A M ERIC AN MUSEUM NOVITATES Number 3790, 46 pp. December 5, 2013 The cranial pneumatic sinuses of the tyrannosaurid Alioramus (Dinosauria: Theropoda) and the evolution of cranial pneumaticity in theropod dinosaurs MARIA EUGENIA LEONE GOLD,1,2STEPHEN L. BRUSATTE,3 AND MARK A. NORELL1 ABSTRACT Archosaurs and mammals exhibit skeletal pneumaticity, where bone is infilled by air- filled soft tissues. Some theropod dinosaurs possess extensively pneumatic skulls in which many of the individual bones are hollowed out by diverticula of three main cranial sinus systems: the paranasal, suborbital, and tympanic sinuses. Computed tomography (CT scan- ning) permits detailed study of the internal morphology of cranial sinuses. But only a few theropod specimens have yet been subjected to this type of analysis. We present CT scans of the remarkably preserved and disarticulated skull bones of the long-snouted tyrannosaurid theropod Alioramus. These scans indicate that Alioramus has extensive cranial pneumaticity, with pneumatic sinuses invading the maxilla, lacrimal, jugal, squamosal, quadrate, palatine, ectopterygoid, and surangular. Pneumaticity is not present, however, in the nasal, postorbital, quadratojugal, pterygoid, or angular. Comparisons between Alioramus and other theropods (most importantly the closely related Tyrannosaurus) show that the cranial sinuses of Alio- ramus are modified to fill the long-snouted skull of this taxon, and that Alioramus has an extreme degree of cranial pneumaticity compared to other theropods, which may be the result of the juvenile status of the specimen, a difference in feeding style between Alioramus and other theropods, or passive processes. Based on these comparisons, we provide a revised terminology of cranial pneumatic structures and review the distribution, variation, and evo- 1 Division of Paleontology, American Museum of Natural History. 2 Richard Gilder Graduate School, American Museum of Natural History. 3 School of GeoSciences, University of Edinburgh, Edinburgh, Scotland, UK. Copyright © American Museum of Natural History 2013 ISSN 0003-0082 2 AMERICAN MUSEUM NOvITATES NO. 3790 lution of cranial pneumaticity within theropod dinosaurs. This review illustrates that most theropods possess a common “groundplan” in which the maxilla and lacrimal are pneuma- tized, and that various theropods modify this groundplan by pneumatizing numerous other bones of the skull. Tyrannosaurids are very pneumatic compared to other theropods, particu- larly in the development of extensive ectopterygoid, quadrate, and palatine sinuses, as well as a pneumatic invasion into the surangular. Tyrannosauroids seem to retain many cranial sinuses, such as the jugal and nasal recesses, which are primitive for coelurosaurs but lost or apomorphically modified in taxa more closely related to birds. INTRODUCTION Archosaurs and mammals are unusual among vertebrates in exhibiting skeletal pneumatic- ity: the invasion of bone by air-filled epithelial diverticula (e.g., Müller, 1908; King, 1966; Duncker, 1971; Witmer, 1990, 1997a, 1997b; Britt, 1993, 1997; Koppe et al., 1999; O’Connor, 2004, 2006; Benson et al., 2012; Butler et al., 2012). Among archosaurs, some theropod dino- saurs possess an extreme degree of cranial pneumaticity, where individual bones of the skull are largely hollowed out by air-filled sinuses. The complex array of theropod cranial sinuses can be organized into several major systems: the paranasal sinuses (which are diverticula of the nasal passage), the suborbital sinuses (which are closely related to the paranasal system), the tympanic sinuses (which originate from the inner ear), (Witmer, 1997b; Witmer and Ridgely, 2008; other sinuses relating to the median pharyngeal system and cervical air sacs, which pneumatize the braincase, are not discussed in detail here). The function of these sinuses is not always clear. The paranasal and suborbital sinuses may influence respiration and physiol- ogy, and the tympanic sinuses may be associated with hearing and balance, but it is also thought that the size, shape, and extent of the sinuses may largely result from opportunistic invasion of bone by soft tissue (e.g., Witmer, 1997a). Regardless of their exact functions, cranial pneumatic sinuses often leave bony correlates on theropod skulls. These correlates include the hollowed out internal sinuses, as well as the foramina and fenestrae that lead into the sinuses and, in some cases, smooth fossae on the external bone surfaces where a soft-tissue sinus (airsac) would have lain against the bone. Because