Tuatara: Volume 24, Issue 1, October 1979
On Two Land-Leeches Labelled as from New Zealand (Hirudinea: Haemadipsoidea)
On Two Land-Leeches Labelled as from New Zealand (Hirudinea: Haemadipsoidea)
Two specimens collected by the U.S. Exploring Expedition, 1832-1842, are assignable to the Chtonobdellinae (Domanib-dellidae). They differ in dorsal pattern from chtonobdellines in eastern Australia. The original labelling of them as from New Zealand is not provably incorrect on zoological grounds or other evidence.
Mason (1976) notes that New Zealand is without representatives of the land-leeches.
Moore (1898) listed without detail two land-leeches in the United States National Museum. He recorded these as ‘Geobdella limbata (Grube Whitman)’, and as collected in the course of the U.S. Exploring Expedition, 1838 to 1842. Each is associated with a label giving the origin as ‘New Zealand’.
These are the earliest known land-leeches collected in the Australian Region. They have not been referred to other than by Moore.
In reply to a letter from Benham (see Benham, 1904, p. 185, footnote p. 186), Moore wrote that he re-examined the specimens and confirmed his identification ‘so far as one can be certain from Grube's description and figures alone’, considered the locality as doubtful, concluded that the specimens probably came from Sydney, ‘one of the stopping places of the Expedition’, and that he regarded them as ‘valueless in establishing the occurrence of the species in New Zealand’.
Moore (1944) notes: ‘The two specimens attributed to New Zealand, U.S. Exploring Expedition, referred to by me (1898), in the collections of the U.S. National Museum, must be labelled incorrectly. as the Wilke's Expedition did not visit New Zealand, and this species has not been reported therefrom.’
Moore was in error on one point.
The narrative of the Expedition (Wilkes, 1845) shows that the vessels of the Expedition departed Sydney late in December 1839, leaving the ‘Scientific Corp’ in Sydney with instructions to ‘join the page 42 squadron on the return of the Expedition, from the Antarctic Ocean, at the Bay of Islands, by the 1st of March next.’
The Scientific Corp arrived at the Bay of Islands on the 24th of February. It is noted: ‘They had been forced to remain inactive at Sydney, in consequence of a change in the destination of the vessel in which they had first taken their passage and … been prevented from pursuing further researches in New South Wales …’ The vessels on the Antarctic cruise reached the Bay of Islands on March 30th, 1840.
This paper describes such detail as can be taken from the U.S.N.M. specimens; compares this with the results from the study of Grube's Type and Paratype specimens from the vicinity of Sydney, N.S.W., preserved for nearly the same length of time as the U.S.N.M. specimens; and discusses the origin of the U.S.N.M. specimens.
Description of Specimens
U.S. Exploring Expedition Specimens
U.S.N.M. 174. Geobdella limbata (Grube) New Zealand? (prob. Australia). U.S. Exploring Expedition (Wilkes Exped.) 1853. Ident. J. Percy Moore, Acc. No. 259718.’ With the specimen, a printed number, 174, and in hand-writing: ‘New Zealand. Ed. Ex.’
One specimen approximately 28.0mm long, in good condition. The body is elongate cylindrical, curving ventrally posteriorly: the anterior sucker everted, thick-rimmed.
The dorsum and venter, and dorsum of the posterior sucker, pale brown; the venter of the body darker than the dorsum.
A clear indication on the left of a narrow pale white contrast stripe on xxiii b2 anteriorly into xx: the nephropores median to dorsal in the stripe, i.e. the stripe in the upper portion of the marginal field, and with no indication of dorsal lobes on the nephroporic annuli. The stripe detectable anteriorly into xvi, and on xiv to ix.
There are no other indications of pattern.
Interannular and intersomital furrows equivalent, well-defined; somital limits not directly recognisable; annuli areolate; dorsal somital sense organs in large areolae, the dorsal paramedians, intermediates, and supramarginals prominent in xii to xxiii; nephropores obvious on b2 in ix to xxiii.
Somites i to iv obscured on the rim of the sucker; v incomplete 2-annulate; vi complete 3-annulate; a1 = a2 with the 5th eyes much < a3; vi a3 < vii a1; vii 3-annulate, a1 = a2 < a3; viii 4-annulate, a1 slightly < a2 = b5 slightly < b6; ix to xxiii 5-annulate (total 15) with a2 a distinctly long annulus, and the paramedian and intermediate somital sense organs unusually large on xvi to xxiii; x to xvi b1 = b2 < a2 > b5 = b6; xvii to xxii b1 = b2 much < a2 much > b5 > b6 (b5 possibly longer than b6 because of the ventral curvature of the body); xxiii b1 = b2 with the 16th nephropores much < a2 > b5 > b6; xxiv 2-annulate, no somital sense organs on a1 a2 = a3, with a3 shortened across the venter; xxv, xxvi, xxvii, uniannulate, fusing to form the white well-developed auricles, each divided into two lobes by a narrow indentation, the ventral lobe continuing ventrally as a thin ridge.
