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A hundred-year-old worm? R. J. Blakemorel and K. L. Elton2 'Department of Zoology, University of Queensland, Brisbane, Queensland 4072 Present Address: CSlRO Entomology, Canberra, ACT 2601 ?Department of Agricultural Chemistry and Soil Science, University of Sydney, New South Wales 2006 ABSTRACT -~~ IS reTd.h .e. . es aA. ubset.d.r a- .iln ia- n~o ce taarl t,h awn.o r ImlL. sS-trorae-tnl oc-ne- r 1si e- lpl-ar o rnv-laa-ce-lde- aa ,vni o (.F ltest clhaex- ro n1o8.m.8-9y,) ,I1sM dw sacsum~s~ls ee~cd~i d~Sa~p~e e: Oc- ~limwecnhsa~w ~ee tca~el . mlleclec from manage0 and at R~chmonaN. SW, a regon wlUlin ifs onginal dlstnbutlon range. I1 was In close assoclatmn wlth the lntrod~cedlu mbrlc d Aponeclodea lrapezwdes (D~gbs)T hls s the flrst cent rmahon of S. macleayr slnce ts deslgnat on over a cenldty ago an0 demonstrates that Ihls natlve species has survived the clearing, cul~vationa nd introduction of exotic species that has accompanied Eum~eans ettlement. INTRODUCTION Type-locality: Sydney (33"53'S, 15lo12'E), "common in the Hon. W. Macleay's garden at Earthworms were collected as part of an Elizabeth Bay" - Fletcher (1889). agronomic study conducted in 1992123 into the distribution of soil fauna in pastures near Distribution Records: Mount Victoria, Blue Richmond, which lies 55 km north-west of Mountains (33"35'S, 15Oo15'E) - Fletcher Sydney and towards the Blue Mountains, (1890, variety c(i)); Raymond Terrace (32'47'S, NSW. There have been few if any systematic 15lo45'E) and Morpeth (32'44'S, 151°38'E) surveys in this region apart from the early - Fletcher (1890, variety c(ii)); Coona- reports of Fletcher (188&9O), consequently, harabran (31°16'S, 14g018'E) and Gunnedah no comprehensive guide to earthworms exists. (30°59'S, 150°15'E) from the banks of the Moreover, earthworm identification usually Namoi - Fletcher (1890, variety ~(iii))a,l l in entails the sometimes laborious task of micro- NSW; ?Deniliquin, NSW (35'33'S,144'58'E) scopic dissection before a specimen can he - Barley and Kleinig (1964). compared with descriptions of species that are known. This paper describes the process of Syntypes: W1344-1345 in the Australian identification of one native species. Museum listed by Reynolds and Cooke (1976). METHODS AND CONVENTIONS Material examined: Richmond, NSW (33"36'S, 15O044'E) from the Yarrumundi Dissections and illustrations were by RJB paddock 3 of the University of Western using a Wild M5 microscope with camera Sydney, Hawkesbury campus. Collected by luclda. In the figure (Fig. l), the clitellum is Karen Elton, January 1993, under grazed two shown shaded. Abbreviations used are: i ii, year-old sown grasslmedic pasture following etc., segments numbered from the peris- oats in a paddock much cultivated previously. tomium; 112 213, etc., intersegmental furrows, Found in association with the lumbricid i.e., between i and ii, etc.; a, b, etc., individual Aponeclodea trapezoides. The site forms part of setae from the most ventral on each side; a Quaternary terrace of the Nepean and ACT, Australian Capital Territory; NSW, Georges Riven with soil, mapped as Blackendon New South Wales; Qld, Queensland; sp.p., sand, a slightly acidic, nutrient-poor red spermathecal pores. podzolic with a well-drained sandy profile to the depth of sampling (0.9 m). SPECIES DESCRIPTION Deposition No.: AM W21710: four mature Spenceriella macleayi (Fletcher, 1889). specimens, two contracted on preservation, Comb. nov. two relaxed hut macerated (one a posterior (Fig. 1) amputee). All inspected, one of each dissected Perichaeta maclea9 Fletcher, 1889: and one contracted specimen drawn (Fig. 1). 