A Novel Antimicrobial Peptide, Dermaseptin-SS1, with Anti-Proliferative Activity, Isolated from the Skin Secretion of Phyllomedusa tarsius.

The emergence of multidrug-resistant bacteria has severely increased the burden on the global health system, and such pathogenic infections are considered a great threat to human well-being. Antimicrobial peptides, due to their potent antimicrobial activity and low possibility of inducing resistance, are increasingly attracting great interest. Herein, a novel dermaseptin peptide, named Dermaseptin-SS1 (SS1), was identified from a skin-secretion-derived cDNA library of the South/Central American tarsier leaf frog, Phyllomedusa tarsius, using a 'shotgun' cloning strategy. The chemically synthesized peptide SS1 was found to be broadly effective against Gram-negative bacteria with low haemolytic activity in vitro. A designed synthetic analogue of SS1, named peptide 14V5K, showed lower salt sensitivity and more rapid bacteria killing compared to SS1. Both peptides employed a membrane-targeting mechanism to kill Escherichia coli. The antiproliferative activity of SS1 and its analogues against lung cancer cell lines was found to be significant.

Distribution and abundance of Peleng Tarsier (Tarsius pelengensis) in Banggai Island group, Indonesia.

The Peleng tarsier (Tarsius pelengensis) is poorly known primate, with a range limited to Banggai island-group, Central Sulawesi, Indonesia. It was classified as "Endangered" by IUCN in 2017 based on extremely limited demographic and distributional data. The aim of this study was to collect and analyze data on the population and distribution of Peleng tarsiers. Surveys were conducted over approximately 5 months in 2017 and 2018 across Peleng and the neighboring islands of Banggai, Labobo, and Bangkurung. We determined that tarsiers only occur on Peleng and Banggai Island. The average population density in Peleng and Banggai was estimated to be 234 individuals/km2. This is comparable to the broad ranges of tarsier densities throughout Sulawesi and offshore islands. Peleng tarsiers were found in all elevations (0-937 m above sea level) and nearly all vegetated habitats in Peleng island. Using the IUCN criteria for determining conservation status, in conjunction with our new data, we believe that the Peleng tarsier population should be classified as "Vulnerable".

NSD1 gene evolves under episodic selection within primates and mutations of specific exons in humans cause Sotos syndrome.

BACKGROUND: Modern human brains and skull shapes differ from other hominids. Brain growth disorders as micro- (ASPM, MCPH1) and macrocephaly (NFIX, GLI3) have been highlighted as relevant for the evolution in humans due to the impact in early brain development. Genes associated with macrocephaly have been reported to cause this change, for example NSD1 which causes Sotos syndrome. RESULTS: In this study we performed a systematic literature review, located the reported variants associated to Sotos syndrome along the gene domains, compared the sequences with close primates, calculated their similarity, Ka/Ks ratios, nucleotide diversity and selection, and analyzed the sequence and structural conservation with distant primates. We aimed to understand if NSD1 in humans differs from other primates since the evolution of NSD1 has not been analyzed in primates, nor if the localization of the mutations is limited to humans. Our study found that most variations causing Sotos syndrome are in exon 19, 22 and 10. In the primate comparison we did not detect Ka/Ks ratios > 1, but a high nucleotide diversity with non-synonymous variations in exons 10, 5, 9, 11 and 23, and sites under episodic selection in exon 5 and 23, and human, macaque/colobus/tarsier/galago and tarsier/lemur/colobus. Most of the domains are conserved in distant primates with a particular progressive development from a simple PWWP1 in O. garnetti to a complex structure in Human. CONCLUSION: NSD1 is a chromatin modifier that suggests that the selection could influence brain development during modern human evolution and is not present in other primates; however, nowadays the nucleotide diversity is associated with Sotos syndrome.

Small, odd and old: The mysterious Tarsius pumilus is the most basal Sulawesi tarsier.

