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Struthio camelus spatzi Stresemann, 1926: 139. Holotype: Egg, Museum für Naturkunde, Berlin, ZMB B.1180a (not examined). Referred specimen: Adult male skeleton ZMB 36879 Stresemann, 1927: 135. Remark: The skeleton ZMB 36879 comes from one of three birds, referred to by Stresemann (1927) as ‘cotypes’, kept at that time in Berlin Zoo. Expanded diagnosis: Smaller than the living subspecies (with most dimensions 88–100%) except for the synsacrum (Supporting Information, Table S3), postacetabular pelvis (Supporting Information, Table S4), distal ends of the humerus (Supporting Information, Table S2), femur except for a smaller corpus diameter (Supporting Information, Table S5), and pedal phalanges III/1 and III/2 (Fig. 2; Supporting Information, Tables S9 and S10). Tarsometatarsus length S. c. syriacus). The ratio of the length of tibiotarsus to tarsometatarsus is> 1.15, compared with 1.11–1.14 in extant ostriches. Phalanges III/1 and III/2 are as in extant ostriches, with the respective total lengths of ~ 90 mm (in the lower range of S. c. camelus) and 60 mm (as in S. c. camelus). The ratio of the length of tarsometatarsus to phalanx III/2 is 7.1, compared with 7.6–9.0 (8.0–9.0 for males) in extant ostriches. The pattern of egg pores of the intermediate type is as described by Schönwetter (1927), Sauer (1968, 1972) and Mikhailov & Zelenkov (2020). crista plantaris mediana; ct, crista trochanteris; dl, foramen dorsale laterale; dm, foramen dorsale mediale; em, epicondylus medialis; fd, facies tibialis distalis; ff, facies femoralis; fg, facies glenoidalis; fl, fovea ligmenti capitis; fm, foramen plantare mediale; fp, facies tibialis proximalis; fpl, foramen plantare laterale; il, impressio ligamenti collateralis medialis longi; l, condylus lateralis; ll, labrum laterale; lm, labrum mediale; m, condylus medialis; p, processus procoracoideus; pc, processus craniolateralis (sternocoracoideus); pm, foramen plantare mediale; sf, semicondylus fibularis; sp, sulcus patellaris; st, semicondylus tibiofibularis; t, trochanter; tf, tuberositas femoralis; ti, tuberculum m. iliofibularis; tIII, trochlea metatarsi III; tIV, trochlea metatarsi IV; tl, trabecula lateralis; tt, tuberositas tibialis. For explanation of measurement tags, see Table 1. SKELETAL VARIATION AND SEXUAL DIMORPHISM Although the coefficients of variation of most measurements in the living ostriches do not exceed 6% (Supporting Information, Tables S2–S 13), some of them are distinctly higher as a result of variable ossification (intrinsic variation) or, in addition, some indeterminacy owing to the absence of exact measurement points when taking minimum width or depth measurements across continuous surfaces. A high intrinsic variability affects single details, such as the sternal interspinal width (CV = 7.4), pelvic angular width (CV = 10.1), the cranial (CV = 8.5) and caudal (CV = 12.1) interforaminal distances of the tarsometatarsus and the proximal width between the cotylar margins of phalanx III/1 (CV = 7.1). The indeterminacy and variability of measurement position might have contributed to the variation of the sternal postcostal width (CV = 9.6), pelvic postacetabular width (CV = 8.9), fibular corpus width (CV = 8.8), tarsometatarsal corpus depth (CV = 8.4) and many transverse measurements of phalanx III/2: corpus width (CV = 9.1), apical width (CV = 7.0), trochlear depth (CV = 7.1) and foveal width (CV = 7.1), with the foveal width also being highly variable in phalanx III/3 (CV = 9.5). The most variable and thus the least usable measurement turned out to be the interischiadic width (ischiadic span) of the pelvis (CV = 19.9), suggesting a strong intrinsic component of this variability in addition to the indeterminacy of the measurement point. With a notable exception for the length of the tibiotarsus (Supporting Information, Table S6) and, especially, the tarsometatarsus (Supporting Information, Table S8), most measurement values (71 of 95medians), including the transverse measurements of the tibiotarsus and tarsometatarsus, tend to be higher in females. However, the differences proved to be statistically significant or near-significant in only 14 measurements (Table 2). In ten of them, the ostrich females have consistently wider and/or deeper ends of limb bones, which suggests that the female skeleton (at least the appendicular skeleton) is more robust even if the males are taller owing to longer tarsometatarsi and tibiotarsi. Only three bones, namely the scapulocoracoid, humerus and pedal phalanx III/3, are significantly longer compared with males., Published as part of Elzanowski, Andrzej & Louchart, Antoine, 2022, Metric variation in the postcranial skeleton of