Holotype Auckland Museum; AMNZ 5254. Single complete specimen, now dissected, from New Zealand, Great Barrier Island, Little Windy Hill (ca. 36°10'S, 175°23'E). Coll: 2.IX.2001, J.W. Early & R.F. Gilbert. “Under rock on forest floor. L11002”. “W-025” on lid. (Small tissue sample was taken for DNA analysis - code RJB09).

Etymology. After Maori name for Great Barrier Island; Aporodrilus is treated as masculine but this place name remains genderless as a noun in apposition.

Diagnosis. Aporodrilus having spermathecal pores paired segmentally in 6, 7 and 8; holandry with seminal vesicles in 9 and 12; oesophageal glands annular in 10–14; large genital markings paired in 17/18 and 18/19 on either side of male pores.

Body circular tapering at both ends. Dark, matt grayish pigment with iridescent cuticular sheen; paler intersegments and setal auriolae. Length 140 mm with 75 segments. Prostomium epilobous. Setae lumbricine, 8 per segment in rows becoming increasingly irregular further back. Clitellum not well marked. Dorsal pores absent. Nephropores not found (meroic). Spermathecal pores segmental, equatorial just below setae a on 6, 7 and 8. Female pores mid-ventral pair anteriormedian to setae a on 14. Male and prostatic pores combined on tiny mounds on 18 in position of deleted setae a. Penial setae not found. Genital markings large, longitudinally symmetrical pads, paired in 17/18 and smaller in 18/19.

Pharyngeal mass to 4. Septa 4/5-10/11 thin, only 11/12/13 with slight thickening and thereafter membranous. Gizzard strong and elongate apparently in 6-7 but discernable in 5 by tracing septum 5/6 to near base despite dorsal-wards displacement. Dorsal blood vessel single; hearts paired and increasingly large in 9-13; supra-oesophageal vessel in 10-13. Nephridia meroic with forests of avesiculate tubules on body wall. Spermathecae in 7, 8 and 9 each with elongate, flaccid ampulla and single, small, clavate diverticulum (inseminated) near base implicated with anterior septum which is transgressed. Holandric: minute funnels in 10 and 11 ventrally; seminal vesicles paired, racemose posteriorly in 9 and anteriorly in 12. Ovaries paired as free egg-string bunches ventrally in 13; ovisacs not found. Prostates tubuloracemose extending to ca. 22 from small flaccid ducts to male pores in 18. Oesophagus with oesophageal glands small in 10 and larger in 11-13 then small again in 14; glands more saccular than composite but dilated compared to extraneous oesophageal width. Intestinal origin in 16. Typhlosole and caeca not found (absent). Gut contains fine colloidal reddish soil.

Lack of dorsal pores is usually associated with aquatic habitat, but possibly also with high rainfall/soil-moisture, however, the strong gizzard suggests a loamy diet. Further ecological and/or behavioural information is wanting.

Aporodrilus aotea compares with Aporodrilus mortenseni (Michaelsen, 1924) that differs, not least, by having its three pairs of spermathecal pores intersegmental in 6/7/8/9 and by lacking genital markings. However, in the review by Lee (1959) that did not routinely note presence or absence of dorsal pores (nor genital markings), the current species keys out nearest to Lee’s Megascolides species now in Notoscolex, viz.: Notoscolex sapidus that differs in its spermatheal pores intersegmental in 6/7/8/9; or to those now in Aporodrilus viz. Aporodrilus equestris (Benham, 1942), and edible Aporodrilus esculentus (Benham, 1904) with which it perhaps comes closest as this too has spermathecae opening on 6-8. Aporodrilus equestris as redescribed below has genital markings elongate in 17 & 19, exceptionally thickened septa, a gizzard in 6 and intestine from 17; while Aporodrilus esculentus has genital markings paired midventrally in 16 and 17, thicker septa, a smaller gizzard, oesophageal dilations only in 15 and its spermathecae of a more spherical and compact form (see figures and compare Benham’s original sketches http://www.archive.org/stream/proceedingsofzoo19042zool#page/240/mode/2up). A more distant contender is Notoscolex urewerae (Benham, 1904) “a short white worm” that has genital marking mid-ventrally in 19/20 and last hearts in 12 amongst other differences (its dorsal pores are unrecorded and possibly it too belongs in Aporodrilus).

