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Galloisiana yezoensis Asahina, 1961 was described on the basis of female specimens from the Ishikari Mountains, Hokkaido, Japan, and has been known as the only species of Grylloblattidae in Hokkaido. In preliminary research, several species closely related to G. yezoensis were collected from various sites in Hokkaido. In the present study, the taxonomic status of G. yezoensis and its relatives was reexamined through molecular and morphological analyses approaches. As a result, the phylogenetic tree of the COII region, and mitochondrial DNA, suggest that G. yezoensis and its relatives form a unique clade distinct from the Galloisiana clade. We therefore establish a new genus, Arctigalloisiana gen. nov., with two new species, A. poropnetopa sp. nov. and A. yubariensis sp. nov. The type species is Arctigalloisiana yezoensis comb. nov., and its male is described here for the first time.

Grylloblatta sp. “Ice crawler” Figures 72 (lateral), 73 (dorsal, ventral) Plates 45 (lateral), 46 (dorsal, ventral) Ice crawlers are remarkably active insects given their ecology, and studies have investigated their tolerance to cold (Schoville et al., 2015). Grylloblatta head morphology, especially musculature, was detailed by Wipfler et al. (2011) using both micro-CT and scanning electron microscopy, but tracheal morphology was notably absent from this study. As the sister group to Embioptera and Phasmatodea, the tracheal morphology of Grylloblattodea was important for assessing homology of these orders. Several head tracheae, including H-Oc, are difficult to localize. Grylloblattodea have very few ommatidia (Wipfler et al., 2011), and perhaps this is why tracheation appears reduced around the eyes relative to other taxa. In the thorax, T 2-DB branches off T 2- L, an unusual condition, as typically T 2-DB extends directly from T 2- S. This branch is very likely T 2-DB, as it leads to T 2- DLT, T 3- DLT, and successive tracheae as seen elsewhere. The branching pattern of T 2,3- AWL and PWL is also notable, in that it is unclear if these tracheae branch from T 2,3-DB or directly from T 2,3- S. Both conditions are present in other taxa, and this could merely be an “intermediate” positioning. DESCRIPTION: HEAD: Major tracheae indicated in plates, refer to figure 74 for head detail. Due to looping, networked 3-dimensional nature of Grylloblatta head tracheae, readers are encouraged to refer to digital supplemental models (see Availability of Digital Data, above). H-DCT slightly ventrad at cervix, curving dorsally on entry into head capsule; H-DCT divides into H-DCT-Vi1, H-DCT-Vi2, and H-Ant anteriad of cervix. H-DCT-V1 lateral and anteriad, following head capsule wall. H-Ant anteriad, curving laterally to extend into antennae. H-DCT-Vi2 anteriad, with each side curving anteriad to meet in X-shaped intersection at H-DX and continuing anteriorly before arcing ventrad to connect with H-VCT-Loop via dorsal-ventral branch H-DVB. H-Ant on specimen left side with link to H-DVB. Just posteriad of cervix, each H-DCT with small branch extending medially, joining in Y at base of head capsule into medial branch, extending anteriad as H-DCMedB (dorsal cephalic medial branch). H-DCMedB splits near vertex into three branches; two extend laterally, looping backward to connect with H-Ant as H-DCT-Loop; last branch directly dorsad, connecting to H-DX. H-VCT directly anteriad, with H-Lbm branching ventrad at posterior margin of submentum. H-VCT continuing anteriad with branches to H-Md and H-Lbr before curving medially to join H-VCT from opposite side in dorsal two branches of X-shaped H-AX. Ventral two branches of H-AX formed by H-Lbm, which continues anteriad before arcing dorsally to H-AX, forming H-Lbm-Loop. H-LbmPlp branching from H-Lbm-Loop near anterior curve of H-Lbm toward H-AX. H-Lbm extending anteriad from H-AX anteriormost point; H-Md extending anteriad from dorsal portion of H-Lbm-Loop before curving ventrad. H-DVB dorsal-ventral connection of H-DCT and H-VCT at branching off point of H-Md. Small H-VC running ventrad near anterior end of H-VCT-Loop. THORAX: T 2- S with four branches: H-DCT, H-VCT, T 2-DB, T 2-VB. T 2- CT absent. H-DCT runs anteriad from T 2- S with slight curve medially; branching of T 1- PL and T 1-Dvi at posterior margin of prothorax, with T 1- PL running laterally and ventrad, proceeding into proleg and T 1-Dvi directly anteriad; T 1-DB beginning just anterior of this split, extending dorsad to join with T 1- DLT in Y-shaped junction. H-DCT compressed laterally anterior of the split; compression likely preservational artifact. H-VCT similar in size to H-DCT, with single T 1- L branching directly ventrad, located mid prothorax. T 2-DB running initially dorsad splitting off large T 2- AWL, smaller T 2-DB continues, extending medially and dorsad, arcing posteriorly to connect to T 3- S via T 2- DLT. T 2- AWL arcing ventrally and posteriad, dividing into T 2- AL and T 2-Awba; T 2- AL continuing posteriorly into midleg, joining with T 2- PL near mesothoracic coxa; small T 2-Awba branching from T 2- AWL, dorsad near apex of arc toward T 2- L. T 2-VB short, proceeding ventrad and medially before splitting into T 1- VLT anteriad and T 2- VLT posteriad and ventrad. T 1- VLT with three ventral commissures: T 1-VC1 most anterior, at the apex of a long oval loop reaching the posterior end of the cervix; T 2-VC2 ventrad, joining at the opposite end of loop to forming X-shaped T 1-VX; T 2-VC3 near the prothoracic coxae; T 1-AL-VC branching ventrad from T 1- AL to connect with loop at halfway between T 1-VC1 and T 1-VC2. T 2- VLT runs ventrad and posteriad, extending toward mesothoracic coxae before turning dorsally to join with T 3-VB. T 2- VLT with three ventral commissures: T 2-VC1 most anterior, branching directly toward midline to join with opposite side; T 2-VC2 branching inward where T 2- VLT arcs dorsally; T 2-VC3, branching from metathoracic portion of T 2- VLT, branching medially and joining with T 2-VC2 at X-shaped intersection T 2-VX. T 2-Cx comprised of two branches, one each extending from T 2-VC2 and T 2-VC3. T 3- S with only two branches, short T 3-DB and T 3-VB. T 3-DB dorsad, splitting into T 2- PWL anteriad and ventrad, T 3- AWL arcing dorsad and posteriorly, and remaining small T 3-DB continuing dorsally to link T 2- DLT and T 3- DLT in Y-shaped junction. T 3-AWba absent; T 3-AWbr absent. Short T 2- PWL runs anteriad, dividing into ventral T 2- PL that links with T 2- AL from anterior, continuning into T 2- L; and small dorsal T 2-PWba. T 2-PWbr not present. T 3-VB short, bifurcating into connection to T 2- VLT from anterior and continuing as T 3- VLT toward posterior. T 3- VLT with two ventral commissures: T 3-VC1 slightly ventrad from T 2- VLT / T 3- VLT intersection; T 3-VC2 where T 3-Cx extends into base of metacoxae. T 3- VLT extends from split of T 3-Cx, continuing in shallow, ventral arc to link with A 1- VLT via A 1-VB. ABDOMEN: A 1..8- S present. All abdominal segments similar, with notable exceptions below. Short A n -SB present, bifurcating into A n -DB and A n -VB. A n -DB runs directly dorsad, linking with A n - DLT in Y-shaped junction; A 5-DB with short A 5-DB-Vi extending mediad. A n - DLT runs in shallow, dorsal arch along tergum; A n -DC absent; short, mediad A 4..6-DLT-Vi present; A 7-DLT-Vi beginning posteriad before arcing medially and then anteriad, extending past A 6- S; A 8- DLT extending posteriad into A-Cr. A 1-SB with two tracheae ventrad: T 3- L ventrad and slightly anteriad, extending into hind leg, along same path as A 1-SB; A 1-VB smaller, extending directly ventrad. A [2,4]-VB extending directly from A [2,4]-SB with no branches. A 3-VB runs directly from A 3-SB, with A 3-VB-Vi continuing ventrad before turning posteriorly while smaller A 3-VB directly ventrad. A [5,6] similar to A 3 with large A [5,6]-VB- Vi anteriad and smaller A [5,6]-VB ventrad. All A n -VB-Vi link with A n - VLT in Y-shaped junction, similar to A n - DLT, but with a less pronounced arch ventrad. A 8-VC present, other ventral commisures not visible and likely absent. Small A 9-VC visible., Published as part of Herhold, Hollister W, Davis, Steven R, Degrey, Samuel P & Grimaldi, David A, 2023, COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA, pp. 1-184 in Bulletin of the American Museum of Natural History 459 (1) on pages 1-184, DOI: 10.5531/sd.sp.55, http://zenodo.org/record/7730159, {"references":["Schoville, S. D., R. A. Slatyer, J. C. Bergdahl, and G. A. Valdez. 2015. Conserved and narrow temperature limits in alpine insects: thermal tolerance and supercooling points of the ice-crawlers, Grylloblatta (Insecta: Grylloblattodea: Grylloblattidae). Journal of Insect Physiology 78: 55 - 61.","Wipfler, B., R. Machida, B. Muller, and R. G. Beutel. 2011. On the head morphology of Grylloblattodea (Insecta) and the systematic position of the order, with a new nomenclature for the head muscles of Dicondylia. Systematic Entomology 36 (2): 241 - 266."]}

Herhold, Hollister W, Davis, Steven R, Degrey, Samuel P, Grimaldi, David A (2023): COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA. Bulletin of the American Museum of Natural History 459 (1): 1-184, DOI: 10.5531/sd.sp.55, URL: http://dx.doi.org/10.5531/sd.sp.55

Order Eoblattida Handlirsch, 1906, sensu Aristov, 2015, comprises 16 extinct families and a single extant one, Grylloblattidae Walker, 1914. Three species of the genus Stereopterum Carpenter, 1950 (Euryptilonidae) from the Elmo, Kansas and Midco, Oklahoma Lower Permian were the only Wellington Formation taxa attributed by Aristov (2015) to the Eoblattida. This assignment was based on work by Carpenter (1950, 1966), for the Kansas records and Beckemeyer and Hall (2007), for the Oklahoma record. In this paper, a new eoblattid genus and species is described based on a specimen collected by Paul Tasch in the 1960′s from the Lower Permian Wellington Formation of Marion County, Kansas and is assigned to family Blattogryllidae Rasnitsyn, 1976.

Grylloblatta campodeiformis campodeiformisWalker (Grylloblattodea: Grylloblattidae) was commonly collected during summer from trees killed by the mountain pine beetle,Dendroctonus ponderosaeHopkins (Coleoptera: Curculionidae), in subalpine forests of Alberta, Canada. Gut content analysis revealed that the grylloblattids fed on subcortical invertebrates. This newly reported habitat association shows that this species is not limited to strictly alpine habitats and glacial margins, and thus may be more widespread and common than suggested by earlier reports.

The enigmatic insect order Grylloblattodea comprises two described species in Canada, which are limited to the Montane Cordillera and Pacific Maritime ecozones. One of the described species has three Canadian subspecies of uncertain taxonomic ranking, and there are two additional undescribed or unreported species known in close proximity to the Canadian border in western Alberta and British Columbia that likely also occur in Canada. Thus, as much as 50% of the total taxonomic diversity of Grylloblattodea in Canada is still undocumented. Targeted surveys and taxonomic work, as well as studies that describe the ecology and conservation status of Grylloblattodea are important goals for future research.

For many terrestrial species, habitat associations and range size are dependent on physiological limits, which in turn may influence large-scale patterns of species diversity. The temperature range experienced by individuals is considered to shape the breadth of the thermal niche, with species occupying temporally and/or geographically stable climates tolerating a narrow temperature range. High-elevation environments experience large temperature fluctuations, with frequent periods below 0 °C, but Grylloblatta (Grylloblattodea: Grylloblattidae) occupy climatically stable microhabitats within this region. Here we test critical thermal limits and supercooling points for five Grylloblatta populations from across a large geographic area, to examine whether the stable microhabitats of this group are associated with a narrow thermal niche and assess their capacity to tolerate cold conditions. Thermal limits are highly conserved in Grylloblatta, despite substantial genetic divergence among populations spanning 1500 m elevation and being separated by over 500 km. Further, Grylloblatta show exceptionally narrow thermal limits compared to other insect taxa with little capacity to improve cold tolerance via plasticity. In contrast, upper thermal limits were significantly depressed by cold acclimation. Grylloblatta maintain coordinated movement until they freeze, and they die upon freezing. Convergence of the critical thermal minima, supercooling point and lower lethal limits point to adaptation to a cold but, importantly, constant thermal environment. These physiological data provide an explanation for the high endemism and patchy distribution of Grylloblatta, which relies on subterranean retreats to accommodate narrow thermal limits. These retreats are currently buffered from temperature fluctuations by snow cover, and a declining snowpack thus places Grylloblatta at risk of exposure to temperatures beyond its tolerance capacity.

