica and they do not appear to be III need of special protective status at this time. This distributional and genetic survey shows these beetles to be restricted in range, although common in appropriate habitats, and represent distinctly separate taxa. Because of their association with unique, highly restricted biological communities, their numbers and populations should be periodically monitored, however, particularly if there are changes associated with ground water usage or in ground water contamination. Decisions to remove P. anteronivea and P. erratica from the C2 list should be made on a specIes by species basis with specific information in hand based primarily upon data generated during this study. The removal of other species from the C2 list should not be made based on information from this study, and these data and conclusions should not be extrapolated to species not includes in this study. Habitat alterations, by water removal, represent the greatest potential threat to the species characterized in this study.

ACKNOWLEDGEMENTS This project was funded by the National Biological Survey (NBS). The tireless efforts of Doug Threloff in logistical support, collecting, and manuscript advise were greatly appreciated. Thanks to Kurt Pindel and Nichole Mortenson for their hospitality and help in the field. Thanks also to Dr. Paul Russell for his help in the field and

25 manuscript advise. A very special thank you to Dr. Sheldon Guttman for his unwavering support, technical advise, and manuscript comments. Finally, a thank you to Jan Trybula and the lab staff for their help in the electrophoresis.

LITERATURE CITED

Acquaah, G. 1992. Practical protein electrophoresis for genetic research. Dioscorides Press. Portland, OR 92 pp.

Collier, M. 1990. Introduction to the Geology of Death Valley. Death Valley Natural History Association, Death Valley, CA. 60 pp.

Dalla Torre, K.W. von. IN: Junk, W. 1912. Coleopterorum Catalogus. V.20, par.49: Scarabaeidae: Melolothinae III. 290 pp.

DeWailley, P. 1948. Etude sur Ie genre Polyphylla Harris in extremeorient (Col. Scarabaeidae). Musee Heude. Notes d'entomologie chinoise 12: 100-121.

------. 1993. Revision des. especes palearctiques du genre Polyphylla Harris (Coleoptera: Melolonthidae). Sciences Nat. Bull. 79:5-14.

Hardy, A.R., and F.G. Andrews. 1978. Studies in the Coleoptera of western sand dunes. I. Five new Polyphylla Harris. Pan.-Pac. Ent. 54: 1-8.

Harris, T.W. 1841. A report on the insects of Massachusetts, injurious to vegetation. Folsom, Wells, and Thurston, Cambridge. 459 pp.

Hebert, P.D.N., and M.J. Beaton. 1989. A practical handbook: methodologies for allozyme analysis using cellulose acetate electrophoresis. Helena Laboratories, 1989.

Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89: 583- 590.

Richter, P.O. 1966. White grubs and their allies. Oregon State University Press, Corvallis, Oregon. 219 pp.

26 Ridley, M. 1993. Evolution. Blackwell Scientific Publications, Inc., New York. 670 pp.

Steenwyk, R.A., and D.Rough. 1989. Biology of the Tenlined June Beetle (Coleoptera: Scarabaeidae). J. Econ. Entomol. 82: 1738-1742.

Swofford, D.L. and R.B. Selander. 1981. BIOSYS-l A FORTRAN program for the comprehensive analysis of electrophoretic data III population genetics and systematics. J. Heredity 72:281-283.

Wright, S. 1978. Evolution and the genetics of populations, Vol. 4. Variability within and among natural populations. Univ. Chicago Press, Chicago. 580 pp.

Young, R.M. 1988. A monograph of the genus Polyphylla Harris in America north of Mexico (Coleoptera: Scarabaeidae: Melolonthinae). Bull. of the Univ. Nebraska State Mus. 11: 1-115.

27 Death Valley National o Park A .<>

~.~~--.~--~~----...... -- .. ------~--~~~~~~~~==::::~~~~~~~~~~~~~~-~~.--.--..-----~-----~-.-----~~-~------.;-.---~.--~-----.-. ------..--.--~- .. -~--.~----..~ .. -~--~--~-~~----

<$.

Figure 1. Location of Death Valley National Park In California.

28 (A)

Figure 2. June Beetle census location along the Amargosa River drainage (A) and the Saline and Eureka Valleys (B) in Death Valley National Park.

29 •

•

Scale (meter.)

o 10000 20000

Key

City/Town •

Roads June ' ~Ues Not round

Figure 3. PolyphyUa erratica survey sites along the Amargosa River • drainage. 30 N

Scale (meters)

o 10000 20000 •

Key

City/To-wn • June BeeUes .A.. Present }4

Roeds -- June Beetles A Nat Found ~

Hi&hY&Y number @ Panamint Springs

Figure 4. Polyphylla anteronivea survey sites in Saline and Eureka Valleys.

3 1 Figure 5. Polyphylla anteronivea male.

32 c!

Figure 6. Polyphylla erratica male and female.

33 A

B Figure 7. Saline Valley Dunes. (A) North side of lake with mesquite covered dunes; (B) West side of lake with creosote and open sand.

34 ' ...... ".

A

B Figure 8. Alkali flats with Distichlis grass. (A) Ash Meadows; (B ) Saratoga Springs.

35 •

Figure 9. Female Polyphylla erratica In her burrow about eight inches below the soil sutface.

36 •

•

Figure 10. Polyphylla erratica impaled on mesquite thorn by loggerhead shrike at Saratoga Springs . • 37 Table 1. Collection sites for june beetle :j5urvey. I ! Site Date Location UTM Collection Polyphylla Northing Easting Method present 1 27-IV-1995 Badwater- West Side Rd-3.9 rd ~i S jct Badwater Rd 4019556 510203 Blacklight N 2 27..:IV-1995 Badwater- West Side Rd-6.0 rd rrli S jct Badwater Rd 4016515 509876 Blacklight N 3 27-IV-1995 Badwater- Tule Springs II 4010820 510705 Blacklight N 4 27-IV-1995 Badwater-West Side Rd-14.4 rd Ti S jct Badwater Rd 4003340 510810 Blacklight N 5 27-IV-1995 Badwater-West Side Rd-18.5 rd mi S jct Badwater Rd 3997820 513905 Blacklight N 6 27-IV-1995 Badwater-West Side Rd-22.9 rd ~i S jct Badwater Rd 3991190 514553 Blacklight N 7 27-IV-1995 Badwater-West Side Rd-35.3 rd rhi S jct Badwater Rd 3979503 522538 Blacklight N 8 27-IV-1995 Badwater-West Side Rd- S jct wi~h Badwater Rd 3976920 526790 Blacklight N 9 27-IV-1995 turn-off to Saratoga Springs I 3944680 555100 Blacklight N 10 27-IV-1995 Saratoga Springs I 3948476 552491 Blacklight N 1 1 28-IV-1995 u.l Saratoga Springs I 3949066 552447 Blacklight N 00 12 28-IV-1995 Saratoga Springs I 3949081 552314 Blacklight N 13 28-IV-1995 Saratoga Springs I 3948887 551943 Blacklight N 14 28-IV-1995 Saratoga Springs I 3948636 552044 Blacklight N 1 5 30-IV-1995 Ash Meadows NWR-Crystal Springs 4030899 561644 Blacklight N 16 30-IV-1995 Ash Meadows NWR-Crystal Sprinbs 4030719 561031 Blacklight N 17 30-IV-1995 Ash Meadows NWR-nr park OffiCil1 4031083 559836 Blacklight N 18 30-IV-1995 Ash Meadows NWR-dry lake bed dl f rd to Peterson Res. 4031095 559847 Blacklight N 19 30-IV-1995 Ash Meadows NWR-Peterson Re' ervoir 4033355 558352 Blacklight N 20 30-IV-1995 Ash Meadows NWR-Peterson Re ilervoir 4033300 558332 Blacklight N 21 30-IV-1995 Ash Meadows NWR-Peterson Re~ervoir 4033262 558312 Blacklight N 22 30-IV-1995 Ash Meadows NWR-Peterson Re~ervoir 4033154 558348 Blacklight N 23 30-IV-1995 Ash Meadows NWR-Peterson Re~1 ervoir 4032931 558335 Blacklight N 24 30-IV-1995 Ash Meadows NWR-Peterson Re [ervoir 4033032 558078 Blacklight N 25 30-IV-1995 Ash Meadows NWR-Peterson Re Ilervoir 4033155 557834 Blacklight N 26 2-V-1995 Stovepipe Wells-1.2 rd mi N Hwy 1 90 4054400 494450 Blacklight N 27 18-V-1995 Saratoga Springs I 3949066 552447 Blacklight N

