Article

Butterflies

From Munshipedia, the MBRAS digital historical encyclopedia

Butterflies

Butterflies of Borneo and the Malay Peninsula have attracted sustained scientific attention since the late nineteenth century, with the region’s extraordinary diversity and the accessibility of its highland and lowland habitats making it a focal point for lepidopterological research. The faunal record, as documented in the Society’s journals, spans from Shelford’s early faunal checklists of Sarawak through Moulton’s comprehensive Bornean catalogues to the mid-twentieth-century taxonomic notes on Peninsular forms, revealing a fauna characterised by high endemism, complex mimicry systems, and notable biogeographic connections to the Philippines and the Sunda Shelf [1, pp. 81–111], [2, pp. 73–177]. The literature also captures dynamic ecological phenomena—mass migrations, seasonal abundance fluctuations, and predator-prey interactions—that illustrate the ecological role of butterflies in tropical forest and lowland environments [3, pp. 203–204], [4, pp. 314–315], [5, pp. 230–231].

Taxonomic Scope and Faunal Composition

The earliest systematic attempt to enumerate Borneo’s butterflies was undertaken by R. S. Shelford, Curator of the Sarawak Museum, who published a series of faunal checklists between 1901 and 1906, each covering a different group of subfamilies [1, pp. 81–111], [6, pp. 29–42], [7, pp. 89–136]. Shelford’s 1904 paper, covering the Danainae through Amathusiinae, enumerated 101 species and described four new species: Lethe cerama, Ypthima abnormis, Erites thetis, and Elymnias brookei, the latter named in honour of the reigning Rajah of Sarawak [1, pp. 96–102]. He identified at least six species confined entirely to Borneo, including Euploea scudderi, Euploea zonata, Euploea masina, Elymnias hecate, Elymnias dara, and Thaumantis aliris [1, pp. 81–111]. Shelford’s central argument was that Borneo, having been “so well explored faunistically,” would yield few further new species, and that a comprehensive list compiled at that juncture could lay claim to “completeness and finality” [1, p. 81]. He justified this ambition by noting that the most recent prior list, Bartlett’s 1896 catalogue, had recorded only 139 Lycaenidae, whereas the number known by 1904 already exceeded 300 [1, p. 81].

Shelford’s 1901 paper on Mount Penrissen, Sarawak, reported butterflies collected in May 1899 at elevations of 2,800 to 4,200 feet, and argued that the mountain’s dense, uncleared jungle produced a comparatively sparse but taxonomically distinctive lepidopteran fauna [6, pp. 29–42]. Several species in the Penrissen assemblage had never been recorded from the more frequently studied nearby peaks of Matang and Santubong despite “the most careful and frequent collecting” there [6, p. 29]. Shelford explicitly framed the mountain fauna as analogous to an insular fauna, a concept that anticipates later island biogeographical thinking [6, p. 29].

J. C. Moulton’s 1911 list of the butterflies of Borneo, spanning 104 pages, represented the most comprehensive treatment of the island’s fauna to that date [2, pp. 73–177]. Moulton, then Curator of the Sarawak Museum, subsequently published a revised and expanded version in 1914 [8, pp. 1–56]. In 1913, Moulton also published taxonomic revisions to a portion of Shelford’s 1904 list, addressing the subfamily Amathusiinae and a single Discophorinae entry, incorporating the nomenclatural changes necessitated by Stichel’s Genera Insectorum (1905–6) [9, pp. 13–18]. Among the species Moulton noted as great rarities was Zeuxidia aurelia aureliana, recorded only from Lawas, Malinau, and Paku [9, pp. 13–18].

In the Malay Peninsula, Moulton’s 1923 paper, published as Director of the Raffles Museum in Singapore, described three new subspecies of Pierine butterflies from Pulo Aor and Pulo Tioman off the East Coast: Appias paulina grisea, Udaiana cynis tiomana, and Gandaca harina aora [10, pp. 233–236]. These island forms were distinguished from their mainland counterparts by specific differences in wing coloration and marking, illustrating the role of small islands as sites of incipient speciation [10, pp. 233–236].

