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Description, Bidirectional flow between surface water and sediment leads to high accumulation of small and lightweight microplastics in rivers., In rivers, small and lightweight microplastics are transported downstream, but they are also found frequently in riverbed sediment, demonstrating long-term retention. To better understand microplastic dynamics in global rivers from headwaters to mainstems, we developed a model that includes hyporheic exchange processes, i.e., transport between surface water and riverbed sediment, where microplastic retention is facilitated. Our simulations indicate that the longest microplastic residence times occur in headwaters, the most abundant stream classification. In headwaters, residence times averaged 5 hours/km but increased to 7 years/km during low-flow conditions. Long-term accumulation for all stream classifications averaged ~5% of microplastic inputs per river kilometer. Our estimates isolated the impact of hyporheic exchange processes, which are known to influence dynamics of naturally occurring particles in streams, but rarely applied to microplastics. The identified mechanisms and time scales for small and lightweight microplastic accumulation in riverbed sediment reveal that these often-unaccounted components are likely a pollution legacy that is crucial to include in global assessments.

DROMOCHORUS VELUTINIGRENS JOHNSON, 1991 (FIG. 9G) Common name Velvet tiger beetle. Type locality ���10 km east of Riviera, Kleberg Co, Texas���. Syntypes (3) in USNM, Washington DC. Distribution Dromochorus velutinigrens is currently known from south and west Texas, from the Gulf Coast south of Corpus Christi, west to Dimmit County. Few localities are known (Fig. 3), and gaps between the known occurrences may be due to a lack of sampling in the intervening areas. It is likely that D. velutinigrens also may be found in adjacent areas of Tamaulipas, Mexico. In the vicinity of LaSalle and Dimmit Counties, D. velutinigrens and D. chaparralensis come in close proximity, and are sympatric in at least one site. Diagnosis This is a very distinctive species of Dromochorus. Dromochorus velutinigrens can be diagnosed by having strong blue, violet or green reflections throughout the entire dorsal surface, especially towards lateral margins, in conjunction with a narrow gracile body form and a pronotum with few to no setae. Males are unique among Dromochorus for having an all dark labrum. The only species that could potentially be confused with D. velutinigrens is D. welderensis. Dromochorus welderensis is black dorsally, with a faint dark-blue sheen. It has scattered to regular white decumbent setae present on the disk of the pronotum, especially along lateral margins. Males have a pale central spot on the labrum. Body form is more robust than D. velutinigrens. Description Medium- to large-sized Dromochorus. Body length 11.4���14.9 mm, mean ♀ 13.3 mm, mean ♂ 12.7 mm. Head slightly wider than pronotum. Head charcoal black with velvety violet to blue sheen, with bright violet, blue or greenish reflections in supraorbital areas and anterior margin. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae bright polished metallic blue to violet, blending to violet posteriorly, with shallow longitudinal striae gradually ending at border of vertex. Clypeus bright metallic blue and violet. Male labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections, rarely with a faint ochre-testaceous spot in centre; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently darkbrown with metallic violet and green reflections; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short, white setae, antennomeres 5���11 dulltextured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.2���3.1 mm in length, mean ♀ 2.8 mm, mean ♂ 2.7 mm; width 2.3���3.2 mm, mean ♀ 2.9 mm, mean ♂ 2.7 mm. Pronotum charcoal black, with velvety violet to blue sheen, slightly wider than long, widest near anterior margin, width to length ratio 0.9 to 1.1, pronotal setae absent or with few, irregular, long setae scattered throughout disk; disc smooth, with thin but distinct median line, shallow sulci present anteriorly, and present but less well-defined posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum metallic blue to violet reflections, glabrous. Elytra elongate, 6.7���9.0 mm length, mean ♀ 8.2 mm, mean ♂ 7.7 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral dorsal surface convex, texture dull throughout, elytral coloration charcoal black with velvety violet to blue sheen, lateral margins and apex with shining blue, violet or green reflections; elytral maculations absent; infuscations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae black with iridescent blue, violet and green reflections, with numerous setae, fewer on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora metallic violet to blue, with green reflections, densely clothed in decumbent white setae; tibiae brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter black with metallic violet to greenish reflections throughout most surfaces. Decumbent white setae present on ventrite 1. Ventrites 2���6 have scattered short brown recumbent setae present throughout, but often abraded. Ecology/natural history Adults are active earlier than other species in the genus. Records are from mid-April through late June, but year to year, emergence dates are highly variable relative to congeners and may be more dependent on spring rainfall patterns. Our limited knowledge of habitat associations for Dromochorus velutinigrens is based on two populations. The type locality is a Texas A&M University��� Kingsville���s Site 55 Biological Research Station (Johnson, 1991) on the northern shore of Baffin Bay in Kleberg County, TX. Much of the habitat along the northern banks of the bay has been destroyed by agricultural use, primarily heavy grazing. Johnson (1991) reported that the species can be found associated with sandy road paths and grassy areas along semi-forested areas through the sites. Our team observed adults in sandy, vegetated backshore regions near the bay just below shrubby clay dunes (lomas). Individuals can be found running in and around bunches of cord grass (Spartina spartinae) and in relatively open sandy areas not far from the water���s edge. The adults of D. velutinigrens are by far the most sand-tolerant of the known Dromochorus, as indicated by their presence also on sandy backshore areas near vegetated dunes. Clay lomas are prevalent along the southern Texas coast from St. Charles Bay through northeastern Mexico along the coast of Tamaulipas to Rancho Tepehauje (Tunnell et al., 2002). Future surveys need to focus around clay dune formations in t h e s e r e g i o n s w h e r e t h e b e e t l e i s c u r r e n t l y undocumented. Dromochorus velutinigrens also occurs more i n l a n d, a s f a r w e s t a s C h a p a r r a l W i l d l i f e Management Area in Dimmit and LaSalle Counties. In these areas, we believe that the species��� presence may be explained by the Dilley soil series. These orange/reddish soils are classified as fine sandy loams and are darker in contrast to the type locality in Kleberg County. Extreme soil colour variability has also been observed across the ranges of D. belfragei and D. pruininus. The Dilley series extends to the North into Zavala County. Although unconfirmed, the range for D. velutinigrens will likely extend into this county as well. This location is part of the greater Gulf Coastal Plain physiographic province, and may explain historical connectivity between inland and coastal populations of D. velutinigrens. It is probable that this inland population is not disjunct but, instead, that there has not been sufficient sampling in the intervening areas. Targeted surveys may yield other populations in sandy formations in Mexico and southern Texas. Adults can be found among ghost crab (Ocypode quadrata) colonies, which D. velutinigrens may use for escape when pursued. The ability to hide in cracks for escape appears to be widespread throughout the genus., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 276-278, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237, {"references":["Johnson WN. 1991. A new species of Dromochorus from southern Texas (Coleoptera: Cicindelidae). Cicindela 23: 49 - 54.","Tunnell JW, Judd FW. 2002. The laguna madre of texas and tamaulipas (Vol. 2). College Station: Texas A & M University Press."]}

DROMOCHORUS CHAPARRALENSIS DURAN, ROMAN, HERRMANN & EGAN SP . NOV . (FIG. 9E) Common name Chaparral tiger beetle. Type locality Carrizo Springs, TX. Holotype (deposited in NMNH): 1 ♀, Carrizo Spgs/Tex. V-27–32 // E. G. Lindsley Collector // M.A. Cazier/Collection. Paratypes: 3 ♂♂, 5 ♀♀, Carrizo Spgs./Tex VI-12–32 // E.G. Lindsley Collector // A. Nicolay collection 1950. 1 ♀, Mexico, Tamaulipas/Nuevo Laredo/20-VI-2010/leg J. Stamatov. 1 ♀, USA: Texas: LaSalle Co./Chaparral W.M.A./19-VI- 2013/E. San Gregario. Fig. 9E, Map Fig. 3 Distribution Inland South Texas (currently known from Dimmit, LaSalle, and Webb Counties) and Tamaulipas, Mexico. This species has not been well sampled and is likely present in adjacent areas within the Gulf Coastal Plain, especially in northern Mexico. It can occur with D. velutinigrens in places where sandy soils mix with the dominant heavier red clays. Diagnosis Dromochorus chaparralensis is a robust, dark beetle, and most specimens are dull black with little colour on dorsal or ventral surfaces. Some specimens may have bluish reflections along the margins. It is restricted to mesquite chaparral in South Texas. This species is most likely to be confused with D. minimus or welderensis. Dromochorus mimimus is usually smaller (Fig. 6; Table 2), and possesses a prominent frosted ashy grey, beige, or blue sheen on the dorsal surface; the pronotum has sparse thin erect setae. Dromochorus welderensis is black with a faint dark blue sheen dorsally, in many individuals. The habitat of D. welderensis is Gulf Coast Prairie grasslands, as opposed to the more inland and forested mesquitechaparral habitat of D. chaparralensis. Description Medium- to large-sized Dromochorus. Body length 12.5–14.5 mm, mean ♀ 13.7 mm, mean ♂ 13.0 mm. Head slightly wider than pronotum. Head black with metallic blue to green reflections mostly limited to the lateral ridge of the supraorbital region. Fine to marked rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black often with weak metallic green to violet reflections, with shallow, longitudinal striae, gradually ending at border of vertex. Clypeus shining black, apparently lacking coloured reflections. Male labrum tridentate with 6–8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-quarter of the total labrum surface; female labrum tridentate with 6–8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently dark brown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2–3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3–4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5–11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.6–3.4 mm in length, mean ♀ 3.1 mm, mean ♂ 3.1 mm; width 2.9–3.5 mm, mean ♀ 3.4 mm, mean ♂ 3.1 mm. Pronotum dull black, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.1, setae sparse, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black, lacking prominent metallic coloured reflections, glabrous. Elytra elongate, dorsal surface convex, 7.2–8.6 mm length, mean ♀ 8.2 mm, mean ♂ 7.7 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration black, may have weak blue reflections along lateral margins; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae black, without or with minimal metallic reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet reflections, densely clothed in decumbent white setae; tibiae brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyishwhite setal pad. Abdomen: Venter mostly black with metallic olive green and violet reflections. Decumbent setae present on ventrite 1. Ventrites 2–6 have sparse short brown erect setae present throughout, but often abraded. Etymology Named for the dominant mesquite-chaparral plant community found throughout the species range. Also, this species has been collected at Chaparral Wildlife Management Area, in Cotulla, Texas. Ecology/natural history Very little is known about this species, as most museum specimens are older, and nothing has been recorded about its ecology. The authors of this study have not observed D. chaparralensis in situ.

