Western North American Naturalist

Volume 74 Number 1 Article 1

5-23-2014

Cave-dwelling and vertebrates of North Rim Grand Canyon, with notes on ecology and management

J. Judson Wynne Northern Arizona University, Flagstaff, AZ, [email protected]

Kyle D. Voyles Bureau of Land Management, St George, UT, [email protected]

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Recommended Citation Wynne, J. Judson and Voyles, Kyle D. (2014) "Cave-dwelling arthropods and vertebrates of North Rim Grand Canyon, with notes on ecology and management," Western North American Naturalist: Vol. 74 : No. 1 , Article 1. Available at: https://scholarsarchive.byu.edu/wnan/vol74/iss1/1

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 74(1), © 2014, pp. 1–17

CAVE-DWELLING ARTHROPODS AND VERTEBRATES OF NORTH RIM GRAND CANYON, WITH NOTES ON ECOLOGY AND MANAGEMENT

J. Judson Wynne1 and Kyle D. Voyles2

ABSTRACT.—Prior to this study, there was no information on arthropods, bats, and other vertebrates of caves in north- westernmost Arizona. Based on invertebrate and vertebrate inventory work conducted during 2005 and 2006, we pro- vide future directions for conservation and management for caves on Grand Canyon–Parashant National Monument, northwestern Arizona. Baseline investigations to find and identify arthropods, bats, and other vertebrates were con- ducted at 7 of the largest known caves on the monument. We identified 52 morphospecies including 44 arthropods, 4 bats, and 4 other vertebrates. Of the cave-dwelling arthropods, we found 10 eisodophiles, 6 troglophiles, 8 questionable troglophiles, 7 trogloxenes, 8 accidentals, 3 taxa of unknown cave affiliations, and 2 mammalian parasites. We made sev- eral contributions to the entomological record including 7 new species (with 2 new genera), 3 possible new species, one range extension, and one possible range extension. Also, we identified 5 bat roosts—1 hibernaculum, 2 night roosts, and 3 summer roosts of unconfirmed use. Observed richness per cave ranged from 1 to 14 morphospecies, and observed bat and other vertebrate (combined) richness was 1–3 morphospecies. We did not detect any cave-adapted arthropods during this investigation. For the caves sampled, we are uncertain whether the lack of cave-adapted taxa is due to (a) low nutrient input and high cryptoaridity associated with many southwestern cave systems or (b) lack of inten- sive sampling. Despite the lack of cave-adapted species, 5 of the 7 caves inventoried are considered of high manage- ment concern. Additional research at these caves will be required to obtain the data necessary to best manage and pro- tect these systems.

RESUMEN.—Anterior a este estudio, no había información sobre los artrópodos, murciélagos y otra fauna en las cav- ernas al noroeste de Arizona. Basados en el inventario de invertebrados y vertebrados realizados durante 2005 y 2006, nosotros proveemos las futuras direcciones para la investigación y gestión de cavernas del Grand Canyon–Parashant National Monument, noroeste de Arizona. Investigaciones iniciales fueron realizados en artrópodos, murciélagos y fauna en 7 de las más grandes cavernas conocidas. Identificamos 52 morfoespecies incluyendo 44 artrópodos, 4 murciélagos y 4 fauna silvestre. De los artrópodos, hubo 10 eisodofilos, 6 troglofilos, 8 pseudo-troglofilos, 7 trogloxenos, 8 acciden- tales, 3 desconocidas y 2 ectoparásitos. Realizamos varias contribuciones al registro entomológico incluyendo 7 nuevas especies (con 2 nuevos géneros), 3 posibles nuevas especies, una expansión distribucional y una posible expansión dis- tribucional. También, identificamos 5 refugios de murciélagos: 1 hibernáculo, 2 dormideros nocturnos y 3 dormideros estivales de uso indeterminado. La riqueza observada de artrópodos oscilo entre 1–14 morfoespecies y la riqueza combi- nada para murciélagos y fauna vario entre 1–3 morfoespecies. Durante este trabajo, no fueron encontrados artrópodos troglomorficos. En las cavernas muestreadas, se desconoce si la falta de taxones adaptados a las cavernas es debido a (i) el bajo aporte de nutrientes y la alta cripto-aridez asociada generalmente con los sistemas de cavernas del suroeste, o (ii) el insuficiente muestreo. A pesar de ello, 5 de las 7 cavernas inventariadas son consideradas como de alto interés de gestión. Investigaciones adicionales en estas cavernas serán necesarios para obtener los datos requeridos para una mejor gestión y protección de estos sistemas.

Information related to the biospeleology desire to provide a regional summary of cave of northern Arizona is limited. Prior to this biology, we present a brief overview of the study, no investigations on cave use by ar - taxa reported during previous work. thropods, bats, and other vertebrates in north- At Wupatki National Monument, Welbourn western Arizona had been undertaken. Based (1976) conducted 1–2 site visits at 5 earth cracks on our literature review, most of the work on (volcano-tectonic fissures) and identified 19 ar- the southern Colorado Plateau (within north- thropods, including 5 troglophiles, 13 troglox- ern Arizona) has been baseline in nature, and enes, and 5 accidentals; no troglobites were these studies were largely focused on Wupatki identified during his work (see below for defi- National Monument and Grand Canyon Na - nitions on cave specificity functional groups). tional Park. Given limited information and the From Welbourn’s inventory (1976), Muchmore

1Department of Biological Sciences, Colorado Plateau Biodiversity Center, Colorado Plateau Research Station and Landscape Conservation Initiative, Northern Arizona University, Box 5614, Building 56, Suite 150, Flagstaff, AZ 86011. URL: http://www.jutwynne.com 2Saint George Field Office, Bureau of Land Management, 345 E. Riverside Dr., St. George, UT 84790.

