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Zootermopsis angusticollis Soldier caste “Dampwood termite” Figures 118 (lateral), 119 (dorsal), 120 (ventral) Plates 73 (lateral), 74 (dorsal), 75 (ventral) DESCRIPTION: HEAD: Major tracheae indicated in plates, due to networked nature of head tracheae, readers are referred to 3D models in supplemental digital data (see “Availability of Digital Data,” above). H-Oc notably absent, as is expected in termite soldiers. H-DCT with Y-shaped branch meeting in midline as H-DCMedB, which loops posteriad to rejoin H-DCT near origin of Y-shaped branch. Two branches at anterior end of H-DCMedB-Loop: H-Ant anteriad and H-DVB directly ventrad, linking with H-VCT on each side. Close to origin of Y-split, H-DV-Loop extends laterally and anteriad in wide arc, following head capsule lateral wall, looping medially and posteriad near base of antenna, connecting with anterior end of H-VCT as H-DV-Loop; additional H-Ant extends from anterior end of H-DV-Loop. H-VCT runs directly anteriad along floor of head capsule, with large H-DVB connecting to H-DCT and H-DCMedB-Loop. Remainder of H-VCT anteriad toward mouthparts, linking with H-DV-Loop, and branches extending to H-Md, H-MxPlp, and H-Lbm along this loop. H-VLT present, extending into labium. H-VLT asymmetric in soldier caste, see figure 121 for ventral view of worker caste showing symmetric H-VLT branches. THORAX: T 2- S with four branches: H-DCT, H-VCT, T 2-DB, and T 2-VB. H-DCT mediad, curving anteriad toward large head capsule; short T 1- DLT from T 2-DB connecting before curve. H-VCT like H-DCT, with ventrad T 1- AL near curve toward anterior. T 2-DB short, with T 2- AWL branch running posteriad close to T 2- S; T 2-DB continues mediad with Y-shaped bifurcation of short T 1- DLT anteriad toward H-DCT and T 2- DLT posteriad toward T 3- S. T 2- AWL arcing posteriad, splitting in to T 2- AL and T 2-Wbr at apex of arc; T 2-Wbr with small T 2-W-c-r and T 2-W-cu-a, connecting with T 2- PWL posteriorly; T 2- AL posteriad and ventral, joining T 2- PL and extending into midleg. T 2-VB runs directly mediad, bifurcating into T 2-VC1 mediad and T 2- VLT arcing ventrally and posteriad along sternite. T 2-VC1, the first of two mesothoracic ventral commissures, proceeding mediad with T 1- PL branch directly ventrad and smaller connection anteriad to T 1-VX. T 1-VX with H-VCT extending anteriad into head; single branch on specimen left side but likely present on right—see T 2-VC2 branching mediad from T 2- VLT anteriad from T 2-VLT’s curve dorsad toward T 3- S; T 2- VLT with additional T 2-Cx branch, larger on right side than on left. T 3- S with four branches: T 2- PWL, T 3-DB, T 3- AWL, and T 3-VB. T 2- PWL from anterior, connecting T 2- PL and T 2-Wbr from T 2- S. T 3-DB mediad, joining with T 2- DLT and T 3- DLT in flat Y-shaped junction. T 3- AWL mediad briefly, arcing posteriad and dorsally, splitting into T 3- AL ventrad into hindleg and T 3-Wbr posteriad, joining with A 1- S via T 3- PWL. T 3-Wbr with small T 3-W-c-r anterior and T 3-W-cu-a posterior. T 3-VB runs directly ventrad, with T 3- VLT splitting posteriorly just ventrad of T 3- S. T 3-VB continuing ventrad, with T 2-VL branch ventrad and posteriad, extending into midleg; remaining T 3-VB arcing medially to form T 3-VC1; T 2- VLT joining with T 3-VB near this arc. T 3- VLT ventrad, arcing posterior along sternite before continuing anteriad toward A 1- S, with T 3-VC2 mediad near apex of this arc. T 2- VLT and T 3- VLT both with two ventral commissures: posterior T 2-VC2 and anterior T 3-VC1 along same section of T 2- VLT; posterior T 3-VC2 and A 1-VC both along same section of T 3- VLT. ABDOMEN: A 1..8- S present. Short