DROMOCHORUS WELDERENSIS DURAN, HERRMANN, ROMAN & EGAN SP . NOV . (FIGS 8B, 9F) Common name Gulf prairie tiger beetle. Type locality Welder Wildlife Foundation, Sinton, TX. Holotype (deposited in NMNH): 1 ♂, Texas: San Patricio Co. / Welder Wildlife Foundation / 11-June-2013 /Coll: A. Mitchell. Paratypes: 18 ♂♂, 15 ♀♀, Texas: San Patricio Co./Welder Wildlife Foundation/11-June-2013/ Coll: A. Mitchell. 2 ♂, 2 ♀, Texas/Buckeye - Matagorda Co./6-8-17 // J.D. Mitchell collector (NMNH). 2 ♀, Texas: San Patricio Co./Sinton/14-V-1966 // leg. W.T. Murray (JSC). 2 ♂, Texas, Victoria/VI-2���06 // C.R. Jones collector. 1 ♀, Texas: Dickinson/May 29 // TAMU-ENTO X0898573 (TAMUIC). 1 ♂, 1 ♀, Texas: Bee Co./Pettus/10.V.1964 // Leg. Pryor (SFASU). 4 ♂, 1 ♀, Texas: Nueces Co./Luetgens Coll. (AMNH). 1 ♀, Texas: Corpus Christi/VI-7-1969/ C.W. Griffin // Nueces River Park (NMNH). Distribution Found in the Gulf Prairie ecoregion of coastal Texas, from Houston area to Corpus Christi area. Diagnosis Dromochorus welderensis is diagnosable by having a black dorsum, often with a faint dark-blue sheen, and no pitting, subsutural foveae or infuscations, in conjunction with all dark maxillary palps and a pronotum with decumbent white setae. This species is most likely to be confused with D. chaparralensis, belfragei, velutinigrens or minimus. Dromochorus chaparralensis may be nearly indistinguishable from D. welderensis morphologically, but is ecologically differentiated. The habitat of D. chaparralensis is forested mesquite-chaparral, unlike the Gulf prairie habitat of D. welderensis. Dromochorus belfragei possesses regular pits on the elytra and often subsutural foveae. Maxillary palpi have a contrasting dark apical segment, with other segments dark yellow-testaceous to dark red-testaceous. Dromochorus velutinigrens has a very prominent green, blue, or violet dorsal sheen. The body is substantially more narrow and gracile, especially in males. Male D. velutinigrens have an all dark labrum, whereas D. welderensis males possess a pale central spot. Dromochorus velutinigrens have few to no setae on disk of pronotum. Dromochorus mimimus is generally smaller (Fig. 6; Table 2), and possesses sparse, thin, erect setae on the pronotum. This species also occurs further inland in forested mesquite-chaparral, unlike the Gulf Prairie grassland habitat of D. welderensis. Description Medium to large-sized Dromochorus. Body length 10.9��� 14.7 mm, mean ♀ 13.7 mm, mean ♂ 12.6 mm. Head slightly wider than pronotum. Head charcoal brownblack with metallic green, green-blue, or bronze reflections mostly limited to the lateral ridge of the supraorbital region. Fine to marked rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black often with metallic green to violet reflections, with shallow longitudinal striae gradually ending at border of vertex. Clypeus black, with metallic violet to green-coloured reflections. Male labrum tridentate with 6���8 setae, central area pale ochre-testaceous, with a thin, dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-quarter of the total labrum surface; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently darkbrown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5���11 dulltextured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.6���3.3 mm in length, mean ♀ 3.1 mm, mean ♂ 2.9 mm; width 2.7���3.4 mm, mean ♀ 3.2 mm, mean ♂ 2.9 mm. Pronotum charcoal brown to black, slightly wider than long, widest near anterior margin, width to length ratio 0.9 to 1.1, setae sparse to regular, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black, with metallic violet reflections, glabrous. Elytra elongate, dorsal surface convex, 6.3���8.6 mm length, mean ♀ 8.1 mm, mean ♂ 7.4 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration charcoal brown to black, often with faint blue reflections throughout elytral surface; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae brown to black, with metallic violet to blue reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet reflections, densely clothed in decumbent white setae; tibiae brown, clothed with setae of two types: sparser brown-testaceous long setae and dense, short, decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter mostly black with metallic olive green and violet reflections. Decumbent setae present on ventrite 1. Ventrites 2���6 have sparse, short, brown erect setae present throughout, but often abraded. Etymology Named for the type locality,WelderWildlife Foundation, in Sinton, Texas, as well as the Foundation���s namesake, Robert H. Welder, who established the foundation with the mission to conduct research and education in wildlife management and conservation. Ecology/natural history Adults have a long activity period, from mid-May through early August (A. Mitchell, pers. comm.). Dromochorus welderensis occurs in the Gulf Prairie ecoregion of the Coastal Plain physiographic province of Texas. This Dromochorus is the least associated with tree cover, and D. welderensis is consistently found in tall grasses in upland prairie habitat. Beetles are often found near heavy clayloam or clay banks and hills. This shade-loving species can be observed in early cool mornings or early evenings, and will try to avoid more open areas on hot, clear days. Even when active, adults are particularly reclusive and tend to stay hidden in tall grasses. They are more reluctant to forage in wide open loam areas, in contrast to D. pruininus and D. velutinigrens. Beetles can also be found on thick grass mats as they forage, mate or disperse. When disturbed, they use these dead grass mats as cover., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 275-276, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237

DROMOCHORUS KNISLEYI DURAN, HERRMANN, ROMAN & EGAN SP.NOV. (FIGS 7C, 8A, 9C) Common name Juniper grove tiger beetle. Type locality Vi c i n i t y o f Pe d e r n a l e s Fa l l s, Te x a s. H o l o t y p e (USNM): 1 ♂, USA: Texas: Blanco Co. / Vicinity of Pedernales / 19-VI-2013 /leg D. Duran. Paratypes: 14 ♂♂, 19 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./19-VI-2013/leg D. Duran. 6 ♂♂, 7 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./08-VI-2015/leg S.J Roman. 5 ♂♂, 3 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./01-VI-2014/leg D. Brzoska. 2 ♂, USA: Texas: Blanco Co./Park E. of Pedernales Falls S.P./20-VI-2014/ leg D. Duran. 2 ♀, USA: TEXAS: Bandera Co./3 miles W of Pipe Creek/22-VI-2013/leg D. Duran. Distribution This species is only known from the Edwards Plateau of central Texas, locally known as the Texas Hill Country. It comes in close geographic proximity to D. belfragei and D. minimus, where all three species distributions converge at the edge of the Balcones Escarpment in Bexar County. Analyses of mtDNA data indicate that hybridization occurs in this contact zone, and apparent hybrid D. knisleyi x belfragei individuals have been found in this area. Diagnosis Dromochorus knisleyi is most easily confused with the sister taxa D. belfragei. For differential diagnosis, see D. belfragei species account. Description Medium-sized Dromochorus. Body length 10.9��� 14.4 mm, mean ♀ 13.0 mm, mean ♂ 12.2 mm. Head slightly wider than pronotum. Head predominantly charcoal black with blue reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black or bright polished metallic violet to blue, with shallow longitudinal striae gradually ending at border of vertex. Clypeus bronze with green to blue reflections throughout. Male labrum tridentate with 6���8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; the pale central area of the labrum may exist as a small spot, up to one-third of the total labrum surface; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently dark-brown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5���11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.4���3.2 mm in length, mean ♀ 2.9 mm, mean ♂ 2.8 mm; width 2.5���3.2 mm, mean ♀ 3.0 mm, mean ♂ 2.9 mm. Pronotum charcoal black, with green to blue or violet reflections, especially along lateral margins, slightly wider than long, widest near anterior margin, width to length ratio 0.9 to 1.1, setae sparse to regularly spaced, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with weak to strong iridescent blue to violet reflections, glabrous. Elytra elongate, 6.8���8.7 mm length, mean ♀ 7.9 mm, mean ♂ 7.7 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral dorsal surface convex; elytral texture dull, with regular small pits present throughout disk, elytral coloration charcoal black, often with blue reflections near humeral region; elytral maculations absent; two dark oblique infuscations present; subsutural foveae prominent, typically with metallic blue, green, or gold reflections. Legs: Pro-, meso-, and metacoxae dark testaceous to black with iridescent blue to violet and cupreous reflections, sparse setae on pro- and mesocoxae, fewer on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyishwhite setal pad. Abdomen: Venter mostly black with occasional metallic green to violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2���6 have sparse short brown erect setae present throughout, but often abraded. Etymology Named for Dr C. Barry Knisley, one of the leading authorities on North American tiger beetle conservation and ecology. D.P. Duran and R.A. Gwiazdowski are greatly indebted to Barry for his mentorship and friendship. Ecology/natural history Dromochorus knisleyi adults have been found from mid-May to late June, but it is likely that they could be active outside of this window. Dromochorus knisleyi is found in upland juniperoak woodlands in the Edwards Plateau, and does not appear to be strongly associated with riparian areas. The preferred habitat is late succession stands of juniper and, as such, it can be difficult for a collector to easily walk through these areas. Adult beetles are active throughout the day and are present in semiopen grassy areas under the cover of juniper trees. The first author observed dozens of beetles over a span of two days, and all adult activity was restricted to these forested areas. Beetles foraged and mated exclusively near or under juniper boughs. Moreover, even during cloudy periods and late in the afternoon, none were observed moving into more open grassy areas outside of the juniper stands. Moreover, beetles were not present in woodlands dominated by oaks. In mixed juniper���oak woodlands, beetles were found exclusively near junipers. This species may be the most ecologically specialized of all Dromochorus. More observations are needed for this rarely collected species. Many aspects of the biology are currently unknown., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 271-272, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237

DROMOCHORUS PILATEI GU��RIN- M��NEVILLE, 1845 (FIG. 9H) Common name Cajun tiger beetle. Type locality ���Velasco, Texas��� (translation). Holotype probably in MHNP, Paris (Bousquet, 2012). Synonymy Cicindela maga LeConte, 1875: 161. Type locatity ���near Lake Ponchartrain, Louisiana���. Syntypes (2) in MCZN. Synonymy established by Sall�� (1877). Taxonomic history This is the type species for the genus Dromochorus, as described by Gu��rin-M��neville (1845). Two remarkably green specimens were collected by LeConte and described as Cicindela maga (1875) from the vicinity of Lake Ponchartrain, LA. Distribution Dromochorus pilatei is known from south-east Texas in the vicinity of the Brazos River east to the Mississippi River in Louisiana, north to Natchitoches, LA. The Lake Ponchartrain record is uncertain. This flightless beetle has otherwise never been found east of the Mississippi River except for LeConte���s specimens. There have been multiple attempts to find the beetle in this area (Graves & Pearson, 1973; D.P. Duran, pers. obs.), but these have been unsuccessful. We regard this record as a potential error until further verification. Diagnosis This species cannot be confused with any other Dromochorus. The distinctive body form, elytral coloration with bronze and green reflections, prominent green subsutural foveae and complex surface texturing are diagnostic. Some individuals have a strong green-bronze sheen over all surfaces, and this trait appears to be more prevalent towards the eastern part of the species range. Description Small- to medium-sized Dromochorus. Body length 10.5���14.7 mm, mean ♀ 13.3 mm, mean ♂ 12.4 mm. Head slightly wider than pronotum. Head predominantly brown with cupreous to brassy reflections, green to blue to violet reflections mostly concentrated near the anterior margin and edges of the supraorbital region. In some specimens, bright green to green-blue reflections present throughout. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in males, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae metallic blue to violet, with shallow, longitudinal striae gradually ending at border of vertex. Clypeus bronze with green to blue reflections throughout; female clypeus more extensively greenblue to blue-violet. Male labrum tridentate with 6���8 setae, entirely pale ochre-testaceous, with a thin darkbrown to black border; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. Maxillary palpi pale yellow-ochre; apical segment dark-brown to black, often with metallic purple and green reflections. Labial palpi coloured similarly to maxillary palpi. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short, white setae, antennomeres 5���11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 1.8���3.2 mm in length, mean ♀ 2.8 mm, mean ♂ 2.6 mm; width 2.3���3.4 mm, mean ♀ 2.9 mm, mean ♂ 2.7 mm. Pronotum brown with cupreous, brassy or violet reflections; some specimens with green to green-blue reflections throughout, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.2, setae sparse to regularly spaced, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with weak to strong iridescent violet reflections, glabrous. Elytra elongate, dorsal surface convex, 6.4���9.0 mm length, mean ♀ 8.2 mm, mean ♂ 7.6 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral surface dull with complex texturing and infuscations, with regular small pits present throughout disk, as well as larger foveae. Bright green or blue reflections generally present in most to all pits and foveae. Legs: Pro-, meso- and metacoxae dark-brown to black, may have iridescent blue reflections, scattered setae on pro- and mesocoxae, fewer on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora dark-brown black with metallic violet and bronze reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter mostly dark-brown to black with faint violet reflections. Erect brown setae present on ventrite 1. Ventrites 2���6 have sparse short, brown, erect setae present throughout, but often abraded. Ecology/natural history Adults appear to be active from mid-May to mid-July. Dromochorus pilatei can be found in significant numbers during peak adult activity (early to mid-June), along shaded, dark soil trails in riparian zones or near the banks of bayous, lakes and salt marshes. Of the Dromochorus, pilatei has the strongest affinity for heavily forested areas. The species is tightly associated with blackish, rich soils with high humus content, which are produced in the forest via decaying vegetation. In contrast, D. pruininus and D. belfragei are associated with disturbed clay deposits that contain less organic matter (red iron oxidized clays or black clay loam). In our observations, D. pilatei apparently avoids the lighter coloured soils that can also be present in its habitat. This species can be found foraging/mating along man-made trails, disturbances or semi-open vegetated areas of applicable forest. Beetles appear to be concentrated on the edges of trails, sometimes with moderate to thick vegetation. Dromochorus pilatei is the only member of the genus that has been collected at lights at night (J. Back, pers. comm.). However, it was collected in small numbers, and it is likely that its presence was due to a high density of prey in the area, created by the lights. Traditionally thought to be crepuscular and perhaps nocturnal (Graves & Pearson, 1973) in this habit, we now know this species is active throughout the day in well-shaded areas., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 278-279, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237, {"references":["Bousquet Y. 2012. Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico. ZooKeys 245: 1 - 1630.","LeConte JL. 1875. Notes on Cicindelidae of the United States. Transactions of the American Entomological Society 5: 157 - 162.","Salle A. 1877. Note sur le genre Dromochorus. Annales de la Societe Entomologique de France (Bulletin des Seances) Ser 5 7: 7 - 8.","Guerin-Meneville FE. 1845. Sur le Dromochorus, nouveau genre de Cicindeletes. Annales de la Societe Entomologique de France (Bulletin Entomologique) Ser 2 7: 1 - 4.","Graves RC, Pearson DL. 1973. The tiger beetles of Arkansas, Louisiana, and Mississippi (Coleoptera: Cicindelidae). Transactions of the American Entomological Society 99: 157 - 203."]}

