Insect Species Described from Alfred Russel Wallace's Sarawak Collections
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Malayan Nature Journal 2006, 57(4), 433 - 462 Insect species described from Alfred Russel Wallace's Sarawak collections ANDREW POLASZEKl* and EARL OF CRANBROOK2 Abstract In just over a year spent in Sarawak (1854-1856) Wallace collected over 25,000 insect specimens; by his estimate, these represented over 5,000 species. About two-fifths of these were beetles, and over 200 of these were formally described in the 1850's and 60's, largely by 1.S. Baly and F.P. Pascoe. Nearly 400 species of moths from Wallace's Sarawak collections, including over 100 new genera, were described by Francis Walker. Fredrick Smith described 150 Hymenoptera from these collections, and taking into account Diptera and Hemiptera from the same collections, this brings the total number of insect species described to well in excess of 1,000. INTRODUCTION Alfred Russel Wallace has a reputation as an extremely prolific collector of plants and animals from many parts of the world, notably the Amazon Basin and the Malay Archipelago. His relatively brief period collecting in a geographically limited area of Sarawak on the island ofBomeo dramatically illustrates his abilities as a co llector and field taxonomist, able to recognise differences between species across a huge range of diverse insect orders. Wallace in Sarawak Alfred Russel Wallace arrived in Sarawak on 1st November 1854 and left on January 25th 1856 (Wallace, 1869). In a little more than a year, and in a rather limited geographical area of southwestern Sarawak (fig. 1), with the aid of Charles Allen and various local assistants, Wallace collected and shipped back to London more than 25,000 insect specimens. By his own estimate these belonged to about 5,000 species, 2,000 of which were beetles, 1,500 moths and 1,500 other insect orders (Wallace, 1856a). 1 Dept. of Entomology, The Natural History Museum, London SW7 SBD, & Department of Biological Sciences, Imperial College of Science, Technology & Medicine, Ascot, Berks SLS 7PY, UK 2 Glemham House, Great Glemham, Saxmundham IP17 UP, UK. * Corresponding Author 433 Figures 1. Map of Borneo showing Wallace's route (dotted lines). 2. Macronota marmorata Wallace (Coleoptera: Scarabaeidae): actual specimen collected by Wallace (Natural History Museum, London). 3. Trogonoptera brookiana (Wallace) (Lepidoptera: Papilionidae). Photo by Sutiono Sarali. 434 Wallace spent the first four months collecting along a frequented reach of the Sarawak River, from Santubong to Bau and Bidi. Despite obtaining "320 different kinds of beetles" during this period, he considered his results to be "comparatively poor and insignificant" (Wallace, 1869). He then moved to the site of the projected coal works near Simunjan on the Sadong River (1100 46'E, 1023'N) where the following nine month period was by far the most productive phase of his collecting in Borneo. He stated " ... I obtained altogether in Borneo about 2,000 distinct kinds [of Coleoptera], of which all but about a hundred were collected at this place, and on scarcely more than a square mile of ground" (Wallace, 1869). Add to this the 3,000 other insect species (Wallace's 1856a estimate), of which a large proportion presumably also came from the same area, and we have at Simunjan a type locality for several thousand insect species. During his last few weeks in Sarawak, in December 1855 and January 1856, Wallace made two visits to Rajah James Brooke's bungalow 'Peninjau', set in forest on the ridge-top ofSerambu hill (1100 13'E, 102S'N). Here, once again, his collector's instincts were rewarded by a rich catch ofnight- flying moths. On good nights, "they would pour in, in a continual stream, keeping me hard at work catching and pinning until midnight. They came literally by thousands", providing a total collection of 1,386 moths (Wallace, 1869). We can suspect that Wallace was as discriminating in his field selection of moth specimens as he was with other insects, and that his collection from Peninjau was therefore varied and biased towards specimens that he recognised as new to him. Although confirmation is not possible, it is likely that this one site provided a high proportion of his total collection of Lepidoptera. The insects described from Wallace's Sarawak material Perhaps unusually in the history of insect taxonomy, a very large proportion of Wallace's insect species collected during his Malay Archipelago period were formally described as new to science within a couple of decades of his voyages. The dispersal and ultimate fate of these specimens has a complex history, unravelled to a considerable extent by Baker (2001). It is unrealistic, therefore, to hope to track down every one of the insect species described from Wallace's Sarawak