pneumatic fossae, foramina, fenestrae, and sinuses are so widespread and variable among theropods, they often form the basis for phylogenetic characters that are instrumental in elucidating the phylogenetic relationships of theropods (e.g., Turner et al., 2012). This is particularly true for the derived theropod subgroup Coelurosauria—birds and their closest relatives—because many pneumatic features originated and changed during the evolution of early birds and their theropod antecedents. Cranial pneumaticity can often be identified on theropod specimens by observation of large foramina or fenestrae leading into internal sinuses. These observations form the basis for most morphological descriptions of theropod pneumaticity, as well as the delimitation of most phylogenetic characters. However, external observations of specimens provide little information on the size, shape, and extent of the internal sinuses. Furthermore, small pneu- matic foramina or other subtle signs of pneumaticity can easily be missed if the external bone 2013 GOLD ET AL.: CRANIAL PNEUMATICITY OF THE TYRANNOSAURID ALIORAMUS 3 surface is poorly preserved. This raises a pressing concern: our understanding of cranial sinus morphology may be extremely limited if we rely on external observation of specimens alone. As a result, phylogenetic character statements and scores based on such observations may be misleading or incorrect. Computed tomography (CT) scanning is a more refined approach, allowing the internal morphology of theropod skulls to be imaged and studied in detail (e.g., Carlson et al., 2003; Witmer et al., 2008). Although CT scanning has become a standard tool in paleontology, most CT-based studies of theropod dinosaurs have focused on the morphology of the brain, inner ear, cranial nerves, and tympanic sinuses of the braincase region (e.g., Rogers, 1998; Larsson et al., 2000; Brochu, 2000, 2003; Franzosa and Rowe, 2005; Sanders and Smith, 2005; Sampson and Witmer, 2007; Balanoff et al., 2009; Norell et al., 2009; Witmer and Ridgely, 2009; Zelenitsky et al., 2009; Bever et al., 2011, 2013; Lautenschlager et al., 2012). Less work has focused on the remainder of skull, including the morphology of the parana- sal, suborbital, and nonbraincase tympanic sinuses. The cranial pneumaticity of a select handful of theropods has been studied in detail (e.g., Brochu, 2003; Witmer and Ridgely, 2008, 2009; Tahara and Larsson, 2011), but additional well-preserved exemplar specimens must be subjected to CT scanning to better understand the range of sinus morphology and variation in theropods. We use CT data to describe cranial pneumaticity in a spectacularly preserved specimen of the long-snouted tyrannosaurid theropod Alioramus altai (IGM 100/1844; Brusatte et al., 2009, 2012; Bever et al., 2011, 2013). The excellent preservation, relative completeness, and complete disarticulation of the cranial bones of IGM 100/1844 allow for a detailed look at the cranial sinuses and thorough comparison to other theropods that have been the subject of CT study. This specimen also offers unique insight into theropod cranial sinus morphol- ogy because it belongs to a subadult individual with a peculiar longirostrine morphology. Therefore, comparisons between the pneumatic patterns in this specimen and other thero- pods may inform how pneumaticity changes through ontogeny and whether taxa that have undergone extreme morphological changes (e.g., evolution of a long snout) also experience changes in internal sinus morphology. We describe the pneumatic features of each bone of the Alioramus altai skull based on a series of CT scans of IGM 100/1844. We make comparisons with other taxa where CT data is available, most notably the closely related Tyrannosaurus rex (Brochu, 2003; Witmer and Ridgely, 2008). We then discuss the distribution and evolution of cranial pneumatic features in theropods within a phylogenetic context. This review illustrates that most theropods share a common “groundplan” in which the maxilla and lacrimal are pneumatized by diverticula from the antorbital sinus of the paranasal sinus system. various theropods exhibit numerous varia- tions on this architecture, including the development of sinuses invading the jugal, nasal, squa- mosal, palatine, ectopterygoid, quadrate, articular, and surangular. Tyrannosaurids such as Alioramus are among the most extensively pneumatic of all theropods, and seem to retain many cranial sinuses that are primitive for coelurosaurian theropods but lost or modified in taxa more closely related to birds. 