! Table 1 (cont) Site Date Location UTM Collection Polyphylla Northing Easting Method present 28 18-V-1995 Saratoga Springs 3949081 552314 Blacklight y 29 18-V-1995 Saratoga Springs 3948887 551943 Blacklight N 30 18-V-1995 Saratoga Springs 3948636 552044 Blacklight N 31 18-V-1995 Saratoga Springs I 3948370 552632 Blacklight Y 32 18-V-1995 Saratoga Springs I 3947889 552645 Blacklight N 33 19-V-1995 Dantes View-Wilderness area spur rd-el 4943 ft 4008465 525550 Blacklight N 34 19-V-1995 rd to Dantes View-el 4896 ft I 4008755 526177 Blacklight N 35 19-V-1995 rd to Dantes View-el 4588 ft I 4009756 527364 Blacklight N 36 19-V-1995 rd to Dantes View-el 3617 ft 4015371 529333 Blacklight N 37 19-V-1995 Furnace Creek 4033130 514499 Blacklight N 38 19-V-1995 Furnace Creek I 4033020 514500 Blacklight N 39 19-V-1995 Stovepipe Wells I 4054380 494479 Blacklight N 40 19-V-1995 Stovepipe Wells I l;.) 4054791 493786 Blacklight N 41 22-V-1995 \0 Saline Valley Dunes-west dunes al~ng Saline Valley' Rd 4069807 421825 Blacklight N 42 22-V-1995 Saline Valley Dunes-west dunes al~>ng Saline Valley Rd 4069252 421976 Blacklight N 43 22-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4068991 422106 Blacklight N 44 22-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4068628 422255 Blacklight N 45 22-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4068648 422422 Blacklight N 46 22-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4068666 422498 Blacklight N 47 22-V-1995 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067619 422980 Blacklight N 48 22-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4068081 422969 Blacklight N 49·22-V-1995 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067857 422889 Blacklight N 5 0 2 2 - V -1 9 9 5 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067978 423054 Blacklight N 51 22-V-1995 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067923 423150 Blacklight N 52 22-V-1995 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067838 423182 Blacklight Y 53 22-V-1995 Saline Valley Dunes-west dunes albng Saline Valley Rd 4067848 422869 Blacklight N 5422-V-1995 Saline Valley Dunes-west dunes al~ng Saline Valley Rd 4067867 422897 Blacklight N 55 25-VI-1995 Death Valley Junction-Jet Hwy 127/Hwy 190 4017864 552000 Blacklight N 56 25-V-1995 South of Death Valley Junction-H~y 127 nr Eagle Mtn 4006480 556000 Blacklight N 57 25-V-1995 10 mi north of Shoshone I 3993213 563506 Blacklight N

