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Volume 80, 1952
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Studies on the Earthworm Fauna of New Zealand. III

Department of Scientific and Industrial Research, Wellington

[Read before the Wellington Branch, May 30, 1951; received by the Editor, May 30, 1951]

Summary

This paper describes the New Zealand species of the Subfamily Megascolecinae, and includes a general discussion on their phylogeny and classification. The following new species are described and figured. Megascolides viridis, Motuhora: M. ruber, Mangamuku Range; M. parvus, Hicks Bay; M. raglani, Raglan Range; M. fuscus, Rotorua district; M. irregularis, Tangowahine; M. alba, Maungakaramea; Megascolex novaezealandiae, Mangonui; Pheretima campestris, Kaitaia; Diporochaeta obtusa, Rotorua; Perionyx egmonti, Mount Egmont.

The Sub-family Megascolecinae

The sub-family Megascolecinae was established by Michaelsen (1900) as a division of the family Megascolecidae. As defined by Stephenson (1930) it includes all those species of earthworms referable to the family Megascolecidae and having the following characters:

“Chaetae either eight per segment, or numerous, and then either in regular chains or approximated in couples. Clitellum beginning with or in front of xiv. Male pores on xviii. Spermathecal pores, if present, one to seven pairs, in front of the testis segments. Usually one gizzard in front of the testis segments, sometimes two or three, exceptionally none. Mega- or micronephridial. Two pairs of testes and funnels in x and xi, or only one pair. Prostates one pair, tubular or racemose (Pheretima—prostates), each prostatic duct uniting with the vas deferens of the same side and opening in common with it (except in Diplotrema).”

The occurrence of the sub-family Megascolecinae in New Zealand raises some interesting zoogeographical considerations. The “root genus” of the sub-family is taken as Diplotrema, which is found in Queensland and New Caledonia, and from Diplotrema all the other genera have been derived. The genera of the sub-family are now found for the most part in the Indian, Malayan, Oriental and Australasian regions and it is reasonable to assume that any general migration of Megascolecinae into New Zealand must have come from the north. A northern origin for the group is also indicated by its distribution within New Zealand, since nearly all the species are found only in the northern regions of the North Island. The entry of the Megascolecinae into New Zealand would require the postulation of a former land connection of the northern portion of the North Island either directly to Australia or to New Caledonia and the Indo-Malayan islands. Such a land connection need not have been continuous at any one time; it would be sufficient that at times portions of the land connection should have been emergent while other portions remained submerged, and the Megascolecinae were able to progress in stages until they eventually reached New Zealand.

Thirty species, belonging to ten genera of the sub-family, have been identified from the New Zealand region by previous workers and eleven new species, belonging to five of the ten genera, are described in this paper. The following is a list of the ten genera known to occur in the New Zealand region, with notes on their distribution within the region:

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Plutellus Perrier (1873) Auckland Islands.
Pontodrilus Perrier (1874) Lake Wakatipu, South Island; Chatham Island.
Megascolides McCoy (1878) northern North Island.
Notoscolex Fletcher (1886) northern North Island (possibly introduced); Poor Knights Island.
Diporochaeta Beddard (1890) North Island; western South Island; Chatham Island.
Spenceriella Michaelsen (1907) northern North Island; Little Barrier Island.
Megascolex Templeton (1844) northern North Island; Norfolk Island.
Pheretima Kinberg (1866) northern and eastern North Island; Raoul Island (probably introduced).
Perionyx Perrier (1872) western North Island; Auckland Islands; Snares Island.
Didymogaster Fletcher (1886) distribution unknown; a single species, probably introduced.

None of these genera is peculiar to the New Zealand region. The genera may be identified from the following key:

a1 Eight chaetae on each segment
b1 Purely meganephridial
  c1 Gizzard vestigial or absent Pontodrilus
  c2 Gizzard well developed Plutellus
b2 Micronephridial, at least in part
  d1 Prostate tubular, with single continuous central duct Megascolides
  d2 Prostate racemose, with branching system of ducts
    e1 Single gizzard in v or vi Notoscolex
    e2 Two gizzards, one in vi and one in vii Didymogaster
a2 More than eight chaetae on each segment
f1 Meganephridial
  g1 Prostate tubular, with single continuous central duct Diporochaeta
  g2 Prostate racemose, with branching system of ducts Perionyx
f2 Micronephridial
  h1 Gizzard in v, vi or vii
    i1 Prostate tubular, with single continuous central duct Spenceriella
    i2 Prostate racemose, with branching system of ducts Megascolex
  h2 Gizzard between septa vii/viii and x/xi Pheretima

The relationships of the New Zealand genera are shown in the diagram below, commencing from the “root genus” Diplotrema:

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Genus Plutellus Perrier

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

“Chaetae eight per segment. Male pores paired or single; female pores mostly paired; spermathecal pores end at furrow 8/9 or on segment ix, a single pair or a series of two to five pairs, or five single pores. A gizzard in the region of segments v–vii. Purely meganephridial. Prostates tubular, with simple unbranched duct.” (Stephenson, 1930.)

A single species, P. aucklandicus Benham (1909), is recorded from the New Zealand region. The species was confirmed by Michaelsen (1923), who noted that short side ducts open into the prostatic duct and consequently the species would perhaps go into the genus Woodwardiella. However, Michaelsen decided that the species should remain in Plutellus, since the side ducts are small and the central duct is of more importance (cf. New Zealand species of Megascolides, discussed below).

Genus Pontodrilus Perrier

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

“Chaetae eight per segment. Male pores paired; female pores paired. Spermathecal pores two to four pairs, the last in furrow 8/9. Gizzard vestigial or absent. Purely meganephridial, nephridia wanting in front of the clitellar region. Two pairs of free testes and funnels. Prostates tubular, with simple unbranched canal.” (Stephenson, 1930.)

The most remarkable feature of this genus is that of the five known species, three have a littoral habitat and live in the intertidal zone, while the fourth is limnic and the fifth terrestrial. This is the only known genus in which Megascolecid worms have a littoral habitat.

Two species are known from the New Zealand region. They are P. matsushimensis var. chathamensis Michaelsen, collected from the shore at Chatham Island, and P. laustris Benham, the only known freshwater species, collected at depths of 300–1,200 feet from the waters of Lake Wakatipu in the western South Island. Benham (1900) raised the Chatham Island form to a species, P. chathamensis, but Stephenson (1930) considers it as a variety of P. matsushimensis Michaelsen.

Genus Megascolides McCoy

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

“Chaetae eight per segment. Spermathecal pores one to five pairs, the last in furrow 7/8 or 8/9 or on segment ix. One gizzard in the region of segments v and vi. Micronephridial at least in the anterior part of the body, often throughout. Prostates tubular, with a simple unbranched canal.” (Stephenson, 1930.)

The species here included in Megascolides have been referred by various authors to Notoscolex, Tokea (a genus which has now been discarded), Megascolides, and in one case (M. morlensem) to Megascolex. The differences between Megascolides and Notoscolex are slight and for diagnostic purposes the form of the prostatic duct is relied upon. A typical Megascolides has simple tubular prostates, each with an unbranched central canal running right through the gland and opening directly to the exterior through the prostatic duct, while a typical Notoscolex has polylobate prostates, each with a diffuse system of ducts ramifying through it and frequently opening into a cavity which is connected with the external pore by the prostatic duct. If all the species of Megascolides and Notoscolex had one or other of these “typical” arrangements of the prostatic ducts, there would be no difficulty in placing a species in its correct genus, but there is a great deal of variation and the two types of prostates grade into one another. It would seem desirable to fuse the two genera, but this would produce a genus

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with a very large number of species, and for convenience it is better that the two genera should be retained.

Benham (1904) established a genus, Tokea, for several species from the North Island of New Zealand which had all the characters of Megascolides, but differed from that genus in one respect. The prostate had a simple central canal, but when sections of the gland were examined, it was found that the gland cells were grouped into small clusters which discharged their secretion into a slight evagination of the central duct. Benham called these small evaginations of the central duct “canalicules”. It was unfortunate that he chose this term, since it really means a small canal and was taken as such by other workers, who placed Benham's species in Notoscolex. However, reference to the figures in Miss Sweet's paper on the structure of prostatic glands (1900), or to Benham's paper (1941) on Megascolides napierensis, in which he figures a cross-section of the prostate of Megascolides (Tokea) esculenta and details of the “canalicules”, will show what Benham meant by the term. I have examined sections of the prostatic gland of other genera of the family Megascolecidae and have found that these “canalicules” are present in the prostates of other genera which have tubular prostates, e.g. in Neodrilus they are present in large numbers. The position of Tokea in relation to Megascolides and Notoscolex has been a subject of much discussion and confusion. Stephenson (1930) summed up the position as it appeared to him, but his statements are in many respects contradictory. He believes that Tokea should go into Notoscolex, a view also adopted by Michaelsen, yet in one part of his monograph (p. 658) he refers to the edibility of certain species of Tokea (Megascolides).” There is one character of Tokea which all the previous workers seem to have overlooked in their discussion. That character is the presence of meganephridia in association with the micronephridia in the posterior segments of the body (see Benham's original description of the genus, 1904). Now, according to the accepted relationships of the genera of the Megascolecinae (as set out above), Megascolides is derived from Plutellus by a change from the meganephridial to the micronephridial condition. In many cases the change is not complete and meganephridia are found in association with micronephridia in the posterior segments of Megascolides. Notoscolex is derived from Megascolides by the branching of the prostatic duct and there are few records of species of Notoscolex with meganephridia and micronephridia co-existent in any of the segments. In view of this fact and the questionable existence of branches in the central prostatic canal of Tokea there can be little doubt that Tokea should be included in Megascolides.

