Denver Museum of Nature & Science Annals

DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER MUSEUMDENVER OF NATURE & SCIENCE ANNALS • NUMBER 5 JULY 15, 2014 NUMBER 5, JULY 15, 2014

WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS Denver Museum of Nature & Science Annals (Print) ISSN 1948-9293 2001 Colorado Boulevard Denver, CO 80205, U.S.A. Denver Museum of Nature & Science Annals (Online) ISSN 1948-9307

The Denver Museum of Nature & Science inspires curiosity and excites minds of all ages through scientific discovery and the presentation and preservation of the world’s unique treasures.

Cover photo: Scott Elias, Royal Holloway University of London (left), and Aaron Spriggs, DMNS collections assistant, excavating site 49, the “Twisty Trench”, at the Ziegler Reservoir for insects.

The Denver Museum of Nature & Science Annals is an Frank Krell, PhD, Editor-in-Chief open-access, peer-reviewed scientific journal publishing Editorial Board: original papers in the fields of anthropology, geology, James Hagadorn, PhD (subject editor, Paleontology and paleontology, botany, zoology, space and planetary Geology) sciences, and health sciences. Papers are either authored Pleistocene Dung Beetles from MIS 5 Bridget Coughlin, PhD (subject editor, Health Sciences) by DMNS staff, associates, or volunteers, deal with DMNS at Ziegler Reservoir, Snowmass John Demboski, PhD (subject editor, Vertebrate Zoology) specimens or holdings, or have a regional focus on the Steve Lee, PhD (subject editor, Space Sciences) Rocky Mountains/Great Plains ecoregions. Village, Colorado (Coleoptera: Frank Krell, PhD (subject editor, Invertebrate Zoology) Scarabaeidae: Aphodiinae) The journal is available online at www.dmns.org/Science/ Steve Nash, PhD (subject editor, Anthropology and Museum-Publications free of charge. Paper copies are Archaeology) exchanged via the DMNS Library exchange program Frank-Thorsten Krell

([email protected]) or are available for purchase Editorial and Production: from our print-on-demand publisher Lulu (www.lulu.com). Frank Krell, PhD: production DMNS owns the copyright of the works published in the Julie Whitman-Zai, PhD: copy editor Annals, which are published under the Creative Commons Attribution Non-Commercial license. For commercial use of published material contact the Alfred M. Bailey Library & WWW.DMNS.ORG/SCIENCE/MUSEUM-PUBLICATIONS Archives at [email protected]. DENVER MUSEUM OF NATURE & SCIENCE ANNALS NUMBER 5, JULY 15, 2014

Frank-Thorsten Krell1 Pleistocene Dung Beetles from MIS 5 at Ziegler Reservoir, Snowmass Village, Colorado (Coleoptera: Scarabaeidae: Aphodiinae)

ABSTRACT—Nine aphodiine dung beetle species are recorded from Pleistocene sediments of the Ziegler Reservoir near Snowmass Village, Colorado, U.S.A. Insect remains from this deposit range from 125,000 to 77,000 yr BP, and are unique in their species richness and in their occurrence at high elevation (2720 meters). Three extant species occur: Aphodius (Dialytodius) decipiens Horn, A. (Planolinellus) vittatus Say, and A. (Planolinoides) duplex LeConte. Six unidentified species are also described, belonging to the genus Aphodius. All species are relatively small coprophagous dwellers known as endocoprids. The faunal richness of this site suggests a speciose dung beetle fauna existed at high elevations in this region, and intimates the importance of this site as one of the richest Pleistocene dung beetle sites in North America.

With mammals come feces, and with feces come dung beetles. When North America hosted an abundant mammal megafauna, the North American dung beetle fauna was supposedly, climate permitting, abundant and diverse, too. Sediments at the Ziegler Reservoir site near Snowmass Village in Colorado, U.S.A. (Johnson & Miller 2012; Johnson et al. 2014; Miller et al., in press), are richly fossiliferous and were deposited during such conditions, approximately 125,000 to 77,000 yr BP. Thus they offer an excellent opportunity to explore the dung beetle fauna of the Rocky Moun- tains before the Last Glacial Maximum. With bison, mammoth, mastodon, ground sloth, sheep, camels, horses, and a speciose small mammal fauna 1Department of Zoology, Denver Museum found at the site (Sertich et al., in prep.), a rich dung beetle fauna can be of Nature & Science, 2001 Colorado expected. Dung beetles (Coleoptera: Scarabaeidae) are a crucial element Boulevard, Denver, Colorado 80205- of most ecosystems with an abundant large herbivore fauna because they 5798, U.S.A. provide ecosystem services such as nutrient cycling and enhancement of [email protected] plant growth, bioturbation, and suppression of parasites and dung breed- ing pests (Losey & Vaughan 2006; Kryger 2009). Here we examine this important element of the Pleistocene Snowmass ecosystems in detail. Krell Pleistocene dung beetles from Snowmass Village, Colorado