Rock-crawlers (Grylloblattodea: Grylloblattidae) in northeastern Asia are low-vagility insects that are restricted to cool temperate forests and mountainous regions. Morphologically distinguishable species are similar ecologically and show narrow endemism and a patchy distribution. As a result, grylloblattids are hypothesized to be relict species that have persisted in situ over long periods of climatic and geological change (Storozhenko and Oliger 1984). We investigate whether the diversification pattern of Asian grylloblattids reflects long-term persistence and divergence due to geological events, or more recent diversification in response to climatic change. Using multilocus genetic data, we examine the phylogenetic relationship to other Asian Grylloblattidae and the geographic pattern of diversification of Korean rock-crawlers, Galloisiana Caudell & King (1924) and Namkungia Storozhenko & Park (2002). Our analysis reveals a monophyletic grouping of Korean species, with multiple cryptic lineages and restricted geographical distributions. Based on genetic data, Korean species are closely related to Japanese Galloisiana. Using a Bayesian relaxed clock model calibrated with a mitochondrial substitution rate, the age of the most recent common ancestor of the Korean—Japanese lineage is estimated within the Miocene epoch. This provides evidence for a diversification event closely tied to the geological events separating the Japanese archipelago from the Korean peninsula.

New members of the family Blattogryllidae are described, including Permoblattogryllus praecox gen. et sp. nov. from the Middle Permian Soyana locality in Russia, Protoblattogryllus nedubrovensis from the Lower Triassic Nedubrovo locality in Russia, Madygenocephalus micropteron gen. et sp. nov. (the second known case of brachyptery in grylloblattids), and Costatoviblatta similis sp. nov. from the Middle Triassic Madygen locality in Kyrgyzstan. Protoblattogryllus zajsanicus Storozhenko, 1990 from the Upper Permian Karaungir II locality in Kazakhstan is redescribed. The genus Microblattogryllus Storozhenko, 1990 from Madygen is considered to be a synonym of Protoblattogryllus Storozhenko, 1990. Protoblattogryllus abruptus Storozhenko, 1990 from the Madygen locality is transferred to the genus Mesoblattogryllus Storozhenko, 1990.

Grylloblatta chintimini, n. sp. (Figs. 2 A���F) Material examined. Holotype male, OSAC #0000583178, with label ��� OR. Benton Co. Marys Peak; on snow bank just below summit; 1m. snow. 2 IV 2006 1200 m.; D. Lytle; 44 �� 30 ��� 15 ��� N 123 �� 33 ��� 18 ������W ���; separate label with code number ��� 2009 ��� written in pen; separate label with ��� HOLOTYPE ��� written in pen. Specimen in ethanol, deposited at OSAC. Extracted genomic DNA OSAC #0000583178: DNA:01. Five paratypes (4 females, 1 male), Oregon, Benton County, Marys Peak, on snow bank just below summit, 44 �� 30 ��� 15 ��� N 123 �� 33 ��� 18 ������W, elevation 1200 m, 31 III 2006, D. Lytle and C. Marshall, colls. Specimen codes: allotype female, OSAC #0000583179// 2012 (OSAC); three females, OSAC #0000583181// 2010 (USNM), OSAC #0000583180// 2011 (CAS), and OSAC #0000583182// 2013 (OSAC); one male, OSAC #0000583183// 2014 (USNM). Four specimens with label data same as holotype: two males, OSAC #0000583187// 2008 (CAS), OSAC #0000583189// 2007 (OSAC); two females, OSAC #0000583188// 2015 (OSAC), OSAC #0000583190// 2016 (OSAC). Three specimens, Oregon, Benton County, Marys Peak on snow, above top parking lot, 2 April 2006, D. A. Lytle, coll., all deposited in OSAC: two males, OSAC #0000583184// 2004, OSAC #0000583185// 2005; one female, OSAC #0000583186// 2006. Description: Neotype 14.6 mm long (measurements taken on ethanol-preserved specimen). Antenna with 33 segments on both left and right. Abdominal tergites and sternites dark, approximately Dresden brown to mummy brown; abdominal sternites finely pubescent, with larger porect setae evenly and prominently distributed across surface; abdominal tergites finely pubescent, with several erect setae along posterior margins. Cerci 9 -segmented, shorter than abdomen; left cercus length 4.8 mm. Head width 2.8 mm, length 2.6 mm. Pronotum length 2.4 mm, width 2.3 mm. Interocular distance 2.0 mm, eye width 0.4 mm. Left hind femur length 4.4 mm, width 0.8 mm. Left hind tibia length 4.6 mm. Supra-anal plate borne asymmetrically, right posterior apical corner less acute; bearing several major setae on surface and lateral edges; right posterolateral angle (angle formed by the right anterolateral edge of supra-anal plate and a line tangent to the curve formed between the right and left posterolateral lobes) obtuse, approximately 100 ��. Right gonocoxite with fine pubescence; single larger seta on ventral edge; anterodorsally with distinct notch anterior of dorsal finger (Fig 2 B). Gonostyli borne laterally on basal half, bearing three or four major setae on posterolateral edge; longest setae exceeding width of gonostyli. Primary copulatory structure with apical beak stout and with posterior face not expanded into a prominent dorsal flange (Fig. 2 D). The secondary accessory sclerite of right phallomere (Fig 2 E,F) distinctly quadrate, appearing chisel-, or tooth-shaped. Size small for genus. Antennal segment counts on specimens range from 31 to 33. Female with similar body coloration to male but generally larger in size. Ovipositor approximately three-fourths the length of cerci; dorsal valvulae slightly longer than ventral valvulae (Fig 2 A). Diagnosis. Although currently G. chintimini is only known from the type locality at Marys Peak in the Oregon Coast Range, this species is similar to G. ro t h i and G. newberryensis from the Oregon Cascades, which would all be identified as G. ro t h i using the key published in Storozhenko (1988), given that they share similar antennal segmentation, relatively short cerci and a strongly asymmetrical male supra-anal plate. However, the species are quite distinct; in addition to having different COII mtDNA (Table 1), G. chintimini has a darker brown, less reddish body coloration and a relatively narrower head and prothorax than either G. rothi or G. newberryensis. The terminus of the male primary copulatory sclerite in G. chintimini differs from that of G. rothi in lacking an expanded apicaldorsal flange, which is also present in G. newberryensis although less pronounced than in G. ro t h i. The secondary accessory sclerite of the right phallomere is also distinct in comparison to both G. ro t h i and G. newberryensis. The cerci and dorsal valvulae of the ovipositor of G. chintimini are comparatively similar to those of G. ro t h i and noticeably shorter than those of G. newberryensis. The only known locality for G. chintimini is Marys Peak (often erroneously referred to as Mary���s Peak). This peak, about 15 miles WSW of Corvallis, Oregon, is the highest point in the Oregon Coast Range. Snow is not uncommon in Oregon���s coast range, but Marys Peak is one of relatively few places high enough in elevation to reliably accumulate snowpack that lasts from fall until spring. Several collecting trips to nearby peaks of lower elevation that might provide suitable grylloblattid habitat have yielded no specimens, so it remains to be seen if G. chintimini is endemic to Marys Peak or has a more widespread distribution. Etymology. The word chintimini is a version of T���cha teemanwi, the Kalapuya tribe place name for Marys Peak (McArthur 2003)., Published as part of Marshall, Christopher J. & Lytle, David A., 2015, Two new species of Grylloblatta Walker, 1914 (Grylloblattodea: Grylloblattidae) from western North America, and a neotype designation for G. rothi Gurney 1953, pp. 408-418 in Zootaxa 3949 (3) on pages 412-414, DOI: 10.11646/zootaxa.3949.3.6, http://zenodo.org/record/234609, {"references":["Storozhenko, S. Y. (1988) A review of the family Grylloblattidae (Insecta). Articulata, 3, 167 - 181.","McArthur, L. A. (2003) Oregon Geographic Names. Oregon Historical Society Press, Portland, OR, 1073 pp."]}

Grylloblatta rothi Gurney 1953 (Figs. 1 A���F) Material examined. Neotype male, OSAC #0000583177, ��� OR: Deschutes Co., Cultus Mtn. on/near summit 6720 ��� 10 T590843, 4852390; Lytle & Marshall, 21 June 2011 ��� with code number ��� 7048 ��� written in pencil on label margin; right metathoracic leg detached, left metathoracic leg missing (removed for sequencing). The UTM coordinates on the label correspond to 43.8190 ��N, - 121.8703 ��W. Specimen in ethanol, to be deposited at USNM. Extracted genomic DNA deposited at Oregon State Arthropod Collection, Corvallis, OR, OSAC #00000583177: DNA:01. Thirteen specimens (3 males, 7 females, 3 juveniles), same collection data as neotype, deposited in OSAC (OSAC _AC_ 2012 - 12 - 12 -001). All specimens were collected on snow fields at night near the summit, except a single juvenile found beneath a rock. Numerous carabid beetles and spiders were active on the snowfields as well. Description: Neotype body length 12.2 mm (measurements taken on ethanol-preserved specimen). Antenna with 24 segments on left and 25 on right [Gurney (1953): 29 on left, 16 on broken right antenna]. Abdominal sternites finely pubescent, with relatively few larger porect setae distributed across the surface. Abdominal tergites finely pubescent, with several erect setae along posterior margins. Cerci with 9 articulated segments, right cercus length 2.2 mm; first two basal segments appear to be comprised of two fused segments (indicated by two rings of trichobothria, which generally occur at the apex of an articulated segment, and faint vestiges of articulation); cerci shorter than the abdomen proper. Head width 2.4 mm, length 2.2 mm. Pronotum length 2.5 mm, width 2.1 mm. Interocular distance 1.8 mm, eye width 0.5 mm. Left hind femur length 3.3 mm, width 0.8 mm. Left hind tibia length 3.3 mm. Male supranal plate strongly asymmetrical; left finger-like lobe approaching the longitudinal midline of the abdomen; right finger-like posterior lobe less developed, at an approximate 60 �� angle. Male gonocoxae (Fig 1 B,C; gcx-l, gcx-r) asymmetrical, covered in fine setae with a few larger ones basomedially. Gonostyli (Fig 1 B,C gst-l, gst-r) articulate medially (not basally), appearing ���hammerhead-like��� when viewed laterally. Right gonocoxa more elongate than left; bearing a distinct dorsal finger that projects medially and opposes the beak-like apex of the primary copulatory sclerite. Primary copulatory sclerite (main phallic sclerite in Gurney 1953) with dorsolateral lobe about half as long as lateral margin below the dorsal cap (Fig 1 E, pcs). Secondary accessory sclerite (apical lobe of accessory sclerite of right phallomere in Gurney 1953) (Fig 1 D,E, sas) rounded and lobe-like. Apex (terminus) of primary male copulatory sclerite expanded along apico-dorsal edge into noticeable flange (Fig. 1 F, pcs ���arrow). Distribution. Kamp (1973) placed G. ro t h i within an assemblage of species inhabiting the high Cascades of Central Oregon, specifically Broken Top and Belknap Crater. Cultus Mountain is within this area, a region largely defined by the peaks of the Three Sisters. In the original description, Gurney also described an allotype, a single female specimen known from Crater Lake. He united these two specimens (the only specimens in the rothi assemblage known to Gurney) based on their relatively short cerci. Later researchers, including the present authors, expressed doubt that these two specimens were actually conspecific. Kamp, who was likely the last person to view the original type material, and who had in his possession specimens from throughout Oregon, believed the population at Crater Lake to be a distinct species (Kamp 1973). Our own research into populations in the vicinity of Crater Lake confirm his suspicions and we do not believe Grylloblatta rothi to be distributed this far south of the central Cascades. Remarks. Gurney (1953) described the holotype male as generally pale ���antimony yellow��� with a ���dark yellow ochre��� dorsum. The neotype and other specimens in our possession are consistent with this general yellowishbrown description and to our eyes their overall color approximates yellow ochre. As in Gurney���s original description, G. ro t h i we have examined are generally small for the genus, with alcohol-preserved male specimens ranging from 11.9 to 12.5 mm and females from 14.3 to 15.0 mm in length. The number of antennal segments in adult Grylloblatta is commonly used in identification keys to species, despite that it is a notoriously variable character between, and even within, individuals in a single population. The neotype differs in its antennal segment count from that reported for the left antenna in Gurney���s holotype. However it must be noted that specimens from the same population as the neotype show a range in antennal counts, ranging from 19 to 29 antennal segments and even display intra-individual discrepancies between right and left antenna as great as 10 segments. Thus, we do not feel that the precise number of antennal segments (in either the holotype or neotype) is of great importance for understanding the species. The male copulatory sclerites were not described in great detail by Gurney (1953) but he does note that the male dorsolateral lobe of the primary copulatory sclerite is about ���about half as long as lateral margin below the dorsal cap������a description that is consistent with the dorsolateral lobe of seen in male Grylloblatta from the Cultus Mountain population (dll, Fig. 1 E). Gurney did not mention that the primary male copulatory sclerite is expanded apicodorsally into a flange, a trait that is shared with G. newberryensis. However, this flange is only visible when viewed from a specific angle (notably an anterior facing view, as shown in the illustrations herein (Fig. 1 F), which was not depicted in Guney���s illustrations of the holotype. Gurney did describe and illustrate the secondary accessory sclerite (referred to as the ���apical lobe of the accessory sclerite of the right phallomere���) as being rounded and lobe like, consistent with the males from Cultus Mountain (Fig 1 D,E, sas). Lastly, the cerci of this species are noticeable short in comparison to many other Grylloblatta species, a trait noted by Gurney in 1953, ���The shortness of the cercal segments separates rothi from all other species except possibly barberi, and the character is so distinctive that the association of sexes [holotype and allotype] is believed correct.��� However, both species described below share this trait as well as several other traits found in G. rothi and a more detailed, comparative study of the species allied to G. ro t h i, is currently being conducted by the authors. Female cerci are only slightly longer than the ovipositor. The ovipositor is ventrally covered in short setae, which is denser at the base and apex than they are medially. The dorsal valvula is slightly longer than the ventral valvula (Fig 1 A). Paratypes (including the allotype) of a species have no special significance in the determination of a neotype. Still, they may provide information to help clarify the species concept being employed by the original author. Unfortunately, in this particular case, the species concept was based on only two specimens (the holotype and allotype) that Gurney treated as conspecific based solely on the short cerci. We searched for the allotype along with the holotype unsuccessfully and we presume they were lost at the same time., Published as part of Marshall, Christopher J. & Lytle, David A., 2015, Two new species of Grylloblatta Walker, 1914 (Grylloblattodea: Grylloblattidae) from western North America, and a neotype designation for G. rothi Gurney 1953, pp. 408-418 in Zootaxa 3949 (3) on pages 410-412, DOI: 10.11646/zootaxa.3949.3.6, http://zenodo.org/record/234609, {"references":["Gurney, A. (1953) Recent advances in the taxonomy and distribution of Grylloblatta (Orthoptera: Grylloblattidae). Journal of the Washington Academy of Sciences, 43, 325 - 332.","Kamp, J. W. (1973) Biosystematics of the Grylloblattodea. Doctoral dissertation, University of British Columbia. [total page number unkown]"]}