Megascolides mortenseni was referred by Michaelsen (1923) to the genus Megascolex, but reference to the original description shows that it is actually a species of Megascolides. M. reptans and M. unipapillatus were referred by Ude (1905) to Notoscolex, but reference to his descriptions shows that they belong to Megascolides, since they have unbranched tubular prostates.

Twelve species of Megascolides have been recorded from the New Zealand region by previous workers and the following is a list of them.

  • M. napierensis Benham (1941).

  • M. (Tokea) maorica (Benham, 1904).

  • M. (Tokea) decipiens (Benham, 1905).

  • M. (Tokea) kirki (Benham, 1904).

  • M. (Tokea) huttoni (Benham, 1904).

  • M. (Tokea) urewerae (Benham, 1904).

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  • M. (Tokea) suteri (Benham, 1904).

  • M. (Tokea) sapida (Benham, 1904).

  • M. (Tokea) esculenta (Benham, 1904).

  • M. (Notoscolex) reptans (Ude, 1905).

  • M. (Notoscolex) unipapillatus (Ude, 1905).

  • M. (Megascolex) mortenseni (Michaelsen, 1923).

Seven new species, M. viridis, M. ruber, M. parvus, M. raglani, M. fuscus, M. irregularis and M. alba are described in this paper.

The two species M. maorica and M. decipiens are distinguished by the number and position of the tubercula pubertatis. I have found these structures to be extremely variable in number and position and consequently the distinction cannot be upheld. and M. decipiens must be considered synonymous with M. maorica.

Three species, M. huttoni, M. urewerae, and M. suteri, cannot be separated according to the descriptions given by Benham. I have discovered two species which agree fairly closely with Benham's descriptions of M. urewarae and M. suteri, and further notes on the two species are included in this paper. I have not found any species which can be referred to M. huttoni, although M. parvus n.sp., described in this paper, is very similar to M. huttoni. M. napierensis was founded on a single specimen collected from a nurseryman's glasshouse at Napier. I have collected many hundreds of specimens of endemic earthworms from the district surrounding Napier and have not found M. napierensis or any species of any other Megascolecine genus among them. I conclude that M. napierensis is to be regarded as introduced into that one glasshouse in Napier with soil or plants and that it may have come from any part of the world where Megascolides is known. M. mortenseni (Michaelsen) was founded on a single incomplete specimen from Palmerston North. No details of its environment are given, so it cannot be definitely decided whether it is introduced or endemic. It has not been recorded again from Palmerston North or from the surrounding district and it is possible that it was, like M. napierensis, an isolated introduced specimen.

The species of Megascolides known from New Zealand may be identified from the following key:

a1 Two pairs of hearts, in xii and xiii M. napierensis
a2 Three pairs of hearts, in x, xi and xii
b1 Two pairs of spermathecae, in viii and ix
  c1 Calciferous glands absent
    d1 Three pairs of vesiculae seminales, in x, xi and xii M. maorica
    d2 Four pairs of vesiculae seminales, in ix, x, xi and xii M. viridis
  c2 One pair of calciferous glands, in xiv M. kirki
  c1 Two pairs of calciferous glands, in xii and xiii M. ruber
  c4 Five pairs of calciferous glands
    e1 Four pairs of vesiculae seminales, in ix, x, xi and xii M. reptans
    e2 One pair of vesiculae seminales, in xii M. unipapillatus
b2 Three pairs of spermathecae, in vii, viii and ix*
  f1 Two pairs of vesiculae seminales. in ix and xii
    g1 Gizzard in v M. parvus
    g2 Gizzard in vi M. suteri
  f2 Four pairs of vesiculae semmales, in ix, x, xi and xii M. urewerae
a2 Four pairs of hearts

[Footnote] * M. huttoni cannot be definitely separated from any species under this heading, due to inadequacy of description.

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h1 Hearts in ix, x, xi and xii M. raglani
h2 Hearts in x, xi, xii and xiii
  i1 One pair of spermathecae M. fuscus
  i2 Two pairs of spermathecae M. irregularis
  i3 Three pairs of spermathecae
    J1 Spermatheca with elongate tubular sac and slender, pyriform diverticulum M. mortenseni
    J2 Spermatheca with stout ovoidal sac and ovoidal or globular diverticulum
      k1 No oesophageal glands M. sapida
      k2 Oesophageal glands as lateral dilations of the oesophagus in xv M. esculenta
a4 Five pairs of hearts, in ix, x, xi, xii and xiii M. alba

The following are descriptions of the seven new species M. viridis, M. ruber, M. parvus, M. raglani, M. fuscus, M. irregularis and M. alba. Types in my collection.

Megascolides viridis n.sp. (Plate 10, figs. 1–3)

Several specimens of this small species were collected from native forest on a mountainside seven miles west of Motuhora, inland from Gisborne. The most striking character of the species is its colour. Dorsally it is reddish-brown and ventrally it is pale bluish-green. The chaetae arise each from the centre of a small raised white spot. The clitellum does not differ in colour from the remainder of the body, but completely surrounds xiv to xvii, obliterating the intersegmental furrows between those segments.

The specimen on which this description is based is 54 mm. in length and 4.5 mm. in diameter and has 66 segments.

The prostomium is prolobous. There are eight chaetae on each segment, arranged in pairs. On xxiv the arrangement is as follows: ab = 1.5 mm.; cd = 2 mm.; bc = 2 mm.; aa = 2.25 mm., dd = 3 mm.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, a pore in line with chaeta a of each side in each intersegmental furrow. The female pores are on xiv, a pore on each side anterior and slightly medial to chaeta a. The male pores open to the exterior on xviii, one on each side, slightly lateral to the line joining the chaetae a of the adjacent segments. Each pore is on the apex of a small, rounded, white papilla. There are two small, oval, unpaired, median ventral tubercula pubertatis, one at 13/14, and the other at 18/19. No nephridiopores nor dorsal pores are visible.

Internal Anatomy (Plate 10, fig. 2)

The septa vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are thickened and muscular.

Alimentary Canal. The pharynx is small and muscular and occupies the first four segments. There is no proventriculus. The gizzard is slender, but has thick muscular walls, and is situated in v. The oesophagus extends from vi to xiv and there are no oesophageal glands. The intestine commences in xv.

Vascular System. The dorsal blood vessel is unpaired throughout its length. A slender, unpaired, supra-intestinal vessel passes through x–xii and from it arise three pairs of dilated hearts, a pair in each of x, xi and xii.

Reproductive System. There are two pairs of small compact testes, a pair in x and a pair in xi, one on each side close to the ventral nerve cord in each segment. A pair of ovaries occurs in a similar position in xiii. Each ovary consists of a cluster of thread-like structures arising from the anterior septum of xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix.

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In the specimen examined both the spermathecae of ix appear to lie on the right hand side of the ocsophagus, although the ducts open one on each side of the segment, as is normal. The left spermatheca has probably been twisted under the oesophagus by the contortion of the earthworm's body when it was killed. Each spermatheca consists of an ovoidal sac, opening by a long, muscular duct to the exterior, and a pyriform diverticulum opening into the medial aspect of the spermathecal duct, close to distal end of the duct. (Plate 10, fig. 3.) The prostates are simple tubular organs with unbranched axial canals. They are of unequal extent, the left prostate commencing from the duct, which passes through xviii and xix, and extending back through xx and xxi into xxii, where it terminates in a dorsally directed portion, and the right arising from its duct in xviii and extending around the lateral surface of the intestine in xix to terminate on the left dorso-lateral surface of the intestine in xix. (See Plate 10, fig. 2.) There are four pairs of vesiculae seminales, a pair in each of ix x, xi and xii Those of ix and xii are much smaller than those of x and xi.

A broad band of small micronephridial tubules extends across the ventrolateral and lateral aspects of the body-wall in each segment.

Remarks. Although the two species are superficially distinct, M. viridis very closely resembles M. maorica in its internal anatomy. It can be distinguished from M. maorica by its possession of four pairs of vesiculae seminales (M. maorica has only three pairs).

