The type specimens and type localities of the orangutans, genus Pongo Lacépède, 1799 (Primates: Hominidae)

ABSTRACT Uncertain type localities undermine orangutan nomenclature. Bequeathed to the British Museum, the holotype of Pongo pygmaeus, according to Hans Sloane’s catalogue, came from Borneo and died in China. The historical evidence makes Banjarmasin its most probable type locality. William Montgomerie, Assistant Surgeon at Singapore from 1819–1827, and Senior Surgeon from 1832, supplied the holotype of Simia morio. In 1836 an adult female orangutan reached Singapore alive from Pontianak, Borneo. The holotypes of S. morio, S. hendrikzii, S. straussii and P[ithecus] owenii probably had the same origin, as pirate attacks endangered visits to other Bornean coasts. Absent from Brunei and north Sarawak, Malaysia, throughout the Holocene, orangutans occur there only as Pleistocene subfossils at Niah. Pan vetus (the Piltdown mandible) probably came from Paku, Sarawak. We identify Pongo borneo Lacépède, 1799 as an objective senior synonym of P. wurmbii Tiedemann, 1808, correcting its type locality from Sukadana to near Pontianak. This is the earliest name for the western subspecies (previously thought nominotypical) unless Pithecus curtus, probably from the Sadong River, Sarawak, represents a separate subspecies. If so, the name Pongo borneo would transfer to the southern population west of the Kahayan River, genetically distinguished at species level from the Sumatran orangutan, P. abelii.


Introduction
Effective international collaboration in wildlife conservation requires an agreed scientific nomenclature. Brandon-Jones (2006) cited the orangutan, genus Pongo Lacépède, 1799 as a high-profile, threatened primate with unsettled nomenclature. Extant orangutans are endemic to Borneo and Sumatra, but which of these SE Asian islands produced the holotype has remained uncertain. Evidence that the orangutan comprises two species of unconfirmed scientific name and uncertain geographic division accentuates this deficiency. Morphological characters have not distinguished whether the two species are parapatric in central Borneo or separated by a sea barrier (Brandon- ). from the bearers, 'the young Gentlemen' James Lyde and Charles Lockyer, their letter of introduction by Charles duBois (c. 1657-1740), dated 18 February 1702. Lyde was sponsored by Robert Harley, Speaker of the House of Commons; Lockyer by another MP, Sir Francis Wyndham (Jessop 1989). This was therefore probably Lockyer's debut with the company and his first opportunity to obtain the holotype.
After a short stay in Aceh in about May 1704, Lockyer (1711), whose book does not mention the orangutan, moved to Malacca (now Melaka in West Malaysia) where he was preoccupied with commerce. Lockyer (1711, p. 180) spent four months in Canton (now Guangzhou, China). On 17 December 1704, he embarked on the Stretham for Mocho (now Al Mukha, Yemen) in the Red Sea (p. 191), briefly calling at Melaka, Anjengo (near Thiruvananthapuram, India), and Callicut Road (now Kozhikode, India). Through poor seamanship or adverse winds (he could not discover which, p. 202) their course was diverted to Gombroon (now Bandar Abbas, Iran), via Muscat (now Masqat, Oman).
On 27 October 1705, heavily laden with freight and passengers, the Stretham sailed for Surat, India, arriving on 24 November. Lockyer (1711, p. 255) had been suffering from 'Quotidian Ague' (malaria with daily bouts of fever) for the previous 10 weeks and on the Malabar Coast contracted 'Flux' (dysentery) for two months, restricting his ability to document local trade. Leaving Surat on 20 December (p. 265), the Stretham reached Goa by Christmas. It departed Goa on 29 December, arrived at Carwar (now Karwar) on 30 December (p. 269), left on 11 January 1706, arrived at Kozhikode on 18 January, then returned to Telichery (now Thalassery) on 25 January, revisiting Kozhikode on 1 February (p. 274). At Thalassery, black monkeys (Macaca silenus or Semnopithecus johnii), if tame, were sold for 10-12 rupees (up to 3 German or Venetian ducats) each (p. 276). (On St. Helena, a Venetian ducat was worth 9 English shillings, p. 307). 'The Monkeys about Calicut are larger than ordinary, and keep in great Companys in the Woods. It is very diverting to see with what Agility they jump from Tree to Tree. I cannot tell if we are allow'd to shoot them; the Gentoos [= Hindus] in other Places having a religious Regard for them ' (p. 282). The Stretham left Kozhikode on 10 February, took on provisions at Cocheen (now Kochi) from 12 to 15 February (p. 283-284), and reached the Cape of Good Hope on 22 May, waiting until 12 July for a convoy to accumulate (p. 288). It stopped at St. Helena from 5 to 14 August (p. 304,307), laying up for two days from 3 September 1706 for wood and water at the small, uninhabited island of John Fernando (perhaps Fernando de Noronha), off the coast of Brazil. Frustratingly, Lockyer's (1711) book ends without mentioning acquisitions, his port of disembarkation, date of arrival, or his occupation in the five years before its publication.
The Journal Book of the Royal Society of 18 May 1727 recorded that on 7 December 1713, a Lockyer (forename omitted) was 'Accomptant to the South Sea Company' when Council approved his election as a Fellow, proposed by Dr (Alexander) Stewart (c. 1673Stewart (c. -1742. On Monday 19 March 1733, the 'Place of Accomptant-General of the S. Sea Company, lately possess'd by Charles Lockyer, Esq; was by the Directors voted useless; and several Clerks, for the same Reason, were discharged' (Anon 1733, p. 154). Beatson (1807, p. 157) recorded that this Charles Lockyer (named 'Chief Accomptant') was Member of Parliament (MP) for Ilchester, Somerset, from the 7th to the 9th Parliaments, i.e. from 28 November 1727 to 18 June 1747 (Beatson 1807, p. vi-vii).
Robert Harley (1661Harley ( -1724, James Lyde's sponsor for his post at Fort St. George, was Speaker from 1701 to 1705, Secretary of State for the Northern Department from 1704 to 1708 and Chancellor of the Exchequer in 1710, when he was created Earl of Oxford and Lord Treasurer in 1711. In May 1711 he co-founded the South Sea Company to fund government debt incurred during the War of the Spanish Succession. British participation (allied with the Netherlands) in the war, begun in 1701, ended in 1713 with the Treaty of Utrecht (Speck 2004). Sir Francis Wyndham (c. 1653Wyndham (c. -1715, Charles Lockyer's sponsor, was MP for Ilchester from 1695 to the parliament commencing 21 July 1702 (Watson and Hanham 2002). Thomas Lockyer (1699-1785 was the leading businessman in eighteenth-century Ilchester. His family had been prosperous mercers and prominent town members since the early seventeenth century (Baggs et al. 1974b). Charles' death notice of 12 February 1752 named Thomas as his brother (Anon 1752). The discrepancy between this date of death and the 13 April 1752 given by the Royal Society might be deemed to refute this Lockyer as the author of An account of the trade in India, but 1752 was the year in which Britain switched from the Julian to the Gregorian calendar (Jessop 1989, p. 15). This or confusion over the date of death and date of notice of death can account for the discrepancy.
In 1722 Charles Lockyer, MP, bought Holly House and in 1728 Coldhall (or West Ealing manor at Little Ealing) and Twyfords. His illegitimate son John Lockyer or Green succeeded him in 1752, and left the estate to his widow Elizabeth and daughter Elizabeth in 1762 (Bolton et al. 1982). Charles Lockyer locally introduced the gypsy moth (Lymantria dispar) (Wilkes 1773, p. 20) and at his Ealing house bred numerous green-brindled crescent moths (Allophyes oxyacanthae) or 'Ealing's Glory', being the only person Wilkes (1773, p. 12) knew who had seen many. In 1718 Charles Lockyer 'of London' bought part of the manor of Charlton Mackrell (near Ilchester), bequeathing it in 1752 to John Lockyer 'of Colehall in Ealing', who sold it in 1759 to (his uncle) Thomas Lockyer 'of London (later of Ilchester)' who had meanwhile purchased the rest of the manor. Thomas bequeathed the reunited manor to his daughter Mary (Baggs et al. 1974a).
John Lockyer is buried at St Mary's parish church, Ealing (Lysons 1795). The last resting place and date of birth of his father remain a mystery. Large age gaps between siblings were not then uncommon, but his description as a young man in 1702 and Thomas' birth in 1699 suggest that Charles was born in about 1685, although he could have been younger, as Sir Thomas Stamford Raffles (1781-1826), for example, joined the East India Company as a clerk aged 14 (Turnbull 2004).
If Lockyer left the East India Company to join the fledgling South Sea Company in 1711, this could explain why his book was issued in a hurried fashion with many loose ends, notably no attempt to bring it up to date. In the interim it had presumably gathered dust and Lockyer realised that he must publish or abandon the project. The South Sea Company's business, exclusively with Central and South America, would provide scant chance to supplement the book's contents. His employment could also explain why so little is known of Lockyer's life after 1711. Initially, as the South Sea Company inflated, he would probably have been inundated with work. After the crash, and perhaps especially after his enforced redundancy in 1733, he probably would have preferred to gloss over his previous occupation, especially as luminaries, such as Sir Isaac Newton (1642-1727), reputedly lost £20,000 in the speculation (Westfall 1981, p. 861). If this supposition is all correct, then the orangutan holotype was probably obtained before 1711.
