identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038D8781FFD72070FCB2FCBBA151FC5E.text	038D8781FFD72070FCB2FCBBA151FC5E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aeshnidae	<div><p>FAMILY  AESHNIDAE</p><p>“Hawkers” or “Darners”</p><p>The adult  Aeshnidae has many air sacs arranged throughout the body, with many thoracic air spaces likely for muscle growth. Preliminary estimates indicate that over 50% of the volume of the specimen is air, either tracheal or in air sacs (Herhold et al., in prep.). Many major tracheae lead into and out of large air sacs, both in the thorax and abdomen, such that determining tracheal pathways and assessing homology is challenging. Most notably, the abdomen possesses four paired longitudinal trunks, a condition not seen in any other insect in this study. This unusual morphology of the adult relative to other winged insects suggested that some of these features may be holdovers from the aquatic immature stage, so a naiad was collected and scanned.</p><p>To facilitate differentiation of tracheae and tracheated air sacs in the plates and figures for the adult, a subset of the air sacs that serve as tracheal pathways are shown in orange and tubular tracheae are shown in yellow, and the specimen body outline in transparent gray (as in other plates and figures). Although the tracheae are presented as if “on top” of the air sacs, they are located deeper inside the body. The inset image in the plates shows all air spaces (tracheae and air sacs) in gray; readers are encouraged to refer to the 3D models in the supplementary digital data, where air sacs, tracheae, and tracheated air sacs can be viewed separately or together interactively.</p><p>X-ray cross sections from CT scan data indicate the likely presence of flow-directing valves between some air sacs and tracheae in the adult, particularly in the thorax and abdomen (see fig. 29). Air-sac flow-control valves have been documented in  Diptera by Wasserthal et al. (2018), and further research into these structures is needed in  Odonata .</p></div>	https://treatment.plazi.org/id/038D8781FFD72070FCB2FCBBA151FC5E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFA02001FEFCFAF8A432FC7A.text	038D8781FFA02001FEFCFAF8A432FC7A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Anisolabis maritima (Bonelli 1832)	<div><p>Anisolabis maritima</p><p>“Maritime earwig”</p><p>Figures 44 (lateral), 45 (dorsal, ventral)</p><p>Plates 24 (lateral), 25 (dorsal, ventral)</p><p>A greater number of smaller tracheae are visible in the  Anisolabis scan than the  Forficula scan. As the CT scans for this study specifically targeted tracheal morphology, visualization of internal morphology of organs and internal structures was generally not possible. Therefore, determining the specific tissues supplied by a given trachea is challenging. However, some structures appear to have a particular tracheal morphology, such as muscle fibers and some other structures, and the detail afforded in the  Anisolabis scan allows for some possibilities. For example, A3-VLT-Vi begins by branching from A3-VLT and proceeding anteriorly. Just before reaching the anterior end of the abdomen; however, A3-VLT-Vi splits into several smaller tracheae that turn posteriorly to form a cone, most likely the tracheation of a portion of the alimentary canal, possibly the proventriculus. Additional detail is observable in the terminalia, specifically the cerci. As mentioned previously, extensive tracheation of A7 and A8 is for musculature controlling the forceps, and two tracheae are seen extending into the cerci. A8-DLT-Cr is supplied by A8-DLT, which arcs ventrad and posteriorly into each side of the forceps, and a second trachea A8-VLT-Cr from A8-VLT, reaching basically straight posteriad to cross over A8-DLT-Cr into the cerci.</p><p>DESCRIPTION: HEAD: H-DCT and H-VCT similar diameter; H-DCTs curve inward slightly such that left and right tracheae nearly touch before turning laterally outward on entry to head capsule; H-VCTs both proceed straight into head. H-DCT with several branches just anterior of cervix: H-DCC, H-DCT-Dvi, H-Oc, and H-Ant. H-DCC present, with several small H-DCC-Dvi fanning laterally and anteriad along head capsule. H-DCT-Dvi runs laterally and dorsad, branching off H-Oc. H-Oc arcing laterally, with several small tracheae extending into eye, then continuing anteriorly and ventrad via H-Oc-Md to link with ventral H-Md; H-Oc-Ant branches off H-Oc into antenna. H-Ant extends anteriorly through head into antenna; H-Ant with multiple tracheae in  Anisolabis (H-Md-Ant, below). H-VCT likewise with several branches just anterior of cervix: H-VCT-Vi, H-Ft-Lbr, H-VC, and H-Mx-Md. H-VCT-Vi laterally and dorsad, like dorsal trachea. H-Ft-Lbr anteriad, with H-VC extending medially to link left and right sides; H-Ft branching laterally and anteriad with H-Lbr running directly anteriad. Both H-VC with H-VC-Dvi running directly anteriad. H-Ft-Lbr runs anteriad, with small H-Ft splitting laterally and anteriad, remaining H-Lbr running anteriad. H-Mx-Md runs anteriad and slightly laterally, with several branches: H-Lbm run ventrad, with short H-LbmPlp; H-Mx running ventrad, with short H-MxPlp. Remaining H-Md branch runs anteriad, with H-Oc-Md connection from H-DCT; H-Md-Ant branching dorsally to parallel H-Ant from H-DCT.</p><p>THORAX: T1-DVi branching from H-DCT closer to T2-S than in  Forficula . T1-MVi visible. Several T1-Fm visible. T3-VL with connection to VLT near A1-VB connection, forming X-shaped chiasma.</p><p>ABDOMEN: A4-DLT-Mvi branching from interior side of A4-DLT, slightly posterior to junction with A4-DB on right side of the body but nearly at Y-shaped junction on the left side. Branching of A n -DLT-Mvi from interior side of the dorsal longitudinal trunk varies between left and right side, but general anterior-posterior direction of tracheae is consistent for each segment (see fig. 46). A4-VLT-Vi anteriad, extending through nearly two body segments. Anterior direction of A n - VLT-Vi consistent for  Anisolabis . A8-VLT-Cr and A8-DLT-Cr extending into forceps.</p></div>	https://treatment.plazi.org/id/038D8781FFA02001FEFCFAF8A432FC7A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFC82068FCA9FAE7A413FFFB.text	038D8781FFC82068FCA9FAE7A413FFFB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Baetidae Leach 1815	<div><p>FAMILY  BAETIDAE</p><p>Two baetids were scanned to investigate developmental changes. These are two separate specimens, not the same specimen scanned twice, as done in beetles by Lehmann et al. (2021). Tracheal morphology between subimago and adult is strikingly consistent, apart from the single alimentary air sac in the adult (vs. divided in subimago), indicating that tracheal development is essentially complete by the subimago stage.</p></div>	https://treatment.plazi.org/id/038D8781FFC82068FCA9FAE7A413FFFB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6920C9FC56FE8CA48CFAF1.text	038D8781FF6920C9FC56FE8CA48CFAF1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Blaptica dubia (Serville 1838)	<div><p>Blaptica dubia</p><p>“Argentinian wood roach”</p><p>Figures 111 (lateral), 112 (dorsal), 113 (ventral)</p><p>Plates 67 (lateral), 68 (dorsal), 69 (ventral)</p><p>The tracheal architecture of B. dubia features wide, bandlike tra- cheae and numerous visceral branches throughout the body; pre- liminary results indicate that tra- cheae may comprise nearly 12% of the volume of the insect (Herhold et al., in prep.). Spiracles and a handful of major tracheae are labeled in the figures and plates; readers are encouraged to view the 3D models provided in the supplementary digi- tal data for further investigation.</p></div>	https://treatment.plazi.org/id/038D8781FF6920C9FC56FE8CA48CFAF1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6120C3FCB3FA06A483FB75.text	038D8781FF6120C3FCB3FA06A483FB75.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Blattodea	<div><p>ORDER  BLATTODEA</p><p>Three roaches were scanned during the study: the common American cockroach  Periplaneta americana, the popular Madagascar hissing cockroach  Gromphadorhina portentosa, and  Blaptica dubia . The tracheal system of  Gromphadorhina has been studied via dissection to determine the physiology of its hissing behavior (Nelson, 1979; Nelson and Fraser, 1980) and was micro-CT scanned in a demonstration of semiautomated segmentation techniques (Lösel et al., 2020). Active tracheal compres- sion in Blattodea was studied via synchro- tron imaging by Westneat et al. (2003).</p><p>Periplaneta is described here in detail. As with other complex specimens, the other two roaches, Gromphadorhina and Blaptica, are only briefly described. Readers are directed to the labeled 3D models in the supplemen- tary digital data for further investigation.</p></div>	https://treatment.plazi.org/id/038D8781FF6120C3FCB3FA06A483FB75	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFD12079FCBFFB0EA11DFD1E.text	038D8781FFD12079FCBFFB0EA11DFD1E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Calopteryx maculata Palisot de Beauvois 1807	<div><p>Calopteryx maculata</p><p>“Ebony jewelwing”</p><p>Figures 38, 39 (lateral, anterior, posterior), 40 (dorsal), 41 (ventral)</p><p>Plates 21 (lateral), 22 (dorsal), 23 (ventral)</p><p>A single damselfly was scanned at 19 µm, which should have been sufficient to capture details of abdominal tracheae; however, this specimen was frozen to -20° C early in the study. As small tracheae were likely infilled, this specimen is not described in detail, but three-dimensional models are included in the online supplementary digital data.</p><p>The thoracic tracheae appear to be very similar to the aeshnid, along with extensive air sacs.</p><p>62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 459</p><p>It is unclear whether there are four paired abdominal trunks as in the dragonfly, but it seems likely. Further studies should focus on Zygoptera in addition to more dragonfly specimens to verify and solidify tracheal patterns in the flight motor and abdomen.</p></div>	https://treatment.plazi.org/id/038D8781FFD12079FCBFFB0EA11DFD1E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFD92000FCC6FACEA178FDCA.text	038D8781FFD92000FCC6FACEA178FDCA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dermaptera	<div><p>ORDER  DERMAPTERA</p><p>Dermaptera have been the subject of recent micro-CT studies, including the discovery of active tracheal compression via synchrotron imaging (Westneat et al., 2003) as well as a detailed analysis of the head of  Forficula auricularia (Neubert et al., 2017) .  Dermaptera phylogeny is considered to be generally unresolved, with distinctions between lower  Dermaptera and higher  Dermaptera considered somewhat arbitrary (Haas, 2018). Recent transcriptomic and morphological studies have refined relationships within  Dermaptera (Wipfler et al., 2020); however, these analyses omit the fossil record (Tihelka et al., in prep). The taxa here consist of one “lower” dermapteran ( Anisolabis maritima,  Anisolabidae, female) and one “higher” ( Forficula auricularia,  Forficulidae, male), and although  Anisolabis lacks wings, they are alike in tracheal architecture. Sex was determined for both species based on sexual dimorphism of the cerci. While the specimens were scanned at similar resolutions, 11.6 mm 3 /voxel for  Anisolabis and 13.5 mm 3 /voxel for  Forficula (see table 3 for all scan parameters), substantially more visceral tracheae are visible in the  Anisolabis scan. This may be the result of slightly higher resolution, but also possibly due to different scanning parameters or preservation artifacts (fluid filling of small tracheae postmortem). Regardless, sufficient detail is present in both specimens to assess tracheal homology.</p><p>The two dermapteran specimens are similar in their overall tracheal layout, with most differences in the heads. Wing base tracheae T2,3-Wbr are well-developed in  Anisolabis, even though apterous; however, wing tracheae T2,3-W-c-r and T2,3-Wcu-a are absent. A1-S is positioned dorsally in both specimens, unlike the remaining abdominal spiracles, possibly a modification for the ability of earwigs to raise the abdomen dorsally and forward to use the forceps for predation and defense. Additionally, the A n -VLT-Vi in  Anisolabis all proceed anteriorly, whereas in  Forficula, there appears to be a “split” where A n -VLT proceeds anteriorly for A2..5-VLT-Vi..5, but posteriorly for A6..7-VLT-Vi.</p><p>DESCRIPTION: HEAD: Majority of tracheal structures similar between  Anisolabis and  Forficula, differences described here for comparative clarity. H-DCT and H-VCT of similar diameter; H-DCTs curve inward slightly such that left and right tracheae nearly touch before turning laterally outward on entry to head capsule; H-VCT proceeds straight into head. H-DCT with several branches just anterior of cervix: H-DCC, H-DCT-DVi, H-Oc, and H-Ant. H-DCC present. H-DCT-Dvi running laterally and dorsad. H-Oc arcing laterally, with several small tracheae extending into eye, then continuing anteriorly and ventrad via H-Oc-Md to link with ventral H-Md; H-Oc-Ant branches off H-Oc in  Anisolabis into antenna. H-Ant extends anteriorly through head into antenna; left side of  Forficula as H-Ant-Ft with H-Ft branching off H-Ant near base of antenna. H-Ant with multiple tracheae in  Anisolabis; likely present in  Forficula but not visible in this scan. H-VCT likewise with several branches just anterior of cervix: H-VCT-Vi, H-Ft-Lbr (absent in  Forficula), H-VC (off H-Ft-Lbr in  Anisolabis), and H-Mx-Md. H-VCT-Vi runs laterally and ventrad, like dorsal trachea. In  Anisolabis, H-Ft-Lbr runs anteriad, with H-VC extending medially to link left and right sides; H-Ft branching laterally and anteriad with H-Lbr directly anteriad. In  Forficula, H-VC branching directly from H-VCT. Both H-VC with H-VC-Dvi running directly anteriad, extending as far as frontal area in  Forficula . H-Ft-Lbr (absent in  Forficula) running anteriad, with small H-Ft splitting laterally and anteriad, remaining H-Lbr anteriad. H-Mx-Md runs anteriad and slightly laterally, with several branches: H-Lbm running ventrad, with short H-LbmPlp; H-Mx ventrad, with short H-MxPlp. Remaining H-Md branch runs anteriad, with H-Oc-Md connection from H-DCT; H-Md-Ant branching dorsally (absent in  Forficula left side) to join H-Ant from H-DCT.