In this study, we present the first genetic evidence of the phylogenetic position of Tarsius pumilus, the mountain tarsier of Sulawesi, Indonesia. This mysterious primate is the only Eastern tarsier species that occurs exclusively in cloud forests above 1800 m.a.s.l. It exhibits striking morphological peculiarities-most prominently its extremely reduced body size, which led to the common name of 'pygmy tarsier'. However, our results indicate that T. pumilus is not an aberrant form of a lowland tarsier, but in fact, the most basal of all Sulawesi tarsiers. Applying a Bayesian multi-locus coalescent approach, we dated the divergence between the T. pumilus lineage and the ancestor of all other extant Sulawesi tarsiers to 9.88 Mya. This is as deep as the split between the two other tarsier genera Carlito (Philippine tarsiers) and Cephalopachus (Western tarsiers), and predates further tarsier diversification on Sulawesi by around 7 Myr. The date coincides with the deepening of the marine environment between eastern and western Sulawesi, which likely led to allopatric speciation between T. pumilus or its predecessor in the west and the ancestor of all other Sulawesi tarsiers in the east. As the split preceded the emergence of permanent mountains in western Sulawesi, it is unlikely that the shift to montane habitat has driven the formation of the T. pumilus lineage.

Identification of multiple TAR DNA binding protein retropseudogene lineages during the evolution of primates.

The TAR DNA Binding Protein (TARDBP) gene has become relevant after the discovery of its several pathogenic mutations. The lack of evolutionary history is in contrast to the amount of studies found in the literature. This study investigated the evolutionary dynamics associated with the retrotransposition of the TARDBP gene in primates. We identified novel retropseudogenes that likely originated in the ancestors of anthropoids, catarrhines, and lemuriformes, i.e. the strepsirrhine clade that inhabit Madagascar. We also found species-specific retropseudogenes in the Philippine tarsier, Bolivian squirrel monkey, capuchin monkey and vervet. The identification of a retropseudocopy of the TARDBP gene overlapping a lncRNA that is potentially expressed opens a new avenue to investigate TARDBP gene regulation, especially in the context of TARDBP associated pathologies.

Identifying genetic relationships among tarsier populations in the islands of Bunaken National Park and mainland Sulawesi.

Eastern tarsiers (Tarsius tarsier complex) are small nocturnal primates endemic to Sulawesi Island and small adjacent islands of Indonesia. In 2004, the hybrid biogeography hypothesis predicted this species complex might contain 16 or more taxa, each corresponding to a region of endemism, based on: 1) geological evidence of the development of the archipelago, 2) biological evidence in the form of concordant distributions of monkeys and toads, and 3) the distribution of tarsier acoustic groups. Since then, 11 tarsier species have been recognized, potentially leaving more to be described. Efforts to identify these cryptic species are urgently needed so that habitat conversion, pet trade, and cultural activities will not render some species extinct before they are recognized. We gathered data to test the hypothesis of cryptic tarsier species on three volcanic islands in Bunaken National Park, North Sulawesi, namely Bunaken, Manadotua, and Mantehage, during May-August 2018. We sequenced individuals at 5 nuclear genes (ABCA1, ADORA3, AXIN1, RAG, and TTR) and made comparisons to existing genotypes at 14 mainland sites. Bayesian phylogenetic analyses revealed that island populations are genetically identical in all 5 genes, and formed a clade separated from the mainland ones. The eastern tarsiers first diverged from the western tarsiers approximately 2.5 MYA. The three island populations diverged from mainland tarsiers approximately 2,000-150,000 YA, due to either human activities or natural rafting. This study provides information for tarsier conservation, advances the understanding of biogeography of Sulawesi, and contributes to Indonesian awareness of biodiversity. Further quantitative genetics research on tarsiers, especially the island populations, will offer significant insights to establish more efficient and strategic tarsier conservation actions.

Living in small spaces: Forest fragment characterization and its use by Philippine tarsiers (Tarsius syrichta Linnaeus, 1758) in Mindanao Island, Philippines.

The Philippine tarsier (Tarsius syrichta) is a charismatic species that is threatened by illegal hunting and deforestation. Although they occur in forest and disturbed habitats, ecological information about them is still considerably lacking, which consequently hampers our ability to effectively protect tarsiers from further endangerment. Here, we characterized a 36-ha forest fragment in Mindanao Island where a population of tarsiers persist, and assessed the factors that could have influenced their distribution within the area. We sampled trees (> 1 cm DBH) within 10 × 10-m sampling plots (N = 54), which were established within 1-ha grids (N = 32) and locations where tarsiers were captured (N = 22). The habitat was characterized as a regenerating forest over limestone, with a generally homogeneous structure in terms of tree species richness, abundance, mean DBH, and height. In both sampling plots, we found an abundance of trees below 5 cm in DBH (> 50%) and between 2.6 and 5 m in height (> 40%), which, accordingly, the tarsiers appeared to prefer to use when foraging or sleeping. Lianas were among the most important features of the forest, possibly being a keystone structure in such habitats. Community assemblage, species richness, and mean height of trees, as well as distance to the forest edge, were found to be significant factors that influenced tarsier distribution in the fragment. Our study provides basic yet critical information on the habitat and ecology of Philippine tarsiers in Mindanao, and highlights the importance of forest fragments with rich flora diversity to the survival of the species.

