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A new species of Macquaridrilus(Annelida: Clitellata: Naididae) fromsubantarctic Campbell IslandAM Pindera & A Jamesb

a Department of Parks and Wildlife, Science and ConservationDivision, Kensington, WA, Australiab EOS Ecology, Christchurch, New ZealandPublished online: 19 Mar 2014.

To cite this article: AM Pinder & A James (2014): A new species of Macquaridrilus (Annelida:Clitellata: Naididae) from subantarctic Campbell Island, New Zealand Journal of Zoology, DOI:10.1080/03014223.2014.884508

To link to this article: http://dx.doi.org/10.1080/03014223.2014.884508

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RESEARCH ARTICLE

A new species of Macquaridrilus (Annelida: Clitellata: Naididae) fromsubantarctic Campbell Island

AM Pindera* and A Jamesb

aDepartment of Parks and Wildlife, Science and Conservation Division, Kensington, WA, Australia; bEOS Ecology,Christchurch, New Zealand

(Received 19 August 2013; accepted 7 November 2013)

Macquaridrilus mcmurtrieae n. sp. is described from Campbell Island. This resembles the only otherspecies in the genus, Macquaridrilus bennettae Jamieson, 1968, in most aspects, but shows significantdifferences in the anatomy of its genitalia. In particular, the spermathecal pores are dorsal rather thanlateral, the spermathecae lack diverticulae, the ejaculatory duct is more stout and muscular, the vasdeferens is shorter relative to other organs and the anterior prostate is compact rather than elongate.The presence of a cuticular sperm canal appears to be an apomorphy for the genus. The new specieswas collected from streams and tarns across the island.

http://zoobank.org/urn:lsid:zoobank.org:pub:652AF61D-CFB2-4D07-94C8-59E6FB549D5F

http://zoobank.org/urn:lsid:zoobank.org:act:984F2456-768D-48A1-87AD-4453768BAB8A

Keywords: Annelida; Clitellata; Naididae; oligochaete; invertebrate; new species; Campbell Island;subantarctic; aquatic; freshwater

Introduction

In 1965 Professor Barrie Jamieson, from theUniversity of Queensland (Brisbane), collectedoligochaetes on Macquarie Island. Amongst thematerial in that collection of mostly terrestrialearthworms was an aquatic oligochaete, collectedfrom a single pond by Isobel Bennett, withsufficiently unusual genital anatomy for Jamiesonto establish the genus Macquaridrilus, withinwhich he placed the new species Macquaridrilusbennettae (Jamieson 1968). This species wassubsequently collected from Macquarie Islandin 1978 (unpublished records from material heldin South Australian Museum, collector PennyGreenslade) and 1992 (Marchant & Lillywhite1994). Collections from Campbell Island, made byan expedition from the Australian Museum andborrowed by the senior author in 1992, includedworms with very similar chaetae to those of

M. bennettae, but these were not examined furtheras the focus at that time was on the Phreodrilidae(Pinder & Brinkhurst 1997). Twenty years later,new collections from Campbell Island alsoincluded specimens resembling M. bennettae andthese are described herein as the new speciesMacquaridrilus mcmurtrieae. The latest collectionswere made during the 2010–2011 CampbellIsland Bicentennial Expedition, a multidisciplinaryexpedition designed to uncover the island’s humanhistory, ecosystem processes and recovery follow-ing eradication of rodents and grazing mammals(www.campbellisland.org.nz).

Material and methods

Specimens were collected by EOS Ecology as partof their survey of the aquatic macroinvertebratecommunities of the island’s streams and tarns in

*Corresponding author. Email: adrianp@dpaw.wa.gov.au

New Zealand Journal of Zoology, 2014http://dx.doi.org/10.1080/03014223.2014.884508

© 2014 The Royal Society of New Zealand

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2010–2011. A total of 25 stream sites and ninetarns were sampled (Fig. 1) and photographsof representatives of these are shown in Fig. 2.At each stream site six Surber samples (0.25 m2,500 µm mesh size, Fig. 2D) and two kick-netsamples (each a composite of five ‘kicks’, 500 µm

mesh size) were collected. At tarn sites four kick-net samples were taken (including benthic habitatsand sweeps of the water column, 500 µm meshsize). All samples were preserved in the field in4% formalin, except for one kick-net sample fromeach stream site, which was preserved in 95%

Figure 1 Map of Campbell Island showing locations of sampled tarns (squares) and streams (circles). Opensymbols are locations where M. mcmurtrieae may be present but from which some or all samples are yet to beprocessed (see Methods) and solid symbols are locations from which M. mcmurtrieae has been identified.