Grylloblatta rothi Gurney, 1953 is redescribed and a neotype is designated from Cultus Mountain in the Oregon Cascades, U.S.A. Two new species of Grylloblatta are described, bringing the total number of Grylloblatta species to 15. Grylloblatta chintimini new species is described from Marys Peak in the Coast Range of Western Oregon, where it occurs on snowpack near the 1250 m summit. Grylloblatta newberryensis new species is described from Newberry Volcano in Central Oregon, where it is associated with snowfields overlying geologically-young lava flows. Morphological characters, primarily derived from male genitalia, are presented to diagnose these species and differentiate them from other Grylloblatta spp. in Oregon, Washington, and California. Molecular sequences from the cytochrome oxidase subunit II gene suggest that significant divergence has occurred among these species and provide a tool to aid identification of juvenile and female specimens.

The presumed phylogenetic link between extant ice-crawlers (Grylloblattidae = 'crown-Grylloblattida') and fossil species of the taxon concept Grylloblattida sensu Storozhenko (2002) is essentially based on postabdominal morphology. However, the fossil data are limited, and the interpretation is open to debate. Here we investigate a sample of a poorly known fossil 'grylloblattidan' family, the Bajanzhargalanidae, collected from the Daohugou locality (Middle Jurassic, China). We describe Sinonele fangi gen. nov., sp. nov., Sinonele hei gen. nov., sp. nov., Sinonele phasmoides gen. nov., sp. nov., and Sinonele mini gen. nov., sp. nov. Thanks to the abundance and exceptional preservation of the material, we could document wing venation intra-specific variability, provide cues to identify male and female individuals, describe and tentatively interpret various body structures of both sexes, and discuss them with a broad pterygotan phylogenetic perspective. The Bajanzhargalanidae exhibit a puzzling combination of postabdominal characters leaving us inconclusive on their affinities, or lack thereof, with crown-Grylloblattida. Our contribution suggests that a substantial effort will be needed to further investigate postabdominal structures from comparatively ancient fossil insects preserved as rock imprints, because of their broad morphological disparity.

An extant species of the relict insect order Grylloblattodea is described from the Ussuri River Basin of southeastern Russia. This species, Galloisiana olgae, is a member of the family Grylloblattidae that probably originated as a lineage during the mid-Cenozoic and experienced subsequent range constriction associated with Pleistocene glaciation. The Grylloblattodea have a richer fossil history in warm-temperate habitats during the Late Paleozoic than the four confamilial genera of today would suggest. These modern taxa represent a specialized Cenozoic lineage that adapted to cool-temperate habitats in northwestern North America and northeastern Asia, and parallel other similar distributions of seed plants and insects.

Schoville, Sean D., Graening, G. O. (2013): Updated checklist of the ice-crawlers (Insecta: Grylloblattodea: Grylloblattidae) of North America, with notes on their natural history, biogeography and conservation. Zootaxa 3737 (4): 351-378, DOI: http://dx.doi.org/10.11646/zootaxa.3737.4.2

