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."]}