, Table 1 (cont) Site Date Location UTM Collection Polyphylla Northing Easting Method present 58 25-V-1995 Shoshone 3980821 566539 Net y 59 25-V-1995 4.5 rd mi South of Shoshone 3975975 565925 Net y 60 25-V-1995 4.5 rd mi South of Shoshone 3975975 565925 Blacklight N 61 25-V-1995 4.5 rd mi South of Shoshone 3975907 566056 Net Y 62 25-V-1995 4.5 rd mi South of Shoshone 3975907 566056 Blacklight N 63 25-V-1995 app. 4.0 rd mi South of Shoshone I 3977170 566045 Net Y 64 25-V-1995 app. 6.5 rd mi South of Shoshone I 3974675 566325 Net Y 65 25-V-1995 Hwy 127-Tecopa Hot Springs turnoff 3975916 566084 Blacklight N 66 25-V-1995 Grimshaw Lake I 3970883 569370 Blacklight N 67 25-V-1995 Hwy 127-Tecopa turnoff I' 3969695 565196 Blacklight N 68 25-V-1995 Ibex Dunes I 3949944 557493 Blacklight N 69 25-V-1995 3949254 557433 Blacklight N 70 25-V-1995 :~:~ ~~~:: i 3948503 557289 Blacklight N .f:o.. 71 25-V-1995 o Ibex Dunes 3948986 557443 Blacklight N 72 26-V-1995 Badwater 3983939 523379 Blacklight N 73 26-V-1995 Badwater 3987053 522232 Blacklight N 74 26-V-1995 Badwater-Trail Canyon 4017057 509870 Blacklight N 75 26-V-1995 Saratoga Spring 3949066 552447 Blacklight Y 76 26-V-1995 Saratoga Spring 3949081 552314 Blacklight Y 77 26-V-1995 Saratoga Spring 3948887 551943 Blacklight Y 78 26-V-1995 Saratoga Spring 3948636 552044 Blacklight Y 79 ·26-V-1995 Saratoga Spring 3948370 552632 Blacklight Y 80 26-V-1995 Saratoga Spring 3947889 552645 Blacklight Y 81 26-V-1995 Dumont Dunes I 3945037 464278 Blacklight N 82 27-V-1995 Ash Meadows NWR-Peterson Reservoir 4033262 558312 Blacklight Y 83 27-V-1995 Ash Meadows NWR-Peterson Res~rvoir 4033154 558348 Blacklight N 84 27-V-1995 Ash Meadows NWR-Peterson Res~rvoir 4032931 558335 Blacklight Y 85 27-V-1995 Ash Meadows NWR-Peterson Res~rvoir 4033032 558078 Blacklight N 86 27-V-1995 Ash Meadows NWR-Peterson Res~rvoir 4033155 557834 Blacklight Y 87 27-V-1995 Ash Meadows NWR-front enterant 4030983 556716 Blacklight Y Table 1 (cont) Site Date Location II UTM Collection Polyphylla Northing Easting Method present 88 27 - V -1 995 Ash Meadows NWR-Crystal Resdvoir 4029591 559392 Blacklight N 89 20-VI-1995 Saratoga Spring I 3948370 552632 Blacklight N 90 20-VI-1995 Saratoga Spring I 3947889 552645 Blacklight N 91 20-VI-1995 Saratoga Spring ! 3948887 551943 Blacklight N 92 21 - VI-1 995 Saline Valley-north side dunes 4070564 428210 Blacklight y 93 2 1 - V 1-1 995 Saline Valley-north side dunes 4070101 428199 Blacklight y 94 2 1 - V 1-1 995 Saline Valley-north side dunes 4069516 428134 Blacklight y 95 2 1 - V 1-1 995 Saline Valley-west side dunes 4069807 421825 Blacklight N 96 2 1 - V 1-1 995 Saline Valley-west side dunes 4069252 421976 Blacklight N 97 2 1 - V 1-1 99 5 Saline Valley-west side dunes 4068991 422106 Blacklight y 98 21 - VI-1 995 Saline Valley-west side dunes 4068628 422255 Blacklight y 99 2 1 - V 1-1 995 Saline Valley-west side dunes 4068648 422422 Blacklight N 100 21-VI-1995 Saline Valley-west side dunes +:- 4068666 422498 Blacklight N 101 21- VI-1 995 Saline Valley-west side dunes· ...... 4067619 422980 Blacklight N 1 02 2 1 - V 1-1 99 5 Saline Valley-west side dunes 4068081 422969 Blacklight y 103 21-VI-1995 Saline Valley-west side dunes 4067857 422889 Blacklight N 104 21-VI-1 995 Saline Valley-west side dunes 4067978 423054 Blacklight N 105 21 - VI-1 995 Saline Valley-west side dunes 4067923 423150 Blacklight y 1 06 2 1 - V 1-1 99 5 Saline Valley-west side dunes 4067838 423182 Blacklight N 107 21-VI-1995 Saline Valley-Inyo Mts 4098420 413419 Blacklight N 108 21-VI-1995 Saline Valley-Inyo Mts 4097786 414102 Blacklight N 109 21-VI-1995 Saline Valley-:lnyo Mts 4097261 415045 Blacklight N 1 1 0 22 - V 1-1 995 Saline Valley-north side dunes 4069446 428036 Mercury Vapor Y 111 22 - VI-1 995 Saline Valley-north side dunes 4070718 426804 Blacklight y 1 1 2 22 - V 1-1 995 Saline Valley-north side dunes 4070878 426834 Blacklight y 113 22 - VI-1 995 Saline Valley-north side dunes 4070820 427105 Blacklight y 1 1 4 2 2 - V 1-1 99 5 Saline Valley-north side dunes 4070744 428095 Blacklight y 1 1 5 22 - V 1- 1 99 5 Saline Valley-north side dunes 4070564 428210 Blacklight y 116 22-VI-1995 Saline Valley-north side dunes 4070101 428199 Blacklight y 1 1 7 22 - V 1-1 995 Saline Valley-north side dunes 4069516 428134 Blacklight y Table 1 (cont) Site Date Location II UTM Collection Polyphylla Northing Easting Method present 1 1 8 2 3 - V 1-1 995 Eureka Valley Dune 4106957 439266 Mercury Vapor N 119 23-VI-1995 Eureka Valley Dune 4106081 439329 Blacklight N 120 23-VI-1995 Eureka Valley Dune 4106116 439231 Blacklight N 121 23-VI-1995 Eureka Valley Dune 4106332 439365 Blacklight N 122 23-VI-1995 Eureka Valley Dune 4106611 439310 Blacklight N 123 23-VI-1 995 Eureka Valley Dune 4107015 440011 Blacklight N 124 23-VI-1995 Eureka Valley Dune 4107023 440156 Blacklight N 125 23-VI-1995 Eureka Valley Dune 4107066 440265 Blacklight N 126 23-VI-1995 Eureka Valley Dune 4107105 440099 Blacklight N 127 23-VI-1995 Eureka Valley Dune 4107051 439319 Blacklight N 128 1 977 Saline Valley 4062500 426000 Blacklight y 129 1 977 Saline Valley 4062650 434250 Blacklight y 130 1977 Saline Valley 4067650 423250 y ~ Blacklight N 131 15-VI-1993 Saline Valley-west side dunes· 4067850 423200 net Y 13226-V-1990 Saline Valley-west side dunes [ 4067850 423200 net Y 133 Ash Meadows NWR-Fairbanks Spting 4038383 559005 Literature Y 134 21-VI-1995 2.9 rd mi E Death Valley Jct on S~ateline Rd 4019980 556385 Net Y 135 30-VI-1 995 nr south enterance Ash Meadows ~WR via State Line Rd 4024315 562011 Net Y 136 30-VI-1995 nr south enterance Ash Meadows ~WR via State Line Rd 4024313 562733 Net Y 137 30-VI-1995 nr south enterance Ash Meadows WR via State Line Rd 4023985 561987 Net Y 138 4-IV-1986 Saratoga Spring 3947889 552645 Seen flying Y 139 30-IV-1986 Saratoga Spring 3947889 552645 Seen flying Y 140 6-V-1986 Saratoga Spring 3947889 552645 Seen flying Y 141 28-VI-1995 Big Spring outflow 4025002 562326 Net Y 14223-V-1995 Big Spring outflow 4025002 562326 Observation N 143 29-IV-1995 Big Dune 4052135 538995 Blacklight N 144 29-IV-1995 Big Dune 4052293 538226 Blacklight N 145 29-IV-1995 Big Dune 4052533 537965 Blacklight N 146 29-IV-1995 Big Dune 4050163 537685 Blacklight N 147 29-IV-1995 Big Dune 4055543 538615 Blacklight N Table 1 (cant) Site Date Location II UTM Collection Polyphylla Northing Easting Method present 1 48 29 -I V -1 995 Big Dune 4056515 539991 Blacklight N 149 29-IV-1995 Big Dune 4055901 540965 Blacklight N 150 26-V-1995 Saratoga Spring 3948320 552790 Net Y 1 5 1 22 -V 1-1 995 near west enterance to refuge 4031139 556656 Net Y 152 23-V-1995 Fairbank Spring 4038383 559005 Emergence holes Y 15323-V-1995 Indian Spring 4031570 561910 Observation N

+:­ VJ Table 2. Enzyme and buffer systems usedllfor electrophoretic analysis of Po/yphylla.

Enzyme # of E.C. # Starch/Cellulose Buffer Loci acetate

+>- AO Aldehyde Oxidase 1 1.2.3.1 Cellulose acetate TG +>- DIA Diaphorase 1 1.6. *. * Cellulose acetate TG GOT Glutamate-Oxaloacetate Transferase 1 2.6.1.1 Cellulose acetate TEB GPDH Glycerol-3-Phosphate Dehydrogenase 1 1.1.1.8 Cellulose acetate TG pH 8.5 GPI Glucose phosphate Isomerase 1 5.3.1.9 Starch CT IDH Isocitrate Dehydrogenase 2 1.1.1.42 Starch TC 7.0 MDH Malate Dehydrogenase 2 1.1.1.37 Starch TC 7.0 MPI Mannose phosphate Isomerase 2 5.3.1.8 Cellulose acetate TEM PEP Peptidase (Leucyl valine) 1 3.4.11 Cellulose acetate TG pH 8.5 6PGDH 6-Phosphogluconate Dehydrogenase 1 1.1.1.44 Starch CT PGM Phosphoglucomutase 1 2.7.5.1 Starch CT I Table 3. Allele frequencies for P. an1eronivea.

LrCUS AND SAMPLE SIZE

AO OlA GOT GPDH GPl lOH1 lDH2 MDH1 MDH2 MPIl MPl2 PEP 6PG PGM

# inds 38 28 38 38 37 167 37 34 37 37 34 38 38 37

ALLELE I A 0.00 0.07 0.00 0.00 0.00 0.00 0.00 1. 00 0.00 0.01 0.99 0.00 0.00 0.00 B 1. 00 0.25 0.00 1. 00 0.00 1. 00 0.00 0.00 0.95 0.97 0.00 0.50 1. 00 1. 00 C 0.00 0.68 1. 00 0.00 0.82 0.00 1. 00 0.00 0.05 0.01 0.01 0.00 0.00 0.00 D 0.00 0.00 0.00 0.00 0.00 00 0.00 0.00 0.00 0.00 0'1 0.00 0.00 0.00 0.00 E 0.00 0.00 0.00 0.00 0.18 0.00 0.00 0.00 0.00 0.00 0.00 0.50 0.00 0.00

~ lJl

I I . Table 4. Allele frequencies for P. errat~ca.

AO DlA GOT GPDH GPl I~Hl lDH2 MOH1 MDH2 MPIl MPl2 PEP 6PG PGM # inds 44 44 44 43 44 413 43 41 44 44 29 44 44 44 I ALLELE I A 0.07 0.00 0.00 0.00 0.03 o .!24 0.44 1. 00 0.00 0.00 0.19 0.01 0.00 0.00 B 0.30 0.73 0.00 0.92 0.00 0)76 0.08 0.00 1. 00 0.16 0.33 0.13 0.59 0.98 C 0.50 0.27 1. 00 0.08 0.96 oJoo 0.48 0.00 0.00 0.84 0.31 0.58 0.40 0.02 D 0.14 0.00 0.00 0.00 0.00 oJoo 0.00 0.00 0.00 0.00 0.17 0.02 0.01 0.00 E 0.00 0.00 0.00 0.00 0.01 0.,00 0.00 0.00 0.00 0.00 0.00 0.18 0.00 0.00 F 0.00 0.00 0.00 0.00 0.00 0.100 0.00 0.00 0.00 0.00 0.00 0.08 0.00 0.00 Table 5. Genetic variability at 14 loci for all populations of P. anteronivea and P. erratica (standard errors in parentheses).

