1\1'10rds ottlle We.s!<'1II AIIs!mlillll MII.seulil 24: 459-461'\ (2001'\).

Phreodrilidae (Clitellata: Annelida) in north-western Australia with descriptions of two new species

Adrian M. Pinder Science Division, I)epartment of Environment and Conservation, PO Box 5\, Wanneroo, Western Australia 6946, Australia

Abstract - Two new species of oligochaete (Clitellata: Annelida: Phreodrilidae) are described from subterranean waters of the Pilbara and Cape Range regions of Western Australia. Pllrcodrilus Iillllllci sI'. novo from the Pilbara, most closely resembles Pllreodrilus penicillus Pinder 2003, also from the Pilbara, but is less widespread. Inslllodrilus IlIlgcla sI'. novo occurs primarily in groundwater, especially within the Ashburton catchment of the Pilbara, but was also collected from two Pilbara springs and from a cave on Cape Range. These, and other undescribed phreodrilids, suggest that north-western Australia, despite being an arid zone, is surprisingly rich in this Condwanan family.

INTRODUCTION aquatic invertebrates of the Pilbara (Pilbara Thirty two species of Phreodrilidae (in 6 Biologica I Su rvey [PBSL Depa rtment of genera) have been described from Australia to Environment and Conservation: Eberhard et al. date (pinder 2003b) but at least as many species (in press) and unpublished data), the Monitoring again await description. The family Phreodrilidae of River Health Initiative (MRHI) and surveys by is common in streams and lentic wetlands of Dr W.F. Humphreys of the Western Australian temperate southern Australia and particularly Museum. diverse in Tasmania and south-western Australia (Pinder and Brinkhurst 1997). The concentration METHODS of species in the most temperate parts of Australia Groundwater specimens were collected using fits with the idea that the family is Gondwanan in haul nets as described in Eberhard et al. (in press) origin (Martin et al. 2008, Pinder and Brinkhurst and preserved in 100'/') ethanol. For the PBS bore 1997). However, while the family is far less sites, oxygen, redox and pH were measured in-situ widespread in more xeric parts of Australia, using a Yeo-Cal meter and ionic composition and new species are increasingly being collected total dissolved solids were analysed from samples from groundwater and groundwater associated collected using a bai ler. Surface water specimens wetlands outside of the temperate zones. Pinder were collected with a 250).101 mesh sweep net (2003a) described the first endemic aquatic and preserved in 100% ethanol. Specimens from oligochaete, Pllreodrillls penicIIIIIs, from the arid New Mowbowra Cave were collected by hand north-west of Western Australia. Examination from floating root mats. Specimens were stained of additional material has revealed the presence in Crenacher's borax carmine, dehydrated in of numerous other oligochaetes in the region, isopropanol, cleared in Histoclear and slide including some new stygobiotic species. These mounted in l'ermount. Drawings were made are part of a regional stygofauna that, with> 350 with the assistance of a drawing tube and species in the Pilbara, is particularly diverse by measurements were made using Auto-Montage world standards (Eberhard et al. 2005; Eberhard Pro 5.01(A) (The Synoptics Croup 2003) with a JVC et al. in press; Humphreys 1999; Humphreys cl KY-F1030 video camera calibrated with a stage al. 2005). In this paper, two new phreodrilids micrometer. are described and the distribution of the family Type material is deposited with the Western in the north-west is discussed. Undescribed Australian Museum (WAM) and additional phreodrilids mentioned in the discussion are material is held by the Department of Environment coded as Plzreodrillls WA12, PIzreodril1l5 WA32, and Conservation (DEC). C;eographic coordinates In5111odril1l5 WA35 and phreodrilid WA36 and will were measured with a hand-held Cl'S using be described at a later stage. datum CDA94, except for New Mowbowra Cave This material was collected during several which was either AGD66 or ACD84. projects: a survey of epigean and hypogean The following abbreviations are used 460 A.M. Pinder