GENUS DROMOCHORUS GUÉRIN- MÉNEVILLE, 1845 Type species Dromochorus pilatei Guérin-Méneville, 1845. By monotypy. Taxonomic history Dromochorus was described by Guérin-Méneville (1845) as a new genus, believed to be most closely related to ‘ Apteroessa and especially Dromica ’. This perceived relatedness was due to the remarkable similarity in gestalt, although several morphological differences were identified, especially with respect to the mouthparts, such as the maxillary and labial palpi, and labrum. Subsequent authors have not shared his view about the systematic placement of the genus within the larger tiger beetle clade. In his revision of the Cicindelinae, Horn (1908) placed Dromochorus in the subtribe Cicindelina, and Dromica in a separate subtribe, Prothymina (Horn 1910). More recent molecular phylogenies have supported Horn’s (1908) treatment, finding Dromochorus to be nested within a clade of other Nearctic Cicindelina (Vogler & Welsh, 1997; Barraclough & Vogler, 2002; Pons et al., 2004; Vogler et al., 2005), and Dromica to be more related to Prothyma (Galián, Hogan & Vogler, 2002). Any similarities between Dromochorus and Dromica must, therefore, be the result of convergent evolution owing to their similar habitats and microhabitats. The Dromochorus species have sometimes been treated as a distinct genus (Guérin-Méneville, 1845; Sallé, 1877; Fleutiaux, 1892; Casey, 1897; Rivalier, 1954, 1963, 1971; Johnson, 1991; Wiesner, 1992; Erwin & Pearson, 2008; Bousquet, 2012; Pearson et al., 2015); or viewed to be a synonym of Cicindela (e.g. LeConte, 1875; Leng, 1902; Harris, 1911; Horn, 1915; Harris & Leng, 1916; Cazier, 1954; Arnett, 1963; Willis, 1968; Bousquet & Larochelle, 1993; Arnett & Thomas, 2000), with some North American workers recognizing the taxon as a subgenus (Boyd, 1982; Freitag, 1999). Despite the variety of treatments, justifications for different taxonomic ranks were almost never given. Recent phylogenies have given support to many of the named clades within the Cicindela sensu lato (Pons et al., 2004; Vogler et al., 2005; Gough et al., in review), and these generally correspond to groups recognized in Rivalier’s revision (1954, 1963, 1971), including the Dromochorus. Based on the aforementioned research, and our own mtDNA tree (Fig. 2), Dromochorus was recovered as a well-supported monophyletic lineage, and sister to another clade of Nearctic tiger beetles. As such, we recognize the group as a distinct genus. Although Dromochorus may appear nested within Cylindera in our tree, this latter group is polyphyletic (Gough et al., 2018), as previously believed by many workers, and the sister clade to Dromochorus is being named as a new genus. Diagnosis Dromochorus are separable from all other North American tiger beetle genera by the following combination of characters, present in the adult stage. Beetles are flightless and lack flight wings, but the elytra are not fused. The elytra are oval shaped, and completely lacking pale maculations. The dorsum is dark, usually black or brown, but may also have a frosted blue, violet, grey or green sheen. The legs and tarsi are clothed in decumbent setae. In subsequent species descriptions, the most salient or diagnostic characters are indicated in bold. Rarely are any of these characters diagnostic by themselves, but the combination of these characters may be. Distribution Dromochorus are geographically restricted to the south-central United States and adjacent Mexico (Fig. 3). Texas is the centre of diversity, and all eight species are found there, at least in part of their ranges. Records from Mexico are few and imprecise. Historically, D. belfragei was the only species recorded from Mexico, although it would appear as if these populations belong to D. chaparralensis sp. nov., described in this treatment. In general, Dromochorus are found at low elevations; apparently, they do not occur in montane environments, such as the Ouachita Mountains of Arkansas and Oklahoma, which border a section of the eastern range of D. pruininus. Most Dromochorus are found below 500 m, with only the westernmost populations of D. belfragei occurring at elevations up to 1000 m. Ecology/natural history Dromochorus appear to have a 2-year life cycle, based on observations of D. pruininus (Herrmann & Duran, unpublished). Adult beetles are terrestrial predators of small invertebrates, and are generally believed to be crepuscular or active in the late afternoon, but we have found them at all hours of the day in more shaded microhabitats or when significant cloud cover is present. Dromochorus are extremely fast runners and may evade capture by darting into dense grasses, or in some species, hiding in cracks in the earth, especially D. belfragei and D. pruininus. Larvae are poorly known, and were only recently described (Spomer, Nabity & Brust, 2008). There are remarkably few specimens of these beetles in major museum collections, even though several species occur near major cities and universities in Texas and Oklahoma. Many species records are based on one or a few specimens, and as such, Dromochorus were reported to be rare, or at low densities. This is likely a consequence of their atypical natural history compared to other diurnal North American tiger beetles, and the fact that their habitats are not as commonly visited by collectors. We have found that Dromochorus can be remarkably abundant in the appropriate habitat during the ideal time of year, rivalling or exceeding densities observed in some common riparian tiger beetle species. These observations are unlikely to represent unusual population explosions, as we have witnessed similarly large numbers of beetles every year in areas that we visited each year between 2012 and 2016. KEY TO THE GENUS DROMOCHORUS 1a. Labial palps at least partly yellow to dark amber, with contrasting darker apical segment (Fig. 7A, B)......................................................................................................................................................2 1b. Labial palps consistently dark brown to black throughout (Fig. 7C)..........................................................4 2a. Elytral surface smooth and finely frosted in texture, without any pitting or subsutural foveae. Dorsum with strong blue to violet reflections throughout. Labial palps yellow, with contrasting darker apical seg- ment. Kansas and western Missouri, south to Central Texas.................................................. D. pruininus 2b. Elytra surface dull, textured with fine to deep pitting throughout. Distinct shallow pits running along elytra suture (subsutural foveae, Fig. 8A) may be present..........................................................................3 3a. Dorsum dark brown with prominent shallow subsutural foveae and irregular pitting throughout ely- tral surface. Foveae and smaller pits with metallic green reflections. Irregular green marbling may be present on elytra, head and pronotum. Labial palps yellow, with contrasting darker apical segment. Louisiana to East Texas.................................................................................................................... D. pilatei 3b. Dorsum black with shallow to deep pitting on elytral surface. Some metallic blue reflections may be pre- sent, especially on the supraorbital region of the head and humeral area of the elytra. Subsutural foveae may be present. Labial palps yellow to dark amber, with apical segment darkest. Oklahoma and Texas panhandle, south to East Central Texas...................................................................................... D. belfragei 4a. Elytral surface rough, with shallow to deep pitting. Subsutural foveae present (Fig. 8A), often with metallic green or blue reflections in pits. ‘Hill Country’ region of Central Texas........... D. knisleyi sp. nov. 4b. Elytral surface smooth, often with velvety or frosted texture. No subsutural foveae present (Fig. 8B)..........................................................................................................................................................5 5a. Pronotum glabrous or with few scattered long thin erect setae irregularly placed, rarely concentrated along margins.................................................................................................................................................6 5b. Pronotum with sparse to regular white decumbent setae, mostly in lateral third....................................7 6a. Dorsum finely velvety black, with strong violet, blue or green reflections, especially along margins. Male labrum entirely dark or very nearly so. Body form gracile. South Texas, coastal areas south of Corpus Christi and inland to vicinity of Dimmit County................................................................. D. velutinigrens 6b. Elytra dark ash-grey, sometimes with frosty blue reflections. South Texas in mesquite chaparral forest. Known only from Bexar, Frio and Atascosa Counties.................................................... D. minimus sp. nov. 7a. Elytra dull black, may have bluish reflections especially near margins. South Texas to Mexico, in mes- quite chaparral. Known from Dimmit, LaSalle, and Webb Counties in Texas, and the state of Tamaulipas, Mexico..................................................................................................................... D. chaparralensis sp. nov. 7b. Elytra finely velvety black, may have a faint dark blue sheen. Found in coastal prairie habitat near Gulf of Mexico...................................................................................................................... D. welderensis sp. nov.