155S558. Perichaeta macleayi var. c (i)-(iii) (?non P. macleayi vars. a and b) Fletcher, 1890: ~ ~features t ~ ~ ~ ~ l I. n. n. -~ -0..0 .7 .. Megoscolex macleayi; Beddard, 1895: 376; Length: 40-75 mm. Width: mean about 4 (?non vars a, b); Michaelsen, 1900: 223. mm. Segments: 90 for all three complete December 1994 Australian Zoologist, Voi. 29(U) 251 Female pores: paired on xiv, just anterior to setal arc nearly the width of the mid-ventral gap. Male pores: largish lateral slitse quatorial on xviii on summits of a pair of opposed raised porophores. Genital Markings: in x and xi, a pair of raised ventral mounds extending from a to c setae,e ach with a trans- lucent, ellipsoid rim to an eyeJike pore on the summit. The anterior pair slightly larger. In xvii and xix are paired genital papillae, just ventrad to the line of the male pores, on each side are a presetal and a postsetal pair; in addition there are smaller mid-ventral genital markings, a pair presetally in xvii and a pair (an analogue in one specimen) postsetallyi n xix. In xviii a single presetal mid-ventral papilla was sometimes present. The anterior annulus of xvii and the posterior annulus of xix are tumid where they accommodate the ventral papillae thus the male field appears as a.basin-shapedd epression. Internal anatomy Septa: 415 weak to anterior of gizzard, 516 and 617 displaced and adpressed by gizzard,t hickening to 9/10 or l0/l I thereafter thinning to become membranous after L4l15. Dorsal blood vessel: single, continuous onto pharynx. Hearts: commissurals viix, hearts x-xii. Gizzard: large barrel or cone-shaped in v, has muscular sheen but can be readily deflected. Calciferous glands: oesophagus Figure 1. SpencerielLam acleayi (a) ventral view with (b) right- dilated in viii-x and in xiii, but more so in hand-side prostate gland and (c) spermathecae shown in each of xi and xii to form a pair of ventro- relative positons; (d) dorsal view of prostomium. laterally sessilec alciferous glands that have internal lamellae (i.e., two pairs). Supra- oesophageal vessel not detected. Intestine specimens.S light secondarya nnulation from origin (caeca, typhlosole): xv, acaecate, vi. The posterior eight segments dorsally typhlosole not found. Nephridia: mero- canaliculatei n one sDecimen.C olour: dorsum nephric with pairs of large tufted nephridia dark-reddish to chbcolate brown especially ventrally in iv-vi, reducing to parietal forests anteriorly, setal auriolae and ventrum pale, of micromeronephridia by the clitellar region clitellum buff. Cuticle not especiallyi ridescent. and thereafter. Pre-septal funnels not seen Prostomium: open epilobous or closed by (possibly numerous per segment?). Sperma- a fine groove parallel and close to inter- thecae: two pairs in viii and ix with large sub- segmental furrow l/2. Peristomium ventrally spherical or elongate ampullae tapering to cleft. First dorsal pore: 5/6 (by mucal ejecta), ducts each bearing near ectal end a shorter, continuous on clitellum. Setae: perichaetine white, digitiform diverticulum. Ovaries: in xiii (numerous) from ii: 2Olvii and t24lxx, as a large pair of palmate strands of numerous with wide mid-ventral and mid-dorsal gaps egg strings with paired diaphanous funnels. which preclude setaeb etween the male pores. Male organs: holandric, paired sacsi n x and Nephropores: not seen (enteronephric). xi contain unusually large palmate or "tufted" Spermathecal pores: in 7lB and 8/9, difficult testes and non-iridescent. nebulous funnels. to determine in deep intersegmental furrows Paired seminal vesicles are lobulate on approximately as wide apart as male pores posterior septum of ix and racemose on (i.e. in line with setae c or d). Clitellum: anterior septum of xii. Prostates: large, multi- annular over r/zxlli-Vzxvii, interrupted lobed racemose glands extending over 24 ventrally in xvii, furrows and setae retained. segments between xvii and xx, with short 252 AustraliaZno ologistV, ol.2 9(34) December1 994 muscular ducts once bent. Gut contents: and (3) position of the two pairs of