U.S.A., Idaho, Boise County, small tributaries of the South Fork of the Payette River near Lowman, ca. 1200 m (3940 ft) elevation, 44.0675°; -115.6822°.

Holotype: a male, deposited in CAS, labeled: “USA: Boise Co., Pine Flat Creek at South Fork Payette River Road, 6 km W of Lowman, 44°04'03"N; 115°40'56"W, T9N, R7E, S31, elev. 1160 m (3805 f), South Fork Payette River Basin, Boise National Forest, 05 May 1999, #26-1999, J. C. Bergdahl, coll.”/ “HOLOTYPE Pterostichus bousqueti Bergdahl designated 2011” [red label]. Paratypes: A total of 11, with 8 males and 1 female with the same data as the holotype, and 2 males labeled: “USA: Boise Co., Archie Creek near confluence with SF Payette R., 8 km E of Lowman, Boise Natn. Forest, 44°04'15"N; 115°31'06"W, T9N R8E S33, elev. 1240 m (4070 ft), South Fork Payette R. watershed, 06 May 1999, #33-1999, J. C. Bergdahl, coll.” The single female paratype is deposited in CAS, the male paratypes in CNC, CMNH, OSU, UICM, WSU, and JCB.

The specific epithet, bousqueti, is a Latinized version (genitive case) of the surname of Dr. Yves Bousquet (Agriculture Canada, Ottawa, Ontario, Canada), in recognition of his outstanding contribution (Bousquet 1985) to our knowledge of Pseudoferonina, which brought order to the chaos in taxonomy of the species known at that time.

A Pterostichus with the characteristics of members of subgenus Pseudoferonina (see Ball 1965, and Bousquet 1999); males easily distinguished from those of all other species of Pseudoferonina by the shape of the median lobe of the aedeagus, especially the apical lamella (apex), which is shaped (in left lateral view) like the blade of a hatchet or tomahawk (Figs 4A and 4C). Also, the shaft of the median lobe in lateral view is thicker in the middle than at either end (where it joins to the base or the apical lamella) and lacks the distinct mid-shaft sinuation (Bousquet 1985, 1992) seen in Pterostichus shulli, Pterostichus spathifer, the following new species(Pterostichus lolo), and, to a lesser degree, Pterostichus smetanai. The small size and thin body form of Pterostichus bousqueti adults are most similar to Pterostichus campbelli adults, which occur only west of the Cascade Range in Oregon.

Size medium-small for subgenus, ABL males 8.5–9.5 mm, female 8.9 mm. Body form slightly slender for subgenus (Fig. 1).

Color. Dorsal body surface black and shiny, pronotal lateral beads piceous, legs, antennae and palpi piceous.

Microsculpture. Head with isodiametric meshes on frons; pronotum with faintly impressed transverse meshes on most of disc, meshes more isodiametric basolaterally; elytra with markedly transverse meshes, faintly iridescent in some areas.

Pronotum. Fig. 3A. Almost as long as wide, lateral margins subparallel anterior to hind angles, slightly sinuate, hind angles rectangular or slightly acute, lateral margins (in lateral view) bent ventrally near hind angles. Anterior and posterior transverse impressions present but faintly impressed, median line distinctly impressed, almost entire, but not quite extended to anterior and posterior margins in most specimens examined; anterior transverse impression faintly punctulate. Lateral margins finely beaded, posterior margin without margination, anterior margin minutely beaded in medial one-third. Basolateral fovea deep and broadly linear, slightly convergent toward midline anteriorly, not or only faintly punctulate.

Elytra.Intervals flat or only slightly convex, but less so than in any other Pseudoferonina species.

Legs. Male mesotibiae slightly curved apically.

Abdomen. Last visible sternite of male with broad, shallow medial indentation, anterior margin of depression not at all carinate, sternite without protuberances; last visible sternite of males with single pair of anal setae, female with two pairs.

Male genitalia. Median lobe of aedeagus as in Figs 4A and 4C. Apical one-third of median lobe (in ventral view) with lightly sclerotized oblique band extended longitudinally next to slight ridge (Fig. 4C); shaft of median lobe (in ventral view) swollen submedially on left, with shaft gradually tapered from swelling to the apical lamella, ventral margin not sinuate subapically (in lateral view); apical lamella average in length for Pseudoferonina species in Idaho, symmetrically hatchet-shaped (in left lateral view). Right paramere short and sharply pointed.

