A Late Miocene Accipitrid (Aves: Accipitriformes) from Nebraska and Its Implications for the Divergence of Old World Vultures

Background

Old World vultures are likely polyphyletic, representing two subfamilies, the Aegypiinae and Gypaetinae, and some genera of the latter may be of independent origin. Evidence concerning the origin, as well as the timing of the divergence of each subfamily and even genera of the Gypaetinae has been elusive.

Methodology/Principal Findings

Compared with the Old World, the New World has an unexpectedly diverse and rich fossil component of Old World vultures. Here we describe a new accipitriform bird, Anchigyps voorhiesi gen. et sp. nov., from the Ash Hollow Formation (Upper Clarendonian, Late Miocene) of Nebraska. It represents a form close in morphology to the Old World vultures. Characteristics of its wing bones suggest it was less specialized for soaring than modern vultures. It was likely an opportunistic predator or scavenger having a grasping foot and a mandible morphologically similar to modern carrion-feeding birds.

Conclusions/Significance

The new fossil reported here is intermediate in morphology between the bulk of accipitrids and the Old World gypaetine vultures, representing a basal lineage of Accipitridae trending towards the vulturine habit, and of its Late Miocene age suggests the divergence of true gypaetine vultures, may have occurred during or slightly before the Miocene.     

Vulture flight behavior and implications for aircraft safety

Growing vulture populations represent increasing hazards to civil and military aircraft. To assess vulture flight behavior and activity patterns at the Marine Corps Air Station in Beaufort, South Carolina, we equipped 11 black vultures (Coragyps atratus) and 11 turkey vultures (Cathartes aura) with solar-powered Global Positioning System (GPS) satellite transmitters during a 2-year study (1 Oct 2006–30 Sep 2008). Turkey vultures had larger seasonal home ranges than did black vultures, and 2 turkey vultures made round-trips to Florida. Black vultures consistently spent less time in flight (8.4%) than did turkey vultures (18.9%), and black vultures flew at higher altitudes than did turkey vultures in all seasons except summer when altitudinal distributions (above ground level) did not differ. Although we recorded maximum altitudes of 1,578 m for black vultures and 1,378 for turkey vultures, most flights were low altitude. A matrix of vulture flight altitude versus time of day revealed that >60% of vulture flight activity occurred from 4 hr to 9 hr after sunrise at altitudes below 200 m. Continuation of aggressive harassment coupled with flexible training schedules to avoid times and altitudes of high vulture activity will decrease hazards to aircraft posed by these birds. © 2011 The Wildlife Society.     

Breeding of the lappet-faced and white-headed vultures (Torgos tracheliotus Forster and Trigonoceps occipitalis Burchell) on the Serengeti Plains, Tanzania

The lappet-faced and white-headed vulture nests in a 1636 km² study area on the Serengeti Plains were kept under observation from June 1972 to August 1973. Active nests were inspected from a light aircraft at intervals of approximately 2 weeks. The inverse nesting densities were estimated as 43 km²/pair for the lappet-faced vulture and 409 km²/pair for the white-headed vulture. Nests were built of sticks in exposed sites on the crowns of trees, Acacia tortilis being the most popular species. Where the previous history was known, 44% of lappet-faced vulture nests were newly constructed, and the rest re-used from the previous year. The incubation period was about 55 days and the fledging period about 100 days in both species. Juveniles continued to frequent their nests after they could fly, sometimes for several months. Most lappet-faced vultures laid their eggs in April-May, the white-headed vultures in June-August. This means that the lappet-faced vultures raised their young during the dry season, when Grant's and Thomson's gazelles were the only animals present in any numbers on the plains. By contrast, the Ruppell's griffons nesting in the nearby Gol Escarpment colony raise their young during February-May, when the wildebeest and zebra herds are normally on the plains. The survival rate of lappet-faced vultures from laying to fledging was about 40%. The chicks are thought to be vulnerable to predation by tawny eagles, and one was known to have been killed by a leopard. Three adult vultures were killed on their nests, from unknown causes. Food remains from lappet-faced vulture nests showed a heavy preponderance of Grant's and Thomson's gazelles in all cases, with golden jackal the next most commonly occurring species. No food remains were obtained from white-headed vulture nests. The possibility of using aerial counts of vulture nests as an indicator of productivity is discussed.     