DROMOCHORUS CHAPARRALENSIS DURAN, ROMAN, HERRMANN & EGAN SP . NOV . (FIG. 9E) Common name Chaparral tiger beetle. Type locality Carrizo Springs, TX. Holotype (deposited in NMNH): 1 ♀, Carrizo Spgs/Tex. V-27���32 // E. G. Lindsley Collector // M.A. Cazier/Collection. Paratypes: 3 ♂♂, 5 ♀♀, Carrizo Spgs./Tex VI-12���32 // E.G. Lindsley Collector // A. Nicolay collection 1950. 1 ♀, Mexico, Tamaulipas/Nuevo Laredo/20-VI-2010/leg J. Stamatov. 1 ♀, USA: Texas: LaSalle Co./Chaparral W.M.A./19-VI- 2013/E. San Gregario. Fig. 9E, Map Fig. 3 Distribution Inland South Texas (currently known from Dimmit, LaSalle, and Webb Counties) and Tamaulipas, Mexico. This species has not been well sampled and is likely present in adjacent areas within the Gulf Coastal Plain, especially in northern Mexico. It can occur with D. velutinigrens in places where sandy soils mix with the dominant heavier red clays. Diagnosis Dromochorus chaparralensis is a robust, dark beetle, and most specimens are dull black with little colour on dorsal or ventral surfaces. Some specimens may have bluish reflections along the margins. It is restricted to mesquite chaparral in South Texas. This species is most likely to be confused with D. minimus or welderensis. Dromochorus mimimus is usually smaller (Fig. 6; Table 2), and possesses a prominent frosted ashy grey, beige, or blue sheen on the dorsal surface; the pronotum has sparse thin erect setae. Dromochorus welderensis is black with a faint dark blue sheen dorsally, in many individuals. The habitat of D. welderensis is Gulf Coast Prairie grasslands, as opposed to the more inland and forested mesquitechaparral habitat of D. chaparralensis. Description Medium- to large-sized Dromochorus. Body length 12.5���14.5 mm, mean ♀ 13.7 mm, mean ♂ 13.0 mm. Head slightly wider than pronotum. Head black with metallic blue to green reflections mostly limited to the lateral ridge of the supraorbital region. Fine to marked rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black often with weak metallic green to violet reflections, with shallow, longitudinal striae, gradually ending at border of vertex. Clypeus shining black, apparently lacking coloured reflections. Male labrum tridentate with 6���8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-quarter of the total labrum surface; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently dark brown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5���11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.6���3.4 mm in length, mean ♀ 3.1 mm, mean ♂ 3.1 mm; width 2.9���3.5 mm, mean ♀ 3.4 mm, mean ♂ 3.1 mm. Pronotum dull black, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.1, setae sparse, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black, lacking prominent metallic coloured reflections, glabrous. Elytra elongate, dorsal surface convex, 7.2���8.6 mm length, mean ♀ 8.2 mm, mean ♂ 7.7 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration black, may have weak blue reflections along lateral margins; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae black, without or with minimal metallic reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet reflections, densely clothed in decumbent white setae; tibiae brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyishwhite setal pad. Abdomen: Venter mostly black with metallic olive green and violet reflections. Decumbent setae present on ventrite 1. Ventrites 2���6 have sparse short brown erect setae present throughout, but often abraded. Etymology Named for the dominant mesquite-chaparral plant community found throughout the species range. Also, this species has been collected at Chaparral Wildlife Management Area, in Cotulla, Texas. Ecology/natural history Very little is known about this species, as most museum specimens are older, and nothing has been recorded about its ecology. The authors of this study have not observed D. chaparralensis in situ., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 274-275, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237

DROMOCHORUS BELFRAGEI SALL��, 1877 (FIGS 7B, 9B) Common name Loamy-ground tiger beetle. Type locality ���Texas���Dallas, Wasco (presumed to be a misspelling of Waco), etc., on the banks of the Trinity River��� (English translation). Syntypes unknown, probably in MHNP (Bousquet 2012). Synonymy Dromochorus bellefragei Heyne, 1893 (unjustified emendation). Dromochorus sericeus Casey, 1897: 294. Type locality ���Texas���. Two syntypes in USNM, Washington DC (synonymy established by Leng, 1902). Taxonomic history Dromochorus belfragei has been a catch-all taxon, and many populations of Dromochorus were lumped under this name in the literature or in museum collections prior to this revision. Dromochorus pilatei and D. velutinigrens are the only nominal taxa that have never been considered conspecific with D. belfragei. Based on morphology, ecology, biogeography, mtDNA genealogy and multilocus genetic data, D. belfragei (sensu stricto) is circumscribed as a separate species from three other distinct species, D. pruininus, D. knisleyi sp. nov. and D. minimus sp. nov. Occasional hybridization with D. knisleyi has been observed where their ranges come in contact, and this is further supported by evidence of mtDNA introgression. Distribution Dromochorus belfragei is known to occur from the north-western panhandle of Texas and southern Oklahoma to south-eastern Texas. Previous literature described a more extensive range from south-eastern Colorado (Michels et al., 2008) to Tamaulipas, Mexico (Cazier, 1954). The former record now appears to be an error (Michels, pers. comm., 2015), and the latter would appear to belong to D. chaparralensis sp. nov. One specimen from the AMNH was labelled ���St. George, Utah���, although this locality would not appear plausible. The range of D. belfragei comes close to several other species. In particular, D. belfragei is nearly sympatric with D. pruininus in several places (Fig. 3; see D. pruininus account). In eastern and southern Bexar County, TX, this species is found sympatrically with D. knisleyi, and possibly D. minimus. Diagnosis This species can be distinguished from all other similar Dromochorus by the presence of finely to coarsely pitted black elytra that are not frosted in texture, in conjunction with maxillary palpi that have the apical segment darkest, with dark yellow-ochre to dark amber coloration in other segments (Fig. 8). Most populations of D. belfragei possess at least small subsutural foveae, but these lack metallic reflections. Southern and eastern populations may have subtle dark infuscations on the elytra. Dromochorus belfragei is most likely to be confused with D. pruininus, knisleyi or welderensis. It can be separated from these taxa by the following: 1. Dromochorus pruininus has a frosted blue sheen on the elytra and the entire dorsal surface, and it lacks any pits or subsutural foveae. The maxillary palpi in D. pruininus are pale yellow-ochre with a dark apical segment, whereas D. belfragei has darker yellow-red to red testaceous palpi with a dark apical segment. 2. Dromochorus knisleyi is similar to D. belfragei, but has more prominent subsutural foveae, often with bright metallic blue, green or gold reflections. In D. knisleyi, the maxillary palpi are always dark in all segments, whereas in D. belfragei, the apical segment is darker than the preceding. Infuscations are always present in D. knisleyi and are usually absent in D. belfragei. Ecologically, D. knisleyi is found in upland juniper woodland in the Edwards Plateau, and D. belfragei is found outside of this region, in clay soils associated with larger riparian systems. The two species appear to hybridize along a narrow contact zone at the edge of the Balcones Escarpment in south-eastern and south-central Texas. Both species and their hybrids may be found in this area. The existence of a hybrid zone was further supported by mtDNA data (Fig. 2). 3. Dromochorus welderensis has a smooth velvety black elytral surface (may have metallic blue-violet reflections), and never has pitting or subsutural foveae. The maxillary palpi are always dark in all segments, whereas in D. belfragei, the apical segment is darker than the preceding. Description Medium- to large-sized Dromochorus. Body length 11.6���15.2 mm, mean ♀ 14.5 mm, mean ♂ 13.2 mm. Head slightly wider than pronotum. Head predominantly black with blue reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae bright polished metallic violet to blue, with shallow, longitudinal striae gradually ending at border of vertex. Clypeus mostly black, with patches of metallic blue or violet reflections; clypeus may be nearly entirely blue to violet in females. Male labrum tridentate with 6���8 setae, central area pale ochre-testaceous, with a thin dark-brown to black bor- der posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, in others the pale area may cover more than two-thirds of the total labrum surface; female labrum tridentate with 6���8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. Maxillary and labial palpi with apical segment darker than other segments; basal to penultimate segments yellow-ochre to dark red-amber, often with metallic purple and green reflections. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5���11 dulltextured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.5���3.4 mm in length, mean ♀ 3.1 mm, mean ♂ 2.8 mm; width 2.8���3.6 mm, mean ♀ 3.4 mm, mean ♂ 3.1 mm. Pronotum black, with some dark-blue reflections, especially in sulci, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.2, setae sparse to regularly spaced, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with weak to strong iridescent blue reflections, glabrous. Elytra convex, elongate, 7.1���9.7 mm length, mean ♀ 8.8 mm, mean ♂ 8.2 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with regular small pits present throughout disk, elytral coloration black, often with blue reflections near humeral region; elytral maculations absent; infuscations rarely present; subsutural foveae, when present, only slightly more prominent than other pits on the elytral disk; subsutural foveae lacking bright metallic reflections, except rarely in basal area. Legs: Pro-, meso- and metacoxae dark testaceous to black with iridescent blue to violet and cupreous reflections, with numerous setae; pro- and meso- trochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous, dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter mostly black with occasional metallic violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2���6 have sparse, short, brown erect setae present throughout, but often abraded. Ecology/natural history Dromochorus belfragei adults are active between mid- May and early July in most of its range (e.g. central to north TX) and late June to late July in northern part of its range (e.g. OK to panhandle of TX). Dromochorus belfragei can be found in natural and managed forested and agricultural areas (e.g. pecan groves) that have semi-open shaded areas or trails beneath the canopy. Soils in these areas can be dark, red to black in color, clay to clay-loam, cracked and sometimes moist in low areas that are heavily trampled by cattle. Adult beetles tend to avoid the lighter coloured sandy areas that are exposed to full sunlight on the forest edges. Despite the common name, this species appears more closely associated with soils that possess high clay content, more so than loam. The preferred habitat of D. belfragei is generally associated with larger riparian systems, although the beetles are not typically found near the water���s edge and we have found them over 3 km from water. They appear to have a wider ecological niche than most Dromochorus; their habitat and geographic distribution encompass a larger number of ecoregions than other species (Fig. 3; Table 1). Dromochorus belfragei and D. pruininus have similar but non-overlapping ranges, and they may be separated by ecological barriers, at least in some areas. In the DFW area of North Texas, the species has been reported a few kilometers from D. pruininus (Pearson et al., 2006), but the two appear to be separated by a narrow extension of the EPA Level IV Eastern Cross Timbers ecoregion, which is not suitable for either species. The forested undergrowth of this area is extremely dense in many areas, and may not possess the necessary surface soil conditions for either species to persist. Dromochorus belfragei may be more susceptible to urbanization than D. pruininus, as none have been collected in the DFW area (Tarrant Co.) since the 1970s, whereas D. pruininus appear much more tolerant to these disturbances and may be abundant in semi-open grassy areas where trails have been established in riparian parks (Dallas, Collin Co.). Thought to be crepuscular, this species is active throughout the day in shady areas or when overcast. Similar in behaviour to D. pruininus and D. pilatei, this species has been observed using soil cracks for escape, especially during dry conditions when virtisolic cracks are pronounced. Like other Dromochorus, D. belfragei frequently moves to vegetated cover to escape when pursued., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 269-271, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237, {"references":["Salle A. 1877. Note sur le genre Dromochorus. Annales de la Societe Entomologique de France (Bulletin des Seances) Ser 5 7: 7 - 8.","Bousquet Y. 2012. Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico. ZooKeys 245: 1 - 1630.","Casey TL. 1897. Coleopterological notices VII. Annals of the New York Academy of Sciences 9: 285 - 684.","Leng CW. 1902. Revision of the Cicindelidae of boreal America. Transactions of the American Entomological Society 28: 93 - 186.","Michels GJ Jr, Newton JL, Lindon HL, Brazille JA. 2008. Invertebrate distribution and diversity assessment at the U. S. Army Pinon Canyon Maneuver Site. Bushland, TX: a report to the US Army and US Fish and Wildlife Service. Texas AgriLife Research.","Cazier MA. 1954. A review of the Mexican tiger beetles of the genus Cicindela (Coleoptera, Cicindelidae). Bulletin of the American Museum of Natural History 103: 231 - 309.","Pearson DL, Knisley CB, Kazilek CJ. 2006. A field guide to tiger beetles of the United States and Canada: identification, natural history, and distribution of the Cicindelidae. New York: Oxford University Press."]}

