Archaehierax sylvestris Mather, Lee, Camens and Worthy, sp. nov.
Creators: name: Mather, Ellen K.
name: Lee, Michael S. Y.
name: Camens, Aaron B.
name: Worthy, Trevor H.
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Aves
Falconiformes
Falconidae
Archaehierax
Archaehierax sylvestris

description: <b> Archaehierax sylvestris <i>Mather, Lee, Camens and Worthy</i> <i>gen. et sp. nov.</i></b> <i>(Figures 3-10)</i> http://zoobank.org/ urn:lsid:zoobank.org:act: 092140CA-E93743B3-B3F1-7127C97094F4 <i>Holotype</i> SAMA P.54998, 63 elements and associated fragments of a single skeleton (see Figure 2) as follows: Fragments of mandible; rostral majority of rostrum; R pterygoid; L quadratojugal; L quadrate; ceratohyal; atlas vertebra; axis vertebra; partial cervical vertebra #3; caudal vertebrae x3 (position in tail indeterminate); cranial part sternum; LR scapulae; cranial and sternal parts LR coracoids; proximal LR humeri; Land distal R ulna; Land distal R radius; L carpometacarpus; LR os carpi ulnare; Los carpi radiale; Rmanual phalanges, proximal fragment MI.1, proximal fragment MII.1 and MII.2; Lmanual phalanges MI.1, MII.2 with distal end eroded, and MIII.1; LR proximal femur fragments; Rtibiotarsus reconstructed in two parts; L tibiotarsus; fragmented LR fibulae; LR tarsometatarsi; LR ossa metatarsalia; pedal phalanges: RI.1, RII.1, RII.2, RIII.1, RIII.2, RIII.3 (partial), RIII.4, LI.1, LI.2 (fragmented), LII.1, LII.2, LIII.1, LIII.2, LIII.3 (partial), LIII.4, LIV.1, LIV.2, LIV.3, LIV.4, LIV.5. The skeleton was found eroding out on the surface with surviving elements recovered in a semi-articulated state from within dolomitic clays (equivalent to layer 5 of Thorn et al. 2021), with most large elements fractured into many roughly articulated pieces, presumably by expansion and contraction associated with the wetting and drying of the clays. The fragments for each element were, where possible, separated, cleaned and reassembled by THW. <i>Measurements (mm)</i> See Appendix 1 Table S1. <i>Etymology</i> The species name ‘sylvestris’ is derived fromthe Greek world ‘sylvas’, meaning forest, and the Latin suffix ‘-estris’, meaning ‘belonging to’. <i>Type locality/Stratigraphy and age</i> As per genus. <i>Diagnosis</i> As for genus. <i>Descriptions</i> <i>Rostrum maxillare</i> (Figure 3 (A, B)). The rostral section of the rostrum maxillare is preserved in reasonably good condition. Morphology of its rostral tip, tomial margin, rostral marginof the nares, and the palatines is visible. The rostrum has a preserved length of 32.6 mm from the rostral tip to the posterior base of the nares, and a preserved depth of 17.3 mm from the tomial margin to the dorsal side of the rostrum taken at the rostral end of the nares. <b>(Trait 1)</b> The rostral tip of the rostrum is hooked, descending below the tomial margin at a 30–40° angle. <b>(2)</b> The lateral tomial margin (Figure 3A: CT), positioned distal to the nares, is ventrally convex. <b>(3)</b> The nasal aperture (Figure 3A: N) is large (height 8.7 mm) and fully open as in most accipitrids, spanning just over half the rostrum depth. <b>(4)</b> The ossified section rostral to the incisura ventromedialis is of small to moderate size (10.1 mm preserved length) relative to the total length of the rostrum. <b>(5)</b> In ventral aspect, an incisura ventromedialis (Figure 3B: IV) (sensu Livezey and Zusi 2007) is present (sediment filled) extending from where damage destroys it rostrally to the preserved caudal end of the palate; it is narrow and widens caudally, rather than being closed forming a fenestra. The pars maxillaris palatini (Figure 3B: PM) are unfused and diverge slightly caudally, the left being least fragmented although it has broken fromthe adjacent lateral margin creating a false incision. Asmall fragment of bone preserved between the pars maxillaris palatini is interpreted as a displaced fragment of the processus maxillopalatinus. <b>(6)</b> Damage precludes ascertaining the presence/form of the fenestra ventrolaterale. Other accipitrid