Genital pores, xi b5 xii b1/b2.
U.S.N.M. 37071. Geobdella sp. New Zealand J. P. Moore Donor.page 43
Acc. No. 259718. Original labels: (a) “New Zealand. Ed. Ex.” — the same script as with No. 174; (b) Geobdella, in Moore's script.’
One specimen, 31.0mm long; partially engorged to be elongated cylindrical, tapering anteriorly. The anterior somites opened by a midventral longitudinal incision, exposing the ventrolateral jaws, i.e. duognathous.
Dark greyish brown above, darker below. There is no indication of pattern.
Annuli flat; interannular furrows distinct from viii to xxvii; no detectable somital sense organs; nephropores distinct in xi to xviii, xx to xxiii, each with a white rim.
Somites i to vii not assessable; viii 4-annulate; ix to xxiii 5-annulate, a2 distinctly a long annulus in xiii to xxiii; ix, x relative lengths not assessable; xi b1 = b2 < a2 = b5 > b6; xii b1 < b2 = a2 = b5 > b6; xiii b1 < b2 < a2 > b5, b6; xiv to xxiii b1 = b2 < a2 > b5 = b6; xxiv a1 a2 slightly > a3; xxv, xxvi, xxvii, auricles, as in specimen 174.
Genital pores, xi b5/b6; xiii b1/b2.
Type and Paratype, Chtonobdella limbata Grube 1866.
The Type: ‘Zool. Mus. Berlin: Kat. nr. 1373. Sydney. Coll. Grube. Typen.’
Total length 25.5mm; widest and deepest posteriorly, tapering gradually anteriorly to the subcylindrical pregenital region. Recognisable as the specimen figured by Grube, pl. 4. fig. 7.
Faded, now almost a uniform pale ashen grey above and below anterior to xxiii a2, pale off-white behind this.
Faintly indicated and recognisable as: (a) a pale narrow short stripe lateral in each paramedian field from in xx b5 to in xxiii b2; (b) an elongate narrow light patch median in each paramedian field extending from xxii b6 on to xxiii b1; (c) a similar but fainter patch lateral to each short stripe on xxii b6 on to xxiii b1; (d) similar paired patches in line with (b) on xxiii b5 and b6; (e) a light stripe in the marginal fields from viii/ix on the left to the auricle, and on the right from ix b2 to xxiii a2, then very faintly indicated to the auricle, the stripes including the nephropores and with indications of dorsal lobes on the nephroporic annuli. Grube shows only (a) and (e).
Strongly contracted, the intersomital and interannular furrows deep, equivalent. Somites i to iii uniannulate; the eyes, each in a large ocular areola, the ocular areolae in ii separated by two pairs of areolae in tandem, in iii by 3 areolae; iv 2-annulate between the oculars; v 2-annulate above, a1 a2 > a3, a1 a2 / a3 extending to the ventral intermediate lines and v uniannulate between these; vi complete 3-annulate a1 > a2 > a3; vii 3-annulate a1 < a2 < a3; viii 4-annulate a1 > a2 = b5 > b6; ix to xxiii 5-annulate (total 15); ix b1 < b2 with the 2nd nephropores < a2 = b5 > b6; x b1 = b2 < a2 > b5 > b6; xi to xxi b1 < b2 < a2 > b5 > b6, the difference between the lengths of b1 and b2, of b5 and b6, are small, a2 is distinctly long; xxii b1 = b2 < a2 > b5 = b6; xxiii b1 = b2 < a2 > b5 > b6, the 16th nephropores on b2; xxiv 2-annulate a1 a2 = a3, no somital sense organs on a1 a2, a3 the last annulus formed on the venter; xxv, xxvi, xxvii, uniannulate, with distinct paramedian sense organs, the lateral ends of the three annuli forming the bilobed auricles, divided by an open arch with a broadly conical median element.
Paratype: with the Type.
Total length 32.0mm; the margin of the anterior sucker turned back dorsally; page 44 the pregenital region depressed oval; the body progressively depressed along the genital and postgenital regions.
The pregenital region pale light brown above, paler to light grey below, darkening posteriorly to xxiii a2 as dark brown above, paler below; then almost off-white to xxvii; dorsum and venter of the posterior sucker, darkish grey.
More distinct than in the Type: (a) a pale narrow short stripe lateral in each paramedian field from xx/xxi posteriorly into xxiii a2; (b) an elongate narrow patch median in each paramedian field from xxii b6 on to xxiii b1; (c) on the right side, a patch in the intermediate field lateral to the short stripe, on xxii b6 and xxiii b1, and lateral to this patch a similar patch on the same annuli, on the left, a single large patch equivalent to the right pair fused; (d) in each paramedian field, a patch in line with (b) on xxiii b5 and b6; (e) a light stripe in each marginal field from x xi on the right, xii/xiii on the left, to the auricles, the stripes including the nephropores, some in a distinct dorsal lobe on the stripe.