4 AMERICAN MUSEUM NOvITATES NO. 3790 Institutional Acronyms AMNH American Museum of Natural History, New York, NY BMR Burpee Museum of Natural History, Rockford, IL CM Carnegie Museum of Natural History, Pittsburgh, PA CMNH Cleveland Museum of Natural History, Cleveland, OH IGM Institute of Geology, Ulaanbaatar, Mongolia ZPAL Institute of Paleobiology, Polish Academy of Science, Warsaw, Poland MATERIALS AND METHODS The disarticulation of the skull of IGM 100/1844 allows each bone to be analyzed separately with high resolution X-ray computed tomography (CT scans). The bones were scanned in a GE phoenix v|tome|x CT scanner at the American Museum of Natural History Microscopy and Imaging Facility. In total, 13 elements were scanned with the following general settings: voltage between 140 and 210 kv, amperage between 125 and 175 µA, and a slice thickness ranging from 0.09 to 0.14 mm (for specific element scan settings, see table 1). Each element was analyzed with visual Graphics Studio Max 2.2, and the region growing and pen tools were used to seperate out the interior pneumatic regions. Those regions were then colored, using the same color scheme as Witmer and Ridgely (2008). Each pneumatic region was compared to corresponding regions in tyrannosauroids and other theropods. Sinus volumes were calculated using the volume calculation tool in visual Graphics Studio Max (table 2). Many of these volumes are slightly overestimated due to the inclusion of pneu- matic foramina (the external openings leading into sinuses) in the volume measurements. The antorbital sinus (which is bounded by many bones and not located within a single bone) was filled in entirely by eye, mainly as a visualization tool; however, its volume is included here for general reference. The squamosal sinus, which is located on the ventral surface of the squamo- sal and not entirely within the bone, was also filled in. The shape of this sinus was inferred from the deep pocket formed by the three processes of the squamosal and the location of the foramina on the ventral side of the dorsal ramus. In rare cases, as with the ectopterygoid, the bone is slightly crushed, so the volume of the internal sinus may represent an underestimate. In such cases, however, differences between the measured volumes and genuine volumes would be minimal, and we hold that the volumes presented here are accurate for the specimen as preserved and are valuable for comparison purposes. TERMINOLOGY In this paper, we use a consistent terminology to refer to the pneumatic space within a bone and the openings on the external surfaces of bones that lead into this internal space. The internal space is referred to as a sinus. For example, the space within the jugal is called the jugal sinus. For descriptive purposes these sinuses are sometimes also referred to as recesses, cavities, or antra, and if they are connected to a larger sinus they may be referred to as a diverticulum 2013 GOLD ET AL.: CRANIAL PNEUMATICITY OF THE TYRANNOSAURID ALIORAMUS 5 TABLE 1. Scanning parameters for each bone of Alioramus altai. Bone Side voltage (kv) Amperage (µA) voxel size Number of slices Angular Right 155 175 0.1107 1913 Ectopterygoid Right 170 160 0.1071 1450 Jugal Left 155 175 0.1416 1774 Lacrimal Left 200 170 0.1224 1804 Larcimal Right 155 125 0.1137 1860 Maxilla Left 190 150 0.1418 1872 Nasals Unpaired 215 180 0.1141 3573 Palatine Left 170 160 0.1037 1860 Pterygoid Right 155 175 0.1348 1897 Quadrate Left 210 150 0.1318 1276 Quadrate Right 190 135 0.1267 1251 Squamosal + postorbital Right 190 135 0.0903 1806 Squamosal Left 210 150 0.0991 1381 Surangular Left 170 160 0.1071 2422 TABLE 2. volume of each cranial sinus in Alioramus altai calculated using vG Studio Max. Bone Sinus volume (mm3) Sinus volume (cm3) Sinus volume in T. rex (cm3)* Maxillary1 23527 23.527 7772.5 Antorbital 229533.6 229.5 6766.1 Lacrimal (L) 11843.28 11.843 1177.1 Lacrimal (R) 15053.64 15.054 1177.1 Jugal 9059.70 9.059 1031.3 Palatine 4146.78 4.146 1082.5 Squamosal 1381.08 1.381 1377.4 Quadrate (L) 23158.917 23.159 482.1 Quadrate (R) 17955.61 17.956 – Ectopterygoid 22184.77 22.185 1641.3 1 Combined volume measurement for promaxillary sinus and maxillary antrum. * From Witmer and Ridgely, 2008. (for example, the jugal sinus is a diverticulum of the larger antorbital sinus). Most sinuses fill the interior of bones, but the antorbital sinus itself lies on the lateral surface of the upper jaw and the squamosal sinus is located on the ventral surface of the bone and extends only slightly into the bone interior. The external openings