Social Behavior of a Reproducing Pair of the Philippine Tarsier (Tarsius syrichta) in Captivity.

Social interactions of the nocturnal primates are not well studied. One of the species for which social behavior is scarcely known is the Philippine tarsier (Tarsius [= Carlito] syrichta). We observed a reproducing pair of captive individuals over two mating seasons for two consecutive years. The tarsiers spent approximately 4% of their activity budget on social interactions; ca. 20% of time in 0-1 m proximity to each other; and shared sleeping sites for half of the study time. The majority of the animals' social interactions were peaceful: affiliative and sexual (83%), and the smallest component of the behavior was agonistic (17%). We witnessed two copulation events (one per estrus day), each lasting ca. 5 min, and both occurring just after waking. We revealed temporal - nightly and hourly - fluctuations in the frequency of social interactions, in the distances the individuals spent from each other and in the number of vocalizations. The results present the first assessment of the social behavior of the Philippine tarsier, much needed to improve the captive breeding management for this highly sensitive species threatened with extinction.

Successful breeding attempt of a pair of Philippine tarsier (Tarsius syrichta) in a conservation center in Bilar, Bohol, Philippines and recommendations for tarsier husbandry.

The Philippine tarsier (Tarsius syrichta) belongs to the least known nocturnal primates. Tarsiers and remaining biodiversity of the Philippines are under tremendous threats from increasing human expansion, with habitat loss and illegal pet trade being the main reasons for tarsier population decline. In addition, even though the attempts were made by western and local facilities, tarsiers have not survived well in captivity. In this paper, I present an example of successful breeding of the Philippine tarsier in captive conditions but in natural climate. As the most important elements of success, I see a large amount of space provided to tarsiers, the climate similar to their natural habitat and the food resembling their natural diet. Our pair of tarsiers were joined during the mating period and held separately outside the mating period, which corresponds with their behavior in the wild and may have played a crucial role in breeding success. Eliminating stress to animals is also important. The study can provide valuable guidelines for other facilities keeping tarsiers in the Philippines and help to improve tarsiers' welfare and in the future help to establish a viable captive population of the Philippine tarsier that will serve as a backup population and also will decrease demand on tarsiers captured from the wild.

Cryptic Communication in a Montane Nocturnal Haplorhine, Tarsius pumilus.

Crypsis, including visual and auditory concealment, usually manifests in primates as an antipredator strategy. Other factors may also influence cryptic communication style, including habitat structure and phylogenetic history. Compared to less cryptic lowland Sulawesian tarsiers, montane pygmy tarsiers (Tarsius pumilus) exhibit a communication style that lacks scent marks and lower-frequency vocalisations. This study examines why auditory crypsis occurs in montane tarsiers more so than in larger tarsier species and presents the only known spectrograms of T. pumilus in the field. T. pumilus regularly exhibited calls with a dominant frequency of 60-80 kHz (n = 4) in both social situations (duet calls) and stressed contexts. These results indicate that highland, smaller-bodied tarsiers habitually communicate at high frequencies in contexts where Sulawesian and Philippine tarsiers use lower frequencies. While predation threats and habitat acoustics may influence the use of high-frequency vocalisations, this study found that T. pumilus shows an expected relationship between vocal frequency and body mass. These traits may represent a retention of primitive haplorhine traits rather than derived adaptations to a montane environment.

Echolocation in a Nocturnal Primate?