The type locality of Pongo pygmaeus determines its application. Edwards (1758) presumed that the holotype came from Africa. Without corroboration, Wallace (1869, p. 144) and Elliot (1913, p. 188) claimed that the orangutan was first discovered in Sumatra. Elliot (1913, p. 188, 192, note 194) contended that if the Sumatran population were separated, the name P. pygmaeus would become geographically indeterminable, but he retained it, believing that the orangutan's exceptional individual variation precludes subspecific recognition (Elliot 1913, p. 195). Groves and Holthuis (1985) were satisfied that the holotype came from Borneo, but Röhrer-Ertl (1988) supposed that intense colonial rivalry between Britain and the Netherlands favours Sumatra as the origin of a British-owned orangutan in 1758.
At that time, however, Britain was not without access to Borneo. English trade with Banjarmasin (now in South Kalimantan province, Indonesia; see Figure 2) began in 1609. About 40 Englishmen and 100 Bugis (a seafaring people from southern Sulawesi, Indonesia, probably ultimately from southern China via Taiwan) settled there circa 1700, sacking five native settlements, 'Banjar, Banjarmassing, Kayu-tangi, Tatas and Martapura', after friction in the first year. A truce reduced personnel to 10 Englishmen and 40 Bugis. Hostilities resumed in 1706, and the English were evicted. The Dutch only occasionally visited from about 1644, with no permanent base there until 1747 (Leyden 1814, p. 23-24).

Captain Daniel Beeckman's voyage
That an eighteenth-century Englishman could have obtained a Bornean orangutan is confirmed by the purchase of one by Captain Daniel Beeckman (1718). Edwards (1758) regarded this animal as 'near of kin' but distinct from Sloane's orangutan. Beeckman's (1718) description being too brief (see below), this inference must derive from Beeckman's (1718) figure (reproduced in Morris and Morris 1966, p. 133), evidently by an artist who had never seen a live orangutan. Instead, it is a reanimated and more anthropomorphic modification of Tulp's (1641, pl. 14) figure (reproduced in Stiles and Orleman 1927, p. 7) of an anthropoid ape presented to stadtholder Frederick Henry (1584-1647), Prince of Orange. This latter ape is usually considered a chimpanzee from Angola, but Rijksen and Rijksen (1988, p. 9) noted that there is an 'Angkola' region (now Mandailing) in north Sumatra. The black pelage and large ears Tulp (1641) mentioned indicate a chimp, although Rijksen and Rijksen (1988) contended that the posture, potbelly, sparse ventral hair and rather flat, virtually browless face all typify an orangutan, an impression which is strengthened by the apparent lack of nails on the foot; also absent are the long pointed fingers and characteristic knuckle-pads of a chimpanzee. (The hair distribution might be unreliable, however, as the heavily depilated limbs indicate that hair has sloughed off from partial decomposition.) Leaving the 'Downs' on 12 October 1713 and returning on 29 October 1715, Beeckman (1718, p. 2, 205) headed fairly directly to and from his destination: Banjarmasin, Borneo. Passing between Teneriffe and Grand Canary Island on 1 November (p. 3), the Eagle-Galley halted at the Cape Verde Islands from 12-16 November for water, supplies and ship repairs (p. 10-15). Sighting ' Table-Land' (now Cape Town, South Africa) on 20 January 1714 (p. 18), Christmas Island on 5 April (p. 19) and Java on 6 April, Beeckman (1718, p. 20-21) anchored on 13 April at the uninhabited Mew Island (now Pulau Peucang in Ujung Kulon National Park (NP)) in the Sunda Straits to restock at a coastal town (see Figure 1). Covertly provisioning and refitting the ship at Batavia, reached on 20 April, took 42 days as the Dutch general singled out the English amongst Europeans for denial of basic necessities (p. 22). On 2 June, in convoy with the Borneo and the Daulbin, the Eagle-Galley sailed along the north coast of Java (p. 27), anchoring at the small island of Carimon-Java (now Karimunjawa) from 14 (p. 31) to 17 June. The Eagle-Galley and Borneo reached Banjarmasin at midnight on 29 June 1714 (p. 32).
From 28 December 1714 to 5 January 1715 Beeckman (1718, p. 155-160) reprovisioned off the eastern tip of Madura. He investigated the Bali Strait as a homeward route, taking 15 days to advance 30 leagues (less than 150 km) owing to the skirting mountains funnelling the southerly winds (p. 164-166). From 16-23 January, Beeckman (1718, p. 167-175)  Reporting on the Bornean fauna, Beeckman (1718, p. 37-38) wrote: The Monkeys, Apes, and Baboons are of many different Sorts and Shapes; but the most remarkable are those they call Oran-ootans, which in their Language signifies Men of the Woods: These grow up to be six Foot high; they walk upright, have longer Arms than Men, tolerable good Faces (handsomer I am sure than some Hottentots that I have seen) large Teeth, no Tails nor Hair, but on those Parts where it grows on humane Bodies; they are very nimble footed and mighty strong; they throw great Stones, Sticks, and Billets at those Persons that offend them. The Natives do really believe that these were formerly Men, but Metamorphosed into Beasts for their Blasphemy. They told me many strange Stories of them, too tedious to be inserted here. I bought one out of curiosity, for six Spanish Dollars; it lived with me seven Months, but then died of a Flux; he was too young to show me many Pranks, therefore I shall only tell you that he was a great Thief, and loved strong Liquors; for if our Backs were turned, he would be at the Punch-bowl, and very often would open the Brandy Case, take out a Bottle, drink plentifully, and put it very carefully into its place again. He slept lying along in a humane Posture with one Hand under his Head. He could not swim, but I know not whether he might not be capable of being taught. If at any time I was angry with him, he would sigh, sob, and cry, till he found that I was recon[ci]led to him; and tho' he was but about twelve Months old when he died, yet he was stronger than any Man in the Ship. Beeckman (1718) alluded no further to the orangutan [unless it was the pet monkey which he claimed (p. 104-105) to have been instantly revived by bloodletting of the forearm when reduced to a stupor by a cannon salute on his ship in the Barito estuary]. He did not explicitly purchase his orangutan on this voyage. The only certainty is that its purchase predated his book. However, the East India Company had commissioned him to try to re-establish trade with Banjarmasin. Former attempts by both the English and Dutch had failed (p. 2). Beeckman's (1718, p. 90) was the first English visit since the eviction. To avoid being refused trade, his party feigned dissociation from the East India Company (p. 49). The occupants of a small proa invited aboard (p. 47) were astonished at the English return after their violent expulsion when many were killed and the fort and accommodation destroyed (p. 2). Beeckman (1718, p. 34) believed Borneo the largest island 'in the whole World, except perhaps California'. Other than 'Banjar Masseen', he knew the other three major Bornean ports: Borneo (now Bandar Seri Begawan, Brunei); Passeer (now Tanahgrogot); and Succadana (now Sukadana; see Figure 3), only by information from the Banjars (p. 45-46). Evidently on his first visit to Borneo, and while a novice with the East India Company (p. 32), Beeckman (1718, p. 49, 102) censured previous English behaviour in Banjarmasin. The prelude to negotiating a contract on 17 July 1714 (p. 65) to buy 4000-5000 peculs of pepper (1 pecul = 60 kg) probably too preoccupied him with diplomacy to buy an orangutan, but the agreed three-month wait for the cargo to be aggregated (p. 67) supplied ample opportunity. The orangutan was therefore probably acquired before 1 November 1714 when the sultan compelled his subjects to meet their obligations and the loaded Eagle-Galley moved to deeper anchorage (p. 100). It might have been bought during the subsequent period of settling their affairs and paying their respects (p. 111-118), before final departure on 21 December (p. 128).
The type localities of [Simia] pygmaeus Linnaeus, 1760 and Simia agrias Schreber, 1799 Lacking 'data which can satisfactorily determine to what subspecies they belong', Rothschild (1905, p. 436) believed the names Pongo pygmaeus and Simia agrias applicable only   Groves et al. (1992, p. 337) treated this as a type locality restriction for Pongo pygmaeus, but as Pontianak (now in West Kalimantan province), situated at the mouth of the Landak River, was not founded until 1770, this river is unlikely to be the type locality of P. pygmaeus. The holotype of Simia agrias Schreber, 1799 is the juvenile female in Schreber's (1799, pl. IIC) plate copied from Camper (1782, pls I-II). It arrived in alcohol in 1770 from Camper's former pupil, (Johannes Paulus) Hoffmann (also spelt Hofman), a physician in Batavia.
After signing a treaty of support for Sambas against Sukadana in return for a trade monopoly, in 1608 the Dutch abandoned their four-year-old trade with Sukadana, vacating their factory there in 1623 (Leyden 1814, p. 25). In 1770 they persuaded 'Abdul Ramman' (= Abdurrahman) of Simpang, son of an Arabian Sayyid Sharif (Hunt 1820, p. 16), 'to settle on the unfrequented shores of the river Pontiana' (Hunt 1820, p. 14), promising to develop it as the trade capital of the kingdom of Sukadana (Hunt 1820, p. 15), sold to them by the Sultan of Bantan (Java) for 30,000 Spanish dollars (Leyden 1814, p. 27). Ramman gathered Bugis, Chinese and Malay settlers to build the town. The Dutch established their factory there in 1786 (Hunt 1820, p. 15) or 1776. In 1786 (Leyden 1814, p. 27, 39-40) or 1787 (Hunt 1820, p. 52) they deposed the chief of Mempawah, entrusting his territory to Ramman. They installed rajas at Landak (now Ngabang) and Sanggau with an export monopoly through Pontianak (Hunt 1820, p. 15). In 1786 (Leyden 1814, p. 27, 32, 40) or 1795 (Hunt 1820, p. 26) they besieged, sacked and burnt the ancient city of Sukadana, transferring the inhabitants to Pontianak or dispersing them and their chief into the interior.