ostriches, Struthio (Aves: Palaeognathae), with new data on extinct subspecies, pp. 88-105 in Zoological Journal of the Linnean Society 195 (1) on pages 95-97, DOI: 10.1093/zoolinnean/zlab049, http://zenodo.org/record/6530641, {"references":["Stresemann E. 1926. Die Vogelausbeute des Herrn Paul Spatz in Rio de Oro. Ornithologische Monatsberichte 34: 137 - 139.","Stresemann E. 1927. Ueber die einstige Verbreitung von Struthio camelus in Algerien. Ornithologische Monatsberichte 35: 135 - 136.","Schonwetter M. 1927. Die Eier von Struthio camelus spatzi Stresemann. Ornithologische Monatsberichte 35: 13 - 17.","Sauer EGF. 1968. Calculations of struthious egg sizes from measurements of shell fragments and their correlation with phylogenetic aspects. Cimbebasia A 1: 27 - 55.","Sauer EGF. 1972. Ratite eggshell and phylogenetic questions. Bonner Zoologische Beitrage 23: 3 - 48.","Mikhailov KE, Zelenkov N. 2020. The late Cenozoic history of the ostriches (Aves: Struthionidae), as revealed by fossil eggshell and bone remains. Earth Science Reviews 208: 103270."]}

Struthio camelus camelus Linnaeus, 1758 LITERATURE. — Dowsett (1993); Portier et al. (2002); Dowsett et al. (2013); Borrow & Demey (2014). STATUS. — Probably extinct in the wild., Published as part of Boano, Giovanni, Belemsobgo, Urbain, Silvano, Fabrizio, Hema, Emmanuel M., Belemsobgo, Aristide, Dimobe, Kangbéni & Pavia, Marco, 2022, An annotated checklist of the birds of Burkina Faso, pp. 27-107 in Zoosystema 44 (2) on page 32, DOI: 10.5252/zoosystema2022v44a2, http://zenodo.org/record/5999691, {"references":["DOWSETT R. J. 1993. - Afrotropical avifaunas: annotated country checklists, in DOWSETT R. J. & DOWSETT- LEMAIRE F. (eds). A Contribution to the Distribution and Taxonomy of Afrotropical and Malagasy birds Tauraco Research Report 5, Tauraco Press, Liege: 1 - 322.","PORTIER B., LUNGREN C. & OUEDA G. H. 2002. - Birding in Burkina Faso, more than just birdwatching. Dutch Birding 24: 127 - 141.","DOWSETT R. J., ATKINSON P. W. & CADDICK J. A. 2013. - Checklist of the birds of Burkina Faso: 1 - 12. https: // www. africanbirdclub. org / [accessed 14 October 2013].","BORROW N. & DEMEY R. 2014. - Field Guide of the Birds of Western Africa. II edition. Christopher Helm, London, 592 p."]}

The avian lung is highly specialized and is both functionally and morphologically distinct 30 from that of their closest extant relatives, the crocodilians. It is highly partitioned, with a 31 unidirectionally ventilated and immobilized gas-exchanging lung, and functionally decoupled, 32 compliant, poorly vascularized ventilatory air-sacs. To understand the evolutionary history of 33 the archosaurian respiratory system, it is essential to determine which anatomical 34 characteristics are shared between birds and crocodilians and the role these shared traits play 35 in their respective respiratory biology. To begin to address this larger question, we examined 36 the anatomy of the lung and bronchial tree of ten American alligators (Alligator 37 mississippiensis) and eleven ostriches (Struthio camelus) across an ontogenetic series using 38 traditional and micro-computed tomography (µCT), three-dimensional (3D) digital models, and 39 morphometry. Intraspecific variation and left to right asymmetry were present in certain 40 aspects of the bronchial tree of both taxa but was particularly evident in the cardiac (medial) 41 region of the lungs of alligators and the caudal aspect of the bronchial tree in both species. The 42 cross-sectional area of the primary bronchus at the level of the major secondary airways and 43 cross-sectional area of ostia scaled either isometrically or negatively allometrically in alligators 44 and isometrically or positively allometrically in ostriches with respect to body mass. Of fifteen 45 lung metrics, five were significantly different between the alligator and ostrich, suggesting that 46 these aspects of the lung are more interspecifically plastic in archosaurs. One metric, the 47 distances between the carina and each of the major secondary airways, had minimal 48 intraspecific or ontogenetic variation in both alligators and ostriches, and thus may be a 49 conserved trait in both taxa. In contrast to previous descriptions, the 3D digital models and CT 50 scan data demonstrate that the pulmonary diverticula pneumatize the axial skeleton of the 51 ostrich directly from the gas-exchanging pulmonary tissues instead of the air sacs. Global and 52 specific comparisons between the bronchial topography of the alligator and ostrich reveal 53 multiple possible homologies, suggesting that certain structural aspects of the bronchial tree 54 are likely conserved across Archosauria, and may have been present in the ancestral 55 archosaurian lung.