Communal farmers of Namibia appreciate vultures and the ecosystem services they provide

Ecosystem services are cited as one of the many reasons for conserving declining vulture populations in Africa. We aimed to explore how communal farmers in Namibia perceive vultures and the ecosystem services they provide, with special focus on cultural and regulating ecosystem services. We surveyed 361 households across Namibia’s communal farmlands and found that over two-thirds of households liked vultures and found them useful, stating that they were harmless and useful for locating dead livestock. The minority of households who disliked vultures believed that they were killing their livestock. Poisoning was the main cause of vulture mortalities reported by farmers. While poisoning appears to be a concern for vultures in the communal farmlands, it appears that cultural use of vulture body parts is a minimal threat. We found that few farmers knew of cultural beliefs about vultures or uses for body parts; most farmers believed these beliefs and practices to be outdated. It is further promising that communal farmers have an overall positive perception of vultures. This highlights the potential for communal conservancies to bring attention to vulture conservation in their constituencies.     

Implications of bacterial,viral and mycotic microorganisms in vultures for wildlife conservation,ecosystem services and public health

The effects that microorganisms (bacteria, viruses and fungi) have on their hosts remain unexplored for most vulture species. This is especially relevant for vultures, as their diet consists of carcasses in various stages of decomposition, which are breeding grounds for potentially pathogenic microorganisms. Here we review current knowledge of bacterial, viral and mycotic microorganisms present in wild vultures. We consider their potential to cause disease in vultures and whether this poses any population-level threats. Furthermore, we address the question of whether vultures may act as disease spreaders or mitigators. We found 76 articles concerning bacterial, viral and mycotic microorganisms present in 13 vulture species, 57 evaluating bacteria, 13 evaluating viruses and six evaluating mycotic microorganisms. These studies come from all continents where vultures are present, but mainly from Europe and North America, and the most studied species was the Griffon Vulture Gyps fulvus. We found that vultures are colonized by zoonotic pathogens, and even host-specific human pathogens. Some recorded bacteria showed multi-antibiotic resistance, especially those that can be associated with anthropogenic food subsides such as supplementary feeding stations. We found evidence that vulture health can be affected by some microorganisms, producing a wide array of clinical alterations that have the potential to influence mortality risk and fitness. We did not find clear scientific evidence that vultures play an epidemiological role spreading microorganisms to humans and other species. However, there is evidence that vultures could prevent the spread of infectious diseases through their removal of decomposing organic material. The evaluation of vulture exposure to microorganisms is of fundamental importance to design better conservation policies for this threatened group, which may serve a key role as ecosystem cleaners.     

A Global Review of Causes of Morbidity and Mortality in Free-Living Vultures

Vulture species worldwide play a key role in ecosystems as obligate scavengers, and several populations have had precipitous declines. Research on vulture health is critical to conservation efforts including free-living vultures and captive breeding programs, but is limited to date. In this systematic review, we determined the reported causes of free-living vulture species morbidity and mortality worldwide. The most commonly reported cause of mortality was from toxins (60%), especially lead and pesticides, followed by traumatic injury (49%), including collisions with urban infrastructure and gunshot. Neglected areas of research in free-living vulture health include infectious diseases (16%), endocrine and nutritional disorders (6%), and neoplasia (<?1%). Almost half of the studies included in the review were conducted in either Spain or the USA, with a paucity of studies conducted in South America and sub-Saharan Africa. The highest number of studies was on Griffon (Gyps fulvus) (24%) and Egyptian vultures (Neophron percnopterus) (19%), while half of all vulture species had five or fewer studies. Future investigations on free-living vulture health should focus on neglected areas of research, such as infectious diseases, and areas with gaps in the current literature, such as South America, sub-Saharan Africa, and under-studied vulture species.

     

Bone crushing carnivores and their significance to osteodystrophy in griffon vulture chicks