Insulodrilus angela from groundwater...or surface water A

Insulodrilus WA35 from groundwater • or surface water. Phreodrilidae WA36 from groundwater * Immature Insulodrilus from groundwater X or surface water x

to

125 km

Phreodrilus peniculus from groundwater • or surface water • Phreodrilus Iinnaei Phreodrilus WA32 A Phreodrilus WA12

to

125 km

Figure 1 Map showing collection localities for lnsulodrilus spp. and an unidentified phreodrilid (top) and Phreodrilus spp. (bottom). Solid lines represent drainage basins as named. Immatures tentatively allocated to lnsulodrilus or Phreodrilus on the basis of the form of the ventral chaetae. Phreodrilidae in north-western Australia 461

throughout the text: a, atrium; bd, broad portion Diagnosis of spermathecal duct; cm, circular muscle The combination of a protrusible pseudopenis on spermathecal duct; ed, ectal portion of and a small pendant penis in P. lillllnei is spermathecal duct; H, female funnel; md, median otherwise found only in P. pelliclllllS, also from portion of spern,athecal duct; mf, male funnel; p, the Pilbara region. However, P. pClliclllllS lacks an pendant penis; pp, pseudopenis; sa, spermathecal enlarged mid-portion of the spermathecal duct, ampulla; sd, spermathecal duct; sv, spermathecal has a broad uncoiled pseudopenis and has a larger vestibule; v, vas deferens. Roman numerals body (up to 520 flm at X), larger anterior ventral denote segment numbers while numbers in chaetae (80-120 pm) and shorter maximum hair Arabic separated by a slash refer to septa between length (up to 325 flm). The complex spermathecal segments. ducts of P. lillllnci, with the widened medial portion, are unique within the family. SYSTEMATICS Description Family Phreodrilidae Beddard 1891 Length unknown, all specimens with posterior Phreodrilus Beddard 1891 missing; width at III 200-210 pm and at XI 310­ 350 flm. Prostomium bluntly conical to rounded. Plireodrillls Beddard 1891: 273. Pharynx heavily glandular and muscular in I and Type species n, oesophagus thin walled in Ill-VIII, enlarging Plireodrillls sllbtcrrnllells Beddard 1891, by into intestine in VIII. Pharyngeal glands not original designation. heavily developed, located ventral and sometimes lateral in IV to V or VI. Remarks Ventral chaetae paired from H, one of each Plireodrillls was established for P. sllbterrllllells by pair bifid, slightly sigmoid, with rudimentary Beddard (1891) from groundwater in New Zealand upper tooth and distinct distal nodulus, the other but is now also known from another groundwater sharply single-pointed, straighter and without species in New Zealand, five from Australia (all nodulus, both 72-85 pm in anterior segments but one epigean) and one groundwater species (Figure 20). Ventral chaetae absent on XII, present from the Arabian Peninsula (Martinez-Ansemil but unmodified on XIII. Dorsal chaetae dorso­ et II/. 2002). The genus is one of the few currently lateral from Ill, each bundle with 1-2 smooth recognised phreodrilid genera with a distinct hairs, each surrounded by a pair of support apomorphy, the coiled protrusible pseudopenes chaetae that do not emerge from the body wall. within a muscular sac, which can be everted to Hairs 180-524 pm long and 1.5 to 2.5 pm wide form a copulatory organ greater than the body at bodv wall, tapering to a fine tip, generally width. Pllreodrillls pelliclllllS was described from increa~ing in size posteriorly over first few springs in the Pilbara (Pinder 2003a) and has segments. subsequently been collected from groundwater in Clitellum from 1/2XIl to end of XIII, c1itellar the same region (Figure 1). epithelium 2-3 times width of somatic body wall and consisting of irregular ovoid to rectangular Phreodrilus linnaei sp. novo cells. All genitalia and pores paired. Male pores Figure 2 ventro-Iateral halfway between 11/12 and 12/13 in line with the ventral chaetae. Female pores Material examined and ducts not visible on types. Dorso-Iateral slit­ Holotlfpe shaped openings of the spermathecal vestibulae Australia: Western Australia: Main Roads in intersegmental furrow 12/13, in line with Department groundwater bore HMBR#1 on dorsaI chaetae. Ilamerslev Mount Bruce Road about half way Testes and ovaries antero-ventro-Iateral in betvveen Karijini Drive and Nanutarra-Munjina XI and XII respectively. Developing sperm Road (site PSS043 of PBS), Fortescue River present from VI to XII. Ventro-Iateral cup-shaped catchment, 2T30'06.3"S 117 c 57'38.3"E, 21 November sperm-funnels on 11/12, each narrowing to a 2002, collected using a haul net, M. Scanlon, J. vas deferens of 8-13 pm width which enters Cocking (WAM V7430; slide mounted). pseudopenial sac entally where sac meets atrium. Atrium (Figure 2A-a) consisting of a short blunt Partlllfpes (85-108 pm x 24-50 pm) 'tail' on the pseudopenial sac with uneven lining tissue, transforming into Australia: lVestern Australia: 2 specimens, a pseudopenis (i.e. with lining tissue dissociated collection details as for holotype (WAM V7431, from muscular sac wall) just before joining the V7432; slide mounted). vas deferens. Eversible pseudopenis (Figure 462 A.M. Pinder