that lead into the sinuses are referred to as pneu- matic fenestrae or pneumatic foramina. The distinction between a fenestra and a foramen is largely subjective, with a fenestra usually referring to a larger, windowlike opening and a fora- men to a smaller hole. Sometimes foramina is specifically used to refer to passages through which structures, like blood vessels and nerves, pass, whereas fenestra indicates a larger open- 6 AMERICAN MUSEUM NOvITATES NO. 3790 ing, but this distinction is often difficult to make without specific knowledge of cranial nerves and vasculature. For the sake of simplicity, the term pneumatic is sometimes dropped when referring to the fenestrae and foramina in this paper (for instance, we use the term maxillary fenestra not maxillary pneumatic fenestra). This terminology is generally consistent to that used by Witmer (1997a, 1997b), Witmer and Ridgely (2008), and previous authors who have studied the cranial sinuses of archosaurs. However, there is some confusion in the literature, especially with the term recess. For example, the Theropod Working Group research team (TWiG) has sometimes written phylogenetic characters in which the external fenestra leading into a sinus has been called a recess (e.g., Brusatte et al., 2010a: char. 67). Usually these characters relate to the position or shape of the fenestra and not to the position or shape of the internal sinus itself, so they can be particularly confusing. We rectify this situation here with consistent terminology. Regions of pneumaticity in the skull take two general forms (Witmer, 1997a). First, some pneumatic air sacs lie against the external surface of the bone, and leave a characteristic fossa or depression on the bone surface. An example of this is the antorbital sinus, which abuts the lateral surface of the snout, within the broad antorbital fossa. Second, and more common, are internal sinuses within bones that are connected to the bone exterior via one or more fenestrae or foram- ina. These sinuses may have one or multiple chambers and are sometimes finely divided by internal laminae. An example of this is the sinus within the quadrate. These regions of pneuma- ticity are identified by the characteristic morphology of large external fenestrae or foramina leading into an interior cavity. This is also the diagnostic combination that allows for the unequiv- ocal identification of postcranial skeletal pneumaticity in the vertebrae, ribs, and other bones of archosaurs (e.g., Britt, 1993; O’Connor, 2006; Wedel, 2007; Benson et al., 2012; Butler et al., 2012). RESULTS General Skull The antorbital sinus—one of the major anatomical features shared by archosaurs—is a large paranasal air sinus positioned between the orbit and the external naris (Witmer, 1997a, 1997b). In Alioramus, as in most other theropods and many other archosaurs, this sinus is an extensive feature that occupies much of the lateral aspect of the snout. The antorbital sinus is not enclosed within bone, but rather is located between the maxilla, lacrimal, jugal, and palatine. It would have been covered laterally by skin (and perhaps other soft tissues) in life, as is normal for most archosaurs (Witmer, 1997a, 1997b). The position of the sinus is marked by the antorbital fossa, as the antorbital air sac (para- nasal air sinus) would have sat against the smooth surface of the fossa in life. The shape of the fossa, and thus the antorbital sinus, mirrors the elongation of the snout in being dorsoventrally low and rostrocaudally elongate. The fossa covers much of the lateral surface of the maxilla, including much of the region rostral to the antorbital fenestra and much of the ascending ramus dorsally and main (alveolar) body ventrally. Therefore, the maxilla would have formed the rostral and much of the dorsal and ventral borders of the antorbital sinus. ventrally, on the main body of the bone, the antorbital fossa is deeply invaginated as a channel that extends 2013 GOLD ET AL.: CRANIAL PNEUMATICITY OF THE TYRANNOSAURID ALIORAMUS 7 rostrocaudally. Thus, the ventral part of the sinus would have sat within this channel in life. The fossa also extensively excavated much of the rostral ramus and main body of the lacrimal and the rostral region of the jugal. Correspondingly, the lacrimal would have comprised its caudoventral border. Finally, much of the lateral surface of the palatine is covered by the ant- orbital fossa, meaning that this bone would have comprised some of the medial and ventral border of the antorbital sinus. The antorbital sinus sends numerous diverticula