subfamilies differ as follows: <b>(Trait 1)</b> Compared to the fossil, the tip of the rostrum is much more sharply hooked ventrally in most subfamilies. Only members of Elaninae, Perninae (except <i>Chondrohierax uncinatus</i>, which is sharper), Buteoninae, and Gypaetinae have similar or shallower angled tips. The rostrum maxillare is overall most similarto that of species of Buteoninae (see SI.2 for more detailed differential comparisons). The left quadrate has considerable breakage affecting the lateral side ventrally, the medial side of the processus oticus, and loss of the processus orbitalis. Preservation of morphological detail isbest medially. On the processus oticus, only about half of the capitulum squamosum is preserved and the dorsal half of the crista medialis is lost. Of the processus orbitalis, only the well-preserved base remains. Both thecondylus pterygoideus and medialisare intact, butthe entire condylus lateralis and caudal half of the condylus caudalis are lost. <b>(Trait 1)</b> The processus oticus (Figure 3D: POt) is short and broad leading up to the capitulum squamosum. <b>(2)</b> The capitulum squamosum (Figure 3D: CS), as preserved is relatively small, and has a tuberculum subcapitulare forming a distinct hook projecting ventrally on its cranial margin. <b>(3)</b> The processus orbitalis (Figure 3C: POr) is dorsomedially oriented. <b>(4)</b> The processus is set entirely in the ventral half of the quadrate, creating a gentle, shallow sloping arc between the base of the processus and the capitulum. <b>(5)</b> Alarge and deep foramen pneumaticum basiorbitale (Figure 3C: FPB) is present between the processus orbitalis and the condylus pterygoideus. <b>(6)</b> Breakage prohibits assessing the status of the foramen rostromediale. <b>(7)</b> The ventral section of the crista medialis is preserved and is quite broad and flat with no projecting ridge. <b>(8)</b> A thin, distinct sulcusruns along the ventral margin of the crista to connect to the foramen pneumaticum caudomediale. <b>(9)</b> A distinct, deep foramen pneumaticum caudomediale (Figure 3C: FPC) is present just medial of the ventral-most point of the crista medialis. <b>(10)</b> The condylus pterygoideus (Figure 3D: CP) is distinct, high-set and well separated from the condylus medialis. <b>(11)</b> The condylus medialis (Figure 3C: CM) is well preserved, showing a large (4.0 mm wide by 2.8 mm deep), semi-ovular facet, with a pointed medial margin that extends further medially than the condylus pterygoideus. <b>(12)</b> The portion of the condylus caudalis (Figure 3C: CC) preserved indicates a facet of a similar size to the condylus medialis, with a semicircular shape. Other accipitrid subfamilies differ as follows (variable characters excluded): <b>(Trait 8)</b> The sulcus running along the ventral crista is broad and indistinct in all other taxa except in species of <i>Milvus</i> and <i>Haliaeetus</i> (Haliaeetinae) where it is narrow and indistinct. <b>(9)</b> The fossa caudomediale is practically absent in the subfamilies Circaetinae and Aegypiinae, as well as the species in <i>Hamirostra, Lophoictinia</i> (Perninae), <i>Neophron</i> (Gypaetinae), and <i>Haliaeetus</i> (Haliaeetinae); shallow in Aquilinae and species of <i>Elanoides, Chondrohierax</i> (Perninae), <i>Polyboroides</i> and <i>Gypohierax</i> (Gypaetinae); and deep in Elaninae, Accipitrinae and Buteoninae, as well as species in <i>Pernis</i> (Perninae). The depressio was indistinctly shaped (i.e. a gradually deepened area rather than a defined pit) in all species, and apneumatic in all species except in <i>Pernis</i> and <i>Haliaeetus</i>. The quadrate is overall most similar to that of species of Aegypiinae (see SI.2 for more detailed differential comparisons). The atlas vertebra of SAMA P.54998 is 10.7 mm wide by 10.7 mm high (from the proximal margin of arcus atlantis to the distal margin of fossa condyloidea). <b>(Trait 1)</b> The arcus atlantis (Figure 3G: Arc. At.) forms a low, flat arch, which dorsally has a maximum proximodistal width