Megascolides ruber n.sp. (Plate 11, figs. 1–3)

A single specimen of this dark red earthworm was collected from Awapuka clay near the summit of the Mangamuku Range, North Auckland, in a large area of native rain-forest. The posterior portion of the specimen is missing, so the length cannot be determined. The average diameter is 4·5 mm.

The prostomium is epilobous. The clitellum covers xiv to xviii and is lighter in colour than the rest of the body. The chaetae are lumbricine in arrangement, eight per segment. On segment xxii their arrangement is as follows:

ab = 1.25 mm.; cd = 2 mm.; bc = 1.5 mm.; aa = 1·5 mm.; dd = 3 mm.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with chaeta b. There is a single, unpaired, median, ventral female pore on xiv slightly anterior to the line of the chaetae on that segment. A single pair of prostatic pores occurs on xviii. Each of the prostatic pores is situated in the centre of a cuplike depression, with a raised tumid edge, in line with the chaetal interval ab on each side. Chaetae a and b are absent on xviii.

Internal Anatomy (Plate 11, fig. 2)

There are no specially thickened septa.

Alimentary Canal. The pharynx is slender and muscular and occupies i to iv. A small spheroidal gizzard lies in v. The oesophagus extends from vi to xv and bears two pairs of prominent calciferous glands, a pair in xii and a pair in xiii. The intestine commences in xvi and is very thin walled.

Vascular System. The dorsal blood vessel is unpaired. There are three pairs of dilated hearts, a pair each in x, xi and xii.

Reproductive System. There are two pairs of small lobate testes, a pair in x and a pair in xi. A single pair of ovaries is present in xiii and these organs are of a peculiar form which serves to distinguish the species from others of the same genus. Each ovary consists of a number of moniliform threads clustered

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together at their attachment to the anterior septum of xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each is in the form of a large, thin-walled, ovoidal sac with a small pyriform diverticulum opening into the anterior aspect of its duct. (Plate 11, fig. 3.) The prostates are simple, unbranched, tubular organs lying ventral to the intestine, as is typical of the New Zealand species of Megascolides (Tokea). Each prostate arises from a narrow muscular duct in xviii. The left prostate is coiled in the form of a reversed S, terminating in xx, and the right prostate extends posteriorly from xviii through xix and xx into xxi, where it curves sharply towards the dorsal aspect of the intestine and terminates. There are two pairs of small racemose vesiculae seminales, a pair in xi and a pair in xii. Segments ix and × contain a mass of loosely packed sperm cells.

Slender micronephridial tubules occur in each segment except the first.

The species is easily distinguishable from other species of Megascolides. No other species has cuplike depressions surrounding the prostatic pores and the form of the ovaries is a diagnostic character found in no other species of the genus.

Megascolides parvus n.sp. (Plate 12, figs. 1–3)

A single specimen of this small species was collected from a small remnant of native forest (kahikatea, tawa, nikau) on a hillside three miles north-west of Hick's Bay, beside the Hick's Bay-Opotiki road. It is 26·5 mm. in length and 3·5 mm. in diameter and has 72 segments It is reddish-brown dorsally, paler ventrally, and has a buff clitellum surrounding xiv–xvii (four segments).

The prostomium is epilobous, without a posterior groove. There are eight chaetae on each segment. On xxiv their arrangement is as follows: ab = 1·25 mm.; cd = 1·75 mm.; aa = 1 5 mm.; bc = 1·75 mm.; dd = 2·25 mm.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

There are three pairs of spermathecal pores, a pair in each of the intersegmental grooves 6/7, 7/8 and 8/9, one on each side slightly medial to the line of chaeta a in each of the grooves. The female pores are on xiv, one on each side, very close to the ventral midline and close to the anterior margin of the segment. The male pores are on xviii, one on each side, in line with chaeta a of the adjacent segments. Each pore opens on the apex of a low, rounded papilla. There is an unpaired tuberculum pubertatis, taking the form of a tumid ridge, extending across the ventral midline in the intersegmental groove 19/20 from the line of chaeta a on one side to the same line on the other side and a pair of small oval tubercula on xviii, one on each side, posterior to the male papilla of the same side. I could find no dorsal pores.

Internal Anatomy (Plate 12, fig. 2)

Septa vii/viii, viii/ix, ix/x, x/xi, xi/xii and xii/xiii are slightly thickened.

Alimentary Canal. The pharynx occupies the first four segments. The posterior dorsal and lateral aspects of the pharynx are covered by masses of salivary gland tissue. There is a thick-walled globular gizzard in v. The oesophagus extends from vi to xiv and there is a pair of prominent rounded calciferous glands in xii. The intestine commences in xv.

Vascular System. The dorsal blood vessel is unpaired. There are three pairs of dilated hearts, a pair in each of x, xi and xii.

Reproductive System. There are two pairs of minute lobate testes, a pair in x and a pair in xi, and a pair of ovaries in xiii. The testes and ovaries are

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situated one on each side, attached to the anterior septum in their respective segments. There are three pairs of spermathecae, a pair in each of vii, viii and ix. Each spermatheca consists of a conical sac, opening by a muscular duct to the exterior, and a small spherical diverticulum opening by a very slender duct into the spermathecal duct. (Plate 12, fig. 3.) The prostates lie one on each side of the ventral nerve cord beneath the intestine and are very long, slender, tubular organs. The left prostate originates from a slender duct passing through xviii and xix and extends back from xx to xxxiv, turns and passes forward again through xxxiii into xxxii, where it terminates. The right prostate passes back only as far as xxxiii, where it terminates. There are two pairs of small, racemose, vesiculae seminales, a pair in ix and a pair in xii.

There are micronephridial tubules in each segment except the first. They form a compact mat of tubules on the lateral aspects of the body-wall in each segment.

Remarks. M. parvus closely resembles and may actually be identical with M. huttoni (Benham, 1904) The inadequacy of Benham's description makes it impossible to positively distinguish the present species from M. huttoni, but the following points of difference are noted:

(i) The male pores of M. parvus are situated on low, rounded papillae on xviii; those of M. huttoni are in slight circular depressions on xviii.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

(ii) M. parvus has a median oval tuberculum pubertatis at 19/20; M. huttoni has a similar tuberculum on the hinder margin of xviii.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

(iii) The spermathecal pores of M. parvus are intersegmental, in the grooves 6/7, 7/8 and 8/9; those of M. huttoni are on the hinder margin of vi, vii and viii.

(iv) The gizzard of M. parvus is short and thick-walled; that of M. huttoni is long, narrow and thin-walled.

(v) The micronephridia of M. parvus commence in ii; those of M. huttoni commence in iii.

(vi) The prostates of M. parvus extend straight back from xviii to xxxiii or xxxiv; those of M. huttoni extend forward from xviii to xvi.

None of these characters is of specific value, since none of them is invariable, but M. parvus does not show sufficient similarity to the scanty description of M. huttoni to be positively identified as M. huttoni, and I prefer to establish a new species for it.

Megascolides raglani n.sp. (Plate 13, figs. 1–3)

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Two specimens were collected, one from a forest remnant near the crest of the Raglan Range, south of the Hamilton-Raglan road, and the other from native forest near Moerangi, beside the Raglan-Kawhia road. Superficially the species is easily mistaken for Spenceriella shakespeari (Benham). It is about the same size and colour and has prominent, transversely disposed tubercula pubertatis, one at 17/18 and another at 18/19, similar to those found in S. shakespeari. However, the possession of only eight chaeta on each segment as compared with 48 in S. shakespeari readily distinguishes the two species. The strong similarity between the two species may be interpreted as an indication that S. shakespeari has been derived from M. raglani as a local variant.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

The specimen on which this description is based is 81 mm. in length and 5 mm. in diameter and has 101 segments. It is brick red dorsally and pale ventrally and has a buff clitellum which completely surrounds xiii–xviii. The clitellum is not greatly developed over xiii and xviii and the intersegmental furrows 13/14 and 17/18 are quite well marked. The prostomium is epilobous.

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There are eight chaetae on each segment, arranged in pairs. On xxiv their arrangement is as follows:

ab = cd = 1.75 mm.; aa = 2·25 mm; bc = 2.25 mm.; dd = 2·5 mm.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with the chaetal interspace ab. There is an unpaired median ventral female pore on xiv, anterior to the level of the chaetae. The male pores are on xviii and their arrangement and the arrangement of the structures associated with them is strikingly similar to the arrangement seen in Spenceriella shakespeari. A pair of small oval depressions with slightly raised white borders lies one on each side of xviii, and in each of these small depressions, in line with chaeta a of the adjacent segments, is a small male pore. There are two very prominent, transversely disposed, oval, glandular tubercula pubertatis, one in each of the intersegmental furrows 17/18 and 18/19, extending across the ventral midline. There are no other tubercula pubertatis.

Internal Anatomy (Plate 13, fig. 2)

The septa ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are muscular and thickened.