The natural removal of 89 ungulate carcasses by predators and scavengers was monitored in various wildlife reserves and ranching areas of South Africa. Carnivores responsible for the bone remains were determined. Spotted hyaenas and, to a lesser extent, Brown hyaenas, were the only carnivores that regularly chewed bones. When hyaenas were absent, months of weathering were required before the smaller bones became disarticulated and able to be removed by vultures. The bone-collecting behaviour and related aspects of breeding of two species of griffon vulture were studied at five different nesting colonies in southern Africa—one Cape vulture colony and one White-backed vulture colony in or near wildlife reserves, as well as two Cape vulture colonies and one White-backed vulture colony in ranching land. A total of 2825 bones was found in or below the vulture nests. These bones were categorized and measured. Hyaena-produced bone fragments were found only in the colonies in the wild areas—none of the 387 chicks examined here had osteodystrophy (metabolic bone disease). By contrast, in the ranching areas, vultures collected larger and less fragmented bones. Many Cape vulture chicks had osteodystrophy (130 of 1917 examined), as did two White-backed vulture chicks (of 196 examined). In 1977, artificial feeding stations, 'restaurants' where carcass skeletons were crushed, were established for Cape vultures. Since then, the incidence of osteodystrophy has declined from 17% to 2–5% in 1983. It is clear that bone fragments are an essential dietary requirement, providing calcium for correct skeletal growth of griffon vulture chicks.     

A new species of Macaranga from the middle Miocene of Fujian,China and its significance

A well-preserved Macaranga leaf fossil from the middle Miocene Fotan Group of Zhangpu County, Fujian, South-eastern China is examined and described as a new species, Macaranga zhangpuensis Z.X. Wang et B.N. Sun sp. nov. The present fossil species represents the highest latitudinal distribution of a reliable Macaranga fossil in the world, and we present the first Macaranga fossil described with detailed cuticular characteristics from China. Based on the global palaeogeographic distribution of Macaranga, we infer that the genus probably originated during the Oligocene in Africa and spread from Africa to India and then to South Fujian, China, further into the Mariana Islands and finally into the Philippine Islands and Malaysia, leading to its present distribution. In addition, the new leaf material described herein is the first Macaranga fossil record with three leaf tips globally. This finding demonstrates that Macaranga with three leaf tips already existed during the Miocene in Fujian, South China, and it provides new information for understanding climatic changes between the Miocene and the present -day.     

Significant population of Egyptian Vulture Neophron percnopterus found in Morocco

The Egyptian Vulture Neophron percnopterus population in Morocco has undergone a marked decline since the 1980s to the point of nearing local extinction in the twenty-first century. A field study of some possible sites for Egyptian Vultures was carried out over six days during June 2014 in the Middle Atlas Mountains, Morocco. We counted a total of 48 Egyptian Vultures at three different localities: two occupied breeding sites and one communal roost that hosted 40 vultures of different ages. A (probable) singe adult bird at the breeding site was located and a previously occupied site was also visited. A preliminary survey amongst local people indicated that threats faced by this species are predator poisoning in some areas, and the use of vulture parts for traditional medicine. Given that the species is considered globally Endangered and populations continue to decline in many areas, the discovered population reported here, although relatively small, is of national and regional (North-west Africa) importance. We expect this new situation will revive the hopes for studying and conserving this and other vulture species in Morocco and North-west Africa in general.     

When hunting benefits raptors: a case study of game species and vultures

Management of hunting activity to serve as a tool for sustainable development has become a key issue in conservation biology. However, little evidence is available showing positive impacts of hunting on ecosystem conservation, limiting its capability to be used as a conservation tool. We analysed hunting and its positive influence on the ecology and conservation of the griffon vulture (Gyps fulvus), a scavenger with a relevant function in the ecosystem, in the Cantabrian Mountains, NW Spain. Use of the area by vultures was addressed by looking for cliffs used as roosts or colonies, and consumption of game species by vultures was evaluated through field surveys and questionnaires to hunters. Results revealed a strong spatiotemporal adjustment in the use of the area by vultures and hunting events, especially of red deer and wild boar. Vultures occupied roosting sites very close to the main hunting sectors of these game species and often were seen consuming their carcasses. The spatiotemporal pattern of roost use by vultures strongly overlapped with hunting of red deer. The numbers of both red deer and wild boar hunting episodes within 3.5 km around the roosts were the best predictors of vulture occurrence and number. Our estimates show that hunting could feed around 1,800 vultures/6 months. Hunting can thus influence species at the top of the ecosystem (scavengers) and could aid sustainable management of griffon vulture populations, reconciling hunting and conservation. However, negative and positive impacts should be taken into account simultaneously for an overall evaluation of hunting on ecosystem conservation.     