A

mf

B

~ md

c

D

Figure 2 Phreodrilus linnaei sp. nov.: A, male genitalia, based mostly on holotype other than ental atrium which is an interpretation from partly obscured and folded atria of the 3 type specimens; B, spermathecal duct of holotype; C, genitalia of holotype in situ, 0, ventral chaetae of holotype. Scale lines: 50pm. Phreodrilidae in north-western Australia 463

2A--pp) 15-30 pm wide, loosely coiled within sac to pendant penes, they nonetheless perform a and with ciliated lumen about' I width of duct. penis-like function of sperm transfer and the term Sac up to 115 pm wide, narrower at either end, protrusible pseudopenis remains apt. ectally forming a duct (20-40 pm wide) containing The small pendant penis present in all the (now unciliated) pseudopenis, the whole duct specimens of P. pellicIIIIIS and P. lillllaei does not protruding into the penial sac to form a short appear to be the same as the partially everted (15-45 pm long x 25-26 pm wide) double-walled pseudopenis that is occasionally seen in other pendant penis occupying 1/; to I of the penis Plireodrillls specimens, e.g. as illustrated for sac Plzreodrillls stocki (Martinez-Ansemil cl al. 2(02). Spermathecal ampullae (Figure 2B-sa) elongate, Water sampled from the type locality was with walls mostly consisting of lining tissue (4-6 neutral freshwater dominated by sodium and pm), muscle not visible, and loosely folded in bicarbonate (Table 1). XIV and XV, filled with sperm in loo~e masses in mated individuals, Ampulla narrowing ectally Etymology to form a short duct (Figure 2B-md) of varying Named in honour of Carolus Linnaeus (1707­ width (19-33 pm) and length (130-160 pm), with 1778) on the 250th anniversary of the publication a narrow lumen, broader lining tissue and a of the 10 th edition of Systelllil Natllme and the 300th circular muscle layer. The latter opens into a anniversary of his birth. broader glandular section of duct (180-270 x 40-85 pm) with large lining cells and a very thin muscle InSlllodrillls Brinkhurst 1991 layer (Figure 2B-bd), which then narrows to form a loose duct (width 25-3.5 pm) (Figure 2B-ed). IIlSlllodrillls Brinkhurst 1991: 2040. This duct covered by longitudinal muscles in its ectal portion which branch off as long thin fibres Type species (Figure 2B-mf) that join the body wall at various P!zreodrillls lacllstris Benham 1903, bv original points. designation.