A n -SB possible on several segments. A 1- S with three branches: T 3- PWL, A 1-DB, and A 1-VB; remaining A 2..8- S with just A n -DB and A n -VB branches. For A 1- S, T 3- PWL from anterior, completing T 3-Wbr link from T 3- S. A n -DB runs mediad and slightly dorsad, meeting A n - DLT branches from anterior and posterior in Y-shaped junction. Large A n -DB-Vi typical for all segments, often asymmetric and extending into various abdominal regions and occasionally spanning several segments. A n -DC absent. A n -VB runs ventrad, following body wall, continuing to form A n -VC; A 8-VB split into A 8-VC1 anterior and A 8-VC2 posterior. A n -VB with A n -MLT branching ventrad from A n - S, directly posteriad toward proceeding posterior segment, linking with A n -VB. A 5..8-MLT slightly modified, connecting nearly directly with subsequent A n - S. As with A n -DB, A n -VB-Vi extend throughout abdomen; A 7-VB-Vi and A 8-VB-Vi extend mediad, crossing middle of body and extending anteriad past A 4- S., 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

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

Minorder Artisoptera Engel, nov. This group includes Tanytermitidae and Icoisoptera (= Kalotermitidae + Neoisoptera) (Engel et al., 2009). Species of this group are quite heterogeneous for most characters, with several reversals in Neoisoptera. Nonetheless, in the groundplan the number of cercomeres is reduced, CuP in the forewing scale termites within the scale rather than on the sutural margin (notably reversed several times in the clade), and the imago pronotal lateral margins are typically subparallel. It remains uncertain whether genera such as Dharmatermes Engel et al., 2007 or Baissatermes Engel et al., 2007 should be included within this group., Published as part of JIANG, RI-XIN, ZHANG, HONG-RUI, ELDREDGE, K. TARO, SONG, XIAO-BIN, LI, YAN-DA, TIHELKA, ERIK, HUANG, DI-YING, WANG, SHUO, ENGEL, MICHAEL S. & CAI, CHEN-YANG, 2021, Further evidence of Cretaceous termitophily: Description of new termite hosts of the trichopseniine Cretotrichopsenius (Coleoptera: Staphylinidae), with emendations to the classification of lower termites (Isoptera), pp. 374-389 in Palaeoentomology 4 (4) on page 384, DOI: 10.11646/palaeoentomology.4.4.13, http://zenodo.org/record/5507925, {"references":["Engel, M. S., Grimaldi, D. A. & Krishna, K. (2009) Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance. American Museum Novitates, 3650, 1 - 27. https: // doi. org / 10.1206 / 651.1","Engel, M. S., Grimaldi, D. A. & Krishna, K. (2007) Primitive termites from the Early Cretaceous of Asia (Isoptera). Stuttgarter Beitrage zur Naturkunde, Serie B, Geologie und Palaontologie, 371, 1 - 32."]}

Hodotermopsinae Engel, subfam. nov. Type genus. Hodotermopsis Holmgren, 1911. Diagnosis. Imago right mandible with apical tooth as long as or slightly shorter than first marginal tooth (apical tooth longer than first marginal tooth in Archotermopsinae s. str.). Imago left mandible with apical short, subequal to first marginal tooth (as long as or longer than first marginal tooth, sometimes apical, first marginal, and second marginal teeth equally developed [i. e., Archotermopsis], in Archotermopsinae s. str.), second marginal tooth forming broad cutting edge, with apex projecting to or nearly to level of first marginal tooth (if second marginal tooth forms broad cutting edge in Archotermopsinae [i. e., Zootermopsis Emerson, 1933], then apex of second marginal tooth projecting to half length of first marginal tooth). Soldier with lateral spines only present on metatibia (present on all tibia in Archotermopsinae s. str.); cerci 1.5× or less length of styli (2× or more in Archotermopsinae