</p><p>THORAX: Although the thoraces of  Anisolabis and  Forficula differ substantially in overall exterior morphology, as seen in figure 43, the tracheal topology of the thorax is retained between the two taxa. T2-S with four tracheae: H-DCT, H-VCT, T2-AWL, T2-DB; T2-CT absent. H-DCT arcs mediad before proceeding anteriorly; T1-Dvi branching close to T2-S, extending anteriad with several smaller branches; T1-DC present. H-VCT similar to H-DCT, arcing medially then anteriad toward head. T1-AWL branching dorsoventrally; T1-VC present, separating from T1-AL while remainder of T1-AL continues into foreleg. T2-AWL extending posteriad and slightly dorsad, bifurcating into T2-AL and T2-Wbr; T2-AL continues ventrad and posteriorly into midleg; T2-Wbr running dorsad and laterally, connecting directly with T2-S. T2-DB extending medially, ventrad and slightly anteriad, with three branches: T1-PL, T2-DLT, T2-VT. T1-PL extending through coxae before joining with T1-AL and continuing into proleg; T1-Fm visible. T2-DLT proceeds anteriorly to connect directly with T3-S via T3-DB. T2-VLT ventrad and posteriad, following mesothoracic sternite before arcing anteriad to link with T3-S. T3-S with four branches: T2-Wbr, T3-AWL, T3-DB, T3-VB. T2-Wbr extends from T2-S linking directly to T3-S. T3-AWL extending slightly dorsad before turning ventrad and laterally, bifurcating into T3-AL and T3-Wbr; T3-AL continues posteriorly into hindleg; T3-Wbr extending posteriad to connect directly to T3-S. T3-DB running mediad, linking with T2-DLT from anterior and T3-DLT continuing posteriorly. T3-VB runs mediad and posteriorly, with T3-VLT branching posteriad; T3-VB subsequently bifurcates into T2-PL and T2-VLT connection from T2-S. T2-PL runs anteriad from T3-S, joining with T2-AL from T2-S, continuing into midleg; several T2-Fm present. T3-VLT with T3-VL into hindleg femur.</p><p>ABDOMEN: A1..8-S present. A1-S highly modified from subsequent segments, placed dorsally with four branches: T3-Wbr, A1-DB, A1-VB, T3-PL. T3-Wbr connecting directly from T3-S; small T3-Wbr-Vi extends dorsad and medially along metathoracic tergite. A1-DB mediad, linking with T3-DLT from anterior and A1-DLT continuing posteriad. A1-VB ventrad, with A1-VLT branching directly posteriad; A1-VB continues ventrally, bifurcating into T3-Fm, extending laterally into hind leg, and A1-VC; T3-VLT connects with A1-VC. T3-PL runs ventrad and posteriad, joining with T3-AL before extending into hindleg. Tracheae from A2..6-S similar, A4-S described here as example. A4-S with three branches: A3-VLT, A4-DB, and A4-VLT. A3-VLT runs anteriad, extending from A3-S. A4-VLT branches medially and dorsad before curving posteriad and laterally toward A5.S. At apex of this arc A4-VLT- Vi extends anteriorly, spanning several segments. A n -VLT-Vi variable, see taxon descriptions below. Midway between A4-S and A5-S, A4-VC branches ventrad at right angle, following abdominal sternite. A4-DB directly dorsad; A3-DLT connecting from anterior and A4-DLT from posterior to form Y-shaped junction. Visceral tracheae extend from either end of A4-DB: A4-DB-Mvi extends laterally and dorsad from start (ventral end) of A4-DB; A4-DB-Dvi from base of Y-shaped junction with A3-DLT and A4-DLT. A4-DLT-Dvi present but A4-DC absent. A n -DLT-Vi numerous and highly variable, see descriptions below. A7-S connections like previous segments but distance between A7-S and A8-S greatly shortened. A8-DB with three tracheae: A7-VLT, A8-DB, and A8-VLT, but with posterior branching varying between taxa, see descriptions below. Extensive tracheation in A7 and A 8 in both genera, no doubt for large muscles that control the forceps.</p></div>	https://treatment.plazi.org/id/038D8781FFD92000FCC6FACEA178FDCA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFCC2068FF11FBD1A3F6FB75.text	038D8781FFCC2068FF11FBD1A3F6FB75.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ephemeroptera Hyatt & Arms 1891	<div><p>ORDER  EPHEMEROPTERA</p><p>Ephemeroptera are unique in having a winged subimago developmental stage; all other pterygotes have lost this molt as winged individuals. Additionally (and relevant to tracheation),  Ephemeroptera, like  Odonata, possess direct flight muscles, where the wing muscles insert directly at the wing bases. Mayfly tracheae were first described in detail by Swammerdam (1737; see fig. 1), and modern treatments include Needham et al. (1935), Landa (1948, 1969), and Soldán (1979). Chapman (1918) included a discussion of wing tracheae of mayflies in his early comparative analysis.</p><p>For this study, three mayfly specimens were scanned: an adult  Ephemera sp. (Ephemeridae) and  Neocloeon triangulifer subimago and adult ( Baetidae). The  Neocloeon specimens afforded the opportunity to compare tracheal architecture between the winged stages. Although scan resolution ranged from 13 µm for  Ephemera sp. to 2 µm for the adult  Neocloeon, sufficient detail was achieved to determine that all three specimens of both taxa are remarkably similar in tracheal architecture, the primary difference being the morphology of the air sac. Adult  Ephemeroptera do not feed, and the alimentary canal forms a large, central air sac (Soldán, 1979), perhaps aiding the emergence of the alate subimago from the water, with visceral trachea either reduced or displaced. While not complete until the adult stage, the subimago possesses smaller but substantial air spaces that fuse to form the single adult air sac (see fig. 21). As air sacs in tracheal visualizations can obscure details of smaller branches, all air sacs are omitted from the  Ephemeroptera plates. Due to the size of the wings,  Ephemeroptera plates are shown with the wings cropped to show as much detail as possible without scaling the image to fit the page. Other minor differences between specimens are described in their respective sections below.</p><p>T2-S is the start point of the longitudinal trunk, which arcs toward the midline of the body for a short distance before turning posteriad and proceeding all the way to the terminalia. While similar structures have been noted in the larvae of many insect orders (Snodgrass, 1935; Whitten, 1960; Wigglesworth, 1972), the presence of a large, nearly linear longitudinal trunk in an adult that spans the length of the body appears to be unique to  Ephemeroptera . This structure is likely homologous with the dorsal longitudinal trunk seen in other taxa, and individual segments of the trachea are referred as such, beginning with T2-DLT as the start. The thoracic portion of the DLT in  Ephemera appears to be two tracheae, arranged dorsoventrally. This is the result of compression in the thorax—distention of the air-filled alimentary canal, compression by packing material for CT scanning, or postmortem settling of tissues can all contribute to deformation of tracheae in soft-bodied insects such as mayflies.</p><p>Noted by Chapman (1918),  Ephemeroptera possess only anterior wing base tracheae (T2,3- AWba), extending into the wing as T2,3-W-c-r with the corresponding PWba and W-cu-a tracheae absent. This lack of a branch leading to the trailing-edge wing tracheae is a condition apparently unique to mayflies.</p><p>The three ephemeropteran specimens display some disparity in the origins of H-Oc branches. These appear to be developmental variations, as larger trunks may supply different regions as smaller tracheae “sprout” from them during ontogeny.</p><p>It remains unclear whether all abdominal spiracles are functional. Needham et al. (1935) indicates that abdominal spiracles may be continuously open, but close examination of the spiracular openings via CT cross section suggests that A2-S through A7-S may be closed or unused.</p><p>DESCRIPTION: BODY: Adults with long central air sac formed from modified alimentary canal, extending from head capsule to end of 7th abdominal segment; subimago with individual air sacs that fuse in the imago (see fig. 21).</p><p>HEAD: Tracheal morphology largely from Landa (1948), but with unified nomenclature here. Palmen’s organ (H-PO) prominent and centrally located. Just after entry into head capsule, thick H-DCT bifurcates into two tracheae; dorsal one of these proceeds anteriad a short distance before dividing into H-Ocel and dorsal branch to H-PO; ventrally H-Lbr branches anteriad shortly with H-DOc branch before remaining trachea turns directly ventrad; H-Lbr continues toward reduced mouthparts with small H-Ant branch at base of antennae. H-VCT thick, splits into three branches at base of head capsule: H-VOc, extending laterally toward eye; H-Lbm directly ventrad, and ventrally supplying H-PO.</p><p>THORAX: T2-S with three branches: T2-CT, T2-SAtr, and T2-DLT. T2-CT very short, before bifurcating into anterior H-DCT and H-VCT at posterior margin of prothorax. T1-L branching directly ventrad from T2-VCT anterior of H-DCT/H-VCT split from T2-CT. T2-SAtr dorsal, with T2-AWL posteriad and multiple T2-FM dorsad: thick T2-FM1 splitting dorsally and subdividing into smaller tracheae that supply flight muscles, thinner T2-FM2 running dorsad and slightly posteriad; T2-AWL arcing briefly dorsad and posteriad before proceeding ventrad into midleg as T2-L, with T2-AWba branching near dorsal apex of arc and continuing to T2-W-c-r. T2-PWba absent. T2-DLT very thick, beginning mediad before arcing directly posteriad and continuing to the terminalia. Small T2-FM3 extends ventrally from T2-DLT midway between T2-S and T3-S. T3-S opens into very thick T3-SAtr, with (effectively) five branches: T3-DB, T3-AWL, and three T3-FM. T3-DB runs directly mediad, linking with T2-DLT from anterior, continuing directly posteriad as T3-DLT. T3-AWL arcing briefly dorsad and then posteriad, similar to T2-AWL, with T3-Wba branching near apex of arc while T3-L continues into hindleg; T3-Wba continues as T3-W-c-r into wing. T3-PWba absent. T3-FM1 very thick, extends directly dorsad and T2-FM2 posteriad and slightly ventrad into flight muscle; T3-FM3 runs ventrad in fanlike extension into flight muscle.</p><p>ABDOMEN: A1-S, A8-S functional; unclear if A2..7-S connect through body wall. A1-S modified, with large A1-FM dorsally; T3-DLT from anterior, continuing posteriad as A1-DLT. Remaining abdominal segments similar, with A n -S linked with neighboring segments via A n - DLT extending linearly through abdomen past A8-S. A8-DLT extending into terminalia as A-Cr. All A n -S with various visceral tracheae extending dorsad and posteriad from A n -S, dorsal and ventral commissures absent.</p></div>	https://treatment.plazi.org/id/038D8781FFCC2068FF11FBD1A3F6FB75	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF9C203CFEF9FB5CA374F8D0.text	038D8781FF9C203CFEF9FB5CA374F8D0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Extatosoma tiaratum (W. S. Macleay 1826)	<div><p>Extatosoma tiaratum</p><p>“Spiny leaf insect”</p><p>Figures 85, 86 (lateral, anterior, posterior); 87, 88 (dorsal, anterior, posterior); 89, 90 (ventral, anterior, posterior)</p><p>Plates 52 (lateral), 53 (dorsal), 54 (ventral)</p><p>The spiny leaf insect  Extatosoma tiaratum features a highly networked tracheal system with many lateral commissures and longitudinal connections. The posterior portion of the abdomen features a dense packing of noodlelike visceral tracheae that has morphological similarities to a package of dried ramen noodles, likely important for ventilation of the ovaries during the con- tinuous egg production of adult females. A n -DLT and A n -VLT are discernible in regions, along with larger anatomical features such as the location of spiracles, but the complexity of the specimen calls for a more in-depth analysis than can be presented here. Interested readers are encouraged to view the 3D models in the supplementary digital data.</p></div>	https://treatment.plazi.org/id/038D8781FF9C203CFEF9FB5CA374F8D0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFA12008FCBEFA28A146FF31.text	038D8781FFA12008FCBEFA28A146FF31.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Forficula auricularia	<div><p>Forficula auricularia</p><p>“Common earwig”</p><p>Figures 47 (lateral), 48 (dorsal), 49 (ventral)</p><p>Plates 26 (lateral), 27 (dorsal), 28 (ventral)</p><p>DESCRIPTION: HEAD: H-DCT and H-VCT thick, of similar diameter; H-DCTs curve inward slightly such that left and right tracheae nearly touch before turning laterally outward on entry to head capsule; H-VCTs both proceed straight into head. H-DCT with several branches just anterior of cervix: H-DCC, H-DCT-Dvi, H-Oc, and H-Ant. H-DCC present. H-DCT-Dvi running laterally and dorsad. H-Oc arcing laterally, with several small tracheae extending into eye, then continuing anteriorly and ventrad via H-Oc-Md to link with ventral H-Md. H-Ant extends anteriorly through head into antenna; left side of  Forficula as H-Ant-Ft with H-Ft branching off H-Ant near base of antenna; H-Ft branches off H-VCTsourced H-Md on right side. H-Ant with multiple tracheae in  Anisolabis; likely present in  Forficula but not visible in this scan. H-VCT likewise with several branches just anterior of cervix: H-VCTVi, H-VC, and H-Mx-Md. H-VCT-Vi running laterally and ventrad, like dorsal trachea. H-VC branches directly from H-VCT. Both H-VC with H-VC-Dvi run directly anteriad, extending as far as frontal area. H-Ft-Lbr absent. H-Mx-Md running anteriad and slightly laterally, with several branches: H-Lbm runs ventrad, with short H-LbmPlp; H-Mx running ventrad, with short H-MxPlp. Remaining H-Md branch runs anteriad, with H-Oc-Md connection from H-DCT; H-Md-Ant branching dorsally on right side to join H-Ant from H-DCT; H-Ft branching from H-Md on right side.</p><p>THORAX: Distance between T2-S and T3-S much shorter in  F. auricularia (See fig. 43).</p><p>ABDOMEN: Abdominal tracheae of  F. auricularia consistent with description of overall dermapteran tracheae above except for direction of visceral tracheae. A6..7-VLT-Vi similar to their anterior counterparts, except A6..7-VLT-Vi extend posteriorly rather than anteriorly, and do not extend beyond segment boundaries; A5-A6 appears to be dividing line for visceral tracheae from median longitudinal trunk, where A2..5- VLT-Vi tracheae extend anteriorly and A6..7- VLT-Vi extend posteriorly. A2..7-VLT-Vi not bilaterally symmetric; right side tracheae larger in cross-sectional area than left. Right side tracheae, while extending into body, also do not supply exact same areas/organs as those from left side. A3-DLT-Mvi and A4-DLT-Mvi on left side extend directly toward the posterior, with A3-DLT-Mvi ending near A7-DB. A3-DLT-Mvi on right side much reduced from left side counterpart, extending posteriorly as far as A4. A4-DLT-Mvi highly modified; after extending toward the middle of the body, continues well into A8 and A9, where it turns toward body wall in large arc that results in trachea pointing anteriorly. A4-DLT-Mvi with similar loop on right as on left, but much shorter, with apex of loop reaching as far as A5-DB. Extensive branching/tracheation in A7 and A8 for cercal (forceps) muscles.