At this time, Pterostichus bousqueti is known only from Pine Flat Creek and Archie Creek near their confluences with the South Fork of the Payette River near Lowman, Boise County, Idaho (Fig. 5). The Payette River is a tributary of the Snake River, which is a major tributary of the Columbia River. To what extent the range of this species is narrowly restricted to this area is not known. Despite sampling many other creeks in this area, JCB has been unable to document any other Pterostichus bousqueti populations. Additional areas, if any, in which this species is most likely to occur include the Big Pine, Deadwood, Clear Creek and Middle Fork of the Payette River watersheds to the west of Lowman and east of the North Fork of the Payette River.

The type locality along the South Fork of the Payette River near Lowman is in a steeply sided, sparsely pine-forested gorge (Fig. 6A) that has experienced intense forest fires. From 1908 to 2000, canopy fires burned more than 50% of the Boise National Forest in central Idaho. Large fires have a huge effect on vegetation, geomorphology, stream hydrographs (Pierce et al. 2004), and undoubtedly Pterostichus bousqueti habitat. Fire legacies of this extent on the landscape probably create severe barriers to dispersal of Pseudoferonina species. Areas at low to mid-elevations along the South Fork of the Payette River form a small, isolated outlier of the Hot Dry Canyons eco-region of the Idaho Batholith (McGrath et al. 2002). Most of this eco-region is associated with the main stem Salmon River, 145 km (90 miles) to the north. McGrath et al. (2002) describe this eco-region as: unglaciated, deep, precipitous canyons; annual precipitation = 305– 889 mm (lowest in deep canyons); mean temperature: Jan. (min/max) = -8.9/0.6°C, July (min/max) = 8.8/31.7°C (warmer with increasing canyon depth). The vegetation is characterized by open ponderosa pine forest and sagebrush scrub on south facing-slopes, and ponderosa pine/Douglas fir forest on north-facing slopes.

Both Pine Flat Creek (Fig. 6B) and Archie Creek are small, perennial streams descending steep canyon slopes. Their lower sections run thru dry open ponderosa pine forests. At higher elevations on sun and wind-exposed south-facing slopes, the vegetation is primarily open scrub. Pine Flat Creek descends a steep scrubby hillside; pine forest is restricted to lower reaches near the Payette River. Archie Creek descends its watersheds at a gentler grade. Both of these creeks near their confluence with Payette River are only about 1.5 m bank-full-width, and are confined primarily to channels with stabilized vegetated banks. Consequently, there are very few gravel bars on which to search for beetles, even at low water. High water for these creeks most likely occurs March-May, during rapid melt of the snow at higher elevations on adjacent hillsides coincident with wet spring storms. Because the watersheds these creeks service are so small, large thunderstorms and cloudbursts, by themselves, also may cause significant peaks in runoff during summer and fall months after the snow pack has melted at high elevation. In exceptionally dry years, both of these creeks may actually dry up to the extent that even lower sections are without surface-flowing water. However, on 25 September (2008) both creeks had nearly one meter of surface water stream width. Given their small watersheds, it is likely that major sections of both streams, especially high up where slopes are steep and the vegetation scrubby, dry up even in average years.

Other carabid species collected by JCB at or near the type locality include: forest habitat - Psydrus piceus LeConte, Pterostichus protractus LeConte; streamside habitats - Platynus brunneomarginatus (Mannerheim), Bembidion iridescens LeConte, Bembidion kuprianovi Mannerheim, Bembidion sp., Diplous aterrimus (Dejean), Nebria sp., Bradycellus californicus (LeConte), Bradycellus nigrinus (Dejean), Bradycellus nubifer LeConte. The trout-stream beetle Amphizoa insolens LeConte (Coleoptera, Amphizoidae) also occurs along creeks in this area.

JCB visited Pine Flat Creek and Archie Creek on two occasions: 05–06 May (1999) and 25 September (2008). In early May, he easily found adults of Pterostichus bousqueti at both sites, however in September collecting along these same stream sections on both creeks yielded no specimens. Pseudoferonina populations in hot dry regions at low elevations appear to aestivate or go into early hibernation, probably in August. Very few teneral specimens have been collected; however, males with soft, poorly chitinized median lobes are common in late summer and fall. This suggests that adults emerge from pupal chambers in late summer through early fall and overwinter as adults. Presumably they escape winter-spring floods by migrating out of the floodplain to higher positions along stream banks to hibernate in the fall before freeze-up, however they probably do not go very far. Once snow packs melt and daytime temperatures approach 10°C (late March and early April at lower elevations in Idaho), adults again become active on the surface. Given the fact that surface-active Pseudoferonina adults are so closely associated with the immediate vicinity of stream margins, the larvae are probably also narrowly restricted to these habitats and may spend a substantial amount of time subsurface at or very near the hyporheic zone.