of 3.2 mm at the centre, overhanging the fossa condyloidea cranially. <b>(2)</b> The incisura fossae (Figure 3G: IF) is shallow and broad, forming a semicircular shape in cranial aspect. <b>(3)</b> The fossa condyloidea (Figure 3G: FC) is 4.3 mm wide by 4.2 mm long. <b>(4)</b> The dorsolateral eminences of the fossa condyloidea are small. <b>(5)</b> The ventral margin of the fossa forms a rounded point in cranial aspect and is prominent cranially in lateral aspect. <b>(6)</b> The zygopophyses caudales are badly worn and difficult to assess, but what is preserved indicates they were distinct and caudally projected from the rest of the arcus atlantis. <b>(7)</b> The incisurae caudales arcus (Figure 3E: IAC) are very shallow. <b>(8)</b> The facies articularis axialis (Figure 3E: FAA) is 5.7 mm wide by approximately 3.8 mm long. <b>(9)</b> The rest of the distal corpus atlantis is worn away and cannot be assessed, but what is present suggests that few additional structures were present, and that there was some asymmetry in the shape of the lateral fossa condyloidea. The axis vertebra is quite fragmented, with most of the neural spine, facies articularis atlantica, dens, facies articularis caudalis, processus ventralis and the area of incisurae caudales arcus broken away. Its width is 12.7 mm across the processus articulares caudales and mid-line length of the corpus vertebra is 8 mm. The facies articularis on zygopophyses caudales are 3.1 mm wide by 3.8–4.1 mm long. <b>(10)</b> Ashort but caudally prominent projection is present dorsal to each facet (Figure 3H: FA), measuring 2–2.5 mm in width. <b>(11)</b> There is no evidence of a bridge enclosing the incisurae caudales arcus (Figure 3F, H: PAC). The three known caudal vertebrae are too broken to identify their position in the caudal series. The vertebrae are overall most similar to that of species of Elaninae (see SI.2 for more detailed differential comparisons). The cranial section of the sternum of SAMA P.54998 is preserved, retaining the structure of the spina externa, pila carinae, crista medialis carinae, apex carinae, the left sulcus articularis coracoideus, and the left labrum internum. It is characterised by: <b>(Trait 1)</b> Aspina interna is absent; a small notch exists in its place. <b>(2)</b> The spina externa (Figure 4B: SE) is 6.4 mm wide at its base, 4.7 mm wide at its blunt tip and 4.2 mm long. In cranial view, the spina externa is triangular as a medial crista forming a lobe projecting 3.6 mm ventrally. The base of the spina externa is broader than the apex carinae (4.0 mm). <b>(3)</b> The crista medialis carinae (Figure 4A: CMC) is short, extending dorsally from the apex carinae to mid-height of the pila carinae with a low profile in lateral view. More dorsad, the pila carinae is smooth to the base of the spina externa. <b>(4)</b> The pila carinae (Figure 4A: PC) is robust for its size, measuring 5.2 mm at mid-depth. It is 23.3 mm long from the ventral margin of the carina sterni to where the pila carinae meets the spina externa. <b>(5)</b> The apex carinae (Figure 4A: AC) at 4.0 mm wide, is noticeably expanded from the width of the pila carinae immediately dorsal to it (3.0). <b>(6)</b> The apex carinae is rounded, with no hooked projection extending cranially. <b>(7)</b> The maximum carina depth below the sternal basin is roughly equivalent to basin depth below the costal margin, typical of most accipitrids except for certain vulture species (Aegypiinae). <b>(8)</b> The apex carinae is set well caudally [assuming the junction of the carina with the pars cardiaca is aligned horizontally] from the base of the spina externa. <b>(9)</b> There are no small, dispersed pneumatic foramina present dorsally in the body of the sternum. <b>(10)</b> The small part of the carina sterni preserved narrows caudally from the pila carinae. <b>(11)</b> The left sulcus articularis coracoideus (Figure 4B: SAC) is 4.3 mm dorsoventrally deep, and extends to the midline of the sternum, directly dorsal to the spina externa, where it does