Alimentary Canal. The pharynx occupies the first four segments and posteriorly it is covered dorsally and laterally by salivary glands. The gizzard has thick muscular walls and lies in v. The oesophagus extends from vi to xv and there are no oesophageal glands. The intestine commences in xvi.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of hearts, a pair in each of ix, x, xi and xii. Those of ix are much more slender than those of the subsequent segments.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Reproductive System. There are two pairs of testes, a pair in x and a pair in xi, and a single pair of ovaries in xiii. The testes and ovaries have a similar, compact, lobate form and are situated one on each side of the ventral midline, attached to the anterior septa of their respective segments. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each consists of a large sac, almost spheroidal, but with a finger-like lobe lying over its upper surface and projecting towards the midline. The sac opens to the exterior by a long, thickwalled duct, and an elongate pyriform diverticulum opens medially into the duct (Plate 13, fig. 3). In one specimen the right anterior spermatheca was situated in vii instead of viii, but its duct passed back through vii and opened to the exterior at the intersegmental furrow 7/8, as is normal in the species. The prostates are short, curved, tubular organs, lying one on each side of the ventral nerve cord. They open to the exterior through a narrow duct, one on each side of xviii, and the organs extend back from xviii to xix. There are three pairs of racemose vesiculae seminales, a pair in each of ix, xi and xii. Those of ix are small rounded organs, attached to the posterior septum of the segment, while those of xi and xii are larger organs, extending around the lateral aspects of the oesophagus and attached to the anterior septa of their respective segments.

Micronephridia occur in every segment except the first, as a mat of small, slender tubules, covering the lateral aspects of the peritoneum. In the eleven most posterior segments there are very small paired meganephridia, in addition to the micronephridia. The meganephridia do not appear to be functional. Each consists merely of a tightly coiled tubule with no obvious funnel or elimination duct, lying close to the ventral nerve cord on its side of the segment.

Megascolides fuscus n.sp. (Plate 14, figs. 1–3)

Several specimens of this large brown earthworm were collected from topsoil

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Fig. 1—Megascolides viridis. Ventral aspect, segments i–xxi.
Fig. 2—M. viridis. Dissection from the dorsal aspect.
Fig. 3—M. viridis. Left anterior spermatheca anterior aspect.

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Fig. 1—Megascolides ruber. Ventral aspect segments i–xxi.
Fig. 2—M. ruber. Dissection from the dorsal aspect.
Fig. 3—M. ruber. Left posterior Spermatheca, lateral aspect.

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Fig. 1—Megascolides parvus. Ventral aspect, segment i–xx.
Fig. 2—M. parvus. Dissection from the dorsal aspect.
Fig. 3—M. parvus. Right spermatheca of viii, posterior aspect.

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Fig. 1—Megascolides raglani. Ventral aspect, segments i–xxiii.
Fig. 2—M. raglani. Dissection from the dorsal aspect.
Fig. 3—M. raglani. Right posteior spermatheca. posterior aspect.

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Fig. 1—Megascolides fuscus. Vential aspect, segments i–xx.
Fig. 2—M. fuscus. Dissection from the dorsal aspect.
Fig. 3—M. fuscus. Left spermatheca, medial aspect.

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Fig. 1—Megascolides irregularis. Ventral aspect, segments i–xx.
Fig. 2—M. irregularis. Dissection from the dorsal aspect.
Fig. 3—M. irregularis. Left posterior spermatheca, anterior aspect.

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Fig. 1—Megascolides alba. Ventral aspect, segments i–xxxi.
Fig. 2—M. alba. Dissection from the dorsal aspect.
Fig. 3—M. alba. Left posterior spermatheca, posterior aspect.

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Fig. 1—Megascolex novae-zealandiae. Ventral aspect, segments i–xx.
Fig. 2—M. novae-zealandiae. Dissection from the dorsal aspect.
Fig. 3—M. novae-zealandiae. Right anterior spermatheca, lateral aspect.

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Fig. 1—Pheretima campestris. Ventral aspect, segments i–xix.
Fig. 2—P. campestris. Dissection from the dorsal aspect.
Fig. 3—P. campestris, Left posterior spermatheca anterior aspect.

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Fig. 1—Diporochaeta obtusa. Ventral aspect, segments i–xxi.
Fig. 2—D. obtusa. Dissection from the dorsal aspect.
Fig. 3—D. obtusa. Left posterior spermatheca, medial aspect.

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Fig. 1—Perionyx egmonti. Ventral aspect, segments i–xx.
Fig. 2—P. egmonti. Dissection from the dorsal aspect.
Fig. 3—P. egmonti. Right posterior spermatheca, medial aspect.
Fig. 4—P. egmonti. Right prostate, ventro-lateral aspect.
Fig. 5—P. egmonti Vertical section through prostate. (Diagrammatic).

acc gl., accessory gland, e.g., cerebral ganglion, ch., chaeta, el, clitellum; c.p.e., central prostatic cavity; d.b.v., doisal blood vessel, d.div., duct of spermathecal diverticulum, div., spermathecal diverticulum, f.p., female pore; g., gizzard, h., heart, int., intestine; l., lobule of piostate; m., mouth; m.f., “male field”, m g., mucus gland, mn., meronephridial tubule, m p., male pore, n, meganephridial tubule; o., ovary, oe., oesophagus; oe.gl., oesophageal gland, o f., ovarian funnel; p., prostate; pap., prostatic papilla, p.d, prostatic duct, peri. peristomium, ph, pharynx, pr, prostomium, pv., proventriculus; s, septum; sal.gl., salivary gland, sh., sheath; s.i.v., supra-intestinal vessel; sp., spermatheca; sp.d., spermathecal duct, sp.p, spermathecal pore; t., testis, t.f., testicular funnel; t.pub., tuberculum pubertatis, v.b.v., ventral blood vessel, v.n.c., ventral nerve cord, v.s., vesicula seminalis.

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and subsoil in Taupo sandy silt in areas of native rain forest and exotic forest (Pinus spp.) in the Rotorua district.

The specimen on which this description is based is 251 mm. in length and 10 mm. in diameter and has 268 segments. The clitellum is greyish-brown and is developed over the entire surface of xv and xvi and on the dorsal and lateral surfaces of xiv, xvii and xviii. The prostomium is epilobous. The first six segments are biannulate, segments vii–xviii are quadriannulate and the post-clitellar segments are biannulate. There are eight chaetae on each segment, arranged in pairs. On xxiv: ab = 1·25 mm.; cd = 2·25 mm.; aa = 4·4 mm.; bc = 4·5 mm.; dd = 10 mm.

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There is only one pair of spermathecal pores, at 7/8 in line with the chaeta b. There is a single pair of female pores slightly anterior and lateral to the chaeta a on xiv. The male pores open on the apices of a single pair of prominent rounded papillae on xviii. The chaeta a is absent on xviii and in its place on each side is the papilla bearing the male pore. There are no tubercula pubertatis. Dorsal pores occur in every intersegmental groove posterior to the clitellum. There are no nephridiopores.

Internal Anatomy (Plate 14. fig. 2)

The first septum occurs at the intersegment iv/v. The septa vii/viii, viii/ix, ix/x, x/xi and xi/xii are muscular and thickened.

Alimentary Canal. A rounded muscular pharynx occupies the first four segments. The pharynx is covered dorsally and laterally by a downlike mass of salivary glands. There is a thin-walled proventriculus in v and a stout muscular gizzard in vi. The oesophagus extends from vii to xvi and there are no oesophageal glands. The intestine commences in xvii, is thin-walled and has a small typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are four pairs of dilated hearts in x, xi, xii and xiii, rising from a supraintestinal vessel which extends only through those four segments.

Reproductive System. There are two pairs of testes in x and xi. The testes have a lobate form and their funnels are on the posterior septa of the segments. The two slender vasa deferentia of each side are prominent, lying side by side on the ventro-lateral aspect of the peritoneum They remain distinct from each other for most of their length, fusing just before they enter the prostatic duct, in xviii. There is a single pair of ovaries in xiii. Each ovary appears as a thin lace-like lamina attached to the anterior septum of the segment. The ovarian funnels are very prominent white structures on the posterior septum of xiii, and the short oviducts, which open to the exterior in xiv, are very much thicker than the vasa deferentia. A single pair of spermathecae occurs in viii. Each is a large, roughly ovoidal, thin-walled sac with a small, thick-walled digitate diverticulum opening into the anterior aspect of its duct. (Plate 14, fig. 3). The whole organ is confined to the eighth segment. The prostates are large, coiled, tubular organs, extending through the segments xviii, xix, and xx and opening to the exterior in xviii by wide muscular ducts. The prostatic duct of each side fuses with the corresponding vas deferens close to the body-wall in xviii. There are two pairs of large, finely racemose vesiculae seminales surrounding the lateral aspects of the oesophagus in xi and xii.

Micronephridia commence in ii. There is a compact cluster of micronephridial tubules on the ventro-lateral aspect of the peritoneum on each side of the segments. The individual tubules are very long and are twisted like corkscrews.