MEAN HETEROZYGOSITY MEAN SAMPLE MEAN NO. PERCENTAGE SIZE PER OF ALLELES OF LOCI DIRECT­ HDYWBG POPULATION LOCUS PER LOCUS POLYMORPHIC* COUNT EXPECTED**

1 erratica 42.5 2.6 64.3 0.18 0.32 (1.1 ) (0.4) (0.05) (0.07)

2 anteronivea 34.1 1.6 28.6 0.13 0.11 (2.4) (0.2) (0.07) (0.05)

* A LOCUS IS CONSIDERED POLYMORPHIC IF THE FREQUENCY OF THE MOST COMMON ALLELE DOES NOT EXCEED 0.95

** UNBIASED ESTIMATE (SEE NEI, 1978)

46 Table 6. Matrix of genetic similarity and/or distance coefficients between P. anteronivea and P. erratica. Above diagonal: Nei (1978) Unbiased Minimum Distance Below diagonal: Modified Roger's Distance(Wright,1978).

POPULATION 1 2

1 erratica ***** 0.17

2 anteronivea 0.42 *****

------~~======~====~------

47 I i i I Table 7. Matrix of genetic similarity and/or dist~nce coefficients for selected Polyphylla taxa. Above Diagonal: Nei (1978) Unbiased Genetic Distance. Below Diagonal~ Modified Rogers' Distance (Wright, 1978). I I POPULATION 1 2 3 5 6 7 8 9 10 11 12 II 4 1 P.nubila ***** 0.18 0.37 0.23 0.65 0.79 0.53 0.54 0.37 0.60 0.39 1.43 Atascadero, CA II 2 P.decemlineata 0.37 ***** 0.14 0.07 0.43 0.50 0.41 0.48 0.37 0.57 0.46 1. 22 Oroville, CA II 3 P.decemlineata 0.48 0.32 ***** 0.15 0.51 0.57 0.44 0.45 0.53 0.65 0.43 0.95 Alamosa, CO I

4 P. decemlinea.ta 0.39 0.25 0.30 JI* * * * 0.51 0.61 0.41 0.38 0.39 0.64 0.47 0.91 Mt.Austin, NV I I 5 P.hammondi 0.61 0.51 0.53 p.53 ***** 0.42 0.41 0.46 0.71 0.77 0.35 1.12 Nogales, AZ .p.. II 00 6 P.hammondi 0.65 0.55 0.56 0.52 ***** 0.82 0.82 1.08 1. 30 0.84 1. 78 Medora, KS r'sa 7 P.diffracta 0.56 0.49 0.50 0.50 0.65 ***** 0.19 0.33 0.58 0.50 1.12 Globe, AZ 1['49

8 P.diffracta 0.55 0.51 0.48 0.51 0.63 0.36 ***** 0.19 0.40 0.35 0.89 Kanab, UT Ir'46 9 P.diffracta 0.51 0.50 0.56 0.64 0.73 0.48 0.38 ***** 0.34 0.38 1.13 Navajo Mt., UT 50 10 P.devestiva 0.60 0.57 0.58 It' 0.64 0.75 0.58 0.49 0.49 0.27 1. 42 I .59 ***** Bruneau Dune,ID

11 P.barbata 0.50 0.53 0.50 0.48 0.66 0.54 0.46 0.51 0.43 ***** 1. 36 Mt. Herman, CA II ,52 I 12 Polylamina 0.76 0.71 0.65 :0.65 0.71 0.80 0.70 0.64 0.73 0.76 0.75 ***** Bay Co., FL Table 8. Estimated acreage for PoJypbylla erratica.

Ash Meadows/Carson Slough: 2,400 acres Tecopa/Shoshone: 1,100 acres Saratoga Springs/lower Amargosa R. 1,250 acres

Total: 4,750 acres

49 Table 9. Current land status for areas with june beetles (1995).

Po/yphyl/a anteronivea Po/yphyl/a erratica

EASTING NORTHING lAND OWNER EASTING NORTHING LAND OWNER

423182 4067838 NPS 552447 3949066 NPS 423150 4067923 NPS 551943 3948887 NPS 426000 4062500 NPS 552044 3948636 NPS 428210 4070564 NPS 566056 3975907 BlM 428199 4070101 NPS 566045 3977170 BlM 428134 4069516 NPS 566325 3974675 BlM 422106 4068991 NPS 552790 3948320 NPS 428036 4069446 NPS 552314 3949081 NPS 426804 4070718 NPS 552632 3948370 NPS 426834 4070878 NPS 566539 3980821 BlM 427105 4070820 NPS 556385 4019980 BlM

~"-"-,-" " _ ~t20R095 4l)LO]~~ _~,~S 56200] 40L4l'LS ~----"----"-""---""""-""-- 434250 4062650 NPS 562733 4024313 BlM 423250 4067650 NPS 561987 4023985 BlM 423200 4067850 NPS 552645 3947883 BlM 422255 4068628 NPS 558312 4033262 USFWS 422969 4068081 NPS 558335 4032931 USFWS 557834 4033155 USFWS 556716 4030983 BlM 559005 4038383 USFWS 562326 4025002 BlM 556656 4031139 USFWS

50 The Pan-Pacific Entomologist

Vol. 54 .JANUARY 1978 No.1

Studies in the Coleoptera of Western Sand Dunes

L Five New Polyphylla Harris

(Scarabaeidae)

Alan R. Hardy and Fred G. Andrews'

Insect Taxonomy Laboratory. California Dept. Food and Agric .. 1220 "N" Street. Sacramento. 95814

Recent (nvestigations of sand dunes in the Western United States have uncovered many species of Coleoptera which are restricted to one or a few dune systems (see Hardy 1971, 1973, 1974,1976, 1977; Hardy and Andrews 1974; Howden and Hardy 1971; Gordon 1974; Gordon and Cartwright 1977). The reasons for this interesting degree of endemism or reduced distribution will be discussed in detail else­ where. One of the most commonly encountered groups adapted to sand niches is the family Scarabaeidae, with many subfamilies re­ presented. Below are described five species of Polyphylla taken in the -===~=~======~le0tH'Se~0·f~~easear,Gh~Qr:t~sar:td~dunes;...,·=,-,.il'''t::.~.,~.c''''-LJ:;';:>''~'''''~~~,!-J~ •.. u ....~_ ...... _ ..... ~ scribed from this niche is PolyphyJla devistiva Young.

Polyphylla avittata Hardy, new species (Figs. 1,2.11)

Holotype male: Length 22 mm, width 7.5 mm. Integumentary color of head piceous; thorax rufous: elytra rufo·testaceus; ventral surfaces (except abdomen), legs rufo·testa­ ceo us; abdomen rufo-piceous. Clypeal margin reflexed, outer apical angles well defined; reflexed margins with dense, close white scales (Fig. 11); few scales medially on clypeal disc, along frcnto-clypeal suture, scattered over front. Ctypeus, front with close to con­ tiguous large punctures, punctures with single scale or suberect halr.'Vertex impunciate, glabrous, shining. Prothorax with close to contiguous punctures, with scales or hairs as above. Scales denser on midline and medio-Iaterally; prcthorax appearing trivittate. Elytra lacking well defined punctures of thorax and head, but sparsely to densely covered with scattered scales, giving appearance of random clumping, without vittae, scales slightly smaller than on prothorax. Pygidium uniformly with scales and short recumbent hairs.

'The authOrs acknowledge the financial assistance of the OHlce of Endangensd Species, U.S. Fish and Wildlife Service. under contract No. 14-16-0008-966 for pan Ions of the fieldwork.