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ethanol. On their return to mainland New Zealandall formalin samples were washed and re-preservedin 70% isopropanol alcohol.

Oligochaetes were identified from 25 samples(from 12 stream sites and six tarns) out of the 218samples collected. Thirteen stream sites and threetarns have not yet had any of their oligochaetesexamined and no sites have had oligochaetesexamined from all samples. All specimens were

examined in ethanol and a selection of matureworms were stained in Grenacher’s borax carmine,partially destained in 3% hydrochloric acid, dehy-drated in isopropanol, cleared in Histoclear™ andthen slide mounted in Permount™. Slide mountedspecimens were either mounted whole or thegenital segments were dissected. Drawings weremade with the aid of a drawing tube. Measurementswere made using an eyepiece graticule or from a

Figure 2 Photographs showing a variety of habitats from which M. mcmurtrieae was collected. A, Site 4, HoneyFalls Stream; B, site 21, Kirk Stream; C, site 12, Middle Bay Stream; D, site 25, Southeast Stream (ShelleyMcMurtrie of EOS Ecology taking a Surber sample); E, site NP1, Tucker Tarn (Jo Hiscock of the New ZealandDepartment of Conservation taking a water sample); F, site BG1, one of the Borchgrevink Bay Tarns. Allphotographs copyright EOS Ecology.

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calibrated measuring tool in the AutoMontage™5.01(A) software produced by Syncroscopy.

Habitats

Macquaridrilus mcmurtrieae is widespread onCampbell Island, where it has been recordedfrom seven stream reaches and two tarn sites todate out of 12 stream sites and six tarns fromwhich oligochaetes were examined (Fig. 1). Thisis almost certainly an underestimate of its occur-rence considering the proportion of samplesexamined to date (see Methods). The stream sitesgenerally had coarse substrates dominated bybedrock and boulders (> 256 mm), large cobbles(128–256 mm) and small cobbles (64–128 mm).Pebbles (2–64 mm) dominated the substratum atone site (site 21: Kirk Stream). Water velocitieswere moderate (0.36–0.56 m/s), with channelmorphology typically being some combinationof pool, step pool, cascade, run and riffle. Thinmats of brown algae covered the substratum at allsites. Riparian vegetation was either dominated byDracophyllum forest (e.g. Fig. 2A, D), tussocks(e.g. Fig. 2B, F) or a mixture of Dracophyllum,ferns and tussocks (e.g. Fig. 2C, E). Canopy covervaried widely from < 5% to 100%. The water ofall streams was tannin stained (1.1–2.5 mg/L)with acidic to neutral pH (5.1–7.0) and specificconductivities that do not indicate any great salineinfluence (158–713 µS/cm). Spot measures of watertemperature (7.4–10.8 °C) and dissolved oxygen(93.8%–98.9%) indicated cool, well-oxygenatedconditions. Tarn NP1 (Fig. 2E) was in a protectedcove and surrounded by Dracophyllum and tus-socks, was approximately 8 m across and 14 m longand ≤ 0.8 m deep. It had steep sides and a peatsubstrate. Tarn BG1 (Fig. 2F) was located near thetop of an exposed high cliff in the north of CampbellIsland, with the adjacent vegetation being tussocksand megaherbs. It was approximately 10 m wide,30 m long and ≤ 0.5 m deep, with a sand and rocksubstrate. NP1 was tannin stained (2.8 mg/L) whileBG1 was not (0.1 mg/L) and both were acidic(pH 5.2 and 4.1 respectively). The specific conduct-ivity of NP1 (574 µS/cm) was within the range of

the stream sites, while BG1 (4810 µS/cm) wasinfluenced by windblown salt spray.

Taxonomic account of the new species

Phylum

Annelida

Class

Clitellata

Family

Naididae Ehrenberg 1828

Genus

Macquaridrilus Jamieson 1968Macquaridrilus mcmurtrieae n. sp.(Figs. 3–5)

Figure 3 Chaeta from segment VII, with enlargementof bifid tip. Scale for whole chaeta 10 µm.