1 2 WESTERN NORTH AMERICAN NATURALIST [Volume 74

(1981) described an endemic pseudoscorpion picture of the natural history of caves on the (Archeolarca welbourni) from 2 earth cracks. southern Colorado Plateau. Bat research has been conducted on a nearly Here we report the first regional all-taxa decadal scale at Wupatki. Gustafson (1964) col- biological inventory of known caves in Grand lected one Townsend’s big-eared bat (Cory - Canyon–Parashant National Monument, Ari- norhinus townsendii) during the summer near zona. Our objectives were to conduct baseline the bottom of an earth crack. Welbourn (1976) inventories by (1) sampling cave-dwelling ar- identified C. townsendii at 4 earth cracks (one thropods; (2) identifying bat use of caves either of the earth cracks was the same visited by directly or by examining previous evidence of Gustafson [1964]). Day roost activity was docu- use; (3) documenting all other vertebrate ac- mented at all features in September, and 2 tivity observed within each cave; and (4) iden- earth cracks were identified as hibernacula. tifying caves of high management concern for Bain (1986) documented winter use of 7 earth further study. cracks (confirming continued use of the one previously visited by Gustafson and 4 visited METHODS by Welbourn [1976]) by C. townsendii. Con- Cave-Dependency Functional Groups cerning other vertebrate species, Welbourn (1976) identified the remains of 2 accidental We divided Grand Canyon–Parashant cave- morphospecies in Wupatki earth cracks: Ro - dwelling taxa into 7 cave-dwelling organism dentia (from 2 earth cracks) and Lagomorpha groups and one special case category. The fol- (from one earth crack). lowing definitions were taken from Barr (1968), Wynne et al. (2007) compiled a review of Howarth (1983), and Wynne (2013): (1) trog- cave-dwelling arthropod research in Grand lobites, obligate cave dwellers who can only Canyon National Park (GRCA) and synthe- com plete their life cycle within the hypogean sized information from 15 caves representing environment; (2) troglophiles, spe cies that oc - 9 studies (conducted between 1975 and 2001). cur facultatively within caves and complete At least 47 cave-dwelling arthropods, includ- their life cycles there, but also exist in simi - ing 3 troglobites, one stygobite (aquatic cave- lar dark and humid epigean microhabitats; (3) adapted ), 7 trogloxenes, 16 troglophiles, trogloxenes, species that frequently use caves and 16 unidentified cavernicoles (presumed for shelter but forage in the epigean realm; (4) troglophiles) were identified (Wynne et al. accidentals, morphospecies that occur in caves 2007). Additionally, bats known to use GRCA but cannot survive within the hypogean envi- caves include the Brazilian free-tailed bat (Ta- ronment; (5) eisodophiles, species that facul- darida brasiliensis) from 2 caves (Pape 1998, tatively use cave entrances and twilight zones Hill et al. 2001) and C. townsendii from 2 and may complete their life cycles there, but caves (Welbourn 1978). Other cave-dwelling also exist in similar partially sheltered epi- vertebrates known to occur in GRCA include gean environments; (6) eisodoxenes, species Neotoma sp. from 4 caves (Peck 1980, Boden- that frequently use cave entrances and twi- hamer 1989, Drost and Blinn 1998, Hill et al. light zones for shelter but forage on the sur- 2001), brush mouse (Peromyscus boylii) from face; (7) unknown, species for which informa- one cave (Drost and Blinn 1998), American tion is lacking to place them within one of the porcupine (Erethizon dorsatum) from one cave 6 aforementioned groups; and, (8) parasites, a (Hill et al. 2000) and ringtail (Bassariscus astu- special-case group, which includes parasitic ar - tus) from 4 caves (Peck 1980, Bodenhamer thropods detected in caves due to the presence 1989, Pape 1998, Hill et al. 2000, 2001). of their host (e.g., bats or other mammals). Peck (1982) conducted invertebrate surveys Study Site and reported on the largely depauperate fauna (n = 2 morphospecies) occurring in 4 lava tube Located in northwestern Arizona, Grand caves at Sunset Crater National Monument and Canyon–Parashant National Monument (here- in the greater Flagstaff region. after shortened to Parashant) is jointly man- Although much of the biospeoleological re - aged by the National Park Service and the search over the past several decades has been Bureau of Land Management. Encompassing baseline in nature, these studies combined with 1.1 million acres, Parashant is characterized our present work begin to produce a regional by rugged terrain containing deeply incised 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 3 canyons, mesas, and mountains. Two geologi- In caves sampled with pitfall traps, we de - cal provinces converge here: the Basin and ployed traps for 4 days, with 2 traps placed Range and the Colorado Plateau. Vegetation within each of the 3 primary zones—light, twi- zones include Mojave Desert at lower eleva- light, and dark. Because the location of the tions, grading through grassland and juniper twilight zone shifts temporally and season - shrubland to ponderosa pine forest on Mt. ally, we estimated the location of this photic Trumbull (elevation 2447 m). zone during summer. Pitfall trap construction Parashant caves were selected for inventory consisted of two 32-ounce stacked plastic if they contained deep zone–like conditions. containers (13.5 cm height, 10.8 cm diameter Cave deep zones are characterized by com- rim and 8.9 cm base). We used approximately plete darkness, stable temperature, water-satu- 4.9 mL of peanut butter as bait and placed it rated atmosphere, and limited to no airflow in the bottom of the exterior container. At the (see Howarth 1980, 1982). This zonal environ- bottom of the interior container, we made sev- ment serves as habitat for cave-adapted arthro- eral dozen holes so the bait could “breathe” pods. Two of 7 caves met all the deep zone cri- to attract . We buried containers to teria. However, caves meeting most of these the rim when possible, built rock ramps to the criteria (i.e., complete darkness, stable tem- trap rim in other cases, and covered all traps perature, limited airflow) were also included with a caprock. Prior to removing traps at the in our study. Because troglomorphic animals end of the sample period, we searched around often represent rare and endemic taxa, caves each trap to identify and capture individuals supporting these animals are considered of attracted to the bait but not captured within high management and conservation value. the trap (Poulson and Culver 1969). Therefore, we sampled caves with deep zones No pitfall traps were placed in PARA 2202 and deep zone–like conditions. due to safety issues associated with a return To safeguard caves and their resources, we visit. Although PARA 0802 lacked deep zone– referred to all of the caves in this study by like conditions and we considered the pres- using an alphanumeric coding system devel- ence of cave-adapted arthropods un likely, we oped by the Na tional Park Service (NPS) rather sampled the cave for arthropods because of than actual cave names. Parashant headquarters the presence of a bat roost (and thus guano). in St. George, Utah, has the cipher table with We searched each cave in areas deemed cave codes and names. A copy of this paper most likely to contain certain arthropods. These with actual cave names is on file with both the areas included detritus deposits, areas with National Park Service in St. George and the condensed water and mud, guano deposits, National Cave and Karst Research Institute, and active speleothems. We searched for ar - Carlsbad, New Mexico. thropods for at least 40 minutes (2 observers We sampled arthropods, bats, and other ver- × 20 min each) in each zone (i.e., light, twi- tebrates in the following date ranges: 4–14 light, and dark). We also opportunistically col- August 2005 and 16–26 September 2005. Win- lected arthropods by traversing the length of ter roost surveys for hibernating bats were the cave and searching for and collecting ar - conducted during 4–10 March 2006. thropods as encountered (2 observers per cave). Arthropod Sampling Arthropod Identification Based on our site visits, we identified 6 We used a combination of existing keys and caves that were believed to support deep zone– worked with staff members at both the Colo - like conditions, and we included one addi- rado Plateau Museum of Arthropod Biodiver- tional cave due to the presence of a summer sity and the Colorado Plateau Research Sta- bat roost. Five caves (PARA 1801, PARA 2602, tion at Northern Arizona University (NAU) to PARA 2204, PARA 1001, and PARA 1401) identify arthropods to the lowest taxonomic were sampled using a combined baited and level possible. For several taxonomic groups, unbaited pitfall trapping approach and direct we sent specimens to experts, including Rolf intuitive searches. Two caves (PARA 0802 and Aalbu (Coleoptera: Tenebrionidae), Thomas PARA 2202) were sampled using direct intui - Barr Jr. (Coleoptera: Rhadine), Earnest Ber - tive searches and opportunistic hand-collect- nard (Col lembola), Theodore Cohn (Orthop - ing throughout each cave’s length. tera: Rhamphidophoridae), Carl Dick (Dip tera: 4 WESTERN NORTH AMERICAN NATURALIST [Volume 74