Struthio cf. karatheodoris Forsyth Major, 1888 TYPE MATERIAL. — Holotypefemur, from Samos 1. Said to be in the Barbey collection (Lambrecht 1933: 100); present location unknown (Mlíkovský 2002). TYPE HORIZON. — Late Miocene (MN 11) (Mlíkovský 2002). TYPE LOCALITY. — Samos 1, Samos Island (southeastern Greece). NEW DIAGNOSIS. — Fossil late Miocene ostrich of large size, only smallerthan S. novorossicus, more bipedal, with better developed fourth toe; more robust pedal phal. in comparison to S. camelus. OTHER MATERIALEXAMINED. — Kalimantsi:tmt. dex. dist. (epiphysis, completearticular end) ad., NMNHS 16371 (Fig. 2); fromthe middle Turolian of the fossiliferous area near Kalimantsi village (Blagoevgrard Region, 43.17’N, 27.44’E; UTM grid: NH 59; c. 300 m a.s.l.). — Hadzhidimovo: phal. 1 dig. 3 ped. sin. (almost complete) ad., NMNHS 16372 (Fig. 2); from the base of the middle Turolian or the early-middle Turolian boundary at Tumbichkite (HD-2), near the town of Hadzhidimovo (Blagoevgrard Region, 41.30’N, 23.52’E; UTM grid: GM 30; 500 m a.s.l.). Both sites represent the richest Bulgarian localities of the so-called “ Hipparion -fauna”. The two specimens are morphometrically similar to each other, and theirrelatively closechronostratigraphic position indicates that they are referrable to the same fossil species. Both specimens were collected in the 1980’s by D. Kovachev. LOCALITIES. — Thefossiliferous area around Kalimantsi village in the Middle Struma basin covers a large region (c. 6 km 2) of late Miocene depositsincluding more than 10 known fossil mammal localities (labelled KM sites). Recentbiostratigraphicrevisiondemonstrates thatall KM sites are of Turolian age. KM-1, belonging to the newly established Gradishte lithocomplex, is likely to yield a fauna of early Turolian age, while other KM faunas (referred to the Strumyani Genetic Lithocomplex) are consideredto belong tothe middle Turolian (Tzankov et al. 2005; Spassov et al. 2006). Themiddle Turolianfauna of KM is presented by c. 10 different localities, which are faunally dominated by Gazella sp., Cremohipparion mediterraneum (Roth & Wagner, 1855) and Hippotherium brachypus (Hensel, 1862), and which also contain Tragoportax cf. amalthea, Palaeoreas lindermayeri (Wagner, 1848), Helladotherium duvernoyi (Gaudry & Lartet, 1856), Bohlinia attica (Gaudry & Lartet, 1856), Adcrocuta eximia (Roth & Wagner, 1854), Mesopithecus pentelicus Wagner, 1839, andother species (Spassov et al. 2006). NMNHS 16371 comes from horizons of middle Turolian age and is probably slightly younger than NMNHS 16372. Hadzhidimovo is a late Miocene site with three fossil vertebrate localities: HD-1, HD-2 (Tumbichkite) and HD-3. The main locality, HD-1 (from which more than 30 mammal species have been reported), is dated to the MN 11/12 boundary (Spassov 2002). The other two localities could be of slightly different age; HD-2 is several tensof metres higher thanthe level of HD-1. The fauna of the locality consists of Gazella sp., Palaeoreas lindermayeri, Tragoportax rugosifrons (Schlosser, 1904), Helladotherium duvernoyi, Hipparion / Cremohipparion spp. (probably H. brachypus and C. mediterraneum), Ancylotherium pentelicum (Gaudry & Lartet, 1856), Deinotherium gigantissimum Stefanescu,1892 and Choerolophodonpentelici (Gaudry & Lartet, 1856). Theoccurrence of P. lindermayeri and C. pentelici further suggeststhat HD-2 is younger than HD-1 (Geraads et al. 2003; Markov 2004)., Published as part of Boev, Zlatozar & Spassov, Nikolaï, 2009, First record of ostriches (Aves, Struthioniformes, Struthionidae) from the late Miocene of Bulgaria with taxonomic and zoogeographic discussion, pp. 493-507 in Geodiversitas 31 (3) on pages 498-500, DOI: 10.5252/g2009n3a1, {"references":["FORSYTH MAJOR C. 1888. - Sur un gisement d'ossements fossiles dans l'ile de Samos, contemporains de l'age de Pikermi. Comptes rendus de l'Academie des sciences, Paris, 107: 1178 - 1181.","MLIKOVSKY, J. 2002. - Order Struthioniformes Latham, in Cenozoic Birds of the World. Part 1: Europe. Ninox Press, Praha: 61 - 62.","TZANKOV TZ., SPASSOV N. & STOYANOV K. 2005. -  Neogene-Quaternary Paleogeography and Geodynamics of the Region of Middle Struma River Valley (South- Western