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Material examined

All collections by Shelley McMurtrie (SM) andAlex James (AJ) of EOS Ecology, or Colin Meurk(CM) of Landcare Research New Zealand. Typematerial has been deposited in Museum of NewZealand Te Papa Tongarewa in Wellington. Alllocations are on Campbell Island and all coordinatesdetermined using the NZ Geodetic 2000 datum.Holotype: W.003476, Southeast Stream (site 25),Surber sample, mature specimen mounted whole onslide, 28 Jan 2011, 52°35.441′S, 169°10.313′E (SMand AJ). Paratypes: W.003483, Southeast Stream(site 25), Surber sample, 1 mature and 2 immature inethanol, 28 Jan 2011, 52°35.441′S, 169°10.313′E(SM and AJ); W.003478 and W.003478/1, GardenStream (site 11), Surber sample, 3 mature on twoslides, 27 Dec 2010, 52°34.205′S, 169°08.339′E(SM and AJ); W.003479, Middle Bay Stream (site12), Surber sample, 1 mature on slide in Permount,29 Dec 2010, 52°34.205′S, 169°08.339′E (SM

and AJ); W.003480, Kirk Stream (site 21), Surbersample, 1 on slide in Permount, 11 Jan 2011, 52°35.044′S, 169°08.329′E (SM and AJ); W.003482(slide) and W.003482/1 (vial), Tucker Tarn (siteNP1), kick-net sample in open water and macro-phytes, 1 mature on slide (head and tail in Kaisersglycerin jelly andmid-body stained in Permount) and2 mature in ethanol, 2 Feb 2011, 52°32.806′S, 169°08.644′E (SM); W.003477, W.003477/1, W.003477/2 and W.003477/3, Borchgrevink Bay Tarns (siteBG1), kick-net sweep, 4 mature on three slides inPermount and 2 mature in ethanol, 31 Jan 2011,52°28.541′S, 169°11.538′E (CM); W.003481 andW.003481/1, Capstan Cove Stream (Site 13), kick-net sweep, 2 on two slides, 31 Dec 2010, 52°33.094′S, 169°05.573′E (SM and AJ).

Additional material

All returned to EOS Ecology, Christchurch.Lookout Bay Stream (site 9), Surber sample, 1 in

Figure 4 Male genitalia (segment XI) of M. mcmurtrieae, based on the holotype: a, atrium; b, bursa; ed, ejaculatoryduct; mf, male funnel; mu, muscle bands covering bursa; pr, prostate gland attached to atrium; sc, sperm canal; v,vas deferens. Scale=50 µm.

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ethanol in mixed species vial, 22 Dec 2010,52°32.609′S, 169°09.423′E (SM and AJ); HoneyFalls Stream (site 4), Surber sample, 1 tail end only, 15Oct 2010, 52°33.263′S, 169°10.581′E (SM and AJ).

Description

All features are as observed for specimens killedin formalin or ethanol and then preserved inalcohol.

Figure 5 Photos of M. mcmurtrieae genitalia. A, Ejaculatory duct (left) entering the copulatory bursa (right) insegment XI: ed, ejaculatory duct; mu, muscle bands covering bursa; cg, copulatory gland cells; sc, sperm canal; B,spermatheca in segment X: sd, spermathecal duct; sa, spermathecal ampulla. Scales: A=30 µm; B=50 µm.

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Dimensions

Length of whole worms 5.5 to 10 mm, width at VIto 0.3 to 0.35 mm, width at XI of mature worms0.35 to 0.45 to mm, number of segments 32 to 38.

External features

Prostomium bluntly conical in dorsal/ventral view,sometimes more squarish in lateral view. Anteriorchaetae bifid crotchets with upper teeth muchshorter than lower, posterior chaetae with uppertooth minute or absent, all chaetae sigmoid andwith indistinct nodulus approximately medial.Anterior chaetae 60 to 70 µm long (measured asa straight line between base and tip) and 3 µmwide in II (at nodulus), increasing in length insubsequent few segments, so that chaetae alongmost of body 85 to 105 µm long and 4 µm wide atnodulus. Number of chaetae 3 to 7 (mostly 4 or 5)per bundle. Clitellum from 10/11 to about thelevel of the chaetae on XII, with about three-quarters of the cells being glandular and the resthyaline. Crescent-shaped ventro-lateral male pores(exterior opening of the bursae) on postero-lateraledges of a broad ventral depression which runs thelength of XI. Opening of bursae with particularlylarge and protruding epithelial cells on the lateralwalls. Female pores within intersegmental furrow11/12. Spermathecal pores above the line of thedorsal chaetae and about two-thirds of the distancebetween 9/10 and 10/11.

Internal somatic features

Pharynx in III with numerous muscular connec-tions to body wall in II–V. Pharyngeal glanddevelopment variable but primarily ventro-lateralto lateral in clumps from IV to posterior of septa5/6. No coelomocytes observed. A layer ofchloragogue cells covering gut from VI. Oeso-phagus expanded and with taller lining cells inVIII, IX and sometimes X. Post-clitellar intestineintrasegmentally constricted (at about a third ofthe distance from anterior to posterior of thesegment), as well as at each septa. In most of thespecimens examined there are parasitic ciliatespartially embedded in the walls of the fore-gut,

especially in segments VIII to X. Paired nephridiapresent in VII to IX with lateral pores anteriorto ventro-lateral chaetae, then again in mostsegments between XV and XVIII.