s. str.). Otherwise typical for Archotermopsidae in presence of cryptically pentamerous tarsi (i. e., second tarsomere reduced and obscured from above giving the false impression of tetramerous tarsi from some angles) and right mandible of image with a subsidiary tooth present at base of anterior margin of first marginal tooth (i. e., between apical tooth and first marginal tooth). While Archotermopsinae tend to have one poorly distinguished inferior branch of Rs apically that parallels the main Rs branch to or just posterior to the wing apex, in Hodotermopsis there may be at times two inferior branches of Rs (Hodotermitidae have more developed, striking straight parallel inferior branches that strike toward the posterior margin (Engel et al., 2009; Krishna et al., 2013). Repeated studies demonstrate the distinctiveness of Hodotermopsis (Bourguignon et al., 2015; Buček et al., 2019). It may be warranted to elevate the rank of this group but given the contradictory placements of several Teletisoptera in recent phylogenetic estimates (Bourguignon et al., 2015; Legendre et al., 2015; Engel et al., 2016; Buček et al., 2019), we are hesitant to make any further refinements to the classification beyond noting the special placement of the genus., Published as part of JIANG, RI-XIN, ZHANG, HONG-RUI, ELDREDGE, K. TARO, SONG, XIAO-BIN, LI, YAN-DA, TIHELKA, ERIK, HUANG, DI-YING, WANG, SHUO, ENGEL, MICHAEL S. & CAI, CHEN-YANG, 2021, Further evidence of Cretaceous termitophily: Description of new termite hosts of the trichopseniine Cretotrichopsenius (Coleoptera: Staphylinidae), with emendations to the classification of lower termites (Isoptera), pp. 374-389 in Palaeoentomology 4 (4) on pages 383-384, DOI: 10.11646/palaeoentomology.4.4.13, http://zenodo.org/record/5507925, {"references":["Holmgren, N. (1911) Termitenstudien: 2. Systematik der Termiten. Die Familien Protermitidae und Mesotermitidae. Kungliga Svenska Vetenskapsakademiens Handlingar, 46 (6), 1 - 86.","Emerson, A. E. (1933) A revision of the genera of fossil and Recent Termopsinae (Isoptera). University of California Publications in Entomology, 6 (6), 165 - 196.","Engel, M. S., Grimaldi, D. A. & Krishna, K. (2009) Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance. American Museum Novitates, 3650, 1 - 27. https: // doi. org / 10.1206 / 651.1","Krishna, K., Grimaldi, D. A., Krishna, V. & Engel, M. S. (2013) Treatise on the Isoptera of the world. Bulletin of the American Museum Natural History, 377, 1 - 2704. https: // doi. org / 10.1206 / 377.1","Bourguignon, T., Lo, N., Cameron, S. L., Sobotnik, J., Hayashi, Y., Shigenobu, S., Watanabe, D., Roisin, Y., Miura, T. & Evans, T. A. (2015) The evolutionary history of termites as inferred from 66 mitochondrial genomes. Molecular Biology and Evolution, 32 (2), 406 - 421. https: // doi. org / 10.1093 / molbev / msu 308","Bucek, A., Sobotnik, J., He, S., Shi, M., McMahon, D. P., Holmes, E. C., Roisin, Y., Lo, N. & Bourguignon, T. (2019) Evolution of termite symbiosis informed by transcriptome-based phylogenies. Current Biology, 29 (21), 3278 - 3734. https: // doi. org / 10.1016 / j. cub. 2019.08.076","Legendre, F., Nel, A., Svenson, G. J., Robillard, T., Pellens, R., Grandcolas, P. (2015) Phylogeny of Dictyoptera: dating the origin of cockroaches, praying mantises and termites with molecular data and controlled fossil evidence. PLoS ONE, 10 (7), e 0130127. https: // doi. org / 10.1371 / journal. pone. 0130127","Engel, M. S., Barden, P., Riccio, M. L. & Grimaldi, D. A. (2016) Morphologically specialized termite castes and advanced sociality in the Early Cretaceous. Current Biology, 26 (4), 522 - 530. https: // doi. org / 10.1016 / j. cub. 2015.12.061"]}