Hematozoa and a new haemoproteid species from Cathartidae (New World vulture) in South Carolina

A survey of turkey vultures (Cathartes aura) in South Carolina revealed the presence of a novel haemoproteid. Haemoproteus catharti n. sp. is described from the slides developed from the blood of this vulture. The new species is a thick, halteridial form with complete margins, and is considered distinct from the other recognized haemoproteids from the diurnal raptors in shape, pigment number, parasite outline, and host-family specificity. While reviewing blood films, a series of unusual immature schizonts of Plasmodium sp. were also observed, and these are illustrated, along with another distinct haemoproteid from old slides of poor quality produced from turkey vulture blood.     

内蒙古中部敖尔班地区的岩石及生物地层(英文)

中国北方新近纪地层中,早中新世的哺乳动物化石地点比较稀少,化石面貌也不十分清楚。在内蒙古中部,尽管通古尔台地上有研究历史近百年的新近纪经典地点,但上世纪90年代发现的嘎顺音阿得格,依然是内蒙古目前惟一产出早中新世动物群的小盆地。同内蒙古诸多其他新近纪哺乳动物群一样,嘎顺音阿得格动物群与相邻动物群彼此缺乏时空上的联系,还不易建立可靠的层序关系。位于苏尼特左旗东南约60 km的敖尔班(曾用名"奥尔班",见Liddicoat et al.,2007),红色地层大面积出露,是2004年发现的一个新的哺乳动物化石地点。敖尔班剖面总厚50余米,大、小哺乳动物化石共生,岩性特征易识别,有利层位对比,是研究生物地层学的理想地点。自2004年以来,我们连续在敖尔班地区采集化石并进行地层工作,建立了一个较完整的哺乳动物序列,时代跨越早中新世晚期到晚中新世晚期。更加难得的是,敖尔班同一剖面上含有4个哺乳动物化石层位,其上下层序关系一目了然,在内蒙古新近纪地层中仅此一例。敖尔班哺乳动物序列的建立,无疑将对整个内蒙古中部新近纪,尤其是早中新世动物群面貌的了解具有促进作用。本文着重对敖尔班岩石地层进行描述,并结合我们4年来对脊椎动物化石的积累与认识,试图进一步完善内蒙古中部地区的生物地层层序,进而建立这一地区的年代地层框架。对化石的详细描述将适时发表。根据岩性及接触关系,敖尔班剖面可分为三大段:敖尔班组、巴伦哈拉根层及必鲁图层。敖尔班组(新建组)由一套红色和绿色的泥岩及粉砂岩组成,发育有成熟的古土壤层,厚约42 m,时代属于早中新世中晚期。敖尔班组的层型剖面建立在敖尔班露头出露最厚的中部,下部未见底,顶部与上覆巴伦哈拉根层呈假整合或不整合接触。该组目前已知分布仅局限于敖尔班露头。敖尔班组可进一步划分出三段:下红泥岩段、中绿泥岩段及上红泥岩段。三段呈连续沉积。敖尔班组与巴伦哈拉根层之间的假整合所代表的沉积间断延续了中中新世的大部分时段。巴伦哈拉根层为一套橘红色砂岩、粉砂岩及底砾岩,时代大致是最晚中中新世至最早晚中新世。不整合于巴伦哈拉根层之上是必鲁图层。两层之间似乎缺失了晚中新世的大部分堆积。必鲁图层的底砾岩是一种切割与充填构造,其中所含钙质结核及相当数量的化石都可能是巴伦哈拉根层原生堆积物再沉积的结果。必鲁图层的分布还需做更多工作,其时代可能是晚中新世晚期。主要依据小哺乳动物的组合,在敖尔班剖面中可建立4个动物群。最早为敖尔班组下红泥岩段产出的下敖尔班动物群。该动物群的特征是,小哺乳动物中在渐新世十分兴旺的一些科,如Ctenodactylidae,Tachyoryctoididae,Aplodontidae和Zapodidae还相当繁荣;中新世出现的属,如Mioechinus,Keramidomys,Heterosminthus和Democricetodon等占动物群总量的半数以上;大哺乳动物中残留有Palaeogale和裂爪兽。比下敖尔班动物群稍晚的是上敖尔班动物群,产出于敖尔班组的上红泥岩段。上敖尔班动物群的特征是,小哺乳动物在渐新世中占统治地位的一些科或完全绝迹,如Ctenodactylidae,或在种类和数量上明显减退,如Aplodontidae和Zapodidae;缺少在下敖尔班动物群中还相当常见的一些古老属,如Amphechinus,Tachyo- ryctoides和Sinolagomys等;出现了下敖尔班动物群中所没有的Megacricetodon,Cricetodon和Alloptox属;大哺乳动物中出现了长鼻类和柄杯鹿(Ligeromeryx/Lagomeryx)。经过相当长的一个沉积间断,敖尔班剖面的上部出现了巴伦哈拉根动物群。该动物群中渐新世常见的小哺乳动物科进一步衰落,同时出现了亚洲古北界晚中新世以后常见的跳鼠科(Dipodidae)和鼢鼠科(Siphneidae),具有明显的中中新世晚期或晚中新世早期生物组合的特点。最后是敖尔班剖面顶部必鲁图层中的必鲁图动物群。必鲁图动物群显然带有中中新世及晚中新世的混合特征,很可能是水流作用再沉积的结果。根据其中最进步分子的成分判断,估计必鲁图动物群的年代是晚中新世的晚期,其特征是,鼠科(Muridae)动物高度分化,而野兔科(Leporidae)尚未出现;含有晚中新世宝格达乌拉动物群或最晚中新世二登图动物群中大量出现的属种,如Lophocricetus grabaui,Paralactaga suni,Dipus fraudator和Hansdebruijnia pusilla等;但二登图动物群中很繁荣的一些属在必鲁图动物群中未被发现或者发现的个体数量很少,如Prospermophilus, Paralophicricetus和Microtodon等。     