GENUS DROMOCHORUS GU��RIN- M��NEVILLE, 1845 Type species Dromochorus pilatei Gu��rin-M��neville, 1845. By monotypy. Taxonomic history Dromochorus was described by Gu��rin-M��neville (1845) as a new genus, believed to be most closely related to ��� Apteroessa and especially Dromica ���. This perceived relatedness was due to the remarkable similarity in gestalt, although several morphological differences were identified, especially with respect to the mouthparts, such as the maxillary and labial palpi, and labrum. Subsequent authors have not shared his view about the systematic placement of the genus within the larger tiger beetle clade. In his revision of the Cicindelinae, Horn (1908) placed Dromochorus in the subtribe Cicindelina, and Dromica in a separate subtribe, Prothymina (Horn 1910). More recent molecular phylogenies have supported Horn���s (1908) treatment, finding Dromochorus to be nested within a clade of other Nearctic Cicindelina (Vogler & Welsh, 1997; Barraclough & Vogler, 2002; Pons et al., 2004; Vogler et al., 2005), and Dromica to be more related to Prothyma (Gali��n, Hogan & Vogler, 2002). Any similarities between Dromochorus and Dromica must, therefore, be the result of convergent evolution owing to their similar habitats and microhabitats. The Dromochorus species have sometimes been treated as a distinct genus (Gu��rin-M��neville, 1845; Sall��, 1877; Fleutiaux, 1892; Casey, 1897; Rivalier, 1954, 1963, 1971; Johnson, 1991; Wiesner, 1992; Erwin & Pearson, 2008; Bousquet, 2012; Pearson et al., 2015); or viewed to be a synonym of Cicindela (e.g. LeConte, 1875; Leng, 1902; Harris, 1911; Horn, 1915; Harris & Leng, 1916; Cazier, 1954; Arnett, 1963; Willis, 1968; Bousquet & Larochelle, 1993; Arnett & Thomas, 2000), with some North American workers recognizing the taxon as a subgenus (Boyd, 1982; Freitag, 1999). Despite the variety of treatments, justifications for different taxonomic ranks were almost never given. Recent phylogenies have given support to many of the named clades within the Cicindela sensu lato (Pons et al., 2004; Vogler et al., 2005; Gough et al., in review), and these generally correspond to groups recognized in Rivalier���s revision (1954, 1963, 1971), including the Dromochorus. Based on the aforementioned research, and our own mtDNA tree (Fig. 2), Dromochorus was recovered as a well-supported monophyletic lineage, and sister to another clade of Nearctic tiger beetles. As such, we recognize the group as a distinct genus. Although Dromochorus may appear nested within Cylindera in our tree, this latter group is polyphyletic (Gough et al., 2018), as previously believed by many workers, and the sister clade to Dromochorus is being named as a new genus. Diagnosis Dromochorus are separable from all other North American tiger beetle genera by the following combination of characters, present in the adult stage. Beetles are flightless and lack flight wings, but the elytra are not fused. The elytra are oval shaped, and completely lacking pale maculations. The dorsum is dark, usually black or brown, but may also have a frosted blue, violet, grey or green sheen. The legs and tarsi are clothed in decumbent setae. In subsequent species descriptions, the most salient or diagnostic characters are indicated in bold. Rarely are any of these characters diagnostic by themselves, but the combination of these characters may be. Distribution Dromochorus are geographically restricted to the south-central United States and adjacent Mexico (Fig. 3). Texas is the centre of diversity, and all eight species are found there, at least in part of their ranges. Records from Mexico are few and imprecise. Historically, D. belfragei was the only species recorded from Mexico, although it would appear as if these populations belong to D. chaparralensis sp. nov., described in this treatment. In general, Dromochorus are found at low elevations; apparently, they do not occur in montane environments, such as the Ouachita Mountains of Arkansas and Oklahoma, which border a section of the eastern range of D. pruininus. Most Dromochorus are found below 500 m, with only the westernmost populations of D. belfragei occurring at elevations up to 1000 m. Ecology/natural history Dromochorus appear to have a 2-year life cycle, based on observations of D. pruininus (Herrmann & Duran, unpublished). Adult beetles are terrestrial predators of small invertebrates, and are generally believed to be crepuscular or active in the late afternoon, but we have found them at all hours of the day in more shaded microhabitats or when significant cloud cover is present. Dromochorus are extremely fast runners and may evade capture by darting into dense grasses, or in some species, hiding in cracks in the earth, especially D. belfragei and D. pruininus. Larvae are poorly known, and were only recently described (Spomer, Nabity & Brust, 2008). There are remarkably few specimens of these beetles in major museum collections, even though several species occur near major cities and universities in Texas and Oklahoma. Many species records are based on one or a few specimens, and as such, Dromochorus were reported to be rare, or at low densities. This is likely a consequence of their atypical natural history compared to other diurnal North American tiger beetles, and the fact that their habitats are not as commonly visited by collectors. We have found that Dromochorus can be remarkably abundant in the appropriate habitat during the ideal time of year, rivalling or exceeding densities observed in some common riparian tiger beetle species. These observations are unlikely to represent unusual population explosions, as we have witnessed similarly large numbers of beetles every year in areas that we visited each year between 2012 and 2016. KEY TO THE GENUS DROMOCHORUS 1a. Labial palps at least partly yellow to dark amber, with contrasting darker apical segment (Fig. 7A, B)......................................................................................................................................................2 1b. Labial palps consistently dark brown to black throughout (Fig. 7C)..........................................................4 2a. Elytral surface smooth and finely frosted in texture, without any pitting or subsutural foveae. Dorsum with strong blue to violet reflections throughout. Labial palps yellow, with contrasting darker apical seg- ment. Kansas and western Missouri, south to Central Texas.................................................. D. pruininus 2b. Elytra surface dull, textured with fine to deep pitting throughout. Distinct shallow pits running along elytra suture (subsutural foveae, Fig. 8A) may be present..........................................................................3 3a. Dorsum dark brown with prominent shallow subsutural foveae and irregular pitting throughout ely- tral surface. Foveae and smaller pits with metallic green reflections. Irregular green marbling may be present on elytra, head and pronotum. Labial palps yellow, with contrasting darker apical segment. Louisiana to East Texas.................................................................................................................... D. pilatei 3b. Dorsum black with shallow to deep pitting on elytral surface. Some metallic blue reflections may be pre- sent, especially on the supraorbital region of the head and humeral area of the elytra. Subsutural foveae may be present. Labial palps yellow to dark amber, with apical segment darkest. Oklahoma and Texas panhandle, south to East Central Texas...................................................................................... D. belfragei 4a. Elytral surface rough, with shallow to deep pitting. Subsutural foveae present (Fig. 8A), often with metallic green or blue reflections in pits. ���Hill Country��� region of Central Texas........... D. knisleyi sp. nov. 4b. Elytral surface smooth, often with velvety or frosted texture. No subsutural foveae present (Fig. 8B)..........................................................................................................................................................5 5a. Pronotum glabrous or with few scattered long thin erect setae irregularly placed, rarely concentrated along margins.................................................................................................................................................6 5b. Pronotum with sparse to regular white decumbent setae, mostly in lateral third....................................7 6a. Dorsum finely velvety black, with strong violet, blue or green reflections, especially along margins. Male labrum entirely dark or very nearly so. Body form gracile. South Texas, coastal areas south of Corpus Christi and inland to vicinity of Dimmit County................................................................. D. velutinigrens 6b. Elytra dark ash-grey, sometimes with frosty blue reflections. South Texas in mesquite chaparral forest. Known only from Bexar, Frio and Atascosa Counties.................................................... D. minimus sp. nov. 7a. Elytra dull black, may have bluish reflections especially near margins. South Texas to Mexico, in mes- quite chaparral. Known from Dimmit, LaSalle, and Webb Counties in Texas, and the state of Tamaulipas, Mexico..................................................................................................................... D. chaparralensis sp. nov. 7b. Elytra finely velvety black, may have a faint dark blue sheen. Found in coastal prairie habitat near Gulf of Mexico...................................................................................................................... D. welderensis sp. nov., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 263-265, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237, {"references":["Guerin-Meneville FE. 1845. Sur le Dromochorus, nouveau genre de Cicindeletes. Annales de la Societe Entomologique de France (Bulletin Entomologique) Ser 2 7: 1 - 4.","Horn W. 1908. Cicindelinae. In: Wytsman P, ed. Genera insectorum. 82 a: 1 - 104.","Horn W. 1910. Cicindelinae. In: Wytsman P, ed. Genera insectorum. 82 b: 105 - 208, plates 1 - 15.","Vogler AP, Welsh A. 1997. Phylogeny of North American Cicindela tiger beetles inferred from multiple mitochondrial DNA sequences. Molecular phylogenetics and evolution 8: 225 - 235.","Barraclough TG, Vogler AP. 2002. Recent diversification rates in North American tiger beetles estimated from a dated mtDNA phylogenetic tree. Molecular Biology and Evolution 19: 1706 - 1716.","Pons J, Barraclough T, Theodorides K, Cardoso A, Vogler A. 2004. Using exon and intron sequences of the gene Mp 20 to resolve basal relationships in Cicindela (Coleoptera: Cicindelidae). Systematic Biology 53: 554 - 570.","Vogler AP, Cardoso A, Barraclough TG. 2005. Exploring rate variation among and within sites in a densely sampled tree: Species level phylogenetics of North American tiger beetles (genus Cicindela). Systematic Biology 54: 4 - 20.","Galian J, Hogan JE, Vogler AP. 2002. The origin of multiple sex chromosomes in tiger beetles. Molecular Biology and Evolution 19: 1792 - 1796.","Salle A. 1877. Note sur le genre Dromochorus. Annales de la Societe Entomologique de France (Bulletin des Seances) Ser 5 7: 7 - 8.","Fleutiaux E. 1892. Catalogue systematique des Cicindelidae, decrits depuis Linne. Vaillant-Carmanne: Imprimerie H.","Casey TL. 1897. Coleopterological notices VII. Annals of the New York Academy of Sciences 9: 285 - 684.","Rivalier E. 1954. Demembrement du genre Cicindela Linne. II. Faune americaine. Revue Francaise d'Entomologie 21: 249 - 268.","Rivalier E. 1963. Demembrement du genre Cicindela L. (fin). V. Faune australienne (et liste recapitulative des genres et sous-genres proposes pour la faune mondiale). Revue Francaise d'Entomologie 31: 127 - 164.","Rivalier E. 1971. Remarques sur la Tribu des Cicindelini (Col. Cicindelidae) et sa subdivision en sous-tribus. Nouvelle revue d'Entomologie 1: 135 - 143.","Johnson WN. 1991. A new species of Dromochorus from southern Texas (Coleoptera: Cicindelidae). Cicindela 23: 49 - 54.","Wiesner J. 1992. Verzeichnis der Sandlaufkafer der Welt (Coleoptera, Cicindelidae). Keltern: Erna Bauer.","Erwin TL, Pearson DL. 2008. A treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of life. Volume II (Carabidae- Nebriiformes 2 - Cicindelitae). Sofia-Moscow: Pensoft.","Bousquet Y. 2012. Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico. ZooKeys 245: 1 - 1630.","Pearson DL, Knisley CB, Duran DP, Kazilek CJ. 2015. A field guide to tiger beetles of the United States and Canada: identification, natural history, and distribution of the Cicindelinae. New York: Oxford University Press.","LeConte JL. 1875. Notes on Cicindelidae of the United States. Transactions of the American Entomological Society 5: 157 - 162.","Leng CW. 1902. Revision of the Cicindelidae of boreal America. Transactions of the American Entomological Society 28: 93 - 186.","Harris ED. 1911. List of North American Cicindelidae in the Harris Collection. Yonkers: Truan Press.","Horn W. 1915. Cicindelinae. In: Wytsman P, ed. Genera insectorum. 82 c: 209 - 486, plates 16 - 23.","Harris ED, Leng C. 1916. The Cicindelinae of North America as arranged by Dr. Walther Horn in Genera Insectorum. New York: The American Museum of Natural History Press.","Cazier MA. 1954. A review of the Mexican tiger beetles of the genus Cicindela (Coleoptera, Cicindelidae). Bulletin of the American Museum of Natural History 103: 231 - 309.","Arnett RH Jr. 1963. The beetles of the United States (a manual for identification). Washington DC: Catholic University of America Press.","Willis HL. 1968. Artificial key to the species of Cicindela o f N o r t h A m e r i c a n o r t h o f M e x i c o (C o l e o p t e r a: Cicindelidae). Journal of the Kansas Entomological Society 41: 303 - 317.","Bousquet Y, Larochelle A. 1993. Catalogue of the Geadephaga (Coleoptera: Trachypachidae, Rhysodidae, Carabidae including Cicindelini) of America north of Mexico. Memoirs of the Entomological Society of Canada 67: 1 - 197.","Arnett RH Jr, Thomas MC, eds. 2000. American beetles, Volume I: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. Boca Raton: CRC Press.","Boyd HP. 1982. Checklist of Cicindelidae: the tiger beetles. Marlton, NJ: Plexus.","Freitag R. 1999. Catalogue of the tiger beetles of Canada and the United States. Ottawa: NRC Research Press.","Gough HM, Duran DP, Kawahara AY, Toussiant EF. 2018. A comprehensive molecular phylogeny of tiger beetles (Coleoptera, Carabidae, Cicindelinae) challenges their current classification. Systematic Entomology (in press).","Spomer SM, Nabity PD, Brust ML. 2008. Larval description of Cicindela (Dromochorus) pruinina (Casey) (Coleoptera: Carabidae: Cicindelinae) with notes on habitat and adult behavior. The Coleopterist's Bulletin 62: 37 - 41."]}