Nycteri biidae), Mark Harvey (Pseudoscorpi- ones: Chernetidae), Robert Johnson (Hyme- noptera: Formicidae), Edward Mockford (Pso- coptera), Pierre Paquin (Arachnida), William Shear (Opiliones), Jon Gelhaus (Diptera: Tipu - lidae), and Chen Young (Diptera: Tipulidae). Voucher specimens for most arthropod groups identified through this work are deposited at the Museum of Northern Arizona, Flagstaff. Vertebrate Sampling

BATS.—During August and September 2005, we surveyed for bats by employing a

onal Monument. combination of techniques including (i) mist- netting and harp-trapping at cave entrances during evening emergence; (ii) hand-netting within one cave; (iii) visually identifying bats as encountered within each cave; and (iv) exam - ining caves for evidence of bat use. For each bat captured, we determined sex, age, weight, and reproductive status (Kunz 1982). Cave entrances were presumed to serve as night roosts if we documented guano and/or parts on the cave floor within 5 m of a cave’s entrance. We set 2 mist nets and a harp trap within the entrances of PARA 1801, PARA 0802, and PARA 1401. Mist nets and harp traps were deployed before sunset and removed 2 hours after bats had started their evening emer- gence. We used 6-m, 2-shelf Avinet mist nets (www.avinet.com) and a G6 Cave Catcher harp trap (Bat Management, www.batmanagement .com; maximum catch area 3.35 m2). We hand- netted bats within PARA 2602. During late winter (4–10 March 2006), each cave was visited once to search for and count hibernating bats. When possible, we visually identified all bats encountered to species. No hibernating bats were handled during this work. OTHER VERTEBRATES.—Within each cave, we searched for and recorded the presence of all other vertebrates. Sign of other vertebrates included direct observation, scat, feathers, and skeletal remains. Eisodophiles Eisodoxenes Troglophiles Troglophiles? Trogloxenes Accidentals Unknown Parasites RESULTS

We detected 52 cavernicolous taxa, including 44 arthropods, 4 bats, and 4 other vertebrates (see the Annotated List of Morphospe cies, page 1. Arthropods, bats and other vertebrates identified across all cave specificity functional groups, Grand Canyon–Parashant Nati 1. Arthropods, bats and other vertebrates identified across all cave specificity functional groups, Grand Canyon–Parashant 8). Of these, arthropods comprised 10 eiso-

ABLE dophiles, 6 troglophiles, 8 questionable troglo - T

ArthropodsBatsOther vertebrates — 10 —philes, 2 — 7 trogloxenes, — 6 18 accidentals, — 3 unknowns — 8 — — 7 4 1 8 — — 3 — — 2 — 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 5

TABLE 2. Total number of arthropods observed per Other notable groups include (7 mor - photic zone by cave. pho species from 5 caves), crickets (5 morpho - Light Twilight Dark species from 3 caves), (3 morpho- PARA 1801 20 16 2 species from 3 caves), and ants (4 morphospecies PARA 2202 — ~1000 4 from 3 caves). All ants that we detected repre- PARA 2602 14 6 13 sent accidental occurrences (R. Johnson, per- PARA 2204 9 ~1000 13 sonal communication, 2006). PARA 0802 — 1 — PARA 1401 — — 1 Across all arthropod groups, most morpho- PARA 1001 >500 >500 6 species were limited to a single observation TOTAL 543a 2523a 39 within a single cave. However, both Leiobunum aThe values were estimated for large populations of crickets (PARA 1001) and townsendii and Entomobrya zona were en- harvestmen (PARA 2202 and PARA 2204) and were likely underestimated for countered across 3 caves. each cave. When we considered total numbers of ob - served individuals per light zone, we found and 2 parasites; all bats were trog loxenes, and that the twilight zone contained the largest other vertebrates included one troglophile, one number of individuals (~2523), followed by accidental, and 2 eisodoxenes (Table 1). Ob- the light (~543) and dark zones (~39; Table served arthropod richness ranged from one to 14 2). PARA 1001 contained a large cricket roost morphospecies per cave. The rank order of caves (estimated to contain >1000 individuals). Al - by arthropod species richness was PARA 1801 though the crickets occurred throughout PARA (14), PARA 2602 (13), PARA 1001 (11), PARA 1001, most were detected within the light and 2204 (8), PARA 2202 (5), PARA 0802 (1), and twilight zones. They were often found den- PARA 1401 (1). We detected bats in most caves ning in large groups along the walls and ceil- (71.4%); PARA 0802, PARA 1401, and PARA ings and within several ceiling fissures. PARA 2602 all contained at least 2 bat morpho species. 2202 and PARA 2204 supported large num- Numbers for other vertebrates ranged from bers of Leiobunum townsendii (estimated in the one to 2 morphospecies per cave; both PARA hundreds) denning in packed clusters in large 1001 and PARA 2202 supported 2 vertebrate overhead fissures within the twilight zones of species. each cave. When we shined our light on these clusters, the harvestmen would rock, sway, Arthropods and often aggregate into tighter clusters. Taxonomically, the most morphospecies- This work resulted in discovery of 7 new diverse groups of arthropods were spiders species (with 2 new genera), 3 possible new spe - (Araneae; n = 14), beetles (Coleoptera: 6 differ- cies, 1 range extension, and 1 possible range ent families; n = 7), cave crickets (Or thoptera: extension. The 2 new genera (comprising 2 new Rhaphidophoridae; n = 5), ants (Hy menoptera: species) are a book louse (Psocoptera: Sphae- Formicidae; n = 4), (Diptera: 4 different ropsocidae: Troglosphaeropsocus voylesi, Mock - families; n = 4), and book lice (Psocoptera: 3 ford 2009) and a cave cricket (Rhaphidophori- different families; n = 3). Other groups of par- dae: cf Ceuthophilus n. gen. n. sp., Cohn and ticular interest due to their ecological and evo- Swanson, unpublished data). Additional new lutionary relationships to cave environments species included the following: a new pseudo- include harvestmen (Opiliones; n = 1), mites scorpion (Chernetidae: Tuberochernes n. sp., (Acari; n = 2), and springtails (Collembola; n = det. M. Harvey), a cave cricket (Ceuthophilus 1). The remaining arthropods detected within n. sp., det. T. Cohn); a carabid (Rhadine caves include true bugs (Hemiptera; Reduvii - n. sp., Perlevis species group, det. T. Barr), and dae; n = 1), fleas (Si phonaptera; n = 1) and 2 tenebrionid beetles (Eschatomoxys pholeter bristletails (Thysanura; n = 1). Thomas and Pape 2007 [Pape et al. 2007] and On a regional level, several taxonomic Eleodes [Caverneleodes] wynnei Aalbu 2012 groups were represented by 4 or more mor- [Aalbu et al. 2012]). Images of select taxa are phospecies. Spiders were the most commonly provided in Fig. 1. Two additional Ceuthophilus detected in all caves visited, represent- sp. (n. sp.? a and n. sp.? b) from PARA 1801 ing ~30% of the arthropods detected. PARA may represent new species; the specimens are 1801 and PARA 2202 contained the highest “distinctive, and both may be undescribed” (T. diversity of spiders, 5 and 4, respectively. Cohn, personal communication, 2006). 6 WESTERN NORTH AMERICAN NATURALIST [Volume 74

Fig. 1. Five new species (including 2 new genera) discovered on Grand Canyon–Parashant National Monument, Arizona: A, Rhadine n. sp., Perlevis species group, on cave floor of PARA 2204; B, high resolution postmortem image of Tuberochernes n. sp.; C, a breeding pair of Eleodes (Caverneleodes) wynnei (Aalbu et al. 2012) from PARA 1001; D, postmortem image of Troglosphaeropsocus voylesi (courtesy of E. Mockford); E, a breeding pair of cf Ceuthophilus n. gen. n. sp. from PARA 1001.