If Beeckman (1718) bought his orangutan on 17 July 1714, it would have died about three weeks into his voyage from Banyuwangi to Cape Town; if on 1 November, it would have died en route from Cape Town to St. Helena. Whatever the date of purchase, it did not die in China, thus discounting the tempting conjecture that Lockyer later acquired it. Beeckman's (1718) account is relevant because, if Lockyer obtained the holotype before 1711, it reduces his ready means of acquiring it by another five years. Lacking contrary evidence, it therefore seems safest to infer that Lockyer obtained the holotype in Guangzhou between about 15 August and 17 December 1704. Its most probable source would have been the East India Company's other trading station at Banjarmasin. Leyden (1814, p. 28) blamed Dutch antipathy towards the bustling trading port of Sukadana on irritation at it 'being the principal haunt of the English and French traders on the island of Borneo'. Sukadana is thus perhaps second favourite as type locality of P. pygmaeus. A third possibility is Mempawah, 'greatly resorted to by traders, especially English and Portuguese' (Leyden 1814, p. 30). However, only in the case of Banjarmasin is there firm evidence of British trade at the appropriate date. This consideration might be irrelevant if Lockyer bought the orangutan from a Chinese trader.
Delighted that Beeckman (1718, p. 67) trusted them sufficiently to venture further upriver than had his English predecessors, the Banjars welcomed his request to pay his respects to Sultan Pannomboang [p. 46; spelt Panambahan by Leyden (1814, p. 24)] at Caytongee (now Kayu Tangi). They revisited on 21 August, this time to meet the sultan of Negarree (now presumably Negara, less than 100 km further north, although the sultan had allegedly travelled 200 miles to see them). On 22 August (p. 93) they visited Pangeran Purba (p. 136), prince of Negarree, at his home in Martapura (p. 93). Beeckman (1718) saw little more of the interior, but on 3 September the gunner from each ship assisted on an expedition to suppress rebels at Negarree (p. 80, 96-97).
If an Englishman at Banjarmasin personally collected the holotype of Simia pygmaeus before 1705, the reluctance to stray restricts its origin to near the estuary. If obtained from an inhabitant, as is more likely, it might have come from as far as Negara or farther. More than a century later, based on reconnaissance by Müller in 1836 while collecting natural history specimens, Schlegel and Müller (1841, p. 12) advised that seeking orangutans near Banjarmasin was generally futile and they were now sporadic farther north on the Doeson or Dusun River, the name Europeans then used for the Barito (Müller 1857, p. 152). Schlegel and Müller (1841) reported that orangutans were less rare some days' journey farther west, especially along the Rivers Kahajan (= Kahayan), Sampiet (= Sampit) and Kotaringin (= Kotawaringin, see Müller 1857, p. 141). The Dayaks of Tanjung-Jawa (1°36'S, 114°50ʹE) told Müller (1857, p. 200) that the otherwise absent orangutan, which they called Keoe, chiefly visited the forest on the opposite or western side of the Barito between their village and Kampong Mampon at the start of the eastern monsoon, i.e. from April to May when its preferred locally abundant figs are fruiting. However, on 17 September 1836, the same Dayaks told Müller (1857, p. 236) that an orangutan had called from the other bank two days earlier. He saw no orangutans, but found nests during his hours of intensive search there next morning in a partly swampy, primary forest habitat. The Ngaju Dayaks called the orangutan Kahieo; distinguishing males (Salamping) from females (Buku). Only the Muslims of Banjarmasin and other coastal towns with foreign residents used the Malay term Orangutan (Müller 1857, p. 201). Dayaks at Longwai confirmed to Bock (1882, p. 28) orangutan frequency in the Dusun district and also their presence on the Teweh River, an eastern tributary of the Barito.
The type specimens and type localities of Pongo abelii Lesson, 1827 Khajuria 1953, p. 138). The right (Pearson 1841;Blyth 1863) hand and foot were preserved in alcohol at the Linnean Society of London (Wilson 1831, p. 25-26, pls 33-34;Blyth 1847, p. 728). If still extant, they were probably sold in 1863 when most of its non-Linnean collection was disbanded (Wheeler 1995). Their subsequent fate is unknown. A cheek-flanged adult male preserved in arrack, sent by Willem Adriaan Palm to the Batavian Society at Batavia for forwarding to the cabinet of the Prince of Orange (William V, the last stadtholder of the Dutch Republic), formed the sole basis for Pongo wurmbii Tiedemann, 1808 (p. 329). Dobson (1953, p. 8), Röhrer-Ertl (1988, p. 9), Groves et al. (1992, p. 337) and Groves (2001, p. 300 Geoffroy and Cuvier (1798, p. 187) repeated this information in the past tense. Geoffroy (1798, p. 343) confirmed that 'Ce squelette du singe de Vurmb [sic], l'un des plus rares & des plus précieux de la collection nationale, va nous fournir des observations assez curieuses' [One of the rarest and most valuable in the national collection, the skeleton of Wurmb's ape allows some intriguing observations]. In late 1795 it was taken to Paris, where it still exists (Barsanti 1989, p. 70, 85). As Paris then led taxonomic research in zoology (Groves 2001, p. 41), the early presence there of this skeleton raises the odds that some names, previously considered mere junior synonyms of Pongo pygmaeus, may also be objective senior synonyms of P. wurmbii. We have detected two such names: Simia pongo Cuvier, 1797 (p. 99) and Pongo borneo Lacépède, 1799.
Simia pongo Cuvier, 1797 is a primary homonym of Simia satyrus pongo Kerr, 1792 whose description is largely a verbatim translation of de Buffon (1766, p. 71) and thus, according to Stiles and Orleman (1927, p. 23), a synonym of Homo sapiens Linnaeus, 1758. Even if partly based on the orangutan, it is not an objective senior synonym of S. pongo Cuvier, 1797. Stiles and Orleman (1927, p. 25) conjectured that as Lacépède (1799) gave a facial angle of 30°for Pongo when promoting it to generic rank, but 65°for Simia, the latter probably denotes the chimpanzee. Lacépède (1799), however, almost certainly employed the anthropoid classification of Geoffroy andCuvier (1795, 1798) and Geoffroy (1798) who, from the facial angle of the skull, inferred that Van Wurmb's (1780) ape, together with mandrills and howler monkeys, is the most brutish of anthropoids (Geoffroy 1798, p. 343), while (immature) orangutans, chimpanzees, gibbons, capuchins and marmosets are the most refined (Geoffroy & Cuvier 1795, p. 461;1798, p. 190). 'Pongo' acquired a derogatory connotation. Hence the general surprise (judging by the wide coverage of the event) on 9 February 1818 when Georges Cuvier announced that an orangutan skull sent from Kolkata by the botanist (Nathaniel) Wallich (1785-1854) bridged the age gap between the 'Pongo' and orangutans, implying they are merely age variants of one species (Cuvier 1818, p. 210;de Blainville 1818). Wallich's skull became the holotype of S[imia] wallichii Gervais, 1838 (p. 379). Cuvier (1818, in de Blainville, 1818 gave no type locality, but Cécile Callou of the Muséum National d'Histoire Naturelle, Paris, kindly photographed the skull ( Figure 4A) in four views which show it to be Bornean (CPG pers. obs.). It is a young juvenile with deciduous dentition, first permanent molars and erupting second ones. Malnutrition indicated by the fragile incisal and canine alveolus, and diastema before the erupting molars, suggests it was of captive origin.
Pongo borneo and P. abelii have the best documented type localities of the speciesgroup names available for the orangutan other than the 10 coined by Selenka (1896). Even Selenka's (1896) 10 may be unreliable, as he shunned the interior, relying on local hunters for most of his specimens (Elliot 1913, p. 183-184). The type localities of other names can only be restricted with a good knowledge of the geographic distribution of the orangutan and insight into where orangutans would most likely have been collected at the appropriate date, or at least a reduction in the options.

The geographic distribution of orangutans in Borneo and Sumatra
Extensive orangutan collections have been made in the Kapuas Valley, West Kalimantan, Indonesia (Selenka 1896;Jentink 1897;Hrdlička 1907;Lyon 1907;Elliot 1913, p. 183-186;Röhrer-Ertl 1984, tables 1-5). Brooke (1841, p. 56) reported that orangutans occur at Pontianak and Sambas 'in considerable numbers, and at Sadung' (the Sadong River), 'but are unknown in the intermediate country which includes the rivers of Sarawak and Samarahan . . . which abound with fruit, and have forests similar and contiguous to the Sadung, Linga and other rivers. The distance from Samarahan to Sadung does not exceed twenty-five miles' (40 km). 'From Sadung, proceeding to the northward and eastward, they are found for about 100 miles, but beyond that distance do not inhabit the forests'. His journal specified Rejang and Sarebas as orangutan localities (Brooke 1848a, p. 228). If reliable, Brooke's (1848a, p. 195) assertion that the 'gigantic ape' is occasionally encountered between Santubong and Tanjong Po, i.e. at Bako (1°40ʹN 110°2 6ʹE) where it no longer occurs, suggests its absence near Kuching may be human mediated. Beccari (1904, p. 200) heard of an orangutan killed at Singhi (= Singai, 1°30ʹN 110°10ʹE) and of the constant presence of orangutans on Gunong Bungo (1°18ʹN 110°1 0ʹE). Brooke (1848a, p. 369) noted a difference in appearance and language between the peoples of the Sarawak River and the adjacent Samarahan. Tradition derived the former from Java, while the latter were originally 'Pegùans' (from Burma).
