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37 BLUMEA tuberous epiphytes The A 335-340 (1993) taxonomic history C.R. Department of Plant Sciences, Oxford of the Rubiaceae 6: of the Hydnophytinae Huxley University, OX1 Road, Oxford, South Parks 3RB, U.K. Summary first described Rumphius recognize the briefly placed further the two elsewhere and genera of the major and Jack described work, confused, and until admirable Beccari’s added. Valeton’s revision genera were Hydnophytinae in 1750, but Hydnophytum was in Myrmecodia Linnaeus 1823. These did not genera were monograph (1884-86) when three sadly incomplete. Introduction The taxonomic history is a presented as this volume), Myrmephytum (Jebb, to the first The first person Rumphius to & Jebb, in Becc. & Jebb, Jack and 1991b), 1991a) background & (Huxley to Jebb, Becc. Squamellaria valid publication who worked for the Dutch East India His classic work 'Herbarium Amboinense' until 1750. published of the work, Joseph did on (ms) (Beaglehole, The third by till names prepared during for the of fire and ant-plants, described the long plant as published, were not the latter halfof blindness, and Nidus formicarum pre-Linnean. Linnaeus, although not niger aware it into his system. after J.D. Epidendroides and the ant-plants tetrandra, of this discovery Hooker had described it encounter with ant-plants descriptions of (1823) Sumatra, described Hydnophytum formicarum niger. No was as in Queens- but his manu- remained plant Myrmecodia described and identified it with specimens have existed, or to of exist also tragic: William Jack's Hydnophytum Island off to was ravages Georg was Ambon in the Moluc- at beccarii 1962). fire and his micarum joint therefore are after his premature death. Jack known the Company the first voyage of the 'Endeavour' found and illustration unrecognized ruber, incorporate not Banks land. Solander delayed by Rumphius' and Nidus formicarum script the concerns prep.), Myrmecodia (Huxley & Jebb, Hydnophytinae (Huxley series since it describe the ant-inhabited tuberous Rubiaceae the seventeenth century, lost in this 1991). Pre-Linnean cas. (Huxley of the genera separate paper Jack Hydnophytum of the and Myrmecodia Myrmecodia Rumphius' Nidus specimens were were published tuberosa from Nias formicarum ruber. He from Sumatra and identified it with Nidus for- ant-plants today. associated with these three authors are BLUMEA 336 Two years after Jack's They neither Rumphius described it as while H. oblong leaves, Jack described H. H. was species a new H. - feature mention- a in western he said had Java and short-petioled leaves. Since, however, ovate the dis- having short-petioled elliptic-ovate leaves, as Later J.D. Hooker tenuous. Java, which he placed western spiny tuber, species had subsessile formicarum Logani- sank H. (1881) into montanum formicarum. Gaudichaud (1830) described collections made in the Moluccas age of the 'Uranie'. He codia he described also sank the genus mis which without reference wrote Mirmecodia [sic] as M. tuberosa Jack the name is A. P. de Candolle H. formicarum Jack and his to indicate a M. species according to H. nizing For a M. armata formicarum while there DC. is Jack and H. was fruit of what is in fact were Becc. later (= as montanum used the name which he identiHe created Rumphius. a of species followed (1833) in in (M. and H. a tuberosa Jack, thinking name recog- but most authors apparently meant he had found numerous seeds Hydnophytum with loranthifolium (Benth.) but Jack, hispida, two pyrenes. He revived in it species two Hydnophytum by Beccari (1885); they oblonga) He Blume. mentioned M. an error with M. iner- De Candolle followed Blume in genus in the Rubiaceae and described a placed L. correct. (1834) M. echinata. Then Bentham made a name unfortunately Myrmecodia ruber of formicarum peace; Henschel followed De Candolle. Richard Lasiostoma of spineless species plant illegitimate. the published voy- Myrme- Blume) and also Gaudichaud's M. echinata. Since Gaudichaud's superfluous, was and hence include Blume's collection from Java armata to Freycinet's on Blume. The collection from the Moluccas. new the genera, but (1830) re-separated or but since he identified the into Mirmecodiaand fied with M. tuberosa Jack and Nidus new echinata, Sprengel to nomenclaturally superfluous Hydnophytum comprised M. inermis Gaudich. in a Hydnophytum This montanum. from species in the position by Blume (1826/27). from Jack. He also found formicarum somewhat transferred both (= Strychnos L.) proper but noted that it had Jack, nor their to of Myrmecodia specimen a in M. tuberosa tentatively tinction 1993 2, Lasiostoma Schreber to however, returned were, Blume also collected by No. publication Sprengel (1825) their genera in the Rubiaceae ed 37, 1824-1872 Confusion aceae. Vol. Becc. H. are (= L. (1843). These oblongum (Benth.) loranthifolia) from New Ireland and New Guinea respectively. Miquel (1857) recognized the pre-Linnean num name Blume's Blume; he also observed that these Myrmecodia name niger for