Our efforts extended the range of the Spe- survey. We trapped one nonreproductive fe- leketor flocki (Psocoptera: ), which male using a harp trap. We were unable to was previously known from 2 other localities in determine the function of this roost based the southwest: Tucson Mountains, southern Ari- upon the one individual captured. Given the zona, and southeastern Nevada. One accidental presence of a fresh guano pile, we ascertained ant species, Paratrechina hystrix?, may repre- the general location of the roost, which was at sent the first record of this species in Arizona the back of the cave. (R. Johnson, personal communication, 2006). We confirmed 2 pallid bat (Antrozous pal- Specimens match the description of P. hystrix lidus; Fig. 2B) roosts during August surveys. In in part; however, because no workers were PARA 0802 and PARA 2602, we observed ~50 collected, this identification cannot be confirmed and ~100 individuals, respectively. We cap- (R. Johnson, personal communication, 2006). tured one postlactating female, 4 nonreproduc- tive females (1 juvenile, 1 adult, and 2 undeter- Bats mined), and one nonscrotal male, using a harp Five of the 7 caves (71.4%) contained bats. trap and mist nets at PARA 0802. These bats Of these, we identified 3 roosts of an unknown were not marked, so there was the possibility of function, 2 night roosts, and a hibernaculum. double counting. In PARA 2602, we captured PARA 1401 serves as both a summer roost and one nonreproductive female and one nonscro- hibernaculum. tal male with handheld nets. We tentatively We documented a fringed myotis (Myotis identified a pallid bat (on the basis of its light thysanodes; Fig. 2C) colony (~30 individuals) brown color and size) flying within PARA at PARA 1801 during the September 2005 2202. 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 7

Fig. 2. Three cave-roosting bat species confirmed on Grand Canyon–Parashant National Monument, Arizona: A, 3 hibernating Corynorhinus townsendii from PARA 1401; B, 3 Antrozous pallidus from PARA 0802; C, Myotis thysanodes in hand during harp-trapping and mist-netting operations at the entrance of PARA 1801. 8 WESTERN NORTH AMERICAN NATURALIST [Volume 74

In August 2005, we counted approximately tion to the back of the cave. We found scat and 10 Myotis sp. emerging from PARA 1401. quills littering the entrance of PARA 1401, as Numerous lepidopteran wings were docu- well as a fully articulated porcupine skeleton mented within the entrance and twilight zones in the cave. Evidence of owl roosting activity of PARA 0802 and PARA 2602. Corynorhinus (i.e., pellets) was documented in the twilight townsendii is one of the more common trog- zone of PARA 2202. loxenic bats in northern Arizona. This bat uses We found scat from a small carnivore (possi- cave entrances as night roosts, where it has bly ringtail) in the dark zone of PARA 2202. been documented removing the wings of moths This could be one of 3 small carnivores known prior to consuming the abdomen (e.g., Lopez- to use caves in the Southwest, including ring- Gonzalez 2005). In West Virginia, Sample and tail (Bassariscus astutus; e.g., Peck 1980, Boden - Whitmore (1993) have shown this species to hamer 1989, Pape 1998, Hill et al. 2000, 2001, preferentially consume moths over other ar - Strong 2006), skunks (Mephitidae), or raccoon thropods. Because of this documented diet pref- (Procoyon lotor; e.g., Winkler and Adams 1972). erence and the large accumulation of moth wings, we suggest these caves are probably ANNOTATED LIST OF MORPHOSPECIES used as night roosts by C. townsendii. During our early March 2006 surveys, we The following is an annotated list of morpho - observed 9 C. townsendii (Fig. 2A) hibernating species detected from 7 caves on Grand Can - within the deep zone of PARA 1401 at mid-cave. yon–Parashant National Monument, Arizona. Other Vertebrates We identified woodrat (Neotoma spp.) mid- Phylum Arthropoda dens in 4 caves, American porcupine use (Er - Class Arachnida ethizon dorsatum) at 2 caves, owl use of one Order Araneae cave, and small carnivore use of one cave. We Family Pholcidae documented Neotoma spp. middens in the en - Undetermined genus and species. trance and twilight zone of 4 caves (PARA Det. P. Paquin. Eisodophile. 2202, PARA 2602, PARA 2204, and PARA 0802). Woodrats and their middens are commonly ob - One juvenile specimen was collected from served along sheltered rock outcrops, crags, rock the twilight zone of PARA 1801. Because the shelters and cave entrances; thus, woodrats are specimen was an immature, it could not be considered troglophiles. Because we did not trap identified beyond family level (P. Paquin, per- small mammals during this study, we were un - sonal communication, 2006). able to provide species-level identifications. Three woodrat species are known to occur Physocyclus sp. within Parashant. The Mexican woodrat (Neo - Det. P. Paquin. Eisodophile. toma mexicana) occurs within rocky outcrops One male specimen was collected from the and slopes and is found in open woodland and light zone of PARA 2204. transition-zone vegetation (Cornely and Baker 1986). Stephen’s woodrat (Neotoma stephensi) Psilochorus sp. is often found in association with juniper trees Det. P. Paquin. Troglophile? ( Juniperus spp.; Jones and Hildreth 1989). Also, One female specimen was collected from the white-throated woodrat (Neotoma albigula) the dark zone of PARA 2602. Given that it was may also occur at the mid-elevations of the collected from the dark zone, we suggest this monument (Macêdo and Mares 1988). Thus, spider may be a troglophile. woodrat activity may represent one or more of these species at any given site. We found significant use by American por- Family Theridiidae cupine in both PARA 1001 and PARA 1401. Achaearanea sp. Det. P. Paquin. Eisodophile. Woods (1973) indicates that porcupines den in caves, hollow trees, and logs. However, we did One female specimen was removed from a not find evidence of recent activity in either web within the light zone of PARA 1801. Peck cave. In PARA 1001, we confirmed use by (1980) suggests this genus occurs in caves, as porcupine based on an extensive deposition of well as in more mesic epigean environments scat and quills, extending from the middle por - in Arizona, California, New Mexico, and Utah. 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 9