</p></div>	https://treatment.plazi.org/id/038D8781FFA12008FCBEFA28A146FF31	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6920CAFC98FC64A1E9F8AE.text	038D8781FF6920CAFC98FC64A1E9F8AE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gromphadorinha portentosa	<div><p>Gromphadorhina portentosa</p><p>“Madagascar hissing cockroach”</p><p>Figures 114 (lateral), 115 (dorsal), 116 (ventral)</p><p>Plates 70 (lateral), 71 (dorsal), 72 (ventral)</p><p>Easily the most popular cock- roach (as far as cockroach popular- ity goes), the physiology of the hissing behavior of G. portentosa was investigated by Nelson and col- leagues (Nelson, 1979; Nelson and Fraser, 1980), with the notable find- ing that the fourth spiracle (A2-S) is the source of the hiss. Heinrich et al. (2013) documented unidirectional airflow in G. portentosa resulting from spiracular valve control. As seen in the plates, the tracheal archi- tecture of Gromphadorhina is remarkably complex, so only spiracles and major tracheae are labeled here. As is common in reared  Gromphadorhina colonies, the individual scanned was infested with mites (see fig. 117), likely  Gromphadorholaelaps schaeferi, found to be possibly beneficial by Yoder et al. (2012).</p></div>	https://treatment.plazi.org/id/038D8781FF6920CAFC98FC64A1E9F8AE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6120C1FD1CFCE8A384FC6F.text	038D8781FF6120C1FD1CFCE8A384FC6F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Idolomantis diabolica (Saussure 1869)	<div><p>Idolomantis diabolica late-stage instar</p><p>“Devil’s flower mantis”</p><p>Figures 104 (lateral), 105 (dorsal), 106 (ventral)</p><p>Plates 61 (lateral), 62 (dorsal), 63 (ventral)</p><p>The tracheal morphology of  Idolomantis is very similar to  Tenodera, in particular the elongate pronotum and the dorsal-ventral switch of H-DCT and H-VCT. The curling of the abdomen over the thorax unfortunately does not lend itself well to diagramming in two dimensions; major tracheae are labeled in the figures and plates, but readers are encouraged to explore the supplementary digital data.</p></div>	https://treatment.plazi.org/id/038D8781FF6120C1FD1CFCE8A384FC6F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6A20CAFF0DFE45A1C7FC6F.text	038D8781FF6A20CAFF0DFE45A1C7FC6F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Isoptera Brulle 1832	<div><p>INFRAORDER  ISOPTERA</p><p>Although the respiratory physiology of termites (especially as it relates to their digestive system) has been studied for decades, micro-CT scans of respiratory architectures have been absent. The tracheal scans of the reproductive caste  Reticulatermes flavipes and soldier caster  Zootermopsis angusticollis shown here detail the remarkable intricacy of tracheal branches, especially in the abdomen, well known for its adaptations for digesting cellulose. Although both specimens were scanned at approximately 5 µm resolution, the worker  Reticulitermes possesses notably smaller tracheae than the soldier  Zootermopsis . Although the two specimens feature many similarities, notably a T1-DLT prothoracic “loop,” the differences are sufficient to describe both in detail.</p><p>Wing-leg branching patterns, referred to here as Chapman’s Triangle (see Discussion section), are evident in both termites, albeit with some minor modifications (Chapman, 1918). In  Zootermopsis, T2-AWL branches from T2-DB rather than directly from T2-S (as in  Dermaptera and others), T3-AWL is off T3-S. In  Reticulitermes, T2-VL branches from T2-VB, rather than T2-VLT, and T3-VL branches off T3-VB. (These could be Tn-Cx and not Tn-VL.)</p></div>	https://treatment.plazi.org/id/038D8781FF6A20CAFF0DFE45A1C7FC6F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6420C0FCB2FE66A126FA3A.text	038D8781FF6420C0FCB2FE66A126FA3A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Mantodea	<div><p>ORDER  MANTODEA</p><p>The tracheal system of mantises in part has been examined in detail since the discovery of the ventrally located “cyclopean” ear by Yager and Hoy (1986). More recent investigations have used micro-CT based methods (Wipfler et al., 2012); however, a complete mapping of mantis respiratory systems has been lacking. Two specimens are included here: the common Chinese mantis  Tenodera sinensis and the popular “Devil’s flower mantis,”  Idolomantis diabolica .  Tenodera is described here in detail but the somewhat more complex  Idolomantis is covered only broadly; 3D models are included in the supplementary digital data for further research.</p><p>The morphology of H-DCT and H-VCT deserves special note here. Nominally, branching patterns starting at spiracle are judged to be more stable, as spiracles are used as the landmarks for assessing homology (as discussed above). The tissues supplied in the mantis prothorax and head, however, indicate a dorsal/ventral swap of H-DCT and H-VCT. For H-VCT in all other taxa, T1-AL branches ventrally from H-VCT in the prothorax. In  Mantodea, however, this branch (H-VCT) begins dorsally at T2-S, and then flips position with the other cephalic trachea at the anterior end of the pronotum and retains the condition of T1-AL extending ventrally. The length of the prothorax in mantises likely plays a role in this dorsal-ventral reversal, but the reasons are unclear. An alternative interpretation is that one of these trunks is T2-CT and splits into H-DCT and H-VCT in the prothorax, in conjunction with dorsal-ventral connections in the head. However, this would also imply that T1-PL splits from T2-CT, a condition not encountered in any specimen.</p><p>Looking toward the posterior end of the thorax, T3-DLT is unusual in that it skips a connection to A1-S, extending from T3-S posteriad to A2-S via A2-DB; A2-DLT continues posteriad as with other taxa. Also curious is T2-PWL branching from T2-PL and connecting with T2-AL. This is similar to what is seen in T 3 in  Orthoptera but appears to be present here in the midleg. The mesothorax is shorter anteriorposteriorly in  Orthoptera, so perhaps this condition is either difficult to discern or simply not present. Another modified branching pattern is T3-VL branching from directly from A1-VC rather than VLT. While the origin (VLT) of T3-VL here is unusual, its extension into the leg and overall pattern appears homologous with T3-VL elsewhere.</p></div>	https://treatment.plazi.org/id/038D8781FF6420C0FCB2FE66A126FA3A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF862021FF0DFC8BA3A2FD3E.text	038D8781FF862021FF0DFC8BA3A2FD3E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Meconema thalassinum (De Geer 1773)	<div><p>Meconema thalassinum</p><p>“Drumming katydid”</p><p>Figures 67 (lateral), 68 (dorsal, ventral)</p><p>Plates 43 (lateral), 44 (dorsal, ventral)</p><p>The female drumming katydid we scanned is likely a late-stage instar due to the reduced wings. T2-DB-Vi and T3-DB-Vi each form a trapezoidal set of double dorsal commissures. It is possible that these may be homologous with sausagelike dorsal commissures seen in other taxa that might be for weight relief or could be involved in stridulatory sound dissemination.</p><p>DESCRIPTION: HEAD: Three pairs of tracheae into head: H-DCT, H-VCT, and H-VLT. Head with several interconnected dorsoventral and lateral loops; readers are encouraged to review the supplementary 3D digital models. Air space, possibly a preservational artifact, present in right side of head capsule (see fig. 69). H-DCT runs anteriad, curving dorsally along head capsule and curving medially to connect in a loop just dorsal of eyes. H-DC present. Anteriad of cervix, H-DCT-Oc branches ventrad, linking with H-VCT-Oc with Y-shaped junction to H-Oc-Md. Several small, visceralike H-Oc run dorsad from H-Oc-Md; H-Oc-Md anteriad, reconnecting to H-VCT with H-Md branching anteriad, with lateral H-FtC branch linking left and right side. H-VCT runs anteriad with ventral curve; T1-AL running posteriad prior to H-VCT entry to head capsule. H-VCT with aforementioned H-VCT-Oc branch running anteriad; H-VCT continuing anteriad with ventral branch, linking with H-VLT before continuing into H-Mx and H-Lbm split.</p><p>THORAX: T2-S opening very large, positioned under pronotum; T2-S with five branches: T2-DB, T2-VB, H-DCT, H-VCT, and T1-PL. Several tracheae behind T1-DVi; see figure 70 for view rotated to see additional detail. Readers are also directed to 3D models from supplemental digital data. T2-DB and T2-VB from short dorsal spiracular branch, extending directly dorsad and ventrad, respectively. T2-DB with T2-AWL split, with T2-DB continuing dorsad, connecting with T2-DLT extending posteriorly. T2-AWL runs dorsad with sharp curve directly ventrad toward midleg; T2-W-c-r positioned posteriad at apex of curve. T2-VB runs directly ventrad from T2-DB/T2-VB split, arcing posteriad to connect with T2-VLT; several small visceral branches forming cagelike network along venter. H-DCT runs dorsad from T2-S, arcing anteriad and continuing through prothorax into head capsule; T1-DVi branches forming cagelike morphology along inner wall of protergum, around H-DCT. H-VCT runs dorsad, arcing ventrad before turning anteriad into head capsule, arranged along outside of enlarged T1-PL; T1-AL with ventral branch anteriad of T1-PL. T1-PL greatly enlarged in hornlike arc, reducing in size and extending into proleg. T1-L-Ty tympanum present on foreleg tibia. T3-S much smaller than T2-S, with five branches: T3-DB, T2-PWL, T3-AWL, T3-VB, and T2-PL. T3-DB and T3-VB split dorsad and ventrad after short dorsal spiracular branch from T3-S. T3-DB running directly dorsad, with T3-AWL branching dorsad and slightly lateral; T3-DB continuing dorsad to join with T2-DLT from anterior and smaller T3-DLT posteriad. T3-AWL runs dorsad, with sharp curve ventrad and posteriad toward midleg. T3-AWL with two connections to tracheae leading to A1-S: T3-Wbr, branching dorsally at apex of curve, with T3-W-c-r and T3-W-cu-a dorsad; and apparent T3-PWL extension, dorsal and anteriad from A1-S, connecting with T3-AWL/T3-AL. (See Discussion on  Orthoptera .) T3-VB runs mediad and ventrad, connecting with T3-VLT toward posterior and T2-VLT anteriad. T2-PL ventrad with curving T2-pf branch to T2-VLT; T2-PL continues into midleg, joining with T2-AL.</p><p>ABDOMEN: A1..8-S present. A1-S modified, positioned slightly dorsad relative to A2..8-S. A1-S with four connections: A1-DB, T3-PWL, A1-VB, and T3-PL. Single A1-DB runs dorsad and slightly anterior, broad and flat; intersecting with T3-DLT anteriad and A1-DLT posteriad. Several small visceral-type tracheae connecting with T3-PWL and and likely extending into flight musculature. T3-PWL running dorsad and anteriad, similar to A1-DB but smaller; T3-PWL splitting slightly dorsad into T3-Wbr anteriad connection to T3-S, and smaller unnamed extension connecting to T3-AL (see THORAX, above, and Discussion section on  Orthoptera). T3-PL thick, running directly ventrad from A1-S, connecting with smaller T3-AL from metathorax and extending into hind leg. T3-VL present, extending into hind leg from T3-VLT. A1-VB runs ventrad and slightly mediad, connecting with T3-VLT from anterior and A1-VLT posteriad. A2..8-S similar, each with anterior and posterior dorsal branches A n -ADB and A n -PDB, both intersecting with A n -DLT along dorsum. A n -VB short, running ventrad and slightly mediad, connecting directly to A n -VLT along venter. X-shaped commissures positioned ventrad to VLT (see fig. 71). Numerous visceral tracheae present in abdomen, often spanning several segments, but none appearing to connect segments longitudinally.</p></div>	https://treatment.plazi.org/id/038D8781FF862021FF0DFC8BA3A2FD3E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF9C20C4FCDDFDB5A37EF84F.text	038D8781FF9C20C4FCDDFDB5A37EF84F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Medauroidea extradentata (Brunner von Wattenwyl 1907)	<div><p>Medauroidea extradentata</p><p>“Vietnamese walking stick”</p><p>Figures 91, 92 (lateral, anterior, posterior); 93, 94 (dorsal, anterior, posterior); 95, 96 (ventral, anterior, posterior)</p><p>Plates 55 (lateral), 56 (dorsal), 57 (ventral)</p><p>Although Strauss (2021) conducted the most recent work on stick insect tracheae, he focused on the prothorax and prolegs, concentrating on hearing. He employed terminology from Ander (1939), also incorporated here, although he left several branches unlabeled. In the  Medauroidea scan here, T1-AL and T1-PL remain separate, at least as far as the distal end of the foretibia. Strauss indicates foreleg (and mid- and hind leg) adaptations for hearing; these cannot be verified from this scan.</p><p>Medauroidea is a good example of assessing homology using secondary criteria (serial homology). The T2-DB branching pattern posteriorly in  Medauroidea is somewhat ambiguous, in particular the placement of T2-AL and T2-Wbr. T2-AL and T2-Wbr could be swapped; however, the decision that the dorsal one is T2-AL is based on serial homology with T3. Additionally, the shorter DB branches, such as T3-DB and T3-VB here in  Medauroidea (but also applicable elsewhere), could be argued to be branching directly from the spiracle. The presence or absence of a spiracular “atrium” is not specific here; these structures are most prominent in groups such as  Orthoptera, where a large “cavity” sits just inside the spiracle with multiple tracheae branching in various directions. In  Medauroidea, it appears that a very short “stub” of DB or VB may have trachea branching from it. T2-CT may be present, but this stub may also be a spiracular atrium, as the closest (phylogenetically) relative with T2-CT is  Plecoptera (rather distant).