Grylloblatta chandleri Kamp, 1963 U.S.A., California. Lassen Co. Ice caves near Eagle Lake, 1.5 miles WSW of Spaulding Tract, elev. 5,180 ft., 5 adult males and 1 adult female coll. by H.P. Chandler, T.L. Rodgers and J.W. Kamp (paratypes), from 12 June to Sep. 1951 (Kamp 1953, 1963); 1 adult female coll. by J.W. Kamp (allotype), 9 July 1951 (Kamp 1963); on moist rocks in cave, 1 adult male coll. by V.D. Roth (holotype), 5 Sep. 1959; 5,000 ft. elev., 1 nymph coll. by H.P. Chandler, 0 2 May 1952 (CAS); Griffith Meadows Ice Caves, 1962 (Kamp 1970), also referred to as Griffith Ranch Ice Caves by Peck (1973); unidentified ice cave (Kamp 1973); Lassen National Forest, Gray's Flat, Township 32 north, Range 7 east, Section 33, northeast 1 / 4, southwest ��, N 40.628 W - 121.18932 1825m elev., on rock talus after onset of snowstorm, 1 nymph coll. by D.R. Cluck, 0 6 Nov. 2005 (Schoville & Roderick 2010); Lassen Peak, NE slope in Devastated Area, 1 adult female coll. by S.D. Schoville and R. Y. Dudko, 17 May 2013. Shasta Co. North Christmas Tree Cave, 1 nymph coll. by G. O. Graening, 21 April 2012; Subway Cave, Lassen National Forest, 1 nymph coll. by G. O. Graening, 0 3 Nov. 2012. Tehama Co. Wilson Lake Ice Cave, Mt. Lassen (Kamp 1970); Wilson Lake Ice Cave, 1615m elev., 1 adult female coll. by M.D. Terry, T.H. Ogden, and K.J. Jarvis, 22 Aug. 2001 (BYU: Jarvis & Whiting 2006); 1 nymph coll. by K.J. Jarvis, G.T. Svenson, and J. Jones, 18 June 2002 (BYU: Jarvis & Whiting 2006). Kamp (1963) suggested G. chandleri is widespread in the Modoc Plateau and Basin Ranges of Eastern California, and later stated that ������almost every ice cave explored between Eagle Lake and the Medicine Lake Highlands contains populations of Grylloblatta. ��� (Kamp 1973). Grylloblatta chirurgica Gurney, 1961 U.S.A. Washington. Skamania Co. Ape Cave, 1 adult male and 3 adult females coll. by H. Reese and family (holotype, allotype and 2 paratypes), 22���28 Nov. 1959 (Gurney 1961); 1 adult female and 1 nymph coll. by R.L. Crawford and C.M. Senger, 14���30 May 1983 (BMUW: Jarvis & Whiting 2006); 2 adults coll. by R.L. Crawford and C.M. Senger, 27 Oct. to 11 Nov. 1983 (BMUW); 4 specimens (CMNH); Bat Cave, 1,160 ft. elev., 1 adult female coll. by W.R. Halliday, 0 1 Nov. 1960 (BMUW); 1 adult coll. by R.L. Crawford and C.M. Senger, 22 July to 0 6 Aug. 1983 (BMUW); 1 adult coll. by R.L. Crawford and C.M. Senger, 18 Nov. to 8 Dec. 1983 (BMUW); H. Reese, 0 1 March 1961; 1 adult male coll. by D. Nielsen, 18 Nov. 1972; Bean Creek, Mt. St. Helens National Volcanic Monument, 1 adult coll. by K. Johnson, 15���28 June 1983 (BMUW); Beaver Cave, 1,520 ft. elev., 1 adult coll. by R.L. Crawford and C.M. Senger, 24 July 1983 (BMUW); R.L. Crawford and C.M. Senger, 27 Oct. to 19 Nov. 1983 (BMUW); Butte Camp A on talus slope, Mt. St. Helens National Volcanic Monument, 1 adult coll., 0 5 Oct. 1983 (BMUW); Cave 6 miles NE Cougar, 3 specimens coll. by R. Zack, 31 Jan. 1976 (WSU); Dollar and A Dime Cave, 1,290 ft. elev., 1 adult female coll. by R.L. Crawford and C.M. Senger, 18 Nov. to 0 8 Dec. 1983 (BMUW); 1 adult female coll. by R.L. Crawford and C.M. Senger, 21 July to 0 6 Aug. 1983 (BMUW); Gremlin Cave, 2,980 ft. elev., 1 adult male coll. by R.L. Crawford and C.M. Senger, 07��� 23 Oct. 1983 (BMUW); 1 adult and 2 nymphs coll. by R.L. Crawford and C.M. Senger, 0 5 July 1983 (BMUW); Little People Cave, 2,840 ft. elev., 1 adult and 1 nymph coll. by R.L. Crawford and C.M. Senger, 23 July to 0 5 Aug. 1983 (BMUW); Manhole Cave, 3,000 ft. elev., 1 nymph coll. by R.L. Crawford and C.M. Senger, 0 8 Oct. 1983 (BMUW); N. Butte Camp, 46.175 N 122.225 W, 4,520 ft. elev., 1 adult coll. by R.L. Crawford, 0 3 Nov. 1982 (BMUW); 1 adult coll. by D. Mann, 1��� 16 Oct. 1981 (BMUW); 2 adults coll. by C.J. Becker, 12���24 June 1983 (BMUW); Lake Cave, 1,900 ft. elev., 1 adult female (paratype) coll. by W.R. Halliday, 28 Dec. 1958 (Gurney 1961); 1 nymph coll. by R.L. Crawford and C.M. Senger, 0 7 Dec. 1983 (BMUW); 1 nymph coll. by R.L. Crawford and C.M. Senger, 25 May to 27 June 1983 (BMUW); 1 nymph coll. by R.L. Crawford and C.M. Senger, 23 Nov. 1973 (BMUW); Mt. St. Helens, Timberline Rd., 3,800 ft. elev., 1 specimen coll. by A. Smetana, 4 July 1978 (CMNH); Ole���s Cave, Mt. St. Helens south end lava beds, 1 specimen coll. by R. Zack, 0 2 February 1978 (WSU); 1 adult female coll. by C.M. Senger, 25 Aug. 1965 (BMUW); 2 adults and 2 nymphs coll. by R.L. Crawford and C.M. Senger, 21 May to 26 June 1983 (BMUW); 3 specimens coll. by R.L. Crawford and C.M. Senger, 27 Oct. to 12 Nov. 1983 (BMUW); Nielsen���s Cave, 2 nymphs coll. by W.R. Halliday, 1958 (Gurney 1961); caves on west flank of Mt. St. Helens (Kamp 1970); NE slope Mt. St. Helens, 1,520���1,620 m elev., 1 adult female and 1 nymph coll. by D.H. Kavanaugh, 5 Aug. 1974 (CAS); Pumice Plain, nymphs coll. by P.S. Sugg and J.S. Edwards, 1984���1985 (Sugg & Edwards 1998); NW side of Lava Dome, Mt. St. Helens, observation of C.H. Anderson, Jr. and M.R. Vining, Sep. 1997 (Anderson & Vining 1999); Prince Albert Cave, 1,200 ft. elev., 1 adult coll. by R.L. Crawford and C.M. Senger, 18 Nov. to 0 8 Dec. 1983 (BMUW); 2 adults coll. by. R.L. Crawford and C.M. Senger, 22 July to 0 2 Aug. 1983 (BMUW); Spider Cave, 5.1 miles NE of Cougar, 2,840 ft. elev., 1 adult and 1 nymph coll. by R.L. Crawford and C.M. Senger, 05��� 31 July 1983 (BMUW); 1 nymph coll. by R.L. Crawford, 0 6 Aug. 1977 (BMUW); 1 adult coll. by G. Ericson, 12 Nov. 1966 (BMUW); Spirit Lake at Bear Creek, 3,200 ft. elev., 1 adult coll. by A. Smetana, 0 6 July 1974 (BMUW); Surprise Cave, 3,240 ft. elev., 1 adult female coll. by R.L. Crawford and C.M. Senger, 23 July 1983 (BMUW; Jarvis & Whiting 2006). Yakima Co. Chinook Pass, 0.25 mi. SE of Chinook Pass Picnic Area, 5,360ft. elev., Z 10 E 613220 N 5192760, 1 adult male coll. by E.A. Lisowski, 10 Nov. 2003 (BYU; Jarvis & Whiting 2006); Goat Peak Trail, 5,880 ft. elev., 1 adult male coll. by E.A. Lisowski, 11 Sep. 2002 (BYU; Jarvis & Whiting 2006); White Pass at summit, under rock in forest north of US 12, specimens coll. by J.C. Bergdahl (J.C. Bergdahl, pers. comm.). Probable records. Washington. Kittitas Co. Mount Baldy, base of large rocks on ridge, 14.3 miles S-SE Ellensburg, UTM 10 Z, 696040 mE, 5186640 mN, 2,200 ft. elev., 1 adult and 3 nymphs coll. by E.A. Lisowski, 21 Dec. 1997; Wilson Creek, 10 miles N. Ellensburg, 1 nymph coll. by S.D. Smith, 17 Dec. 1982 (BMUW). Lewis Co. Eagle Peak Trail, 11.7 miles NW Packwood, UTM 10 Z, 591340 mE, 5177730 mN, 2,960 ft. elev., coll. by E.A. Lisowski, 11���25 February 1995 (BMUW); Laughing Water Creek, 46.753 ' N 121.548 'W, 2,600 ft. elev., 9 nymphs coll. by J. Putera, 08��� 17 February 1988 (BMUW); 1 adult female and 9 nymphs coll. by J. Putera, 17���24 February 1988 (BMUW); 1 adult male, 7 adult females and 10 nymphs coll. by J. Putera, 24 February to 0 5 April 1988 (BMUW); 1 adult coll. by J. Putera, 15 April to 0 6 May 1988 (BMUW); 1 adult female coll. by J. Putera, 28 Sep. to 27 Oct. 1988 (BMUW); 2 adult males, 2 adult females and 40 nymphs coll. by J. Putera, 27 Oct. to 0 1 Dec. 1988 (BMUW); coll. by J. Putera, 11 Dec. 1988 to 0 5 Jan. 1989 (BMUW); 2 adult males and 1 nymph coll. by J. Putera, 15���22 Jan. 1989 (BMUW); 1 nymph coll. by J. Putera, 15���24 Jan. 1989 (BMUW); 3 adults and 3 nymphs coll. by J. Putera, 21 Feb. to 23 March 1989 (BMUW); 5 adults coll. by J. Putera, 22 Jan. to 21 Feb. 1989 (BMUW); Tongue Creek Trail, 46.426 N 121.780 W, 2,900 ft. elev., 1 nymph coll. by J. Putera, 17���28 Jan. 1988 (BMUW). Pierce Co. 0.1 mi. East Route 410 Entrance, 13.5 miles S SE Greenwater, Mt. Rainier National Park, UTM 10 Z, 611640 mE, 5203560 mN, 2,800 ft. elev., 2 nymphs coll. by E.A. Lisowski, 22 Dec. 1995 (BMUW); Bear Head Mountain Summit, 47.023 N 121.814 W, 6,000-6,089 ft. elev., 1 adult coll. by R. Crawford, 15 Aug. 1982 (BMUW); Carbon River, 14.7 miles SE Wilkeson, UTM 10 Z, 590400 mE, 5201740 mN, 2,600 ft. elev., 1 specimen coll. by E.A. Lisowski, 0 4 Feb. 1995 (BMUW); Upper Spray Park, 1 adult coll. by J.S. Edwards, 0 2 June 1979; Westside Road, 2.7 miles N-NE main road and 7.1 miles E-NE Ashford, UTM 10 Z, 585120 mE, 5181140 mN, 2,920 ft. elev., 1 adult coll. by E.A. Lisowski, 0 7 Jan. 1996 (BMUW); 14.7 miles SE Wilkeson, UTM: 10 Z, 590400 mE, 5201740 mN, 1 adult coll. by E.A. Lisowski, 4 Feb. to 11 March 1995 (BMUW). Yakima Co. 0.5 mi SE Windy Point, 7.5 miles W Tieton, UTM 10 Z, 660000 mE, 5172300 mN, 2,520 ft. elev., 1 specimen coll. by E.A. Lisowski, 27 Feb. 2000 (BMUW); Forest Service Road 417 at elev. 2,960 ft., 9.2 miles W- SW Tieton, UTM 10 Z, 657560 mE, 5171040 mN, 1 specimen coll. by E.A. Lisowski, 24 June 2001; Little Rattlesnake Creek., middle of section 16, 11.9 miles S-SE Cliffdell, UTM 10 Z, 656080 mE, 5183700 mN, 2,480 ft. elev., 1 specimen coll. by E.A. Lisowski, 10 March 2001; Tieton Reservoir Road near Goose Egg Mountain, 1.7 miles SE Rimrock, UTM 10 Z, 645720 mE, 5167940 mN, 2,760 ft. elev., 1 specimen coll. by E.A. Lisowski, 31 July 1997; Thunder Creek, 2.9 mi. S-SE Goose Prairie, UTM 10 Z, 633560 mE, 5190360 mN, 5,480 ft. elev., 2 specimen coll. by E.A. Lisowski, 14 Oct. 2001; 2.7 mi. S-SE Goose Prairie, UTM 10 Z, 633460 mE, 5190660 mN, 1 specimen coll. by E.A. Lisowski, 0 3 Nov. 2001; US 12 between 1 st and 2 nd Tieton River Bridge, 7.9 miles W Tieton, UTM 10 Z, 659300 mE, 5173060 mN, 2,280 ft. elev., 1 specimen coll. by E.A. Lisowski, 24 Jan. 1998 (BMUW). Surveys showed that G. chirurgica occupied disturbed habitat in Mt. St. Helens within 4 years of the eruption in 1980 (Sugg & Edwards 1998). This species is sympatric with G. sp. nov. ���Sawyer���s Ice Cave # 1 " at Chinook Pass (Jarvis & Whiting 2006). Its range also approaches G. sp. nov. ���Mt. Rainier��� and G. sp. nov. ���Trout Lake Caves.��� Grylloblatta gurneyi Kamp, 1963 U.S.A. California. Siskiyou Co. All the following records are from lava tubes in or near Lava Beds National Monument: Merrill Ice Cave, 4,740 ft., 1 adult male and 1 adult female coll. by J. W. Kamp and D. H. Kistner (holotype and allotype), 20 Feb. 1960 (USNM; Kamp 1963); 1 adult male coll. by V.D. Roth (paratype), 1954 (Kamp 1963); R. Crawford, 24 Nov. 1989 (BMUW); 1 adult female and 3 nymphs coll. by K.J. Jarvis, G.J. Svenson, and J. Jones, 0 2 June 2002 (BYU; Jarvis & Whiting 2006); 1 adult female coll. by Fuhrman, K.J. Jarvis, G.J. Svenson, and J. Jones, 0 1 Jan. 2003 (BYU; Jarvis & Whiting 2006); 1 adult female and 1 nymph coll. by S.D. Schoville and R. Y. Dudko, 12 May 2013; 3 -level Ice Cave, 1 adult female coll. by C.M. Senger, 23 June 1984 (BMUW); 2 adults coll. by C.M. Senger, 23���29 June 1984 (BMUW); 1 adult male coll. by T. Briggs, 19 June 1973 (CAS); 1 specimen coll. by H. MacDonald et al., 1997; Angleworm Cave/Lost Pinnacle Cave, 1 nymph coll. by R.L. Crawford, 27 May to 23 June 1990 (BMUW); R. Crawford, 24 Nov. 1989 (BMUW); Arch Cave, R. Crawford, 20 Nov. 1989 (BMUW); Catacombs Cave, 5,000 ft. elev., 5 adults coll. by R.L. Crawford, 21 Nov. to 9 Dec. 1989 (BMUW); Cox Ice Cave, 4,580 ft. elev., 1 adult female coll. by R.L. Crawford, 27 May 1990 (BMUW); 1 adult female coll. by K.J. Jarvis, G.J. Svenson, and J. Jones, 20 June 2002 (BYU; Jarvis & Whiting 2006); Deep Ice Cave, NE of McCloud, 5,560 ft. elev., 1 specimen coll. by D. Cowan, and 3 nymph coll. by C.M. Senger, 23���29 June 1984 (BMUW); Frozen River Ice Cave, coll. by S.B. Peck (Peck 1973); Heppe Cave, SE grotto, 5,280 ft. elev., 1 nymph coll. by C.M. Senger, 22���29 June 1984 (BMUW); Indian Wells Ice Cave, 3 miles south of Merrill, 2 adult males and 4 adult females coll. by J.W. Kamp (paratypes), Sep. 1951 and 1954 (Kamp 1963); 2 nymphs coll. by S.D. Schoville and R. Y. Dudko, 12 May 2013; Mushpot Cave, 1 nymph coll. by R.L. Crawford, 21 Nov. to 9 Dec. 1989 (BMUW); Skull Cave, D. Cowan, 23 Nov. 1989 (BMUW). Probable records. California. Siskiyou Co. Bray Ice Cave, 2 adult females coll. by T. Briggs, 19 June 1973; Lava cave near Hambone, 1 adult male coll. by T. Briggs, 28 May 1977 (CAS); Jack Jones Ice Cave (Halliday 1962); Jake Bell Ice Cave(s) (Halliday 1962); Mayfield Ice Cave (Halliday 1962); Mt. Shasta, ���high elevations on snowbanks,��� coll. by S.B. Peck (Peck 1973); Mt. Shasta, 1 specimen, coll. unknown, 21 May 1978 (BME: not identified to species); Constance Creek, Mt. Shasta, 1 adult male, 1 adult female and 5 nymphs coll. by S.D. Schoville and R. Y. Dudko, 13 May 2013; Starr Cave (Halliday 1962, Peck 1973). Grylloblatta marmoreus Schoville, 2012 U.S.A. California. Siskiyou Co. Bigfoot Cave, Marble Mountains Wilderness Area, 1 adult male G. O. Graening and Guy Graening (holotype), 0 3 Sep. 2010 (Schoville 2012); Brokedown Palace Cave, collections in 1977 and 1978, B. Her (Suggett 1982); Marble Valley Guard Station, 16 specimens coll. by D. Hemphill, M. Winning, and R.L. Suggett, 0 1 Feb. 1980 (Suggett 1982); 3 specimens coll. by M. Winning and R.L. Suggett, 15 Feb. 1981 (Suggett 1982); trail to Marble Valley from Lover���s Camp, 1 specimen coll. by R.L. Suggett, 23 Feb. 1980 (Suggett 1982); Marble Valley, 30 + specimens coll. by S.D. Schoville and R. Y. Dudko, 14 May 2013; Planetary Dairy Cave, 1 nymph coll. by G. O. Graening and D. Weaver, 4 July 2010 (Schoville 2012). Grylloblatta oregonensis Schoville, 2012 U.S.A. Oregon. Josephine Co. Oregon Cave, Oregon Caves National Monument, 1 adult female coll. by J. Roth (holotype), 15 June 2010 (Schoville 2012); 1 adult, 10 Oct. to 15 Nov. 1992 (BMUW); one nymph, 30 Nov. 2006 (Schoville 2012); one nymph, 11 July 2008 (Schoville 2012); one nymph, 14 Jan. 2009 (Schoville 2012); one nymph coll. by S.D. Schoville, 20 March 2010 (Schoville 2012); 1 specimen coll. by C. Davis, April 1983; Kincaid���s dancehall, 5 nymphs, 31 May 1993 (BMUW); Wedding Cake room, 15 nymphs, 31 May 1993 (BMUW); 1 adult, 10 May 1993 (BMUW). This species is currently a single-site endemic and potentially cave-restricted (Schoville 2012). It is parapatric or sympatric with G. siskiyouensis at Oregon Caves National Monument. Grylloblatta rothi Gurney, 1953 U.S.A. Oregon. Deschutes Co. Happy Valley, 15 to 20 miles south of Sisters on Century Drive, 6,450 ft. elev., 1 adult male coll. by V.D. Roth (holotype), 12 Sep. 1918 (Gurney 1953); McKenzie Pass, Willamette National Forest, 1 specimen coll. by P.S. Ward, 19 May 1985 (BME); 2 specimens (CMNH; Kamp 1973); 44.25 ˚ N 121.81 ˚W, 5,220 ft. elev., 1 adult female coll. by J.S. Edwards, 24 June 1985 (BMUW); N 44.260 W -121.809, 1 specimen coll. by SSE, 23 June 1981; 3 adult males, 12 adult females and 8 nymphs coll. by K.J Jarvis, G.J. Svenson, and J. Jones, 24 June 2002 (BYU; Jarvis & Whiting 2006); Edison Ice Cave (Kamp 1970). Hood River Co. Mt. Hood, junction of Phlox Point and Timberline roads, 5,900 ft. elev., 2 adult males and 5 adult females coll. by J.W. Kamp, 18 June 1970 (Kamp 1973); mileage post 68 on Mt. Hood Loop Road, one nymph (NMNH); Tilly Jane Creek, Mt. Hood National Forest, N 45.39896 W - 121.64813 1,772 m elev., 1 adult male coll. by S.D. Schoville, 18 June 2009 (CAS). Klamath Co. Crater Lake, 2 adult females coll. by J.E. Elsea (paratype), 27-28 Nov. 1936 (CAS; USNM; Elsea 1937, Gurney 1953). Probable records. Oregon. Deschutes Co. 1 mi. NW Mt. Bachelor, 1 specimen coll. by J.G. Todd, 12 March 1983 (CMNH); Bachelor Butte (Kamp 1973); Belknap crater lava fields (T. 15 S, R. 8 E, USGS Three Sisters Quad.), 5,384 ft. elev., 5 adult males and 12 adult females coll. by J.W. Kamp, 18 June 1970 (Kamp 1973); one adult female coll. by J.M. Taylor, 18 July 1971 (NMNH); Crossbill Ice Cave, ~ 20 mi. SE of Bend, 2 adult females coll. by B. McGregor, 5 May 2013; Ice Cave N Newberry Crater; caves near Bend (Peck 1973); Lava Top Butte Ice Cave, 1 adult obs. by B. McGregor, April 2012; South Ice Cave at Fort Rock, 1 adult female coll. by J.W. Kamp (Kamp 1970). Linn Co. Lebanon (Kamp 1963). Marion Co. Boca Cave near Triangulation Peak, one adult obs. by B McGregor, Nov. 2011; Silver Falls State Park near Stayton, N 44.87877 W - 122.6590, several larvae obs. by M. Leppin, 13 Dec. 2008. There appears to be extensive sympatry of G. rothi and G. sculleni, including sites at Edison Ice Cave (Kamp 1970) and throughout the Three Sister region (Kamp 1973). Kamp (1973) identified a third species at the Belknap lava fields that is sympatric with both G. sculleni and G. rot hi at this site. However, the species description was never published. He also suggested samples from Mt. Hood form a new species (Kamp 1973), but our genetic and morphological analysis assign this population to G. rot hi. Grylloblatta scudderi Kamp, 1979 Canada. British Columbia. Whistler Mountain, Garibaldi Provincial Park, 1,951 m elev., 1 adult male, 1 adult female, and 9 nymphs coll. by L. Bartlett (holotype and allotype), 13 July 1970 (Kamp 1979). Probable records. Canada. British Columbia. Blowdown Pass in Stein Valley, near Pemberton, N 50.3626 W- 122.15, 1 adult coll. by R. Bennett and D.R. Copley, 15 Sept. 2012 (RBCM); Grouse Mountain (Spencer 1945); Mission Ridge above Seton Portage, near Lillooet, N 50.76356 W- 122.169806, 2,174m elev., 4 specimens coll. by R. Bennett, C.R. Copley and D.R. Copley, 3 August 2011 (RBCM); Mount Garibaldi (Kamp 1979). Kamp (1973) lists the Grouse Mountain record as ���doubtful��� due to extensive glaciations in this area. Grylloblatta sculleni Gurney, 1937 U.S.A. Oregon. Benton Co. Mary���s Peak (T. 12 S., R. 7 W., Section 28 NW 1 / 4 of the NE 1 / 4 of the NW 1 / 4), 3,960 ft. elev., 1 adult female coll. by J. LaBonte, 3 April 1993 (Jarvis & Whiting 2006); 1 adult female coll. by J. LaBonte, 3 April 1995; 3 adult females coll. by J. LaBonte, 10 Feb. 1996; Mary���s Peak, 14 miles west of Corvallis (Kamp 1973). Deschutes Co. Scott Camp, Three Sisters, 6,600 ft. elev., 3 adult females and 6 nymphs coll. by H.A. Scullen and R. Rider (holotype and paratypes), 12 July 1936 (CAS; USNM; Gurney 1937); 1 adult female and 2 nymphs coll. by H.A. Scullen and G. Ferguson (paratype), 0 6 Aug. 1935 (Gurney 1937); 1 adult female coll. by H. Scullen, 1 Aug. 1959 (CAS; Gurney 1961); Sunshine Meadow, 6,600 ft. elev., 1 adult female coll. by H. Crowell, 1 Aug. 1959 (Gurney 1961); Belknap crater lava fields, Scott Pass, south slope of North Sister, 6,500 ft. elev., 4 adult males coll. by J.W. Kamp, 22 July 1970 (Kamp 1973); Edison Ice Cave, Deschutes National Forest, 5,200 ft. elev., 2 adult males and 6 adult females (Kamp 1973); McKenzie Pass, 2 adult males coll. by J.W. Kamp, 18 July 1971 (CMNH; Kamp 1973). Probable record. Linn Co. Quartzville Creek �� mile E of Green Creek (Sec. 26, T 11 S, R 4 E), 1,900 ft. elev., 1 nymph coll. by H. Hacker, 14 Nov. 1959 (Gurney 1961). See comments on sympatry with G. rot hi. Kamp (1973) proposed subspecific status to a morphological variant of G. sculleni from Edison Ice Cave, but the description was never published. Grylloblatta siskiyouensis Schoville, 2012 U.S.A. Oregon. Josephine Co. Big Tree Loop Trail, Oregon Caves National Monument, 1 adult female coll. by A. Y. (h, Published as part of Schoville, Sean D. & Graening, G. O., 2013, Updated checklist of the ice-crawlers (Insecta: Grylloblattodea: Grylloblattidae) of North America, with notes on their natural history, biogeography and conservation, pp. 351-378 in Zootaxa 3737 (4) on pages 356-360, DOI: 10.11646/zootaxa.3737.4.2, http://zenodo.org/record/223034