Remarks Remarks The term protrusible pseudopenis was used by Illslllodrillls was established following a Brinkhurst (1965) to describe the portion of th~ phylogenetic analysis by Brinkhurst (1991) for male genital ducts that is a slightly to strongly species with pendant penes and with female coiled and ciliated tube within a thin-walled sac pores not located within the (generally small) of P!zreodrillls species, This tube is everted from spermathecal vestibulae. However, Brinkhurst the male pore during copulation. This has been noted that this grouping was probably thought of as a modification of the ectal part paraphyletic and only an interim 'conservative of the atrium, which is normally a glandular measure'. Species of the otherwise similar organ consisting of an inner layer of glandular Astacopsidrillls (Coddard, 19(9) are distinguished lining cells around a lumen and an outer layer of from ll1slllodrillls by the presence of female muscle cells. Such a modification from atri~lm to pores that are located within greatly enlarged protrusible penis could have arisen by separation spermathecal vestibulae. However, these are of the lining tissue from the muscle tissue, characters of degree and some ll1slllodrillls have leaving a tube (protrusible penis) within a tube well developed spermathecal ampulla and some (sac) (Brinkhurst 19(5). However, the lumen of Astacopsidrillls have female pores spermathecal the standard atrium is never ciliated vvhereas pores only just within or co-incident with the vasa deferentia (which normally feed into the openings of the vestibulae. Future analyses of the atria) are ciliated. This suggests that the mostly family Phreodrilidae will most Iikelv result in ciliated protrusible penes of P!Zreodrillls may i;, these genera being synonymised or (perhaps more fact be highly modified vasa deferentia (I'. Ma~tin, likely) split into multiple smaller genera. The IW\,y Institut Royal des Sciences Naturelles de Belgique, species below has moderately well developed pers. comm.) rather than atria. Cilia are absent vestibulae with separate female pores. The genus from the most ectal part of the pseudopenis, but occurs in New Zealand, South America. Africa modified vasa deferentia of some of the species and Australia. Eight species have previously been lack cilia ectally (Custav'sson and Erseus 1999). described from Australia, with most known only In most other aquatic oligochaetes the penes are from Tasmania. . conical to elongate structures located within an invagination of the body wall at the external Inslllodrillls angcla sp. novo opening of the male duct and often referred to Figure 3 as pendant penes. While the coiled structures present in Pilreodrillls are clearly not homologous 464 A.M. Pinder