Genitalia

Small antero-ventral testes anteriorly in X. Ovarieslarger than testes and present anteriorly in XI. Eggsacs extending posterior as far as XIV. Sperm sacsnot observed but a few sperm morulae sometimespresent in coelom of IX and X.

Sperm funnels 50–65 µm across at the flare(with cilia projecting forward by up to 40 µm),narrowing to 20–30 µm behind the flare, thennarrowing further to 10 µm by the time it becomesthe ental vas deferens. The ciliated vas expandingto the width of the atrium as it passes (or passesthrough) the prostate to join the atrium. Atrium170–190 × 20–30 µm, consisting of a thin layerof peritoneal cells around a strongly vesiculatedepithelial lining with cell walls often not visible,around a narrow ill-defined lumen. A compactprostate gland appears to be broadly attached tothe ental portion of the atrium, although theprecise nature of the connection(s) could not bedetermined. Atrium joining the ental end of anejaculatory duct. Entally, the latter is a duct ofabout 80 µm in length and 15–20 µm in width,with well developed circular muscle and epithelialcells around a narrow lumen. This duct thenexpands into a spindle-shaped organ 180–240 µmlong and 55–70 µm wide with a thick (12–15 µm)layer of spiral muscle fibres between a layer ofperitoneal cells and epithelial cells with cuticularwalls: the basements of the latter forming a well-defined line inside of the muscle layer. Epithelialcells directed posteriad and strongly elongated andtapered at their extremity, especially from themiddle to ectal part of the ejaculatory duct, so thattheir apical walls combine to form a cuticular tube(‘sperm canal’ sensu Jamieson 1968) with medialwidth of 13–16 µm that projects into the anterio-medial wall of the dorsal chamber of the bursae.The epithelial lining cells of the ectal-most partof the ejaculatory duct are not elongated and arecontinuous with the epithelial tissue of the bursa to

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which it connects. There are muscle fibres encirc-ling the region on the ejaculatory duct where itenters the bursa. The nature of the ectal end of thesperm canal cannot be discerned in the typematerial.

The internal structure of the bursa is difficultto discern from dissected or whole-mountedworms, but it appears to be at least two-chambered(or at least with the walls strongly folded inwardsto form constrictions). It has a layer of epithelialcells surrounded by multiple layers of musclefibres differing in their orientation. Additionally,there are muscle fibres that run in parallel from theventral to dorso-lateral body wall and anchor theentire bursa to the lateral body wall. The upperhalf of the bursa is covered by diffuse accessorycopulatory gland cells (sensu Erséus & Jamieson1981) underlying the muscle fibres. In some of theexamined specimens the bursa are protruded fromthe body a distance of about half of the bodywidth. Within this protruded bursa, the ectal partof the muscular ejaculatory duct can be seen,though it is not clear how close the cuticularsperm canal then is to the opening of the bursa.

Spermathecae short (115–155 µm × 35 µm),hardly differentiated along their length other thana constriction at the pore and thinner epithelium atits ental extremity. Each spermatheca entirelycovered by a thick (5–6 µm) layer of longitudinalmuscle covered by a single layer of low peritonealcells. Glandular cells are clustered around the baseof the duct at the pore. For the ectal two-thirds tothree-quarters of the spermathecae (this part beingequivalent to a duct) the epithelial layer is broad,leaving only a small lumen. The epitheliumbecomes much thinner in the ental portion (theampulla). Sperm in the ampulla is loose andoccupies the entire lumen and there are oftensperm lying along the lumen of the duct. Thespermathecae lack a diverticulum.

Etymology

Named for Shelley McMurtrie, from EOS Eco-logy, New Zealand; leader of the freshwater teamon the 2010–2011 Campbell Island BicentennialExpedition.

Discussion

With its elongate atria, each with a prostategland and well developed muscular ejaculatoryducts terminating in a cuticular sperm canal whichprotrudes into copulatory bursae covered byglandular cells, the new species appears to bemost closely related to M. bennettae Jamieson1968. The general arrangement and histology ofthe genitalia of the Campbell Island worms is verysimilar to that of M. bennettae, but there aresufficient differences in the genitalia to be certainthat they are a separate species. In particular, theprostate runs along the length of the atrium inM. bennettae, whereas it is compact and ovoid totriangular in M. mcmurtrieae and certainly notpresent all along the atria. The ejaculatory ductis much broader in relation to its length inM. mcmurtrieae. The most conspicuous differenceis the much simpler spermathecae. WhereasM. bennettae has a small diverticulum joiningthe spermathecal duct near its base, this is lackingin M. mcmurtrieae. Furthermore, the spermathecalduct is much narrower than the ampullae inM. bennettae and leads to lateral pores, whereasin M. mcmurtrieae the ampulla is hardly widerthan the duct and the latter leads to dorsal pores.