Recently it has been noted that Gursky's spectral tarsier emits ultrasonic calls. Because high-frequency sounds are capable of travelling only short distances, their function is limited by the distance the sound can be discerned. One possible function for these short-distance, high-frequency sounds is that they might be used by the tarsiers to navigate throughout their environment. I conducted this study at Tangkoko Nature Reserve in Sulawesi, Indonesia, from May to August 2017. Using the focal follows, I recorded the start of each locomotor activity to the nearest second. I also simultaneously recorded all ultrasonic vocalisations that the focal individual emitted using a Wildlife Acoustics Ultrasonic Song Meter BAT3. Approximately 42% of the ultrasonic calls that I recorded during this study were given within 1 s prior to engaging in locomotor activity. The tarsiers gave statistically more ultrasonic calls during locomotion than during other behaviours (rest, forage and social). Four types of ultrasonic calls were given while locomoting: whistles, doubles, trills and frequency-modulated calls. Of these, the trill and the frequency-modulated call were only given during locomotion, and trills were only given prior to leaping. These results preliminarily suggest that the tarsiers do use ultrasonic calls for navigation. Studying echolocation in organisms closely related to bats is critical for understanding the evolution of echolocation.

Activity Patterns of Captive Philippine Tarsiers (Tarsius syrichta): Differences Related to Sex and Social Context.

Among tarsiers, nocturnal, obligatory faunivorous primates inhabiting islands of South-East Asia, the Philippine tarsier (Tarsius [= Carlito] syrichta) is one of the least studied. To date, activity patterns of this threatened species have not been the subject of any investigation. In the present study, we provide the first quantitative data on how captive male and female T. syrichta apportion their time for various activities in two social contexts: solitary and paired. We found that the sexes do not differ in activity budgets during the non-mating season, both spending most of their time scanning, resting, foraging and travelling. Comparison of activity budgets of the sexes between the mating and non-mating seasons revealed that although both tarsiers noticeably increased travelling time at the expense of time spent resting, the male changed his behaviour to a much greater extent than the female. We also report on fluctuations in the tarsiers' activities throughout a night and compare time budgets of T. syrichta with available data on the western and eastern species of tarsiers. The results extend the current knowledge of tarsier behaviour and may also assist in practical considerations for keeping this highly sensitive, difficult-to-breed species in captivity.

Spatial and Spectral Mapping and Decomposition of Neural Dynamics and Organization of the Mouse Brain with Multispectral Optoacoustic Tomography.

In traditional optical imaging, limited light penetration constrains high-resolution interrogation to tissue surfaces. Optoacoustic imaging combines the superb contrast of optical imaging with deep penetration of ultrasound, enabling a range of new applications. We used multispectral optoacoustic tomography (MSOT) for functional and structural neuroimaging in mice at resolution, depth, and specificity unattainable by other neuroimaging modalities. Based on multispectral readouts, we computed hemoglobin gradient and oxygen saturation changes related to processing of somatosensory signals in different structures along the entire subcortical-cortical axis. Using temporal correlation analysis and seed-based maps, we reveal the connectivity between cortical, thalamic, and sub-thalamic formations. With the same modality, high-resolution structural tomography of intact mouse brain was achieved based on endogenous contrasts, demonstrating near-perfect matches with anatomical features revealed by histology. These results extend the limits of noninvasive observations beyond the reach of standard high-resolution neuroimaging, verifying the suitability of MSOT for small-animal studies.

Two new species of Eimeria (Apicomplexa: Eimeriidae) in Philippine tarsier (Tarsius syrichta).

Philippine tarsier (Tarsius syrichta) is a small nocturnal primate from the Philippines. Little is known about tarsier parasites, including coccidia (Apicomplexa: Eimeriidae), a highly prevalent parasitic protist group in all vertebrate classes. Only 7 valid species of the genus Eimeria, seven species of Isospora and 5 species of Cyclospora have been described in Primates. This study extends the number of coccidia known in primates by two new species obtained from faeces of Philippine tarsiers from Bohol Island. The newly described Eimeria syrichta n. sp. and Eimeria boholensis n. sp. differ morphologically from each other as well as from other coccidia reported from primates. Partial DNA sequences of three genes were obtained from oocysts of E. syrichta n. sp. and E. boholensis n. sp., and formed clusters according to their host specificity; however, there are no other sequentional data of coccidia from primates, except for the genus Cyclospora, which clusters inside the chicken eimerians, and Cystoisospora belli, which is phylogenetically related to Sarcocystidae. More molecular data on coccidia infecting primates are needed for further discussion.

Assessing female reproductive status of spectral tarsier (Tarsius tarsier) using fecal steroid hormone metabolite analysis.