Family Linyphiidae Family Uloboridae Undetermined genus and species. Uloborus sp. Det. P. Paquin. Eisodophile. Det. P. Paquin. Eisodophile. Two juvenile specimens belonging to the One female specimen was collected from same genus and species were collected from the light zone of PARA 1001. the light zone of PARA 1801. Because they were juveniles, they could not be identified Family Oecobiidae beyond family level. Oecobius sp. Det. P. Paquin. Eisodophile. Agyneta sp. One juvenile wall spider was collected from Det. P. Paquin. Troglophile? the twilight zone of PARA 2602. We collected one female specimen from the dark zone of PARA 2204. Given that it was Order Opiliones collected from the dark zone, we suggest this Family Sclerosomatidae spider may be a troglophile. Leiobunum townsendii Weed, 1893. Det. W. Shear. Trogloxene. Family Tetragnathidae We collected 3 specimens in the light zone Metellina sp. Det. P. Paquin. Eisodophile. of PARA 1801. In the twilight zones of both PARA 2204 and PARA 2202, several hundred Two juvenile specimens were collected from individuals were observed within ceiling fis- the light zone of PARA 1001. sures. These individuals were denning close together in large masses. We suspect these Family Tengellidae opillionids den in fissures during the day, and exit to forage at night. This species was also Undetermined genus and species a. observed throughout the length of PARA 1001. Det. P. Paquin. Troglophile? One juvenile specimen was collected from Order Pseudoscorpiones the dark zone of PARA 2202. Given that it was Family Chernetidae collected from the dark zone, we suggest this spider may be a troglophile. Tuberochernes n. sp. Det. M. Harvey. Troglophile? Undetermined genus and species b. Det. P. Paquin. Troglophile? We collected 3 specimens from the dark zone of PARA 1001. These specimens are with We collected 2 juvenile specimens of a a taxonomist and will ultimately be described. genus and species different from the afore- mentioned specimen from the dark zone of PARA 2202. Given that these specimens were Subclass Acari collected from the dark zone, we suggest this Acari species a. spider may be a troglophile. Specimens misplaced by taxonomist. Unknown. Undetermined genus and species c. Two specimens were collected from the Det. P. Paquin. Eisodophile. twilight zone of PARA 2602. One juvenile specimen was collected from Acari species b. the light zone of PARA 1801. Specimens misplaced by taxonomist. Unknown. Two specimens were collected from the Family Plectreuridae light zone of PARA 1801. Plectreurys sp. Det. P. Paquin. Eisodophile. Class Entognatha We collected one specimen from a web Subclass Collembola within the twilight zone of PARA 2202. Order Entomobryomorpha 10 WESTERN NORTH AMERICAN NATURALIST [Volume 74

Family Entomobryidae Family Leiodidae Entomobrya zona Christiansen and Bellinger, Undetermined species. 1980. Det. E. Bernard. Troglophile. Det. J. Wynne. Troglophile? Specimens from this family were collected Twelve specimens were collected from 2 in the light (n = 3) and twilight (n = 10) zones Parashant caves. The specimens consist of one of PARA 1801; the light (n = 4), twilight (n = from the light zone of PARA 1001 and 10 from 1), and dark (n = 7) zones of PARA 2602; and PARA 2204 (4 from the light zone and 6 from the light zone (n = 4) of PARA 2204. We ob- the twilight zone). This beetle species was ob - served this collembolan in the light zones of 3 served denning in large numbers within the caves and the twilight zones of 2 caves. We sug- fissures in the twilight zone and in association gest this animal is a troglophile because (1) it with Leiobunum townsendii. Most leiodid bee- was found throughout the lengths of 2 caves; (2) tles are habitat generalists and feed as both it is a largely nonvagile edaphic organism, and adults and larvae on certain fungi or microor- (3) collembolans are routinely found in caves. ganisms associated with decaying organic mat- ter (e.g., Majka and Langor 2008). We suggest Class Insecta these beetles may have a commensal relation- Order Coleoptera ship with harvestmen (opilionid spiders) and Family Anobiidae may feed on fungi growing on spiders’ feces. There is a literature, rich with examples of Undetermined species. troglomorphic species, that confirms that leio- Det. C. Drost. Accidental. dids use caves (e.g., Peck 1974, 1978). We sug- We collected 3 specimens from the light gest this morphospecies may be a troglophile. zone of PARA 1801. Most members of the family of death watch or spider beetles occur Family Mordellidae in dry vegetation (Triplehorn and Johnson 2005). Members of this family are occasionally Undetermined species. noted in caves where they may occur in pack- Det. R. Delph. Accidental. rat midden material. Fifteen specimens were collected including 3 specimens from PARA 2602 (2 from the dark Family Carabidae zone and one from the light zone) and 12 speci - mens from PARA 2204 (2 from the dark zone Rhadine n. sp. (Perlevis species group). and 10 from the twilight zone). Mordellid lar- Det. T. Barr. Troglophile. vae feed on decaying wood and vegetation, A total of 6 specimens were collected, con- while adults feed on flowers (Triplehorn and sisting of 3 males and one female from PARA Johnson 2005). Because both of these caves 2204 and one male and one female from PARA were dry and detritus deposition was low, we 2202. These specimens represent an unde- suggest this morphospecies is accidental. scribed species (T. Barr, personal communica- tion, 2007). Family Tenebrionidae Eschatomoxys pholeter Thomas and Pape Family Bruchidae 2007 (new species). Troglophile. Undetermined species. This species was newly discovered and de - Det. R. Delph. Accidental. scribed through this research. This beetle was We collected 2 specimens from the light collected from the twilight zone of PARA 2602. zone of PARA 1801. This seed-boring beetle Pape et al. (2007) considered this beetle a trog- was detected within the cave entrance but lophile. Representing the fourth documented is not expected to use caves unless seeds locality, this species was also collected in Ram- are transported into the cave by aeolian part, Bat, and Christmas Tree Caves, Grand currents. Canyon National Park (Pape et al. 2007). 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 11

Eleodes (Caverneloedes) wynnei Aalbu, Smith communication, 2013). Peck (1981) and Reeves & Triplehorn 2012 (new species). Troglophile. et al. (2000) considered Tipulids detected in This species was newly discovered and de - caves in the SE United States to be trogloxenes. scribed through this research. Three speci- mens were collected from the twilight zone of Order Hemiptera PARA 1801 and the light zone of PARA 1001. Family Reduviidae Also, we collected tenebrionid larvae within PARA 1801. All Caverneleodes species are con- Triatoma cf rubida. sidered troglophiles (Aalbu et al. 2012). Det. J. Wynne. Accidental. We collected one specimen from the light Order Diptera zone of PARA 1001. Given that this species is Family (formerly sanguinivorous and that we didn’t observe any known as Hippoboscidae) vertebrates within this cave, we suggest the occurrence is accidental. Basilia antrozoi Townsend, 1893. Det. C. Dick. Parasite. Order Hymenoptera One female specimen was removed and col - Family Formicidae lected from a captured Antrozous pallidus at Note: All ant morphospecies were detected within cave the entrance of PARA 0802. entrances and in only association with baited pitfall traps. We consider all of these ants to be accidental.

Family Phoridae Solenopsis xyloni McCook, 1879. Det. R. Johnson. Accidental. Undetermined species. Det. J. Wynne. Troglophile? Five specimens were collected from the en- trance of PARA 2602. We collected 2 specimens from the dark zone of PARA 1001. This cave contains cricket Pheidole vistana Forel, 1914. guano and decaying vegetation. Adults of this Det. R. Johnson. Accidental. species may be feeding on the fungus growing We collected 3 specimens from the en- on these 2 nutrient sources. trance of PARA 2602.