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Remarks. This species most closely resembles M. (Tokea) sapida Benham (1904), collected from Ruatahuna, between Rotorua and Lake Waikaremoana. It agrees with Benham's species in size, colour, and in many points of its anatomy, but the number and form of the spermathecae in the two species are markedly different. M. sapida has three pairs of spermathecae, each with a small globular diverticulum, while M. fuscus has only one pair of spermathecae and their diverticula are digitate. The number and form of the spermathecae and their diverticula are very reliable systematic characters and serve to distinguish the two species one from the other.

Megascolides irregularis n.sp. (Plate 15, figs. 1–3)

One specimen of this large worm was collected near Tangowahine, North Auckland. Other specimens have been collected from localities in the same area. The specimen on which the description is based is 117 mm. in length with 120 segments, and has an almost uniform diameter of 7·5 to 8 mm. It is purplish-red in colour dorsally, and pale ventrally. The clitellum is buff in colour and completely surrounds xiv to xvii, except for a small area on the ventral surface of xiv where the female pores occur.

The prostomium is epilobous. There are eight chaetae on each segment, but their arrangement is most irregular. They are not arranged in pairs, but occur four on each side of each segment, scattered indiscriminately. The distance between corresponding chaetae a on each segment is more or less uniform anterior to the prostatic pores, but varies considerably posteriorly.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9. On each side the pores lie in line with chaeta a. A single pair of female pores is present on xiv, very close together in a small pale area covering the ventral mid-line. On xviii is a pair of prostatic pores surrounded by tumid lips. Each pore is in line with the chaetae a on the same side. Across the ventral mid-line, at the anterior margin and at the posterior margin of this segment is a small tuberculum pubertatis.

Nephridiopores are not visible externally.

Internal Anatomy (Plate 15, fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi and xi/xii are thickened and muscular.

Alimentary Canal. A large muscular pharynx, to which is attached dorsally a mass of small salivary glands, occupies the first four segments. A small muscular gizzard lies in v. The oesophagus extends from vi to xv and has a single pair of large calciferous glands in xiii. The intestine commences in xvi, is fairly thick-walled and has a narrow typhlosole on its median dorsal aspect.

Vascular System. The dorsal blood vessel is single throughout and bears four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. Two pairs of small lobate testes lie close to the ventral mid-line, a pair in x and a pair in xi, and a pair of very small lobate ovaries lies close to the ventral mid-line in xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each spermatheca takes the form of an ovoidal sac with a wide muscular duct leading to the exterior. A small rounded diverticulum opens into the medial aspect of the duct close to its distal extremity (Plate 15, fig. 3). A single pair of small tubular prostates is present. Each prostate opens by a narrow muscular duct in xviii, but the organs are of unequal extent. The left prostate extends through xviii and xix to terminate in xx in

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a coiled portion. The right prostate extends posteriorly, without coiling, through xviii, xix and xx to xxi, where it terminates. The prostates are noteworthy in that they lie ventral to the intestine and close to the ventral nerve cord, thus disposed similarly to those of M. (Tokea) esculenta Benham, 1904. There are two pairs of racemose vesiculae seminales, a pair in ix and a pair in xii.

A band of micronephridial tubules is present in each segment except the first, but on each side of the first segment there is a cluster of tubules larger than those of the micronephridia which probably function as a slime gland, as in some species of Octochaetus (e.g. O. multiporus Beddard, 1892).

Remarks. The asymmetrical arrangement of the chaetae in this species is an unusual character, and one of which the author can find no previous record in this genus. It would seem to be a secondary modification of the paired arrangement normally found in the genus. The disposition of the vesiculae seminales is also unusual, and assists in distinguishing the species from others of the same genus.

Megascolides alba n.sp. (Plate 16, figs. 1–3)

Several specimens of this large earthworm were collected from Motatau clay in a remnant of native bush about a mile south-east along the main Whangarei-Auckland highway from the Maungakaramea road junction. The specimen on which this description is based is 90 mm. long, has 141 segments, and is 7.5 mm. in diameter for the greater part of its length It is completely unpigmented except for a slightly yellowish clitellum surrounding xiv to xvii. The transparency of the body-wall makes the species readily recognizable, since the prostates are visible through the body-wall as a pair of yellowish streaks lying close to the ventral nerve cord and extending from xviii to xxviii or xxix.

The prostomium is epilobous. Chaetae are lumbricine in arrangement, there being eight on each segment. In the mid body:

ab = 1·5 mm.; cd = 1·25 mm.; aa = 2·5 mm.; dd = 9 mm.; bc = 2 mm.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with chaeta a. A single pair of female pores is present on xiv, each pore situated slightly medial to chaeta a. There is a single pair of prostatic pores on xviii. Each pore is situated in the centre of an oval tumid area which lies in line with the chaetal interval ab on each side. Chaetae are absent on xviii. Tubercula pubertatis are present. Each is in the form of a large, oval, white pad which lies across the ventral mid-line, one at 16/17 and one at 19/20.

Nephridiopores are not visible externally.

Internal Anatomy (Plate 16, fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi and xi/xii are muscular and thickened.

Alimentary Canal. A small muscular pharynx occupies the first four segments. There is a strongly muscular gizzard in v. The oesophagus extends from vi to xvi and lacks oesophageal glands. The intestine commences in xvii.

Vascular System. The dorsal blood vessel is single throughout. There are five pairs of hearts, a pair each in ix to xiii. the pair in ix being much smaller than the others.

Reproductive System. There are two pairs of lobate testes, a pair in x and a pair in xi, and a single pair of ovaries in xiii. Two pairs of spermathecae are present, a pair in viii and a pair in ix. Each is an ovoidal sac with a small medial

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digitiform diverticulum opening into the duct distally. (Plate 16, fig. 3.) There is a single pair of tubular prostates arising from narrow muscular ducts in xviii. They lie close to the ventral nerve cord ventral to the intestine and are of unequal length, the right prostate terminating in xxvii and the left in xxix. Two pairs of vesiculae seminales are present, a pair in x and a pair in xi.

Micronephridia occur in each segment except the first, in the form of a band of slender tubules on the lateral aspects of the body-wall.

The following are supplementary notes on the two species M. suteri (Benham, 1904) and M. urewerae (Benham, 1904).

Megascolides suteri (Benham), 1904

Syn. Tokea suteri Benham, 1904

Megascolides (Tokea) suteri Benham, 1941

The type specimen of M. suteri was collected from Auckland and Benham did not record the species elsewhere It has now been collected from the Rotorua-Taupo district and the East Coast district of the North Island. All the specimens have been collected in or under rotten logs. This habitat preference was not noted by Benham in the case of the type specimen.

The description given by Benham is insufficient to distinguish the species from M. huttoni (Benham) and M. urewerae (Benham), so some additional notes on the characters of the species are set out below.

External Features. There is a pair of female pores on xiv, one on each side, anterior and slightly medial to chaeta a. The tubercula pubertatis may be absent.

Intenal Anatomy. The pharynx occupies the first four segments. There is a short proventriculus in v, opening into the small, feebly developed gizzard in vi. The oesophagus extends from vii to xv and has a pair of slight lateral dilatations in xii. The intestine commences in xvi.

The dorsal blood vessel is unpaired throughout its length. A very slender, unpaired, supra-intestinal vessel commences in x and extends back to xiii. From the supra-intestinal vessel arise three pairs of hearts, a pair in each of x, xi and xii. The supra-intestinal vessel terminates in a number of slender branches, running across the dorsal aspect of the oesophagus in xiii.

There are two pairs of testes, a pair in x and a pair in xi. Each testis is a small, folded, laminar organ. A pair of ovaries occurs in xiii. The ovaries are larger, thicker, laminar organs. There are two pairs of racemose vesiculae seminales, a pair in ix and a pair in xii.

Micronephridia are found in each segment except the first. The tubules form a desnely packed mass over the ventro-lateral aspects of the peritoneum, but are less numerous laterally and absent dorso-laterally.

Megascolides urewerae (Benham), 1904

Syn. Tokea urewerae Benham, 1904

Megascolides (Tokea) urewerae Benham, 1941

A specimen which agrees fairly closely with Benham's description of M. urewerae was collected from native forest on a hillside about seven miles west of Motuhora. Some additional notes on the species are necessary, since the description given by Benham is insufficeint to distinguish the species from M. suteri (Benham) and M. huttoni (Benham). The following additional notes are therefore given.

External Features. The female pores are on xiv, one on each side, close to the ventral mid-line, anterior and medial to chaeta a. The spermathecal pores are

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on vi, vii and vii, slightly anterior to the furrows 6/7, 7/8 and 8/9. A pair of small oval tubercula pubertatis occurs on xviii, one on each side posterior to the pit containing the male pore, extending slightly over the intersegmental furrow 18/19.

Internal Anatomy. The intestine commences in xvi.