</p><p>The tracheal branching in first abdominal segment is very unusual and assessing its homology calls for some explanation. A1-DLT is quite clearly absent, as no tracheae arc posteriorly in the form of a DLT; T3-DLT connects directly to the dorsum of A1-S with no continuation. A1-VLT initially appears to be A1-MLT, but the ventral branch of T3-VL into the hind leg is indicative of it being a VLT, not MLT. Additionally, A1-VC branches from T3-VL, rather than A1-VB, an unusual arrangement. This branch is not a section of A1-VB, as A1-VLT connects A1-S and A2-S. The remaining branches are typical.</p><p>The second abdominal segment is also slightly modified from the remaining segments extending posteriad. A2-DB and A2-VB are basically absent—while A2-DC and A2-VC are present, they branch directly from A2-DLT and A2-VLT. The primary connection between A2-S and A1-S anteriad is A1-VLT. A2-DC extends a little anteriad of A2-S; A2-DC could arguably be A1-DC but the branching pattern is from A2-DLT, so homologizing based on connectivity or branching pattern seems more reasonable.</p><p>DESCRIPTION: HEAD: The head tracheal morphology of  M. extradentata features a network of loops interconnecting both dorsoventrally and laterally. Exploration of the 3D models in the supplementary digital data is encouraged. Three sets of tracheae into head: H-DCT, H-VCT, and additional H-VLT. H-DCT dorsad, proceeding anteriorly and forming a prominent H-DCTVLT-Loop anteriad and ventrad, connecting directly with H-VLT. H-Lbm anteriad from ventral apex of H-DCT-VLT-Loop. H-VCT runs anteriad, dividing into H-Ant and two branches, one looping posteriad to connect with H-VCT, forming H-Ant-Loop; second branch looping ventrad and posteriad to join H-VLT.</p><p>THORAX: T2-S with four connections: possible T2-CT, T2-DB, T2-VB, and T1-PL. T2-CT short and running directly anteriad, bifurcating into H-DCT and H-VCT near posterior margin of prothorax; as T2-CT absent in other  Phasmatodea, this T2-CT is possibly a deeper spiracular atrium rather than T2-CT. H-DCT runs anteriad, extending through prothorax into head. H-VCT anteriad, with T1-AL splitting off into foreleg; short connection to T1-VLT at this branching point. T2-DB runs posteriad, curving slightly ventrad before splitting into two pairs: T2-DLT/T2-AL dorsal branch and T2-Wbr/ T2-lvl ventral branch; T2-AWL notably absent. For dorsal T2-DLT/T2-AL pair, T2-DLT as with other specimens, positioned along dorsum with connection posteriad to T3-DB; T2-AL extending posteriad, connecting with T3-S via T2-PWB. Ventral T2-Wbr/T2-lvl pair with T2-AL posteriad with shallow arc dorsad to connect with T2-PWL; T2-lvl along venter, connecting with T3-S via T2-VLT connection just anteriad of T3-S. T2-VB short and directly ventrad, linking with T2-VLT posteriad and T1-VLT anteriorly. Both T1-VC and T2-VC present. T1-PL runs anteriad, linking with T1-VLT via short T1-VL; T1-PL extends into foreleg without joining T1-AL. T1-AL and T1-PL do not join and remain separate at least until the distal end of the foretibia. Two small, visceral medial A1-DVi-Med and A1-VVi-Med extend through mesothorax, originating at A1-S and extending into head. T3-S with four branches: T2-VLT, T3-DB, T3-VB, and T2-PL. T2-VLT from anterior, connecting T2-S to T3-S. T3-DB short and mediad, quickly branching into T3-AL posteriorly and remaining T3-DB dorsad; T3-DB joining with T2-DLT anteriorly and T3-DLT posteriorly in Y-shaped junction, linking to T2-S and A1-S. T3-AL posteriad, joining with T3-PL to form T3-L, extending into hindleg. T3-VB short, similar to T3-DB, quickly splitting into T3-VLT posteriad and remaining T3-VB ventrad. T3-VLT runs directly posteriad to A1-S; T3-DC present toward posterior margin of metathorax. T3-VB continues to T3-PVC, which forms posteriad segment of hexagonal network comprised of T3-VC anteriad and lateral sections connected to T2-VLT. T2-PL anteriad, joining T2-AL and extending into mideg. T2-VL branching from T2-VLT, also extending into midleg; T2-VL and T2-L remain separate to distal end of midleg tibia. A2-DVi-Med and A2-VVi-Med extending through metathorax, with laterally asymmetric connections: A2-DVi-Med to T2-VLT on specimen’s left side, A2-DVi-Med to T2-VLT on specimen’s right side.</p><p>ABDOMEN: A1..8-S present. First abdominal segment approximately half as long as remaining abdominal segments. A1-S and A2-S branching patterns highly modified from remaining abdominal segments. A1-S with four branches: T3-DLT, A1-VLT, T3-VLT, and A1-PL. A1-S with additional A2-VVi-Med connection on left side only, see description of A2-VVi-Med below. T3-DLT runs dorsad, connecting in anterior arc from T3-S; A1-DLT notably absent. A1-VLT runs directly posteriad to A2-S; T3-VL directly ventrad from A1-VLT; A1-PVC branching from T3-VL. T3-VLT from anterior, with two ventral commissures A1-AVC1 and A1-AVC2. T3-PL ventrad, linking with T3-AL before arcing posteriad to extend into hind leg. A2-S with seven branches: A1-VLT, A2-DVi-Med, A2-VVi-Med, A2-DLT, A2-DVi, A2-VVi, and A2-VLT. A1-VLT runs directly anteriad from A1-S. A2-DVi-Med beginning as sinusoidal, looping branch, extending anteriorly and medially, each side combining in Y-shaped junction in metathorax and proceeding anteriorly along venter; A2-DVi-Med asymmetric, with sinusoidal form on both sides but only right side leading to Y-shaped join, with left side coming from A1-S. A2-VVi-Med similar, arranged along venter. A2-DLT runs dorsad and posteriad in arc connecting to A3-S; A2-DC present off A2-DLT. A2-DVi and A2-VVi ventrad, connecting with A3-S. A2-VLT likewise in ventral arc, also connecting with A3-S; four tracheae connect A2-S with A3-S. Remaining A3.. A8 segments similar, with varying degrees of A n -DB; A3-DB short, A4-DB not present, etc. A n -DLT and A n -VLT present on all, connecting segments longitudinally. A3..8-S with anteriad visceral tracheae generally dorsad, posteriad visceral tracheae generally ventrad. A4-Vi-VC present, formed from visceral tracheae and not from VLT as is typical.</p></div>	https://treatment.plazi.org/id/038D8781FF9C20C4FCDDFDB5A37EF84F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFCA206AFF3EFF7DA239FA7C.text	038D8781FFCA206AFF3EFF7DA239FA7C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Neocloeon triangulifer	<div><p>Neocloeon triangulifer subimago</p><p>“Triangle small minnow mayfly”</p><p>Figures 24 (lateral), 25 (dorsal, ventral)</p><p>Plates 14 (lateral), 15 (dorsal, ventral)</p><p>DESCRIPTION: BODY: Several alimentary canal air spaces present, fusing into single one in adult.</p><p>HEAD: H-DCT and H-VCT similar in size, extend into head capsule from prothorax.</p><p>THORAX: T1-DVi branches from H-VCT on left side, H-DCT on right.</p><p>ABDOMEN: A2..7-Gi gill attachments visible laterally on A n -DLT, midway between spiracles; A1-Gi on posteroventral extension from A1-S, reduced in adult (no other vestiges of gill attachment sites present in adults). Several dorsal visceral tracheae extend anteriorly beyond segment boundaries; ventral Vi (retained) within segment.</p></div>	https://treatment.plazi.org/id/038D8781FFCA206AFF3EFF7DA239FA7C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFCA206AFF25FC0EA1F3FC8B.text	038D8781FFCA206AFF25FC0EA1F3FC8B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Neocloeon triangulifer	<div><p>Neocloeon triangulifer adult</p><p>“Triangle small minnow mayfly”</p><p>Figures 26 (lateral), 27 (dorsal, ventral)</p><p>Plates 16 (lateral), 17 (dorsal, ventral)</p><p>DESCRIPTION: H-DCT, H-VCT, T n -DLT, A n - DLT, and several T n -FM compressed laterally along some sections; since subimago maintains these as broad tracheal lumena, this is likely a preservation artifact in the adult.</p><p>BODY: Single air sac, coopted from alimentary canal, extending from head capsule to end of 7th abdominal segment (see fig. 21).</p><p>HEAD: As above, but with H-DOc extending from trachea leading to H-PO.</p><p>THORAX: T1-DVi from H-VCT.</p></div>	https://treatment.plazi.org/id/038D8781FFCA206AFF25FC0EA1F3FC8B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFCA2077FEE8FA64A431FBDB.text	038D8781FFCA2077FEE8FA64A431FBDB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Odonata Several D Tillyard 1914	<div><p>ORDER  ODONATA</p><p>Respiration in  Odonata, especially aquatic naiad immatures, has been the subject of investigation for some time, although much of the morphological work was performed more than a century ago. Scott (1905) presented one of the earliest detailed studies on dragonfly tracheae using a libellulid immature, highlighting notable characteristics such as the crossover of longitudinal trunks leading from the thorax to the abdomen, a trait retained in the adult. Tillyard (1917) remains the standard for dragonfly morphology, and he carefully diagrammed the tracheal architecture via dissection, although also with an emphasis on immatures. Chapman (1918) included both  Anax and  Lestes in his comparative study of leg-wing tracheal morphology. Kennedy (1922) built on Chapman’s work, extending it to abdominal tracheae; his extension to multiple orders insects using only odonate specimens is shown here to be insufficient. Recent studies have investigated physiological aspects, including active tracheal compression (likely related to respiration) in  Odonata observed through synchrotron imaging by Westneat et al. (2003). For this study, three odonates were scanned, an aeshnid dragonfly larva at 30 µm resolution, an adult aeshnid at 21 µm resolution, and a calopterygid damselfly at 19 µm. As noted above, the wings of the adult aeshnid were removed and scanned separately; wing scans are not included here, though wing-base tracheae are retained and labeled.</p><p>Tillyard (1917) identified three pairs of longitudinal trunks in his figure 75 (included here as fig. 28), whereas the aeshnid and calopterygid specimens here possess four, a condition not seen in any other order. A n -DLT is comprised of elongated and compartmentalized air-saclike tracheae connected segmentally or perhaps intermittently by regular tubular tracheae, and these were likely misidentified by Tillyard as air sacs rather than tracheae, which were discernible as such with micro-CT.</p><p>Scott (1905) indicated several tracheae present in the mid and hind legs of immatures but absent in adults, also noted by Chapman (1918), although with his own terminology. It is possible that these extra tracheae develop into the air-sac tracheae, which seem to be largely absent overall in immatures (otherwise they would have difficulty swimming underwater). Future studies should investigate this proliferation of tracheae and their possible development into the numerous air-sac tracheae.</p><p>The thorax of both specimens features a large number of air spaces. The flight musculature of odonates is thought to grow substantially during maturation of the adult (Marden, 1989; Anholt et al., 1991; Marden, 2000). It could be that these air spaces allow expansion of flight muscle—as the adult exoskeleton is fixed in size, empty volume must be allocated in advance to allow for an increase in flight muscle mass. Interestingly, this would be the inverse of the “gas bag in the gut” phenomenon (see Discussion), where the alimentary canal is coopted into a large air space in the adult.</p></div>	https://treatment.plazi.org/id/038D8781FFCA2077FEE8FA64A431FBDB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF8F2034FCA0FBA4A250F86C.text	038D8781FF8F2034FCA0FBA4A250F86C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Oligotoma negra Hagen 1885	<div><p>Oligotoma negra</p><p>“Black webspinner”</p><p>Figures 75 (lateral), 76 (dorsal, ventral)</p><p>Plates 47 (lateral), 48 (dorsal, ventral)</p><p>The respiratory system of  Embioptera remains rather understudied, with the most recent detailed analysis of tracheal anatomy by Lacombe (1958, 1971). The specimen of  Oligotoma shown here is a male and is notable because of the air-filled alimentary canal (fig. 77). Male webspinners do not feed, and like  Ephemeroptera (fig. 21), the alimentary canal is coopted as a large air space, spanning the length of the body. In this specimen, the distention of the alimentary canal compressed tracheae against the inner body wall, making determination of morphology and assessment of homology challenging (particularly in the abdomen). Some notable differences were observed between Lacombe (1958) and the specimen here, particularly in the thorax, but her work was useful in mapping abdominal tracheae.</p><p>The meso- and metathoracic wing base T2,3- Wbr are likely present but partial in this scan, likely due to fluid infilling of this specimen as it was frozen to -20° C rather than -80° C.</p><p>Although a tympanal hearing organ in the femur (of pro- and mesoleg, and occasionally hind leg) of webspinners was described by Szumik et al. (2019), there is no evidence of an air space in femur of any leg (as seen here in the foretibia of  Gryllus, for example).</p><p>DESCRIPTION: HEAD: H-DCT extending anteriad, curving laterally along head capsule; H-VCT slightly ventrad. H-DC present, slightly anterior of cervix. H-DCT with H-DCT-DVi running dorsad anterior of cervix, following head capsule. H-DCT runs directly anteriad, dividing into H-DVB ventrad and H-Ant. H-VCT with H-VCT-DVi extending dorsally, nearly in contact with H-DCTDVi and following head capsule laterally. H-VCT continues anteriorly, with dorsal H-DVB connection to H-DCT. After short extension, H-VCT divides into H-Lbm, and subsequently into H-MxPlp, H-Md, and H-Oc. H-Lbm branch connecting mediad as H-VC. Branch leading to H-Oc continues to H-Ft.