Family Sciaridae Paratrechina cf hystrix. Det. R. Johnson. Accidental. Undetermined species. Det. J. Wynne. Troglophile? We collected 2 specimens from the entrance of PARA 1801. R. Johnson (personal commu- We collected one specimen from the dark nication, 2006) suggests it may be P. hystrix. If zone of PARA 1001. This cave contains cricket so, this record is the first for Arizona (R. John- guano and decaying vegetation. Larvae of this son, personal communication, 2006). Paratre- morphospecies are likely feeding on the fungus china hystrix is a northern species known to growing on these 2 nutrient sources. Addition- occur in Nevada and Utah. R. Johnson (per- ally, larvae of some of these sciarid species are sonal communication, 2006) suggests the spec- known to be predaceous (Cole and Schlinger imens match the description in part; however, 1969, Triplehorn and Johnson 2005). No larvae because we did not collect any workers, a reli- were de tected within PARA 1001. able identification is not possible. Pheidole sp. Family Tipulidae Det. R. Johnson. Accidental. Tipula kaibabensis Alexander 1946. One specimen was collected from the en - Det. J. Gelhaus and C. Young. Trogloxene. trance of PARA 2204. R. Johnson indicated One specimen was photographed in the twi- the specimen was a single minor worker. Mi - light zone of PARA 1001, and one was collected nors are often difficult to identify to species, from PARA 0802. Tipulids use dark damp places and the specimen was not examined further for dens during the day (J. Gelhaus, personal (R. Johnson, personal communication, 2006). 12 WESTERN NORTH AMERICAN NATURALIST [Volume 74

Order Orthoptera Order Psocoptera Family Rhaphidophoridae Family Sphaeropsocidae cf Ceuthophilus n. gen. n. sp. Troglosphaeropsocus voylesi Mockford 2009 Det. T. Cohn and A. Swanson. (new genus and species). Trogloxene. Det. E. Mockford. Unknown. Nine specimens (8 females and one male) This animal represents both a new genus were collected from the light and twilight zone and species discovered through this research. of PARA 1001. T. Cohn (personal communica- We collected one specimen from the twilight tion, 2006) indicated that this new genus has zone of PARA 2602 (located in the Mojave clasping cerci with unique short spines on Desert). The cave is dry and dusty with little their base, a subgenital plate that matches no to no aeolian-deposited detritus; however, it is other specimens in his collection, and a dis- used as a summer roost by Antrozous pallidus. tinctive last tergite. Until we have additional information on this psocid’s occurrence within caves, we consider Ceuthophilus n. sp. its cave affiliation to be unknown. Det. T. Cohn. Trogloxene. Family Psyllipsocidae One male was collected from the twilight zone of PARA 2204. T. Cohn (personal com- ramburii Sélys-Longchamps, munication, 2006) suggests that the structures 1872. Det. E. Mockford. Troglophile. on this specimen are unique and that the speci- This species was trapped in the light, twi- men represents an undescribed species. light, and dark zones of PARA 1801 and the Ceuthophilus n. sp. a? dark zone of PARA 2602. Because psocids are Det. T. Cohn. Undescribed? routinely found living in caves, we suggest it is Trogloxene. a troglophile. We collected one female from the entrance of PARA 1801. Although we will require adult Family Prionoglarididae males for confirmation, T. Cohn (personal com- Speleketor flocki Gurney, 1943. munication, 2006) indicated that the female has Det. E. Mockford. Troglophile. distinctive characters and may be undescribed. We collected a nymph of this species from Ceuthophilus n. sp. b? the dark zone of PARA 2602. E. Mockford Det. T. Cohn. Undescribed? (personal communication, 2006) indicated that Trogloxene. although the specimen was a nymph, its head Another female was collected from the en- markings are unmistakable. This specimen trance of PARA 1801. T. Cohn (personal com- represents the third locality for this species in munication, 2006) indicated that the female the western United States. It has been con- has an “extraordinarily elongate and curved ovi- firmed from a cave in the Tucson Mountains positor” and is distinctive and may be unde- and Gypsum Cave, southeastern Nevada. scribed. Male specimens will be required to This psocid routinely lives in caves, and be - confirm this. cause a nymph was found in the dark zone of this cave, we suggest it is probably a troglophile. Undetermined Ceuthophilus sp. Det. T. Cohn. Order Siphonaptera Trogloxene. Undetermined family, genus, and species. There was at least one undetermined im- Det. J. Wynne. Parasite. mature female Ceuthophilus species collected in PARA 1401. Because the specimen was an One specimen was collected from the dark immature, it was not possible to identify it zone of PARA 2602. This cave supports a pos- beyond genus. However, it may also represent sible maternity roost for Antrozous pallidus, a new species (T. Cohn, personal communica- which is likely the host of this siphonapteran tion, 2006). and explains its occurrence. 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 13