Material examined litoralis Michaelsen 1903 (from the subantarctic Campbell Island, with long slender atria, Holotype smaller spermathecal vestibulae and ventral Australia: Western Australia: bore WAWB51, chaetae as described for lacllstris). West Angela mine site, approximately 100 km NW of Newman (PBS site PSSI67), 23°09'11.6"S, Description 118°44'51.3"E, 14 October 2004, H. Barron, J. Cocking Body of slide-mounted specimens 170-290 J.lm (WAM V7433, slide mounted (left coverslip). wide at Ill, 225-375 J.lm wide at X, most specimens missing posterior but one complete specimen Para types 6.05mm. Prostomium rounded and conical, Australia: Western Australia:, 1 specimen pharynx in Ill, strongly muscular posteriorly with on same slide (right coverslip) as holotype, thick muscle connections to dorso-lateral body collection details as for holotype (WAM V7434); 2 wall of II-V. Pharyngeal glands on lower half of specimens, Nyeetbury Spring (site PSW016 of PBS), anterior and posterior sides of septa 4/5 to 6/7 21°51'29"S, 116°30'57"E, 15 May 2005, A. Pinder, J. or 7/8 and laterally on oesophagus in V to VII. McRae (WAM V7435, slide mounted); 1 specimen, Oesophagus widening into gut in VIII-X. Nyeetbury Spring (site ONS03 of MRHI), 21°51'36"S, Ventral chaetae (55-80 J.lm) from II, all paired 116°30'42"E, 11 May 1996, W. Kay, M. Smith (WAM and bifid with small but distinct upper teeth V7436; slide mounted); 1 specimen, Bore WB32, (Figure 3A). Ventral chaetae of XIII modified: one West Angelas Mine, Pilbara, 23°09'48"S, 118°41'05"E, long (85-105 J.lm) and hollow tipped, protruding 6 October 1998, S. Eberhard (WAM V7437; slide either end of the large spermathecal chaetal mounted); 1 specimen, Bore YR15/73 (site PSS132 gland, the other short and not protruding. Ventral of PBS), Yule River borefield, Pilbara, 20 0 32'19.2"S, chaetae of XII absent. Dorsal chaetae from III, 1 118°13' 02.1"E, 12 October 2004, J. Cocking, H. to 2 smooth hairs, each surrounded by a pair of Barron (WAM V7438; slide mounted); 1 specimen, support chaetae, mostly 110-480 J.lm long (rarely Limestone Bore (site PSS394 of PBS), Pilbara, shorter), tapering evenly to a fine tip. 22°43'32.6"S, 118°48'02.4"E, 20 August 2004, M. Genitalia paired. Male pores ventro-lateral Scanlon, H. Barron (WAM V7439; slide mounted); in posterior third of XII in line with the ventral 1 specimen, floating root mat in New Mowbowra chaetae. Testes antero-ventral in XI, ovaries Cave, a non-anchialine cave on the coastal plain antero-ventral in XII. Female ducts penetrate on the eastern side of Cape Range, 21°59'34"S, 12/13 to exit at pores immediately anterior of the 114°07'19"E, 26 June 1993, W. Humphreys (WAM opening of the spermathecal vestibule antero­ V7440; slide mounted). ventral on XIII in line with the ventral chaetae (Figure 3B-ff). Thin clitellum from V2 XII to end of Other material examined XIII, not developed ventrally on XII. Australia: Western Australia: 2 specimens, Narrow (7.5 to 9.5J.lm) ciliated vas deferens Millstream Delta (PBS site PSW011, a warm (Figure 3B-v) joins elongate (35-44 J.lm wide, spring), Millstream National Park, Pilbara, 270-540 J.lm long) unciliated atrium (Figure 3B-a) 21°34'58"S, 117°04'08"E, 24 August 2003, A. Pinder, as the latter narrows just before entering penis J. McRae (DEC slide 68); 1 spedmen, Limestone (join not visible on holotype but seen on some Bore (site PSS394 of PBS), Pilbara, 22°43'32.6"S, paratypes). Atrium with lumen of variable width, 118°48'02.4"E, 20 August 2004, M. ScanIon, H. probably reflecting reproductive stage. Slender Barron (DEC slide 188); 1 specimen, Bore TCS002 pendant penis occupying most of the penis sac. (site PSS388 of PBS), 23°33'06.8"S, 118°15'16.7"E, 9 Spermathecal vestibule vase-shaped ectally May 2005, M. Scanlon, H. Barron (DEC slide 202). (Figure 3B-sv) with longitudinal muscle layer and numerous muscle filaments extending