Site ticulated fragments, particularly their much reduced The Ziegler Reservoir is situated east of Snowmass stiffness, small size, and the complete absence of setae. Village, in Pitkin Country, Colorado, at 39°12’28”N, 106°57’54”W, 2720 m asl. It is the only one of the few high elevation Pleistocene sites in the Rocky Mountains Systematic Paleontology that preserved fossil insect faunas from before the Last Glacial Maximum, from the period between 125,000 Family Scarabaeidae Latreille 1802 and 77,000 yr BP. For geologic setting and stratigraphy Subfamily Aphodiinae Leach 1815 see Pigati et al. (2014), for geochronologic framework Mahan et al. (2014). Tribe Aphodiini Leach 1815

Methods Aphodius Hellwig 1798 Scott Elias and his team extracted beetle frag- Note: I do not follow the controversial upgrading of all ments as described in Elias (2014) from 25 samples Aphodius subgenera to genera as practiced by Gordon & (7.5 l each) taken from the Ziegler Reservoir site in Skelley (2007), because it lacks any phylogenetic foun- June, 2011. Those fragments tentatively assigned to dation, hence being premature. Scarabaeoidea were received from S. Elias for further identification. The specimens were identified by using published descriptions and extant material from Aphodius (Dialytodius) decipiens Horn 1887 the collections of the Denver Museum of Nature & (Figs 1–5) Science and the Field Museum of Natural History, Chicago. All the scarab specimens, and all beetle Material. MIS 5a: 2 elytral fragments: DMNS ZE.15259 samples from the Ziegler Reservoir site are depos- (Fig. 1; 49.16/15–20 cm; 84–77ka), DMNS ZE.15260 ited in the Entomology Collection, Department of (Fig. 2; 49.16/20–25 cm; 84–77ka). MIS 5c: 1 elytral Zoology, Denver Museum of Nature & Science (DMNS). fragment: DMNS ZE.15261 (Fig. 3; 49.11/30–35 cm; 103–101ka); 2 presumably associated heads: DMNS Results ZE.15262 (Fig. 4; 49.11/20–25 cm; 103–101ka), width From a sample volume of 287.5 l (Elias 2014), 1.1 mm; DMNS ZE.15263 (Fig. 5; 49.11/15–20 cm; 32 fragments of scarab beetles were extracted. 103–101ka). Twenty-nine fragments could be assigned to Description. The elytral sculpture of Aphodius decipi- presumably 9 species. All species belong to the ens with deeply impressed, largely punctured striae subfamily Aphodiinae, and to the tribe Aphodiini and tectiform intervals is unique, at least amongst of the predominantly coprophagous dweller guild North American species. The shape of the two heads (endocoprids). The three other fragments, two front with slightly denticulate, emarginated, laterally legs and one elytral part, likely belong to Aphodiinae convex clypeus, only slightly protruding lateral lobes also, but did not allow association with one of the (“genae”), densely punctured surface, and distinct established species. All specimens are from Marine frontal suture without tubercles fits this species (cf. Isotope Stage (MIS) 5 and range in age from 125,000 Gordon & Skelley 2007: fig. 447). to 77,000 years (Elias 2014), covering the whole Discussion. Aphodius decipiens is an extant species period from which fossil insects were obtained at the currently known from Alberta, Idaho, Montana, site. Although some Aphodiinae species are associ- Oregon, and Wyoming (Gordon & Skelley 2007), and ated with rodent burrows, we can exclude modern in Colorado from Mineral and San Juan Counties contamination because of the condition of the disar- (Krell 2010). It is associated with rodents, having been

2 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 3 Krell Pleistocene dung beetles from Snowmass Village, Colorado