The dorsal blood vessel is unpaired throughout its length. Three pairs of hearts arise from the dorsal vessel, a pair in each of x, xi and xii.

There are four pairs of racemose vesiculae seminales, a pair in each of ix, x, xi and xii. The prostates lie under the intestine, one on each side of the ventral nerve cord, and extend straight back through the segments.

The remainder of the anatomy of the specimen agrees closely with the description given by Benham.

Genus Notoscolex Fletcher

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“Chaetae eight per segment. Spermathecal pores one, two or three pairs, the last in furrow 8/9 (in certain abnormal species 7/8). One gizzard in segment v or vi. Micronephridia present, sometimes with meganephridia also. Prostates with branched system of ducts.” (Stephenson, 1930.)

Only two species from the New Zealand region, N. equestris Benham (1942) and N. hakeaphilus Benham (1949) are assigned to the genus Notoscolex. Of these two it is possible that N. hakeaphilus is introduced. The two species are readily distinguished by the folloiwng differences:

(i) N. hakeaphilus is a large worm (up to 650 mm. in length); N. equestris is about 200 mm. in length.

(ii) N. hakeaphilus is brick red in colour dorsally, pale ventrally; N. equestris is coloured alternately dark brown and pale cream in narrow bands.

(iii) N. hakeaphilus has two pairs of spermathecae, a pair in each of viii and ix; N. equestris has three pairs, a pair in each of vii, viii and ix.

(iv) N. hakeaphilus has three pairs of hearts, a pair in each of viii, ix and x; N. equestris has four pairs, in x, xi, xii and xiii.

There are many other points of difference which are obvious from Behnam's descriptions of the species (q.v.).

Genus Spenceriella Michaelsen

“Chaetae more than eight per segment. Spermathecal pores one to three pairs. One gizzard in segment v. Micronephridial. Prostates tubular, with simple unbranched duct.” (Stephenson. 1930.)

Two species of the genus were described by Benham (1905(a)) and referred by him to the genus Diporochaeta. Both species were collected from Little Barrier Island. Michaelsen (1907) established the genus Spencieriella for some Australian species, and included Benham's species, in it, since they differ from Diporochaeta in being micronephridial while Diporochaeta is meganephridial. The two New Zealand species are Spenceriella gigantea and S. shakespeari. They may be identified from the following key:

a1 One pair of vesiculae seminales in xii. Last heart in xii. S. gigantea

a2 Two pairs of vesiculae seminales, in ix and xii. Last heart in xiii S. shakespeari

S. gigantea is a very large worm, measuring about 4 ft. 6 in. (1,370 mm.) in length and half an inch (11 mm.) in diameter. It has been recorded only from Little Barrier Island.

S. shakespeari is a small, stout, brick-red worm (about 100–150 mm. in length and 4–5 mm. in diameter). It is readily distinguished in the field by the unusual

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form and disposition of its tubercula pubertatis and the large number of chaetae on each segment. It was collected from Little Barrier Island by Benham, and I have collected it from the western coastal ranges of the Waikato and Taranaki districts.

Genus Megascolex Templeton

“Chaetae, at least in the middle and hinder regions of the body, numerous (more than eight) in each segment. Spermathecal pores usually one to five pairs, between segments iv and ix (the exceptions are constiuted by the few cases where the pores are fused in the middle line, or where they are numerous on each side in each segment) One gizzard in v, vi or vii. Micronephridial. Prostates with brached system od ducts.” (Stephenson, 1930.)

Of about 120 known species only two have been recorded from the New Zealand region. They are M. laingii Benham (1902) and M. novae-zealandiae, a new species, described below. M. laingii is known only from Norfolk Island and M. novae-zealandiae has been found only in the extreme northern portion of the North Island of New Zealand. A species recorded by Baird (1871) as Megascolex antarctica may have been intended as a species of Megascolex, but Baird's description is insufficient for the recognition of the species or genus. There is no other record of the genus from the mainland of New Zealand.

The following is a description of the new species M. novae-zealandiae. Type in my collection.

Megascolex novae-zealandiae n.sp. (Plate 17, figs. 1–3)

Five specimens of this large red earthworm were collected, three from Awapuka clay loam six miles from Mangonui, and two from a steep bush-clad hillside two miles and a half south-east of Ahipara. The specimen on which this description is based is 179 mm. in length and 9·5 mm. in diameter, and has 120 segments.

The prostomium is epilobous. Chaetae average about 36 per segment, but there may be more or less. They are perichaetine in arrangement and commence on ii. Their spacing on the segments is not regular. The clitellum is darker in colour than the adjacent segments, and surrounds xiv to xviii, and the anterior portion of xix.

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Spermathecal pores are not visible externally. A single pair of female pores lies adjacent to the ventral midline on the anterior aspect of xiv. They cannot be seen with the naked eye, and are very close together. A single pair of male pores is present on xviii, each pore situated 1·75 mm. from the ventral midline and occupying the centre of a small oval light brown area. A prominent oval tuberculum pubertatis lies in the intersegmental groove 17/18, and another similar tuberculum in the groove 18/19. There are no nephridiopores.

Internal Anatomy (Plate 17, Fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are thickened and muscular.

Alimentary Canal. A muscular pharynx occupies the first four segments. The gizzard, which has thick muscular walls, is elongate and is confined to v. The oesophagus extends from vi to xv and lacks oesophageal glands. The intestine commences in xvi, is thin walled and has a marked typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. There are two pairs of free testes, a pair in × and a pair in xi, and a pair of ovaries in xiii. Two pairs of spermathecae are present, a pair

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in viii and a pair in ix. Each spermatheca is in the form of a large pyriform sac opening ventrally to the exterior by a wide muscular duct. A small ovoidal diverticulum opens by a slender duct into the anterior aspect of the spermathecal duct. (Plate 17, Fig. 3.) The spermathecal pores are in 7/8 and 8/9. A single pair of prostates is present. The left prostate commences from a short duct in xviii and extends anteriorly through xvii/xviii to terminate in xvii. The right prostate commences from a similar duct in xviii and extends posteriorly through the septa to terminate in xxiii, lying close to the ventral nerve cord throughout its length. Examination of sections of the prostate shows that the axial duct receives small ductules from the surrounding prostatic tissue.

Micronephridial tubules occur in a wide band on the lateral walls of each segment except the first. Anterior to the gizzard the tubules are very numerous and completely cover the lateral walls of the segments.

This species is distinct from M. laingii, since it has four pairs of hearts, while M. laingii has only three pairs; it has large prostates while M. laingii has small compact prostates, and in a number of other points of internal and external anatomy it is also markedly distinct from M. laingii.

Genus Pheretima Kinberg

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“Chaetae numerous in each segment. Spermathecal pores one to six pairs, between segments iii and ix One gizzard between 7/8 and 10/11. Micronephridial. Testes and funnels enclosed in testis sacs; prostates with branched system of ducts; penial chaetae almost always wanting.” (Stephenson, 1930.)

There are about three hundred and fifty known species, of which only two are recorded from New Zealand. The two New Zealand species are P. clerica Benham (1947), collected from the Gisborne district, and P. campestris n.sp., collected from North Auckland and Raoul Island and described in this paper. P. campestris may be regarded as introduced into Raoul Island, probably with plants and seeds taken from New Zealand within the last few years. P. campestris may not actually be a new species, but since I cannot obtain literature concerning most of the great number of species belonging to the genus and it does not coincide with any of the descriptions that I am able to find. I have established a new species for it.

The following is a description of P. campestris. Type in my collection.

Pheretima campestris n.sp. (Plate 18, figs. 1–3)

Two specimens were collected from Mangakahia loam in a top-dressed pasture on the Victoria Valley road near Kaitaia, and two specimens from Otonga peaty loam in a top-dressed pasture near Awanui, North Auckland.

In both cases the species was found together with the introduced species Octolasium cyaneum and Allolobophora caliginosa, living in the upper inch of the topsoil among the roots of the pasture grasses. It is easily distinguished from the introduced species by its yellow-brown colour and its rapid movement when disturbed.

The specimen on which this description is based is 87 mm. in length, has 100 segments and is 3·75 mm. in diameter. The chaetae are perichaetine in arrangement, about 48 on each segment in the midbody, and evenly spaced around the perimeter of each segment. The clitellum is darker in colour than the adjacent segments and completely surrounds xiv to xvi. There are no chaetae on the clitellar segments.

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Spermathecal pores are not visible externally. There is a single median ventral female pore, in the centre of a median pigmented area on xiv. A pair of male pores is present on xviii. Each of the male pores occupies the centre of a round white area on a small papilla and is 1·5 mm. from the ventral midline. There are twelve chaetae between the two male pores.

Internal Anatomy (Plate 18. Fig. 2)

There are no thickened septa.