from the Diagnosis vestibule to the dorso-lateral body wall and septa. Insulodrilus angela differs from other described The vestibule dilated at its dorsal extremity and 1l1slllodrilus primarily in the size and form of surrounded by circular muscle tissue (Figure 3B­ the spermathecal vestibule and the un-modified cm), bending posteriorly to join a spermathecal somatic chaetae. Other species lacking modified duct (Figure 3B-sd) above the spermathecal somatic chaetae are l. lacustris s. str. (see chaetal gland. Ducts joining voluminous ampullae Remarks), l. martel1si Martin and Giani 1995 in XIV, which extends into at least Xv. Sperm in (from Africa, with alveolate penes, thickened loose oval bundles in ampulla. septa in the genital region and short stiff hairs), l. gel1italisetifera Martin and Brinkhurst 1994 Remarks (also from Africa, with genital chaetae in XI These specimens were at first identified as and XIII and an alveolate penis) and lnsulodrillls 1l1sulodrillls lacustris s. I., the suffix indicating that Phreodrilidae in north-western Australia 465 B A cm

sd

v ff sv

Figure 3 111SIIlodrillls ililXflil sp. nov.: A, ventral chaeta; B, genitalia, excluding the spermathecal ampulla. Scale lines: A, 20Mm; B, 100Mm.

this name has been given to what now appears to distributions (pinder and Brinkhurst, 1997; Pinder be a complex of similar species in Australia. 2001) and the recognition that the Australian Despite indications to the contrary by Pinder and "laclIstris" are in fact multiple endemic species fits Brinkhurst (1997), the Australian specimens differ this pattern. from l. lacllstris s. str. (as described by Benham, All specimens were collected in circum-neutral 1903 from New Zealand specimens) in a number fresh to slightly brackish G;2.4g/L) water with of respects. In contrast to all Australian forms, variable ionic composition, including dominance I. laclIstris has narrow and simple spermathecal by sodium with bicarbonate, or magnesium with vestibulae, hardly differentiated from the duct chloride, sulphate or bicarbonate. apart from the presence of longitudinal muscles in the former and circular muscles in the latter. They Etymology also have dorsal chaetae that project only V4 to This species is named after the type locality, a 1/3 the body height and scarcely half the segment bore near West Angela I-lill. length (whereas most Australian forms have the hair chaetae as long as or longer than the body DISCUSSION height) and have anterior ventral chaetae with The global distribution of the Phreodrilidae one chaeta of a pair clearly bifid and the other indicates a Condwanan origin (Martin cl al. 2008. almost simple (just a rudimentary hint of a tooth) Pinder and Brinkhurst 1997) and, to date, the whereas in Australian specimens both chaetae in majority of species have been found in places all ventral pairs are always distinctly bifid. that seem to offer refugia from past or present The several forms of lacllstris-like lnslllodrillls aridity: alpine areas, high southern latitudes, in Australia differ in the extent of development groundwater or deep lakes. cLhl'ardi of the spermathecal vestibule and ducts, length Pinder 2003, which occurs in ephemeral seepages of the atrium, body size, chaetal size and a on granite outcrops, is an apparent exception, number of other features. The states of these though nothing is known of where it Slll'\,ives characters seem be consistent and distinct within when the seepages are dry. The occurrence of geographic regions and sufficient to recognise a seven (mostly still undescribed) phreodrilids number of species, including l. angcla and another in the arid Pilbara is further evidence that the undescribed Pilbara species IllslIlodrillls WA35. familv. occurs more widelv. within Australia than Almost all phreodrilids have limited geographic previously thought. Their occurrence mostlv 466 A.M. Pinder