found in burrows of the Columbian ground squirrel 93–87ka). 1 presumably associated elytron: DMNS (Urocitellus columbianus (Ord)), marmot (Marmota ZE.15283 (Fig. 11; 49.14/83–90 cm; 93–87ka), with sp.), and pocket gopher (Geomyidae sp.) (Gordon & humeral and apical red spot. MIS 5c: 1 head: DMNS Skelley 2007). The Wyoming ground squirrel (Urocitel- ZE.15267 (Fig. 12; 49.11/30–35 cm; 103–101ka), width lus elegans (Kennicot)) found at the site in layers of 1.2 mm. similar age as the A. decipiens fragments (Sertich et al., Description. The following head characters correspond in press; Mahan et al. 2014) was likely one of several to the extant species Aphodius vittatus: head uniformly possible hosts available to this species. finely punctured, alutaceous, frontal suture with three tubercles, clypeus laterally slightly longitudinally wrinkled, medially very slightly emarginated, sides broadly Aphodius (Planolinellus) vittatus Say 1825 rounded, lateral lobes rounded and slightly protrud- (Figs 6–12) ing. The elytral intervals of this species are finely, but distinctly punctured and alutaceous; punctures notch Material. MIS 5a: 1 elytral fragment: DMNS ZE.15286 the thin striae; the color varies from reddish to dark, (Fig. 6; 49.16/15–20 cm; 84–77ka). MIS 5b: 3 heads: normally being two-tone. One elytron and one elytral DMNS ZE.15264 (Fig. 7; 49.14/78–83 cm; 93–87ka), fragment of the right size show these characters and width 1.2 mm; DMNS ZE.15265 (Fig. 8) and DMNS probably belong to this species. One pronotum of the ZE.15266 (Fig. 9; 49.14/83–90 cm; 93–87ka), width 1.15 fitting size shows double punctation of the right density mm and 1.0 mm, respectively. 1 presumably associated and is basally and laterally marginated, corresponding pronotum: DMNS ZE.15284 (Fig. 10; 49.14/83–90 cm; to A. vittatus.

1 2

3 4 5

Figures 1–5. Aphodius (Dialytodius) decipiens Horn 1887 from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 1. Elytral fragment DMNS ZE.15259, 49.16, 84–77ka. Fig. 2. Elytral fragment DMNS ZE.15260, 49.16, 84–77ka. Fig. 3. Elytral fragment DMNS ZE.15261, 49.11, 103–101ka. Fig. 4. Head DMNS ZE.15262, 49.11, 103–101ka. Fig. 5. Head DMNS ZE.15263, 49.11, 103–101ka.

2 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 3 Krell Pleistocene dung beetles from Snowmass Village, Colorado

Discussion. Aphodius vittatus is a currently wide- Aphodius (Planolinoides) duplex LeConte 1878 spread and common species, occurring from southern (Figs 13–19) Canada to northern Mexico, and from eastern Europe to central Asia (Gordon & Skelley 2007). In Colorado Material. MIS 5a: 1 head: DMNS ZE.15268 (Fig. 13; it has been recorded from several western counties, 49.16/10–15 cm; 84–77ka), width 1.55 mm. 1 pos- eastward up to Weld and El Paso Counties (Krell 2010). sibly associated elytral fragment (rolled up; not It is a generalist dung feeder occurring in a variety of photographed): DMNS ZE.15276 (49.16/20–25 cm; habitats (Gordon & Skelley 2007). 84–77ka). MIS 5b/5a: 1 elytral fragment: DMNS ZE.15285 (Fig. 14; 49.16/5–10 cm; 84–77ka). MIS 5b:

6 7

8 9 10

11 12

Figures 6–12. Aphodius (Planolinellus) vittatus Say 1825 from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 6. Elytral fragment DMNS ZE.15286, 49.16, 84–77ka. Fig. 7. Head. DMNS ZE.15264, 49.14, 93–87ka. Fig. 8. Head DMNS ZE.15265, 49.14, 93–87ka. Fig. 9. Head DMNS ZE.15266, 49.14, 93–87ka. Fig. 10. Pronotum DMNS ZE.15284, 49.14, 93–87ka. Fig. 11. Elytron DMNS ZE.15283, 49.14, 93–87ka. Fig. 12. Head DMNS ZE.15267, 49.11, 103–101ka.

4 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 5 Krell Pleistocene dung beetles from Snowmass Village, Colorado

1 head: DMNS ZE.15269 (Fig. 15; 49.14/83–90 cm; males, clypeus and frons (except the middle) strongly 93–87ka), width 1.5 mm. 3 possibly associated elytral punctured, vertex with finer, irregular punctation, and fragments: DMNS ZE.15270 (Fig. 16) and DMNS lateral lobes only slightly protruding. As Horn (1887) ZE.15271 (Fig. 17; 49.14/78–83 cm; 93–87ka); DMNS already realized, “The presence of a transverse clypeal ZE.15272 (Fig. 18; 49.14/83-90 cm; 93–87ka). MIS 5e: carina is less of a peculiar character than supposed by 1 head: DMNS ZE.15273 (Fig. 19; 66.7; 125ka), width Dr. LeConte” [the author of the species] but taking the 1.1 mm. other characters mentioned above into consideration, Description. The head of Aphodius duplex has a trans- the identification is likely. The size difference between verse clypeal ridge, the frontal suture tuberculated in the two heads is remarkable, but Aphodius species