Holotype Auckland Museum; AMNZ 5255. Single mature, posterior amputee rather poorly preserved from Waitakere Ranges, Waiatarua. Coll: 9.V.1995, G. Ripley. “Nikau/Ponga forest L761”; “W-012” on lid. (Small tissue sample was taken for DNA analysis coded RJB10). [Two other specimens from the same jar are a posterior portion of a worm (AMNZ 5256) matching the dimensions and frayed edge of the current specimen is itself missing its tip; the other (AMNZ 5254) is a large mature, anterior amputee that is certainly different and probably a new species but which is inadequate for formal description here].

After Maori name for silver fern Cyathea dealbata (G. Forster) Swartz, 1801, from the habitat detail and also the symbol commemorating the All Blacks victory in 2011 Rugby World Cup; Aporodrilus is masculine, but a noun in apposition is genderless.

Aporodrilus having spermathecal pores paired intersegmentally in 7/8/9; metandric with seminal vesicles in 12; no oesophageal glands; genital marking as a distinct pad in 17/18 with male pores on lower rim replacing setae a.

Body robust, dorsally canaliculated in parts before amputation. Pale putty coloured in alcohol. Length 220+ mm anterior portion (a posterior fragment in jar is also 220mm and if from same specimen would give length = 440 mm). Prostomium much wrinkled prolobous. Setae lumbricine, obscure in anterior and mostly occluded on clitellum apparently converging towards male pores; further back the rows except for setal a lines become progressively irregular. Clitellum slightly more tumid and yellowy in ½13–17 (or thereabouts). Dorsal pores absent. Nephropores absent (meroic). Spermathecal pores intersegmental, detected by probe from interior and approximately in setal a lines in 7/8/9. Female pores large paired on 14 (setae obscure) in line with setae a of 13. Male pores superficial on 18 in place of deleted setae a on bottom rim of pad (detected by probe internally). Penial setae not found. Genital marking as a large pad in 17/18 distending both adjacent segments.

Septa and pharyngeal mass absent before 5, septa 5/6–12/13 greatly thickened, thereafter membranous. Gizzard mucular barrel in 5. Dorsal blood vessel single; hearts sinuous in 9–13. Nephridia meroic forests on body wall. Spermathecae paired in 8 and 9 each with flask-shaped ampulla on equally long flat duct with multilocular diverticular frill (inseminated) near base. Probably metandric as paired seminal vesicle seen in 12 only. Testis and ovaries not located, probably minute and lost in musculature of septa and body wall. Prostates rounded but finely incised throughout so not as found in Acanthodrilidae and Octochaetidae (cf. Exxidae), i.e. tubuloracemose with small flaccid ducts in 18. Oesophagus without noticeable dilations (what I initially took as a hemispherical thickening of posterior of 9 was determined as a septum). Intestine substantial yet dilated and easily ruptured, origin appears in 15 or 16. Gut contains finely ground organic matter, organic soil plus coarse multi-coloured grits.

Anterior musculature and thickened septa are associated with strong burrowing, and lack of (anterior) dorsal pores may aid maintenance of hydroskeletal turgor pressure.