All known individuals are brachypterous (flightless) and restricted to the margins of small, isolated, forested streams, so dispersal power of Pterostichus bousqueti adults is expected to be extremely low. Like Pterostichus shulli, Pterostichus spathifer and Pterostichus lolo sp. n., the geographic range of Pterostichus bousqueti (Fig. 5) appears to be highly restricted.

Among Idaho’s Pseudoferonina species, Pterostichus bousqueti appears to be a member of the lanei species-group (with Pterostichus lanei), as opposed to the shulli species-group (Pterostichus shulli, Pterostichus spathifer and Pterostichus lolo sp. n.). Idaho species in the shulli group have a distinct ventral sinuation (bulge and concavity) on the ventral surface of the median lobe of the aedeagus near the middle of the shaft (easily seen in lateral view). Species in the lanei group (and Pterostichus vexatus) do not have this feature. To what extent this feature, or its absence, is indicative of phylogenetic relationship is presently unknown.

Property ownership in the South Fork Payette River area in vicinity of the type locality is primarily the U.S. Forest Service (Boise National Forest), however according to the Boise National Forest map there are many small in-holdings along the river. Two of these in-holdings include both lower Pine Flat and lower Archie creeks, near where the type specimens were collected. Both of these creeks probably have experienced prospecting or mining activity. There is a small Forest Service campground and hot springs very near Pine Flat Creek on the north bank of the Payette River, but visitors seem to have little impact on the creek.

Because Pterostichus bousqueti is known from only two easily identifiable localities, we hope that collectors will exercise restraint when sampling at these sites, and instead focus on discovering new sites for this species in the immediate vicinity, such as in those watersheds listed above.

U.S.A, Idaho, Clearwater County, Cottonwood Creek near the confluence of Orogrande Creek and the North Fork of the Clearwater River, ca. 870 m (2860 ft) elevation, 46.5528°; -115.5522°.

Holotype, a male, deposited in CAS, labeled: “USA: IDAHO, Clearwater Co.; Cottonwood Creek @ USFS Rd 250; Orogrande Ck. Rd.; NF Clearwater R. watershed; elev. ~870 m (2860 ft); 12.5 MI. ESE Headquarters, 3.5 MI. SW of Bungalow; 46°33'10"N, 115°33'08"W; T38N, R7E, S34; 17 July 2009; #73-2009; J. C. Bergdahl, coll.”/ “HOLOTYPE Pterostichus lolo Bergdahl designated 2011” [red label]. Paratypes: A total of 22 paratypes from same locale as the holotype (7 males and 5 females from 17 July 2010; 2 males and 9 females from 03 July 2010) deposited in CAS, CMNH, CNC, JCB, OSU, UICM and WSU.

The specific epithet, lolo, is a noun in apposition, and refers to the Lolo Trail, an ancient hunting and trade route developed by the Nez Perce tribe to cross the Bitterroot Mountains from their homelands on the Clearwater River near Kamiah, Idaho, to the buffalo herds of Montana. The beginning of this trail in its western foothills starts in the upper Lolo River watershed, near the type locality of Pterostichus lolo. The Lolo Trail is also the route the Lewis and Clark Expedition took over the Bitterroot Mountains heading west in September 1805 and east in June 1806.

A Pterostichus with the characteristics of members of subgenus Pseudoferonina (see Ball 1965; and Bousquet 1999); males easily distinguished from those of all other species of Pseudoferonina by the shape of the median lobe of the aedeagus, especially the form of the mid-shaft sinuation and associated convexity (Figs 4B and 4D), which is markedly and abruptly convex, nearly tuberculate (in lateral view), and the shape of the apical lamella (apex), which is broader in ventral view than in Pterostichus shulli males (see Bousquet 1992, Fig. 16) and narrower in lateral view than in Pterostichus spathifer males (see Bousquet 1992, Fig. 15).