13 14

15 16 17

18 19

Figures 13–19. Aphodius (Planolinoides) duplex LeConte 1878 from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 13. Head DMNS ZE.15268, 49.16, 84–77ka. Fig. 14. Elytral fragment DMNS ZE.15276, 49.16, 84–77ka. Fig. 15. Head DMNS ZE.15269, 49.14, 93–87ka. Fig. 16. Elytral fragment DMNS ZE.15270, 49.14, 93–87ka. Fig. 17. Elytral fragment DMNS ZE.15271, 49.14, 93–87ka). Fig. 18. Elytral fragment DMNS ZE.15272, 49.14, 93–87ka). Fig. 19. Head DMNS ZE.15273, 66.7, 125ka.

4 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 5 Krell Pleistocene dung beetles from Snowmass Village, Colorado

are known for intraspecific size variation. The frontal 20 suture of the smaller head has no tubercles, indicating that it belonged to a female. The larger heads are male. Four elytral fragments show characters similar to A. duplex elytra (Gordon & Skelley 2007: fig. 170) with thin striae the punctures of which notch the borders of the striae, and with obsolete punctation on the smooth, slightly convex intervals. Those fragments could belong to this species. Discussion. Aphodius duplex is a rodent associate having been collected from burrows of Richardson’s ground squirrel (Urocitellus richardsonii) in Alberta Figure 20. Aphodius sp. 1, subgenus and Wyoming. The species is currently distributed from Agoliinus, aleutus-group, from MIS 5 of the southern Manitoba to Alberta and eastern Washington, Ziegler Reservoir site, Snowmass Village, south to Montana and Colorado (Gordon & Skelley Colorado. Head DMNS ZE.15274, 49.11, 2007). In Colorado, it has been recorded from Fremont 103–101ka. and Larimer Counties (Krell 2010). The Wyoming ground squirrel (Urocitellus elegans) found at the site in layers of similar age as the A. duplex fragments (Sertich et al., in prep.; Mahan et al., 2014) was likely Aphodius sp. 2, subgenus Agoliinus one of several possible hosts available to this species. (Figs 21–22)

Aphodius sp. 1, subgenus Agoliinus, Material. MIS 5a: 1 head: DMNS ZE.15275 (Fig. 21; aleutus-group 49.16/20–25 cm; 84–77ka), width 1.5 mm. MIS 5b: (Fig. 20) 1 head: DMNS ZE.15277 (Fig. 22; 49.14/83–90 cm; 93–87ka), width 1.35 mm. Material. MIS 5c: 1 head: DMNS ZE.15274 (Fig. 20; Description. The following combination of characters 49.11/30–35 cm; 103–101ka), width 1.3 mm. points to the subgenus Agoliinus: Head alutaceous, Description. A punctured head with an emarginated punctures of clypeus and frons moderately strong, finer clypeus, obtusely angled at the sides of the emargina- at the margin of the clypeus, punctures on vertex of dif- tion, straight towards the well pronounced, rounded ferent sizes, frontal suture without tubercles (ZE.15277; triangular lateral lobes, with a relatively flat frons, female) or with three weak tubercles (ZE.15275; male), and without tubercles on the frontal suture is present clypeus emarginated, sides broadly rounded, lateral in several extant species of the subgenus Agoliinus of lobes slightly protruding, narrow, rounded, sinuated the aleutus-group. It also resembles heads of Dialytes, close to the eye (resembling more Agoliinus than the but the relatively flat frons and the shape of the lateral somewhat similar Planolinoides). lobes do not fit extant Dialytes species. Discussion. Agoliinus species generally inhabit higher Discussion. Overall, the aleutus-group contains 18 elevations or higher latitudes and are either surface species in North America. All species are surface dung dung feeders or associated with rodents (Gordon & feeders, mostly generalists (Gordon & Skelley 2007) Skelley 2007). which would have been readily supported by the large mammal fauna of the site (Sertich et al., in prep.).

6 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 7 Krell Pleistocene dung beetles from Snowmass Village, Colorado

21 22

Figures 21–22. Aphodius sp. 2, subgenus Agoliinus, from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 21. Head DMNS ZE.14275, 49.16, 84–77ka. Fig. 22. Head DMNS ZE.15277, 49.14, 93–87ka.