Subclass Apterygota the entrance of PARA 2602. At PARA 0802, Order Thysanura we used a combined mist-netting / harp-trap- Family Lepidotrichidae ping approach to capture 7 individuals (2 post- lactating adult females, one nonreproductive Undetermined species. juvenile female, 2 nonreproductive undeter- Det. J. Wynne. Eisodophile. mined females, one nonscrotal juvenile male, We trapped one silverfish within the en - and one nonreproductive adult female). This trance of PARA 2602. roost contained ~50 individuals and was lo - cated in the rock fissures in the ceiling within the cave’s light zone. In PARA 2202, we ob - Phylum Chordata served a bat flying in the twilight zone whose Class Mammalia pelage was consistent with A. pallidus. Al - Order Chiroptera though we did not confirm it was a pallid bat, Family Vespertilionidae the observation likely represented this spe - Myotis sp. cies. While pallid bats are found roosting in Det. J. Wynne. Trogloxene. caves, a majority of data suggest they roost primarily in rock crevices and outcrops (Her- Approximately 3 Myotis sp. were observed manson and O’Shea 1983). during exit counts at PARA 1401. According to Arizona Game and Fish Department’s (AGFD) Corynorhinus townsendii Cooper 1837. Heritage Data Management System, this ob- Townsend’s big-eared bat. servation is likely either M. thyasanodes or M. Det. J. Wynne. Trogloxene. yumanensis. Only M. yumanensis is known to Nine individuals were observed hibernat- roost both in caves and human-made struc- ing in the twilight zone of PARA 1401. This tures (AGFD 2003a, 2003b). Williams et al. was the only documented hibernaculum on (2006) suggest this species is commonly found the Grand Canyon–Parashant National Monu- near small- to moderately sized bodies of wa- ment. Lepidopteran wings were also observed ter, and of the 4 habitat types investigated, this within the entrance of PARA 2602 and PARA species occurs in riparian woodland more than 0802, suggesting night use by this species of in all other habitats combined. Though there bat. Corynorhinus townsendii is a cave-roost- are 2 water tanks within 3 miles of PARA ing bat, but it also roosts in mines and build- 1401, these tanks are intermittent water sources. ings (Kunz and Martin 1982). We suggest the best candidate water source is probably Imlay Reservoir, approximately 8.85 km from the cave. Order Rodentia Family Cricetidae Myotis thysanodes Miller 1897. Fringed myotis. Det. J. Wynne. Trogloxene. Neotoma sp. Packrat or woodrat. Det. J. Wynne and K. Voyles. Troglophile. One nonreproductive female was captured in a harp trap at the entrance of PARA 1801. Midden activity was documented in the en - We estimated roost size between 20 and 30 trances and into the light zones of PARA 2202, individuals. We suggest PARA 1801 may be a PARA 2602, PARA 2204, and PARA 0802. maternity roost. This bat roosts in caves, mines, Woodrats use cave entrances rock outcrops, and buildings (O’Farrell and Studier 1980). rock fissures, and other suitable features for establishing dens. Antrozous pallidus Allen, 1862. Pallid bat. Det. J. Wynne. Trogloxene. Family Erythizontidae Pallid bats were identified in PARA 2602, PARA 0802, and possibly PARA 2202. Two Erethizon dorsatum Cuvier 1822. North individuals (one adult female and one non- American porcupine. scrotal male) were captured and identified in a Det. J. Wynne. Eisodoxene. handheld net in PARA 2602. During our sur- Both PARA 1001 and PARA 1401 have been vey in August 2005, we observed ~100 indi- used extensively by porcupine. In PARA 1001, viduals roosting in the boulder breakdown at guano deposition of 2–10 cm was ob served 14 WESTERN NORTH AMERICAN NATURALIST [Volume 74 throughout the cave; however, there were no tained roosting bats, which provided nutrients signs of recent porcupine use. A fully articu- via guano. PARA 1001 (11 morphospecies) con - lated porcupine skeleton was photo graphed tained water condensation on the ceiling and amid a deep deposition of guano in PARA walls and supported the largest known cricket 1401. Though the North American porcupine den in Arizona (likely on the order of thou- dens in caves, it also will den in other features sands of individuals). Such a large number of offering similar microhabitats (e.g., hollow logs crickets generate a significant nutrient load in and trees). Strong (2006) indicates that this the form of cricket eggs, nymphs, and guano. species is commonly documented in caves in The ecological importance of cave crickets the Chihuahuan Desert, New Mexico. has been widely documented (e.g., Barr 1967, Howarth 1983, Taylor 2003, Culver 2005, Poul - son 2005), and we suggest that the presence Order Lagomorpha of crickets was why this cave supported a Family Leporidae higher arthopod diversity. Additionally, given Lepus californicus Gray 1837. Black-tailed the significant nutrient loading provided by jackrabbit. Det. J. Wynne. Accidental. crickets and the presence of a cave deep zone, PARA 1001 may also support cave-adapted A black-tailed jackrabbit skeleton was found arthropods. approximately 5 m from the entrance of PARA No troglomorphic taxa were identified dur- 1001. This animal likely fell into the cave, ing this survey. Detections of cave-adapted became trapped, and eventually died. The car- taxa are reportedly low for northwestern Ari- cass provided a pulsed food resource for scav- zona. However, there are cave-adapted taxa enging cavernicoles. known regionally. For example, 3 troglobites and one stygobite are known from Grand Can - Class Aves yon National Park (Wynne et al. 2007), and a Order Strigiformes cave-limited millipede (Shear et al. 2009) and a cave-adapted copepod have been collected Undetermined family, genus and species. from the BLM–Arizona Strip lands adjacent to Det. K. Voyles and J. Wynne. Eisodoxene. Parashant (J. Wynne unpublished data). We documented owl pellets within the twi- Concerning more regional patterns, we light zone of PARA 2202. Owls routinely roost identified at least 18 arthropods with strong in cave entrances and other suitable habitats cave affinities (e.g., trogloxenes or troglophiles) both during the day, and also during the night and 21 accidentals or eisodophiles from 7 between hunting outings. caves. Given the arid conditions of the desert Southwest, we suggest that few ground-dwell- DISCUSSION ing arthropods in the Southwest are genetically predisposed to colonizing the often more mesic This study represents the first regional all- cave environment. Barr (1968) suggests that taxa biological inventory of caves in the south- most trog lobites were preadapted to the cave western United States. Our study resulted in environment in that they previously inhabited discovery of at least 7 new species (with 2 new similar mesic habitats such as leaf litter, moss, genera) and 3 potential new species (which or deep soils. By extension, given that these will be described in future publications), as more mesic habitats are nonexistent in the well as 2 range extensions and one possible Mo jave Desert and juniper shrublands within range extension of arthropods. Additionally, the study area, a contemporary preadapted we identified 5 bat roosts and cave use by sev- pool of cave colonists seems to be lacking, and eral other vertebrates. Though our study has thus may be reflected by the low number of contributed significantly to the natural history ground-dwelling cavernicoles observed. of the region, cave biological research within Similarly, Peck (1978, 1980) suggests that Parashant remains incomplete. the low numbers of cave-adapted taxa in Two of the most morphospecies-rich caves, southwestern U.S. caves may be due to the PARA 1801 (14 morphospecies) and PARA low nutrient input and high aridity associated 2602 (13 morphospecies), supported a largely with southwestern cave systems. At Wupatki epigean arthropod community. Both caves con- National Monument, Welbourn (1976) indicated 2014] CAVE DWELLERS OF NORTH RIM GRAND CANYON 15 that moisture was the most limiting abiotic genus, cf Ceuthophilus n. gen. n. sp. Though factor in the earth cracks he sampled. While this genus may occur elsewhere on and off not completely excluding the likeli hood that Grand Canyon–Parashant National Monument, troglomorphic arthropod diversity will be low this has not been confirmed. To increase our in northern Arizona, Wynne et al. (2007) sug- knowledge of the current population and to gested that the low numbers of cave-adapted better define its range, we recommend (1) taxa in Grand Canyon caves may reflect lim- conducting a multiyear census of the popula- ited sampling effort and perhaps inappropriate tion during the most productive time of year— techniques for detecting cave-adapted taxa. between the