Biogeographic Patterns and Mimicry

Shelford identified a distinct “Philippine element” in the North Bornean fauna, citing species such as Euploea leucostictos kadu, Euploea dufresnei tyrianthina, and Clerome phaon as evidence of biogeographic infiltration from the Philippines [1, pp. 92–93]. This observation established a pattern of faunal exchange across the South China Sea that subsequent workers would refine. Moulton’s 1923 paper extended this theme to the Malay Peninsula, where Hebomoia glaucippe aturia from Pulo Aor showed a pale lemon tinge distinguishing it from the dead-white Peninsula form and possibly linking it to the yellower Philippine forms [10, pp. 233–236].

Mimicry complexes received particular attention in Shelford’s 1904 paper, which documented extensive mimicry among the Danainae and Elymniinae, listing the specific mimetic species for each model [1, pp. 81–111]. In his 1901 Mount Penrissen paper, Shelford proposed a theoretical framework for Lycaenid underside wing-pattern evolution, arguing that “the more highly organised the butterfly, the more the underside wing-pattern tends to disappear from the discs of the wings and concentrate at the margins and at the anal angle of the hind-wing” [6, p. 34]. He linked this to a general principle of defensive specialisation: as discal patterns fade, the marginal eyespot becomes relatively more conspicuous, directing predator attacks away from vital body parts, while the unornamented disc reduces the risk of attracting attention to the body itself [6, pp. 34–35]. He illustrated the continuum with Loxura atymnus, in which the eyespot is replaced entirely by hindwing tails [6, p. 35].

Shelford also made the notable observation that Bornean butterflies do not exhibit the seasonal (dry- and wet-season) morphological variation well known from Indian populations; all Mycalesis species in his collection corresponded to the wet-season phase, and both seasonal forms of Melanitis ismene flew simultaneously regardless of the season [1, pp. 98]. This suggested that the island’s more uniform rainfall regime eliminated the selective pressure for seasonal polymorphism [1, pp. 98].

A further biogeographic anomaly was documented by Moulton in 1923, who recorded the occurrence of two Palearctic species in the tropics: Pieris canidia, known from India through Burma to China and the Philippines, appeared in Singapore in 1922–1923 with no records from Peninsular Siam or the Malay Peninsula, strongly suggesting introduction by human agency, possibly via larvae transported on vegetable matter to a major port; and Pieris rapae, a species ranging from Europe to Afghanistan and Kashmir, was recorded from Kuala Lumpur in December 1917, an occurrence Moulton considered explicable only by human-mediated introduction [10, pp. 233–234].

Ecological Dynamics: Migration, Predation, and Behaviour

The phenomenon of mass migration was documented on several occasions. Shelford’s 1903 note recorded an abnormal swarm of Cirrochroa bajadeta in Sarawak, which he attributed to a comparative drought (rainfall well below the 75.17-inch average) and the exceptionally favourable monsoon conditions for insects [3, pp. 203–204]. He corroborated the broader insect-favourable conditions with three additional observations: an increased number of social wasp and bee swarms over the preceding three months; a swarm of dragonflies reported by Captain A. Balser aboard the s.s. Rajah of Sarawak on 20 January, approximately 50 miles west of St. Pierre Island, moving northward in unsteady winds; and a swarm of Pierine butterflies (probably Catopsilia crocale) observed by Mr. H. B. Crocker at Paku, Upper Sarawak, on 27 January, flying in a solid phalanx roughly 20 fathoms long by 8 fathoms wide in a westerly direction [3, p. 204].

A more detailed migration account was provided by the Rev. R. Cardon in 1927, who documented a mass migration of Catopsilia pomona in Upper Perak in May 1927, moving from north to south [4, pp. 314–315]. Cardon described encountering the migration between Lenggong and Lawin, where thousands of the butterflies flew in dense swarms along the road, with some resting so thickly on wet ground that they were at risk of being run over by vehicles [4, p. 314]. He confirmed through netting that the migration was composed almost exclusively of C. pomona, with only a few C. pyranthe, C. chryseis, and C. scylla present as non-migratory associates [4, p. 314]. He identified two female forms among his captures: a new form bidotata with two patches on the underside of the posticae, and the common catilla form with a large discal reddish patch [4, pp. 314–315]. The migration lasted approximately four days, from 26 to 29 May, after which only isolated individuals remained [4, p. 315]. Cardon briefly compared this to a 1902 migration of Delias aglaia parthenope in Singapore and to Rev. L. C. Biggs’s 1881 account of D. aegialea migrations in Java [4, p. 315].