into the surrounding elements (Witmer, 1997a, 1997b; Witmer and Ridgely, 2008). These accessory sinuses often hollow out large por- tions of these bones. Among these are two discrete sinuses in the maxilla (the promaxillary sinus and the maxillary antrum), a chamber within the lacrimal (which may be confluent with a second, smaller medial lacrimal sinus), the jugal recess, and a sinus within the palatine (which is a caudal extension of the maxillary antrum). Additionally, as discussed by Witmer (1997a, 1997b) and Witmer and Ridgely (2008), there is probably a suborbital sinus that extends cau- dally and ventrally from the antorbital sinus into the orbit. This sinus is difficult to confirm, much less describe, because it likely was bounded almost entirely by soft tissue. However, it is possible that the suborbital sinus may be the source of diverticula leading into the squamosal and/or ectopterygoid (Witmer and Ridgely, 2008). Maxilla Both left and right maxillae are preserved as disarticulated bones in the IGM 100/1844, but only the left maxilla was CT scanned here, as it is more complete, better preserved, and substantially less deformed than the right (fig. 1; Brusatte et al., 2012). The left maxilla is nearly complete, and even retains most of the thin, fragile medial bounding wall of the maxillary antrum and promaxillary sinus (the two main accessory chambers of the antorbital sinus that extend into the maxilla, rostral to the antorbital fenestra: Witmer, 1997a, 1997b). In contrast, this thin, bony flange is eroded or broken in nearly every other known tyrannosaurid maxilla (Witmer and Ridgely, 2008). Because this wall encloses the maxillary antrum and promaxillary sinus medially, the shape and volume of these sinuses can uniquely be studied in complete detail in IGM 100/1844. As in Tyrannosaurus and most other nonavian tetanuran theropods, Alioramus possesses two separate diverticula extending from the antorbital sinus into the interior of the maxilla. These are the promaxillary sinus and maxillary antrum, which together are often referred to as the maxillary sinus (Witmer, 1997a, 1997b). More basal theropods, on the other hand, pos- sess only a very small, single sinus within the maxilla. This condition is exemplified by the large abelisaurid Majungasaurus (Sampson and Witmer, 2007; Witmer and Ridgely, 2008). In Alio- ramus, Tyrannosaurus, and most other nonavian tetanurans, the diverticulum leading into the promaxillary sinus passes through the promaxillary fenestra and the connection with the max- illary antrum passes through the maxillary fenestra. The promaxillary and maxillary fenestrae are features characteristic of the maxilla in most tetanurans and usually are small, subsidiary openings within the antorbital fossa, rostral to the antorbital fenestra (Witmer, 1997a). Their presence indicates that the promaxillary sinus and maxillary antrum, or some homologous feature, is also present even if CT data are not available 8 AMERICAN MUSEUM NOvITATES NO. 3790 e n atm mu xiol oc prht pg n aRi ntorbital sinus intorbital sinus. ws ahe a hout t n sho umwit ft colrent Lepa ws. rans viemit ) e Ds (a ral xill ntma e d vws no ah C), n s (m orsal colu B), dddle mm. medial (rent. Miar = 45 eral (A), mitranspas. Scale b n lata sesinu axilla imaxillxillary eft mwith d ma 1. LURE position, ws isolate Ge o FIlifsh 2013 GOLD ET AL.: CRANIAL PNEUMATICITY OF THE TYRANNOSAURID ALIORAMUS 9 to confirm this. Alioramus has large, distinctive promaxillary and maxillary fenestrae (Brusatte et al., 2012), as do other tyrannosauroids (e.g., Molnar, 1991; Brochu, 2003; Currie, 2003; Hurum and Sabath, 2003). Therefore, all known tyrannosauroids likely possess a promaxillary sinus and maxillary antrum, based on their possession of the corresponding fenestrae. The size, shape, and position of the fenestrae are highly variable among tyrannosauroids. This variability has been differentiated into several characters used in phylogenetic analyses (e.g., Holtz, 2001; Currie et al., 2003; Holtz, 2004; Sereno et al., 2009; Brusatte et al., 2010a; Carr and Williamson, 2010). These characters are not reviewed in detail here, but are discussed more thoroughly in Brusatte et al. (2012). The promaxillary sinus of Alioramus is a large space occupying much of the interior of the maxilla rostral to the antorbital fossa. There is a wall of bone dorsal to the roots of the maxillary teeth that prevents the promaxillary sinus from communicating with the tooth crypts. From the dorsal edge of that bone wall, the promaxillary sinus extends