Aphodius sp. 3, subgenus Agoliinus/ protruding, narrow, rounded, sinuated close to the eye Planolinoides (more of the Agoliinus type). (Figs 23–24) Aphodius sp. 4 Material. MIS 5a: 2 heads: DMNS ZE.15278 (Fig. 23), (Fig. 25) DMNS ZE.15279 (Fig. 24; 49.16/15–20 cm; 84–77ka), right side damaged, but ca. 1.9 and almost 2 mm wide, Material. MIS 5b/5a: 1 head: DMNS ZE.15280 (Fig. 25; respectively. 49.16/5–10 cm; 84–77ka), width 1.3 mm. Description. The following combination of characters Description. The characters of the head do not allow points to the subgenera Agoliinus or Planolinoides: the assignment to a subgenus (head orange brown, Vertex with dense fine punctation, frons and clypeus frons and vertex finely and sparsely punctured, clypeus with larger, moderately dense punctures, frontal more strongly, but equally sparsely punctured, frontal suture with strong central tubercle and weaker lateral suture with three weak tubercles, rounded lateral lobe tubercles, clypeus not emarginated, lateral lobes slightly well pronounced on left side, clypeus not emarginated).

23 24

Figures 23–24. Aphodius sp. 3, subgenus Agoliinus or Planolinoides, from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 23. Head DMNS ZE.15278, 49.16, 84–77ka. Fig. 24. Head DMNS ZE.15279, 49.16, 84–77ka.

6 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 7 Krell Pleistocene dung beetles from Snowmass Village, Colorado

25 suture visible, without tubercles, lateral lobes weak, broadly rounded, hardly any angle between lobes and clypeus, clypeus weakly emarginated, sides broadly rounded).

Aphodius sp. 6 (Fig. 27)

Material. MIS 5a: anterior half of 1 head: DMNS ZE.15282 (Fig. 27; 49.16/30–35 cm; 84–77ka), fragment width ca. 1.1 mm. Figure 25. Aphodius sp. 4 from MIS 5 of Description. The characters of the head do not allow the Ziegler Reservoir site, Snowmass Village, the assignment to a subgenus (alutaceous, clypeus with Colorado. Head DMNE ZE.15280, 49.15, fine, dense punctures, slightly wrinkled anterolaterally, 87–84ka. clypeus hardly emarginated, sides broadly rounded, frontal suture with a central tubercle and weak lateral Aphodius sp. 5 tubercles). (Fig. 26)

Material. MIS 5a: 1 head: DMNS ZE.15281 (Fig. 26; 27 49.16/15–20 cm; 84–77ka), width 1.1 mm. Description. The characters of the head do not allow the assignment to a subgenus (moderately strong, deep punctures, interspersed with finer punctures, frontal

Figure 27. Aphodius sp. 6 from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Head DMNS ZE.14282, 49.16, 84–77ka..

Unidentified Aphodiinae fragments (Fig. 28–30)

Figure 26. Aphodius sp. 5 from MIS 5 of A front leg (DMNS ZE.15291; MIS 5b, 49.14/83–90 the Ziegler Reservoir site, Snowmass Village, cm; 93–87ka; Fig. 28), a front tibia (DMNS ZE.15292; Colorado. Head DMNS ZE.15281, 49.16, MIS5b, 49.14/78–83 cm; 93–87ka; Fig. 29) and an 84–77ka.. elytral fragment (DMNS ZE.15293; MIS 5c, 49.11/20–25

8 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 9 Krell Pleistocene dung beetles from Snowmass Village, Colorado

cm; 103–101ka; Fig. 30) belong to Aphodiinae, but number for the North American and Eurasian Pleisto- could not be assigned to one of the species above. cene (Kiselev & Nazarov 2009; Zinovyev 2011). In a few Pleistocene sites in Europe, 10 to 15 dung beetle species Note. Elias (2014 and pers. comm.) mentioned Aegialia were found (Coope et al. 1961; Coope & Angus 1975; (Psammoporus) opaca Brown 1931 (Tribe Aegialiini Bidashko & Proskurin 1987). With nine dung beetle Laporte 1840) from MIS 5a, represented by a frag- species of the genus Aphodius, the Ziegler Reservoir site ment of a right elytron from sample 49.16/20–25 cm is one of the most speciose sites for dung beetles in the (84–77ka). I could not find this specimen in any of North American Pleistocene. the samples and cannot confirm this species from the Most Pleistocene scarab specimens can be assigned Ziegler Reservoir fauna. to extant species (Krell 2006). With 79 extant species of aphodiine dung beetles recorded from Colorado (Krell 2010) and over 430 in the Nearctics (Smith 2009), Discussion many having a rather uniform appearance, we need While beetles are generally abundant in Pleistocene unique structures or sculptures to reliably identify fossil deposits (Buckland & Coope 1991), Scarabaeidae are fragments to species. Only a portion of the specimens represented less frequently and are rarely species rich from Ziegler Reservoir provides us with species-specific (Kiselev and Nazarov 2009). Other Colorado Pleisto- characters. Even then, as with all fossil fragments, the cene sites revealed only one to four dung beetle species identification is very likely to be correct, but not abso- (Short & Elias 1987; Elias & Nelson 1989), being a usual lutely certain since we do not know possible extinct