The Pan-Pacific Entomolog.st 5-1:1-8. January 1978 2 THE PAN-PACIFIC ENTOMOLOGIST

Antennal club (measured in straight line, not along curve) twice length 01 three basal segments (Fig. 11). Apical segment 01 maxillary palp slightly less than length 01 two basal segments: flattened on dorsal surlace. Anterior tibia bidentate. Ventral surface 01 thorax with long grayish hairs. Variation in paratypes: Length 19.5·22 mm, width 7·7.5 mm. Most paratypes (9 of 12) have a lalnt indication 01 a third anterior tibial tooth, but this is never developed into more than just a slight flange on the tibial edge. Paratypes agree well in other respects. Female: Unknown. The name relers to the splotched, rather than vittate, elytra Holotype male. (CAS# 1309B) and 12 male paratypes: Utah, Washington Co., 6 mi. S. Hurricane. Hurricane Dunes, VII·15·1975, Fred G. Andrews, Alan R. Hardy, collected at blacklight. (Deposited in USNM, CAS, Howden, CDA and Hardy.) Diagnosis: P. avittata keys to either hammondi or the diffracta com­ plex (couplet #4) in Fall's Key (1923:34), depending upon interpreta­ tion of the bidentate vs. tridentate condition of the anterior tibiae. When scaled dorsally, P. hammondi is unmistakably vittate, unlike the "blotched" elytral pattern of avittata (Figs. 1, 2). P. avittata is also smaller than hammondi. In the diffracta complex (see Young 1976), avittata keys to diffracta, but differs in the "blotched" verses vittate elytra (diffracta occasionally lack vittae, but are never "blotched"). The "blotched". elytra of avittata resemble the new world species vari%sa or nubi/a, or several old world speCies, such as fullo, al­ though not as densely scaled as the old world species. P. avittata differs from nubi/a by the lack of erect elytral hairs which are present in nubi/a. From vari%sa, avittata may be easily distinguished by the dorsal squamae, which are thin and hair-like in vari%sa, but broad and scale-like in avittata, and the sharp clypeal corners of avittata compared with the round clypeal corners of varioiosa. Remarks: This species was collected at blacklights placed upon the dunes. The area is also known as the "Warner Valley Dunes" by local residents, and is visible to the south from State Highway 17, to ~._~_... the east of the Virgin River. The dunes are evidently derived from ·~~======i===~N~a~va;;J~·o~S;"a';n;a"Ts:;'t~o~n~e;,~a~na·are'th·ele·dl:otoMypical=0f4he=san€l=an€l=r,QCkS . .. 1.__ ...... _ ...... _.. . 'in that part of Utah. Vegetation on the dunes is primarily Artemesia filiform/s. while surrounding vegetation is Larrea divaricata Cav.

Polyphylla anteronivea Hardy, new species (Figs. 3, 4, 12)

Holotype male: Length 26 mm, width B.B mm. Integumental'{ color 01 head piceous to rulo.piceous; thorax rulo·piceous; elytra, ventral body surfaces. appendages testaceous. Clypeal angles well dell ned (Fig. 12); reflexed margin, disc. Iront with close to contiguous round to oval. cream to white scales. Front with lew semierect cream colored· bristle­ like hairs. Vertex smooth, glabrous, thorax with close to contiguous to densely overlap-

Figs. '·10, Po/yphy/la new species. Figs. 1 and 2. P. avlITara. Figs. 3 and 4, P. anreromvea. Figs. 5 and 6, P. erratica. Figs. 7 and B. P. pottsorum. Figs. 9 and 10. P. mona­ nansensls. All to same scale. VOL 54, NO.1, JANUARY 1978 3

./~~:;. • '7':"' • If_ ~~. 4 THE PAN-PACIFIC ENTOMOLOGIST

ping oval white scales, which obscure virtually entire thoracic integument. Few scattered erect hairs at anterior margin of prothorax. Scutellum contiguously covered with oval white scales. Elytra anteriorly, along suture. lateral margin and apically, with scattered to contiguous oval white scales. Ventral thoracic segments with sparse, medium, gray to white erect hairs. less dense than in most PolyphylJa. Apical segment of maxillary palp short (.7 times length of basal two segments), stout (length 3.2 times width), with small, slightly concave area dorsally. Antennal club 2 (linear measurement) or 2.8 (along curve) times length of basal segments. Anterior tibiae strongly tridentate (Fig. 12). Variation in paratypes: Length 21·29 mm. width 7·10 mm. Except for specimens which have been abraded or greased, the only notable difference is in the elytral scalation, which may be reduced, or may not appear vitt.ate. Female: Unknown. The name refers to the snow covered appearance of the anterior portion at the insect (Fig. 4). Holotype male (CAS ~13097) and two male paratypes: California Inyo Co., Saline Valley Dunes, VI·13-1976 D. Giuliani call. Thirty·three additional paratypes. all Saline Valley, as follows: Dunes Blackllght. ~8, VIl-B·1976 D. Giuliani (3); Dunes ~11. D. Giuliani VIlf.10-1976 (1); VI·'·1974 N. Rulien (1); Dunes. VI-10-1973. J. M. Cicero (21); Dunes, V·18·1973, 'J. M. Cicero (6); Lake. "·18·1973, J. M. Cicero (l).IDeposited in USNM. CAS, CDA, Cicero, How­ den, Nelson, Potts. Rulien.) Diagnosis: P. anteronivea. by virtue of the strongly tridentate anterior tibiae (Fig. 4), would key to couplet two in Fall's 1928 key, which in­ cludes the species cavifrons and hammondi. P. squamiventris Cazier would also run to that portion of the key. P. cavifrons has the antennal club less than twice as long as the basal segments, hair-like dorsal scales, less acute clypeal angles, and a more densely pubescent ventral thoracic area than anteronivea. P. hammondi has a vittate thorax. the anterior angles of the clypeus are narrower than the lateral margins (clypeus of anteronivea is widest at angles), and the front and clypeus are more pubescent than in anteronivea. (It should be noted

.-"-~"-'---.-. that although both of the keys by Casey and Fall use the tridentate ==~ aMerier=tihiae oJ hammondi as significant, in series, Arizona speci­ mens range from strongly triaerira:te-tc)~eakly=t·rictel'ltate=to.. bjd.. entate. There is, additionally, an undescribed species near hammondi from central Mexico with strongly tridentate anterior tibiae.) P. squamiven­ tris has an antennal club less than 1.5 times the length of the basal segments, a pronounced pronotal vittae, and unimpressed maxillary palp. which distinguishes this species from anteronivea. P. anter­ onivea is easily recognized at first glance by the unique vestiture (Figs. 3, 4). 0. anteronivea is related to the following species.