Aporodrilus ponga differs from Aporodrilus aotea on almost each specific point. According to Lee (1959), who often took tubuloracemose prostates to be tubular, this specimen keys to genus Megascolides but fails to match any known taxa from there. If more properly allowed into Lee’s Notoscolex the similarity with Notoscolex hakeaphilus Benham, 1949 is remarkable: viz. large size (650–950 mm) with irregular setae, male pores on a median oval depression, septa absent before 5, spermathecae in 8 and 9, and metandry. Presumed differences however, are darker colour (current specimen bleached in alcohol?), epiloby and again much furrowed as here, tufted nephridia (how tufted?) in anterior, a thick-walled enlargement of oesophagus in 8 (possibly as I initially thought was in 9), prostates claimed as flat rather than rounded (although figured as rounded), spermathecae with “a minute globular diverticulum” (variation?), and male pores shown laterally within pad on 18 rather than on its rim as here. It is possible Benham mistook some of these points. His report of last hearts in segment 10 for this species is undoubtedly anomalous as invariably they are in either segments 12 or 13 in normal Megascolecidae; and intestine in 12 is also anterior to what is usual. No mention was made of dorsal pores by him. Benham’s type was collected in 1946 from Kerikeri (A.48.31 – supposedly in poor condition but confirmation from Otago museum unforthcoming) that he thought imported from Australia as was the plant it was found under. This seems unlikely for such a large species: even if its cocoons were introduced, large species often have particular habitats unlike most small to medium cosmopolitans. Lee (1959: 318), presumably accepting Benham’s characterization, has another specimen (current location unknown) from Pukehohe, suburb of Auckland, from subsoil collected by W. Cottier in 1951. (An online GBIF record of Australian Museum AM W.29352 at Taupo is unconfirmed http://data.gbif.org/occurrences/237279142 accessed November, 2011). A much smaller species but with remarkable superficial similarity of marking to Benham’s Notoscolex hakeaphilus is his Notoscolex maoricus (Benham, 1904) (syn. Tokea decipiens Benham, 1905) that also comes from “Waitakerei Bush” (= Waitakere), near Auckland.

Without information to the contrary we must reluctantly accept the balance of Benham’s earlier diagnosis, in which case a new name for this specimen has merit. Confirmation of independence of either species now depends on reinspection of Benham’s type, apparently beyond the brief, budget and resources of successive workers for the last 62 years, including the present one.

Holotype Auckland Museum; AMNZ 5253. Single complete specimen, now dissected, from New Zealand, Cuvier Island (36°26'S, 175°46'E) SE catchment 40–60 m. Coll: 3.IV.2000, J.W. Early & R.F. Gilbert. “Under rock in stream bed. L8229” “W-024” on lid. (Small tissue sample was taken for DNA analysis coded RJB07).

After Maori name for Cuvier Island; Notoscolex is treated as masculine but this place name remains genderless as a noun in apposition.

Notoscolex having spermathecal pores paired posteriorly in 7 and 8 but with clavate spermathecae anteriorly in 8 and 9; holandry with seminal vesicles in 9 and 12; oesophageal gland annular in 12; genital markings mid-ventral in 16/17/18/19.

Body circular. Pale unpigmented in alcohol. Length 135 mm with 149 segments. Prostomium prolobous. Setae lumbricine, 8 per segment in mostly regular rows and almost equidistant throughout. Clitellum not marked. Dorsal pores present but minute and difficult to detect, possibly commencing from 9/10 or 10/11. Nephropores not found (meroic). Spermathecal pores segmental, posteriorly just above intersegments in setal a lines on 7 and 8. Female pores difficult to detect with certainty, possibly mid-ventral pair anterio-median to setae a on 14. Male and prostatic pores combined on small tumescences on 18 in position of deleted setae a. Penial setae not found. Genital markings mid-ventral eye-shaped sucker pads in 16/17, 17/18 and 18/19; a yellowy midventral patch from ½14-16/17 may be artefactual.

Septa increasingly thickening from 4/5–10/11; 11/12 thin and thereafter membranous. Gizzard large but weak in 5. Dorsal blood vessel single; commissurals in 5–8; hearts paired and small in 9, much larger in 10–13; supra-oesophageal vessel not found. Nephridia meroic with several avesiculate tubules almost evenly spaced in several rows on body wall in each segment. Spermathecae in 8 and 9 each with saccular ampulla and single, small clavate diverticulum (non inseminated). Holandric: testes and funnels minute in 10 and 11 ventrally; seminal vesicles paired, racemose in 9 and, larger, in 12. Ovaries paired as fine string masses ventrally in 13; ovisacs not found. Prostates flattened, tubuloracemose extending to ca. 24 from small ducts to male pores in 18. Sessile tumidity associated with genital markings internally. Oesophagus large and folded in on itself in anterior in 6–9 at least; with annular dilated oesophageal gland in 12. Intestinal origin in 16. Typhlosole and caeca not found (absent). Gut contains fine silt with few organic fragments. Intestine paler and concertinaed between 35–40 where several (gregarine?) parasitic cysts on stalks attach to it (see figure).