28 29

Figures 28–30. Unassigned fragments of Aphodiinae from MIS 5 of the Ziegler Reservoir site, Snowmass Village, Colorado. Fig. 28. Front leg DMNS ZE.15291, 49.14/83–90 cm; 93–87ka. Fig. 29. Front tibia DMNS ZE.15292, 49.14/78–83 cm; 93–87ka. Fig. 30. Elytral fragment DMNS ZE.15293, 49.11/20–25 cm; 103–101ka.

8 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 9 Krell Pleistocene dung beetles from Snowmass Village, Colorado

similar species, recent speciation events, extensive range beetle species in Colorado (Krell 2010) because of the shifts, or a possibly different morphological variation of plentiful food supplied by a more diverse Pleistocene the species a hundred thousand or more generations ago. mammal fauna. Nevertheless the higher-than-average A third of the beetle species identified from the Ziegler number of dung beetle species found at the site indicates Reservoir fauna no longer occurs in Colorado (Elias a higher-than-average dung beetle diversity of the Pleis- 2014), but the three identified Aphodius species of dung tocene dung beetle communities in this high elevation beetles still do, or occur here again. Colorado is outside part of the Rocky Mountains. the current range of the here unconfirmedAegialia opaca (reported by Elias 2014) which has its southern- most limits in northern California and Montana. Acknowledgments Not all sites with recorded megafauna bear a diverse I am grateful to Kirk Johnson, now at the Smithson- dung beetle fauna (Barnosky et al. 1988; Coope 2007), ian, and Ian Miller, DMNS, for the opportunity to work sometimes not containing any dung beetles at all (Med- on Snowmass material. Scott Elias, Royal Holloway, vedev and Voronova 1977; Kiselev et al. 1982); but some University of London, for providing me with possible reveal them in abundance such as the Trafalgar Square scarabaeoid fragments from the Ziegler Reservoir site. site (Franks et al. 1958; Coope 2000), the late-glacial Chris Grinter, DMNS, took the photographs. woolly mammoth discoveries at Condover, Shropshire (Allen et al. 2009), and the mammoth channel at Lynford, Norfolk (Coope 2012), as we would expect. The Ziegler Reservoir fauna also met our expectations and Literature Cited adds to our deficient knowledge of Pleistocene North Allen, J.R.M., Scourse, J.D., Hall, A.R. & Coope, G.R. American dung beetles. Despite numerous megafaunal 2009. Palaeoenvironmental context of the sites, the only dung beetle having been found associated Late-glacial woolly mammoth (Mammuthus with Pleistocene mammoth dung is the large, Holarctic primigenius) discoveries at Condover, Shropshire, Aphodius fossor (Elias 2010: 190). Although this species UK. Geological Journal 44: 414–446 + electronic is currently widespread in Colorado (Krell 2010), it is not supplement. represented in the Snowmass samples. The only insects Barnosky, A.D., Barnosky, C.W., Nickmann, R.J., having been found associated with ground sloth dung Ashworth, A.C., Schwert, D.P. & Lantz, S.W. 1988. are Diptera larvae and pupae of the family Sciaridae. No Late Quaternary paleoecology at the Newton site, beetle was found (Waage 1976). Bradford Co., northeastern Pennsylvania: Mam- muthus columbi, palynology, and fossil insects. Conclusion Bulletin of the Buffalo Society of Natural History The dung beetle specimens represented in Pleistocene 33: 173–184. samples of the Ziegler Reservoir site contain both rodent Bidashko, F.G. & Proskurin, K.P. 1988. The entomo- associates and general surface feeders. All of them belong logical and carpological reconstruction of the to the smaller species of the dung beetle communities, bio-environment of the Singilian (Middle Pleisto- and to the dweller guild. Rollers and tunnelers which cene) of the Lower Volga. Paleontological Journal form a significant part of extant temperate dung beetle 1987 (4): 66–72. faunas were not found. Despite the relative richness for a Buckland, P.C. & Coope, G.R. 1991. A Bibliography and Pleistocene site, the nine aphodiine species are likely to Literature Review of Quaternary Entomology. represent only a small portion of the Pleistocene dung Sheffield: J.R. Collins Publications. 85 pp. beetle fauna of Colorado. This fauna is unlikely to be Coope, G.R. 2000. The climatic significance of coleo- much poorer than the over 90 currently occurring dung pteran assemblages from the Eemian deposits