Polyphylla erratica Hardy, new species (Figs. 5, 6, 14)

Holotype male: Length 26.5 mm, width 9.7 mm. Integument at head rufous, except piceous vertex. anterior, lateral clypeal margins and angles. Thoracic integument rufous. Elytra, ventral surfaces, pygidium. appendages rufo·testaceous; antennal club testa­ ceous. Clypeus quadrate, margins strongly retlexed, anteriorlv bisinuate. outer angles sharp, nearly spinose (Fig. 14). Clypeal disc with close to contiguous oval white scales; VOL. 54, NO.1, JANUARY 1978 5

Figs. 11·15, Polyonvlla. new species. Fig.11. P a,'mara Fig. 12, P. anteron/vea. Fig.13, P. ;:JOtlsorum Fig. 14. P errallca. Fig. 15, P mor.a·~a~sensls. Fig.11·14 to same scale.

front with close to contiguous to "shingled" oval white scales, with semierect to erect testaceous hairs; vertex shining, impunctate. glaorous. Prothorax with close, to "shin· gled" oval white scales, slightly denser medially. medio·laterally, forming trivittate pro· thoracic pattern normal for genus: few erect to semlerect white to testaceous hairs at anterior margin. Scutellum "shingled" with oval white scales. Elytra with oval white scales, which are dense and "shingled" anteriorly: becoming scattered laterally, pos· terlorly; forming broken "vittae". Elytra lacking hairs. Pygidium with close white oval scales. Ventral surfaces nearly completely covered (except sutures) with oval white scales. Thorax ventrally with erect white hairs. Antennal club 2 (linear) to 3.3 (along curve) times length basal segments. Antenor tibiae strongly tridentate. Variation in para types; Length 23·27 mm. width 8·9.5 mm. Most pronounced difference is in the squamal vesllture, which may be nearly aosent (aorasion?) to the semivittate con­ dition described above (Fig. 6). Female: Unknown. The name refers to the erratic elytral pattern. Holotype male, (CAS #13099) and three male paratypes: California, San Bernardino Co. [Death Valley National Monument], Saratoga Springs IV·1974, DUSk, D. Giuliani collr. 62 6 THE PAN·PACIFIC ENTOMOLOGIST additional paratypes as follows: California, Death Valley, Inyo Co. [sic.I, Saratoga Springs VI·12·1964, Fred G. Andrews (3); California, Death Valley, V·27/29·1955 Belkin et al., (57); California. Inyo Co .. Tecopa VI·17·54, Belkin and McD. (1); California. Inyo Co., Tecopa VII· 11-1953 (1). (Paratypes in USNM, LACM, CAS. Howden, Hardy.) Diagnosis: Po/yphyl/a erratica, because of the tridentate anterior tibia, would key to couplet two in Fall's key. Species which also have this tridentate characteristic are P. hammondi, cavifrons, squamiven­ tris, and anreronivea. P. hammondi has shorter antennal club (1.5 to 2 times length of basal segments, erratica usually greater than 2 times [Fig. 14]), less pronounced clypeal angles, both white and yellow pro­ thoracic scales (erratica has white only), testaceous (as opposed to white) ventral hairs, and usually at least some hairs on clypeus and thoracic disc behind anterior margin (erratica has hairs confined to front and anterior prothoracic margin). P. cavifrons lacks the clypeal angles, has an anteriorly expanded clypeal shape (quadrate in erra­ tical a smaller antennal club (less than twice length of basal seg­ ments) and fine hair·1 ike scales (oval in erratica). P. squamiventris has a short antennal club (shorter than cavifrons), and an unimpressed maxillary palp (small concave area in erratica). P. erratica is evidently most closely related to the preceding species. P. anteronivea has slightly less well developed clypeal angles, an apically broadened clypeus (quadrate in erratica), less elongate and apically less pointed scales, and an anterior elytral impression which results in a trans­ verse carina from the scutellum towards the humeral area. Addition­ ally, anterom'/ea lacks any trace of a pattern or vittae in the posterior half of the elytra (erratica usually displays at least some evidence of fragmented vittae posteriorly). Remarks: Some specimens from Saratoga Springs were taken at dusk flying low over Distich/is grass in the salt encrusted bottom of the Amargosa River. These specimens were not attracted to black­ light after dark.

Polyphylla pottsorum Hardy, new species

(Figs. 7, 8, 13)

HOlotype male: Length 20 mm, width 7 mm. Integument of head, prothorax. scutellum piceous; elytra antennal club. ventral body surfaces pale testaceous; pygidium, basal segments of antennae, legs testaceous. Clypeus widened apically, anterior margin reflex­ ed. nearty linear. angles sharp, not produced above anterior margin (Fig. 13). Clypeus, front with elongate, pointed white scales. intermixed with semierect gray hairs. Pro­ thoracic disc With few erect hairs close to anterior margin; with elongate white scales U closely to densely 'over suriace; scales intermixed with recumb'ent white scale-like hairs I' medially; Prothorax appearing faintly trivittate (Fig. 8). EIYlra sparsely covered with small. elongate, pointed white scales. faintly vittate at elytral base. Pygidium sparsely scaled. Antennal club approximately 2 (linear) to 2.5 times (along curve) length of basal segments (Fig. 13). Ventral abdominal segments glabrous basally near suture, densely scaled api. cally; covered with sparse to dense, long, fine pubesence. Ventral suriace of thorax den· VOL. 54, NO.1, JANUARY 1978 7

sely covered with long, white hairs, except glabrous patch at midline of metasternum. Anteriortibia distinctly tridentate, not as strongly as previous species (Fig. 13). Variation in paratypes: Length 16·22 mm, width 5.5-7.6 mm. There is some variation in the development of the third (basal) tibial tooth, but in all cases, there is clearly a third tooth present. Variation in scale density varies the elytral appearance from that of gla­ brous (although scales are present) to distinctly vittate, with vittae extending nearly to elytral apex. Female: Unknown. The name is in recognition of the contributions of Mr. and Mrs. R. W. L Potts, who first drew my attention to this species. Holotype male (CAS #13101) and 145 male paratypes: Texas, Ward Co., Monahans Sand­ hills State Park, VI·6/7- 19n, Potts and Potts collrs.; 184 additional male paratypes, same locale, as follows: V-28-1975, Potts and Potts (29); VII-19n, R. Lenczy (S): VII-15/16-1975, dead at light, G. H. Nelson (36); VII·23/24-1976, ultraviolet light, G. H. Nelson (III). (Depo­ sited in CAS, USNM, Lenczy, Nelson, Potts, Hardy.) Diagnosis: P. pottsorum is another of the species with tridentate anterior tibiae (hammondi, cavifrons, squamiventris, anteronivea and erratica, above). From all of these species except squamiventris, pott­ sorum may be readily separated by the small size (all other species greater than 22 mm in length). P. pottsorum may be distinguished from the closely related squamiventris, by the longer antennae (1.5 times basal segments, in squamiventris vs. 2 times, or more, in pott­ sorum), presence of clypeal setae, lighter elytral color, impressed maxillary palp, longer, thinner squamae on prothorax, and testaceous . .1 legs.

Polyphylla monahansensis Hardy, new species (Figs. 9, 10,15) Holotype male: Length 26 mm, width 8.5 mm. integumentary color testaceous to rUlo­ testaceous except for piceous vertex, ocular canthi, clypeal suture, lateral clypeal margins. Apical clypeal margin reflexed; laterally, medially angulate. Reflexed clypeal margin, lateral marg ins, ocular canthi, lateral portions of front, vertex with close, recumbent elongate white scales. Medial portions of clypeus, front with few scattered I coarsely punctured, each puncture with a recumbent scale or single long erect testa­ ceous hair. Punctures finer. closer anteriorly at midline. Elytra noticably vitiate (Fig. 10), with scales smaller than thoracic scales. Pygidium without scales, with numerous short, semirecumbent testaceous hairs. Apical pygidial margin reflexed. Antennal club nearly 3 times length of basal segments (straight line mflasure). Maxillary palp cylindrical in cross ... section, without flattened area. Anterior tibia bidentate. Thorax ventrally with dense, erect .... testaceous hairs. Ventral abdominal segments with sparse, recumbent, small white scales. Variation in paratypes: Length 21-27.5 mm, width 7.5-9.5 mm. Most conspicuous variation from the description above is in the shape of the reflexed anterior clypeal margin, which may lack the medial clypeal angulation, and have the lateral angles more rounded; and the anterior tibial teeth, which may be bidentate (as abOve) to strongly tridentate. The thoracic vittae may be more clearly developed than the type, and the elytral vittae occasionally may be faint, but are, in all specimens examined, clearly discemable throughout their length. Female: Unknown. The name refers to the type locality, Monahans, Texas. Holotype male (CAS #13100) and 39 male paratypes: Texas, Ward Co; Monahans Sandhills State Park, VI-&7-19n, Potts and Potts. 98 additional paratypes, same ·8 THE PAN-PACIFIC ENTOMOLOGIST