Habitat location (under rocks in stream) would indicate an aquatic or semi-aquatic life style, a conclusion supported by the pale colouration plus reduced dorsal pores and gizzard; while the folded oesophagus in the anterior would allow considerable extension (for movement and feeding) and the gut contains silty (alluvial?) soil. Alternatively, this specimen may be an unintentional interloper washed into the stream from adjacent soil; more ecological information is needed to confirm or disconfirm this.

Notoscolex is primarily an Australian genus with representatives in Sri Lanka and southern India as well as NZ. The current specimen although large is possibly subadult (it has genital markings but lacks a distinct clitellum and spermathecae uninseminated) yet appears to be a distinct species. Its morphology is comparable to the nine previously known regionally compatriot Notoscolex species, all confined to the north of the North Island, many of which were at some time placed in the cohesive genus Tokea Benham, 1904. This latter genus was made junior synonym of Notoscolex following Michaelsen (1916), Stephenson (1930: 837) and as herein compared to Lee (1959) who placed it within Megascolides.

Of the nine or so New Zealand species now known, Notoscolex repanga differs: from Notoscolex sapidus, Notoscolex urewerae, Notoscolex huttoni and Notoscolex suteri, all by Benham (1904) and each having three pairs of spermathecae; and from Benham’s Notoscolex kirki and Notoscolex maoricus which share two pairs but differ in their arrangements of genital markings and spermathecae. Whereas Notoscolex kirki has intersegmental spermathecal pores in 7/8/9, Notoscolex maoricus has them segmentally but in posterior of 7 and 8, and not 8 and 9 as he originally stated and as inadvertently retained by Lee (1959: 302). [This correction according to Benham (1905: 240, pl XL, figs. 1–2, 8–9) – see http://www.archive.org/stream/transactionsproc38newz#page/240/mode/2up or http://rsnz.natlib.govt.nz/volume/rsnz_38/rsnz_38_00_002970.pdf where he unconventionally records segments 7 & 8 as “7/8” and 8 & 9 as “8/9”; see also http://rsnz.natlib.govt.nz/image/rsnz_38/rsnz_38_00_0736_0000f_ac_01.html for his original figures of Tokea maorica and its junior synonym Tokea decipiens Benham, 1905]. Notoscolex hakeaphilus (Benham, 1949) does have spermathecae in 8 and 9 (with pore locality indeterminate) but differs not least in its large (650 mm) dark body with irregular setae and metandry. Notoscolex napierensis (Benham, 1941) has its four pairs of spermathecal pores equatorial on 6–9, and was probably misdescribed by Benham as having two pairs of tubular prostates on 17 and 19, where the genital markings lie, while Lee (1959: 304), who thought it introduced, accords it a single pair of lobate prostates in 18; dorsal pores were unstated by both authors. A previous Notoscolex member, Notoscolex mortenseni (Michaelsen, 1924) is now moved to Aporodrilus.

Superficially, Notoscolex repanga is somewhat similar to several Megascolides spp., such as Megascolides viridis, Megascolides raglani, Megascolides irregularis, Megascolides alba and Megascolides novaezealandiae, but it differs from all generically by its non-tubular prostates, and specifically by virtue of combination of segmental spermathecal pores and three mid-ventral genital markings, plus an oesophageal gland in 12 only and its last hearts in 13 rather than 12.

