2. Springs Dependent Species (SDS)

Springs types have been defined on the basis of gal’s flaveria; – specuicola their sphere of discharge by Springer and Stevens cave-dwelling primrose). More than 170 spe- (2008), who described 12 active springs types cies of Pyrgulopsis springsnails are known from and 13 geomorphologically-defined springs western North America, the majority of which microhabitats. Many microhabitats with asso- occur at the sources of individual springs, and ciated biological assemblages co-occur within among which at least 17 are or have been consid- some individual springs, greatly contributing to ered for listing as endangered species. Leopard biodiversity and species packing (Hallam 2010). frogs (Ranidae: Lithobates spp.) in the Southwest We define springs-dependent species (SDS) as are commonly found at springfed cienegas, fens, those that require, or most often occur in, springs and other helocrene and low gradient hillslope or springs-supported habitats during one or springs (Clarkson and Rorabaugh 1989). Nu- more phases of their life cycles. Some springs merous native fish species in the DLCC are SDS types support endemic (unique) taxa or other (Beckman 1952, Minckley 1973, Sublette et al. SDS, particularly limnocrene (pool- forming), 1990, Sigler and Sigler 1996, Wallace and Zaro- mound-forming, helocrene (marsh-forming), ban 2013). We exclude as SDS those generalist and gushet springs (e.g., Spamer and Bogan species that occasionally (non-obligatorily) occur 1994, Stevens and Meretsky 2008, Stevens and in springs-generated habitats, including non-na- Polhemus 2008, Stevens and Bailowitz 2009). tive crayfish and non-native fish. However, some For example, several endemic subspecies of generalist taxa occur obligatorily in springs in speckled dace (Rhinichthys osculus) and many particular regions and such populations will be pupfish (Cyprinodon spp.) are SDS, occurring in regarded as being conditionally SDS. For exam- hillslope or limnocrene springs run-out channels, ple: American Dipper (Cinclus mexicanus) nest which are springs-supported habitats (Minckley virtually obligatorily in cool spring-supported 1973; Minckley and Unmack 2008). Rheocrene streams when they nest at low elevations in the (in-channel) springs are somewhat less likely to arid Southwest (Stevens et al. 1997); and north- support endemic or rare SDS due to increased ern water shews (Sorex palustris) may occur flooding disturbance. In addition to springs type, primarily along high elevation springfed streams several other factors influence species richness and in wet meadows (Hoffmeister 1986). and the occurrence of SDS at springs, including: springs habitat area, isolation, and geomorphic complexity (Hallam 2010, Schaller 2013). SDS include a wide array of aquatic springs source biota [e.g., Hydrobiidae – most Pyrgulop- sis springsnails; many elmid riffle beetles; many stoneflies (Plecoptera); some freshwater amphi- pods and isopods (e.g., Socorro isopod - Thermo- sphaeroma thermophilum); and many cyprino- dontid pupfish], as well as wetland species (e.g., Orchidaceae – Epipactis gigantea heleborine orchid; Cyperaceae – Carex specuicola Navajo sedge; Asteraceae – Flaveria mcdougalii McDou- 27 Springs-Dependent Species of Grand Canyon

Rotund Velvety Shore Bug Herbert ‘s and Breviceps Giant Water Bugs (: Ochteridae (Hemiptera: Ochterus rotundus Polhemus and Polhemus) herberti herberti Hidalgo and Abedus breviceps Stål) The rotund velvety shore bug is a small (4.5-5.5 mm long), oval-flattened predatory that lives on wet These are two of seeping moss walls at springs in the upper half of the three giant water bug Colorado River corridor. As a member of the order species known from Hemiptera (true bugs, with sucking mouthparts), it Grand Canyon, all of is distantly related to water striders, backswimmers, which occur in desert water boatmen, kissing bugs, and bed bugs. The Pol- springfed streams, and hemus brothers first collected it from Grand Canyon all of which are aquatic at tributary springs during their river trip in 1975, and predators. Herbert’s gi- we have subsequently found it at a few other springs ant water bug is found from Buck Farm downstream to Royal Arch Creek. in lower Grand Can- yon in streams within the ancient drainage of the early Cenozoic “California River”, a river that flowed east and north across the northwestern corner of Arizona. Herbert’s giant water bug is widespread through the Southwest. In contrast, the abedus water bug is found only in a single stream in central Grand Canyon, the only population of that species on the southern Colorado Plateau. Behavioral O. rotundus is predatory, likely feeding on fly and observations on abedus water bugs in southern Ari- other soft-bodied insect larvae that crawl around in zona revealed that it senses on-coming flashfloods, and the moss. It is quite cautious around humans, detecting climbs out of the water and above the flood waters to approaching visitors and adroitly slipping into crevices avoid the scouring effects of summer floods. out of sight until the danger has passed. Hence, it is very easily overlooked. Its habitat—wet, near-vertical, dripping moss walls— is an ancient one that supports some of the most ancestral semi-aquatic on Earth. It is also often relatively protected from floods, which roar down the canyons but leap over falls, leavingthe mossy vertical backwalls unscathed. This species is one of several Grand Canyon springs- dependent insects that are known from central and southern Mexico into Guatemala. This “Guatemala biogeographic connection” means that sometime in the past few thousands or millions of years, similar habitats existed, either in continuous connection or as “stepping stones” of habitat between Central America and Grand Canyon. Abedus breviceps 28 Four-spotted Saldid Shore Bug Apache Spiketail (Hemiptera: Saldidae (: Codulegastridae Micracanthia quadrimaculata (Champion) diadema diadema Selys)