Alfred Russel Wallace (1823Wallace ( -1913 suspected no European had seen more free-living orangutans than he did as Brooke's guest in Sarawak. Shooting 16 from April to August 1855, he accumulated about 21 specimens (e.g. Figure 4B), most without precise localities, but probably from at or near the specified ones of Semabang (1°14ʹN, 110°5 3ʹE), Menyille (up another tributary of the Simunjan River) and Simunjan (1°23ʹN, 110°4 5ʹE) (Wallace 1869;Cranbrook et al. 2005). In October 1878 in the same vicinity, William Temple Hornaday (1885, p. 405) and his hunters killed 43 orangutans. These figures pale against the 300 orangutan skulls Selenka (1896) acquired in the region of northwest Kalimantan east of Pontianak, many of them immature (Jentink 1897, p. 28;Elliot 1913, p. 183). Both Wallace (1869) and Hornaday (1885) fancied they were collecting two species. In March-May 1867, Beccari (1904, p. 194-200) collected 24 specimens at Marup (1°08ʹN 111°39ʹE). None of his females had cheek flanges, but natives and a missionary assured him others did. He speculated that the two orangutan morphs, more distinct than a Bactrian camel is from a dromedary, are species merged naturally by hybridisation, the flanged one from a region more fertile than the other. Charles Hose (1893) had ample opportunity and motive to collect orangutans in northern Sarawak, but recorded only those collected by G. D. Haviland in 'Sarawak' (= Kuching, but probably collected elsewhere); by C. S. Pearse on the Sadong River; and by J. Revilliod at Simunjan. The annual report of the Sarawak Forest Department for 1959 (in Medway 1965, p. 88) stated, Until recently there were considerable numbers in the upper Sadong, Undup, and the border range as far east as the Balui, and also in the Binatang District. It appears probable now that there are barely forty left in the border hills, and only scattered pairs elsewhere. The only remaining concentration . . . appears to be in the swamps between the Sadong and Batang Lupar (even here it is doubtful whether there are more than 200-300 left), and south of the Serian-Simmanggang Road in the Balai Ringin and Sabal Forest Reserve.
The estimated 350 orangutans in 1961 between the Sadong and Lupar rivers had declined by 1988 to only 20-100 in three separate, doubtfully viable populations, but at that time over 500 probably still occurred in the Lanjak-Entimau Wildlife Sanctuary and the contiguous Batang Ai NP (Bennett 1991).
Despite a Pleistocene subfossil presence at Niah (Harrisson 1959;Hooijer 1960;Medway 1965), Banks (1931) confirmed that Sarawak orangutans now occur only between the Sadong and Rejang rivers. Noting that 'the early Sea Dayaks' share this distribution, Banks (1931) inferred that orangutans colonised Sarawak from Indonesia. A single large male skull was reputedly collected in the Sungai Silat region (c. 2°56ʹN, 114°55ʹE) two generations ago and it is difficult to see how it got there for Mias are unknown and always have been in the Baram district. Some Ulu Baram Punans, the nomad hunters, also had a story of how they once saw a big bundle of sticks in a tree from which a Mias emerged; the Punans ran helter-skelter dropping 'parang' and blow-pipe quivers in their terror, a fact which indicates the animals rareness to even the keenest hunters. (Banks 1931, p. 117) Although 'several were seen in July 1945' in the Sembakung (River) area of 'north-east Dutch Borneo', Harrisson (1949) agreed that there are no orangutans in northern-Sarawak or north-western Dutch Borneo, but numbers further south. Mr. Ian Urquhart reports from Binatang on the lower Rejang River, Sarawak, that the ape is said to have been present in the immediate vicinity until fairly recent times. Here again, the increase and expansion of the Sea Dayaks has driven them further back from the navigable rivers.
Urquhart (in Harrisson 1949) added: 'The whole area was completely uninhabited, and it is the Chinese from 1870 onwards who started developing from the Rejang River towards the Second Division boundary and the Dayaks from the Second Division who came into the Julau River' after 1861, 'and from there into Binatang District who drove these animals into the only remaining areas of virgin jungle'. Harrisson (1949) feared orangutans were now 'confined to the headwaters of the smaller streams, Darau, Narasit and Ilang'.
Although in 1993 still retaining about 80% of land under forest cover (Booth 1993), Brunei is evidently outside the Holocene range of the orangutan. St. John's (1863, p. 240) implausible tale of a Murut hunter being abducted by a female orangutan is probably a reliable indicator of orangutan presence on the Padas River, Sabah, Malaysia. An isolated population occurs in the Crocker Range NP (Bennett 1991). Mt Kinabalu, where the species has apparently always been scarce, seems its western limit farther north. On 9 February 1888, Whitehead (1893, p. 169) related how 'Tungal came back to camp [at 2240 m] in a great fright one dayhe had seen a devil looking at him out of a thick tree in the forest; for some time he sat quiet, but at last began to tell us about this "hantu" ( . . . doubtless a large Orang-Utan or Mias). He had met with it close to the camp, but when I went in pursuit the devil had disappeared'. Dusun call Mias 'Kāgeau'. Like Whitehead (1893), Griswold (in Allen & Coolidge 1940, p. 149) in 1937 saw none. Labuan saw one at 6000 ft (= 1800 m), just below Griswold's camp.

Pre-twentieth-century Borneo
Three kingdoms ruled Borneo from early times: Brunei, Sukadana and Banjarmasin (Hunt 1820, p. 7). The first Europeans, survivors of Ferdinand Magellan's exploration, visited in 1521. The 25,000 dwellings of Brunei, the magnificent court and luxuriant, fertile country so impressed Antonio Pigafetta (Magellan's chronicler and guest on the voyage) that, presuming it also belonged to the Raja, he mistakenly extended the city name (mispronounced Borneo) to the whole island, locally known as Kalimantan from an indigenous sour fruit (Hunt 1820, p. 2).
In return for the cession of north Borneo, the Sultan of Sulu assisted the Raja of Brunei in suppressing a Chinese and Murut insurrection. The Spanish subsequently conquered the Sulu Archipelago, imprisoning the Sultan and his court. On capturing Manila (during the Seven Years' War), the British offered to reinstate him in exchange for the transfer to the East India Company of north Borneo, south Palawan and the intervening islands: terms the Sultan accepted in 1763 (Hunt 1820, p. 10-11). In 1775, Sulus expelled the two-year-old British colony from the previously uninhabited island of Balambangan (off north Sabah), killing the garrison (Hunt 1820, p. 44).
As European trade grew with China, fixed prices undercut Chinese junks. Europeans forced Bornean ports to divert their produce to Jakarta and Melaka. The additional transport costs stifled trade in 'gruff goods' (cassia, ebony, pepper, rattan, sago and wax), creating unemployment (Hunt 1820, p. 9). Bornean ports consequently degenerated, their inhabitants resorting to piracy. By 1809 Brunei was a glimmer of its former glory, with fewer than 3000 homes and fewer than 6000 Chinese settlers (Hunt 1820, p. 8). Acts of piracy had for years deterred Chinese junks from visiting. The Portuguese at Macau (China) occasionally attempted to renew trade but their agent withdrew in 1808 after pirates attacked a large ship in 1807, killing its crew (Hunt 1820, p. 58).
James Brooke (1803Brooke ( -1868 first visited Borneo on 1 August 1839 (Brooke 1848a, p. 14), staying less than two months in Sarawak, as a Dayak rebellion made the interior inaccessible (Mundy 1848a, p. 28). In Singapore on 31 March 1843, after almost two years of uninterrupted stay in Sarawak (Brooke 1848a, p. 334), Brooke (1848a, p. 338) wrote, 'I have received long arrears of correspondence from England'. Although the Sultan of Brunei had confirmed Brooke as Rajah of Sarawak on 1 August 1842 (Mundy 1848a, p. 322), commerce between Singapore and Sarawak evidently remained minimal. The Sarawak coast was virtually 'blockaded by the fleets of Sulu Pirates' (Brooke 1848a, p. 196). Commencing an expedition against the Sarebas Dayaks on 10 June 1843, Keppel (in Mundy 1848a, p. 349) described them as 'a race of savage pirates, whose country no force had ever yet dared to invade, and who for more than a century had been inflicting, with impunity, every sort of cruelty on all whom they encountered'.
On 1  Osman or Usman died in the jungle of grape-shot wounds inflicted during the British destruction of his stronghold on 19 August 1845 (Mundy 1848b, p. 202, 205;St. John 1863, p. 192).
Concluding in 1848 that antimony deposits at Bintulu were too far inland to be extracted economically, Robert Burns, a huckster known for his quick temper, continued trading along the north-west coast of Borneo. In 1851, alert to the risk of pirate attack, he resolved to explore the trade options in camphor and edible bird-nests on the Kinabatangan River. Captain Robertson grounded their schooner near Marudu Bay. Their acquaintance, the Marudu chief, Sharif Hussein mediated in the ensuing row. On 11 September, Memadam, an Iranun, and Ibrahim, a Sulu, boarded the schooner, purportedly to display their wares. Hussein and several others later joined them. Ibrahim drew a kris from a rolled mat passed to Burns and beheaded him. Speared, Robertson fell into the sea. Hussein sustained minor injuries; several of the native crew were killed; the rest decamped. Memadam persuaded them to rejoin the ship, sailing for Tungku, an infamous pirates' nest east of Lahad Datu. Nine days later Memadam visited the chief, Sharif Yassin, at Bengaya in Labuk Bay. On learning the vessel's ownership, Yassin decided to apprehend the pirates and report the murders to Labuan. Memadam escaped into the jungle but two of his crew were killed. The acting Consul-General, St. John, arrived to recover the schooner. In 1852, Captain Massie with three steamers was sent to apprehend the culprits. Finding Tungku deserted, he razed it; the stronghold there being finally eradicated in 1879. Lack of evidence exonerated Hussein (King and Talib 1995) but, being Usman's son (St. John 1863, p. 215), he may well have engineered the crime.