The wild population of spectral tarsier is declining and attempts to breed the species in captivity have been of limited success. One possible reason for this is that information on the reproductive biology of Tarsius tarsier is extremely limited and data on the species reproductive physiology are completely lacking. We validated fecal estrogen (E-total) and progesterone metabolite (5-P-3OH) measurements for monitoring female ovarian activity and pregnancy. We used this approach to provide the first data on cycle and pregnancy length based on endocrine information in this species. We collected regular fecal samples in combination with observations on socio-sexual behaviors for a maximum of 15 months from three females maintained at Primate Research Center of Bogor Agricultural University, Indonesia. Hormonal profiles indicated that behavioral estrus was associated with marked elevations in fecal E-total concentrations followed by increases in 5-P-3OH levels indicating luteal function. Pregnancy was characterized by low levels of E-total and 5-P-3OH during the first month and markedly rising concentrations thereafter. An ovarian cycle length of 21.7 ± 5.7 days was found. Gestation length was 128d (live infant), 131d (stillbirth), and 164d (death of mother and infant due to dystocia). Despite the small sample size, the study demonstrates the overall validity of fecal sex hormone metabolite measurements for reproductive monitoring in female T. tarsier, as such, the methods described here may ultimately help to improve the breeding management of the species in captivity. They may also offer new opportunities for investigating basic questions of tarsier reproductive biology in the wild by using fecal hormone metabolite analysis to diagnose pregnant animals and determine reproductive rates in relation to ecological and other factors influencing tarsier reproduction. Thus, non-invasive assessment of female reproductive condition as described here may ultimately contribute to facilitate in and ex situ conservation efforts of this endangered primate species.

Sexual Behaviour of the Spectral Tarsier (Tarsius spectrum) in Captivity.

Tarsius spectrum is a primate species endemic to Sulawesi. Populations of the species have decreased due to habitat destruction and hunting. The sexual behaviour of T. bancanus and T. syrichta are known, but that of T. spectrum has not been reported until recently. The aim of this research was to study the sexual behaviour of T. spectrum in captivity. We observed 3 pairs of T. spectrum at the captive breeding facility of the IPB Primate Research Centre for 9 months using focal animal sampling. We showed that principal courtship behaviours were scent marking (36.7%) and genital marking for females (16.2%) and genital inspection for males (16.0%). Copulations lasted between 3 and 4 min, starting with the male mounting the female and thrusting quickly as many as 168-236 times followed by slow thrusting 9-20 times. When slow thrusting occurred, females vocalized up to 6 times. At the end of the copulation sequence, males remained motionless with their penis inserted within the female's genitalia for about 31 s. Copulation occurred only once for each pair during the observation period. Our results should be useful to support breeding programmes and conservation actions for tarsiers.

Impact of Typhoon Haiyan on a Philippine Tarsier Population.

Over the last 2 decades the Philippine tarsier (Carlito syrichta aka Tarsius syrichta) has had its conservation status revised from Endangered to Data Deficient to Near Threatened. The last status change was based on a study of the species' population density, which suggested that a single natural catastrophe could potentially wipe out the Philippine tarsier. In 2013 typhoon Haiyan hit Bohol, one of the island strongholds for this species. In this study we compare the density of the Bohol tarsier population within the Philippine Tarsier and Wildlife Sanctuary before and after the typhoon. We demonstrate that the typhoon significantly affected the density of the Philippine tarsier in the sanctuary. Before the typhoon, tarsier density was approximately 157 individuals/km2 whereas after the typhoon the density was a mere 36 individuals/km2. Prior to the typhoon, more Philippine tarsiers were found in older secondary forest than in younger secondary forest, whereas after the typhoon all observed individuals were found in relatively younger secondary forest. Vegetation plots where we observed Philippine tarsiers prior to the typhoon contained a mean of 33 trees/m2, with a mean diameter at breast height (DBH) of 24 cm, and a mean height of 4 m. After the typhoon vegetation plots contained an average of 156 trees, had a mean DBH of 6 cm, and a mean height of 2 m. Based on the IUCN Red List criteria, the reduction and fluctuation in the density of this species suggests that the conservation status of the Philippine tarsier should be changed to Vulnerable. This study indicates natural disasters can have a significant effect on the extinction risk of primates, with implications for future effects of anthropogenic climate change.