Bulgaria) . Publishing House of the South- Western University \" N. Rilski \", Blagoevgrad, 199 p. (in Bulgarian, with English summary).","SPASSOV N., TZANKOV T. & GERAADS D. 2006. - Late Neogene stratigraphy, biochronology, faunal diversity and environments of South-West Bulgaria (Struma river valley). Geodiversitas 28 (3): 477 - 498.","SPASSOV N. 2002. - The Turolian Megafauna of West Bulgaria and the character of the late Miocene \" Pikermian biome \". Bollettino della Societa Paleontologica Italiana 41 (1): 69 - 81.","GERAADS D., SPASSOV N. & KOVACHEV D. 2003. - Palaeoreas lindermayeri (Wagner, 1848) from the upper Miocene of Bulgaria, and a revision of the species. Geodiversitas 25 (2): 405 - 415.","MARKOV G. 2004. - [The Fossil Proboscideans (Proboscidea, Mammalia) of Bulgaria]. Ph. D. Thesis. NMNHS, Sofia: 1 - 217 (in Bulgarian with English summary)."]}

The morphogenesis of the crocodilian and avian columella auris is analysed, using Alligator and Struthio as models that are studied in detail, and Crocodylus, Gallus, and Coturnix as models for comparative purposes. This investigation was undertaken to address controversies emanating from previous studies concerning the identity of archosaurian hyal elements, and to clarify confounding nomenclature that has been used for both topographic and phylogenetic purposes. We found that confusion in the identification of the reptilian and avian hyoid arch derivatives is attributable to: (1) redundant and/or inconsistently applied topographical terminology; (2) mixing of topographical and phylogenetic terminology; (3) interpretation in a recapitulatory framework; and (4) failure to consider that developmental pathways of recent archosaurians (reptilians and birds) might be divergently modified. Our analysis identifies errors in previous interpretations, misinterpretations of prior work, and histological observations that have been idealized. The most controversial issues are addressed – the derivation of the supracolumellar process; the concept of the laterohyal component; and segmentation of the distal part of the reptilian and avian columella auris. Based on new data, we suggest that the osteichthyan infrapharyngohyal, suprapharyngohyal, epihyal, and ceratohyal are not applicable terms for reptilian and avian columellar components from a phylogenetic perspective, and should not be used as topographical terms either. We further propose that the original morphogenetic program of the hyoid arch has been overwritten in newly derived evolutionary pathways of crocodilians and birds, in which a new functional and morphological patterning of the hyoid arch, as a specific sound-transmitting apparatus, have evolved. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 384–410.

Struthio casuarius [spec. nov.] S. pedibus tridactylis, vertice palearibusque nudis. Casuarius. Olear. mus. 23. t. 13. f. 2. Alb. av. 2. p. 56. t. 60. Emeu. Dodart. mem. 377. Raj. av. 36 Clus. exot. 98. Bont. jav. 71. Will. ornith. 105. t. 25. Worm. mus... Jonst. av. 172. t. 56. Aldr. orn. l. 20. t. 541. Habitat in Asia, Sumatra, Molucca, Banda. Callus elevatus quasi diadema verticis. Palearia 2 sub collo; Collum a tergo nudum. Ungue intermedio majore ferit; ova punctis excavatis., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 155, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Struthio americanus [spec. nov.] S. digito postico rotundato mutico. Struthio Camelus americanus. Raj. av. 36. Nhanduguacu brasiliensibus. Marcgr. bras. 190. Pis. bras. 84. Habitat in America australi., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 155, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Struthio [gen. nov.] Rostrum conicum. Alae ad volandum ineptae., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 155, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Struthio cucullatus [spec. nov.] S. pedibus tetradactylis, omnibus unguiculatis. Gallus gallinaceus peregrinus. Clus. exot. 99. t. 100. Olear. mus. 23. t. 13. f. 5. Cygnus cucullatus. Nieremb. nat. 231. Raj. av. 37. Dronte. Bont. jav. 70. Habitat in India., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 155, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206