.3 in groundwater associated habitats fits with a .8 Qj > general pattern of taxa of presumed Gondwanan '." ro 1-. CC origin occurring preferentially in refugial/ C,) b!) > ::l C ro ~ CC "1J 1-. relictual habitats when they occur in arid zones rt: > c bl). > E 1 re ro I 1-. >. [e.g., some water mites (Harvey 1998) and ~ :a .:oL "1J ~ V u C ,;; rJl 0 phreatoicid isopods (Knott and Halse 1999)]. Three 1-. ro ~ "1J rJl q; of the Pilbara phreodrilids have been collected 0 rJj co only in groundwater, and another two have been "1J q; collected only in groundwater plus springs and 2: N LD OCJ l'-. O~ '0 G! \0 0 trj 1 1 I lr) spring fed pools. Only the undescribed Pl1reodrilus 'Jj ,.... 00 ,....cr-: LD ,.... N,.... ".; .;a rJl tr, "1J WA12 has not been found in groundwater to date

N but its only known locality is a spring-fed creek. ] if) c:: \0 G! Q\ N LD l{''j LD Pl1reodrilus peniculus is particularly widespread in .8 0 0 ."r tr, I ".; 1 1 rr) 0 0 00 U Lt) \0 M N l'-. 11 '2 :I: "'" "'" the region's surface waters, including deep gorge rJl ~ Cl "tl pools, springs, spring fed pools and river pools t- c:: ~ \0 M ~ N OCJ I I without a flowing spring but which are probably ~ if) c:;; "'"0 0 "'"0 0 0 0 N N N M N ,.... Q\ c:: permanent as a result of hyporheic inflow. There ,....Q\ "'" .=-:u is a single record of an immature specimen from if) u >. if) ~ \0 ,.... N ~ a seasonal turbid creek pool with no obvious q; ~ ,.... 1 N,.... N 1-. C N 0 ."r M .c ~ groundwater influence. 0... iii S > Eberhard et al. (in press) discuss the difficulty of ::l 'S + LD OCJ ~ ,.... l'-. :::t C'"! N determining the ranges of groundwater species, 0" 'iu 00 0 I LD N ,.... 0 r' r'"'" S u N M N N M N N particularly in relation to sampling effort. Their 0 a 1-. '"- '6 analyses indicated that 70 'X, of stygal species q; + ,.... M M l'-. OCJ 2 > ~ l'-. may have ranges <10,000km (which Harvey ro 0 'eo ."r u:; I N"'" ,.... I c:;; N to 00 U ::; M M N M N (2002) proposed as an upper threshold for short ro "'" ;:1-. range endemism). This has impact assessment 0 + l'-. N M N \0 M N~ Q\ Q\ implications for the many resource developments .!:l to r' 1 to N ".; ."r 0 ;: Z M N M in the Pilbara that are dewatering aquifers. Three 0 "'" "'" "'" ::; phreodrilids may have small ranges (Figure ;: q; 1): P. limzaei is presently known only from its rJl .... z , N lr) N l'-. ...,l \0 I € ,.... ,.... groundwater type locality, the undescribed 1-. Cl 0 N"'" "'"0 0 t- tib 0 " C'"!,.... 0 0 '"- Phreodrilus WA12 is known only from two springs ro -:u (one on the Robe River and one on the adjacent Cl x 0 ;> M \0 OCJ M ,.... lower Fortescue River) and Phreodrilus WA32 is "tl l'-. I 1 M N OCJ OCJ ~ ~ E LD M lr) t", known from two groundwater bores in close 1:! ... "'" q; proximity on the coastal plain near the Cane ~ £ 0 River. Phreodrilid WA36 is more widespread c:: c?- lr) .5 ~ 0 G! C'"! l'-. Q\ 1 I Lr) but also infrequently collected and known only 00 0 M N "1J M LD l() q; >. to X N from groundwater. 1I1sulodrilus angela and the C 0 0 undescribed Insulodrilus WA35 are widespread, "-§ q; Q.. common and found in groundwater and (less OCJ OCJ N l() l'-. 0; S u Q\ OCJ 1 ,.... E 0 N c:;; N 0 00 r' r' "'"0 frequently) surface waters (Figure 1). 1I1sulodrilus if) ,.... N N N N M M q; ~ :-e angela occurs in at least four drainage basins in ro the Pilbara plus a cave on Cape Range. Insulodrilus u OCJ OCJ LD Q\ M N M .2 :I: M OCJ "'"N I Q\ \0 Q\ l'-. Q.. r' r'"'" r' r' r' r' WA35 occurs mostly in groundwater of two C,) 0 0 0 £ basins (DeGrey and Port Hedland Coast) whose 1-. N LD \0 M M OCJ l() 0 0 Q\ Q\ 0 0 f100dwaters occasionally merge in their lower 2 ...... 0 ...... Q\ ...... "'" ...... "'"0...... 0 ...... "'" ,.... trj lr) OCJ \0 0 0 OCJ LD 0 reaches, but one record in a groundwater-fed .8 ~ ,.... 0 0 0 0 ,.... ,.... 0 0 ,.... ro -:u ...... "1J ,.... l() ,.... \0 \0 0 Q\ N pool is a geographic outlier separated from Cl N ,.... ,.... N N 0 ,.... N 0 ,.... ro riJ "'" "'" other records by the eastern Hamersley Ranges. :2 q; cq; 0 Phreodrilus peniculus appears to be the most if) > ro s q; C u common and widespread species (Figure 1). 0 ::0 bJ) 1-. ro ro 1-. .5 .8 1-. q; Immature specimens with chaetae resembling ";; 1-. ;: ,.... 1-. .8 :; 0... o:J l() 0 M C rJl Cl 0 l'-. Insulodrilus (both chaetae of a pair bifid) or W-l ,:.c c:<: .!:l co co ...... co >, N q; tr, 1-. E 3: M 3: ,.... Pl1reodrilus (one simple and one bifid chaeta in ~ ::l ro 0 co C ,...., q; N c:<: .!:l 1-. ~ .8 each bundle) have been found across the Pilbara. .£ ~ if) 0 ~ :::t Q) 3: >- q; q; 0 iii q; ~ ;: q; q; rJl q; These may just be immatures of the already :c U 1-. q; 1-. ~ 1-. 0 C >, 0 E u 0 ~ ...,l co Z ~ Z co cO :J t- co common species but their wide distribution Phreodrilidae in north-western Australia 467 indicates that the full picture of phreodrilid Oliguchacta from New Zealand. Pro(eedlll:\'; ot till' distributions in the Pilbara is yet to be revealed. 