Predation on butterflies was investigated by C. L. Collenette in two short notes published in 1922. In the first, Collenette argued that the eye-spots and tails of Lycaenid butterflies in Malaya function primarily as protection against lizards rather than birds [5, pp. 230–231]. He observed that Lycaenids in Malaya live close to the ground, alighting at heights of 4–6 ft. when disturbed, and that birds are uncommon in the lower jungle while lizards are abundant at low elevations [5, p. 231]. His observations of settling behaviour—butterflies choosing exposed upper-leaf positions with the head lower than the tail—suggested the protective apparatus is oriented toward the branch side, i.e., toward the direction from which a lizard would approach [5, p. 231]. To test the hypothesis experimentally, Collenette liberated 17 Lycaenids in a room in Penang in January 1921 alongside Hemidactylus frenatus (Chi-chah). Three butterflies were attacked: one was taken by the head and eaten, while two were attacked from the tail and escaped when the lizard’s snap missed entirely [5, p. 231]. He further noted that the broken portions of worn hind wings resemble the rounded shape of a lizard’s mouth rather than a bird’s bill, and that the proportion of Lycaenids with damaged hind wings may reach 10 per cent in specimens that have been flying for several days [5, p. 231].

In his second note, Collenette documented field observations of predation in Singapore, made in January–February 1922 [11, pp. 268–269]. He described observing a Praying Mantis (Rhombodera basalis) perched on a Papaya tree, where it successfully captured a Skipper (Hidari irava) and attempted to seize a Hawk Moth (Hippotion boerhaviae), missing on both occasions [11, p. 268]. He noted the Mantis’s camouflage and its apparent inability to reason that prey was available on nearby flowers, remaining just out of reach despite repeated visits over three weeks [11, p. 269]. A second observation recorded a Chameleon (Calotes cristatellus) in the Botanic Gardens positioned near a cluster of Ypthima butterflies, remaining motionless for twenty minutes [11, p. 269]. Collenette also reported finding a powerful Euthalia merta caught in a spider’s web across a path in the Changi jungle, undamaged but helpless [11, p. 269].

Moulton’s 1912 expedition to Mount Batu Lawi in the Limbang district of Sarawak provided valuable field observations of butterfly behaviour in a highland environment. At the kuala Madalam, Kallima inachis buxtoni was observed and noted as “not uncommon in the Limbang district but rarely met with in other parts of Sarawak,” indicating a degree of local restriction [12, p. 13]. Papilio payeni brunei and Amblypodia narada were both recorded as rare species in Sarawak; the latter was captured feeding on animal excrement on a sunny gravel bed in mid-stream [12, p. 13]. At the kuala Madihit, Moulton observed a Lycaenid butterfly (Allotinus nivalis) in the act of attending a Homopteran larva in the presence of two small black ants, a trophallaxis-like interaction he had not previously witnessed personally [12, p. 23]. At the summit of Mt. Selinguid (4,850 ft.), butterflies were described as “plentiful” and the party spent most of a full day collecting there [12, p. 41]. By contrast, on the upper slopes of Batu Lawi itself, where the party reached an altitude of 5,660 ft., Moulton noted but one butterfly (an Hesperid, probably Bibasis uniformis), and attributed the dearth to the particularly wet conditions and dense humidity [12, pp. 40–41].

Research and Documentation

The Society’s literature on butterflies reveals a clear evolution in scope and methodology across the period from 1901 to 1927. The earliest contributions, by Shelford in the first decade of the twentieth century, were primarily faunal checklists and species descriptions, reflecting the colonial-era practice of government officials and museum curators producing natural-history contributions from material accumulated in institutional collections [1, pp. 81–111], [6, pp. 29–42]. Shelford’s work was characterised by an ambition towards completeness, a methodological commitment to the trinomial system, and an interest in theoretical questions of mimicry and seasonal polymorphism [1, pp. 81–111], [6, pp. 34–35].