</p><p>THORAX: T2-S with four branches: H-DCT, H-VCT, T2-DB, and T2-VB. T2-CT absent. H-DCT runs directly anteriad, with T1-DVi branching dorsad before continuing dorsally along prothoracic tergum. T1-DC present, extending mediad near branching of T1-DVi. H-VCT runs anteriad, with T1-L branching ventrad just prior to a nearly 90° turn ventrad, subsequently turning anteriad along prothoracic sternite. T1-VC present, branching off T1-L. T2-VB runs posteriad and slightly ventral briefly before curving dorsad; small T2-AWL branching anteriad at apex of curve, dividing into T2-AWba dorsad and T2-AL continuing posteriorly toward midleg. T2-DB continues, curving medially with several branches, likely for flight muscle, before extending posteriad as T2-ADLT. Full DLT not visible but likely present, possibly fluid infilled; Lacombe (1958) indicates presence of dorsal connective. T2-VB runs posteroventrad, with large T2-FM branching dorsad and laterally. T2-VB continues as T2-VLT, connecting with T3-S. T2-VC present, branching mediad approximately halfway between T2-S and T3-S. T3-S with four branches: T3-AWL, T2-PWL, T3-DB, and T3-VB. T3-AWL running dorsad, curving posteroventrad where T3-W-c-r splits dorsally and remaining branch continues as T3-L. T2-PWL connecting directly from anterior; T2-PL branching ventrally, joining with T2-AL after a short distance and continuing as T2-L; T2-Wbr continues anteriad from T2-PWL; T2-Wbr partial likely due to fluid infilling. T3-DB runs mediad, dividing after a short distance into T2-PDLT anteriorly and T3-DLT posteriad. T3-VB runs posteroventrad, with a connection to T2-VLT very close to T3-S; T3-VB continues posteriorly as T3-VLT, linking up with A1-S. T2-VLT-Vi on right side, extending past T3-S into abdomen; unclear if T2-VLT-Vi links with an abdominal spiracle. T3-S on right side slightly different from left, possibly due to displacement of tracheae by distended alimentary canal air space; T3-S on right side with T3-VLT positioned posteriad as with right side, but with short, curving spiracular branch dorsad and anteriad, with T2-VLT connecting from anterior; branch continues dorsad to T2-AWL, T2-DB, T2-PWL split as with left side.</p><p>ABDOMEN: A1..8-S present. A1-S modified from subsequent abdominal segments; A1-S with five branches: A1-DB, T3-PWBa, T3-VLT, and T3-PL. A1-DB short and running mediad, connecting with T3-DLT from anteriad and continuing as A1-DLT posteriad. T3-WBr (also partial, likely due to fluid infilling) runs anteriad, continuing with small T3-W-cu-a into trailing edge of hind wing. T3-VLT from directly ventrad. T3-PL runs ventrad, lateral from T3-VLT, joining with T3-AL and extending into hind leg as T3-L. Segments A2..7 likely similar but morphology difficult to determine due to distention of air-filled alimentary canal. A2..7-DLT present, arcing slightly dorsad and usually sinuous. A n - VLT present. A6-VC visible; other A n -VC likely present but displaced against body wall, see figure 78. Several visceral tracheae visible, but dif- ficult to determine morphology; A3-Vi and A4-Vi large and directly posteriad, spanning sev- eral segments. A-Cr visible at base of cerci.</p></div>	https://treatment.plazi.org/id/038D8781FF8F2034FCA0FBA4A250F86C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFB12013FCC5FC74A1ACFFFB.text	038D8781FFB12013FCC5FC74A1ACFFFB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Orthoptera	<div><p>ORDER  ORTHOPTERA</p><p>Orthopteran anatomy (and some tracheae) has been diagrammed in classic works, but the first proper treatments of respiratory systems were by Vinal (1919: for  Dissosteria carolina), Carpentier (1927: for  Phasgonura viridissma), and Ander (1939: for  Ensifera in general). Several tracheal terms were introduced using  Orthoptera, particularly those involving auditory adaptations and saltatorial legs, and the work of these researchers (particularly Ander) was instrumental in mapping the tracheae here. More modern studies have focused on physiology, including ventilation in  Schistocera gregaria in influential experimental works by Miller (1960a, 1960b, 1960c), active tracheal compression in  Orthoptera observed via synchrotron imaging by Westneat et al. (2003), and the distribution of air spaces in  Schistocera americana by Shaha et al. (2013).</p><p>Four orthopteran specimens were scanned:  Gryllus sp. (Gryllidae),  Romalea microptera (Romaleidae),  Tachycines asynamorous (Raphidophoridae), and a  Meconema thalassinum (Tettigoniidae) .  Orthoptera tracheal morphology is best represented here in the scans of  Gryllus and  Meconema, and these are described here in detail. Notable features are described for the remaining specimens, but they are presented in a more basic manner due to the suboptimal quality of the scan of  Tachycines and the substantial complexity of  Romalea .</p><p>The most comprehensive work to date on tracheal morphology of  Orthoptera is by Ander (1939). His work formed the basis for many of the homology statements here, although we reinterpret some of his tracheae, in particular the supraventral (suv) in the prothorax and mesothorax. Ander indicates that the suv is absent in the metathorax in all  Ensifera (his fig. 81). Interestingly, Ander cites Carpentier (1927) as the source for supraventral, where it does not go into the leg (Carpentier’s figure 3). However, Ander’s figure 83 details  Gryllidae (fig. 81 is his overall view of  Ensifera) and identifies three tracheae going into the proleg: pa, pp, and suv (supraventral), an apparent departure from Carpentier’s suv. Herein, T n -AL refers to Ander’s pa, T n -PL for pp, and T n -VL for pve. (Refer to table S 1 in the online supplement for all reassignments of labels and terms from previous studies.) Ander’s suv, more consistent than Carpentier’s, is likely T1-Cx (and even T1-PL in some taxa), designated here as such. Ander’s suf (suprafurcal), lvl (lateral lateroventral), and pf (postfurcal connective, apparently absent in  Gryllus but present in other  Ensifera) tracheae were not found to be homologous outside  Orthoptera . His terminology is retained for descriptions of  Orthoptera for convenience.</p><p>The dorsum of the metathorax in both  Gryllus (Gryllidae) and  Meconema (Tettigoniidae) both possess T3-DB-Vi, paired air sacs reminiscent of a root vegetable, and ending blind (fig. 59, pl. 36). These tracheae are similar in morphology to those found in the ventral thorax of  Dictyoptera for hearing, and their possible function as such should be investigated further.</p></div>	https://treatment.plazi.org/id/038D8781FFB12013FCC5FC74A1ACFFFB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6320CDFC9EFDE0A4A9FFFB.text	038D8781FF6320CDFC9EFDE0A4A9FFFB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Periplaneta americana	<div><p>FAMILY  BLATTIDAE Periplaneta americana</p><p>“American cockroach”</p><p>Figures 107 (lateral), 108 (dorsal), 109 (ventral)</p><p>Plates 64 (lateral), 65 (dorsal), 66 (ventral)</p><p>The P. americana specimen here was scanned early in the study, frozen to -20° C rather than -80° C, and at relatively coarse resolution (38.6 µm, specimen approximately 3.5 cm in length). Some fluid infilling likely occurred due to the freezing to -20° C, but sufficient detail is present to assess homology of major tracheae and discern substantial visceral tracheal anatomy. Although not described in detail, Blaptica was used as a comparison to infer the likely presence of thoracic tracheae in Periplaneta, in particular branching patterns from the mesothoracic spiracle. The thorax features a network of bandlike visceral tracheae along the dorsum and venter, connecting in various locations throughout. This morphology, combined with the long, broad coxae reminiscent of silverfish, obscures internal thoracic tracheal structures in two-dimensional views. The hypognathous position of the head also results in ventral tracheal views of the thorax being obscured; see figure 110 for a ventral view with head and leg tracheae removed.</p><p>Periplaneta has large air spaces in the coxae, and jumping has been filmed at high speed by Smith (2022). The scan here is of relatively coarse resolution (38 µm); it is possible muscle tracheae could be observed at higher resolutions or with contrast-enhancing stains such as iodine or phosphotungstic acid (Gignac et al., 2016).</p><p>DESCRIPTION: HEAD: H-DCT and H-VCT present, branching into head capsule in highly networked cagelike arrangement. Mouthparts, H-Ant, and H-Oc determined, but networked nature of head morphology, combined with some apparent fluid infilling, makes assessing homology difficult. Readers are directed to 3D models in the supplementary digital data.</p><p>THORAX: T2-S with five branches: H-DCT, H-VCT, T2-DB, T2-AWL, and T2-VB. H-DCT and H-VCT both anteriad, curving slightly medially, with H-DCT proceeding along dorsum and H-VCT along venter. T1-AL branching ventrad off H-VCT. T2-DB short, splitting anteriad into T2-DLT and posteriad as network of cagelike T1-DVi along pronotum. T2-AWL posteriad, with T2-W-c-r splitting dorsad and posteriorly; T2-AWL splits into T2-AL extending into midleg and T2-Wbr, partial in this scan but likely present between T2-S and T3-S. T2-VB likewise short, bifurcating into T2-VC directly ventrad and T2-VLT posteriad. T2-VL branching posteriad close to T2-S, extending into midcoxa and T2-L; T2-VLT continuing posteriad to link with T3-S. T3-S with n branches: T2-PWL, T3-DB, T3-AWL, T2-VLT, and T3-VLT. T3-VB not visible but may be very short. T2-PWL anteriad from T2-S, bifurcating into T2-Wbr anteriad and curving posteriad as T2-PL, joining with T2-AL from anteriad and extending into mid-leg. T3-DB mediad and slightly ventrad, curving posteriorly before extending along dorsum to join with small T2-DLT anteriad and larger T3-DLT posteriad; T3-DC visible at this junction. T3-AWL posteriad, with T3-W-c-r splitting dorsally; T3-AWL continues as T3-AL posteriorly and ventrad into hind leg. T2-VLT mediad, connect- ing anteriorly to T2-S. T3-VLT mediad and slightly posterior, connecting to A1-S via A1-VB.</p><p>ABDOMEN: Abdomen featuring numerous visceral tracheae throughout. A1..8-S present. A1-S modified from A2..8-S, with three branches: T3-PWL, A1-DB, A1-VB. T3-PWL anteriad and slightly mediad, curving posteriorly and joining with T3-AL and extending into hind leg; T3-Wbr branching from T3-PWL at apex of curve and extending anteriad to T3-S. A1-DB runs mediad, linking with T3-DLT anteriad and A1-DLT posteriad; A1-DB partial in this scan, likely due to fluid infilling (A1-DB present in B. dubia and G. portentosa). A1-VB mediad and slightly ventrad, extending along venter to A1-VC, connecting to A1-VLT posteriad; A1-MLT posteriad from A1-VB. A n -MLT runs straight, extending posteriad and much smaller than thoracic T3-VLT. A2..7-S branching all similar: A n -DB, A n -VB, and A n -MLT. A n -DB beginning small, expanding into bandlike trachea and intersecting with A n -DLT sections anteriorly and posteriorly in Y-shaped junctions; A n -DC absent. A n -MLT wide and almost bulblike, connecting abdominal spiracles along lateral margin of body wall. A n -VB much smaller and apparently fluid infilled in several spiracles, extending along venter to join with straight A n -VLT anteriad and posteriad in T-shaped junction. A4..8-S with numerous visceral tracheae; readers are encouraged to view 3D models in supplementary digital data.</p></div>	https://treatment.plazi.org/id/038D8781FF6320CDFC9EFDE0A4A9FFFB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFAB2016FEF8FA9DA1FEFD45.text	038D8781FFAB2016FEF8FA9DA1FEFD45.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Perlodidae	<div><p>FAMILY  PERLODIDAE</p><p>“Stripetails” or “Springflies”</p><p>Figures 51 (lateral), 52 (dorsal), 53 (ventral)</p><p>Plates 29 (lateral), 30 (dorsal), 31 (ventral)</p><p>The scanned perlodid is a gravid female specimen, with numerous eggs visible (see section on abdomen). These have been omitted from visualizations for clarity; visceral tracheae anastomose among eggs. Large central air sac begins in mesothorax and extends as far as A3-S in abdomen (also omitted from visualizations for clarity, see fig. 50). As mentioned above, adult perlodids have reduced mouthparts and generally do not feed; the head tracheal system is described in full here.</p><p>DESCRIPTION: HEAD: H-DCT and H-VCT elongate and winding through prothorax, bifurcating from T2-CT well posteriad of cervix. H-DCT begins straight on entry to head capsule with short H-DVT-DVi before turning medially toward eye, with three branches: H-Oc, H-FtAnt, and H-Ft. H-Oc short, extending laterally into eye. H-Ft-Ant dorsoventral, linking H-DCT with H-Ant from H-VCT. H-DCT blind ending at H-Ft. H-DCC absent. H-VCT generally straight and running anteriad, with anterodorsal H-VCT-DVI and ventral H-Lbm before continuing anteriad to split into H-Ant, H-VC, and H-Md-Lbr. H-Ant lateral, with connection to H-DCT via H-Ft-Ant. H-VC running directly mediad, linking with opposite side of head. H-Md-Lbr continues anteriad, arcing medially with branch into H-Md before continuing anteriorly and blind ending at H-Lbr.</p><p>THORAX: T2-S with four tracheae: T2-CT, T2-DB, T2-VB, T2-AWL. T2-CT large in diameter, proceeding anteriorly in sinusoidal curve, much shorter than in  Nemouridae, dividing into H-DCT and H-VCT in mid-prothorax. Single T1-Vi present. T1-L running ventrad from H-VCT, with tibial trachea greatly thickened relative to femoral trachea. T1-VC present, extending mediad from each T1-L to connect with opposite side; T1-VC-Avi extending anteriad toward head, looping posteriad near cervix; T1-VC-Pvi running directly posteriad. T2-DB running directly mediad, arcing posteriorly to continue as T2-DLT, with several T2-DLT-Dvi branching dorsally, likely into flight muscle; T2-FM off T2-DLT and T2-DC not visible but likely present. T2-VB posteriad and ventrad, ending blind near mesocoxae; T2-VC present, positioned approximately halfway between forecoxae and mesocoxae; continuation of T2-VB as T2-VLT toward T3-S (as in  Nemouridae) likely present but not visible in this scan. T2-VB-Vi not visible but likely present. T2-AWL begins dorsally, turning posteriorly and ventrad toward midleg, joining T2-PL ventrally from T3-S; T2-AWL with a sharp turn posteriad, likely where T2-AL and T2-Awba (or T2-W-c-r) would split but not visible in this scan; T2-AL tracheal lumen not visible and likely fluid filled in right side; left side of same specimen with gap and smaller trachea connecting. T2-W-c-r not visible but likely present. T3-S with four connections: T3-DB, T3-VB, T2-PWL, T3-AWL. T3-DB short and running directly inward, linking with T2-DLT running anteriad and T3-DLT posteriad; large T3-FM dorsad where T3-DB joins T2-DLT and T3-DLT. T3-FM running dorsad with several tracheae extending into flight muscles; T3-FM continues dorsad, arcing medially to join with opposite side via T3-DC, not visible but likely present. T3-VB running ventrad and posteriad, with numerous branches extending into flight muscule; T3-VB with T3-VC branching inward, meeting opposite side, near ventral sternite. T3-VLT not visible but likely present. T2-PWL running directly anteriad, bifurcating into T2-Pwba extending dorsally and anteriad and T2-PL, mediad. T2-PWBa continues dorsally, with several branches into flight muscle and single, small T2-W-cu-a extending into trailing edge of forewing. T2-PL arcing medially and ventral before turning laterally, joining with T2-AL anteriorly and extending posteriorly into T2-L. T2-L with tibial trachea greatly thickened relative to femoral trachea. T3-AWL running ventrad, just medial from T3-S, with small T3-W-c-r branching dorsally where remaining trachea turns posteriad as T3-AL. T3-W-c-r not visible but likely present.