these two In 1858 Asa American M. echinata Gaudich. He described Myrmecodia no small two was an of Arts and new species, imberbis which mellaria. Gray is Gray presented account one about the by one or the to be to this by following In Jack de Candolle's name M. was not published wrote Fiji to the until 1860. and the other to a new and ar- followed Blume. longiflorum Beccari species monta- include Blume's collection existing literature old H. distinguished. M. inermis, (1861) Miquel Hydnophytum two but he transferred of Seemann's collections from later transferred rather confused obscurity was to ignoring Sciences, though was scarcely name collections. In his flora of Sumatra Academy Hydnophytum are superfluous from Java and Gaudichaud's from the Moluccas, mata of from H. formicarum Jack species two he removed De Candolle's instead. He retained the species two Nidusformicarum genus sadly of this genus and Squa- "There respecting C.R. the distinction between it and Seemann, apparently up." vitiensis, but M. name More confusion on of still was collections arise. to be to lected an error by Zippelius removed it from the tion of Jack and H. Baron next von From the for H. Another red montanum. and referred ant-plant montanum by referring by Zippelius report Mueller was of (1871) collected ant-plants referred them to ‘Genera Plantarum’ the distinction between well floral as M. imberbis A. (1873) ones. They In 1874 Beccari entered the (1884) to herring and also a by with arena to a new Myrmecodia again myrmecophytic he the saw genus and that the zippelianum. and M. Australia. armata. succeeded in (1873) stem clarifying and tuber char- that M. inermis A. Gray (meaning describe it. not of Myrmecodia selebica which Myrmephytum Becc. Von Mueller in them to Papua, assigning on 1875 H. formicarum the Andaman Is- Jack. (1880) suggested should be two no flowers. No-one ideas. In 1880 Britten mentioned M. mistake for M. inermis. In the (1886) that regarded Hydnophytum as be sunk section of the a non- L. He also mentionedLasiostoma Bentham of which Uragoga but found specimens were (1884-86) description a genus Beccari Cape York, Myrmecodia, noting recognized H. formicarum as ovatum Blume which ap- called H. (1885) Nares from distinct genus, but did Baillon in his 'Histoire des Plantes' into that he his H. lanceolatum col- was echinata. In 1877 Kurz found Hydnophytum lands and identified it H. species ellipticum H. formicarum listed D'Albertis' collections from southern Myrmecodia a new Cephaelis laevigatum; Beccari to Hydnophytum Gray) belongs he later transferred and plant same clear 1861 the also mentionedanother incomplete collec- Miquel genera. to H. Bentham and Hooker in their 'Genera Plantarum' acters as in Blume and concluded they to from ?New Guinea which Beccari Zippelius' The the was named Miquel (1869) distinct. He confused later writers pears out, this to ought published and De Vriese from Temate. He discussed the sim- by Teijsmann ilarity between H. formicarum not Gray's description, Gray (1862) pointed as 337 Hydnophytinae which the Dutch botanists Hydnophytum unaware the history of longiflorum. had called H. based Taxonomic Huxley : same glabra seems to have followed Baillon's from Borneo, but this year Beccari cited two species was of probably Myrmecodia a in Count d'Alberts' book 'New Guinea': M. albertisii and M. muelleri. J.D. Hooker in carum lected in that 1881 took the step of Jack. Home in the one to species, omissions, morphology of the of these Odoardo Beccari were not H. wilsonii, Squamellaria sinking year listed Fiji, though descriptions of these transferred it merous same was two H. montanum species published not a of the Blume into Hydnophytum till 1883 by group, in gave a a paper H. formi- he had col- Baker who realized typical Hydnophytum in 1886. Franz Antoine taxonomy of ; history, indeed Beccari albeit with mainly devoted nu- to the as the plants (1882). published his monumental study of ant-associated plants second part of his three-volume work 'Malesia' between 1884 and 1886. This volume is subtitled 'Piante Ospitatrici; ossia piante descritte ed illustrate da O. Beccari'; plants, but the Hydnophytum was greatest part was based familiar with the concerns on type formicarie della Malesia it deals extensively e della Papuasia with other ant-associated the Rubiaceae. His revision of Myrmecodia and considerable material in knowledge Europe. of the He added plants 16 in the field and he species to the existing BLUMEA 338 of Myrmecodia and 29 two he removed form the to the six of to new Vol. genus phytum accommodate his to not believe six-merous corolla and ters, the of species ants From Beccari Valeton to After Beccari there species but with a species species of in finds. of Archipelago. from the British of specimen ted new and of piecemeal work with in began colony of (1927). But he of J.D. species. new plant distinctive a which had been and moved the an error a careless and colony two of as of Drake H. and Myrmecodia species led exploration new Hydnophytum