The 1910s saw a shift towards more comprehensive regional catalogues. Moulton’s 1911 and 1914 lists of Bornean butterflies represented the most extensive treatments of the island’s fauna, drawing on the Sarawak Museum collection and field material from expeditions such as the 1911 journey to Mount Batu Lawi [2, pp. 73–177], [8, pp. 1–56], [12, pp. 77–88]. Moulton’s 1913 revisions to Shelford’s list demonstrated the ongoing process of taxonomic refinement as new nomenclatural authorities were published [9, pp. 13–18]. The Batu Lawi expedition report, a collaborative effort involving H. N. Ridley, E. B. Copeland, J. J. Smith, A. Griffini, and F. F. Laidlaw, reflected the multi-disciplinary approach typical of Sarawak Museum expeditions of this period, with the butterfly collection forming one component of a broader faunal and ethnographic survey [12, pp. 77–88].

By the 1920s, the focus had broadened to include the Malay Peninsula and to address ecological and behavioural questions. Moulton’s 1923 paper on Pierine butterflies new to Malaysia, published from the Raffles Museum in Singapore, extended the taxonomic work to Peninsular island forms and raised the question of human-mediated introduction of Palearctic species [10, pp. 233–236]. Collenette’s 1922 notes on predation and protective devices represented a shift towards experimental and observational ecology, moving beyond mere enumeration to address the functional significance of morphological features [5, pp. 230–231], [11, pp. 268–269]. Cardon’s 1927 migration report and Brooks’s 1923 note on the early stages of a Danaine butterfly from West Sumatra [13, pp. 260–261] further illustrate the diversification of interests, encompassing population dynamics and life-history studies.

The archival record also reveals the institutional infrastructure that supported this research. The Sarawak Museum collection, built through colonial-era collecting expeditions, served as the primary source for Shelford’s and Moulton’s Bornean lists [1, p. 81], [9, pp. 13–18]. The Raffles Museum in Singapore, the Selangor Museum, the British Museum (Natural History), the Hill Museum at Witley, and the Oxford collection formed the standard network of colonial and metropolitan entomological repositories from which comparative material was drawn [10, pp. 233–236]. The collaboration between local curators and metropolitan authorities—Shelford’s reliance on determinations by Dr. A. G. Butler and Mr. F. A. Heron at the British Museum, and Moulton’s use of comparative material from multiple institutions—was a defining feature of the period’s taxonomic practice [1, p. 81], [10, pp. 233–236].

MBRAS Sources

References

  1. R.S. Shelford (1904). A list of the butterflies of Borneo with description of new species JSBRAS 41: 81–111.
  2. J.C. Moulton (1911). A list of the butterflies of Borneo with descriptions of new species JSBRAS 60: 73–177. Read on JSTOR
  3. R.S. Shelford (1903). A swarm of butterflies in Sarawak JSBRAS 39: 203–204.
  4. R. Cardon (1927). On a migration of Catopsilia pomona (F. JMBRAS 5(2): 314–315. Read on JSTOR
  5. L. Collenette (1922). Protective devices by Lycaenid butterflies against the attacks of lizards and birds JSBRAS 85: 230–231. Read on JSTOR
  6. R.S. Shelford (1901). A list of the butterflies of Mt. Penrissen, Sarawak, with notes on the species JSBRAS 35: 29–42.
  7. R.S. Shelford (1906). A list of the butterflies of Borneo, and Nymphalinae JSBRAS 45: 89–136.
  8. J.C. Moulton (1914). A list of the butterflies of Borneo JSBRAS 67: 1–56.
  9. J.C. Moulton (1913). Notes on a portion of the late Mr. Shelford’s list of Bornean butterflies, Part 1, published in the Society’s journal No. 41 JSBRAS 65: 13–18.
  10. J.C. Moulton (1923). Some Pierine butterflies new to Malaysia JMBRAS 1(1): 233–236.
  11. L. Collenette (1922). Notes on the enemies of butterflies JSBRAS 86: 268–269.
  12. J.C. Moulton (1912). Butterflies taken on the Batu Lawi expedition JSBRAS 63: 77–88.
  13. C.J. Brooks (1923). Early stages of a Danaine butterfly JMBRAS 1(1): 260–261. Read on JSTOR