</p><p>ABDOMEN: Numerous eggs present, see figure 54. T 3 -Pwba branching directly from A1-S. A n -VB, A n -SB, A[1..7]-VC absent (or not visible), A8-VC present. A[1..8]-DLT-Dvi present, possibly supplying dorsal vessel. Several segments with elongate visceral tracheae, spanning several segments, including: A2-DLT-Vvi2, extending posteriorly past A5-S; A7-DLT-Vvi, extending anteriorly past A5-S; A6-DLT-Dvi, extending dorsad and anteriorly past A4-S on left side only. Remaining visceral tracheae elongate but winding, often extending past segment boundary before reversing direction, occasionally several times; visceral tracheae often with asymmetric left/right branching patterns. Noticeable lack of left-right commissures with exception of A8-S and right side A8-DLT-Dvi extending mediad, crossing center line and proceeding anteriorly on left side toward A7-S; A8-DLT-Dvi on left side extending anteriorly, not crossing to opposite side. FAMILY  NEMOURIDAE</p></div>	https://treatment.plazi.org/id/038D8781FFAB2016FEF8FA9DA1FEFD45	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF942034FF0CFE07A246FD89.text	038D8781FF942034FF0CFE07A246FD89.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phasmatodea Jacobson & Bianchi 1902	<div><p>ORDER  PHASMATODEA</p><p>Tracheal architecture of stick insects has most recently been investigated by Strauss (2021), with an emphasis on the prothorax and possible adaptations for sound reception. Here, three specimens were scanned: the basal  Timema cf. californicum, the popular Australian stick insect  Extatsoma tiaratum, and the Vietnamese stick insect  Medauroidea extra- dentata. Although many basal taxa exhibit synapo- morphies found in sister taxa of a given monophyletic group,  Timema exhibits numerous apomorphies and its tracheal system is quite distinct from  Extatosoma and  Medauroidea .  Medauroidea appears to be more representative of the larger phas- mids, with good scan quality and characters that appear to be common to  Extatosoma (which is more complex and somewhat harder to homolo- gize).  Timema and  Medauroidea are described here in detail;  Extatosoma is covered here in a more basic fashion due to the substantial complexity in the abdomen. Future studies should focus on the unique morphology of the abdominal tracheae—resem- bling the jumble of dried, packaged ramen noo- dles—of  Extatosoma and its possible functions.</p></div>	https://treatment.plazi.org/id/038D8781FF942034FF0CFE07A246FD89	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFA8200BFEF8F9A4A137FD88.text	038D8781FFA8200BFEF8F9A4A137FD88.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Plecoptera	<div><p>ORDER  PLECOPTERA</p><p>Respiration in stonefly naiads via gills has been studied for decades, partly because of the significance of stoneflies as indicators of water quality. Two plecopteran specimens were scanned, one from  Perlodidae and one from  Nemouridae . The  Nemouridae scan, at 3 µm resolution, shows substantially more smaller tracheae than the  Perlodidae, scanned at 6 µm. Cromulent detail is present in both specimens to assess homology.</p><p>Numerous visceral tracheae branch from A4-DLT-Dvi and extend throughout the abdomen. Tracheae from A4-DLT-Dvi are pervasive throughout the abdomen, leading to the question of why this particular spiracle is so important. Other insects have adapations for individual spiracles, such as the single abdominal “hissing” spiracle in  Gromphadorhina (Nelson, 1979; Nelson and Fraser, 1980; Heinrich et al., 2013), but the presence of tracheae throughout the abdomen indicates a likely respiratory function. Future studies should investigate O 2 input versus CO 2 output, for example.</p><p>The perlodid specimen scanned here is another example of cooption of the gut as an air space in nonfeeding adults. Likely for weight reduction, this condition also seen in mayflies and male  Embioptera .</p><p>Other notable open issues include ventral visceral VVi tracheae in  Perlodidae that are likely VC, as these are present in  Nemouridae . Additionally, T2,3-VLT are present in  Nemouridae but not  Perlodidae . It is possible that the absence of T2,3-VLT in  Perlodidae is a preservational artifact and scanning of further specimens is indicated to verify these features.</p><p>DESCRIPTION: HEAD: The two specimens differ in life history: adult  Nemouridae, in the Group Euholognatha, have functional mouthparts adapted for feeding on algae, lichens, or soft pollen; while  Perlodidae, in Systellognatha, is known to have adults with reduced mouth parts that typically do not feed. Consequently, head morphology is different enough to describe each family in the sections below.</p><p>THORAX: Thoracic tracheal morphology is largely similar between the two species scanned but with substantive differences; thoraces for both are described here for comparative clarity and also detailed in family sections below. Large thoracic air spaces present in both species, but spherical nature of air spaces in  Nemouridae indicates possibility of preservational artifact (see fig. 50). Putative air sac in  Perlodidae begins in thorax, extending into abdomen. T2-S with four tracheae: T2-CT, T2-DB, T2-VB, T2-AWL. T2-CT very thick, proceeding anteriorly in sinusoidal curve;  Nemouridae with two branches off T2-CT: T1-L extending ventrad, and T1-Gi just posteriad of cervix. T1-Vi present. T1-L with tibial trachea greatly enlarged relative to femoral trachea. T2-DB running directly mediad, arcing posteriorly to continue as T2-DLT, with a number of T2-DLT-Dvi branching dorsally, likely into flight muscle;  Nemouridae with large T2-FM extending dorsad close to T2-S; T2-Fm arcs medially to connect to opposing side as T2-DC; T2-FM off T2-DLT and T2-DC not visible in  Perlodidae . T2-VB posteriad and ventrad; in  Perlodidae, blind ending near mesocoxae; T2-VC present, positioned approximately halfway between procoxae and mesocoxae. T2-VB asymmetric in  Nemouridae; on right side, extending ventrad and posteriad, arcing dorsally just anterior to mesocoxae to link with T3-S via T2-DLT, while on left side, T2-VB mirrors right side but turns abruptly mediad to link with T2-VB on left side, forming T2-VC. T2-VB in  Nemouridae with multiple T2-VB-Vi that likely supply flight muscle; single large T2-VB-Vi extending directly from T2-S on left side but mirroring similar T2-VB-Vi from T2-VB on right side. T2-VB-Vi not visible in  Perlodidae scan. T2-AWL begins dorsally, turning posteriorly and ventrad toward midleg, joining T2-PL ventrally from T3-S; T2-AWL with a sharp turn posteriad in  Perlodidae, likely where T2-AL and T2-Awba (or T2-Wc-r) would split but not visible in this scan; in  Nemouridae T2-AWL short, with T2-AL and T2-W-c-r bifurcating just dorsal to T2-S. T2-AL tracheal lumen not visible and likely fluid filled in right side of  Perlodidae; left side of same specimen with short gap and smaller trachea connecting. T2-AL complete in  Nemouridae . T2-W-c-r extending dorsally and posteriad into wing. T2-W-c-r not visible in  Perlodidae . T3-S with four connections: T3-DB, T3-VB, T2-PWL, T3-AWL. T3-DB short and directed inward, linking with T2-DLT anteriorly and T3-DLT posteriorly; large T3-FM runs dorsad where T3-DB joins T2-DLT and T3-DLT. T3-FM running dorsad with several tracheae extending into flight muscles; T3-FM continues dorsad, arcing medially to join with opposite side via T3-DC; T3-DC not visible in  Perlodidae but likely present. T3-VB runs ventrad and posteriad, with numerous branches extending into flight muscle; T3-VB with T3-VC branch inward, meeting opposite side, near ventral sternite. In  Nemouridae, T3-VB on left side continues as T3-VLT, linking with A1-S; T3-VLT absent on right side. T3-VLT not visible in  Perlodidae . T2-PWL running directly anteriad, bifurcating into T2-Pwba extending dorsally and anteriad and T2-PL, mediad. T2-PWBa continues dorsally, with several branches into flight muscle and single, small T2-W-cu-a extending into trailing edge of forewing. T2-PL arcing medially and ventral before turning laterally, joining with T2-AL from anterior and extending posteriorly into T2-L. T2-L with tibial trachea greatly enlarged relative to femoral trachea. T3-AWL running ventrad, just medial from T3-S, with small T3-W-c-r branching dorsally where remaining trachea turns posteriad into T3-AL. T3-W-c-r not visible in  Perlodidae but likely present.</p><p>ABDOMEN: Abdominal morphology largely similar between  Perlodidae and  Nemouridae, with overall structure described here and specific differences given below. Air sac in  Perlodidae extends as far as A3-S (fig. 50). A[1..8]-S present. A1-S branching modified from remaining abdominal segments, with slight differences between  Perlodidae and  Nemouridae . Both specimens with A1-DB and A1-VB;  Perlodidae with T3-Pwba directly from A1-S,  Nemouridae with T3-Pwba from A1-VB slightly ventrad from A1-S. Both T3-Pwba anteriad and slightly dorsad, splitting into several smaller visceral tracheae likely supplying flight muscle and single T3-W-cu-a into trailing edge of hind wing. A[1..8]-DB and A[1..8]-DLT present in both taxa, with numerous visceral tracheae from DLT detailed in sections below. A[1..8]-VB, A[2..8]-SB, A[1..8]-VC present in  Nemouridae but absent (or not visible) in  Perlodidae . See family-level sections for descriptions of visceral tracheae.</p></div>	https://treatment.plazi.org/id/038D8781FFA8200BFEF8F9A4A137FD88	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF7220DCFCC6FD94A35CFB75.text	038D8781FF7220DCFCC6FD94A35CFB75.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Reticulitermes flavipes (Kollar 1837)	<div><p>Reticulitermes flavipes</p><p>“Eastern subterranean termite”</p><p>Figures 122 (lateral), 123 (dorsal, ventral)</p><p>Plates 76 (lateral), 77 (dorsal), 78 (ventral)</p><p>DESCRIPTION: HEAD: H-Oc absent, as is typical in termite workers and soldiers. H-DCT runs slightly dorsad anterior of cervix, with visceral branches dorsad and lateral. H-DCT arcing anteriad, with H-Ant branching laterally where H-DCT meets H-DVB join with H-VCT. H-VCT running directly anteriad, with split into H-MdMx branch, H-Lbm, and third branch that splits into H-DVB dorsad, H-Ft anteriad, and H-VC medially. H-Md-Mx anteriad, with H-Mx dividing ventrad. H-Md runs anteriad with short branch anteriad as second H-Ant.</p><p>THORAX: T2-S with four branches: H-DCT, H-VCT, T2-DB, and T2-VB. H-DCT runs mediad, curving anteriad toward head capsule. H-VCT curving similar to H-DCT, but with single T1-L directly ventrad; T1-AL and T2-PL absent, but T1-VC present, branching from T1-L and joining medially. T2-DB short, with T2-AWL splitting dorsally and posteriad; T2-DLT continues medially, splitting into Y-shaped junction with T2-DLT posteriad and a pronounced, looping T1-DLT joining with H-DCT. T2-AWL arcing dorsad, bifurcating into T2-Wbr and T2-AL at apex of curve. T2-Wbr runs posteriad, linking with T3-S via T2-PWL; T2-AL runs ventrad, proceeding into T2-L. T2-VB short and running ventrad, splitting into anterior visceral branch and remaining T2-VB, running ventrad and toward posterior as T2-VLT. T2-Wbr with small T2-W-c-r and T2-W-cu-a present. T2-VC present, extending off T2-VLT. T3-S with three branches: T2-PWL, T3-DB, and T3-VB. T2-PWL anteriad from T3-S, splitting into T2-Wbr toward T2-S and T2-PL running ventrad, joining with T2-AL and proceeding into midleg as T2-L. T3-DB runs directly mediad, with T3-AWL branch arcing posteriorly close to T3-S; remining T3-DB runs further mediad, joining with T2-DLT from anterior and proceeding as T3-DLT toward posterior. T3-AWL arcing posteriorly, splitting into dorsal T3-Wbr and ventrad T3-AL. T3-W-c-r and T3-W-cu-a not visible off T3-Wbr but likely present and very small. T3-VB runs ventrad with anterior branch leading to T2-VL. T2-VB continuing ventrad, joining with T2-DLT anterior and T3-DLT posterior in Y-shaped junction. T3-VC present, branching medially off T3-DLT.</p><p>ABDOMEN: A1..8-S present. Short A n -SB possible on several segments. Nearly all A n -DB and A n -VB with visceral branches that occasionally span several segments. A1-S with three branches: T3-PWL, A1-DB, and A1-VB. T3-PWL joining medially from anterior, completing link between T3-S via T3-Wbr. A1-DB runs mediad and slightly ventrad, with A1-DB- Vi branching ventrally while A1-DB, smaller, continuing medially to join T3-DLT from anterior and A1-DLT to posterior in Y-shaped join. A1-VB runs ventrad, splitting off A1-VB-Vi with A1-VB continuing ventrad with T3-VL lateral before A1-VB splits to join T3-VLT from anterior and A1-VLT to posterior; A1-VC present, branching off A1-VB. Remaining A2..8-S with just A n -DB and A n -VB branches. A n -DB runs mediad and slightly ventral, meeting A n - DLT branches from anterior and posterior in Y-shaped junction. A n -DLT typically small and sinuous. 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. Large A n -VB-Vi also typical for all segments; directional notation in 3D supplemental models for visceral tracheae is to denote relative directions for clarity and not an assessment of homology. A n -VB with small A n -MLT branching ventrad from A n -S, directly posteriad toward proceeding posterior segment, linking with A n -VB. As with A n -DLT, A n -MLT often small and hard to distinguish, occasionally not visible on one side of the specimen but likely present.</p></div>	https://treatment.plazi.org/id/038D8781FF7220DCFCC6FD94A35CFB75	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFB92019FEF3FD10A147FDA6.text	038D8781FFB92019FEF3FD10A147FDA6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Romalea microptera (Palisot de Beauvois 1817)	<div><p>Romalea microptera</p><p>“Eastern lubber grasshopper”</p><p>Figures 63 (lateral), 64 (dorsal, ventral)</p><p>Plates 38 (lateral), 39 (dorsal), 40 (ventral)</p><p>The tracheal structure of the lubber grasshopper  Romalea microptera features a substantial number of air sacs, integrated into the tracheal system, and discrete air cells distributed throughout the body, especially the head and thorax. Due to the complexity of the tracheal architecture of  Romalea, only spiracles and major tracheae are labeled in the plates. 