Hunt (1820, p. 64) reported that although Sandakan was technically a British port, the Sulus distrusted and would attack any vessel entering it. St. John (1863, p. 244) recorded that the Bornean elephant had not yet been seen by Europeans. When ascending the river Baram, in the north-west coast, one of the guides I had with me said he had frequently traded in the country where elephants abounded, and that was in the interior of the Kina Batañgan river, on the northeast coast. When we went round to look for that district we failed to find the entrance of the river.
The Müller mountain range in central Kalimantan is named after a major of the Royal Dutch Indian Army, said to have been its first European traverser. In reality he probably never set foot on it. Arriving at Samarinda in 1825 with 24 Javan soldiers, George Müller negotiated with Subahdar Abdullah an annual fee from the Dutch government for rights to the duty revenue of the Sultan of Kutai. The Sultan's family and other dignitaries lambasted this agreement, convinced that for permission to sell Kutai to the Dutch, Müller had bribed Abdullah. Müller announced that his expedition would proceed overland to Pontianak and Sambas. He had intended to travel to Berau (now Tanjungredeb) to make Dutch government contact with the Sultan there, but to increase the opportunity for an assault, Abdullah misrepresented Berau as impassable owing to Dayak unrest (Dalton 1831b).
Abdullah recommended Sa Jarring (Dalton 1831a, 1831c) as a guide. This notoriously dissolute and violent young man was widely acknowledged as Müller's killer, either while Müller was asleep or with a parang while walking behind him at the rear of the party. The panicked soldiers were speared while trying to flee across the river. Dalton (1831b) learnt that this occurred near Muarakaman (at least 300 km east of the Müller Mountains) after six days' travel. Three days later, culprits returned to Tenggarong with spoils, including European arms, navigational instruments and a drum. The excellent, rust-free condition of the London-made chronometer belied the Sultan's claim that it was recovered after days in the river. Sa Jarring owned Müller's French shotgun and had melted upcountry at the news of Dalton and his Danish companion, Mr Hecksher arriving in Kutai. Dalton (1831b) believed Abdullah, the Sultan and others premeditated the murder, foisting the blame on Dayaks within 160 km of Pontianak to avert punitive action. One of the death squad, Seredin, a freedom-seeking Javan slave the Sultan bought from pirates, named Abdullah as the ringleader. Dalton (1830) further reported that off Pulau Laut in 1819 (Buckley 1902, p. 208), Raga, a brother of the Rajah of Pagatan, waylaid the English Captain Gravessome, apparently sailing under a Dutch flag after successfully trading at Sambas and Pontianak. Enthusing about trading prospects at Tenggarong, Raga accompanied him to Samarinda where, in collusion with the Sultan of Kutai, he killed him and most of his crew, sharing the booty (Dalton 1829). Dalton (1831a) spent 11 months in Kutai, eight of them on starvation rations after the Sultan stole all his property including his jacket. Dalton escaped on 2 October 1828 only by pretending that on returning to Singapore he would advance the avaricious Sultan cash and goods for produce. Even then the crew of the proa planned to murder Dalton and Hecksher for what little they retained. They survived by sleeping in turns and because the increasingly waterlogged proa distracted the crew. On beaching on 11 October at Mamuju, Sulawesi, they escaped, the Bugis village chief supplying hospitality and protection (Dalton 1831c, d).

The holotype of Simia morio Owen, 1837
Dr. W. Montgomerie of Singapore sent the holotype of Simia morio Owen, 1837 with another Bornean orangutan skull, and the skin and skull of a Sumatran juvenile that had died in his possession soon after erupting its second permanent molars. All three were exhibited at a meeting of the Zoological Society on 25 October 1836 (Owen 1837(Owen , 1838. Although in 1855 the British Museum purchased the mammal catalogues of the Zoological Society museum and much of its collection (Wheeler 1997), the fate of the holotype is unknown. Richard Owen was Hunterian professor at the Royal College of Surgeons, London. Of the orangutans in catalogues of its museum by Owen (1853, p. 762) and Flower and Garson (1884, p. 10), only a female 'Mias Rambi' skull (O.C.5056) donated by Owen might be the holotype. It probably is not, as Owen (1853) classified it, 'Pithecus Satyrus', while still recognising 'Pithecus Morio' as a distinct species ). Blyth's (1836) letter to G.R. Waterhouse, the newly appointed curator of the Zoological Society museum (Wheeler 1997, p. 91), concerning 'the age of the Ourang whose skin I forwarded', corroborates Blyth's (1853, p. 369) claim to have drawn Owen's attention to the specimens. Blyth (1836) wrote, 'the lady to whom it belongs . . . favoured me with the enclosed extract from her brother's letter' (now lost), adding that her brother 'had the animal for some years in his possession'. This indication that Montgomerie's sister kept the specimens after their exhibition could explain their disappearance.
Dr. J. William Montgomerie (1797-1856) arrived at Singapore in its founding year, 1819, as Assistant Surgeon with the Bengal Native Infantry (Buckley 1902, p. 60), transferring in 1827 to a post in India (Bastin 1981, p. 51). He returned in 1832 as Senior Surgeon when Singapore became capital of the Straits Settlements (Lee 1973, p. 18). In 1843, he alerted the Bengal Medical Board to the surgical application of guttapercha (Buckley 1902, p. 402), a tree sap and forerunner of rubber, used in cable insulation. He apparently never visited Borneo or Sumatra. Bugis traders informed him that gutta-percha grew at Coti (now Tenggarong on the Mahakam River, then known as the Coti or Kutai River), and (James) Brooke told him of its occurrence in Sarawak (Buckley 1902, p. 404).

Strangers from Borneo
In a letter of December 1822, Stamford Raffles mentioned sending home the skeleton of a 5.5-foot Bornean orangutan (Buckley 1902, p. 77). It may be the skeleton (O.C.5050) he donated to the museum of the Royal College of Surgeons ( [Owen] 1853, p. 759), if it was shipped before 1824 when much of the rest of his scientific collection and historical archive was destroyed by fire with the ship Fame shortly after it left Bengkulu, Sumatra (Buckley 1902, p. 10-12). The only other reference to the orangutan in Buckley's (1902, p. 304) history of Singapore from 1819-1867 is his report that in March 1836 'two strangers arrived from Borneo, being a pair of Orang Outans, brought from the interior, which excited a good deal of curiosity'. His sources evidently were two articles in the weekly Singapore Chronicle and Commercial Register. The first, entitled 'Strangers from Borneo', in the 5 March 1836 issue, p. 3, reads: During the week two Orang Outangs male and female, have been imported from the interior of Borneo. They are, in the words of the advertisement, beautiful, although in our optics rather in the ill favored [sic] sense of the term. We do not remember having ever seen a larger or a more hideous looking animal than the female one, which besides possessing a most loathsome physiognomy, has very many satanic propensities about it. In a recumbent position which she assumed to display her unequalled charms, we were much reminded of some, weatherbeaten, withered, old flat nosed hag which, among the poorer and lower description of Bugis and Malays may occasionally be seen crawling about, without however the pilosity of the ape species. To any one proceeding on a long voyage, and exceedingly fond of having something to do, an animal of this description just from the woods, would afford an inexhaustible source of amusement by breaking the almost uninterrupted monotony of a sea passage, in taming its present savage disposition, training it to some manners, and dispose it to acquire some habits of refinement and civility, so that its value on arrival in Europe might be materially enhanced. Under proper tuition such an animal would be accounted cheap at a couple of hundred pounds sterling. The male is still young and gentle, though somewhat sneakishly [sic] shy, but this by a moderate supply of modest assurance, which could well be spared to him by his Grand-dame, he would in all probability prove an animal of no small value and sagacity.
The advertisement on p. 1 of the same issue reads: 'FOR SALE. One small, and one very large and beautiful female Orang Outang, just imported from Borneo. Apply at the Chronicle Office. Singapore, 5th March, 1836'. This advert was repeated verbatim in the issues for 12, 19, 26 March and 2 April 1836.
The second article, this one entitled 'The Borneo Strangers', in the 12 March 1836 issue, p. 3, reads: The Orang Outangs of which we made mention last week have attracted considerable attention, for, we understand, the greater portion of the ladies and gentlemen of the settlement have been to visit them. The female Orang Outang, we have been told, impatient probably of confinement and by gnawing the stout bottom bars of her enclosure and afterwards cunningly contriving to upset the same from its perpendicular, effected a temporary escape. Perched upon some crates of earthenware next morning, contemplating possibly the felicities of freedom or the misery of the late invasion of it, she was discovered in that situation by some servants, who without bidding good morning to the self liberated dame, fled with the utmost precipitation from her presence and ran to awake their dormant master who learning the cause of alarm, collected a posse and after a great deal of trouble succeeded at length in securing the dame, and reconducted her to her abode to which she returned evidently with all the symptoms of chagrin and disappointment clearly impressed upon her physiognomy. We had hoped as there were two vessels lately for Europe when these animals arrived, that they would have been purchased at once on speculation, and, we think, had they lived home that the speculation would not have been unremunerative, but as the hazard of a long voyage might have deterred all from such a purchase, it may be an equally lucrative adventure, if not more so, to consign them to Bengal where any of the rich natives establishing, or in possession of, a menagerie, would consider it a fit opportunity of being lavish of his money to call himself master of a female of the magnitude, strength and beauty of the animal who is yet, at this settlement, but a comparative stranger, having been only recently imported from its native woods in the interior of Borneo. Evidently in 1836 the capture and successful sea transport from Borneo to Singapore of a live adult female orangutan were rare, if not unique, occurrences. Even today (if permissible) they would constitute quite a feat. Other than their simultaneous presence in Singapore, we have established no closer link between Montgomerie and this orangutan. The coincidence of events might seem too extraordinary for the adult not to be the holotype of Simia morio but, astonishingly, the Bornean strangers probably had a different fate.