DROMOCHORUS KNISLEYI DURAN, HERRMANN, ROMAN & EGAN SP.NOV. (FIGS 7C, 8A, 9C) Common name Juniper grove tiger beetle. Type locality Vi c i n i t y o f Pe d e r n a l e s Fa l l s, Te x a s. H o l o t y p e (USNM): 1 ♂, USA: Texas: Blanco Co. / Vicinity of Pedernales / 19-VI-2013 /leg D. Duran. Paratypes: 14 ♂♂, 19 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./19-VI-2013/leg D. Duran. 6 ♂♂, 7 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./08-VI-2015/leg S.J Roman. 5 ♂♂, 3 ♀♀, USA: Texas: Blanco Co./Vicinity of Pedernales Falls St. Pk./01-VI-2014/leg D. Brzoska. 2 ♂, USA: Texas: Blanco Co./Park E. of Pedernales Falls S.P./20-VI-2014/ leg D. Duran. 2 ♀, USA: TEXAS: Bandera Co./3 miles W of Pipe Creek/22-VI-2013/leg D. Duran. Distribution This species is only known from the Edwards Plateau of central Texas, locally known as the Texas Hill Country. It comes in close geographic proximity to D. belfragei and D. minimus, where all three species distributions converge at the edge of the Balcones Escarpment in Bexar County. Analyses of mtDNA data indicate that hybridization occurs in this contact zone, and apparent hybrid D. knisleyi x belfragei individuals have been found in this area. Diagnosis Dromochorus knisleyi is most easily confused with the sister taxa D. belfragei. For differential diagnosis, see D. belfragei species account. Description Medium-sized Dromochorus. Body length 10.9– 14.4 mm, mean ♀ 13.0 mm, mean ♂ 12.2 mm. Head slightly wider than pronotum. Head predominantly charcoal black with blue reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black or bright polished metallic violet to blue, with shallow longitudinal striae gradually ending at border of vertex. Clypeus bronze with green to blue reflections throughout. Male labrum tridentate with 6–8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; the pale central area of the labrum may exist as a small spot, up to one-third of the total labrum surface; female labrum tridentate with 6–8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently dark-brown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2–3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3–4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5–11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.4–3.2 mm in length, mean ♀ 2.9 mm, mean ♂ 2.8 mm; width 2.5–3.2 mm, mean ♀ 3.0 mm, mean ♂ 2.9 mm. Pronotum charcoal black, with green to blue or violet reflections, especially along lateral margins, slightly wider than long, widest near anterior margin, width to length ratio 0.9 to 1.1, setae sparse to regularly spaced, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with weak to strong iridescent blue to violet reflections, glabrous. Elytra elongate, 6.8–8.7 mm length, mean ♀ 7.9 mm, mean ♂ 7.7 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral dorsal surface convex; elytral texture dull, with regular small pits present throughout disk, elytral coloration charcoal black, often with blue reflections near humeral region; elytral maculations absent; two dark oblique infuscations present; subsutural foveae prominent, typically with metallic blue, green, or gold reflections. Legs: Pro-, meso-, and metacoxae dark testaceous to black with iridescent blue to violet and cupreous reflections, sparse setae on pro- and mesocoxae, fewer on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyishwhite setal pad. Abdomen: Venter mostly black with occasional metallic green to violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2–6 have sparse short brown erect setae present throughout, but often abraded. Etymology Named for Dr C. Barry Knisley, one of the leading authorities on North American tiger beetle conservation and ecology. D.P. Duran and R.A. Gwiazdowski are greatly indebted to Barry for his mentorship and friendship. Ecology/natural history Dromochorus knisleyi adults have been found from mid-May to late June, but it is likely that they could be active outside of this window. Dromochorus knisleyi is found in upland juniperoak woodlands in the Edwards Plateau, and does not appear to be strongly associated with riparian areas. The preferred habitat is late succession stands of juniper and, as such, it can be difficult for a collector to easily walk through these areas. Adult beetles are active throughout the day and are present in semiopen grassy areas under the cover of juniper trees. The first author observed dozens of beetles over a span of two days, and all adult activity was restricted to these forested areas. Beetles foraged and mated exclusively near or under juniper boughs. Moreover, even during cloudy periods and late in the afternoon, none were observed moving into more open grassy areas outside of the juniper stands. Moreover, beetles were not present in woodlands dominated by oaks. In mixed juniper–oak woodlands, beetles were found exclusively near junipers. This species may be the most ecologically specialized of all Dromochorus. More observations are needed for this rarely collected species. Many aspects of the biology are currently unknown.

DROMOCHORUS MINIMUS DURAN, ROMAN, HERRMANN & EGAN SP.NOV. (FIG. 9D) Common name Pygmy dromo tiger beetle. Type locality SE of Pleasanton, TX. Holotype (deposited in U S N M): 1 ♂, U S A: Te x a s: A t a s c o s a C o./S E o f Pleasanton / 19-VI-2014 /leg D. Duran. Paratypes: 3 ♂ ♂, 3 ♀ ♀, U S A: Te x a s: A t a s c o s a C o./S E o f Pleasanton/19-VI-2014/leg D. Duran. 3 ♂, 4 ♀♀, USA: Texas: Atascosa Co./SE. of Pleasanton/30-V- 2014 / leg D. Sunberg. 1 ♂, 1 ♀, USA: Texas: Atascosa Co./SE. of Pleasanton/02-VI-2015/leg S.J. Roman. 17 ♂♂, 7 ♀♀, USA: Texas: Atascosa Co./SE. of Pleasanton/29-VI-2014/leg D. Brzoska. 1 ♂, USA, Texas: Bexar Co./7 miles S. San Antonio/06-VI-2010/ leg G. Waldren. 1 ♂, USA: Texas: Bexar Co./Loop 1604 Hwy/12-VI-2016/leg J. Back. Distribution Central/south Texas, south of the Balcones Escarpment. Currently known only from Bexar, Atascosa and Frio Counties. Diagnosis Dromochorus minimus can be separated from all other species, by the presence of a frosted or ashy grey to beige dorsum, sometimes with blue reflections, in conjunction with labial palpi that are all dark (apical segment is not darker than other segments), and sparse erect setae on the pronotum, often irregularly placed throughout. This species is most likely to be confused with D. pruininus, chaparralensis or welderensis. Dromochorus pruininus is generally larger (Fig. 6; Table 2), has pale maxillary palps with a contrasting dark apical segment and is also separable by geographic range. Dromochorus chaparralensis is usually larger (Fig. 6; Table 2), and lacks any prominent frosted texturing on the dorsal surface. The pronotum of D. chaparralensis has setae more regularly arranged, mostly along lateral third. Dromochorus welderensis is usually larger (Fig. 6; Table 2), and its pronotum has decumbent white setae, mostly along lateral third. The habitat of D. welderensis is Gulf Coast Prairie. Description Small- to medium-sized Dromochorus. Body length 10.5���13.7 mm, mean ♀ 12.7 mm, mean ♂ 11.7 mm. Head slightly wider than pronotum. Head predominantly charcoal black with blue to green reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black with bright polished metallic green to violet reflections, with shallow longitudinal striae gradually ending at border of vertex. Clypeus shining black with blue to violet reflections throughout. Male labrum tridentate with 6���8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-third of the total labrum surface; female labrum tridentate with 6���8 setae, entirely dark brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently darkbrown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2���3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3���4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5���11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.4���3.1 mm in length, mean ♀ 2.8 mm, mean ♂ 2.7 mm; width 2.5���3.3 mm, mean ♀ 3.0 mm, mean ♂ 2.7 mm. Pronotum charcoal black, typically with frosty pale grey to brown, or blue to violet sheen, especially along lateral margins, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.1, thin erect setae sparse to irregularly spaced on pronotum; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with iridescent olive green to violet reflections, glabrous. Elytra elongate, dorsal surface convex, 6.4���8.0 mm length, mean ♀ 7.6 mm, mean ♂ 7.1 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration charcoal black, typically with grey, brown or blue-grey frosted texture along lateral margins, apex with blue or grey frosted texture; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae dark brown with iridescent violet and cupreous reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter black with metallic olive green and violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2���6 have sparse short brown erect setae present throughout, but often abraded. Etymology Dromochorus minimus is named for its smaller size. On average, this species is the smallest in the genus (Table 2). Ecology/natural history Little is known about this species��� natural history. Adults have been collected from mid-May until late June, but it is possible that the species may be active outside of this window. Dromochorus minimus occurs in mesquite-chaparral savannah in central/south Texas, just south of the Edwards Plateau, part of the larger Gulf Coastal Plains physiographic province. It has been found in open grassy areas interspersed between mesquite trees and clumps of Opuntia cactus. Adult beetles may be found venturing into the open spaces between clumps of grass, and will rapidly run into vegetation if pursued. Dromochorus minimus appears to be remarkably swift, even compared to other species of Dromochorus. In direct sunlight, live specimens appear beige-grey to smoky blue-grey., Published as part of Duran, Daniel P., Herrmann, David P., Roman, Stephen J., Gwiazdowski, Rodger A., Drummond, Jennifer A., Hood, Glen R. & Egan, Scott P., 2019, Cryptic diversity in the North American Dromochorus tiger beetles (Coleoptera: Carabidae: Cicindelinae): a congruence-based method for species discovery, pp. 250-285 in Zoological Journal of the Linnean Society 186 on pages 272-274, DOI: 10.1093/zoolinnean/zly035, http://zenodo.org/record/3089237