fragments on a 2.5% agarose gel. The two primers used were C2-J-3400 (COII-specific) and TK-N-3785 (tRNAZY~-specific); for primer details see [14]. 1 Drosophila melanogaster (Diptera: Drosophilidae), 2 Grylloblatta rothi (Grylloblattoidea: Grylloblattidae), 3 Ruspolia nitidula (Orthoptera, Ensifera: Tettigonioidea), 4 Cyphoderis monstrosus (Orthoptera, Ensifera: Haglidae), 5 Locusta migratoria (Orthoptera, Caelifera: Acrididae), 60edipoda coerulescens (Orthoptera, Caelifera: Acrididae), 7 Prosphena scudderi (Orthoptera, Caelifera: Pyromorphidae) es that the rearrangement has occurred within Orthoptera, unquestionably a monophyletic group with respect to the other taxa in our sample. These two sets of observations indicate that the KD~DK rearrangement has occurred at least twice during insect evolution; once in the lineage leading to Hymenoptera, and once in the lineage leading to the orthopteran suborder Caelifera. However, the overall divergence of the Apis genome, including the apparently high number of tRNA rearrangements (11 with respect to Drosophila), indicates that the observed gene orders are probably the result of quite separate rearrangement events in the different lineages. Therefore, the observations support a previous study of mtDNA gene order evolution which stressed the importance of considering rearrangements of groups of tRNA genes rather than of individual genes [13]. This has important consequences for any future studies of insect evolution using tRNA gene orders and is, to our knowledge, the first reported case of an apparently homoplastic mtDNA gene rearrangement in metazoans. The result emphasizes the need to sample both a representative number of insects, and a large enough sample of tRNA genes in order to make a correct interpretation of the data. Received March 1 and May 8, 1995

Grylloblatta siskiyouensis new species (Figs. 1 B, 2 C���D) Type material. Holotype female in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M., Big Tree Loop Trail N 42.098 W- 123.407 WGS 84, Note: near water station 06-III- 2007 9: 15 pm AY SDS 10 - 004��� [white label]/ ��� HOLOTYPE Grylloblatta siskiyouensis Schoville det. S.D. Schoville 2011 ��� [red label]; Big Tree Trail of Oregon Caves National Monument, 42.098 o North and - 123.407 o West, Josephine County, Oregon, U.S.A. Other specimens. Three juveniles in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M., Blind Leads Cave, N 42.098 W- 123.407 WGS84, 20-III- 2010. Note: twilight zone under rocks��� [white label]/ ��� Grylloblatta siskiyouensis Schoville det. S.D. Schoville 2011 ��� [white label]. Two juveniles in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M., N 42.098 W- 123.407 WGS84, 20-III- 2010. Note: on snow surface��� [white label]/ ��� Grylloblatta siskiyouensis Schoville det. S.D. Schoville 2011 ��� [white label]. Etymology. The specific epithet, siskiyouensis, is named for the Siskiyou Mountains in which the type locality, Oregon Caves National Monument, is located. Diagnosis. Adult females of this species can be distinguished from all other Nearctic Grylloblatta species by the following combination of character states: color amber brown on dorsum and white ventrally with light amber yellow patches; small body size; more than 30 antennomeres; ratio of antennomere lengths 2 and 1 1.2; head and interocular distance wider than pronotum; compound eyes large; pronotum short and narrow with a notable point formed at posterior edge; ventral valve of ovipositor broad and slightly up-curved; dorsal valve of ovipositor projects beyond ventral valve one-fifth total length; ovipositor nearly as long as cercus; cercomeres short, distally brown and proximally white. Description of holotype female. Antennomere number on right antennae 32 and left antennae (broken) 19. Compound eyes well developed, large and black. Interocular distance slightly larger than pronotum width. Head width slightly greater than head length. Pronotum short and narrow, slightly convex, anterior end wider than posterior end, transverse sulcus at anterior end curving slightly towards the posterior end, a weak mid-dorsal line visible, and a notable point formed at the posterior edge. Leg coloration distinctly yellow compared to dorsal body and abdomen, and brown setae distinctive. Ovipositor nearly as long as cercus, extending to eighth cercomere. Ventral valve of ovipositor broad in the middle and slightly up-curved, roughened distally (possible scarring), with noticeable long setae. Dorsal valves of ovipositor extend beyond ventral valve by one-fifth total length and at midpoint nearly straighten out. Cercal segments short, distally brown and proximally white. Dorsal body color amber brown (55), legs light amber yellow (83), abdomen white laterally and light amber yellow patches ventrally, antennomeres and cercomeres amber brown fading to white at distal ends, ovipositor salmon-orange (50). Measurements of holotype female (cm). BL 1.295, HL 0.294, HW 0.326, EL 0.071, EW 0.0476, ID 0.223, Ant 1 L 0.0397, Ant 2 L 0.0476, AntRatio 1.2, PL 0.3176, PW 0.183, FL front 0.333, FL mid 0.397, FL rear 0.508, TL front 0.238, TL mid 0.349, TL rear 0.453, TrL front 0.071, TrL mid 0.103, TrL rear 0.175, PTrL front 0.270, PTrL mid 0.318, PTrL rear 0.318, OvL 0.437, OvW 0.135, CL 0.476. Description of other specimens. Juveniles are amber brown (55) on the dorsal surface of the head and pronotum. Comparisons. Grylloblatta siskiyouensis n. sp. can be distinguished from all Grylloblatta species by dark dorsal coloration and dark abdominal color patches, as well as notable point on posterior end of pronotum. It can be distinguished from G. oregonensis n. sp. by a smaller adult body size and limb length, notably narrower pronotum and head, antennomere count higher (> 30) and ratio of antennomere 2: 1 lower (G. ro t h i by a smaller body size, higher antennomere count (> 30), and long, setaceous ovipositor. It can be distinguished from G. sculleni by a cercus nearly equal in length to ovipositor. It can be distinguished from G. barberi by more reddish dorsal color, narrower head and pronotum, fewer antennomeres, a cercus nearly equal in length to ovipositor, and a less up-curved dorsal valve of ovipositor. It can be distinguished from G. gurneyi in having fewer antennomeres and ovipositor nearly equal in length to cercus. It can be distinguished from G. chandleri by fewer antennomeres and broader ventral valve of ovipositor. Habitat distribution. The type specimen was found walking on snow at night in pine forest. I collected two juvenile specimens on the snow at night on March 20, 2010 and three more in the twilight zone of Blind Leads Cave under stones. The adult female specimen examined by Jarvis and Whiting (2006) was collected in a cheesebaited trap near the entrance of a cave. Geographical distribution. Grylloblatta siskiyouensis n. sp. is presently known only from Oregon Caves National Monument (Fig. 3). Due to its surface activity, it is potentially more widespread in the Siskiyou Mountains. Kamp (1973) refers to a grylloblattid population near Mt. Ashland that might represent the same species. Phylogenetic relationships. Based on mitochondrial cytochrome oxidase subunit II data, G. siskiyouensis n. sp. (GenBank Accession JN 612961) is distinct from (minimum ~ 10 %) other nearby grylloblattids (Table 1). This species is represented by the Oregon Caves sample deposited in GenBank (Jarvis & Whiting 2006), from which it differs by a single base (0.1 %). In a neighbor-joining analysis, it is grouped with G. oregonensis n. sp. and G. m a rmoreus n. sp. in an unresolved polytomy (Fig. 4)., Published as part of Schoville, Sean D., 2012, Three new species of Grylloblatta Walker (Insecta: Grylloblattodea: Grylloblattidae), from southern Oregon and northern California, pp. 42-52 in Zootaxa 3412 on pages 47-48, DOI: 10.5281/zenodo.212975, {"references":["Jarvis, K. J. & Whiting, M. F. (2006) Phylogeny and biogeography of ice crawlers (Insecta: Grylloblattodea) based on six molecular loci: Designating conservation status for Grylloblattodea species. Molecular Phylogenetics and Evolution, 41, 222 - 237.","Kamp, J. W. (1973) Biosystematics of the Grylloblattodea. Department of Zoology, University of British Columbia, Vancouver, 275 pp."]}