3D models are provided in the supplementary digital data for further investigation.</p></div>	https://treatment.plazi.org/id/038D8781FFB92019FEF3FD10A147FDA6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFB92026FF0AFA9BA126FB99.text	038D8781FFB92026FF0AFA9BA126FB99.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tachycines asynamorous Adelung 1902	<div><p>Tachycines asynamorous</p><p>“Greenhouse camel cricket”</p><p>Figures 65 (lateral), 66 (dorsal, ventral)</p><p>Plates 41 (lateral), 42 (dorsal, ventral)</p><p>A single camel cricket,  Tachycines asynamorous, was scanned, but substantial fluid infilling of tracheae resulted in assessing homology of many branches difficult. Although of suboptimal quality, this specimen was included because of its placement within  Orthoptera and the relative modifications of the mesothoracic tracheae, specifically how they relate to modifications for sound reception. Identifiable tracheae are labeled and described here but should be considered preliminary.</p><p>Our mesothorax assessments differ from Ander (1939), particularly with H-DCT and H-VCT. Ander identifies most of the larger tracheae extending into head as multiple branches of H-VCT and the small branch as H-DCT, whereas we have determined the small dorsal branch is T2-VB, the next ventral branch is H-DCT, followed by H-VCT, which does have a split into H-VCT and a smaller anterior trachea that has T1-AL ventrad.</p><p>DESCRIPTION: HEAD: T1-DLT partially visible and likely extending into head capsule. H-DCT and H-VCT both thick, with dorsal-ventral connection likely but not visible. Several air spaces visible, likely preservational artifacts. H-VCT splits into two branches, large dorsal one and smaller ventral, which proceeds anteriorly with T1-AL ventrad.</p><p>THORAX: T2-S with five branches: T2-DB, T2-VB, H-DCT, H-VCT, and T1-PL. T2-DB running dorsad and anteriad, extending into head capsule after broad curve along prothorax. H-DCT thick, extending anteriad through prothorax into head capsule. H-VCT splitting into two branches: large dorsal trachea, extending anteriad into head, and a smaller ventral branch, extending anteriad into head with T1-AL ventrad. T2-VC not visible but likely present; many ventral thoracic tracheae infilled with fluid. T2-DB runs dorsad, with T2-AL branching in hairpinlike turn ventrad into T2-L. Remaining branches of T2-DB not visible. T2-VB partial, with T-shaped intersection with T2-VLT. H-DCT and H-VCT both large, of similar size, extending into head as described above. T1-PL running directly ventrad, partial, likely joining with T1-AL but not visible in this scan. T3-S branches largely infilled and difficult to differentiate, appearing to include: T3-DB, T3-VB, T3-lvl, T2-PL, and T3-suf. T3-DB partial, with T3-AWL visible, leading to T3-AL, dorsad before curving sharply ventrad and posteriorly into hind leg. T3-VB running ventrad, joining with T3-VLT. T3-lvl extending mediad, joining with T3-VLT, with small branch posteriad connecting with T3-PL. T3-suf partial, extending mediad and slightly anteriorly. T3-VL possibly visible on specimen left side; T3-VL likely present but infilled.</p><p>ABDOMEN: A1..8-S present. A1-S modified from subsequent abdominal segments, with large ventral T3-PL branch. A n -VB visible on most segments, leading to partially infilled A n -VLT. A n -DB likely present but infilled and not visible. A n -DLT likewise probably present but not visible, likely infilled. Several visceral tracheae present from all A n -S.</p></div>	https://treatment.plazi.org/id/038D8781FFB92026FF0AFA9BA126FB99	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6020C1FEEDFCE8A3CAFA3A.text	038D8781FF6020C1FEEDFCE8A3CAFA3A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tenodera sinensis Saussure 1871	<div><p>Tenodera sinensis</p><p>“Chinese mantis”</p><p>Figures 97, 98 (lateral, anterior, posterior); 99, 100 (dorsal, anterior, posterior); 101, 102 (ventral, anterior, posterior)</p><p>Plates 58 (lateral), 59 (dorsal), 60 (ventral)</p><p>DESCRIPTION: Pronotum, metathorax, and anterior portion of abdomen with air-filled spaces, likely alimentary canal and transient as not connected to tracheae (see fig. 103).</p><p>HEAD: Tracheae leading to appendages are labeled; however, the head of  Tenodera features a complex network of intersecting tracheae and air sacs. Readers are encouraged to review the 3D models in the supplementary digital data.</p><p>THORAX: Thorax with numerous elongate, bandlike visceral tracheae throughout, often obscuring views. Exploration of 3D models in digital supplementary data is encouraged. T2-S positioned posterior to characteristically elongate pronotum, with four branches, crowded close together: T2-DB, T2-VB, H-DCT, H-VCT. T2-DB runs mediad and slightly dorsad, with split off to T2-AWL posteriad; T2-DB continues briefly before joining T1-DLT anteriad and T2-DLT posteriad in Y-shaped junction. T2-DLT running in shallow arc ventrad, with unusual condition of linking with T3-S via connection to T2-PL. T2-AWL arcing dorsally and posteriad, with T2-Wbr branching dorsad at apex of curve; T2-AWL continues as T2-AL, extending posteriad into midleg; T2-Wbr with dorsad T2-W-c-r branch, remaining T2-Wbr connecting to T2-PWL from T3-S. T2-VB short, running directly anteriad, bifurcating into T1-PL anteriad and T2-VLT posteriad; T2-VC likely present off T2-VLT; T1-PL extending through mesothorax and prothorax with ventral curve into foreleg at base of forecoxa. T2-VL branching close to origin of T2-VLT, extending straight through coxa into midleg. H-DCT beginning ventrad and H-VCT dorsad, opposite of usual arrangement; both tracheae extending directly anteriad, switching dorsoventrally to typical positions at anterior margin of mesothorax. (See Discussion section for homology assessment of these tracheae.) H-VCT with T1-AL running ventrad just anterior of dorsoventral H-DCT/H-VCT switch. T3-S with n connections: T3-DB, T3-AWL, T3-VB, T2-PWL. T3-DB mediad, curving dorsally and posteriad as T3-DLT, connecting directly to A2-S with no connection to A1-S; A1-DB absent. T3-AWL runs just lateral of T3-DB, arcing posteriad, with T3-Wbr branch; remaing T3-AWL continuing as T3-AL into hind leg. T3-Wbr with T3-W-c-r, continuing posteriad to connect to A1-S via T3-PWL. T3-VB runs ventrad and posteriad, connecting with A1-S via T3-VLT; T3-VL branching from T3-VLT close to T3-S. T2-PWL short, extending anteriad, splitting into T3-PL ventrad and T2-Wbr anteriorly. Metathorax with several networked visceral tracheal branches, especially along venter; most notable is T3-Ty for hearing. (Additional tracheae of unknown homology enter the legs, similar to  Blattodea [roaches], which require further investigation.)</p><p>ABDOMEN: A1..8-S present and functional. A1-S connections atypical from remaing spiracles; A1-S with only two connections: T3-PWL and A1-VB. A1-DB absent; A1-MLT possible but not discernible in this scan. T3-PWL runs from anterior, linking with T3-S via T3-Wbr. A1-VB ventrad, following sternite, forming A1-VC; T3-VLT joining with A1-VB, A1-VLT extending posteriad from T3-VC. T3-VL branching from A1-VC. Remaining A2..8-S connections largely similar, with A n -DB extending dorsad, joining with A n -DLT; A n -MLT extending anterior-posteriorly through length of abdomen; and A n -VB ventrad, joining with A n -VLT. Nearly all tracheae bandlike, with visceral branches throughout abdomen. A4-S and A5-S with numerous sinusoidal visceral tracheae.</p></div>	https://treatment.plazi.org/id/038D8781FF6020C1FEEDFCE8A3CAFA3A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF94203CFF0BFAB9A17AFD64.text	038D8781FF94203CFF0BFAB9A17AFD64.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Timema californicum S. H. Scudder 1895	<div><p>Timema cf. californicum</p><p>“ California timema (walkingstick)”</p><p>Figures 79, 80 (lateral, anterior, posterior); 81, 82 (dorsal, anterior, posterior); 83, 84 (ventral, anterior, posterior)</p><p>Plates 49 (lateral), 50 (dorsal), 51 (ventral)</p><p>The tracheal morphology of  Timema is nota- ble for the proliferation of comparatively thin tracheae that form networks throughout the body. These networks make assessment of homology difficult especially in the thorax, where differentiating similarly networked tracheae from other taxa is unclear. Identifiable homologous tracheae are labeled here, and incorporation of data from some of the 20 or so other  Timema species could help to resolve ambiguous structures. Notably, T2,3-DLT are absent in  Timema, with the additional absence of the corresponding T2,3-DB; however, the networked nature of tracheae elsewhere likely compensates for the lack of dorsal longitudinal connections. Although distantly related, the  Timema thorax is rather reminiscent of termites, with (possibly) dual ventral commissures, which may be convergent.</p><p>DESCRIPTION: HEAD: Head tracheae with several networked interconnections dorsoventrally; due to three-dimensional nature of head tracheal architecture, readers are encouraged to refer to models in supplementary digital data. Prominent H-DCT-VCT-Loop at anterior margin of prothorax, with smaller H-DCT and H-VCT branching from dorsal and ventral portions of loop. H-DCT extending anteriad and slightly dorsad, splitting into dorsal branch to H-DX intersection near vertex; and anteriad H-DCT-Ant branch, joining with ventral H-VCT-Ant branch and extending into H-Ant. H-DX with anteriad branches extending laterally and ventrad, reconnecting with H-VCT-Ant as H-DX-VCT-Loop. H-VCT runs anteriad, with H-VCT-Ant branching anterodorsally to join with H-DCT-Ant. H-VCT continues anteriad with H-Mx branching ven- trad; H-Mx with H-VC before branching to H-MxPlp. H-VCT with continued anterodorsal curve, with H-Md branching anteriad; H-VCT joining with H-DCT via H-VCT-Ft-Loop; H-FtLbr branching ventrad near anterior apex of H-VCT-Ft-Loop.</p><p>THORAX: T2-S with three branches: H-DCT, H-VCT, and T2-AWL. H-DCT runs directly anteriad toward head; small T1-Cx ventrad in middle of prothorax. H-VCT likewise running anteriad, with T1-AL ventrad at posterior margin of prothorax; T1-PL not present. T1-AL with medial branch to T1-VX. T1-VX branching into three anteriad tracheae, extending toward head as H-VLT, with right side H-VLT fusing with H-VLT-Med anteriad of cervix. T2-AWL runs ventrad and mediad before turning dorsally and posteriad in an S-shaped curve; small T2-DB and T2-VB branching dorsad and ventrally (respectively) from T2-AWL. T2-DB linking with apparent T1-DLT anteriad; T2-DLT not visible. T2-VB runs ventrad at bottom of T2-AWL S-curve, bifurcating into anterior and posterior branches, both leading to network of tracheae forming numerous ventral commissures. Remainder of T2-AWL bifurcating into T2-Wbr and T2-AL; T2-Wbr directly posteriad, linking with T3-S via T2-PWL; T2-AL posteriad and ventrad, linking with T2-PL and extending into midleg. T3 with three branches: T2-PWL, T3-AWL, and T3-VB. T2-PWL runs anteriad, splitting into T2-Wbr dorsally and T2-PL ventrad; T2-PL joining with T2-AL and extending into midleg. T3-AWL runs dorsad, curving posteriorly and splitting into T3-Wbr posteriad and T3-AL ventrad; T3-AL joining with T3-PL and extending into hind leg. T3-VB splitting into four tracheae close to T3-S; at least two of these extending into network of ventral commissures; several X-shaped commissure intersections present.</p><p>ABDOMEN: A1..8-S present, short A n -SB spiracular branch present on all A1..8-S. A1-S branching pattern slightly modified from remaining A2..8-S, with T3-PWL running anteriad, splitting into T3-Wbr anteriad and T3-PL ventrad, with T3-PL joining with T3-AL and extending into T3-L. A1..8-MLT present, with A1..5-DB branching dorsad from A n -MLT; A6..8-DB branching directly from A6..8-SB. All A1..6-DB linking with thin, sinuous A1..6-DLT along dorsum; A n -DC not present. A7,8-DLT substantially larger, with fanlike morphology expanding into highly tracheated hind- and midgut “appendices” (Shelomi et al., 2015). A1..8-VB present, extending to link with sinuous A n -VLT along venter; A2,3,6,7,8-VC present; given distribution, other A n -VC likely present but not visible. Numerous visceral tracheae, most notably forming asymmetric connectives A4-A5-Vi-AsymC and A5-A6-Vi-AsymC; A7-Vi-VC also present.</p></div>	https://treatment.plazi.org/id/038D8781FF94203CFF0BFAB9A17AFD64	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFC42064FF00FA55A221FF46.text	038D8781FFC42064FF00FA55A221FF46.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tricholepidion gertschi Wygodzinsky 1961	<div><p>Tricholepidion gertschi</p><p>“Relic Silverfish”</p><p>Figures 12 (lateral), 13 (dorsal, ventral)</p><p>Plates 4 (lateral), 5 (dorsal, ventral)</p></div>	https://treatment.plazi.org/id/038D8781FFC42064FF00FA55A221FF46	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFFF205AFF2DFB7CA232FD64.text	038D8781FFFF205AFF2DFB7CA232FD64.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trigoniophthalmus alternatus (Silvestri 1904)	<div><p>Trigoniophthalmus alternatus</p><p>“Jumping bristletail”</p><p>Figures 8 (lateral), 9 (dorsal, ventral)</p><p>Plates 1 (lateral), 2 (dorsal), 3 (ventral)</p><p>Trigoniophthalmus, as the representative of the most basal order in Class  Insecta, is notable in its tracheal architecture by a complete lack of longitudinal connections between spiracles. This taxon is critical to understanding the apparent insect ground-plan tracheal structure.