Alimentary Canal. An elongate muscular pharynx lies in i to iv. The gizzard, as in P. clerica, lies posterior to vii/viii and there is no septum between viii and ix, nor between ix and x. The oesophagus commences slightly anterior to x/xi and extends to xiv. There are no oesophageal glands. The intestine commences in xv and the junction between the oesophagus and the intestine is quite distinct. A pair of flattened cornuiform intestinal diverticula arise from the ventral surface of the intestine in xxvi and extend anteriorly, one on each side of the ventral nerve cord, through xxv into xxiv.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. There are two pairs of testes, a pair in × and a pair in xi, each testis lying in a testis sac. There is a pair of small lobate ovaries in xiii. Although the specimen is obviously sexually mature, there are no prostatic glands, although there is a pair of coiled muscular ducts in xviii resembling prostatic ducts. The vas deferens of each side opens to the exterior by the corresponding “prostatic” duct. The author dissected another mature specimen and found the same condition of the male genital organs. The vas deferens simply expands in xviii into what appears to be a prostatic duct, which coils before opening to the exterior, and there is no sign of a prostatic gland. This may be due to a seasonal regression of the prostates, but it is similar to the condition described by Stephenson (1930, p. 370) in P. heterochaeta. There are four pairs of spermathecae, a pair each in vi, vii, viii and in ix. Each spermatheca is in the form of a flattened cornuiform sac which opens to the exterior by a wide duct. A long, very narrow duct arises laterally from the spermathecal duct close to the body-wall and terminates in a small spheroidal diverticulum. (Plate 18, Fig. 3.) There are two pairs of lobate vesiculae seminales, a pair in xi and a pair in xii.

Micronephridia commence in ii and occur in every other segment except x. Septa viii/ix and ix/x are absent, but the nephridial tubules corresponding to viii and ix are referable to the spermathecae corresponding to those segments. There are, however, no nephridial tubules corresponding to x.

Remarks. This species shows a close similarity to P. clerica Benham (1947), but can be distinguished from P. clerica by the following points of difference:

(1) There are 48 chaetae on each segment in the midbody; P. clerica has 36.

(2) The colour of this species is yellowish-brown; P. clerica is purplish-brown in colour.

(3) There are no small tubercula pubertatis beside the male pores; tubercula pubertatis are present in P. clerica.

(4) The vesiculae seminales lie in xi and xii; in P. clerica they lie in x and xi.

(5) The transition from the posterior region of the oesophagus into the intestine is quite distinct, the oesophagus terminating in xiv and the intestine

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commencing in xv; in P. clerica it is difficult to determine the limits of the oesophagus and the intestine.

(6) The intestinal caeca arise in xxvi; in P. clerica they arise in xxvii or xxviii.

(7) Benham does not mention the absence of the micronephridia corresponding to × in P. clerica; these micronephridia are absent in P. campestris.

(8) There are no prostates in P. campestris; prostates are present in P. clerica.

It is the accumulation of such small differences which serves to distinguish one species from another, and the differences in this case are of an order which leaves little doubt that P. campestris is a species distinct from P. clerica.

Genus Diporochaeta Beddard

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“Chaetae, at least in the middle and hinder parts of the body, numerous (more than eight) per segment. Spermathecal pores two to five pairs, the last in furrow 8/9. One gizzard in the region of segments iv–vi, seldom vestigial. Purely meganephridial. Prostates tubular, with simple unbranched duct.” (Stephenson, 1930.)

Of the forty-eight known species, three have been recorded from the New Zealand region by previous workers and one more (D. obtusa) is described below. The three previously recorded species are D. intermedia Beddard (1890), an aquatic species, from the west coast of the South Island, D. aquatica Benham (1903), an aquatic species from Lake Manapouri in the south-western region of the South Island, and D. chathamensis Benham (1900), from Chatham Island. D. intermedia and D. aquatica are the only known aquatic species of Diporochaeta. D. aquatica was found at depths of 350–500 feet beneath the lake surface (see Benham, 1903a).

The New Zealand species may be identified from the following key:

a1 Two pairs of spermathecae D. aquatica
a2 Three pairs of spermathecae
b1 Spermathecae without diverticula D. chathamensis
b2 Spermathecae with diverticula D. obtusa
a3 Four pairs of spermathecae D. intemedia

The following is a description of the new species D. obtusa. Type in my collection.

Diporochaeta obtusa n.sp. (Plate 19, figs. 1–3)

Two specimens of this species were collected from Taupo sandy silt in an area of exotic forest two miles south of Rotorua. Specimens have also been collected from Stokes Valley (Wellington), the East Cape district, Norsewood, Waiouru, Lake Waikaremoana, Awakino, Kawhia, and Raglan. Such a wide distribution is remarkable for an endemic earthworm species, and it is difficult to imagine how such a distribution could have been achieved in one species while other species have generally a very limited distribution.

The specimen on which this description is based is 21 mm, in length, with 65 segments. The diameter is 1·5 mm. and the body terminates bluntly at the mouth and anus. The prostomium is tanylobous. The chaetae are perichaetine in arrangement, eighteen per segment, and are evenly spaced around the body except that the ventral gap is slightly greater than any other. The clitellum covers only segments xiv–xvi and is continuous right around these segments. Dorsal pores are obvious, commencing at iv/v and occurring in each intersegmental groove posterior to this.

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Three pairs of spermathecal pores occur at 6/7, 7/8 and 8/9 in line with chaeta b. A pair of small round white markings lies close together, one on each

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side of the mid-ventral line on the anterior margin of ix, in close association with the spermathecal pores, and a single similar marking is present on viii on the right, and on vii on the left. There is a single pair of female pores on xiv, slightly anterior to chaeta a. There is a very prominent pair of white papillae on xviii, between the chaetae a and b on each side, and on the apices of these papillae the common male and prostatic pores open to the exterior. Tubercula pubertatis occur on xvii, xviii, and xix. Those of xvii and xix take the form of a pair of small papillae anterior to the chaetae a and b on xix and posterior to those chaetae on xvii. Those of xviii are large transverse bands on the anterior and posterior margin of the segment, extending from the line of chaeta a of one side to the same line on the other side. There is also on this segment a pair of small papillae immediately anterior to the chaeta c of each side.

The nephridiopores are lateral and occur in a single series on each side in line with the chaeta e.

Internal Anatomy (Plate 19, Fig. 2)

Alimentary Canal. The pharynx occupies the segments i–v, is not very muscular, and is covered dorsally by salivary glands. There is a small gizzard in vi. The oesophagus extends from vii to xvi and has three pairs of calciferous glands in xiii, xiv, and xv. The intestine commences in xvii and lacks a typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are three pairs of hearts in x, xi, and xii, those of xii being extremely dilated.

Reproductive System. There is one pair of free polylobate testes in xi, close to the mid-ventral line, and a single pair of ovaries close to the mid-ventral line in xiii. There are three pairs of spermathecae in segments vii, viii, and ix. Each consists of a lobulated sac opening by a narrow duct to the exterior, with a small, smooth-walled diverticulum lying close to the medial aspect of the sac and opening into the spermathecal duct. (Plate 19, Fig. 3.) One pair of very large prostates occurs. Each of the prostates is a wide, convoluted, tubular structure extending through the segments xviii to xxiii, closely applied to the lateral wall of the gut and opening to the exterior by a narrow duct in xviii. There are no penial chaetae. There is only one pair of vesiculae seminales, in xii. There is, however, a diffuse mass of sperms lying loose in the coelomic cavity of xi, the same as was recorded by Benham in D. chathamensis.

Remarks. D. obtusa shows very close affinities to D. chathamensis and also closely resembles the species D. maplestoni Spencer and D. walhallae Spencer from Australia. The differences between D. obtusa and these three species are set out in the accompanying table:

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D. obtusa D. chathamensis D. maplestoni D. walhallae
No. of chaetae on each segment 18 16 14 20–24
No. of spermathecae 3 pairs 3 pairs 2 pairs 5 pairs
Spermathecal diverticulum present absent absent absent
No. of testes 1 pair 1 pair 1 pair 2 pairs
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Genus Perionyx Perrier

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“Chaetae numerous (more than eight) per segment, in rings which are often almost closed. Male pores often approximated in greater or less degree, and may be very close to the middle line. Female pores unpaired (? always). Spermathecal pores, like the male pores, often very near the middle line, the last pair in 7/8 or 8/9. Gizzard very frequently more or less vestigial, in v or vi. Meganephridial. Two pairs of testes and funnels. Prostates with branched system of ducts.” (Stephenson, 1930.)

Of about fifty species, four have been recorded from the Auckland Islands and one (P. heterochaeta) from Snares Island, within the New Zealand region. These five species are P. heterochaeta Benham, 1909), P. brachysoma (Benham. 1909), P. perionychopsis (Benham, 1909), P. helophilus (Benham, 1909), and P. duodecimalis Michaelsen (1923). The first four of these species were placed in the genus Diporochaeta by Benham, but were transferred by Michaelsen (1923) to Perionyx. One new species, P. egmonti, is described in this paper. It is the first species of the genus to be recorded from the mainland of New Zealand. When I first discovered it, on the slopes of Mt. Egmont, I thought that it might have been introduced with some plants from Australia or India, but I have subsequently found it in areas of native forest inland from Mt. Egmont, as far east as Raurimu, in the central North Island near Mt. Ruapehu, so it is unlikely that it has been introduced. It does not coincide with any of the descriptions of species from the Auckland Islands, Snares Island, Australia or India.