sper- when the fine, yellow soil and occasional mathecal pores. Variety c from three locations organic "husks" were flushed away, large given in the distribution records above), how- quartz crystals (up to 0.5 mm diameter) ever, were of similar size and consistently had remained. These were present the length of the distinctive papillae in x and xi with the gut and may have been selectively "opposite the interval between the first and ingested to serve as "crop stones" or, alterna- second setae a pair of fossettes". Consequently, tively, they may represent involuntary ingestion only variety c is included here in the of a sandy medium. These worms are probably synonomy of S. macleayi whilst the other geophagous. varieties are provisionally excluded (cf. Michaelsen 1900). The arrangement of paired genital mark- Taxonomic Remarks ings on x and xi, characteristic of S. macleayi, This species is clearly assignable to the is found also in S. fecunda (Fletcher 1887a: genus Spenceriella, Michaelsen (1907), as it 401-02) from the Blue Mountains, NSW, falls within the range of the generic definition which nevertheless has four pairs of sper- as emended by Jamieson (1974) (following the mathecae, calciferous pouches in x-xiv and discovery by Jamieson, 1972, that neotypic last hearts in xiii. Unravelling the relationship material of the type-species, Spenceriella of S. macleayi with the S. fecunda "variety" from notabilir (Spencer 1900), had racemose Burrawang, NSW, briefly described by prostates, rather than tubular prostates as Fletcher (1890: 1007-008) as having only two originally described by Spencer). The only pairs of spermathecae, is not attempted here. notable departure from the generic definition is that there are two, rather than three or Diagnosis: Spenceriella macleayi, as described four, pairs of calciferous glands. above, is distinguished from other members of Spenceriella by the combined characters of The above description agrees on each point widely paired genital markings, each with a with the original description of external central pore-like depression ventrally in x and features of Perichaeta (=Spenceriella) macleayi xi; paired papillae in xvii and xix; tufted (Fletcher 1889: 1556558), especially with nephridia anteriorly; two pairs of regard to biometry and the arrangement of spermathecal pores in 718 and 819; calciferous genital markings. They also agree in having pouches in xi and xii and last hearts in xii. two pairs of calciferous pouches in xi and xii, which were later described as "so incompletely pinched off as to be little more than dilations" DISCUSSION (Fletcher 1890: 1004). The internal anatomy concurs with Fletcher's (1989) statement that J. J. Fletcher was one of the earlier Aust- in other respects this species was like ralian workers who named about 20 native Perichaeta (=Spenceriella) australis (Fletcher Perichaeta species, besides S. macleayi, in a 1886: 561-65), notwithstanding his mislocating series of papers (188C90). His "varieties" of the gizzard in vi rather than v (corrected in S. macleayi are perhaps part of a species com- Fletcher 1887a: 399) and his mistaking the plex, but, as with S. macleayi proper, none seminal vesicles for testes (corrected in have subsequently been "re-discovered". A Fletcher 1887b: 602603). Fletcher (1889) footnote by Barley and Kleinig (1964) makes differentiated his dozen specimens of a passing reference, without further detail, to Spenceriella macleayi from Sydney from those their having collected Megascolex macleayi of S. awtralis from Burrawang, NSW, by their (Fletcher) along watercourses near Deniliquin, smaller size, the presence of genital markings NSW. The above account, then, probably and possession of only two pairs of calciferous represents the first confirmation of this pouches rather than three pairs (in x to xii). species since it was