01 London 2: 202-232 Springs fed by discharging groundwater Brinkhurst, R.O. (1965). 1\ taxonomic revision of the I'hreodrilidae (Oligochaeta). Iou mill of 147: or hyporheic flow a re present t h roughout the 363~386 Pilbara region and the occurrence of the same Brinkhurst, R.O. (]991). A phylogenctic analysis of phreodrilid species in both these springs and in the I'hreodrilidae (i\nnelida, Oligochacta), with a groundwater suggests that the springs are a portal description of a new species. Canadian 10ll/'llal of for movement between surface and groundwater. Zoology 69: 2031~2040. Epigean movement within catchments, and Eberhard, S.M., I!dlse S.A. and Humphreys, W.I:. (2005). between catchments when coastal floodwaters Stygofauna in the I'ilbara region, north-west Western merge, would also be enhanced by the regular Australia: a review. 10ll/'llal of the I\oyal Socicly of and severe floods resulting from cyclonic rains. Weste/'ll Allstralia88: ]67-]76. Changes to flow paths and isolation of ancient Eberhard, S.M., lIaise, S.A., Williams, M., Scanlon, M.D., river tributaries by rising sea levels have perhaps Cocking, IS and Barron, 11.1. (in press) Exploring the also contributed to the occurrence of these relationship between sampling efficiency and short species across present catchment divides. For range endemism for groundwater fauna in the I'ilbara example, the Fortescue River formerly flowed region, Western Australia. Freshwater Biologl/. to the coast via what is now the lower Robe Coddard, E.I. (1909). Contribution to a further knowledge River but is now entirely separate. Similarly, the of Australasian Oligochaeta. Part 1. Description of two Ashburton River, which now tlows to the coast species of a new genus of Phreodrilidae. Proceedings of the Linnean Socicly of Nei/' SOllth Wales 33: 768-793. near Onslow, is believed to have been in close proximity to Cape Range between the Middle Custavsson, L.M. and Erseus, C (1999). Development of the genital ducts and spermathecae in the Miocene and Late Pleistocene (Wyrwoll et Ill. rhyacodrilines Rhyacodri/lls coccinells and 1993), bringing it much closer to the record of I. lvlollOpylephoms rtllmmii!ells (Oligochaeta, Tubificidae). IIllgclll (1993) from the Cape. Knott advanced the 1011 mal of Morphology 242: 141-156. latter concept as an explanation for the presence Harvey, MS (1998). Unusual new water mites (Acari: of the Iimnostygobiont shrimp Stygiocllris stl/lifL'r11 Hydracarina) from Australia, I'art 1. [{ecords of the and two subterranean fish species on the Cape, Western Australian Muselllli 19: 91-106. all three of which co-occurred with I. IIllg('/1I in Harvey, MS (2002) Short-range endemism among the New Mowbowra Cave (W.E. Humphreys, Western Australian fauna: some examples from non marine Australian Museum, pers. comm.). environments. Invertebrate Sl/stclliatics 16: 555-570. Humphreys, W.E (1994). The sllbterranean/i1llna of the Cape ACKNOWLEDGEMENTS Range coastal plain, IlOrthi/'estem Allstralia. Report to The Pilbara Biological Survey was funded by the Australian Heritage Commission and the Western the Department of Environment and Conservation Australian Heritage Committee. 202 pp. Western Australian Museum, Perth. Western Australia, the Australian Government Humphreys, W.E (1999). Relict stygofaunas living in sea through the Natural Heritage Trust, and Straits salt, karst and calcrete habitats in arid northwestern Resources. The Monitoring of River Health Australia contain many ancient lineages. In: W. Ponder Initiative was funded by the Commonwealth and D. Lunney (eds). The other 99'/0. The consermtion Government through the National Heritage Trust and Iliodiversity or invertebrates: 219-227. Royal and Land & Water Australia and by the State Zoological Society of New South Wales, Mosman. Government through Department of Environment Humphreys, W.E, Watts CHS and Bradbury pI. (2005) and Conservation and the Water and Rivers Emerging knowledge of diversity, distribution and Commission. William Humphreys of the Western origins of some Australian stygofauna. In: I. Cilbert Australian Museum kindly provided specimens (ed.). Proceedings o(all Intematiollal Syllipo..;iulIi on World and data collected with financial assistance SII/!tcrraneall 57~60. Villeurbanne, France, from the National Estates Program, the l;eritage 8-10 December 2004. Universitv of Lyon, France. Council of Western Australia, the Water Authority Knotl, B. (1993). Stygofauna from Cape Range peninsula, of Western Australia and the Western Australian Western Australia: Tethyan relicts. Records of tlie Museum. Susan Jones helped prepare the map. Westc/'ll Allstralll/n Ivlllselllli SlIpplellient 45: ]09-127. Mike Scanlon assisted with slide mounting and Knotl, B. and lIaise, S.A. (]999). 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