dorsally, following the dorsal cur- vature of the maxilla until it reaches the interfenestral strut. The sinus is rostrocaudally broad, extending caudally from the rostral margin of the maxilla to the rostral edge of the antorbital fossa, where the promaxillary fenestra is positioned. Therefore, the promaxillary sinus is located strictly rostral and medial to the promaxillary fenestra. The medial surface of the rostral margin of the maxilla curves as it projects dorsally and is a very thin bone layer that has many medial openings. These openings may allow the promaxillary sinus to send narrow extensions into the premaxilla or the nasal passage. Neither premaxilla is preserved in IGM 100/1844, so any poten- tial extensions of the promaxillary sinus into the premaxilla can only be inferred. The long and extensive maxillary antrum is located ventral and caudal to the promaxillary sinus. The ventralmost edge of this sinus appears to penetrate the root of each tooth, extending into the alveoli in many cases along the tooth row. This may be an artifact of CT scanning, however, where adjacent structures look continuous. Therefore, the maxillary antrum may terminate prior to each tooth root. This is similar to the condition in Tyrannosaurus, in which “interalveolar recesses” extend ventrally from the maxillary sinus between the tooth crypts (Witmer, 1997a; Witmer and Ridgely, 2008). ventrally, the maxillary antrum continues along the tooth row until the last maxillary tooth, penetrating into each alveolus. The maxillary antrum extends through the medial surface of the maxilla through an opening above the ros- tromedial process (which currently is unnamed), and also through an open channel ventrome- dial to the antorbital fossa. Nasals The nasals (fig. 2) are elongate and fused into a single, vaulted unit, as is normal for tyran- nosaurids (e.g., Holtz, 2001; Snively et al., 2006). The ventral face forms the dorsal surface of the nasal airway. Both the internal and external morphology of the bone, however, demonstrate that the antorbital sinus and any associated diverticula were not associated with the nasal. The internal region of the bone lacks any pneumatic pockets, chambers, or sinuses. The external surface lacks any extension of the smooth antorbital fossa and any foramina or fenestrae (Bru- satte et al., 2012). Both of these conditions are generally shared with other tyrannosaurids. Brochu (2003) and Witmer and Ridgely (2008) used CT data to confirm the lack of any internal 10 AMERICAN MUSEUM NOvITATES NO. 3790 FIGURE 2. Nasals in lateral (A), dorsal (B), and ventral (C) views. Scale bar 45 mm. cavities within the nasal of Tyrannosaurus. Most tyrannosaurid nasals also lack any trace of the antorbital fossa on the external surface, although some juvenile tyrannosaurids do exhibit a very narrow extension of the fossa (Carr, 1999). The condition in more basal tyrannosauroids, however, is drastically different. The basal (non-tyrannosaurid) taxa Dilong (Xu et al., 2004), Eotyrannus (Hutt et al., 2001), Guanlong (Xu et al., 2006), and Yutyrannus (Xu et al., 2012) exhibit pneumatized nasals with internal cavities. These cavities communicate with the antorbital sinus via one or more pneumatic fenestrae, which are housed within an extension of the antorbital fossa that excavates part of the lateral surface of the nasal. The basal tyrannosauroid Proceratosaurus also appears to have internal nasal pneumaticity, based on CT data presented by Rauhut et al. (2010). Nasal pneumaticity is rare in coelurosaurs, but present in many non-coelurosaurian outgroups, such as allosauroids (e.g., Allosaurus: Madsen, 1976; Neovenator: Brusatte et al., 2008; Sinraptor: Currie and Zhao, 1993), ceratosaurs (e.g., Majungasaurus: Sampson and Witmer, 2007; Witmer and Ridgely, 2008), and Monolophosaurus (Brusatte et al., 2010b). The distribution of pneumaticity in the- ropod nasals indicates that it evolved early in theropod evolution and was secondarily lost in tyrannosauroids around the base of Tyrannosauridae (see Discussion). This indicates that the lack of pneumaticity in more-derived coelurosaurs is an independent loss and not primitive for Coeulurosauria. Lacrimal The lacrimal (figs. 3, 4) is extensively pneumatized internally by a single, large chamber, the lacrimal sinus (Witmer, 1997a, 1997b; Witmer and Ridgely, 2008). Numerous foramina on the lateral surface lead into this recess, including several within the antorbital fossa on the rostral ramus (the portion of the lacrimal roofing the antorbital fenestra, and thus the antorbital sinus