not overlap the right sulcus. <b>(12)</b> The left labrum internum (Figure 4A: LI) is maximally 4.2 mm deep and 2.8 mm wide in dorsal view as preserved. <b>(13)</b> Apila medialis (Figure 4C: PM), 2.1 mm wide, on the dorsal face of the sternum separates two deep, pneumatic fossae in the pars cardiaca. Extant accipitrid subfamilies differ as follows: <b>(3)</b> The crista medialis extends to the base of the spina externa in membersof allsubfamilies except Aegypiinae. <b>(8)</b> Theapex carinaelies directly ventral to the base of the spina externa, or projects more craniad, in species in all subfamilies except for Gypaetinae and Aegypiinae. <b>(11)</b> The sulci articularis coracoidei overlap in all species except for <i>Gampsonyx swainsonii</i> (Elaninae) and <i>Sarcogyps calvus</i> (Aegypiinae). <b>(13)</b> No species in any subfamily has a distinct pila medialis separating pneumatic fossae in the pars cardiaca, which is thus an autapomorphy suggesting subfamilial distinction for the new species. The sternum is overall most similar to that of species of Aegypiinae (see SI.2 for more detailed differential comparisons). The well-preserved left and right omal ends and fragments of both sternal ends of the coracoids of SAMA P.54998 were recovered. They reveal the following: <b>(Trait 1)</b> Aforamen nervi supracoracoidei (Figure 4G: FoNS) is present and located adjacent to the shaft rather than near the medial margin of the processus procoracoideus; <b>(2)</b> The foramen lacks an opening into the corpus; and <b>(3)</b> it is small, about 1 mm in width, and positioned just sternal of the cotyla scapularis. <b>(4)</b> Alarge (6 mm wide) pneumatic foramen is present in the sulcus m. supracoracoidei (Figure 4F: SMS). The width of the sulcus is approximately 14.5 mm fromthe ventrosternal corner of the facies articularis clavicularis to the facies articularis humeralis, and 12.8 mm from the medial margin to the laterodorsal margin immediately cranial to the cotyla scapularis. <b>(5)</b> The facies articularis clavicularis (Figure 4F: FAC) is large, broad, and clearly delineated sternally by a crista that dorsally overhangs the aforementioned foramen, and ventrally is a low non-overhanging crista. The sternal margin of this facet is straight with no notch nor dorsal or ventral projections directed sternally. <b>(6)</b> The cotyla scapularis (Figure 4G: CtS), preserved on the right omal fragment, is deep and large (6.7 mm wide by 5.7 mm long) in relation to the processus procoracoideus and triangular shaped. <b>(7)</b> The facies articularis humeralis (Figure 4G: FAH) is 7.7 mm wide and 12.8 mm long. <b>(8)</b> The impressio lig. acrocoracohumeralis (Figure 4H: ILA), best seen on the left specimen, forms a distinct sulcus ~ 7 mm wide by 18.1 mm long on the processus acrocoracoideus, although this may be exaggerated by damage to the fossil. <b>(9)</b> The processus procoracoideus (Figure 4I: P. Procor.) forms a short projection medially, barely as long again as the cotyla scapularis width, with its tip sharply angling ventrally towards the medial face to partly enclose the triosseal canal. <b>(10)</b> The best preserved sternal-end fragment shows that the angulus medialis is acute, forming a 30–45° angle. <b>(11)</b> The medial side of the facies articularis sternalis is 6.5 mm wide at its broadest point, and shallow, with little deepening towards the dorsal margin. Extant accipitrid subfamilies differ as follows: <b>(Trait 5)</b> The sternal margin of the facies articularis clavicularis does not form a crest overhanging the sulcus supracoracoideus in Elaninae, Perninae (except <i>Chondrohierax uncinatus</i>), Gypaetinae (except <i>Polyboroides typus</i>), Aegypiinae, Haliaeetinae, Accipitrinae, and Buteoninae. <b>(8)</b> The impressio lig. acrocoracohumeralis is shallow in Elaninae, Perninae, Gypaetinae, Accipitrinae, and Buteoninae. <b>(9)</b> In all subfamilies, the processus procoracoideus does not, or barely, angles ventrally towards the medial face. The coracoid is overall most similar to