Grylloblatta marmoreus new species (Figs. 1 C, 2 E���H) Type material. Holotype male in ethanol, deposited in CAS, labeled: ���Big Foot Cave, Marble Mountain Wilderness Area, Siskiyou Co., CA, G. O. Graening & Guy Graening, 3 Sept. 2010 ��� [white label]/ ��� HOLOTYPE Grylloblatta marmoreus Schoville det. S.D. Schoville 2011 ��� [red label]; Big Foot Cave, located in Marble Valley, approximately 41.5628 o North and - 123.1899 o West, elevation 5,935 ft, Marble Mountain Wilderness Area, Siskiyou County, California. Other specimens. Juvenile in ethanol, deposited in CAS, labeled: ���Planetary Dairy Cave, G. O. Graening and David Weaver, 4 July 2010 ��� [white label]/ ��� Grylloblatta marmoreus Schoville det. S.D. Schoville 2011 ��� [white label]; Planetary Dairy Cave, approximately 41.560 o North and - 123.197 o West, elevation 5,621ft, Marble Mountain Wilderness Area, Siskiyou County, California near the headwaters of the Canyon Creek watershed. Etymology. The specific epithet, marmoreus, means ���of marble��� in Latin and is named for the Marble Mountains of California where this specimen was collected. Diagnosis. Adult males of this species can be distinguished from those of all other Nearctic Grylloblatta species by the following combination of character states: color buff yellow; small body size for genus; relatively short limbs; fewer than 30 antennomeres; ratio of antennomere lengths 2 and 1 1.5; head wider than pronotum; compound eyes well-developed and medium size; pronotum trapezoidal with rounded proximal end and forming soft triangular point at posterior end; subgenital plate with slight rounded tooth; right coxopodite with sharply curved process; left coxopodite with short, stubby stylus; cercus short with short cercomeres. Description of holotype male. Antennomere count on right antennae 27 and broken left antennae 25. Antennomere 2 1.5 X larger than antennomere 1. Compound eyes well developed, length greater along body axis than width, and pigmentation black. Head slightly broader than pronotum but narrower than interorbital distance. Pronotum gently curved in anterior end and triangular at posterior end, overall trapezoidal as width at anterior end greater than posterior end, posterior end forming soft triangular point, transverse sulcus at anterior end curving slightly towards anterior end, length greater than width. Leg coloration amber yellow, slightly lighter than dorsal body color, and small brown setae present. Legs relatively short. Portion of right mid leg below femur removed and destroyed during DNA extraction. Cercus length short with short cercomeres. Subgenital plate more than twice as wide as long, distal edge hardened, distal end forming very slight rounded tooth. Right coxopodite longer than wide with a strong curve towards the ventral side of the body, a short process strongly curved towards anterior on dorsal side near base, stylus borne directly on distal end approximately 30 % of the length of coxopodite, and heavily pubescent. Left coxopodite width greater than length, stylus at distal end short and stubby (0.021cm). Dorsal and ventral body color buff yellow (87), appendages lighter amber yellow (83), antennomeres and cercomeres amber yellow fading to white at distal ends, subgenital plate and coxopodites amber yellow. Measurements of holotype male (cm). BL 1.256, HL 0.228, HW 0.255, EL 0.054, EW 0.034, ID 0.208, Ant 1 L 0.02, Ant 2 L 0.03, AntRatio 1.5, PL 0.235, PW 0.214, FL front 0.295, FL mid 0.268, FL rear 0.3886, TL front 0.235, TL mid 0.275, TL rear 0.402, TrL front 0.067, TrL mid 0.1005, TrL rear 0.154, PTrL front 0.201, PTrL mid 0.201, PTrL rear 0.214, SbgL 0.0536, SbgW 0.1273, RCxL 0.1005, RCxW 0.0804, LCxL 0.1139, LCxW 0.134, CL 0.4104. Description of other specimens. Juvenile specimen is amber yellow (83) on the head and pronotum, with a white abdomen and appendages. Comparisons. Grylloblatta marmoreus n. sp. can be distinguished from G. oregonensis n. sp. in having smaller body size, pronotum narrowing at posterior end and forming soft triangular point, and smaller antennomere 2: 1 ratio. It can be distinguished from G. siskiyouensis n. sp. in having fewer antennomeres, buff yellow coloration and weaker point at posterior end of pronotum. It can be distinguished from G. ro t h i in having a subgenital plate without a lobelike projection on distal end and right coxopodite with a strongly down-curved process. It differs from G. chandleri in having a subgenital plate with a more rounded tooth at distal end and right coxopodite with strongly down-curved process. It can be distinguished from G. sculleni in having smaller body size and fewer antennomeres; from G. gurneyi in having a subgenital plate with a single rounded tooth at distal end and right coxopodite with strongly down-curved process; and from G. b a r b e r i in smaller body size and having less than 30 antennomeres. Habitat distribution. The type specimen was found in the twilight zone of a cave habitat, on a rock near ice. The juvenile was found on a rock at the bottom of a pit near the cave entrance. Geographical distribution. Grylloblatta marmoreus n. sp. is known from multiple cave sites in the Marble Valley of Marble Mountains, California (Fig. 3). Phylogenetic relationships. Based on mitochondrial cytochrome oxidase subunit II data, G. marmoreus n. sp. (GenBank Accession JN 612963 & JN 612964) is distinct from (minimum ~ 10 %) other nearby grylloblattid species (Table 1). The adult type specimen and the juvenile from Planetary Dairy Cave differ by 0.4 %. In a neighbor-joining analysis, it is grouped with G. oregonensis n. sp. and G. siskiyouensis n. sp. in an unresolved polytomy (Fig. 4)., Published as part of Schoville, Sean D., 2012, Three new species of Grylloblatta Walker (Insecta: Grylloblattodea: Grylloblattidae), from southern Oregon and northern California, pp. 42-52 in Zootaxa 3412 on pages 49-50, DOI: 10.5281/zenodo.212975

Grylloblatta oregonensis new species (Figs. 1 A, 2 A���B) Type material. Holotype female in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. N 42.098 W- 123.407 WGS 84, Note: in dark zone of cave 15 -VI- 2010 J. Roth SDS 10 -007��� [white label]/ ��� HOLOTYPE Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [red label]; dark zone of cave, Oregon Caves National Monument, 42.098 o North and - 123.407 o West, Josephine County, Oregon, U.S.A. Other specimens. Juvenile in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. 20 -III- 2010. Note: in the dark zone, drowned in a pool of water.��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Juvenile in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. 30 -XI- 2006 ��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Juvenile in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. 11 -VII- 2008 ��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Juvenile in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. 14 -I- 2009 ��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Juvenile in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M.��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Penultimate male in ethanol, deposited in CAS, labeled: ��� USA OR: Josephine Co., Oregon Caves N.M. SDS 10 -003��� [white label]/ ��� Grylloblatta oregonensis Schoville det. S.D. Schoville 2011 ��� [white label]. Etymology. The specific epithet, oregonensis, is named for the state of Oregon in which the type locality, Oregon Caves National Monument is located. Diagnosis. Adult females of this species can be distinguished from all other Nearctic Grylloblatta species by the following combination of character states: color buff yellow on dorsum and white ventrally; large body size (> 1.4cm); long legs; fewer than 30 antennomeres; ratio of antennomere lengths 2 and 1 2.67; broad head; head slightly wider than pronotum; compound eyes large; pronotum square-shaped with flat posterior edge; long, narrow and weakly up-curved ovipositor; dorsal valve of ovipositor noticeably down-curved at tip; ovipositor half as long as cercus; cercomeres long. Holotype specimen with slight damage limited to the juncture of the pronotum and head on the left side (caused during transfer to alcohol). Description of holotype female. Antennomere number on right antennae 28 and on left antennae 26. Compound eyes well developed, black. Interocular distance slightly smaller than pronotum width. Head width less than head length. Head slightly damaged by split in the exoskeleton and swelling tissue in the posterior end. Pronotum square-shaped with rounded corners, surface broad and flat, length greater than width, width at anterior and posterior ends similar, slight inward curve laterally, transverse sulcus at anterior end straight, posterior edge flat. Slightly damaged along anterior and left sides by a split in the exoskeleton and swelling tissue. Legs long and robust, same yellowish color as dorsum with distinctive brown setae. On abdomen, ovipositor half as long as cercus. Ventral valve gradually becoming narrower and curving gently upwards towards distal end, lightly setaceous with short setae. Dorsal valves barely extending beyond ventral valve (Measurements of holotype female (cm). BL 1.905, HL 0.476, HW 0.378, EL 0.103, EW 0.064, ID 0.325, Ant 1 L 0.024, Ant 2 L 0.064, AntRatio 2.67, PL 0.405, PW 0.349, FL front 0.564, FL mid 0.476, FL rear 0.635, TL front 0 445, TL mid 0.476, TL rear 0.714, TrL front 0.119, TrL mid 0.095, TrL rear 0.278, PTrL front 0.476, PTrL mid 0.357, PTrL rear 0.397, OvL 0.516, OvW 0.1588, CL 1.032. Description of other specimens. Juveniles share the characteristic pale dorsal coloration and are sometimes completely white. Comparisons. Grylloblatta oregonensis n. sp. can be distinguished from G. siskiyouensis n. sp. by larger body size, buff yellow dorsal coloration, fewer number of antennomeres and larger antennomere 2: 1 ratio (> 2), longer cercus to ovipositor ratio, longer and slightly tapered ventral valve of ovipositor, and slightly down-curved distal end of the dorsal valve of ovipositor. It can be distinguished from G. m a r m o re u s n. sp. by the large, square pronotum with flat posterior edge, and relatively longer limbs. It can be distinguished from G. ro t h i by its wider head and pronotum, longer limbs, and slightly less up-curved ventral valve of ovipositor. It can be distinguished from G. sculleni by smaller body size, fewer antennomeres, and longer ovipositor. It can be distinguished from G. gurneyi by buff yellow dorsal coloration, fewer antennomeres, and longer ovipositor and cerci. It can be distinguished from G. barberi by smaller body size, buff yellow dorsal coloration, larger pronotum, and slightly tapered ventral valve of ovipositor. It can be distinguished from G. chandleri by having fewer antennomeres, a large, square pronotum with flat posterior edge, and slightly down-curved distal end of the dorsal valve of ovipositor. Habitat distribution. The type specimen was found on the floor in a dark zone of the main cave system, drowned in a pool of water on June 15, 2010. This pool was located approximately 160 meters from the cave entrance and 43 meters below the surface. This cave is surveyed daily and the specimen was found in good condition (vivid coloration, robust exoskeleton), suggesting that it was freshly deceased. One juvenile specimen used for genetic study was found in the dark zone, drowned in a pool of water on March 20, 2010. Four other juveniles and one penultimate male were found at other times of the year, including November 30, 2006, July 11, 2008, and January 14, 2009. This species is a suspected troglophile, as searches in above-ground habitats surrounding the caves and twilight zones within the caves were unsuccessful. Geographical distribution. Grylloblatta oregonensis n. sp. is presently known only from Oregon Caves National Monument (Fig. 3). It may be a limited endemic in this cave system. Phylogenetic relationships. Based on mitochondrial cytochrome oxidase subunit II data, G. oregonensis n. sp. (GenBank Accession JN 612962) is distinct from (minimum ~ 10 %) other nearby grylloblattids (Table 1). No genetic variation was evident in the two samples. In a neighbor-joining analysis, it is grouped with G. siskiyouensis n. sp. and G. marmoreus n. sp. in an unresolved polytomy (Fig. 4)., Published as part of Schoville, Sean D., 2012, Three new species of Grylloblatta Walker (Insecta: Grylloblattodea: Grylloblattidae), from southern Oregon and northern California, pp. 42-52 in Zootaxa 3412 on pages 43-47, DOI: 10.5281/zenodo.212975

The taxon Blattogryllopterida tax.n., whose distinctive character is 'in forewing, M + CuA splitting into MA and MP + CuA' (as opposed to 'into M and CuA'), is erected to make possible a taxonomic assignment of isolated grylloblattidan wings which cannot be conclusively assigned either to the Blattogryllidae or the Plesioblattogryllidae. Material of two blattogryllopteridan species collected from the Daohugou locality (Middle Jurassic; Northeast China) is described. The limited available data on the species Plesioblattogryllus minor Ren & Aristov, 2011 are complemented by descriptions of four wellpreserved specimens, some exhibiting variation in wing venation. Duoduo qianae gen.n. et sp.n. is erected on the basis of an isolated wing. Characters useful for the identification of members of the families Blattogryllidae and Plesioblattogryllidae as well as characters possibly linking these taxa with the extant Grylloblattidae are discussed.