This tiny shore bug is known in the United States The Apache spiketail dragonfly is the largest only from a single alkaline spring in the Inner Gorge and one of the most brilliantly colored creatures on the of Grand Canyon. It is a tiny predator, Colorado Plateau: a jet black body with bright yellow likely preying on soft-bodied inver- banding, and enormous, startlingly brilliant blue eyes. tebrates, such as fly larvae. It patrols One of two western species, the spiketails are particu- steeply sloping bedrock faces, watching larly adapted to life in steep, narrow canyons. This spe- for prey and enemies. M. quadrimacu- cies flies in mid- to late summer and into autumn, and lata is otherwise reported from Central adults are most often found close to montane springs. America and is yet another example of Females lay their eggs in mud at the water’s edge, often the “Guatemala biogeographic connec- at springs sources. They remain in the larval stage for tion” in Grand Canyon. three years, growing to nearly 5 cm (2”) in length, and live in stream bottom mud. In 1917 Clarence Kennedy Grand Canyon Masked Clubskimmer suggested that spiketail larvae likely wash downstream (Odonata: Libellulidae in floods over their multi-year larval lifetime, so start- Brechmorhoga pertinax (Hagen) ing life at the headwaters makes sense. Watch for these remarkable in Deer Creek from August This large, showy dragonfly was discovered in 2003 through October. and the Grand Canyon population of masked club- skimmer dragonfly is the only breeding population in the United States. It lives along flowing springfed streams in Grand Canyon from Nankoweap down- stream to Stone Creek. Its larvae are predators, con- suming mayfly larvae and other soft-bodied stream invertebrates. The type specimen of masked clubskim- mer was collected in Central America, and prior to its discovery in Grand Canyon, this species was only known from two specimens that were thought to have blown into the US from Mexico. Thus, the Grand Can- yon population represents an extremely isolated popu- lation, and genetics analyses are being undertaken to determine whether the Grand Canyon population represents a new subspecies.

Photo courtesy of Arizona Dragonflies Website

29 Springs-Dependent Species of Grand Canyon Southwestern Viceroy Butterfly found in Grand Canyon only in the ancient path of the (Leptidoptera: Nymphalidae early Cenozoic “California River”, a river that flowed Limenitis archippus obsoleta W.H. Edwards) east and north across the northwestern corner of Ari- zona. Nerthra lives at the margin of springfed streams In 1934, David Rockefeller, Sr., then a teenager, in the lower Canyon, crossed Grand Canyon on a rim-to-rim hike on the where it feeds on first major entomological collecting expedition in damselfly larvae and Grand Canyon. Fortunately, he deposited specimens at other soft-bodied in- Grand Canyon and at several other museums, includ- vertebrates. It is rarely ing the American Museum of Natural History in New seen, but is related to York. Mr. Rockefeller captured a southwestern vice- the toad bugs (Gelas- roy butterfly at Phantom Ranch, as well as a number tocoris)—two species of aquatic beetles (he went on to assemble the largest of diminutive but personal beetle collection in the world). However, the much cuter bugs that viceroy butterfly disappeared from Phantom Ranch, hop along the shore- and for many decades was assumed to have been extir- lines of desert streams pated from Grand Canyon. In the Southwest, viceroy in Grand Canyon. butterflies are most often associated with coyote willow (Salix exigua), and that willow species may have been The Grand Canyon Wetsalts Tiger Beetle eliminated from lower Bright Angel Creek by flood- (Coleoptera: Carabidae/Cicindelinae ing in the 1960s. Larry Stevens found viceroys living Cicindela hemorrhagica arizonae Wickham) in Deer Creek Valley in 2009, confirming its continued existence in Grand Canyon. Look for it there around The Grand Canyon wetsalts tiger beetle is endemic the willows in August and September. to Grand Canyon. It is a bronze species with cream In the eastern US the viceroy is renown as a markings, and is reddish underneath. The adults are Műllerian mimic of the bad-tasting monarch butter- agile and found at the edge of springfed streams, such fly (Danaus plexippus), but in the West it mimics the as Hermit Creek, in mid-summer. They sit upright, at- equally bad-tasting queen butterfly (Danaus gilippus). tentively searching for prey and enemies, but are easily alarmed and fly and hide when one approaches closer than about 1.5 meters. The larvae live in tunnels and lunge out at passing prey—mostly soft-bodied insects. This subspecies occurs upstream to Cliff Dwellers Lodge, and downstream to the Virgin River, although the of the downstream population is not clear. It was first discovered in Grand Canyon by Row- land Hayward and C.H. Townsend, who collected it in July 1892 near Hance’s in-canyon cabin site, and David Rockefeller, Sr. collected it at Phantom Ranch in 1934.

Photo courtesy of Tom Checknis Martin’s Nerthra (Hemiptera: Gelastocoridae Nerthra martini Todd)

One of the strangest-looking springs bugs in the Southwest, this 1 cm (1/2”) semi-aquatic predator is 30