</p><p>The thoracic tracheae of most taxa are supplied by both T2-S and T3-S, with T2-DLT connecting longitudinally. In  Trigoniophthalmus, however, T3-S appears to only supply T3-L and its associated coxa. Branches from T2-S extend posteriorly into the metathorax, nominally supplied by T3-S, including an apparent T3-DLT. The naming of T3-DLT here is an instance of using the positional criterion in homology (sensu Remane, 1952), in that the connectivity of this trachea is not consistent with other taxa; however the position suggests its identification as T3-DLT (de Pinna, 1991). T2-VB extends posterior, almost in the opposite direction of the anterior-reaching cephalic branches. While the position of T2-VB may not appear to be “ventral” in nature, a comparison of its relative position in the hump-backed  Trigoniophthalmus and in particular its connection to T2-L with apterygote taxa from  Zygentoma and  Dermaptera demonstrates its homology as a ventral branch. Several tracheae range anteriorly or posteriorly beyond segment boundaries, most prominently A5-DB-DVi, which extends posteriorly past A6-S, reaching A7-S; and A7-Cr, which likewise extends posteriorly from A7-S into both the cerci and terminal filament. However, tracheae for most segments, particularly in the abdomen, remain restricted to their individual segment, placing  Trigoniophthalmus among the simplest tracheal body plans in this study and unique in its lack of longitudinal connections. This corroborates observations made by Palmén (1877) and reviewed by Dittrich and Wipfler (2021).</p><p>DESCRIPTION: HEAD: Characteristically arched thorax, pronotum covering much of head, making boundary between thoracic tracheae and head tracheae rather indistinct. T2-S at anteriormost margin of mesothorax, covered by overhanging tergum, shown in figure 10. H-DCT very thick, with three branches: H-Ic, extending anteriorly and dorsad; H-Oc-Ant anterior and medially before dividing into H-Oc toward midline and H-Ant laterally; H-Mx proceeds anteriad with H-MxPlp branching ventrally and anteriad. H-VCT very thick, with two branches: H-Lbm ventrad, extending into H-LbmPlp; H-Md anteriad, crossing over H-Md from opposite side of head but not connecting.</p><p>THORAX: T2-S present, with five tracheae: H-DCT and H-VCT leading anteriad directly into head, beginning at T2-S and running parallel for length of prothorax; T2-DB leading dorsad; T2-VB and T2-L directly posteriad. H-DCT with no thoracic branches; H-VCT with T1-L branching ventrally; T1-VC present, extending from T1-L. T2-DB-Vi branches just dorsal of T2-S, extending anteriorly; T2-DB continuing dorsad, dividing into what appear to be anterior T1-DLT and posterior T2-DLT near midline; T1-DLT and T2-DLT extend into viscera with no connections to neighboring spiracles. T2-L running posteriad and ventrally, into midleg; large T2-L-Vi and smaller T2-VC divide from T2-L posterior from T2-S. T2-VC joins with anterior-arching T3-VC to form T2-VX intersection. T2-L-Vi extends posteriad into metathorax, with apparent dorsal T3-DLT, T3-Cx ventral, and numerous small visceral tracheae. T3-S ventral to and much smaller than T2-S, with two tracheae: T3-L and T3-VB. T3-L extending dorsad with small T3-L-Vi before arcing ventrad into hind leg; T3-VB directly toward midline with small T3-VC branch that arcs anteriorly to join T2-VC at T2-VX.</p><p>ABDOMEN: A1..7-S present, all located ventrally. No longitudinal connections present; figure 11 with representative abdominal segment. A n -S with A n -DB running dorsad along arc of body wall, turning inward toward midline of body but not forming DC; A n -VB branching dorsally for a short distance before arcing ventrad toward midline. A n -VC absent. A n -DB with visceral tracheae A n -DB-MVi, branching anterior and medially halfway up body; A n -DB-DVi branches dorsally toward tergal wall. A5-DB-DVi extends posteriorly to 8th abdominal segment; A7-DB-DVi extends posteriorly past 8th abdominal segment, dividing into A-TF and A-Cr; neither A5-DB-DVi nor A7-DB-DVi connect to spiracles of other segments. A-TF with two tracheae per side (4 total); A-Cr single trachea per cercus.</p></div>	https://treatment.plazi.org/id/038D8781FFFF205AFF2DFB7CA232FD64	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FF6A20D2FF64FA54A474F8AE.text	038D8781FF6A20D2FF64FA54A474F8AE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zootermopsis angusticollis (Hagen 1858)	<div><p>Zootermopsis angusticollis Soldier caste</p><p>“Dampwood termite”</p><p>Figures 118 (lateral), 119 (dorsal), 120 (ventral)</p><p>Plates 73 (lateral), 74 (dorsal), 75 (ventral)</p><p>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.</p><p>THORAX: T2-S with four branches: H-DCT, H-VCT, T2-DB, and T2-VB. H-DCT mediad, curving anteriad toward large head capsule; short T1-DLT from T2-DB connecting before curve. H-VCT like H-DCT, with ventrad T1-AL near curve toward anterior. T2-DB short, with T2-AWL branch running posteriad close to T2-S; T2-DB continues mediad with Y-shaped bifurcation of short T1-DLT anteriad toward H-DCT and T2-DLT posteriad toward T3-S. T2-AWL arcing posteriad, splitting in to T2-AL and T2-Wbr at apex of arc; T2-Wbr with small T2-W-c-r and T2-W-cu-a, connecting with T2-PWL posteriorly; T2-AL posteriad and ventral, joining T2-PL and extending into midleg. T2-VB runs directly mediad, bifurcating into T2-VC1 mediad and T2-VLT arcing ventrally and posteriad along sternite. T2-VC1, the first of two mesothoracic ventral commissures, proceeding mediad with T1-PL branch directly ventrad and smaller connection anteriad to T1-VX. T1-VX with H-VCT extending anteriad into head; single branch on specimen left side but likely present on right—see T2-VC2 branching mediad from T2-VLT anteriad from T2-VLT’s curve dorsad toward T3-S; T2-VLT with additional T2-Cx branch, larger on right side than on left. T3-S with four branches: T2-PWL, T3-DB, T3-AWL, and T3-VB. T2-PWL from anterior, connecting T2-PL and T2-Wbr from T2-S. T3-DB mediad, joining with T2-DLT and T3-DLT in flat Y-shaped junction. T3-AWL mediad briefly, arcing posteriad and dorsally, splitting into T3-AL ventrad into hindleg and T3-Wbr posteriad, joining with A1-S via T3-PWL. T3-Wbr with small T3-W-c-r anterior and T3-W-cu-a posterior. T3-VB runs directly ventrad, with T3-VLT splitting posteriorly just ventrad of T3-S. T3-VB continuing ventrad, with T2-VL branch ventrad and posteriad, extending into midleg; remaining T3-VB arcing medially to form T3-VC1; T2-VLT joining with T3-VB near this arc. T3-VLT ventrad, arcing posterior along sternite before continuing anteriad toward A1-S, with T3-VC2 mediad near apex of this arc. T2-VLT and T3-VLT both with two ventral commissures: posterior T2-VC2 and anterior T3-VC1 along same section of T2-VLT; posterior T3-VC2 and A1-VC both along same section of T3-VLT.</p><p>ABDOMEN: A1..8-S present. Short A n -SB possible on several segments. A1-S with three branches: T3-PWL, A1-DB, and A1-VB; remaining A2..8-S with just A n -DB and A n -VB branches. For A1-S, T3-PWL from anterior, completing T3-Wbr link from T3-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; A8-VB split into A8-VC1 anterior and A8-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. A5..8-MLT slightly modified, connecting nearly directly with subsequent A n -S. As with A n -DB, A n -VB-Vi extend throughout abdomen; A7-VB-Vi and A8-VB-Vi extend mediad, crossing middle of body and extending anteriad past A4-S.</p></div>	https://treatment.plazi.org/id/038D8781FF6A20D2FF64FA54A474F8AE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFD92079FEF1FB4EA434FC27.text	038D8781FFD92079FEF1FB4EA434FC27.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zorotypus hubbardi Caudell 1918	<div><p>Zorotypus hubbardi</p><p>“Hubbard’s angel insect”</p><p>Figure 42 (lateral, dorsal)</p><p>Several  Z. hubbardi specimens were collected from Florida. Unlike the other taxa, where specimens were frozen alive, these specimens died during transport and were frozen upon arrival. Likely due to their untimely demise, tracheae suffered from fluid infilling before scanning, resulting in obscured morphology. Several specimens were scanned, and the best example was chosen for inclusion here.  Zoraptera are an excellent candidate for further micro-CT study of better-preserved specimens.</p><p>DESCRIPTION: HEAD: H-DCT and H-VCT in close contact, proceeding anteriad. H-Ant off H-DCT, no other head tracheae visible in this scan.</p><p>THORAX: T2-S with several branches visible on specimen right side but only two determinable: T2-VB and T2-CT. T2-VB ventrad; T2-VC present. T3-S visible with T3-DLT extending dorsad and posteriorly to link with A1-S. Other thoracic tracheae partial, determined from examining volume cross sections to establish position in the body.</p><p>ABDOMEN: A1..7-S discernible.</p></div>	https://treatment.plazi.org/id/038D8781FFD92079FEF1FB4EA434FC27	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
038D8781FFFA2064FEFEFB10A144FC6F.text	038D8781FFFA2064FEFEFB10A144FC6F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zygentoma	<div><p>ORDER  ZYGENTOMA</p><p>There are very few micro-CT studies of  Zygentoma to date; most notably, synchrotron radiation micro-CT of  Tricholepidion gertschi heads has been used to investigate  Zygentoma morphology for systematic study (Blanke et al., 2014), but not tracheae. This genus is particularly significant given the disputed placements within and near  Zygentoma (Engel, 2006; Blanke et al., 2014).  Zygentoma are relevant in general as a sister group to the pterygotes, and indeed they share several tracheal synapomorphies. Three specimens from two orders were scanned here:  Thermobia domestica and  Lepisma saccharinum from  Lepismatidae, and the relict silverfish  Tricholepidion gertschi from  Lepidotrichidae . Scan resolutions were within a 2.4 µm 3 /voxel range, with  Lepisma at the highest resolution of 4.8 µm 3 /voxel and  Thermobia with the lowest resolution at 7.2 µm 3 /voxel. Resolving the smallest tracheae at these resolutions tends to be more a function of specimen preservation rather than scanning parameters. Although the level of detail varies slightly between these scans, sufficient resolution was achieved to locate small visceral tracheae in all three specimens, indicating suitability for comparative purposes. Although the zygentoman head morphology is similar among the specimens, in the sections below  Lepidotrichidae thoracic and abdominal detail are described separately due to substantial differences.</p><p>Tricholepidion appears to possess only two pairs of functional abdominal spiracles, A7-S and A8-S. The elongate, sinusoidal trunks beginning at A7-S and extending anteriorly to the thorax are not present in the two lepismatid specimens. If  Tricholepidion is indeed basal to  Lepismatidae, it is possible that these trunks are plesiomorphic, and a loss of these tracheae in  Thermobia and  Lepisma could be derived.</p><p>In both species scanned from  Lepismatidae, leg tracheae branch from T2-S and T3-S in an arc leading posteriorly into the legs, each with a small branch extending dorsad. Here we interpret these as T2-AWL and T3-AWL, with T2-AWba and T3-AWba branching dorsad. An alternative pattern is to designate these dorsal branches as simply T2,3-L-DVi, but the similarity of the branching pattern with both apterous and winged taxa included herein (e.g.,  Grylloblatta) strongly suggests homology with these wing base tracheae; Šulc (1927) interpreted these as possible wing tracheae.</p><p>DESCRIPTION: HEAD: H-DCT with two branches: H-Ft dorsal; H-Ant-Lbr anteriad and slightly ventral, dividing into H-Ant and H-Lbr close to base of antenna. H-VCT with three branches, dividing at approximately same location: small H-Lbm extends ventrally; H-Mx continues anteriorly before proceeding ventrad; larger H-OcMd extends anteriorly. H-VC absent in  T. gertschi but present in both  Lepismatidae . H-Oc branching dorsally, H-Md continues anteriorly and ventrad, following shape of mandibular sclerite.</p><p>THORAX:  Tricholepidion with significant differences among  Zygentoma, notably T2-DLT and T3-DLT elongate, with dorsal visceral branches similar to T2-DB and T3-DB; a complete description of the thorax of  Tricholepidion thorax is below. For both  Lepismatidae: T2-S with three branches: T2-CT, T2-AWL, and T2-VB. T2-CT very thick, running directly anteriad with T2-DB just anterior to T2-S; T2-CT dividing into H-DCT and H-VCT in prothorax. Thinner T2-AWL arcing posteriorly, slightly dorsad, with thin but prominent dorsad and posteriorly arcing T2-AWba. T2-ACx present with T2-VC, remainder of T2-AWL extending into midleg as T2-L. T2-DB dividing into T2-DLT, extending in posterior arc and T1-DLT, ending blind in prothorax. Small T1-Pn present just anterior of H-DCT/H-VCT split. T1-L branching ventrally from H-VCT; T1-L with T1-VC branch present; T1-ACx from T1-VC; T1-PCx from T1-L. Small T2-VB with T2-VB-Vi. T2-L two branches: prominent, dorsad and posteriorly arcing T2-L-DVi.  Thermobia with T2-PCx from T2-L, note that T2-PCx is from T2-VB in  Lepisma . T2-VC with connection from T3-AsymC medially, originating from left side in  Thermobia and right side in  L. saccharinum . T3-S with three branches: T3-DB, T3-AWL, and T3-VB. T3-DB directly dorsal with Y-shaped bifurcation to T2-DLT from anterior and T3-DLT toward A1-S. T3-AWL posteriad with T3-AWba, longer in  Lepisma than  Thermobia . T3-PCx from T3-L in  T. domestica, T3-VC in  L. saccharinum . T3-VB ventral and slightly inward with AsymC from left side of  T. domestica, right side of  L. saccharinum, anteriad to connect with T2-VC.</p><p>ABDOMEN:  T. gertschi with significant differences to other  Zygentoma, including apparent lack of functional abdominal spiracles (except A7-S and A8-S), and a sinusoidal A7-VLT extending anteriorly over several segments; a complete description of abdomen is below. For  Lepismatidae: A1..8-S present, all located laterally. A n -SB present for segments in middle of abdomen but variable; see detailed descriptions. A n -S (or A n -SB) with A n -DB and A n -VB present. A n -DB running dorsad, bifurcating into Y-shaped junction with A n -DLT anteriad and posteriad; all abdominal segments connected via dorsal trunk. A n -VB running ventrad, connecting with opposite side via thin A n -VC; A1..7-VC present in  T. domestica, A4..7-VC not visible in  L. saccharinum scan but likely present; A8-VC absent. A n -DB-Vi and A n -VB-Vi numerous, see sections below for details. A8-DB and A8-VB large, curving medially before proceeding posteriad; A8-DB with A-TF and A-Cr in  T. domestica but not visible in  L. saccharinum scan (presence/ absence ambiguous).</p></div>	https://treatment.plazi.org/id/038D8781FFFA2064FEFEFB10A144FC6F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Herhold, Hollister W;Davis, Steven R;Degrey, Samuel P;Grimaldi, David A	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