Jumping in the Night: An Investigation of the Leaping Activity of the Western Tarsier (Cephalopachus bancanus borneanus) Using Accelerometers.

Accelerometers enable scientists to quantify the activity of free-living animals whose direct observation is difficult or demanding due to their elusive nature or nocturnal habits. However, the deployment of accelerometers on small-bodied animals and, in particular, on primates has been little explored. Here we show the first application of accelerometers on the western tarsier (Cephalopachus bancanus borneanus), a nocturnal, small-bodied primate endemic to the forests of Borneo. The fieldwork was carried out in the Lower Kinabatangan Wildlife Sanctuary, Sabah, Malaysian Borneo. We provide guidelines for the deployment of accelerometers on tarsiers that might also be applied to other primate species. Our collected data on 2 females show levels of leaping activity comparable to those previously described using direct observation of wild or captive individuals. The 2 females showed different patterns of leaping activity, which calls for work to explore individual differences further. Our work demonstrates that accelerometers can be deployed on small primates to acquire body motion data that would otherwise be demanding to collect using classic field observations. Future work will be focused on using accelerometer data to discriminate in more detail the different behaviours tarsiers can display and to address the causes and consequences of individual variations in activity.

Functional preservation and variation in the cone opsin genes of nocturnal tarsiers.

The short-wavelength sensitive (S-) opsin gene OPN1SW is pseudogenized in some nocturnal primates and retained in others, enabling dichromatic colour vision. Debate on the functional significance of this variation has focused on dark conditions, yet many nocturnal species initiate activity under dim (mesopic) light levels that can support colour vision. Tarsiers are nocturnal, twilight-active primates and exemplary visual predators; they also express different colour vision phenotypes, raising the possibility of discrete adaptations to mesopic conditions. To explore this premise, we conducted a field study in two stages. First, to estimate the level of functional constraint on colour vision, we sequenced OPN1SW in 12 wild-caught Philippine tarsiers (Tarsius syrichta). Second, to explore whether the dichromatic visual systems of Philippine and Bornean (Tarsius bancanus) tarsiers-which express alternate versions of the medium/long-wavelength sensitive (M/L-) opsin gene OPN1MW/OPN1LW-confer differential advantages specific to their respective habitats, we used twilight and moonlight conditions to model the visual contrasts of invertebrate prey. We detected a signature of purifying selection for OPN1SW, indicating that colour vision confers an adaptive advantage to tarsiers. However, this advantage extends to a relatively small proportion of prey-background contrasts, and mostly brown arthropod prey amid leaf litter. We also found that the colour vision of T. bancanus is advantageous for discriminating prey under twilight that is enriched in shorter (bluer) wavelengths, a plausible idiosyncrasy of understorey habitats in Borneo.This article is part of the themed issue 'Vision in dim light'.

Ontogeny of the Postorbital Region in Tarsiers and Other Primates.

Bony structure of the postorbital region is a key trait distinguishing major clades of primates. Strepsirrhines share a postorbital bar, and anthropoids share a complete postorbital septum. At issue is whether the partial postorbital septum of tarsiers unites living tarsiers more closely with anthropoids than with certain large-eyed Eocene fossils. Previously we reported incomplete postorbital closure in tarsiers at birth. In this article, we document comparative analyses of the postorbital region in a broad range of perinatal primates. Virtual reconstructions of microCT data were used to study three-dimensional structure of the perinatal cranium in these taxa. We also describe and illustrate formation of the tarsier partial postorbital septum through the perinatal period using a growth series of Tarsius syrichta. Our results support the hypothesis that partial postorbital septation in the tarsier is secondary to eye hypertrophy. Based on these observations, we propose a structural hypothesis for phylogenetic differences observed in the primate postorbital region. Specifically, we propose that key postorbital traits, including the frontal spur in strepsirrhines and the posterior lamina of the zygomatic in anthropoids, develop as a result of the spatial relationships of brain, eyes, and teeth. Haplorhines are united by expansion of the anterior cranial fossa and loss of the frontal spur. Anthropoids are further united to the exclusion of tarsiers by expansion of the temporal lobes and associated formation of the posterior lamina of the zygomatic. Mechanical forces related to these spatial relationships may be modulated by deep fascia of the orbit to induce formation of the postorbital septum. Anat Rec, 299:1631-1645, 2016. © 2016 Wiley Periodicals, Inc.