Size medium-large for subgenus, ABL males = 9.5–10.5 mm, females = 9.0–10.3 mm. Body form average for subgenus (Fig. 2).

Color. Dorsal surface black and shiny, pronotal lateral beads piceous, antennae and palpi rufous or rufopiceus, femora piceous except distally rufopiceous, tibiae and tarsi rufopiceus or rufous.

Microsculpture. Head with faintly impressed isodiametric meshes on frons; pronotum with moderately impressed transverse meshes on most of disc, meshes more isodiametric posteriorly; elytra with moderately impressed transverse meshes and very slight iridescence. Pronotum. Fig. 3B. Almost as long as wide, widest point approximately at anterior one-third, lateral margins slightly sinuate in posterior half, almost parallel before hind angles, hind angles slightly obtuse to subrectangular, lateral margins (in lateral view) bent ventrally near hind angles, basal one-quarter of pronotum finely but distinctly punctate. Anterior and posterior transverse impressions faintly or not at all impressed, median longitudinal impression distinctly impressed, nearly extended to anterior and posterior margins in most specimens examined. Lateral margins finely beaded, posterior margin without margination, anterior margin minutely beaded in lateral thirds, without margination in medial one-third. Basolateral fovea deep and broadly linear, slightly convergent toward midline anteriorly, finely but distinctly punctate, areas between basal fovea and lateral margins convex, subcarinate.

Elytra. Intervals nearly flat, striae entire, moderately impressed, not or only very faintly punctulate. Legs. Male mesotibiae slightly curved apically. Abdomen. Last visible sternite of male with broad, shallow medial indentation, anterior margin of depression slightly carinate, sternite without protuberances; last visible sternite of males with single pair of anal setae, female with two pairs.

Male genitalia. Median lobe of aedeagus as in Figs 4B and 4D. Apical one-third of median lobe (in ventral view) with lightly sclerotized oblique band extended longitudinally next to slight ridge (Fig. 4Da); shaft of median lobe (in ventral view) markedly swollen submedially on left, with shaft abruptly tapered from swelling to the apical lamella, ventral margin distinctly sinuate subapically (in lateral view) with markedly abrupt, subtuberculate convexity basad of sinuation (Fig. 4Db); apical lamella average in length for Pseudoferonina species in Idaho, apex evenly rounded, very slightly swollen in ventral view (Fig. 4Da), narrow and very slightly reflexed ventrally at tip in lateral view (Fig. 4Db). Right paramere as illustrated by Bousquet (1992, Figs 15b and 16b) for Pterostichus spathifer and Bembidion shulli.

Currently, Pterostichus lolo is known only from the type locality on Cottonwood Creek (at 870 m), which is a small, low-order tributary of Orogrande Creek, which is itself a tributary of the North Fork of the Clearwater River. In turn, the Clearwater is a tributary of the Snake River, which is a major tributary of the Columbia River. To what extent the range of this species is narrowly restricted to this area is not known. Despite sampling many other creeks in this area over the past decade, JCB has been unable to document any other Pterostichus lolo populations.

The type locality is in the Clearwater Mountains and Breaks section of the Northern Rockies eco-region (Nesser et al. 1997). This area has mountains of moderate relief with rounded landform, and steep breaklands. Elevations range between ca. 610 and 2070 m (2000 and 6800 ft.). The climate is generally cool, with a significant maritime influence. Summers are warm and dry, winters cool and moist. The soils are mantled by a thick layer of Quaternary volcanic ash and colluvium, underlain by Tertiary granitic rocks of the Idaho Batholith and Precambrian gneiss and schists (McGrath et al. 2002). Annual precipitation is ca. 889– 2032 mm, with 40–50% falling as snow. Snow packs at higher elevations can become very deep and linger late into spring. Rain-on-snow events are common below 1370 m (4500 ft.); January min/max temperatures: -8.3/0.6°C; July min/max temperatures are 7.2/27.2°C.