In 1836, R.W.G. Frith donated to the museum of the Asiatic Society of Bengal an adult female skeleton (4A of Blyth 1863, p. 4; 3ww of Anderson 1881, p. 14) of 'the Sumatran Orang-otang which lately died in Calcutta' ( [Prinsep] 1837, p. 833). Khajuria (1953, p. 139) reported that only its skull (ZSI.11713) then remained at the Indian Museum, Kolkata. The skin, including the hand and foot bones, was discarded before Frith secured the rest, rescuing a few 'digital bones'. Blyth (1853, p. 371-372, pls 7-8) clarified that, probably fully adult when caught, it was actually one of a pair purchased in Singapore by the Society's joint-Secretary, (Arthur) Grote (1814-1886). As Grote arrived in Kolkata in 1833 (Prior 2004), his purchase opportunity was short. Grote told Blyth (1853, p. 371, footnote): 'They were not from Sumatra, but from Borneo. At least I am pretty sure that my memory does not deceive me on this point'.
That Montgomerie's juvenile orangutan derived from Sumatra (Owen 1837) could be attributed to a failure of memory or communication, but that two adult female orangutans were brought alive from Borneo in about 1836 is implausible. Grote therefore probably purchased the Bornean strangers. Montgomerie evidently sourced his orangutans elsewhere. He did acquire a live male orangutan Bugis traders brought from Pontianak to Singapore in July 1826. In July 1827 he gave it, still with no permanent teeth, to George Swinton in Kolkata. By then Montgomerie had seen 'at least ten or twelve' evidently live Bornean orangutans, the largest being 'about three feet in length from the crown to the heel', including a female he owned for 12 months which died before its first permanent molars had penetrated the gum (Grant 1828, p. 1-2, 12, 15). Montgomerie (in Grant 1828, p. 16-17) also described an adult (probably male, judging by his description of the canines) Bornean orangutan skull, collected for Raffles on his last visit to Singapore (in 1823), but believed lost in the destruction of the Fame. Its collector, 'a country-born Frenchman, (a native I believe of Chandernagore,)' was undoubtedly Pierre-Médard Diard (1794-1863, see Brandon-Jones 2004, p. 1567, who is known to have collected near Pontianak in about 1826 (Müller and Schlegel 1841, p. 71), sending the National Museum of Natural History (now Naturalis) at Leiden in the Netherlands four mounted orangutan specimens and two adult male skeletons (Temminck 1841, p. 135).
The holotype of S[imia] hendrikzii Wagner, 1839 is the plaster cast (no. 7197) at the Königliche anatomische Museum in Berlin of an (adult) orangutan skull owned by Professor Hendrikz. Its type locality is unrecorded, but its availability to Müller (1836, p. xlvii) establishes that it was collected before 1836.
The holotype of Simia straussii Wagner, 1839 is an adult skull with significant dental attrition, brought from Borneo by Dr. Strauss and sent by Professor Rudolph Wagner for Wagner (1839, cols 417-419) to examine.
The historical evidence indicates that, like the holotype of Simia morio Owen, 1837, these three holotypes probably came from the Bornean south or west coast, as other coasts were then dangerous destinations. If operational at all, Mempawah and Sukadana no longer functioned as major ports, but orangutans could have originated from Banjarmasin or Sambas. By then the Dutch had probably monopolised trade with Banjarmasin via Jakarta so, even if Pontianak did not export the holotypes, the most likely alternative is the not-too-distant Sambas.

James Brooke's orangutan collection of 1840
James Brooke (1841) sent the Zoological Society five live orangutans: one from Sadong (Sarawak), three from Pontianak and one from Sambas, with instructions to Captain Swan that they were to be preserved in alcohol if they died on his ship, the Martin Luther. Brooke (1841) did not recount how he obtained his Pontianak and Sambas orangutans, but he knew the Kapuas River as the Pontiana or Kappawas and reported second-hand that a half-day or daylong mountain crossing from the headwaters of the Sadong or Sarawak rivers brought you within two days' river journey of Sanggau (Brooke 1848a, p. 255). The Sambas adult female may have been obtained from Chinese of the San Ti Qu kunsi (company) from Sambas who mined gold and antimony on the Sarawak River under licence from the Sultan of Brunei (Brooke 1848a, p. 289). However, Brooke (1848a, p. 304) reported that turtle eggs from the islands of Talang Talang were 'exported to Sambas and Pontiana, and all along the S.W. coast' and that a fish spawning in April attracted people from far afield, including Sambas (Brooke 1848a, p. 305). There was evidently ample intercourse between these areas.
Twenty-five orangutan skulls and 14 skeletons donated by Brooke (1842) were displayed at the Zoological Society's meeting of 11 January 1842. Most of this 'fine collection . . . from the north-west coast of Borneo, either shot by myself or brought by the natives' (Brooke 1841, p. 55) is now preserved at the Natural History Museum, London. The accession numbers allocated to the series, ZD.1855.12.26.2-21, were never applied to the specimens. They are now registered as ZD.1986.1092, ZD.1986.1095-1109, ZD.1986.1112-1114and ZD.1986.1118-1120(Jenkins 1990. Most skulls of this series are numbered from 1 to 25 in the mammal catalogues of the Zoological Society museum. Evidently this was Brooke's (1841, p. 59) sequence with which he specified the five orangutans he shot, and the two that 'died aboard', presumably en route from Sarawak to Singapore where Brooke (1841, p. 55) wrote his letter. Brooke (1848a, p. 241) left for Singapore on 14 February 1841. With the 18 specimens procured from natives (Brooke 1848a, p. 227), some of them at Sangi (= Kampong Ensengei Melayu, 1°25ʹN, 110°38ʹE) on 19 September 1840 (Brooke 1848a, p. 229), the total of 25 seems accurate. Most specimens retain their Zoological Society numbers in ink on the skull or skeleton. These accession numbers diverge from Brooke's sequence. Brooke apparently also instigated an uncorrelated numerical sequence from at least 29 to 43 expressly for skeletons. Two specimens without Brooke skull numbers, ZSL.1098 (ZD.1986.1100, probably the Sambas adult female) and ZSL.1131(ZD.1986.1114) may be two of the five sent home alive. Brooke (1848a, p. 224) shot his second adult female orangutan (probably ZSL.1096, ZD.1986.1119) near Kundah hill (= Bukit Punda, 1°11ʹN 110°51ʹE) by the Sadong River on 16 September 1840, less than three weeks into his second period in Sarawak after a six-month visit to Sulawesi (Brooke 1848a, p. 172, 176). Brooke (1841, p. 57) claimed he shot his first specimen, an adult male (ZSL.1087, specimen untraced) some days before, 'about thirty miles distant', but his journal records that he actually shot it very near the other ones, apparently (unless there is a hiatus in the journal extracts) on the previous day (Brooke 1848a, p. 218). The infant taken alive from the first adult female (probably ZSL.1089, ZD.1986.1102, shot on the same day as the male (Brooke 1848a, p. 222), is probably ZSL.1090 (ZD.1986(ZD. .1106, which 'died aboard'. Brooke (1841, p. 55) believed that, based on the presence or absence of cheek flanges and the development of the cranial sagittal ridge, his material and native reports could 'beyond a doubt, prove the existence of two, if not three distinct species in Borneo'. Brooke (1841, p. 56) recorded cheek flanges of 'nearly two inches in thickness' in the first male (seemingly qualifying it as a Mias Pappan) whilst he incongruously asserted its 'wide distinction from the pappan, or, as the Dyaks call it, chappin' (Brooke 1848a, p. 222) and that he had 'not seen the largest mias pappan' (p. 225).
Datu Maraja, of Simunjang, had often seen the female pappan in the woods about his residence, but the rembi was not found there. The females had the cheek callosities as well as the males. At Sangi a Dyak came to inform me he had killed a large mias, and wanted to know if I would buy it. I inquired what sort it was: he answered by putting his hands on each side of his face to intimate it was a pappan. I then asked was it a male or female. He replied, a female. How did he know? Because she had a young one in her arms. I promised to buy the head, but unfortunately he never returned. I may add, that in some places the rembi alone is found; at Linga the pappan is unknown. (Brooke 1848a, p. 227) The type specimens and type localities of Pithecus brookei Blyth, 1853, P. owenii Blyth, 1853 andP. curtus Blyth, 1856b Concurring that there are at least three distinct species of orangutan, Blyth (1853, p. 375, footnote) proposed that the Mias Rambi should be named Pithecus brookei in honour of Brooke (1841), its first discriminator. One syntype is the skull of a Bornean adult male (Blyth 1853, p. 369, 383, pls 3-4;3D of Blyth 1863, p. 3;3nn of Anderson 1881, p. 13), donated to the museum of the Asiatic Society of Bengal in July 1838 by Major (William Michael) Gregory (1789-1842), and now apparently consisting only of a mandibular ramus (ZSI.11723) at the Indian Museum, Kolkata (Khajuria 1953, p. 139). The other syntype is the skull of a Sumatran adult male (Blyth 1853, p. 369, footnote, 383, pls 1-2) simultaneously loaned by Gregory to the museum. Both are cranially larger than the holotype of Pongo abelii. As the Society already owned that Sumatran specimen, Gregory retained possession of his (E[vans] 1838), but evidently he did not reclaim it, as it apparently still exists at the Indian Museum (3 C of Blyth 1863, p. 3;4b of Anderson 1881, p. 25;ZSI.11727 of Khajuria 1953, p. 138). Blyth (1853, p. 369) categorised the holotype of Simia morio as a Mias Kassar. He presumed it male as its alveolus is deeper, its maxillary premolar and molar row longer, and its zygomatic arch sturdier (Blyth 1853, p. 371) than in adult female ZSI.11713 (donated by Frith; see Strangers from Borneo, above). The canine morphology in Owen's (1838, pls 33-34) figures, however, unequivocally diagnoses the holotype as female. The permanent canine of a female orangutan is spearhead-shaped and almost mesio-distally symmetrical. In the male it is more fang-like, often with a distal spur at the crown-root junction. Ironically, this (sexual) dimorphism was Owen's (1837Owen's ( , p. 95, 1838) chief diagnostic feature of S. morio. Brooke (1841, p. 59) misdiagnosed his adult female no. 9 (ZSL.1099, ZD.1986.1098) as an adult male. Blyth (1853, p. 369, 383) mistakenly regarded the Bornean syntype of Pithecus brookei as female. Confusion between sexual dimorphism and speciation in the orangutan was thus inevitable.