that of species of Aegypiinae, Accipitrinae and Buteoninae (see SI.2 for more detailed differential comparisons). Both the left and right scapulae of <i>Archaehierax sylvestris</i> gen. et. sp. nov. are almost complete, lacking only the distal third or less of the corpus scapulae. In total, the preserved craniocaudal length of the scapulae is 56.8 mm (left) and 53.3 mm (right). The proximal dorsoventral width of the scapula is 14.4 mm from the acromion to the ventral side of the facies articularis humeralis. <b>(1)</b> The tuberculum coracoideum (Figure 4E: TC) is low and barely cranially prominent dorsal of the facies articularis humeralis. <b>(2)</b> The acromion (Figure 4D: Ac) has a distinct cranio-laterally oriented crista lig. acrocoracoacromiali dorsally, and a robust rounded medial prominence. <b>(3)</b> There are no pneumatic foramina or fossae present in the acromion cranially, <b>(4)</b> nor on the lateral or medial facies between the acromion and the facies articularis humeralis. <b>(5)</b> The facies articularis humeralis (Figure 4D: FAH) is quite large and broad, measuring 6.3 mm dorsoventrally by 10.8 mm craniocaudally on the left specimen; 6.3 mm by 10.4 mm on the right. <b>(6)</b> The acromion barely projects proximally/cranially of the tuberculum coracoideum. <b>(7)</b> Minimum dorsoventral depth of the collum scapulae (Figure 4E: ColS) is 6.3 mm. <b>(8)</b> While the extremitas caudalis is broken off, the corpus scapulae is elongate and moderately narrow, <b>(9)</b> but it greatly increases in depth caudal to the margo dorsalis ridge (Figure 4D: MD), attaining a maximum dorsoventral depth of 8.1 mm. <b>(10)</b> The lateral face dorsal to and immediately posterior to the facies articularis humeralis is flat. <b>(11)</b> The ligamental attachment on the margo dorsalis has a very small prominence and is not elevated dorsally above the rest of the margo dorsalis. Extant accipitrid subfamilies differ as follows: <b>(Trait 6)</b> The acromion strongly projects cranially in all species except in pernines (<i>Chondrohierax uncinatus, Pernis apivorus, Aviceda subcristata</i>), and <i>Ictinia mississippiensis</i> (Buteoninae). The scapula is overall most similar to that of species of Elaninae (see SI.2 for more detailed differential comparisons). The humeri are poorly preserved in SAMA P.54998. Only the caput humeri, crus dorsale fossae, fossa pneumotricipitalis, and the incisura capitis of the proximal end of the right humerus is preserved. The left humerus is more complete, preserving about 60mm of proximodistal length of the proximal end including the caput humeri, crus dorsale fossae, fossa pneumotricipitalis, incisura capitis, sulcus lig. transversus, facies bicipitalis, crista deltopectoralis, and some of the proximal shaft. However, there is also significant breakage and fracturing of the bone surface in this specimen, which has resulted in the loss of the tuberculum dorsale, the ventral marginof the crista bicipitalis, tuberculum ventrale, and sulcus n. coracobrachialis. These specimens reveal the following: <b>(1)</b> The incisura capitis (Figure 5A: IC), as best observed in the left humerus, is shallow and lacks secondary deepening. <b>(2)</b> There is no visible ligamental scar in the distal incisura capitis. <b>(3)</b> The caput humeri (Figure 5A: CH) is quite flattened, projecting proximal to the incisura capitis only a few millimetres. <b>(4)</b> The fossa pneumotricipitalis (Figure 5A: FP) was large and deep, although breakage precludes assessing its former width. The betterpreserved left specimen shows it was minimally 7.7 mm w
description: Published as part of <i>Mather, Ellen K., Lee, Michael S. Y., Camens, Aaron B. &amp; Worthy, Trevor H., 2021, An exceptional partial skeleton of a new basal raptor (Aves: Accipitridae) from the late Oligocene Namba formation, South Australia, pp. 1175-1207 in Historical Biology 34 (7)</i> on pages 7-22, DOI: 10.1080/08912963.2021.1966777, http://zenodo.org/record/5534487
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