DROMOCHORUS WELDERENSIS DURAN, HERRMANN, ROMAN & EGAN SP . NOV . (FIGS 8B, 9F) Common name Gulf prairie tiger beetle. Type locality Welder Wildlife Foundation, Sinton, TX. Holotype (deposited in NMNH): 1 ♂, Texas: San Patricio Co. / Welder Wildlife Foundation / 11-June-2013 /Coll: A. Mitchell. Paratypes: 18 ♂♂, 15 ♀♀, Texas: San Patricio Co./Welder Wildlife Foundation/11-June-2013/ Coll: A. Mitchell. 2 ♂, 2 ♀, Texas/Buckeye - Matagorda Co./6-8-17 // J.D. Mitchell collector (NMNH). 2 ♀, Texas: San Patricio Co./Sinton/14-V-1966 // leg. W.T. Murray (JSC). 2 ♂, Texas, Victoria/VI-2–06 // C.R. Jones collector. 1 ♀, Texas: Dickinson/May 29 // TAMU-ENTO X0898573 (TAMUIC). 1 ♂, 1 ♀, Texas: Bee Co./Pettus/10.V.1964 // Leg. Pryor (SFASU). 4 ♂, 1 ♀, Texas: Nueces Co./Luetgens Coll. (AMNH). 1 ♀, Texas: Corpus Christi/VI-7-1969/ C.W. Griffin // Nueces River Park (NMNH). Distribution Found in the Gulf Prairie ecoregion of coastal Texas, from Houston area to Corpus Christi area. Diagnosis Dromochorus welderensis is diagnosable by having a black dorsum, often with a faint dark-blue sheen, and no pitting, subsutural foveae or infuscations, in conjunction with all dark maxillary palps and a pronotum with decumbent white setae. This species is most likely to be confused with D. chaparralensis, belfragei, velutinigrens or minimus. Dromochorus chaparralensis may be nearly indistinguishable from D. welderensis morphologically, but is ecologically differentiated. The habitat of D. chaparralensis is forested mesquite-chaparral, unlike the Gulf prairie habitat of D. welderensis. Dromochorus belfragei possesses regular pits on the elytra and often subsutural foveae. Maxillary palpi have a contrasting dark apical segment, with other segments dark yellow-testaceous to dark red-testaceous. Dromochorus velutinigrens has a very prominent green, blue, or violet dorsal sheen. The body is substantially more narrow and gracile, especially in males. Male D. velutinigrens have an all dark labrum, whereas D. welderensis males possess a pale central spot. Dromochorus velutinigrens have few to no setae on disk of pronotum. Dromochorus mimimus is generally smaller (Fig. 6; Table 2), and possesses sparse, thin, erect setae on the pronotum. This species also occurs further inland in forested mesquite-chaparral, unlike the Gulf Prairie grassland habitat of D. welderensis. Description Medium to large-sized Dromochorus. Body length 10.9– 14.7 mm, mean ♀ 13.7 mm, mean ♂ 12.6 mm. Head slightly wider than pronotum. Head charcoal brownblack with metallic green, green-blue, or bronze reflections mostly limited to the lateral ridge of the supraorbital region. Fine to marked rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black often with metallic green to violet reflections, with shallow longitudinal striae gradually ending at border of vertex. Clypeus black, with metallic violet to green-coloured reflections. Male labrum tridentate with 6–8 setae, central area pale ochre-testaceous, with a thin, dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-quarter of the total labrum surface; female labrum tridentate with 6–8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently darkbrown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2–3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3–4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5–11 dulltextured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.6–3.3 mm in length, mean ♀ 3.1 mm, mean ♂ 2.9 mm; width 2.7–3.4 mm, mean ♀ 3.2 mm, mean ♂ 2.9 mm. Pronotum charcoal brown to black, slightly wider than long, widest near anterior margin, width to length ratio 0.9 to 1.1, setae sparse to regular, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black, with metallic violet reflections, glabrous. Elytra elongate, dorsal surface convex, 6.3–8.6 mm length, mean ♀ 8.1 mm, mean ♂ 7.4 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration charcoal brown to black, often with faint blue reflections throughout elytral surface; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae brown to black, with metallic violet to blue reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet reflections, densely clothed in decumbent white setae; tibiae brown, clothed with setae of two types: sparser brown-testaceous long setae and dense, short, decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter mostly black with metallic olive green and violet reflections. Decumbent setae present on ventrite 1. Ventrites 2–6 have sparse, short, brown erect setae present throughout, but often abraded. Etymology Named for the type locality,WelderWildlife Foundation, in Sinton, Texas, as well as the Foundation’s namesake, Robert H. Welder, who established the foundation with the mission to conduct research and education in wildlife management and conservation. Ecology/natural history Adults have a long activity period, from mid-May through early August (A. Mitchell, pers. comm.). Dromochorus welderensis occurs in the Gulf Prairie ecoregion of the Coastal Plain physiographic province of Texas. This Dromochorus is the least associated with tree cover, and D. welderensis is consistently found in tall grasses in upland prairie habitat. Beetles are often found near heavy clayloam or clay banks and hills. This shade-loving species can be observed in early cool mornings or early evenings, and will try to avoid more open areas on hot, clear days. Even when active, adults are particularly reclusive and tend to stay hidden in tall grasses. They are more reluctant to forage in wide open loam areas, in contrast to D. pruininus and D. velutinigrens. Beetles can also be found on thick grass mats as they forage, mate or disperse. When disturbed, they use these dead grass mats as cover.

DROMOCHORUS BELFRAGEI SALLÉ, 1877 (FIGS 7B, 9B) Common name Loamy-ground tiger beetle. Type locality ‘Texas–Dallas, Wasco (presumed to be a misspelling of Waco), etc., on the banks of the Trinity River’ (English translation). Syntypes unknown, probably in MHNP (Bousquet 2012). Synonymy Dromochorus bellefragei Heyne, 1893 (unjustified emendation). Dromochorus sericeus Casey, 1897: 294. Type locality ‘Texas’. Two syntypes in USNM, Washington DC (synonymy established by Leng, 1902). Taxonomic history Dromochorus belfragei has been a catch-all taxon, and many populations of Dromochorus were lumped under this name in the literature or in museum collections prior to this revision. Dromochorus pilatei and D. velutinigrens are the only nominal taxa that have never been considered conspecific with D. belfragei. Based on morphology, ecology, biogeography, mtDNA genealogy and multilocus genetic data, D. belfragei (sensu stricto) is circumscribed as a separate species from three other distinct species, D. pruininus, D. knisleyi sp. nov. and D. minimus sp. nov. Occasional hybridization with D. knisleyi has been observed where their ranges come in contact, and this is further supported by evidence of mtDNA introgression. Distribution Dromochorus belfragei is known to occur from the north-western panhandle of Texas and southern Oklahoma to south-eastern Texas. Previous literature described a more extensive range from south-eastern Colorado (Michels et al., 2008) to Tamaulipas, Mexico (Cazier, 1954). The former record now appears to be an error (Michels, pers. comm., 2015), and the latter would appear to belong to D. chaparralensis sp. nov. One specimen from the AMNH was labelled ‘St. George, Utah’, although this locality would not appear plausible. The range of D. belfragei comes close to several other species. In particular, D. belfragei is nearly sympatric with D. pruininus in several places (Fig. 3; see D. pruininus account). In eastern and southern Bexar County, TX, this species is found sympatrically with D. knisleyi, and possibly D. minimus. Diagnosis This species can be distinguished from all other similar Dromochorus by the presence of finely to coarsely pitted black elytra that are not frosted in texture, in conjunction with maxillary palpi that have the apical segment darkest, with dark yellow-ochre to dark amber coloration in other segments (Fig. 8). Most populations of D. belfragei possess at least small subsutural foveae, but these lack metallic reflections. Southern and eastern populations may have subtle dark infuscations on the elytra. Dromochorus belfragei is most likely to be confused with D. pruininus, knisleyi or welderensis. It can be separated from these taxa by the following: 1. Dromochorus pruininus has a frosted blue sheen on the elytra and the entire dorsal surface, and it lacks any pits or subsutural foveae. The maxillary palpi in D. pruininus are pale yellow-ochre with a dark apical segment, whereas D. belfragei has darker yellow-red to red testaceous palpi with a dark apical segment. 2. Dromochorus knisleyi is similar to D. belfragei, but has more prominent subsutural foveae, often with bright metallic blue, green or gold reflections. In D. knisleyi, the maxillary palpi are always dark in all segments, whereas in D. belfragei, the apical segment is darker than the preceding. Infuscations are always present in D. knisleyi and are usually absent in D. belfragei. Ecologically, D. knisleyi is found in upland juniper woodland in the Edwards Plateau, and D. belfragei is found outside of this region, in clay soils associated with larger riparian systems. The two species appear to hybridize along a narrow contact zone at the edge of the Balcones Escarpment in south-eastern and south-central Texas. Both species and their hybrids may be found in this area. The existence of a hybrid zone was further supported by mtDNA data (Fig. 2). 3. Dromochorus welderensis has a smooth velvety black elytral surface (may have metallic blue-violet reflections), and never has pitting or subsutural foveae. The maxillary palpi are always dark in all segments, whereas in D. belfragei, the apical segment is darker than the preceding. Description Medium- to large-sized Dromochorus. Body length 11.6–15.2 mm, mean ♀ 14.5 mm, mean ♂ 13.2 mm. Head slightly wider than pronotum. Head predominantly black with blue reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae bright polished metallic violet to blue, with shallow, longitudinal striae gradually ending at border of vertex. Clypeus mostly black, with patches of metallic blue or violet reflections; clypeus may be nearly entirely blue to violet in females. Male labrum tridentate with 6–8 setae, central area pale ochre-testaceous, with a thin dark-brown to black bor- der posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, in others the pale area may cover more than two-thirds of the total labrum surface; female labrum tridentate with 6–8 setae, entirely dark-brown to black with polished metallic cupreous to green reflections. Maxillary and labial palpi with apical segment darker than other segments; basal to penultimate segments yellow-ochre to dark red-amber, often with metallic purple and green reflections. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2–3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3–4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5–11 dulltextured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.5–3.4 mm in length, mean ♀ 3.1 mm, mean ♂ 2.8 mm; width 2.8–3.6 mm, mean ♀ 3.4 mm, mean ♂ 3.1 mm. Pronotum black, with some dark-blue reflections, especially in sulci, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.2, setae sparse to regularly spaced, mostly present along lateral third of dorsal surface; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with weak to strong iridescent blue reflections, glabrous. Elytra convex, elongate, 7.1–9.7 mm length, mean ♀ 8.8 mm, mean ♂ 8.2 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with regular small pits present throughout disk, elytral coloration black, often with blue reflections near humeral region; elytral maculations absent; infuscations rarely present; subsutural foveae, when present, only slightly more prominent than other pits on the elytral disk; subsutural foveae lacking bright metallic reflections, except rarely in basal area. Legs: Pro-, meso- and metacoxae dark testaceous to black with iridescent blue to violet and cupreous reflections, with numerous setae; pro- and meso- trochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous, dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter mostly black with occasional metallic violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2–6 have sparse, short, brown erect setae present throughout, but often abraded. Ecology/natural history Dromochorus belfragei adults are active between mid- May and early July in most of its range (e.g. central to north TX) and late June to late July in northern part of its range (e.g. OK to panhandle of TX). Dromochorus belfragei can be found in natural and managed forested and agricultural areas (e.g. pecan groves) that have semi-open shaded areas or trails beneath the canopy. Soils in these areas can be dark, red to black in color, clay to clay-loam, cracked and sometimes moist in low areas that are heavily trampled by cattle. Adult beetles tend to avoid the lighter coloured sandy areas that are exposed to full sunlight on the forest edges. Despite the common name, this species appears more closely associated with soils that possess high clay content, more so than loam. The preferred habitat of D. belfragei is generally associated with larger riparian systems, although the beetles are not typically found near the water’s edge and we have found them over 3 km from water. They appear to have a wider ecological niche than most Dromochorus; their habitat and geographic distribution encompass a larger number of ecoregions than other species (Fig. 3; Table 1). Dromochorus belfragei and D. pruininus have similar but non-overlapping ranges, and they may be separated by ecological barriers, at least in some areas. In the DFW area of North Texas, the species has been reported a few kilometers from D. pruininus (Pearson et al., 2006), but the two appear to be separated by a narrow extension of the EPA Level IV Eastern Cross Timbers ecoregion, which is not suitable for either species. The forested undergrowth of this area is extremely dense in many areas, and may not possess the necessary surface soil conditions for either species to persist. Dromochorus belfragei may be more susceptible to urbanization than D. pruininus, as none have been collected in the DFW area (Tarrant Co.) since the 1970s, whereas D. pruininus appear much more tolerant to these disturbances and may be abundant in semi-open grassy areas where trails have been established in riparian parks (Dallas, Collin Co.). Thought to be crepuscular, this species is active throughout the day in shady areas or when overcast. Similar in behaviour to D. pruininus and D. pilatei, this species has been observed using soil cracks for escape, especially during dry conditions when virtisolic cracks are pronounced. Like other Dromochorus, D. belfragei frequently moves to vegetated cover to escape when pursued.