designated over 100 years ago. Fletcher (1890: 1003407) went on to describe three "varieties" of S. macleayi which Spenceriella, of which S. macleayi is a he noted separately from each locality. representative example, will probably become Varieties a and b (from the Blue Mountains, one of the most numerous native genera by NSW, and Burrawang, NSW, respectively) revision of the heterogeneous congeries of differed from the "typical form", not only by Perichaeta and Megascolex (see Jamieson and having 20-60 more segments, but also in (1) Wampler 1979). It is also likely to receive the number and character of the genital mark- many new species from further collections. ings (2) the position of the first dorsal pore Sharing several characteristics with the allied Australian Zoologist, Vol. 29(34) 253 genus +colex Edmonds and Jamieson 1973 Beddard, F. E., 1895. A mgraph o/the &dn Ollgochela. Oxford: Clarendon Press. (see Jamieson 1974), it is partially differentiated Edmonds, S. J. and Jamieson, B. G. M.. 1973. A new genus by having segmental rather than intersegmental and species of earthworm (Megascolecidae: genital markings (the unpaired female pore Oligochaera) from South Australia. Tranr. R. Soc. S. and lack of calciferous glands of Gemascolex Au1. 97: 23-27. are occasionally met in Spenceriella). Fletcher. J. J.. 1886. Notes on Australian earthworms. Part 1. Proc Linn Soc. NSW l(2): 523-76. The current study demonstrates that S. macleayi persists despite cultivation and intro- Fletcher, J. J., 1887a. Notes on Australian earthworms. Pan 111. Prac. Linn. Sol. NSW. 2(2): 37742. duction of exotic species that followed Fletcher, J. J., 1887h. Notes on Australian earthworms. Part European settlement. It was found in close 1V. Proc. Linn. Soc NSW. 2(2): 601-20. association with Apmectodea trapendes suggest- Fletcher, J. J., 1889. Notes on Australian earthworms. Pan ing that this exotic lumbricid is not displacing V. Pror. tin". Soc. NSW. 3(2): 1521-558. the native species. Interestingly, this was also Fletcher, J. J., 1890. Nates on Australian earthworms. Part the case in Fletcher's original accounts (e.g., V1. Proc. tinn Sac. NSW. 4(2): 987-1019. 1886, 1887a) where he recorded the presence Jamieson, B. G. M., 1972. The Australian earthworm genus Spenceriella and description of two new genera of both native and exotic groups, with A. (Megascolecidae: Oligochaeta). Mm. Natl. Mw., Vic. 33: lrapezoides from almost every locality from 73-68. which he obtained earthworms. Fletcher Jamieson, B. G. M., 1974. Earthworms (Megascolecidae: (1886) noted that the Hon. W. Macleay's Oligochaeta) from South Australia. Tronr. R. Soc. S. Awl. 98(2): 79-1 12. garden at Elizabeth Bay, whence he procured Jamieson, 8. G. M. and Wampler, J. E., 1979. Bio- S. macleayi as well as several exotic species, was luminescenr Australian Earthworms 11. Taxonomy and then one of the oldest established gardens in preliminary repon of bioluminescence in the genera Australia to which plants had been brought Spenc~No, F~lchcrodn'Iu( and Ponrodnlw (Mega- from many parts of the world. scolecidae: Oligochaeta). Aurt. j. Zwl. 27: 63769. Michaelsen, W., 1900. Dm TIm&h Vol 10: Oligochelo. ACKNOWLEDGEMENT FriedlPnder & Sohn: Berlin. Michaelsen, W., 1907. Oligochaeta in Die Fauna Siidwest- Dr Pat Hutchings is thanked for refereeing Ausrraliens. l(2): 117-232. Jena: Gusrav Fischer. the original manuscript. Reynolds. J. W. and Cook, D. G., 1976. Nmclalvra Oligochoerologica: a Calalogue of Names, Dwc"p1ias and REFERENCES Type Specim~lr of the Oligmhekz. University of New Brunswick: Frederiction. Pp. 217. Barley, K. P. and Kleinig, C. R., 1964. The occupation of newly inigated lands by earthworms. Awl. j. Sci. 26: Spencer, W. B., 1900. Further de-ptions of Australian 29&91. earthworms. PTM.R . SOC.V iCI. 13(l): 29-67. 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