As in the Type, excepting vi 3-annulate above. 2-annulate below.
Chtonobdella limbata Grube 1866 is based on the two specimens collected at Sydney during the Novara Expedition, 1857-1859.
There was available to Moore (1898): (a) Grube's account (1868) of the Novara specimens, describing colour, pattern which is in general terms and not recognisably defined, the annulation showing them to have 5-annulate somites in the middle series with 7 annuli between the genital pores; and (b) the brief note in Whitman (1886, p. 322) that a specimen sent by Haswell was duognathous, and proposing Geobdella as a generic name. At that time, no other land-leech was known with these features.
The U.S.N.M. specimens agreeing with the details given by Grube and Whitman, Moore identified them as Geobdella limbata. Blanchard (1917) assigned Australian land-leeches with 5-annulate somites in the middle series and 7 annuli between the genital pores to Haemadipsa limbata (Grube, 1866). The g. Haemadipsa is trignathous.
The characteristics of limbata as used by Moore and Blanchard, i.e. a general somital annulation of vii 3-annulate, viii 4-annulate, ix to xxiii 5-annulate, xxiv 2-annulate, the genital pores posterior in xi and anterior in xiii, are present in species in the Chtonobdellinae in genera other than Chtonobdella (Richardson, 1975).
Land-leeches from southern Queensland recognised by Blanchard as ‘H. limbata’ are Jaabdella whitmani (Lambert, 1899); others from northern New South Wales, Quaesitobdella bilineata Richardson 1975, as also specimens from the same area assigned to C. limbata by Moore (1944), v. Richardson (1974, 1975).
C. limbata is known now as restricted to central New South Wales cast of the Dividing Range, and south from Sydney.
The U.S.N.M. specimens are assignable to the Chtonobdellinae.
Preserved, the general colour is much the same in Grube's Type page 45 and Paratype as in the U.S.N.M. specimens, excepting the latter are slightly darker.
As described above, the Type and Paratype of C. limbata show a characteristic topographically defined dorsal contrast pattern restricted to the posterior body somites. This is unique in the Chtonobdellinae, others in the subfamily having dorsal contrast stripes in or anterior to ix, extending posteriorly into xxiii or further.
The pattern in limbata (Richardson, 1974), is a dark broad band within the ocular arch posteriorly to xx; the band divided on each side in the posterior annuli of xx to in the anterior annuli of xxiii, by a narrow contrast stripe, fragmented in some few specimens. On both sides of each stripe, elongate contrast patches as transverse and longitudinal rows, the patches centred on xxi/xxii and xxii/xxiii, with another transverse row posterior in xxiii; a contrast stripe in each marginal field from the anterior sucker to the auricle, lobed dorsally on each nephroporic annulus, the nephropores in the lobes; the venter, a dark band between the marginal stripes.
Excepting engorged specimens, the dorsal pattern has been recognisable in detail in other specimens of limbata preserved in the latter part of the last century.
There is no indication of contrast stripes or patches anywhere on the dorsum of the U.S.N.M. specimens or of dorsal lobes on the marginal stripes. I have not seen an adult Australian chtonobdelline from the Sydney area or elsewhere with an undivided, patchless dorsal pattern as in No. 174.
The U.S.N.M. specimens are assessable as a species distinct from C. limbata.
A relationship between the freshwater dependent leech faunas in New Zealand and eastern Australia is demonstrable in the aquatic hirudiniform leeches and the barbronids. The present systematisation of the freshwater rhynchobdellids is zoologically inadequate, such that the species described from these countries are not usable for zoogeographic purposes.
The Richardsonianidae, a family of aquatic jawed sanguivores in Australia, Tasmania, Lord Howe Island, with a torresian genus in Queensland, the Northern Territory, New Guinea, and the Kei Islands, is represented in New Zealand by ‘Richardsonianus’ mauianus (Benham, 1907). The Ornithobdellidae, based on Ornithobdella edentula Benham 1909 of the Snares Islands, with a second genus and two species in New Zealand, is represented in eastern Australia from Victoria to Queensland by the g. Aetheobdella Moore 1935 (Richardson, 1969; Mason, 1976).
Mason (1976) assigns a barbronid in New Zealand to Barbronia weberi (Blanchard, 1897) as in Moore (1927). Moore describes weberi, and later others in the Oriental Region, as having a higher level of annulation on the somites of the middle series than as described by Mason, and also the median region of the male system page 46 as recognisably myomeric mesomorphic, not amyomeric micromorphic as shown by Mason, Fig. 29.