10 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 11 Krell Pleistocene dung beetles from Snowmass Village, Colorado

in southern England. Geologie en Mijnbouw 79: Horn, G.H. 1887. A monograph of the Aphodiini 252–267. inhabiting the United States. Transactions of the Coope, G.R. 2007. Coleoptera from the 2003 excavations American Entomological Society 14: 1–110. of the mammoth skeleton at Niederweningen, Johnson, K. & Miller, I. 2012. Digging Snowmastodon. Switzerland. Quaternary International 164–165: Discovering and Ice Age World in the Colorado 130–138. Rockies. Denver and Aspen, CO: Denver Museum Coope, G.R. 2012. The insect remains from the of Nature & Science and People’s Press. mammoth channel at Lynford, Norfolk. Pp. 75–95 Johnson, K.R., Miller, I.M., Pigati, J.S. & Snowmastodon in: Boismier, W.A., Gamble, C. & Coward, F. (eds) Project Science Team (Ager, T., Anderson, S., Neanderthals among Mammoths. Excavations Carrara, P., Fisher, D., Graham, R., Holen, S., at Lynford Quarry, Norfolk. Swindon: English Jackson, S., McDonald, G., Sertich, J., Shapiro, B., Heritage. Stucky, R., Baker, D., Bright, J., Brown, P., Bryant, Coope, G.R. & Angus, R.B. 1975. An ecological study B., Cherney, M., Cwynar, L., Demboski, J., Elias, of a temperate interlude in the middle of the S., Gray, H., Hadley, E., Hansen, K., Haskett, D., last glaciations, based on fossil Coleoptera from Honke, J., Jiménez-Moreno, G., Kline, D., Krell, Isleworth, Middlesex. Journal of Animal Ecology F., Lifton, N., Lucking, C., Mahan, S., Miller, D., 44: 365–391, 2 pls. Muhs, D., Nash, S., Newton, C., Paces, J., Poinar, Coope, G.R., Shotton, F.W., Strachan, I. & Dance, S.P. H., Porinchu, D., Roundtrey, A., Sharpe, S., 1961. A late Pleistocene fauna and flora from Spaulding, S., Stiller, M., Street, J., Strickland, Upton Warren, Worcestershire. Philosophical L., Thompson, B., Fox, N., Freierman, A., Grimm, Transactions of the Royal Society London B 714: B., Kerr, T., MacCracken, G., O’Neill, H., Petrie, 379–421. L., Richards, S., Stanley, K. & Troll, C.) 2014. The Elias, S.A. 2010. Advances in Quaternary Entomology. Snowmastodon Project. Quaternary Research, doi: Developments in Quaternary Sciences 12: i–xiv, 10.1016/j.yqres.2013.12.010 1–288. Kiselev, S.V. & Nazarov, V.I. 2009. Late Cenozoic insects Elias, S.A. 2014. Environmental interpretation of fossil of northern Eurasia. Paleontological Journal 43: insect assemblages from MIS 5 at Ziegler Res- 723–850. ervoir, Snowmass Village, Colorado. Quaternary Kiselev, S.V., Druk, A.Ja. & Krivolutskii, D.A. 1982. O Research, doi: 10.1016/j.yqres.2014.01.005 faune zhukov i pantsirnykh kleshchei zakhorone- Elias, S.A. & Nelson, A.R. 1989. Fossil invertebrate nii mamontov. Pp. 44–53 in Anon.: Yupibeiskii evidence for Late Wisconsin environments at the Mamont. Moskva: Nauka. [in Russian] Lamb Spring site, Colorado. Plains Anthropologist Krell, F.-T. 2006. Fossil record and evolution of Scara- 34: 309–326. baeoidea (Coleoptera: Polyphaga). Coleopterists Franks, J.W., Sutcliffe, A.J., Kerney, M.P. & Coope, G.R. Society Monograph 5: 120–143. 1958. Haunt of elephant and rhinoceros: the Krell, F.-T. 2010. Catalogue of Colorado scarab and stag Trafalgar Square of 100,000 years ago–new dis- beetles (Coleoptera: Scarabaeoidea), based on coveries. The Illustrated London News 1958: 1101. literature records. Denver Museum of Nature & Gordon, R.D. & Skelley, P.E. 2007. A monograph of Science Technical Report 2010-4: 1–84. the Aphodiini inhabiting the United States and Kryger, U. 2009. Conservation of dung beetles. Pp. Canada (Coleoptera: Scarabaeidae: Aphodiinae). 387–483 in Scholtz, C.H., Davis, A.L.V. & Kryger, Memoirs of the American Entomological Institute U.: Evolutionary Biology and Conservation of 79: 1–580. Dung Beetles. Sofia: Pensoft.