A further Megascolides species occurring in NZ and now possibly extinct was described by Schmarda in 1861 under the title of “Hypogaeon orthostichon, ” that is subjected to separate treatment in a forthcoming publication (Blakemore submitted.).

From Unuwhao Mt., nr Spirits Bay, northern extremity of Northland, NZ.

AMNZ 5038 for Megascolex animae has the following label in jar: “Unuwhao nr. Spirits Bay, N.Z. Coll: A.W.B.P. Feb 1946 AM8 1039”. It is a substantial specimen – 196 mm × 13.5 mm – collected by former director of Auckland Museum, A.W.B. Powell in Feb. 1946. Although the specimen in alcohol is now bleached of its earlier dark brown dorsal pigmentation and is wrinkled and hardened, it is yet well preserved. Reinspection of the type largely conforms to Lee’s original except that his figured spermathecae (from 9rhs that he included in a separate vial) is broken off after the diverticulum, and diverticula are newly found to be present in the remaining in situ spermathecae (see figure). The peristomium was not obviously cleft ventrally and genital markings, rarely described by Lee, were not found, but dorsal pores were: small in 4/5 and more obvious afterwards. Specimen AMNZ 5038 is the monotypic holotype fixed by original designation under (ICZN, 1999: Art. 73.1.1); the only slight ambiguity is that Lee (1959: 282) also noted another specimen with “Same data as type material” that has not been subsequently located.

Known only from Great Island, Three Kings Islands. Buckley et al. (2011) claim to have “Megascolides tasmani” (WM82), but this must be a mistake if they could not identify it as clearly belonging in Notoscolex with non-tubular prostates as herein from type inspection (or perhaps they failed to attempt specimen dissections?).

Specimen AMNZ 5039 has the following labels in its jar: “Auckland Museum Coll No. 13 Wet Greywacke gravel. Tasman Stm. Great Island. Three Kings 31.xii.52 J.S. Edwards”; “HOLOTYPE Megascolides tasmani Lee Tasman Stm, Great Is. Col. J.S. Edwards 30/12/52 1021. 3KI3” [note slightly different dates], a further label is blank. Somewhere are two other non-type specimens from the same locality according to Lee’s account (Lee 1959: 314) with one collected “5.I.53”.

The type specimen is dark and brittle, shrunken to 53 mm long. Lee has 67.5 mm with 132 segments but it is a posterior amputee so must naturally be greater (there is also a slight possibility it acquires more setae posteriorly). Dorsal pores, not noted by Lee, are present from 10/11, at least. It appears lumbricine with widely spread setae that converge slightly towards male pores. Lee has overlooked the penial setae which are protruding (due to shrinkage?) from each of the male pores on 18. In 17 in ab lines is a reddish patch that may be a residual genital artefact. The female and spermathecal pores are no longer obvious. Vascularization is mostly as described by Lee, i.e., commissurals are in 6–9, hearts are in 10–12 from dorsal blood vessel that loops between septa in 11, 12, (not 13) 14, 15 and 16 thereafter single (Lee 1959: fig. 327 shows only in 14 and 15). Gizzard appears in 6 rather than 5 but being overlain by tufted pharyngeal glands it is difficult to discern. Seminal vesicles are in 9 and 12 as described. Prostates differ significantly as they are flattened tubuloracemose structures, rather than “tubular, convoluted” as Lee has them; the duct is not traceable in the delicate specimen and, moreover, there is a bundle of long reddish penial setae more ventrally. An excised spermatheca is in a separate vial in the jar and, apart from being desiccated, complies with Lee’s (1959) account and figure. The wide intestine contains organic matter and is full of coarse grits of various minerals.

Having non-tubular prostates qualifies this taxon as a new combination in Notoscolex. The remote chance it acquires extra setae posteriorly after cut would permit it in Anisochaeta. Penial setae are unusual for New Zealand Megascolecidae, but it is interesting that they do not correspond well to the length of the spermathecal diverticula (see Blakemore 2000c; 2008). Further investigation is required for confirmation.