Fragment islands, viewed from the paradigm of island biogeographic theory, depend on continual immigration from continental sources to maintain levels of species diversity, or otherwise undergo a period of relaxation where species diversity declines to a lower equilibrium. Japan is a recently derived fragment island with a rich endemic flora and fauna. These endemic species have been described as paleoendemics, and conversely as recently derived Pleistocene colonists. Geological events in the Miocene period, notably the fragmentation and collision of islands, and the subsequent uplift of mountains in central Japan, provided opportunities for genetic isolation. More recently, cyclical climatic change during the Pliocene and Pleistocene periods led to intermittent land bridge connections to continental Asia. Here we investigate the pattern and timing of diversification in a diverse endemic lineage in order to test whether ongoing migration has sustained species diversity, whether there is evidence of relaxation, and how geological and climatic events are associated with lineage diversification. Using multi-locus genetic data, we test these hypotheses in a poorly dispersing, cold-adapted terrestrial insect lineage (Grylloblattodea: Grylloblattidae) sampled from Japan, Korea, and Russia. In phylogenetic analyses of concatenated data and a species tree approach, we find evidence of three deeply divergent lineages of rock-crawlers in Japan consistent with the pattern of island fragmentation from continental Asia. Tests of lineage diversification rates suggest that relaxation has not occurred and instead endemism has increased in the Japanese Grylloblattidae following mountain-building events in the Miocene. Although the importance of climate change in generating species diversity is a commonly held paradigm in Japanese biogeography, our analyses, including analyses of demographic change and phylogeographic range shifts in putative species, suggests that Pleistocene climatic change has had a limited effect on the diversification of rock-crawlers.

The rectal pads of the primitive insect Grylloblatta compodeiformis (Orthoptera : Grylloblattidae) were studied using light and electron microscopy. In this species, the rectal epithelium is thickened to form 6 prominent rectal pads, each of which is composed of tall columnar epithelial cells and laterally placed slender junctional cells, but is devoid of secondary cells. The rectal pads are interconnected by simple rectal epithelium, and are lined by a thin cuticular intima. They are surrounded by an extensive connective tissue space, which contains bundles of delicate connective tissue fibers, neurosecretory axons, and tracheae and tracheoles, which do not penetrate into the pads. The epithelial cells exhibit extensive infoldings of the apical plasma membranes that are closely associated with mitochondria. The lateral membranes are also highly folded around large mitochondria that possess longitudinally oriented cristae. These membrane folds form mitochondrial-scalariform junctional complexes and enclose intercellular channels and spaces. The apical cytoplasm of the epithelial cells contains numerous coated vesicles, dense tubular elements, multivesicular bodies and lysosomes, which suggests receptor-mediated endocytosis of macromolecules. The presence of large whorls of rough endoplasmic reticulum and abundant free ribosomes in the cytoplasm and nuclei with multiple, well-developed nucleoli indicate that the epithelial cells are actively engaged in protein synthesis. The ultrastructural features were examined in relation to their role in fluid transport in a cold habitat.

Background Climate in alpine habitats has undergone extreme variation during Pliocene and Pleistocene epochs, resulting in repeated expansion and contraction of alpine glaciers. Many cold-adapted alpine species have responded to these climatic changes with long-distance range shifts. These species typically exhibit shallow genetic differentiation over a large geographical area. In contrast, poorly dispersing organisms often form species complexes within mountain ranges, such as the California endemic ice-crawlers (Grylloblattodea: Grylloblattidae: Grylloblatta). The diversification pattern of poorly dispersing species might provide more information on the localized effects of historical climate change, the importance of particular climatic events, as well as the history of dispersal. Here we use multi-locus genetic data to examine the phylogenetic relationships and geographic pattern of diversification in California Grylloblatta. Results Our analysis reveals a pattern of deep genetic subdivision among geographically isolated populations of Grylloblatta in California. Alpine populations diverged from low elevation populations and subsequently diversified. Using a Bayesian relaxed clock model and both uncalibrated and calibrated measurements of time to most recent common ancestor, we reconstruct the temporal diversification of alpine Grylloblatta populations. Based on calibrated relaxed clock estimates, evolutionary diversification of Grylloblatta occurred during the Pliocene-Pleistocene epochs, with an initial dispersal into California during the Pliocene and species diversification in alpine clades during the middle Pleistocene epoch. Conclusions Grylloblatta species exhibit a high degree of genetic subdivision in California with well defined geographic structure. Distinct glacial refugia can be inferred within the Sierra Nevada, corresponding to major, glaciated drainage basins. Low elevation populations are sister to alpine populations, suggesting alpine populations may track expanding glacial ice sheets and diversify as a result of multiple glacial advances. Based on relaxed-clock molecular dating, the temporal diversification of Grylloblatta provides evidence for the role of a climate-driven species pump in alpine species during the Pleistocene epoch.

Publisher Summary Theis chapter discusseses Grylloblattodea, or ice crawlers, as small group of softbodied, apterous, terrestrial, and termite like insects confined to the Northern Hemisphere. The Grylloblattodea comprise a single extant family, Grylloblattidae, which includes only five genera. Grylloblatta occurs in western North America and Canada, where at least 11 species are known. Grylloblattina is known from one species from Russian Far East. It has two known species from Siberia. Galloisiana contains at least 12 species from Japan and east continental Asia. Nambungia has two known species, including the largest-known grylloblattid. Grylloblattids are cryptic, ground-dwelling insects that prefer wet habitats and cool temperatures. They shun light and occur under stones or in dense leaf litter. Species of Grylloblatta that live under rocks under or near snowbanks emerge after dark and feed as scavengers or predators on dead or dying insects that have been blown onto the snow from lower elevations.

The postembryonic development of genital and postgenital segments of Galloisiana nipponensis (Notoptera : Grylloblattidae) have been described in detail. The coxite, stylus, epiproct and paraproct are already observed in the first nymphal instar, but the sexual characters are almost indistinguishable in the 1st to 4th nymphal instars, because the genital segments in these stages are not yet differentiated. In the 5th nymphal instar in the male, the paired primary phallic lobes start to appear at the hind margin of the 9th sternum. The rudiments of ventral valvulae in the female originate at the hind margin of the 8th sternum. In the 6th nymphal instar, the rudiments of inner valvulae begin to appear on the 9th sternum, in addition to the ventral valvula rudiments. The long dorsal valvulae are formed from the extension of nymphal coxite, while the nymphal styli are more reduced than those of younger instars. The ventral valvulae of the 8th abdominal segment are thought to be homologous with the inner valvulae of the 9th segment, judging from their site and mode of development. Further, the inner valvulae may also be homologous with the primary phallic lobes (phallomeres) of the male. The external genitalia in G. nipponensis originate from 2 structures: the dorsal valvulae (considered of appendicular origin), and the phallomeres; ventral and inner valvulae are regarded as mere integumentary outgrowths.

Kim, Byung-Woo, Lee, Woncheol (2006): Redescription of the largest ice bug, Namkungia magnus com. nov. (Grylloblattodea, Grylloblattidae) from Korea. Zootaxa 1359: 57-66, DOI: 10.5281/zenodo.174644