species a new moment new species activity 1915) of one from Hydnophytum described (1895) costatum; this was a spot- to of Hydnophytum at an elderly Kew, and and Elmer (1911, of man; he and he left undescribed ant-plants saw cies in New Guinea followed of three and Hydnophytum fared a has taken up of two are little better. His two of by descriptions photographs of these from New Guinea in his of key were not Myrmecodia von (1911, Papuasien' seven and consists of new a and H. key to of which species, 16 but spe- frag- species new have survived and P. (H. vaccinifolium specimens. on only fragmentary, all that survived. His manuscripts names from the the material neither at Florence names many of the collections he described account 1913, 1934) Myrmecodia be sunk. species ailing in 1905 and ceased but American authors and four of Hydnophytum named several later lost in Berlin. Thus his (1983) the was species named (1907, 1908, 1913, eight species already was nor Unfortunately, ments of when he undertook the Rubiaceae for Lauterbach's 'Flora (Beccari's) were German added (1892) In Papua. a (1898) Hooker Almost all of these will have Valeton had 1912a, b) Merrill total of Philippines. charac- Beccari also recorded description named three 1894) in 1913. Then German two more a had Myrmedoma, vegetative 1924. earnest: described genus Myrmecodia beccarii, but found that he had made Schumann and Lauterbach described a Rechinger genus, Myrmecodia. Scyphiphora hydrophyllacea (Rubiaceae) Pittard in by monotypic in Australia. This while Schumann Myr-mecodia, as Myrme- 1898. Warburg (1891, the Bismarck new of the tubers. Northeastern New Guinea had become to new a Banks and Solander. Schumann in 1888 honoured Beccari Hydnophytum, Psychotria to phase a vegetation published by not ?wil- (1927) was from mangrove seen origin honoured Beccari's (1886) Hooker and H. be ant-associated to recognized from each genus Gray present in the ant-plants, discussed theirrelationship with and considered the plants, Squamellaria but these differed in genera large bracts, resembling respectively Hydnophytum the different species imberbis A. previously described Myrmecodia selebica. This satellite with his other genus Myrmephytum, along a One Hydnophytum. observation of tubers. He also Gray's 1993 2, Squamellaria (M. sonii Home ex J.G. Baker). He did he missed No. 37, van of Royen crassicaule). After Valeton In the species same These three viously In year that Valeton's work collected by Brass species described by of on one Hydnophytum Hooker and 1940 Fosberg described a appeared, of the many and were, Beccari, new Moore (1927) important described three Archbold however, all very close which Moore did species of not Hydnophytum to Expeditions. species apparently from the pre- examine. Solomon Huxley : C.R. the history of Taxonomic 339 Hydnophytinae Islands, and in 1942 Bremekamp published H. inerme (Gaudich.) Bremek. gaudichaudii combinationfor H. type for H. lections came of these species Finally, of from areas species see missed. were a existing types, num- of the col- Many and variability range therefore often under-estimated. was in 1967 A.C. the flora of Smith, studying and called it S. Squamellaria the and 15 of Hyd- Myrmecodia not before and little since, the explored not did they as of species seven and identities with earlier relationships a new was Jack. Perry published from the Brass collections. But nophytum specimen as which the type for M. inermis Gaudich. since this not was formicarum In 1945 Merrill and ber of Becc. gaudichaudii included H. species Becc. But the Gaudichaud found Fiji, distinct a species major. CONCLUSION The taxonomic was the This led him to the available its types. His plants as the styly of only the commonest existing specimens synthesis or of these first-hand over much of theirrange and at is the only which one Beccari understanding'; unique symbiosis interest in the ochlospecies H. formicarum, edge to an covers subgeneric classification, attempts and the association with led by treatment discusses interpret we the study distribution, what of the group shows little history author stimulated only to to the group plants. examine all throughout define the variation of and which considers hetero- Valeton, Merrill, and Perry had first-hand knowl- ants. with their limited acquaintance species; this, together their work little more than being listing a of new with collections. ACKNOWLEDGEMENT I would like to thank F. White. supervisor my REFERENCES d'Albertis, L.M. Antoine, F. Zeitschr. 32: Baillon, what I did 1883. On des Plantes the flora of 1962. New South Wales The and Angus & 1874. Descrizione O. 1884-86. Malesia, vol. 2, 1843. Enumeration C.L. 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Lauterbach, Beitrage zur Flora 127-150. zur Kennmis Hellwigianae. der from 503-514. Hydnophytum spathulatum. von noteworthy flowering plants Systema vegetabilium Nova Guinea Rubiaceae & 90-91. plants. ... 256. 18. New papuanischen Flora. Bot. Jahrb. 18: Bot. Jahrb. 209-211. 13: 440-441. von