Poor Knights Islands, New Zealand [an online report, along with several other New Zealand earthworms, as a Marine invertebrate from Mexico - http://mexinverts.lifedesks.org/pages/1545 (Oct. 2011) is clearly a mistake].

Two specimens in jar: AMNZ 5040 a larger ~200mm specimen dissected previously, and AMNZ 5280 a smaller complete mature 140 mm long. Labelled “TYPES Notoscolex equestris Benham 1942”; “Notoscolex Equestris Benham 1942”; “TAWHITI RAHI ISLAND, POOR KNIGHTS ISLANDS 26 November 1940 G.A. Buddle, R.A. Wilson, E.G. Turbott”.

There is some slight confusion with Notoscolex equestris Benham, 1942, in that Lee (1959) erroneously placed it in Megascolides, and Lee (1959: 296) said types were in Otago Museum (No. A.43.52 - two specimens in fair condition but confirmation from Otago museum unforthcoming), yet he also gives “4 specimens. (Auckland Museum Collection)”. There are indeed two specimens in the Auckland Museum (pers. obs.) viz.: AMNZ 5040 with labels as above. Benham (1942: 220 - see http://rsnz.natlib.govt.nz/volume/rsnz_72/rsnz_72_03_002070.pdf) actually stated that Mr R.G. Turbott of the Auckland Memorial Museum had sent him two phials, one from Chatham Island that contained four earthworms, and these other two larger specimens collected from Poor Knights Islands by Majors G.A. Buddle and R.A. Wilson.

Both specimens are here inspected and described: the larger one – that entirely agrees superficially with Benham’s figures – had been previously dissected with the 8lhs spermatheca, 18rhs prostate, and the anterior of the intestine removed and missing from the jar. Additions to Benham’s and Lee’s earlier descriptions are that the highly wrinkled prostomium is construed as pro-epilobous rather than prolobous, and no ventral cleft is present on the peristomium. Benham was “unable to detect the dorsal pores owing to the strongly contracted state of the body” and, for some reason, Lee omitted mention of them entirely except for exotic Lumbricidae. They are here confirmed as being absent throughout the body in both specimens (i.e., qualifying for Aporodrilus). Setae c and d are increasingly irregular. Spermathecae are in 7–9 but for 8lhs only the stub remains with the small diverticulum still attached (hence overlooked by earlier workers who also mistook slight folds in the soft duct as “excrescences”); as for other spermathecae, the small diverticula are visible by their slight iridescence just above the body wall at the base of the duct [see Fig. 6 and cf. Benham (1942: fig. 5), Lee (1959: fig. 308)]. Only the prostate 18lhs remains and is here construed as cylindrical tubulo-racemose i.e. non-tubular [see Fig. 6 and cf. Benham (1942: fig. 4)]. Genital markings agree as per original (Benham 1942: fig. 3) and the smaller undissected specimen (AMNZ 5280) is provided with a rough sketch showing how it conforms too. The gizzard appears more in 5 than 6 and oesophageal dilations are increasingly large in 10–14 (at least) but, as gut is removed, the intestinal origin cannot be confirmed. Although a typhlosole is absent and it is noted that the intestine below the break is filled with particularly coarse plant fragments only (no soil).

Both specimens are surely syntypes (one dissected agrees and key organs removed suggests they were figured by Benham, although Lee also dissected a prostate) and, under ICZN (1999: Art. 74) I hereby expressly designate the larger dissected specimen AMNZ 5040 the lectotype of Aporodrilus equestris (Benham, 1942) leaving the remaining undissected specimen as paralectotype (AMNZ 5080). In compliance with “Declaration 44 – Amendment of Article 74.7.3 of ICZN” (1999 – see http://iczn.org/content/declaration-44-amendment-article-7473), this act is in order to provide stability in its taxonomic name coupled with the augmented description provided herein. Enquiries made to verify Otago Museum material (Email: cody.fraser@otagomuseum.govt.nz 15^th Oct., 2011) were fruitless, but it is probable Lee (1959: 296) in his account confused the two lots that were sent to Benham, as commented on above.