!ocale; except, VII·1977, R. Lenczy (8); VII·28-7S, Potts and Potts (23); VII·1S/16-7S, G. H. Nelson (S); VII-23124·1976, G. H. Nelson (62). (Deposited in CAS, USNM, Howden, Nelson, Lenczy, Potts, Hardy.) Diagnosis: P. monahansensis keys to hammondi or diffracta. depending upon the dentition of the anterior tibiae (bidentate or tri­ dentate). From hammondi. monahansensis may be distinguished by the longer antennal club (length 2.75 times or less length of basal segments in hammondi. 2.75 times or greater [usually ± three times] in monahansensis), position of the tee.th on the anterior tibiae (all more apical in monahansensis); and suberect to erect long hair over disc of prothorax. From diffracta. monahansensis is distinguished by the long antennal club (2 times or less in diffracta). evenly cylindrical maxillary palp (flattened dorsally in diffracta) and apically more expanded clypeus. Polyphylla speciosaCasey

Polyphylla rufescenta Tan ner 1928:276.N EW SYNO NYMY We have recently seen a series of specimens taken at Zion National Park, which display in every respect the characters described by Tanner for his unique male. The "edentate" condition is believed to be derived from wear, as is the case in some individuals in the series above.

Literature Cited Fall, H. C., 1928. A Review of the Genus Pofyphylfa. Proc. Entomol. Soc. Wash. 30(2): 30-35. Gordon, R. D~ 1974. Additional Notes on the Genus Glaresis. Proc. BioI. Soc. Wash. 87(12): 91 ·94. Gordon, R. D. and O. L. Cartwright, 1977. Four New Species of Aegialia (s.str.) (Coleoptera: Scarabaeidae) from California and Nevada Sand Dunes. J. Wash. Acad. Sci.: 67(1): 42·48 . .~=~======,H:!.!a~rd!tY~,,f,A.';J:!R~ .• ~19~7~1~.~T~h~e~N:r;0rth American Areodina with a Description of a New Genus from . . California. pan~Paclf~ortrol~7(3):=Q3S-242:o=.======~ Hardy, A. R., 1973. A New Species of Phobetus. Pan·Pacific Entomol. 49(2): 127·131. Hardy. A. R., 1974. A New Species of Cyc/ocepnala Latreille from Galifornia Sand Dunes. Pan·Pacific Entomol. SO: 160·161. Hardy. A. R., 1974. Two New Species of Pseudocotalpa Hardy. Pan·Pacific Entomol. 50: 243-247. Hardy, A. R., 1974. Observations on the Mating Behavior of Pseudocotalpa giulianii Hardy. Coleopt. Bull. 30(3): 301·302. Hardy, A. R., 1976. A New Species of Anomala Samouelle from California Sand Dunes. Coleopt. BUll. 30(4): 36S·367. Hardy, A. R~ 1977. Observations on Some Rare Scarabaeidae Mainly from California. Coleopt. Bull. 31(1): 91·92. Hardy, A. R. and F. G. Andrews, 1974. Observations on Megasoma with Behavioral Notes on Some Lameilicorn Beetles Associated with Sand Dunes. Pan·Pacific· Entomol. SO(2): 124·128. Howden. H. F. and A. R. Hardy, 1971. Generic Placement and Adult Behavior of the Genus Lepronoplia. Proc. Entomol. Soc. Wash. 83(3): 337·341. Tanner, V. M., 1928. The Coleoptera of Zion National Park, Utah. Ann. Entomol. Soc. Amer. 21: 269·290. Young, R. M~ 1976. Polyphylfa Harris in America, North of Mexico. Part I: The Diffracta Complex. Trans. Amer. Entomol. Soc. 93: 279·318.

Ronald M. Young.

A Monograph of the Genus Polyphylla Harris . A . . In m lCO______(Coleoptera: Scarabaeidae: Melolonthinae)

BULLETIN OF The University ~f Nebraska State Museum VOLUME 11. NUMBER 2 FEBRUARY. 1988 \.

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A MONOGRAPH OF THE GENUS POLYPHYLLA I 75

species cluster not having elytral hairs it is the only (length 3.2 times width), with small, slightly concave species exceeding 22 mm length which is deep brown area dorsally. Antennal club 2 (linear measurement) to black and devoid of formed vittae. or 2.8 (along curve) times length basal segments. lale The paratypes range in overall length from 22.1- Anterior tibiae strongly tridentate." (Hardy and An­ :lIes 23.3 mm, in width from 10.1-11.0 mm. Elytrallength drews 1978). ype ranges from 14.3-15.7 mm. Overall they show very "mi- little variation in external morphology. In two para­ Female.-Unknown. types the pronotal disc is devoid of squamae, having only a light covering of recumbant hairs. In another Distribution.-Specimens examined: the holo­ ed. the elytral squamae are missing centrally, giving the type male and one paratype male. Known only from lwn animal a simple deep brown appearance. Protibiae the Saline Valley Dunes (Fig. 29). 3ad vary from barely to deeply tridentate. Once seen, this MAY (7), JUNE (25), JULY (3), AUGUST (1). ral­ distinctive species should be easy to recognize. The nc­ spotted or speckled aspect of the elytra (hence the Biology.-Directly associated with a sand dune se; name stellata) , with each spot being a single isolated complex and taken at blacklight. rior white scale, is most unique. ilar Remarks.-P. anteronivea is distinguished very ing easily from all its congeners by the snowy white pron­ :In- otum and scutellum; the covering composed of solid 3p­ THE HAMMONDI COMPLEX white scales. Hardy and Andrews (1978) note that m­ overall length ranges from 21-29 mm., and width from I at Polyphylla anteronivea Hardy and Andrews 7-10 mm. They also record that elytral scalation may ~e- (Figs. 8a. 29, 68) be reduced or not appear vittate. 3S, This species is very closely related to P. erratica, ng Polyphylla anteronivea Hardy and Andrews 1978: 2. so closely in fact that I doubt that both names rep­ lit- (Holotype male CASC 13097. ThirtY-five male par­ resent biological species. This matter is discussed atypes in CASC, Ca. Dept. Agriculture, USNM and more fully under P. erratica. Jr- private collections of Cicero, Howden, Nelson, ird Potts, Rulien. Type locality: CALIFORNIA: Inyo Co.: Ja __. .ol~~=~S~~,!!lin~e~Va~I~le~'i~g~~~_n~_~~~)~. ~-====-~-~~=··~·-=··~-=====~~PolypfiYlla bfdwnae young------_· g- (Figs. 8b, 30, 69--70) e, Holotype Male.-"Length 26 mm, width 8.8 mm. Jr- Integumentary color of head piceous to rufo-piceous; P. brownae Young 1986: 47. (Holotype male ANSP :s, thorax rUfo-piceous: elytra, ventral body surfaces! 8395, paratype males CASC, MCZC. Type locali­ e, appendages testaceous. Clypeal angles well de­ ty: ALABAMA: Washington Co.: Calvert). ~h fined; reflexed margin. disc, front with close to con­ tiguous round to oval. cream to white scales. Front Holotype Male.-Long, narrow. Length 29.3 mm, with few semierect cream colored bristle-like hairs. width 12.1 mm. Elytra light brown, head and prono­ Vertex smooth, glabrous, thorax with close to contig­ tum deeper, richer brown. Clypeus moderately con­ uous to densely overlapping oval white scales, which cave, lateral margins contracted for posterior one­ obscure virtually entire thoracic integument. Few fourth; surface entirely, deeply punctate, each punc­ scattered erect hairs at anterior margin of prothorax. ture giving rise to a broad, yellowish, suberect hair, Scutellum contiguously covered with oval white hairs becoming erect in posterio-lateraJ comers. Head scales. Elytra anteriorly, along suture, lateral margin with large, dense, shallow punctures, covered with and apically, with scattered to contiguous oval white widely spaced, erect hairs throughout, hairs becom­ e scales. Ventral thoracic segments with sparse, me­ ing shorter and more concentrated along epistomal e dium, gray to white erect hairs, less dense than in suture; scales lacking. Antennal club one and one­ 11 most Polyphylla. Apical segment of maxillary palp third longer than combined length of head and cly­ e short (.7 times length of basal two segments), stout peus, distal one-third gradually curved outwards.