The following is a description of P. egmonti. Type in my collection.

Perionyx egmonti n.sp. (Plate 20, figs. 1–5)

This a very large and distinctive species of earthworm, fairly common on the eastern slopes of Mt. Egmont. After heavy rain specimens are frequently found crawling on the surface of the ground. The specimen described below was found on a path near the Stratford Mountain House after heavy rain. It is brick-red dorsally, paler ventrally, and the pink clitellum surrounds xiii to xvii. It is 138 mm. in length and 10 mm. in diameter and has 156 segments. The prostomium is epilobous. There are about 80 chaetae on each segment, evenly spaced except that the ventral gap is about twice that between the other chaetae and the dorsal gap is about twice the ventral gap.

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The spermathecal pores are not visible externally. A pair of female pores occurs on xiv, a pore slightly anterior to the ventral-most chaeta of each side of the segment. The male pores are on xviii, one on each side in line with the second from ventral-most chaeta of the adjacent segments. Each pore is situated on the apex of a small papilla. The papillae lie on the lateral boundaries of an oval “male field” bounded anteriorly and posteriorly by the margins of xviii. Within this “male field” are three pairs of small, rounded tubercula pubertatis, a pair adjacent to one another across the ventral midline on the anterior border of the “male field”, a pair similarly placed on the posterior border of the “male field”, and a third pair lying one on each side, immediately anterior to the prostatic papillae. Another pair of smaller tubercula pubertatis lie one on each side of the ventral midline at 19/20. (Plate 20, Fig. 1.)

The nephridiopores cannot be seen externally.

Internal Anatomy (Plate 20, Fig. 2)

The septa vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii, and xiii/xiv are thickened and muscular.

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Alimentary Canal. The pharynx is large and muscular and occupies the first four segments. A thin, lace-like salivary gland is attached to each side of the pharynx near its posterior margin. The gizzard is very well developed and strongly muscular and lies in v and vi. The oesophagus extends from vii to xvi. There are five pairs of dilated calciferous glands, a pair in each of vii, viii, ix, xiii, and xiv, and in vii there is a pair of accessory glands, similar in form to the salivary glands, but smaller in size, pressed against the lateral walls of the oesophagus and opening by a narrow duet into the ventro-lateral aspect of the oesophagus, independently on each side. The function of these accessory glands is unknown, but they probably secret some enzyme important in the metabolism of the worm. The intestine commences in xvii. It has thin, almost transparent walls and is very wide.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of dilated hearts, a pair in each of x, xi, xii, and xiii, arising from a slender supra-intestinal vessel which extends only from x to xiii.

Reproductive System. There are two pairs of testes, a pair in x and a pair in xi. The testes are small lobate organs situated close to the ventral midline on the anterior septa of their respective segments. A pair of ovaries occupy a similar position in xiii. The ovaries take the form of a small flattened sheet, attached to the anterior septum of the segment, one on each side. The free edge of the ovary, which projects into the coelomic cavity of the segment, is fimbriate. There are four pairs of spermathecae, a pair in each of v, vi, vii, and viii. Each consists of a simple ovoidal sac without diverticula, opening by a narrow duct to the exterior 0·75 mm. from the ventral nerve cord and close to the anterior septa of its segment. (Plate 20, Fig. 3.) The prostates are dorso-ventrally flattened, mushroom-shaped organs lying beneath the intestine, one on each side of the ventral nerve cord in xviii. Each prostate is enclosed in a transparent sheath and is divided, within the sheath, into a number of small lobules. (Plate 20, Fig. 4.) The prostate opens to the exterior by a narrow duct, arising from the mid-ventral region of the organ. In sections it is seen that the prostatic duct communicates with a central prostatic cavity and that from each lobule of the glandular tissue a small ductule opens into the central cavity. (Plate 20, Fig. 5.) In the region of the prostates are three pairs of rounded glandular masses, attached to the ventral aspect of the peritoneum. They are situated one on each side, at the anterior and posterior borders of xviii and at xix/xx. These positions correspond with the positions of the external tubercula pubertatis and the glands probably secrete a mucous substance which holds the worms together when they are in copulo. There are four pairs of botryoidal vesiculae seminales, a pair in each of ix, x, xi, and xii.

There is a pair of slender meganephridial tubules in each segment except the first. The nephridia open to the exterior through a small vesicle about 1 mm. from the ventral midline on each side, close to the anterior septum of each segment.

Genus Didymogaster Fletcher

“Chaetae eight per segment. Spermathecal pores three pairs, on segments ix, x, and xi. Two gizzards, in vi and vii. Micronephridial. Prostates with branched system of ducts.” (Stephenson, 1930.)

There is only one species of the genus (D. sylvaticus) and it has been recorded from Australia (New South Wales) and from New Zealand. Stephenson (1930) regards it as introduced into New Zealand.

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References

Baird, W., 1871. Megascolex antarctica, an Earthworm from New Zealand. Proc. Linn. Soc., vol. 11, p. 96.

Beddard, F. E., 1890. Proc. Zool. Soc., London, 1890, p. 56.

— 1892. Further Notes on New Zealand Earthworms. Proc. Zool. Soc., London. 1892, p. 668.

Benham, W. B., 1900. On Some Earthworms from the Islands Around New Zealand. Trans. N.Z. Isnt., vol. 33, p. 129.

— 1902. On a New Species of Earthworm from Norfolk Island. Trans. N.Z. Inst., vol. 35, p. 273.

— 1903. On Some New Species of Aquatic Oligochaeta from New Zealand. Proc. Zool. Soc., London, 1903, pt. 2.

— 1903a. A note of the Oligochaeta of New Zealand Lakes. Trans. N.Z. Inst., vol. 36, p. 192.

— 1904. On Some Edible and Other Species of Earthworms from the North Island of New Zealand. Proc. Zool. Soc., London, 1904, pt. 2, p. 220.

— 1905. Additional Notes on the Earthworms of the North Island of New Zealand. Trans. N.Z. Inst., vol. 38, p. 239.

— 1905a. An Account of Some Earthworms from Little Barrier Island. Trans. N.Z. Inst., vol. 38, p. 248.

— 1909. Report on the Oligochaeta of the Subantarctic Islands of New Zealand. Subantarctic Islands of New Zealand. Government Printer, Wellington.

— 1941. Megascolides napierensis a New Species of Earthworm. Trans. Roy. Soc. N.Z., vol. 71, p. 27.

— 1942. Notoscolex equestris, an Earthworm from the Poor Knights Island. Trans. Roy. Soc. N.Z., vol. 72, p. 220.

— 1947. Studies in Earthworms. XLII. The Occurrence of the Genus Pheretima in New Zealand. Trans. Roy. Soc. N.Z., vol. 76, p. 423.

— 1949. A Yard-long Earthworm, Notoscolex hakeaphilus. Trans. Roy. Soc. N.Z., vol. 77, p. 346.

Fletcher, J. J., 1886. Notes on Australian Earthworms. Proc. Linn. Soc. N.S.W., vol. 1, p. 554

Kinberg, J. G. H., 1866. Annulata nova. Kongl. Vetenskap. Akad. Forhandlingar. Stockholm, vol. 23, p. 102.

McCoy, F., 1878. Prodr. Zool. Victoria, vol. 1, dec. 1, p. 21.

Michaelsen, W., 1900. Das Tierreich, 10—Oligochaeta. Berlin, 1900.

— 1907. Oligochaeta. Die Fauna Sudwest Australiens, vol. 1. Jena, 1907.

— 1923. Oligochaeten von Neuseeland und den Auckland-Campbell Inseln nebst einigen anderen Pacifischen Formen. Dr. Th. Mortensen's Pacific Expedition, 1914–16. No. 17.

Perrier, E., 1872. Nouv. Arch. Mus. Hist. nat. Paris, vol. 8, p. 126.

— 1873. Arch. Zool. Exp. gen. Paris, vol. 2, p. 250.

— 1874. C. R. Acad. Sci., Paris, vol. 78, p. 1582.

Stephenson, J., 1930. The Oligochaeta. Clarendon Press, Oxford, 1930.

Sweet, Georgina, 1900. On the Structure of the Spermiducal Glands and Associated Parts in Australian Earthworms. Linn. Soc. Journ. Zool., vol. 29, p. 109.

Templeton, R., 1844. Proc. Zool. Soc., London, vol. 12, p. 89.

Ude, H., 1905. Terricole Oligochaeten von den Inseln der Sudsee und von verschiedenen andern Gebieten der Erde. Zeitt. fur Wiss. Zool., vol. 83.