Mason's barbronid has the characteristics as known to me of barbronids of eastern Australia, Tasmania, the New Guinea Archipelago, New Caledonia, and Palmyra Island in the Outlying Group of the Pacific Islands, i.e. the New Zealand barbronid belongs to an Australian/Oceanian group, and not an Oriental group (Richardson, 1971).
The land-leeches are assembled in the Haemadipsoidea. The distribution of the Superfamily includes many and some wide oceanic separations. There are discrete faunas: Idiobdellidae, Seychelle Islands; Haemadipsidae s.s., the Orinetal Region, with one species in eastern Wallacea; the Domanibdellidae: Domanibdellinae, New Guinea Archipelago, with extensions across Wallacea into the Oriental Region, the Australian Northern Territory, New Caledonia, Fiji, Palmyra Island; Malagabdellinae, Madagascar; Leiobdellinae, New Guinea Archipelago, Cape York Peninsula, Samoa; Chtonobdellinae, 5-annulates, and Philaemoninae, 4-annulates, of continental Australia east of the Dividing Range from North Queensland to central New South Wales, with the Philaemoninae continuing into Tasmania, and on Lord Howe Island (Richardson, 1975; 1978).
As above, many land-leeches are tropical, but not all. The Chtonobdellinae in eastern Australia are tropical, subtropical, to cool temperate. In the subtropical and cool temperate, they are present from sea-level to altitudes above 1700m, including areas snow covered for a week or longer in some winters. The Bay of Islands is central in the eastern edge of an area with warm humid summers and an annual rainfall in excess of 200mm. The area cannot be assessed climatically as unsuitable for land-leeches. It should be noted that the Narrative records Dr William Pickering of the Expedition as having crossed this area from the Bay of Islands to Hokianga, and return.
The typical land-leech habitat is a forested area or forest fringe. The gently sloping floor provides a rapid run-off during rain, and is free from standing water. Above this area, there is a catchment area providing seepage maintaining soil moisture in the habitat area over extended periods. Persisting small pools and/or moss are common below the habitat area.
Land-leeches are so highly selective in habitat that a species is discontinuous within its distribution. Habitat areas are commonly small, some no more than 30m by 2m, and may be separated from other land-leech habitats by up to 20km to 50km. Land-leeches are active when ground moisture is suitable, inactive at other times. There is a tendency to be seasonal, but they may be inactive for long periods during the suitable season (Moore, 1932; Richardson, 1968; Bhatia and Bora, 1973).
For these and other reasons, land-leeches can be difficult to find. The one small specimen known for Fiji was included in a collection page 47 of planarians. Only two very small specimens are known from the Samoan Islands, as also Réunion. Eucryptobdella parva (Moore, 1944), the New Hebrides, is based on one small specimen. Originally, only four specimens were known for the Seychelles. My correspondents in Fiji and the New Hebrides do not know of land-leeches in these islands.
|(a)||The U.S.N.M. specimens are assignable to the Chtonobdellinae.|
|(b)||The distinctive dorsai pattern of C. limbata remains fully recognisable in the Type and Paratype specimens collected at Sydney between 1857 and 1859.|
|(c)||The U.S.N.M. specimens show no trace of the dorsal pattern of C. limbata.|
|(d)||The dorsal pattern in U.S.N.M. No. 174, a wide intact dark band, is not known in adult chtonobdellines from the vicinity of Sydney, or elsewhere in eastern continental Australia.|
|(e)||The examples of the Richardsonianidae, the Ornithobdellidae, and barbronids, show a relationship between the freshwater dependent leech faunas of New Zealand and eastern Australia.|
|(f)||The distribution of the Domanibdellidae and of the barbronids is indicative of an antiquity as great as or greater than the antiquity of the Richardsonianidae and Ornithodbdellidae.|
|(g)||The presence of a chtonobdelline land-leech in New Zealand is in agreement with the antiquity of the Domanibdellidae and the demonstrable relationship of the freshwater dependent leech faunas in New Zealand and Australia.|
|(h)||The locality, New Zealand, as given on the original labels with the U.S.N.M. specimens, is not provably incorrect on zoological or other grounds.|
I express my thanks to Dr A. Soos, the Hungarian Natural History Museum, who arranged with Dr G. Hartwich, Museum für Naturkunde, to make available to me the Type and Paratype of Chtonobdetla limbata; to Dr Marian H. Pettibone, the Smithsonian Institution, for the privilege of studying the two specimens collected by the U.S. Exploring Expendition; and to Dr J. C. Yaldwyn, the National Museum, Wellington, for a copy of the relevant portions of the Narrative of the Expedition.
This study is collateral to researches on the zoology of the Australian freshwater dependent leeches, supported by awards from the Australian Research Grants Committee.
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