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78 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Biology.-A species of the true xeric desert, often Broad, robust, much wider posteriorly. Length 26.5 on add attracted to lights. mm., width 8.5 mm. Pronotum deep reddish-brown, work 0 elytra medium brown. Clypeus, head, and pronotum do not Remarks.-P. cavifrons is distinguished from all totally devoid of squamae except for a few minute, tionsa others by its southwestern distribution in combination white scales in posterior pranotal angles: clypeus, labels with its pruinose. avittate appearance. It should not head, and pronotum with extremely deep, dense " replacE be confused with samples of small, avittate P. ham­ punctures, nearly rugose. Elytra unicolorous brown labelle mondi, which have deeply yellow elytral squamae. to the naked eye except for a few very widely scat­ As f- tered scales, scales more concentrated lateral to hu­ and P. , meral umbones and posterior to apical umbones. Py­ readily Po/yphy/la erratica Hardy and Andrews gidium with a sparse covering of minute, white tellum (Figs. ad, 29, 72) squamae and semierect, yellowish, stout hairs. combir occur i Po/yphy/la erratica Hardy and Andrews 1978: 4. (Hol­ Paratypes. (Variation).-Length 23-27 mm, width onivea otype male CASC 13099, and 65 male paratypes 8-9.5 mm. Scaled vestiture on the pronotum and ely­ P. erra, in CASC, LACM, USNM and private collections tra varies from nearly absent to semivittate. It is noted early st Hardy, Howden. First known female here discov­ in the above description of the holotype that the oval no col ered. homotype, UCDC. Type locality: CALIFOR­ white scales are closely enough packed on the pron­ heavily NIA: San Bernardino Co.: Death Valley National otum, scutellum, and anterior elytra to be character­ tellum, Monument, Saratoga Springs). ized as "shingled." This character brings P. erratica gradin£ " very close indeed to P. anteronivea. onivea Holotype Male.-"Length 26.5 mm, width 9.7 mm. is so f Integument of head rufous, except piceous vertex, Distribution.-Specimens examined, 71. This pearan anterior, lateral clypeal margins and angles. Thoracic number includes the holotype, 55 paratypes, and 15 two wr. integument rufous. Elytra, ventral surfaces, pygi­ homotypes here designated. The homotypes include ~ lection dium, appendages rufotestaceous; antennal club tbe QI1L}( kI1OWI1JemaLe,~d~tscrjbcSJ:I~hoJLe*aDoJbejir;sLo-o-o_o_

126.5 on additional specimens collected during his contract Polyphylla hammondi leConte -tp mwn, :,.. work on the fauna of western sand ~u~e habitats. I (Figs. 8e, 8f, 30-32, 73-75, Table 4) tum ... do not know if he had seen my material In the collec- Polyphyl/a hammondi LeConte 1856: 228; LeConte inute, tions at Davis, California. I have now remo~ed all type 1863: 39; Crotch 1873: 61; Horn 1881: 73; Hen- peus, labels from my original series of 14 specimens and shaw 1885: 92; Bates 1888: 215; Casey 1891: 18; iense replaced them with homotype labels; I have also so Dalla Torre 1912: 259; Casey 1914: 325; Leng lrown labelled one additional male in the AMNH. 1920: 257; Dawson 1922: 118; Fall 1928: 31; scat- As Hardy and Andrews (1978) noted, P. erratica Kuntzen 1933: 463; Leng and Mutchler 1933: 39; to hu- and P. anteronivea are closely related. They are most Cazier 1938: 163; Blackwelder 1939: 53; Cazier 3. Py- readily separated by the snowy-white pr~notum, sc~- 1939: 200; Brown 1940: 185; Cazier 1940: 135; white tellum and anterior elytra of P. anteromvea used In Blackwelder 1944: 227; Blackwelder and Black- combination with the collecting locality. Both species welder 1948: 33; Young 1967: 284; Young 1972: occur in Inyo Co., California (Fig. 29), with P. anter- 31; Blackwelder and Arnett 1974: R30.68; Hardy width onivea recorded from the north (Saline Valley), and and Andrews 1978: 2; Lago 1979: 63; Young 1986: j ely- P. erratica from the south (Death Valley). At this very 49. (Holotype male and four paratype males MCZC 10ted early stage in our knowledge of these demes: we have 3792. Type locality: KANSAS: Fort Riley). . oval no collecting records between these POints. The Polyphyl/a diffusa Casey 1914: 329. (Holotype and Jron- heavily scaled (in some specimens) pr?no~um, scu- paratype males USNM 35622. Type locality: ARI- .cter- tellum, and anterior elytra of P. erratlca IS clearly ZONA) . v. atica grading towards the solid white covering of P. anter- Polyphyl/a latifrons Casey 1914: 340. (Holotype male ;:, onivea. Though I have not seen any P. erratica which USNM 35637. Type locality: NEW MEXICO: Je- I; is so fully scaled as to give an opaque white ap- mez Springs). NEW SYNONYMY. This pearance, there certainly are gradations between the Polyphylla oblita Casey 1914: 326. (Holotype and two d15 two which can only be clearly identified by the col- paratype males USNM 35618. Type locality: TEX- lude , lection locality. In his descriptions of these two spe- AS: near EI Paso). i first cies, Hardy did not extract aedeagi. I have done so Polyphyl/a oblita impigra Casey 1914: 326. (Holotype also and find no differences considered significant. In sum, and two paratype males USNM 35619. Type lo- we have at this writing, insufficient data to decide cality: NEW MEXICO). ~ino whether these two names represent valid, somewhat Polyphyl/a oklahomensis Hatch 1926: 145. (Holotype Jnu- cryptic species, or a single, perhaps clinal, taxon. As female originally in Hatch's personal collection. As the nomenclatural history of this genus is one of ex- of late 1985-early 1986, it is not.iQJ~~_~JL~<:;!i..Q.!:1~ __ ._. lnks~---o ~ --,. ~~~tre~m...... ,e~J.agmenta.ti.QD~-'JJa\!(e_triEl:dJb.rou.gb_ocuUbis~WQJ:k'~~~or the Oniversity Of ORlahoma [personal commu- . to be conservative with names. In this case, for now, nication, H.P. Brown], Oregon State University I believe these two names are best left standing as [personal communication, G.L. Parsons], Wash- we do not have enough field data to justify a change. ington State University [personal communication, ded Hardy and Andrews (1978) distinguish P. R.S. Zack], CASC [personal communication, N.D. } a~ter- lere onivea from P. erratica as follows: "P. anteromvea Penny] or USNM (personal communication, R.D. the has slightly less well developed clypeal angles, an Gordon]. Not located. Type locality: OKLAHOMA: ese r;,.. apically broadened clypeus (quadrate in erratica), less Choctaw Co., 4 mi. S. of Grant along Red River, ark ~ : elongate and apically less pOinted scales, and an an- on grass). . terior elytral impression which results in a transverse Polyphyl/a pimalis Casey 1914: 330. (Holotype and carina from the scutellum towards the humeral area. paratype males USNM 35623. Type locality: ARI- lOS, ... hol- Additionally, anteronivea lacks any trace of a patt~rn ZONA: probably southern). 10S- , of vittae in the posterior half of the elytra (erratlca Polyphyl/a proba Casey 1914: 329. (Holotype male lme usually displays at least sonie evidence of frag- USNM 35621. Type locality: ARIZONA). ~M. mented vittae posteriorly)." Polyphyl/a rufescenta Tanner 1928: 276. (Holotype rip- male BYUC. Type locality: UTAH: Saint George). iy's NEW SYNONYMY. led