Namkungia magnus (Namkung, 1986) com. nov. (Figures 1���2) Material examined. Holotype. 1 ��, Balgudeok cave, Munung-ri, Jeongseong-gun, Gangwon-do, 4 May 1986, collected by Y.K. Kim. The holotype specimen has been deposited in the Arachnological Institute of Korea. Diagnosis. This species resembles to the N. biyongensis (Namkung, 1974) in having large body length in size (longer than 30 mm), eyeless condition, more than 47 antennomeres, and 12 cercomeres but is distinguished by asymmetrical supra-anal plate of the bulging 10 th abdominal tergite with truncate projection (dull in N. biyongensis), 52 antennomeres in left (48 in N. biyongensis), the cervical sclerite with nine (left) or 11 (right) setae (10 or 9 in N. biyongensis) on lateral margin, and several copulatory sclerites of the genital structure; accessory copulatory sclerite composed of three parts, principal copulatory sclerite composed of two parts surrounded with left phallomeres and 1 st accessory copulatory sclerites, and genital opening situated in the middle part of principal copulatory sclerite. Also, this species is the largest body length (43.3 mm) (38.5 mm in the original description) known and has the greatest 12 cercomeres (10 in the original description) like N. biyongensis. Male: Total body (Fig. 2 A) length 43.3 mm (measured from the anterior margin of rostrum to the posterior margin of tenth abdominal segment), found wandering on ground amongst stones in cave. Head (Figs. 1 A, 2 B) attached obliquely to pronotum. Cranium wider than long (length 6.1 mm, width 6.4 mm), with ten setae on each lateral margin, four setae around the antennal socket; epicranial suture distinct Y-shaped, reaching the circumantennal suture, and a pair of short parietal sutures extend from the occipital foramen over to the vertex. Clypeus distinctly 1.8 times wider than long (length 1.2 mm, width 2.1 mm), projected on its anterior middle part. Anterior tentorial pits sclerotized on their upper lateral margins. Eye absent, composed of the ocular seta. Antennae filiform composed of 49 antennomeres (52 in original description) on left part broken, 16 antennomeres on right broken, the 3 rd segment (0.8 mm) 2.7 times as long as the 2 nd segment (0.3 mm). Labrum 1.5 times wider than long (length 1.1 mm, width 1.7 mm), broadly crescentshaped, covered with many setae on the middle and lower parts. Labial palp 2.6 mm (0.8 + 0.7 + 1.1), with three segments including the longest distal segment, symmetrical, many scattered setae on the 1 st and 2 nd, and absent on the 3 rd. Paraglossa and glossa symmetrical, pointed, with many minute hairs on their apical parts. Maxilla symmetrical, hooked. Maxillary palp, 4.9 mm (0.4 + 0.6 + 1.3 + 1.2 + 1.4), with five segments including the longest distal segment, symmetrical, many scattered setae on the 1 st and 2 nd. Lacinia pointed, with two preapical teeth, a row of sparsely arranged many long setae on its inner margin. Pronotum (Figs. 1 B, 2 A) 1.1 times as long as wide (length 6.2 mm, width 5.7 mm), with six setae on the anterolateral, and clearly vertical suture on the middle. Mesonutum (Figs. 1 C, 2 A) 1.2 times as wide as long (length 4.5 mm, width 5.3 mm), with many scattered hairs on the posterior, slightly concave in the posterior part, with five setae on the anterolateral part, with a long longitudinal suture on the middle. Metanotum (Figs. 1 D, 2 A) 1.2 times as wide as long (length 4.1 mm, width 5.0 mm), distinctly concave in the posterior, with five setae on the middle lateral part, and two median sutures on the anterior part. Cervical sclerites (Fig. 1 J) 1.6 times as long as wide (length 2.3 mm, width 1.4 mm), symmetrical, with nine setae on the lateral margin of left (11 on right), and four small setae on the posterior of right part and five on the left. Basisternum of prothoracic segment 1.1 times as long as wide (length 2.0 mm, width 1.8 mm), symmetrical, triangular expanded on the anterior part, with two long median sutures, with some scattered hairs on the anterior. Katepisterna of prothorax distinctly situated between trochantin and cervical sclerite. Trochantin of prothorax (length 0.86 mm) crescent-shaped, symmetrical, with two setae on the distal part. Basisternum of mesothorax (Fig. 1 K) 1.5 times as wide as long (length 1.8 mm, width 2.7 mm), symmetrical, many scattered setae on anterior part, with two median sutures on the middle originated in the posterior part with distinctly big sternal apophyses. Katepisterna of mesothorax symmetrical, composed in two parts jointed on the lateral margin (upper part length 1.8 mm, width 0.6 mm, with ten setae on the middle and anterior parts; lower part length 1.5 mm, width 0.9 mm, diamond shaped, with two median sutures and one seta on the mid-lateral part). Trochantin of mesothorax circle, with distinct big sternal apophysis and situated between coax of 2 nd leg. Basisternum of metathorax (Fig. 1 L) 2.1 times as wide as long (length 1.4 mm, width 3.0 mm), many scattered setae on its anterior part, with two short median suture on the posterior. Katepisterna of metathorax symmetrical, composed in two parts jointed on the lateral margin part (upper part length 1.8 mm, width 0.9 mm, long rectangular, many scattered setae; lower part length 1.4 mm, width 0.8 mm, diamond shaped, with two median sutures. Trochantin of mesothorax circle, with two sternal apophyses and situated between coax of the 2 nd legs. Legs (Table 1) elongated, leg formula 321, with each two setae on all trochanter, two large spines on the ventral end of the tibia, developed white pulvilli, two setae each on the apical of the 2 nd and 4 th tarsi, many strongly long hairs on all tarsi, and two strong tarsal claws without teeth; 1 st leg 21.8 mm, femur ratio 3.0, some scattered setae on coax, trochanter smaller than others, 25 setae (nine on the dorsal, 16 on the ventral) on femur, ten setae in tibia, 12 in tarsi (six on the 1 st tarsus, two each on the 2 nd and 3 rd); 2 nd leg 23.1 mm, femur ratio 4.2, some setae scattered on coxa, three rows composed of 18 setae (ten on the dorsal, two on the prolateral, six on the ventral) on femur, 20 setae on tibia, 12 on tarsi (six in the 1 st tarsus, two each in the 2 nd and 3 rd); 3 rd leg 29.7 mm, femur ratio 4.7, many scattered setae on coax, three rows composed of 29 setae (seven on the dorsal, four on the prolateral, 20 on the ventral) on femur, 26 setae (12 on the dorsal, five on the prolateral, nine on the ventral) on tibia, 12 on tarsi (six on the 1 st tarsus, two each on the 2 nd and 3 rd). Abdominal tergites (Figs. 1 F, 2 E) slightly convex in the middle, symmetrical, with lateral margin flexed to the posterior, ten-segmented, a pair of median suture on the lateral of 1 st segment to 6 th, supra-anal plate of 10 th segment with truncate projection, one pair of seta on the middle lateral side of 2 nd to 8 th, eight setae on the posterior of the 1 st, and the 3 rd to 8 th, six on the 2 nd and 9 th, four on the 10 th; 1 st tergite 2.8 times as wide as long (length 1.6 mm, width 4.5 mm), 2 nd three times as wide as long (length 1.6 mm, width 4.8 mm), 3 rd 2.8 times as wide as long (length 1.7 mm, width 4.8 mm), 4 th 2.5 times as wide as long (length 2.0 mm, width 5.0 mm), 5 th 2.6 times as wide as long (length 1.9 mm, width 4.9 mm), 6 th 2.5 times as wide as long (length 1.9 mm, width 4.7 mm), 7 th 2.7 times as wide as long (length 1.6 mm, width 4.3 mm), 8 th 2.6 times as wide as long (length 1.4 mm, width 3.6 mm), 9 th 2.1 times as wide as long (length 1.4 mm, width 2.9 mm), 10 th 1.5 times as wide as long (length 1.6 mm, width 2.4 mm). Abdominal sternites (Figs. 1 G, 2 C) slightly convex, symmetrical, with the lateral margin flexed to the posterior part, distinct eversible sac on the mid-posterior margin of the 1 st sternite, two rows (one row composed of eight to twelve setae on the middle part, another composed of 14 to 18 on the posterior margin) on the 3 rd to 7 th sternite; 1 st sternite two times as wide as long (length 1.3 mm, width 2.6 mm), 2 nd about two times as wide as long (length 1.7 mm, width 3.4 mm) with four on the middle and five on the posterior margin, 3 rd 1.9 times as wide as long (length 1.9 mm, width 3.7 mm) with 18 setae on the middle and nine on posterior margin, 4 th 2.1 times as wide as long (length 2.0 mm, width 4.1 mm) with 15 setae on the middle and 11 on the posterior margin, 5 th about two times wider than length (length 2.1 mm, width 4.1 mm) with 15 setae on the middle and nine on the posterior margin, 6 th 1.8 times as wide as long (length 2.2 mm, width 4.0 mm) with 14 setae on the middle and seven on the posterior margin, 7 th about two times as wide as long (length 1.9 mm, width 3.7 mm) with 15 setae on the middle and eight on the posterior margin, 8 th 2.4 times as wide as long (length 1.3 mm, width 3.1 mm) with seven setae on the middle and four on the posterior margin. Ninth abdominal segment (Figs. 1 E, 2 D) indistinctly separated from 9 th abdominal sternite, slightly projected in the middle and with many scattered hairs. Styli twosegmented, with strongly asymmetrical coax, left (length 1.3 mm, width 1.5 mm) about 1.5 times as wide as right (length 1.9 mm, width 1.0 mm) wrapped round process on the mid-prolateral part; distal segment of styli, length about 4.5 times as long as wide (length 0.9 mm, width 0.2 mm), cylindrical, symmetrical, two setae each on the middle and distal parts. Cercomeres (Figs. 1 H���I, 2 D���E) 12 -segmented, some broken on the last segment, symmetrical, cylindrical, with ring patterns of trichobothria (sound sensory organ) and setae on the distal part of segments except for the terminal segment, and the 1 st cercomere without seta; 1 st cercomere (length 0.3 mm) with one ring pattern composed of 15 trichobthria; 2 nd cercomere (length 0.3 mm) shorter than others, with three setae (1 d, 2 v), one ring pattern composed of seven trichobothria; 3 rd cercomere (length 0.3 mm), with one ring pattern composed of five setae (2 p, 1 r, 2 v), one ring pattern composed of nine trichobothria; 4 th cercomere (length 0.6 mm), with eight setae (0-2 d, 0-3 p, 0-2 r, 0- 1 v), one ring pattern composed of four trichobothria; 5 th cercomere (length 0.7 mm), with eight spines (0-2 d, 0-2 p, 0-3 r, 0- 1 v), one ring pattern composed of six trichobothria; 6 th cercomere (length 0.9 mm), with eight setae (0-2 d, 0-2 p, 0-2 r, 0- 2 v), one ring pattern composed of five trichobothria; 7 th cercomere (length 1.1 mm), with eight setae (0-2 d, 0- 2 r, 0-2 r, 0- 2 v), one ring pattern composed of six trichobothria; 8 th cercomere (length 1.3 mm), with seven setae (0-2 d, 0-3 p, 0-1 r, 0- 1 v), one ring pattern composed of six trichobothria; 9 th cercomere (length 1.4 mm), with five setae (0-1 d, 0-2 p, 0-1 r, 0- 1 v), one ring pattern composed of five trichobothria; 10 th cercomere (length 1.5 mm), with four setae (0-1 d, 0-2 p, 0-1 r, 0- 1 v), with six trichobothria; 11 th cercomere (length 1.4 mm), three setae (0- 0-1 d, 0- 0- 2 v), three trichobothria; 12 th cercomere broken in about one thirds. Genital organ (Figs. 1 M���O, 2 F) situated between the ventral parts of the 9 th and the 10 th segment, strong asymmetrical, eversible sac of phallomere, 2.5 times as long as wide (length 2.0 mm, width 0.8 mm), wrinkled cylindrical shape, wound to the upper left cercus and originating from the lateral base of the left phallomere; left phallomere slightly as long as wide (length 1.0 mm, width 0.9 mm), L-shaped with broad base, situated in between eversible sac of phallomere and principle copulatory sclerite and many scattered hairs; right phallomere situated between left phallomere and right stylus coax, semicircular, with round distal part; accessory copulatory sclerite composed of three parts (1 st part reverse ��shaped, 2 nd reverse L-shaped, 3 rd elongated oval-shaped situated in between eversible sac of phallomere and the ventral of the 10 th abdominal segment); principal copulatory sclerite composed of two parts (higher and lower with respect to genital opening) surrounded by left phallomere and the 1 st accessory copulatory sclerite (higher part, with semicircular expanded process, largely depressed in the right; lower part, triangularly concave process in the top); genital opening situated in the middle part of principal copulatory sclerite., Published as part of Kim, Byung-Woo & Lee, Woncheol, 2006, Redescription of the largest ice bug, Namkungia magnus com. nov. (Grylloblattodea, Grylloblattidae) from Korea, pp. 57-66 in Zootaxa 1359 on pages 58-64, DOI: 10.5281/zenodo.174644, {"references":["Namkung, J. (1986) One new species of the genus Galloisiana (Notoptera: Grylloblattidae) from Korea. The Korean Journal of Systematic Zoology, 2 (2), 53 - 58."]}

A new ice bug species, Galloisiana odaesanensis sp. nov., is described from Mt. Odae, Pyeongchang-gun, Gangwon-do, South Korea. This species resembles the G. yuasai group in having a short body (less than 20 mm), distinct compound eyes, fewer than 35 antennomeres, and seven or eight cercomeres. The species is, however, clearly distinguished from the G. yuasai group by the following characters: slightly curved supra-anal plate with dull apical margin; rectangular pronotum that is straight on the posterior; 3rd antennomere 1.7 times as long as the 2nd. The distribution of the genus Galloisiana is discussed and a key for Galloisiana species is presented.

We report the 3.74-Mb genome sequence of Galloisinimonas intestini B14 T , isolated from the gut of one of the world's rarest insect species, Galloisiana sp., collected at a Mosan cave, Moonkyung, Gyungsangbook-do, South Korea. Strain B14 T is a novel genus candidate of the family Enterobacteriaceae .

Grylloblatta oregonensis new species, Grylloblatta siskiyouensis new species, and Grylloblatta marmoreus new species are described from the Klamath Mountains region of Oregon and California, U.S.A. Two species were found at OregonCaves National Monument, with G. oregonensis inhabiting the dark zone of caves and G. siskiyouensis inhabiting surfacehabitat and the twilight zone of caves. The third species, G. marmoreus, is known from Big Foot Cave and Planetary DairyCave, Marble Mountains, California. These three species are recognized on the basis of morphological characters anddistinguished from nearby Grylloblatta species. Analysis of genetic data from the cytochrome oxidase subunit II genesupports the morphological diagnosis of these three species as unique lineages and confirms their genetic divergence from other ice-crawler populations found in Oregon and California.

The eruption of Mount St Helens in 1980 eliminated all biota from an area of ca. 80 km2 to the north of the crater. The recolonization of the mineral surface of the pyroclastic flows by arthropods was observed from 1981 to 1985. Diverse dispersing arthropods reached the site immediately after the eruption. Representatives of at least 150 families from 18 orders arrived in the devastated zone during the first 3 yr following the eruption. Among these immigrants 19 species showed evidence of established breeding populations on the primary mineral surfaces by 1985. Several carabid beetles in the genus Bembidion, notably B. planatum, B. improvidens, and B. obscurellum were the first arthropods known to breed on the pyroclastic flow; this occurred in spring 1981. Beetles predominated among the pioneers with 15 species of Carabidae, and one each of Agyrtidae, and Tenebrionidae. The hemipteran families Lygaeidae and Saldidae were also early pioneers. Pedestrian taxa (Grylloblattidae and Gryllacrididae) arrived on the pyroclastic flow after 3 yr. Primary succession at Mount St. Helens began with the development of an aeolian community of arthropod predators and scavengers, all of which were supported by an allochthonous resource of arthropod fallout.

This paper describes the fine structure of the developing oocytes of the grylloblattodean, Galloisiana nipponensis by transmission and scanning electron microscopy. Particular attention was given to the peripheral ooplasm, the surrounding follicular epithelium and the surface structure of the micropylar area. An ovariole contains a linear array of about 12 developing oocytes in different stages; the anterior 9 oocytes (1–9) show previtellogenic stages, the 10th and 11th oocytes the vitellogenic stages, and the 12th oocyte at the posterior end of the ovariole is fully grown and black. In the previtellogenic stages when oocytes grow in volume, the follicular cells change their shape from flattened to cuboidal, and then to columnar. During the vitellogenic stage, lamellae or whorls of rough endoplasmic reticulum and Golgi bodies develop considerably in the cytoplasm of columnar follicular cells. In the final stage of oogenesis, a thin vitelline membrane (0.4–0.5 μm) and a thick, highly electrondense chorion (9–9.3 μm) with disc-shaped projections are deposited rapidly by the follicular cells that concurrently develop rough endoplasmic reticulum and Golgi bodies. There are 8–10 micropyles (1.1–1.2 μm dia), somewhat circularly distributed around the anterior end of the egg.

After repeated attempts to collect an adult of Grylloblatta barberi Caudell, which was described from immature specimens more than 35 years ago, one was found at the type locality in Plumas County, California. Two new species here described from California ice caves are G. chandleri from Lassen County and G. gurneyi from Modoc County. The distribution of Grylloblatta , especially the Pacific Coast species, coincides closely with areas subject to glaciation during the Wisconsin period. Maps demonstrate this correlation and also show topographic barriers between species groups.

The Grylloblattidae, a small family comprised of only two genera, possibly but one, apparently belong to the order Notoptera. The two species and one variety in the genus Grylloblatta occur in North America, the other genus, Calloisiana, with one species and one variety, is native of Japan. All of the above species are represented by very few specimens and many of these are nymphs. The fact that one species and one variety have been described from nymphs only, together with the unusual scientific interest which this family of insects has aroused due to their primitive structure and peculiar life habits, seems just cause to set out the facts pertaining to the collecting of six adults and numerous nymphs of this species, on Mt. Baker, Wash., in 1931.

A numerical taxonomic analysis was performed using eight exemplars of the orthopteroids. Two suites of attributes were used in a non-weighted dissimilarities analysis. The same suites of attributes were analyzed for a similarity index with dendrograms based on the shortest connection network of Prim. The Tettigoniidae and Gryllidae are in agreement with current systematic placement within the Ensifera. The Caelifera diverge early from ancestral stem lines and suggest a greater separation from the Ensifera within the Orthoptera than generally accepted for their systematic position. The dendrograms suggest that the Blattaria and Mantodea show a relationship closer than the present subordinal rank permits. In the dendrograms the Grylloblattodea and Dermaptera form a remote complex from the other orthopteroids and each qualifies as a separate order. Modern classification of the orthopteroids considers the Dictyoptera, Dermaptera, Orthoptera, and Phasmida as orders. My analysis elevates the Grylloblattidae to the ordinal level as Grylloblattodea.