Toxicity of non-steroidal anti-inflammatory drugs to Gyps vultures: a new threat from ketoprofen

Three Gyps vulture species are on the brink of extinction in South Asia owing to the veterinary non-steroidal anti-inflammatory drug (NSAID) diclofenac. Carcasses of domesticated ungulates are the main food source for Asia''s vultures and birds die from kidney failure after consuming diclofenac-contaminated tissues. Here, we report on the safety testing of the NSAID ketoprofen, which was not reported to cause mortality in clinical treatment of scavenging birds and is rapidly eliminated from livestock tissues. Safety testing was undertaken using captive non-releasable Cape griffon vultures (Gyps coprotheres) and wild-caught African white-backed vultures (G. africanus), both previously identified as susceptible to diclofenac and suitable surrogates. Ketoprofen doses ranged from 0.5 to 5 mg kg⁻¹ vulture body weight, based upon recommended veterinary guidelines and maximum levels of exposure for wild vultures (estimated as 1.54 mg kg⁻¹). Doses were administered by oral gavage or through feeding tissues from cattle dosed with ketoprofen at 6 mg kg⁻¹ cattle body weight, before slaughter. Mortalities occurred at dose levels of 1.5 and 5 mg kg⁻¹ vulture body weight (within the range recommended for clinical treatment) with the same clinical signs as observed for diclofenac. Surveys of livestock carcasses in India indicate that toxic levels of residual ketoprofen are already present in vulture food supplies. Consequently, we strongly recommend that ketoprofen is not used for veterinary treatment of livestock in Asia and in other regions of the world where vultures access livestock carcasses. The only alternative to diclofenac that should be promoted as safe for vultures is the NSAID meloxicam.     

山东山旺中中新世鳅科化石

本文描述的较完整的花鳅 Cobitis 化石在我国尚属首次发现.花鳅属是花鳅亚科(Cobitinae)中较进步的鱼类,广布于欧洲、亚洲和非洲,已知化石则产自欧洲和亚洲的第三系.长胸鳍花鳅 Cobitis longipectoralis sp. nov. 的发现为探讨鳅科起源及其地理分布提供了新的资料.     

Lappet-faced vultures (Torgos tracheliotus) hatch and rear a White-headed vulture (Trigonoceps occipitalis) in the wild

In Gonarezhou National Park, southeast Zimbabwe Rhodesia, White-headed vultures laid their egg in a nest of Lappet-faced vultures in June 1978. Lappet-faced vultures then incubated the egg and reared the nestling successfully to fledging. The chick's feathers however had "fault bars", and during seven days of observation it received less than one visit per day by its foster parents. In the same area in 1973 a White-headed vulture was found incubating its egg in a Lappet-faced vulture style nest. The sizes of these eggs (811 times 650 mm and 79–9 times 65–8 mm) and their estimated dates of laying (24 June 1973 and 21 June 1978) were almost identical, suggesting that the same pair of White-headed vultures was involved.     