Placing this new species in Macquaridrilusrequires two modifications to the definition ofthe genus as defined by Jamieson (1968): Thespermathecae can possess (bennettae) or lack(mcmurtrieae) diverticulae and have pores thatare lateral (bennettae) or dorsal (mcmurtrieae).

Erséus & Jamieson (1981) described twoAustralasian genera Heronidrilus and Macquari-driloides and allied these with Macquaridrilus onthe basis of similarities in the genitalia. Erséus(1990) later synonymised Macquaridriloides andHeronidrilus and suggested that Macquaridrilusbennettae could belong in the same group. There iscertainly little to distinguish Macquaridrilus fromHeronidrilus other than thatMacquaridrilus speciespossess the cuticular sperm canals and have prostatetissue that is more restricted in its covering of theatria. Both characters may be synapomorphies forthe two species irrespective of what larger groupthey belong to.

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Jamieson (1968) suggested that M. bennettaemight only recently have reachedMacquarie Island,probably via other southern landmasses such asCampbell Island. This was on the basis that: (1) itwas a freshwater species; (2) it appeared at that timeto be related to freshwater genera such as Tubifexand Limnodrilus; (3) it seemed to have a restricteddistribution on Macquarie Island (it is now knownto be widespread; Marchant & Lillywhite 1994);and (4) there may not have been time since retreat ofPleistocene glaciers to have allowed speciationfrom a colonising ancestor (the island is nowknown to have been partly ice-free during thelast glacial maximum; Selkirk et al. 1990).Canvassed dispersal mechanisms were rafting andavian transport.

Subsequent discovery of the clearly alliedmarine worms mentioned above led Erséus &Jamieson (1981) to suggest that M. bennettae mayhave had a marine ancestry. There are certainlyother tubificoid Naididae in the subantarctic/ant-arctic marine benthos, including Torodrilus Cook,1970, which Erséus & Rota (1996) suggested mayalso be related to Heronidrilus. The MacquarieIsland and Campbell Island Macquaridrilus may,therefore, have had a common marine ancestorthat colonised the limnic habitats of each island.Marchant & Lillywhite (1994) noted that multipleinvasions of subantarctic island freshwaters areknown for some other marine lineages, such assome janirid isopods (Wilson & Wagele 1994) andplanarians (Ball & Hay 1977). A hybrid hypothesisis that a marine ancestor of these Macquaridrilusspecies made a single invasion of freshwater anddescendants have then island hopped and speciated.For some other oligochaetes, non-marine dispersalbetween subantarctic islands seems much morelikely. Astacopsidrilus campbellianus Benham,1909 belongs to a family (Phreodrilidae) that isalmost entirely restricted to freshwater but occurson both Campbell Island and Macquarie Island(Pinder & Brinkhurst 1997). Similarly, the cosmo-politan naidid Nais elinguis Müller, 1774, whichalso occurs on both islands (Marchant & Lillywhite1994 and unpublished data), often occurs in brack-ish water but rarely in marine waters. There isgrowing evidence for avian transport of aquatic

invertebrates (Green & Figuerola 2005; Green et al.2008) and given the large numbers of seabirdsusing oceanic islands this method of dispersal foroligochaetes seems likely.

AcknowledgementsThanks to Shelley McMurtrie and Amber Sinton at EOSEcology for making the Campbell Island worms availableto AMP and for providing funds for their identification.Thanks also to Professor Christer Erséus of the Universityof Gothenburg in Sweden for attempting to obtainmolecular genetic information that would have helpedunderstand the phylogenetic affinities of this species.The Campbell Island Bicentennial Expedition was madepossible by the 50° South Trust (www.50south.org.nz),established to further research and education on NewZealand’s subantarctic islands. Processing of the benthicaquatic invertebrate samples was supported by EOSEcology and a Terrestrial and Freshwater BiodiversityInformation System grant (TFBIS 278).

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new monocelid flatworm from Macquarie Island(Platyhelminthes, Turbellaria). Bijdragen tot deDierkunde 47: 205–214.

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Erséus C, Jamieson BGM 1981. Two new genera ofmarine Tubificidae (Oligochaeta) from Australia’sGreat Barrier Reef. Zoologica Scripta 10: 105–110.

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