late monsoonal season and the While troglomorphic taxa in the South- early post-monsoonal season (mid-August to west may be depauperate in comparison to mid-September); (2) sampling additional caves cave-obligate communities in the mesic cen- within Parashant to search for this animal; and tral and eastern portions of the United States, (3) conducting surface surveys in the region to we maintain that sampling effort and ineffec- estimate the actual geographic distribution of tive sampling techniques may still explain the this new cricket. Additionally, we know of only lack of troglomorphic taxa detected in Para- one other cave in Arizona that supports such a shant caves. large cricket population. Because cave crickets For future work, we recommend a multi- contribute important nutrients via guano, eggs, year systematic study design, including an in- nymphs, and cricket carcasses and are a prey creased number of sampling stations per cave source for predaceous arthropods (e.g., Barr for timed searches and baited pitfall trapping 1967, How arth 1983, Taylor 2003, Culver 2005, (see Wynne 2013), as well as direct intuitive Poulson 2005), the cricket population is likely searches and bait stations (provisioned with an important component of this ecosystem. chicken liver, mushrooms, blue cheese, and Given the population size within PARA 1001, sweet potato) within deep cave zones (e.g., we suggest that the presence of crickets likely Howarth et al. 2007). This approach would (a) has an important bottom-up effect on ecologi- provide us with a more thorough inventory cal structure and species richness and also sug - of cave-dwelling arthropods, (b) provide for gest that the actual number of species will be inferential statistical comparisons across sites, higher than the observed richness presented and (c) increase the likelihood of detecting in this study. cave-adapted organisms. Secondly, we suggest We recommend that all caves containing that sample size (i.e., the number of caves both summer roosts and hibernacula (PARA inventoried) may also be an issue. We provide 0802, PARA 1401, PARA 1801, and PARA 2602) the results from only 7 caves within Parashant. be closed to recreational use until these sites In the adjacent BLM–Arizona Strip lands and can be thoroughly studied and their functions Grand Canyon National Park lands, there are determined. There is little argument that hu - approximately 250 and >400 known caves, man disturbance to bat roosts is detrimental respectively. As we expand our efforts into (e.g., Humphrey 1969, Mohr 1972, McCracken other management units in Arizona and obtain 1988, 1989, Harnish 1992, Brown et al. 1993, a larger sample of study sites, we hypothesize Boyles and Brack 2009). Additionally, there that more troglomorphic taxa will be discov- is a growing threat of white-nose syndrome ered. Only through increased sampling effort (WNS), a disease caused by Pseudogymnoas- and use of systematic techniques will we be cus destructans (refer to Minnis and Lindner able to make ecological comparisons to other 2013), which will likely spread through the regions in the southwestern United States and western United States. This psychrophilic fun- infer whether the American Southwest truly gus has resulted in the “most precipitous de - supports low troglomorphic diversity. cline of North American wildlife in the past century” (BCI 2010). Furthermore, WNS has Management Implications resulted in the mortality of over 7 million bats Through this research, we tentatively iden- and has been detected in 23 states and 5 tified one cave as a high management priority Canadian provinces (Wynne 2014). because of the presence of cave-dwelling Little is known concerning the habitat char - arthropods. PARA 1001 supports the only acteristics of bat hibernacula in the Ameri can known type localities for the new cricket Southwest. In Arizona alone, we lack baseline 16 WESTERN NORTH AMERICAN NATURALIST [Volume 74 information regarding the locations of cave- National Monument and vicinity, Arizona. Unpub- roosting hibernating bats (A. McIntire, Ari- lished report to National Park Service, Wupatki Na- tional Monument, Flagstaff, AZ. 18 pp. zona Game and Fish, personal communication, BARR, T.C., JR. 1967. Observations on the ecology of 2010). Thus, resource managers can best pre- caves. American Naturalist 101:475–491. pare for the westward advance of WNS by ______. 1968. Cave ecology and evolution of troglobites. intesifying their efforts to establish population Pages 35–102 in T. Dobzhansky, M. Hecht, and W. estimates of nonmigratory cave-roosting bats Steere, editor, Evolutionary Biology. Volume 2. Appleton-Century-Crofts, New York, NY. and characterize habitat of cave roosts in the [BCI] BAT CONSERVATION INTERNATIONAL. 2010. White nose western United States. syndrome: a crisis for America’s bats, BATFAQ. BCI, To improve our knowledge regarding bat Austin, TX. Available from: http://www.batcon.org/pdfs distributions in northwestern Arizona, we rec- /whitenose/WNSFACTSSHEET_Mar15_2010.pdf BODENHAMER, H. 1989. Impacts to caves within Grand ommend the following: (1) additional surveys Canyon National Park, preliminary observations and of summer roosts (at PARA 0802, PARA 1801, research proposal. Unpublished report to National and PARA 2602) during the middle of the Park Service, Grand Canyon National Park, Grand nursery period (mid- to late June) to deter- Canyon, AZ. 100 pp. BOYLES, J.G., AND V. B RACK JR. 2009. Modeling survival mine whether these roosts are actually mater- rates of hibernating mammals with individual-based nity/nursery sites; (2) establishment of annual models of energy expenditure. Journal of Mammal- or biennial winter bat censuses of the hiber- ogy 90:9–16. naculum cave (PARA 1401); and (3) expanded BROWN, P., R. BERRY, AND C. BROWN. 1993. Bats and searches to identify additional cave roosts, mines: finding solutions. Bats 11:12–13. COLE, F.R., AND E.I. SCHLINGER. 1969. The flies of west- with follow-up inventories, roost monitoring, ern North America. University of California Press, and habitat characterization as appropriate. Berkeley, CA. xi + 693 pp. CORNELY, J., AND R. BAKER. 1986. Neotoma mexicana. ACKNOWLEDGMENTS Mammalian Species 262:1–7. CULVER, D.C. 2005. Species interactions. Pages 539–543 in D.C. Culver and W.B. White, editors, Encyclope- We extend special thanks to Darla Sidles, dia of caves. Elsevier, Burlington, MA. Mark Sogge, Karen Yanskey, and Carol Cham- DROST, C.A., AND D.W. BLINN. 1998. Invertebrate com- bers for their logistic support and adminis - munity of Roaring Springs Cave, Grand Canyon trative guidance. Ben Solvesky provided field National Park, Arizona. Southwestern Naturalist 42: support and assisted with bat identifications. 497–500. GUSTAFSON, T.A. 1964. The bats of Wupatki National The image of Troglosphaeropsocus voylesi was Monument, Coconino County, Arizona. Unpublished provided by Edward Mockford, University of report on file with National Park Service, Wupatki Illinois, Normal. Eric Alejandro Pinto gra- National Monument, Flagstaff, AZ. 12 pp. ciously translated our abstract to Spanish. Jeff HARNISH, H. 1992. Protecting the bats of Devil’s Den: when endangered bats needed special protection Bradybaugh, Charles Drost, Jennifer Fox, Stew - from disturbance, a cave alarm system provided an art Peck, and 2 anonymous reviewers pro- effective and creative solution. Bats 10:13–18. vided valuable suggestions leading to the im - HERMANSON, J.W., AND T.J. O’S HEA. 1983. Antrozous pal- provement of this manuscript. This work was lidus. Mammalian Species 213:1–8. funded by the National Park Service, Grand HILL, C.A., V.J. POLYAK, D.C. BUECHER, R.H. BUECHER, AND C.G. GRAF. 2001. Trip report: Bat Cave, Grand Can- Canyon–Parashant National Monument, ARCS– yon National Park, Arizona, June 11, 2001. Unpub- Phoenix Chapter, and the American Museum lished report on file with National Park Service, Grand of Natural History, Theodore Roosevelt Memo- Canyon National Park, Grand Canyon, AZ. 48 pp. rial Fund. HILL, C.A., V.J. POLYAK, D.C. BUECHER, R.H. BUECHER, AND P. P. P ROVENCIO. 2000. Trip report: Tse’an Kaetan (Cave of Prayer Sticks), Grand Canyon National LITERATURE CITED Park, Arizona, February 11, 2000. Unpublished re - port on file with National Park Service, Grand Can- AALBU, R.L., A.D. SMITH, AND C.A. TRIPLEHORN. 2012. A yon National Park, Grand Canyon, AZ. 46 pp. revision of the Eleodes (subgenus Caverneleodes) HOWARTH, F.G. 1980. The zoogeography of specialized with new species and notes on cave breeding Eleodes cave animals: a bioclimatic model. Evolution 34: (Tenebrionidae: Amphidorini). Annales Zoologica 62: 394–406. 199–216. ______. 1982. 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