African Vultures Don’t Follow Migratory Herds: Scavenger Habitat Use Is Not Mediated by Prey Abundance

The ongoing global decline in vulture populations raises major conservation concerns, but little is known about the factors that mediate scavenger habitat use, in particular the importance of abundance of live prey versus prey mortality. We test this using data from the Serengeti-Mara ecosystem in East Africa. The two hypotheses that prey abundance or prey mortality are the main drivers of vulture habitat use provide alternative predictions. If vultures select areas based only on prey abundance, we expect tracked vultures to remain close to herds of migratory wildebeest regardless of season. However, if vultures select areas where mortality rates are greatest then we expect vultures to select the driest regions, where animals are more likely to die of starvation, and to be attracted to migratory wildebeest only during the dry season when wildebeest mortality is greatest. We used data from GSM-GPS transmitters to assess the relationship between three vulture species and migratory wildebeest in the Mara-Serengeti ecosystem. Results indicate that vultures preferentially cluster around migratory herds only during the dry season, when herds experience their highest mortality. Additionally during the wet season, Ruppell’s and Lappet-faced vultures select relatively dry areas, based on Normalized Difference Vegetation Index, whereas White-backed vultures preferred wetter areas during the wet season. Differences in habitat use among species may mediate coexistence in this scavenger guild. In general, our results suggest that prey abundance is not the primary driver of avian scavenger habitat use. The apparent reliance of vultures on non-migratory ungulates during the wet season has important conservation implications for vultures in light of on-going declines in non-migratory ungulate species and use of poisons in unprotected areas.     

Demographics of Black Vultures in North Carolina

Abstract: Understanding the contributions of vital rates to species population growth is critical to developing new management protocols. We constructed a model population for black vultures (Coragyps atratus) in North Carolina, USA, based on demographic data from a 14-year study. The model population was similar in stage structure to the reference population, and adult survival was the primary contributor to the annual rate of increase (10.6%). We suggest that the North Carolina black vulture population is experiencing high rates of survival and fertility, potentially breeding at an age younger than previously assumed, and growing rapidly.     

The diversity and paleoenvironmental significance of Calophyllum (Clusiaceae) from the Miocene of southeastern China

Abstract

Three species within the genus Calophyllum collected from middle Miocene Fotan Group sediments in Zhangpu County, Fujian, southeastern China are described in this paper. These fossils include Calophyllum zhangpuensis sp. nov., Calophyllum striatum, and Calophyllum suraikholaensis. The new fossil species C. zhangpuensis sp. nov. is oval, possesses entire leaves with closely spaced parallel secondary veins and has a round, or slightly retuse, apex. These specimens represent the first known fossil records of this relative wide leaf-type form of Calophyllum from China and have a length:width (L:W) ratio less than 3:1. In combination with the known modern geographic distribution and habitats of this wide leaf-type Calophyllum and other plants, data suggest that the middle Miocene Fotan flora is indicative of a warm climate. Thus, based on available fossil data, we speculate that this genus probably originated in India during the Paleocene before spreading from India to Bangladesh and into China, Sumatra, Malaysia, Indonesia, and Java during the Neogene, leading to its modern distribution. At least, the 3 fossil species in this region can explain floristic exchange between India, Fujian, and South China, which is consistent with previous studies; the occurrence of these 3 species indicates that Calophyllum began to diversity in China no later than the Miocene.     

Rapid population declines of Egyptian vulture (Neophron percnopterus) and red-headed vulture (Sarcogyps calvus) in India

Since the early 1990s, large and rapid population declines of three species of vulture ( Gyps spp.) endemic to south Asia have occurred on the Indian subcontinent and have led to these species being listed by IUCN as critically endangered. Evidence of rates of population decline, cause of death and toxicity is consistent with these declines being caused by poisoning of vultures through the ingestion of tissues from livestock treated with the anti-inflammatory drug diclofenac. In this paper, analysis of repeated surveys in and near protected areas widely spread across India shows that populations of two other vulture species, Egyptian vulture Neophron percnopterus and red-headed vulture Sarcogyps calvus , have also declined markedly and rapidly, but probably with a later onset than Gyps vultures in the same region. The declines continued at least up to 2003. It is recommended that these two species are considered for inclusion in the IUCN Red List and for urgent remedial conservation measures. Research is needed to determine whether or not the principal cause of these declines is diclofenac poisoning and to establish population trends in other scavenging birds in the Indian subcontinent.