DROMOCHORUS MINIMUS DURAN, ROMAN, HERRMANN & EGAN SP.NOV. (FIG. 9D) Common name Pygmy dromo tiger beetle. Type locality SE of Pleasanton, TX. Holotype (deposited in U S N M): 1 ♂, U S A: Te x a s: A t a s c o s a C o./S E o f Pleasanton / 19-VI-2014 /leg D. Duran. Paratypes: 3 ♂ ♂, 3 ♀ ♀, U S A: Te x a s: A t a s c o s a C o./S E o f Pleasanton/19-VI-2014/leg D. Duran. 3 ♂, 4 ♀♀, USA: Texas: Atascosa Co./SE. of Pleasanton/30-V- 2014 / leg D. Sunberg. 1 ♂, 1 ♀, USA: Texas: Atascosa Co./SE. of Pleasanton/02-VI-2015/leg S.J. Roman. 17 ♂♂, 7 ♀♀, USA: Texas: Atascosa Co./SE. of Pleasanton/29-VI-2014/leg D. Brzoska. 1 ♂, USA, Texas: Bexar Co./7 miles S. San Antonio/06-VI-2010/ leg G. Waldren. 1 ♂, USA: Texas: Bexar Co./Loop 1604 Hwy/12-VI-2016/leg J. Back. Distribution Central/south Texas, south of the Balcones Escarpment. Currently known only from Bexar, Atascosa and Frio Counties. Diagnosis Dromochorus minimus can be separated from all other species, by the presence of a frosted or ashy grey to beige dorsum, sometimes with blue reflections, in conjunction with labial palpi that are all dark (apical segment is not darker than other segments), and sparse erect setae on the pronotum, often irregularly placed throughout. This species is most likely to be confused with D. pruininus, chaparralensis or welderensis. Dromochorus pruininus is generally larger (Fig. 6; Table 2), has pale maxillary palps with a contrasting dark apical segment and is also separable by geographic range. Dromochorus chaparralensis is usually larger (Fig. 6; Table 2), and lacks any prominent frosted texturing on the dorsal surface. The pronotum of D. chaparralensis has setae more regularly arranged, mostly along lateral third. Dromochorus welderensis is usually larger (Fig. 6; Table 2), and its pronotum has decumbent white setae, mostly along lateral third. The habitat of D. welderensis is Gulf Coast Prairie. Description Small- to medium-sized Dromochorus. Body length 10.5–13.7 mm, mean ♀ 12.7 mm, mean ♂ 11.7 mm. Head slightly wider than pronotum. Head predominantly charcoal black with blue to green reflections mostly concentrated near the anterior margin and edges of the supraorbital region. Fine rugosity often present on the frons and vertex. All head portions glabrous except for two supraorbital setae next to each eye. Frons concave in median area, especially in male, bulging towards slightly convex near anterior margin, clearly delimited from clypeus, gradually blending into vertex. Genae black with bright polished metallic green to violet reflections, with shallow longitudinal striae gradually ending at border of vertex. Clypeus shining black with blue to violet reflections throughout. Male labrum tridentate with 6–8 setae, central area pale ochre-testaceous, with a thin dark-brown to black border posteriorly and sometimes anteriorly, dark-brown to black laterally; in some populations, the pale central area of the labrum may exist as a small spot, up to one-third of the total labrum surface; female labrum tridentate with 6–8 setae, entirely dark brown to black with polished metallic cupreous to green reflections. All segments of maxillary and labial palpi consistently darkbrown; apical segment is not darker than other segments. Antennae normal length, reaching back to humerus and basal third of elytron, slightly longer in male than female; scape dark testaceous to black with metallic reflections of violet, cupreous and green, with 2–3 apical setae; pedicel dark testaceous with metallic reflections of violet, cupreous and green, lacking any setae; flagellum dark testaceous, antennomeres 3–4 with metallic violet and green reflections, densely clothed in short white setae, antennomeres 5–11 dull-textured without metallic reflections and possessing erect setae in apical rings only, covered with fine pubescence throughout. Thorax: Pronotum 2.4–3.1 mm in length, mean ♀ 2.8 mm, mean ♂ 2.7 mm; width 2.5–3.3 mm, mean ♀ 3.0 mm, mean ♂ 2.7 mm. Pronotum charcoal black, typically with frosty pale grey to brown, or blue to violet sheen, especially along lateral margins, slightly wider than long, widest near anterior margin, width to length ratio 1.0 to 1.1, thin erect setae sparse to irregularly spaced on pronotum; disc finely rugose, with thin but distinct median line, with well-defined shallow sulci present anteriorly and posteriorly; notopleural sutures clearly defined, not visible from dorsal view; proepisternum black with iridescent olive green to violet reflections, glabrous. Elytra elongate, dorsal surface convex, 6.4–8.0 mm length, mean ♀ 7.6 mm, mean ♂ 7.1 mm, shape similar in both sexes, but slightly wider in female, especially toward apical third; sutural spine absent, microserrations not present on elytral apices; elytral texture dull, with no pitting present, elytral coloration charcoal black, typically with grey, brown or blue-grey frosted texture along lateral margins, apex with blue or grey frosted texture; elytral maculations absent; subsutural foveae absent. Legs: Pro-, meso- and metacoxae dark brown with iridescent violet and cupreous reflections, numerous setae on pro- and mesocoxae, sparse on metacoxae; pro- and mesotrochanters with a single erect seta, metatrochanter glabrous, trochanters dark brown-testaceous; femora black with metallic violet and green reflections, densely clothed in decumbent white setae; tibiae testaceous brown, clothed with setae of two types: sparser brown-testaceous long setae and dense short decumbent white setae; two tibial spines present; tarsi brown-testaceous, first three dilated protarsomeres in male with dense greyish-white setal pad. Abdomen: Venter black with metallic olive green and violet reflections. Decumbent white setae present on ventrite 1. Ventrites 2–6 have sparse short brown erect setae present throughout, but often abraded. Etymology Dromochorus minimus is named for its smaller size. On average, this species is the smallest in the genus (Table 2). Ecology/natural history Little is known about this species’ natural history. Adults have been collected from mid-May until late June, but it is possible that the species may be active outside of this window. Dromochorus minimus occurs in mesquite-chaparral savannah in central/south Texas, just south of the Edwards Plateau, part of the larger Gulf Coastal Plains physiographic province. It has been found in open grassy areas interspersed between mesquite trees and clumps of Opuntia cactus. Adult beetles may be found venturing into the open spaces between clumps of grass, and will rapidly run into vegetation if pursued. Dromochorus minimus appears to be remarkably swift, even compared to other species of Dromochorus. In direct sunlight, live specimens appear beige-grey to smoky blue-grey.

Este artículo contiene 15 páginas, 7 figuras, 3 tablas., Benthic (streambed) biofilms metabolize a substantial fraction of particulate organic matter and nutrient inputs to streams. These microbial communities comprise a significant proportion of overall biomass in headwater streams, and they present a primary control on the transformation and export of labile organic carbon. Biofilm growth has been linked to enhanced fine particle deposition and retention, a feedback that confers a distinct advantage for the acquisition and utilization of energy sources. We quantified the influence of biofilm structure on fine particle deposition and resuspension in experimental stream mesocosms. Biofilms were grown in identical 3 m recirculating flumes over periods of 18–47 days to obtain a range of biofilm characteristics. Fluorescent, 8 mm particles were introduced to each flume, and their concentrations in the water column were monitored over a 30 min period. We measured particle concentrations using a flow cytometer and mesoscale (10 mm to 1 cm) biofilm structure using optical coherence tomography. Particle deposition-resuspension dynamics were determined by fitting results to a stochastic mobile-immobile model, which showed that retention timescales for particles within the biofilm-covered streambeds followed a power-law residence time distribution. Particle retention times increased with biofilm areal coverage, biofilm roughness, and mean biofilm height. Our findings suggest that biofilm structural parameters are key predictors of particle retention in streams and rivers., This study was funded by a Marie Curie Intra- European Fellowship to WRH (FP7- PEOPLE-2011-IEF-302297) and an Austrian Science Fund grant to T.J.B. (START Y420-B17). K.R.R. was supported by a CUAHSI Pathfinder fellowship and U.S. NSF Graduate Research Fellowship. J.D.D. was supported by a Fulbright-Spain fellowship. The modeling effort was supported by U.S. NSF grants EAR- 1215898 and EAR-1344280 to AIP. Supporting data are provided at doi:10.6084/m9.figshare.4252193.

Rivers are important ecosystems under continuous anthropogenic stresses. The hyporheic zone is a ubiquitous, reactive interface between the main channel and its surrounding sediments along the river network. We elaborate on the main physical, biological, and biogeochemical drivers and processes within the hyporheic zone that have been studied by multiple scientific disciplines for almost half a century. These previous efforts have shown that the hyporheic zone is a modulator for most metabolic stream processes and serves as a refuge and habitat for a diverse range of aquatic organisms. It also exerts a major control on river water quality by increasing the contact time with reactive environments, which in turn results in retention and transformation of nutrients, trace organic compounds, fine suspended particles, and microplastics, among others. The paper showcases the critical importance of hyporheic zones, both from a scientific and an applied perspective, and their role in ecosystem services to answer the question of the manuscript title. It identifies major research gaps in our understanding of hyporheic processes. In conclusion, we highlight the potential of hyporheic restoration to efficiently manage and reactivate ecosystem functions and services in river corridors.

To improve early enrollment in HIV care, the Swaziland Ministry of Health implemented new linkage procedures for persons HIV diagnosed during the Soka Uncobe male circumcision campaign (SOKA, 2011–2012) and the Swaziland HIV Incidence Measurement Survey (SHIMS, 2011). Abstraction of clinical records and telephone interviews of a retrospective cohort of HIV-diagnosed SOKA and SHIMS clients were conducted in 2013–2014 to evaluate compliance with new linkage procedures and enrollment in HIV care at 92 facilities throughout Swaziland. Of 1,105 clients evaluated, within 3, 12, and 24 months of diagnosis, an estimated 14.0%, 24.3%, and 37.0% enrolled in HIV care, respectively, after adjusting for lost to follow-up and non-response. Kaplan-Meier functions indicated lower enrollment probability among clients 14–24 (P = 0.0001) and 25–29 (P = 0.001) years of age compared with clients > 35 years of age. At 69 facilities to which clients were referred for HIV care, compliance with new linkage procedures was low: referral forms were located for less than half (46.8%) of the clients, and few (9.6%) were recorded in the appointment register or called either before (0.3%) or after (4.9%) their appointment. Of over one thousand clients newly HIV diagnosed in Swaziland in 2011 and 2012, few received linkage services in accordance with national procedures and most had not enrolled in HIV care two years after their diagnosis. Our findings are a call to action to improve linkage services and early enrollment in HIV care in Swaziland.