The type locality of Pithecus brookei Blyth, 1853 depends on a lectotype designation. Since 2000, a lectotype designation is invalid unless its taxonomic purpose is specified (International Code of Zoological Nomenclature, Article 74.7.3). As the origin of the syntypes remains inexact, a designation here would achieve nothing. If a syntype proved to represent a taxonomically distinct unnamed Bornean population, then its designation as the lectotype would be appropriate. According to Helfer (1838, p. 858), however, Gregory brought both skulls from Sumatra. Should a second Sumatran subspecies be recognised, designation of the Sumatran syntype, depending on its precise locality, might be appropriate.
Being published before 1961, the conditionally proposed P[ithecus] owenii Blyth, 1853 (p. 375, footnote, 381-383, pls 9-10) is an available name (International Code of Zoological Nomenclature, Article 11.5.1). The holotype is the mounted skin and skeleton of a female that had lived for 12 years in captivity with J. Apcar in Kolkata; Apcar donated it in 1846 to the museum of the Asiatic Society of Bengal. Both Blyth (1853) and Anderson (1881, p. 12, specimen 3bb) attributed its abnormal morphology to captive conditions. Blyth (1853, p. 372) designated it as adolescent, specifying that its maxillary and mandibular third molars 'had just pierced the gums'. It was six months old when Apcar obtained it. Blyth (1847, p. 729, footnote) had often seen it alive, seemingly always mild and good-natured. Its age indicates that it was a subadult, probably with dietary deficiency causing retarded dental eruption. Blyth (1863, p. 4, specimens 4D and E) later referred it to Simia morio. It now remains at the Indian Museum, Kolkata, apparently as a skull only (ZSI.11805). Khajuria (1953, p. 142, mistakenly) credited W. Rutledge as its donor. This female, captured before 1835, probably originated from Pontianak or Sambas.
Writing to Charles Darwin for his opinion of Wallace's (1855) paper on speciation, Blyth (1855) famously referred to the author as 'friend Wallace'. Blyth (1856b, p. 526, footnote) cited an unnamed 'friend who has resided long in Borneo, and has examined numerous skulls of Orang-utans (including those which have passed through the hands of Sir J. Brooke)'. Wallace had been in Sarawak since 1 November 1854 (Wallace 1869, p. 46) and is probably this friend who conjectured that, 'P[ithecus] OWENII represents, in the southern part of the great island, the P. MORIO of the northern part' (Blyth 1856b, p. 526, footnote). No evidence of a southern origin for P. owenii was specified, but this brings Banjarmasin into contention as the type locality.
The holotype of Pithecus curtus Blyth, 1856b (p. 520, 525-526) is the skeleton of an adult Bornean orangutan which James Brooke donated in 1855 to the museum of the Asiatic Society of Bengal. Khajuria (1953, p. 138) credited Brooke as donor to the Indian Museum, Kolkata, of only one orangutan skeleton (ZSI.11709). Whether it is the holotype or, if not, whether the holotype still exists there but with mislaid data, should be determinable by the limb brevity that prompted the name P. curtus. Blyth's (1856b, p. 526, footnote) observations that the canines are eroded to incisor height and that especially the mandibular canines of most males and some females exceed its in circumference, discredits Blyth's (1856bBlyth's ( , p. 520, 1863) and Anderson's (1881, p. 14, specimen 3vv) diagnosis of the holotype as male.
The type locality of Pithecus curtus can be provisionally restricted to Sadong, the only more precise locality specified for any of the seven orangutan skeletons Brooke donated in 1855 (Blyth 1856b, p. 522-524;Anderson 1881, p. 14) in two consignments: two skeletons arriving before July (Blyth 1856a); the rest in July-September (Blyth 1856c). From unspecified characteristics (probably their colour or incompleteness), Blyth (1856b, p. 524) inferred that the three Sadong skeletons had been prepared by burial. Their donation in the year the Zoological Society museum disbanded may not be coincidental. Rather than collecting fresh skeletons, Brooke may only have redistributed his existing material. The presence of only seven of his orangutan skeletons at the Natural History Museum, London (Jenkins 1990), substantiates this possibility. ZSL numbers in ink on the bones would confirm it. The only two of the series (ZD.1986.1100and ZD.1986.1114 which are possibly from Pontianak or Sambas (see Brooke's 1840 orangutan collection, above) are at the Natural History Museum, so the type locality is fairly secure. The details of the seven skeletons do not match records of Wallace's collection (Wallace 1869;Cranbrook et al. 2005).
The holotype and type locality of Pan vetus Miller, 1915Gerrit Smith Miller, Jr. (1869-1956 instigated the name Pan vetus Miller, 1915 for the Piltdown mandible, believing it a fossil chimpanzee contemporaneous with the more human cranial fragments allegedly found with it. The holotype (also that of Boreopithecus dawsoni Friederichs 1932) is an orangutan right mandibular ramus (accession number E.594) with first and second molars in situ. Despite protesting to Miller that the scientific name of Piltdown man 'was based solely on the mandible' (Spencer 1990, p. 134), Woodward (1913) associated together the mandible and brain-case fragments 'without any hesitation'. His uncertainty was only that the cranial characters might not be diagnostic (Woodward 1913, p. 135). He specified no holotype. Miller (1915, p. 19) was therefore justified in designating the left temporal bone (E.591) as the lectotype of Eoanthropus dawsoni Woodward, 1913. As first reviser, his designation is definitive (International Code of Zoological Nomenclature, Article 74.1).
De Beer (1955) requested the International Commission on Zoological Nomenclature to reject all scientific names (including Homo piltdownensis Keith 1931) instigated for 'Piltdown Man', claiming that its hypothetical basis excludes such names from zoological nomenclature (International Code of Zoological Nomenclature, Article 1.3.1), but this contention is specious, as the authors treated the finds as authentic relics of real animals, and their type specimens are tangible skeletal parts. The Commission cannot force a zoologist to abandon a conviction, however improbable, that all or part of the assemblage was genuine and the described taxa recognisable. Now that the remains have been re-identified, the names attached to them are available should they prove to represent recognisable taxa, such as a subspecies of orangutan. The remains are preserved at the Natural History Museum, London (Oakley 1971, p. 35). Edgar William Willett (1856-1928, an early-retired anaesthetist, was a friend of Charles Dawson (1864Dawson ( -1916, the solicitor and antiquarian who brought the Piltdown finds to public attention (Spencer 1990, p. 18-19, 23, 33, 87-88, 222). Willett or his father, the wealthy brewer and businessman Henry Willett , probably acquired the Piltdown mandible from Alfred Hart Everett (1848-1898) whose cave explorations in Sarawak they had sponsored (Sherratt 2002, fig. 1). This provenance is corroborated by the mandible not having been boiled (Oakley 1959, eliminating most research specimens), its radiocarbon date of 90 ± 120 years BP (Hedges et al. 1989, p. 210), its undiminished nitrogen content and its finely crackled bone surface (not replicable by Oakley 1959) resembling that of an adult male orangutan skull ZD.1884.10.30.2 Everett collected. Everett (1879, nos. 100 and 111) doubted the cave origin of ZD.1884.10.30.2. It and the Piltdown mandible were probably trophy skulls (perhaps with surface altered by proximity to a fire). The subsequently announced first molar, apparently the left one from the same mandible (Clark 1955, p. 239;Hrdlička in Spencer 1990, p. 162;Oakley in Spencer 1990, p. 210-211;C. Stringer, I. De Groote and C. Dean, e-mail, 16 April 2013), indicates the fraudsters broke a complete mandible. Its small size, virtually unworn molars (before artificial abrasion), incompletely erupted missing third molar, and immature canine (Clark 1955) diagnose the mandible as from a subadult female. Miller (1915, p. 12) discussed the canine, but omitted it from his paradigm of Pan vetus.