material. However, a review of the better-known monographs and papers published within the two decades following Wallace's travels has provided a list of more than 1,000 species described from Sarawak (below). Wallace himself originally intended to work on a much larger proportion of the material he collected than was eventually the case (Wallace, 1905). It is evident from his notebooks of this period (1855-1859) that among the groups he planned to work on were Coleoptera and Lepidoptera. In the end he described comparatively few species from his Sarawak material, but these did include some striking cetoniine scarab beetles. He mentions specifically Macronota marmorata: "A single specimen ofthis fine species was taken on foliage in the swampy forests ofthe Sadong River" (see fig. 2) (Wallace, 1868). Among the butterflies, Wallace famously described the spectacular Ornithoptera brookiana (Papilionidae, now Trogonoptera) named in honour of Rajah (Sir) James 435 Brooke (fig. 3) "whose benevolent government of the country in which it was discovered every true Englishman must admire" (Wallace, 1856b). Wallace described three other butterflies from Sarawak, only one of which remains valid (at species level) today. Recognising Wallace's specimens Many of Wallace's insect specimens from the Malay Archipelago collections were labelled either by Wallace himself, or by his assistant Charles Allen, with a small, round handwritten label. This single label generally bore a three-letter abbreviation ofthe locality, which in the case of the Borneo collections was almost always "SAR" (see Baker, 1996, p. 190), often with Wallace's own collection number. Unfortunately, labels were variously either removed or replaced over the years, by dealers in specimens, collectors and taxonomists, but much of Wallace's material has retained these original labels, and is still readily identifiable. Coleoptera I.S. Baly was the foremost expert on what was known at the time as the "Phytophaga", approximating to the superfamily Chrysomeloidea as understood today. He published a great many descriptive papers, and an attempted synthesis of the Southeast Asian species in his 300-page monographic "Phytophaga Malayana" (Baly, 1865). His contemporary coleopterist F.P. Pascoe also worked on the phytophagous beetles, particularly Cerambycidae (longhorn beetles) and Curculionidae (weevils). Pascoe published an even more extensive monograph, "Longicornia Malayana" (Pascoe, 1864-1869) treating about 1200 species, of which around 280 species were collected by Wallace from Sarawak, and of these, 250 were new to science. Between them, J.S. Baly (1859, 1862a, 1862b, 1864a, 1864b, 1865) and F.P. Pascoe (l857a, 1857b, 1858, 1862a, 1869, 1874) described more than 300 beetle species from Wallace's Sarawak material, belonging to the leaf beetles (Chrysomelidae), longhorn beetles (Cerambycidae) and weevils (Curculionidae). Other coleopteran groups, in particular the jewel beetles (Buprestidae) and click beetles (Elateridae) apparently reached several European continental entomologists, including E. Fleutiaux and R. Oberthuer (Hom & Kahle, 1936). Wallace (1856a) estimated that he collected almost 800 new species in these groups. How many more of these were formally described we have not researched, but a figure of about a thousand new species of beetles from Sarawak seems reasonable. Lepidoptera Most of Wallace's Sarawak Lepidoptera were described by the notorious Francis Walker (Walker 1862-1863). Walker has been described as "having done an amount of injury to entomology almost inconceivable in its immensity" (Anonymous, 1874). However, broader views of Walker 's life and work (e.g. Doncaster, 1961; Graham, 436 1976) reveal many P?sitive aspects, and it seems unlikely that he was as mercenary as has often been depIcted. Doncaster (1961) credits Walker with accurate observation of ap.hids, including .the correct ~ss~ciation of the various morphs that occur during the hfe-cycle. By hIS own admIsSIOn Walker was unable to bring this degree of acc~rate obs~rvati?n to hi~ copious studies of Diptera and Microhymenoptera, but persIsted agaInst hIS better Judgement for reasons of vanity and enjoyment (Graham, 197~). F~om W~llace's Sarawak material Walker described nearly 400 Lepidoptera speCIes, IncludIng more than 100 new genera. Based on Walker's reputation one might expect many of these to have since fallen into synonymy, but according to Dr lD. Holloway, the foremost expert on Lepidoptera of Borneo, most of Walker's Sarawak taxa have stood the test of rigorous taxonomic investigation. Hymenoptera, Diptera and remaining orders Fredrick Smith, another British Museum entomologist, described 150 Hymenoptera species collected by Wallace from Sarawak (Smith, 1857-58; 1873). The taxonomic history ofthe bees from this collection has been treated