These mountains have been largely unglaciated and support a highly dissected stream network of ancient narrow valleys and canyons. Drainage density (length of stream/area) is high. The natural vegetation is primarily a very species-rich mosaic of mesic conifer forest, including exceptional conifer species diversity (grand fir, Douglas fir, western red cedar, western hemlock, mountain hemlock, ponderosa pine, lodgepole pine, white pine, western larch, subalpine fir, Englemann spruce and yew). Ponderosa pine/Douglas fir and cedar/hemlock/pine forests occur at lower elevations, and spruce/fir or mountain hemlock forests at higher elevations. Some of the forest types in this area are the best examples of inland temperate rainforest in the Rocky Mountain region (Bergdahl 2008), including western red cedar trees estimated to be more than 3000 years old (Smith and Fischer 1997). The flora of this region has many unique coastal disjunct species and the area is often referred to as the Clearwater Refugium (Brunsfeld et al. 2001, Carstens et al. 2005, Brunsfeld and Sullivan 2006).

Like all other species of Pseudoferonina, Pterostichus lolo appears to be a habitat specialist, primarily on the wet margins of small, low (0-2) order forest streams descending hillsides in mountainous country. These habitats are often isolated in headwater basins, or on breaks and canyon walls at lower elevations. These beetles are not aquatic, but they are strict hygrophiles. Surface-active adults are usually found within two meters of wet stream channels, primarily within one meter of the strandline and especially right at the water’s edge. When disturbed during warm weather, they will often run into the water and crawl down into the gravel or float away.

Other carabid species collected by JCB at or in the vicinity of the Cottonwood Creek locality include: Scaphinotus (Pseudonomaretus) regularis (LeConte), Scaphinotus (Pseudonomaretus) relictus (Horn), Scaphinotus (Pseudonomaretus) merkelli (Horn), Zacotus matthewsii LeConte, Trechus coloradense Schaeffer, Bembidion kuprianovi, Bembidion iridescens, Bembidion breve (Motschulsky), Pterostichus (Hypherpes) ecarinatus Hatch, Pterostichus (Leptoferonia) idahoae Csiki, Pterostichus (Leptoferonia) beyeri Van Dyke, Pterostichus (Pseudoferonina) shulli (Hatch) and Pterostichus (Pseudoferonina) vexatus Bousquet.

Adults of Pterostichus lolo are probably active from the first warm days of spring after snowmelt to the first days of hard frosts in October. A single pair of adults was observed in copulo, on 17 July 2009 [JCB]. Pterostichus lolo, and all other Pseudoferonina species, are probably “spring breeders” (Bousquet 1986)—that is, they breed early in the active season, larval development takes place in late spring-summer, new adults emerge from pupal chambers in late summer-early fall, and individuals overwinter primarily as immature adults. Males collected in fall are usually immature, suggesting they may die after a single breeding season. These beetles probably vacate streambeds and floodplains in the fall so as to escape winter or spring freshets of high water. Flash floods from thunderstorms during the active season may be a major cause of mortality. The ecology of the larvae is poorly known because they are rarely seen on the surface. The extent to which larvae and adults of Pseudoferonina utilize the subterranean hyporheic zone may be substantial. As mentioned above, adults do not hesitate to crawl down into saturated streambed gravels to escape capture. The highest density of adults is often found along small creeks and reaches that have recently experienced small debris slides with rocks embedded in substantial amounts of wet organic mud. Presumably these muddy microhabitats also support the highest larval density and survival.

All known individuals are brachypterous (flightless) and restricted to the margins of a small, isolated, forested stream, so dispersal power of Pterostichus lolo adults is expected to be extremely low. Like Pterostichus shulli, Pterostichus spathifer and Pterostichus bousqueti sp. n., the geographic range of Pterostichus lolo (Fig. 5) appears to be highly restricted.

Based on the form of the median lobe of the male aedeagus, members of Pterostichus lolo appear to be most closely related to those of Pterostichus shulli and Pterostichus spathifer, (i.e., the shulli species-group). It is perhaps significant that both of these species, along with the more distantly related Pterostichus vexatus, occur in areas near the single known locality for Pterostichus lolo. The North Fork Clearwater River basin has the highest diversity of Pseudoferonina species.

Property ownership in the vicinity of type locality is primarily the U.S. Forest Service (Clearwater National Forest). Potlatch Corporation, Bennett Tree Farms, Inc. and the State of Idaho also have significant property ownership in this region. The primary land use impacts in this area are timber harvesting, roads, mining and off-road vehicle use. There is a large, poorly managed campground on U.S. Forest Service property along both sides of Cottonwood Creek at the type locality, with substantial off-road vehicle impacts.

Because Pterostichus lolo is known from only one, easily identifiable locality, we hope that collectors will exercise restraint when sampling at this site, and instead focus on discovering new sites for this species in the immediate vicinity.