Two of the 32 caves Everett (1880, p. 311) examined are in Gunong Subis on the Niah River. As discussed above, orangutans no longer occur there, so the Piltdown mandible probably came from one of the other 30 caves, all in 'Upper Sarawak Proper'. Fourteen are in the Bidi and Paku districts. The caves were assigned numbers but few were named in Everett's (1879Everett's ( , 1880 published reports: Ahup, on the Sambas-Sarawak frontier; Guah Kokan in the Kapoh hill at Bidi; Kawa cave near Bidi; a cave in the Eusunah gorge, near Paku; an old gold cave at Piat; cave No. 14 at the second milestone on the Tagora road; Gunong Jawang; and two on Jambusan hill. It being the only specified locality from which Everett (1879) recovered orangutan material, the type locality of Pan vetus is here provisionally restricted to (Gunong) Paku (1°26ʹN 110°11ʹE), Sarawak, Malaysia. For the approximate location of this site, 6 km along a tramway from Busau (= Buso), see Hornaday (1885, p. 477). Warren et al. (2001) determined and analysed mitochondrial DNA (mtDNA) control region sequences from six Bornean orangutan populations, identifying four regional groups: an eastern one from Kutai NP; a southern one from Gunung Palung NP and Tanjung Puting NP; a western one from Danau Sentarum, Kalimantan, and Semongok Wildlife Rehabilitation Centre, Sarawak; and a northern one from Sepilok Orangutan Sanctuary, Sabah. The western and northern groups are indistinct at the more stringent 1% significance level. Pairwise group sequence divergence ranges from 2.1% within the southern group, to 6.5% between the southern and western groups, rising to 16.5% to 19.2% between Bornean and Sumatran populations. Four orangutans of uncertain origin falling within the Bornean clade but outside the regional groups imply more such groups exist. Consistent western group mtDNA affiliation of hair samples from the nest of a wild Kutai NP individual indicates it was a released ex-captive.

Subspeciation in the orangutan
Multivariate analysis by Groves et al. (1992) cranially pooled north and west Bornean adult female orangutans. Adult males from these regions are more distinct, especially when mandibular dimensions are included. Sumatran and south Bornean adult orangutan skulls are almost completely distinct from north Bornean ones (which had the smallest sample size), but partially overlap the west Bornean ones (with the largest sample size) and one another. Measurements of two adult male skulls confiscated in Samarinda and thought to be from Kutai NP assigned them to the northern group (Groves 2001). Uchida (1998) found south Bornean orangutans dentally resemble Sumatran ones more closely than do west Bornean orangutans, although in her canonical analysis the Bornean populations mutually overlap but not with the Sumatran population.
A spatial analysis of molecular variance (SAMOVA) corroborates the seven main starlike geographic clusters in the phylogenetic network Arora et al. (2010) generated from the mitochondrial haplotypes of 211 orangutans from 10 sites in Borneo. Except for Danum Valley clustering with south Kinabatangan about 90 km away, and Gunung Palung sharing its only haplotype with Sabangau, this indicates almost complete separation of all sampled sites. Their recent mean coalescence date of 176 ka (95% highest posterior density, 72-322 ka) is compatible with post-glacial Bornean orangutan dispersal from a rainforest refugium in north central Borneo, roughly midway between the sampling sites. If Brandon-Jones' (1996) inference is correct that a dry period provisionally dated at 190 ka eradicated all primates from Sumatra, the much earlier coalescence of Bornean and Sumatran haplotypes (mean, 3.6 Ma; 95% highest posterior density, 2.3-5.0 Ma; as calculated by Arora et al. 2010) perhaps would indicate that orangutans recolonised Sumatra from Indochina rather than from Borneo (Brandon-Jones 1998). This alternative might be testable if the late Pleistocene orangutan skeleton from Vietnam (Bacon and Vu 2001) can supply DNA. In all events, if both morphological and molecular evidence correctly indicate that Pongo pygmaeus and P. abelii are separate species, the affiliation of the Indochinese population requires clarification.
Concluding that currently recognised Bornean orangutan subspecies are (contrary to expectations, by definition) not reciprocally monophyletic, Arora et al. (2010) linked the north and south regional groups. Within this aggregation, the southern population west of the Kahayan River is most distinct. The east and west regional groups are interlinked on a separate branch, but otherwise are no more distinct than the trans-Kahayan population. This perhaps indicates that a latitudinal dispersal preceded a longitudinal one and that the most recent spread is across or round the Kahayan River, creating the illusion that the Kapuas River is a taxonomic boundary. Based on this analysis, if Banjarmasin is accepted as the type locality of Pongo pygmaeus, at least three Bornean orangutan subspecies might be recognised: a western one for which the name P. borneo seems the earliest, an eastern one (apparently unnamed, unless one of the Pithecus brookei syntypes came from east Borneo), and the nominotypical northsouth one. The cranial evidence favours splitting the latter. The molecular evidence of Arora et al. (2010) and Nater et al. (2011) seems to preclude referring the trans-Kahayan population to P. abelii. Their evidence strengthens the case (discussed in depth by Steiper [2006]) for treating P. abelii as a separate species. Nater et al. (2011) found Sumatran orangutans form four strong mtDNA geographic clusters, with the southernmost one at Batang Toru (1°40ʹN, 99°05ʹE) genetically closest to the Bornean sampled sites. The Sumatran orangutan Y-chromosomal haplotype network is much tighter, but subspecies may be recognisable.

Discussion
A problem in orangutan nomenclature is that Pontianak is a probable port of export, rather than a collecting locality. At the time the holotypes of Simia morio, Simia crossii, S. hendrikzii, S. straussii and P[ithecus] owenii were collected, Pontianak was also a synonym for the Kapuas River, the main artery for trade with the interior (Brooke 1848a, p. 370). Skull and dental measurements seem insufficiently discriminatory to determine whether the holotypes were collected north or south of the Kapuas.
Even if it is assumed that the holotypes originated from the immediate vicinity of Pontianak, it remains unconfirmed that the Pontianak and Sadong River populations pertain to the same subspecies. If the populations did attain their recent distributions in the above-described manner, rainforest regeneration and orangutan dispersal probably hugged the mountains before descending into the lowlands. Pontianak is about midway between the two highland areas of west Borneo, so orangutan populations could have reached it from either direction. The intervening Kapuas River perhaps favours a northern route but, without unequivocal contrary evidence, Pongo pygmaeus borneo may include the trans-Kahayan population. The National Museum of Natural History, Smithsonian Institution, Washington, DC, has three orangutan skins with skulls from Sungai Sama (Lyon 1907, p. 548, 571), close enough to Pontianak to act as a proxy for this now possibly extinct population. DNA extracted from tissue samples might settle this issue.

Conclusions
We conclude that the holotype of Pongo pygmaeus (Linnaeus 1760) is a Bornean orangutan, almost certainly from the south or west coast, obtained in China by Charles Lockyer, probably at Guangzhou in late 1704. It probably originated from Banjarmasin, though possibly from Sukadana or Mempawah. In about 1740, Lockyer gave it to Hans Sloane who bequeathed it to the British Museum, London. Most of the holotype was destroyed there in about 1850, but its skull may be the one preserved at Uppsala, Sweden.
We consider Pontianak the most probable type locality for Simia morio Owen, 1837, Simia crossii Wagner, 1839, S. hendrikzii Wagner, 1839, S. straussii Wagner, 1839 and P[ithecus] owenii Blyth, 1853. P. brookei Blyth, 1853 has two syntypes, one from Sumatra, the other probably from Borneo. Unless more syntype history comes to light, attempting to settle its type locality with a lectotype designation would currently be pointless and therefore invalid. We provisionally restrict the type locality of P. curtus Blyth, 1856b to the Sadong River, and that of Pan vetus Miller, 1915 to Paku, Sarawak. The type localities of some species-group names available for the Bornean orangutan remain imprecise but, if Banjarmasin is accepted as the type locality of Pongo pygmaeus, then Pongo borneo Lacépède, 1799 is the earliest available name for the Pontianak population.
Regrettably, this reverses the subspecific nomenclature advocated by Groves (2001) but, as we here subsume both P. wurmbii Tiedemann, 1808 and S. morio into P. borneo, the new nomenclature is unlikely to be confused with the old (Figure 5A and B).
In recent times, most institutions housing captive specimens have scrupulously avoided interbreeding Bornean and Sumatran orangutans. If Borneo proves to hold several subspecies, then it is equally important to try to avoid interbreeding them, either in captivity or by reintroduction or translocation into the wrong area. We therefore advocate tentative recognition of the four subspecies outlined above. We baulk at instigating a new scientific name for the eastern subspecies until its credentials are more Figure 5. The nomenclature and geographic distribution of the subspecies of Pongo pygmaeus, (A) following Groves (2001), and (B) as revised in this paper firmly established and the availability of an existing name is more comprehensively excluded. We recommend that the name Pongo pygmaeus borneo is provisionally adopted for the western subspecies, but that an urgent attempt is made to confirm that it is a senior synonym of Pithecus curtus, which otherwise should replace it for that subspecies, releasing the name Pongo borneo for the trans-Kahayan population. If P. borneo is unavailable for the trans-Kahayan population, Simia agrias may be available. If orangutans were already scarce at Banjarmasin by 1770, the holotype may derive from the nearby trans-Kahayan population. Available evidence indicates that Pongo borneo predates Simia agrias Schreber, 1799 by at least a few weeks. If the two names prove to have been published simultaneously and to represent the same subspecies, as first revisers we award priority to Pongo borneo, the one with an unequivocal type locality.

Systematics
Pongo pygmaeus pygmaeus (Linnaeus, 1760) [  (Arora et al. 2010), and formerly farther south to Banjarmasin (the type locality). If this is the most archaic Bornean subspecies, it is probably the closest living relative of the north Sarawak population, extinct throughout the Holocene, and known only from Niah subfossil cave material. Cranially resembles Pongo pygmaeus borneo but adult male skulls smaller, especially in mandibular dimensions. These cranial characters are unconfirmed in the southern population.