10 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 11 Krell Pleistocene dung beetles from Snowmass Village, Colorado

Losey, J.E. & Vaughan, M. 2006. The economic value of Short, S.K. & Elias, S.A. 1987. New pollen and beetle ecological services provided by insects. BioScience analyses at the Mary Jane site, Colorado: Evidence 56: 311–323. for late glacial tundra conditions. Geological Mahan, S.A., Gray, H.J., Pigati, J.S., Wilson, J., Lifton, Society of America Bulletin 98: 540–548. N.A., Paces, J.B. & Blaauw, M. 2014. A geochrono- Smith, A.B.T. 2009. Checklist and nomenclatural logic framework for the Ziegler Reservoir fossil authority file of the Scarabaeoidea of the Nearctic site, Snowmass Village, Colorado. Quaternary Realm. Version 4. http://museum.unl.edu/ Research, doi: 10.1016/j.yqres.2014.03.004 research/entomology/SSSA/nearctic.htm [accessed Medvedev, L.N. & Voronova, N.V. 1977. Koleoptero- 29 Dec. 2013] logicheskii analiz geologicheskikh razrezov Waage, J.K. 1976. Insect remains from ground sloth mamontovykh kladbishch v severnoi Jakuitii. dung. Journal of Paleontology 50: 991. Trudy Zoologicheskogo Instituta 72: 72–77. [in Zinovyev, E. 2011. Sub-fossil beetle assemblages associ- Russian] ated with the “mammoth fauna” in the Late Miller, I.M., Pigati, J.S., Anderson, R.S., Johnson, K.R., Pleistocene localities of the Ural Mountains and Mahan, S.A., Ager, T.A., Baker, R.G., Blaauw, M., West Siberia. ZooKeys 100: 149–169. Bright, J., Brown, P.M., Bryant, B., Calamari, Z.T., Carrara, P.E., Cherney, M.D., Demboski, J.R., Elias, S.A., Fisher, D.C., Graham, R.W., Gray, H.J., Haskett, D.R., Honke, J.S., Jackson, S.T., Jiménez- Moreno, G., Kline, D., Leonard, E.M., Lifton, N.A., Lucking, C., McDonald, H.G., Miller, D.M., Muhs, D.R., Nash, S.E., Newton, C., Paces, J.B., Petrie, L., Plummer, M.A., Porinchu, D.F., Rountrey, A.N., Scott, E., Sertich, J.J.W., Sharpe, S.E., Skipp, G.L., Strickland, L.E., Stucky, R.K., Thompson, R.S. & Wilson, J. (in press). A high-elevation, multi-proxy biotic and environmental record of MIS 6-4 from the Ziegler Reservoir fossil site, Snowmass Village, Colorado, USA. Quaternary Research. Pigati, J.S., Miller, I.M., Johnson, K.R., Honke, J.S., Carrara, P.E., Muhs, D.R., Skipp, G. & Bryant, B. 2014. Geologic setting and stratigraphy of the Ziegler Reservoir fossil site, Snowmass Village, Colorado. Quaternary Research, doi: 10.1016/j. yqres.2013.12.011 Sertich, J.J.W., Graham, R.W., Stucky, R.K., McDonald, H.G., Newton, C., Fisher, D.C., Scott, E., Demboski, J.R., Lucking, C., McHorse, B.K. & Davis, E.B. (in prep.). High-elevation Late Pleistocene (MIS 5-6) vertebrate faunas from the Ziegler Reservoir fossil site, Snowmass Village, Colorado. Quaternary Research.

12 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 DENVER MUSEUM OF NATURE & SCIENCE ANNALS | No. 5, July 15, 2014 PB DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER MUSEUM OF NATURE & SCIENCE ANNALS DENVER MUSEUMDENVER OF NATURE & SCIENCE ANNALS • NUMBER 5 JULY 15, 2014 NUMBER 5, JULY 15, 2014

The Denver Museum of Nature & Science inspires curiosity and excites minds of all ages through scientific discovery and the presentation and preservation of the world’s unique treasures.

Cover photo: Scott Elias, Royal Holloway University of London (left), and Aaron Spriggs, DMNS collections assistant, excavating site 49, the “Twisty Trench”, at the Ziegler Reservoir for insects.