European Journal of Taxonomy 72: 1-17

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2Q14 • Lorz A-N., Myers A. & Gordon D.

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Research article

urn:lsid:zoobank.org:pub:94AB5EEl-49F4-4398-B081-C0189747CF23

An inquiline deep-water bryozoan/amphipod association

from New Zealand, including the description of

a new genus and species of Chevaliidae

Anne-Nina LORZ Alan MYERS & Dennis GORDON

^ National Institute of Water & Atmospherie Researeh, Private Bag I4-9QI,

Kilbimie Wellington, New Zealand

^ Sehool of Biologieal, Earth and Environmental Seienees, University College, Cork,

Cork Enterprise Centre, Distillery Fields, North Mall, Cork, Ireland

* Corresponding author. E-mail: [email protected]

^ E-mail: [email protected]

E-mail: [email protected]

^ um:lsid:zoobank.org:author:9442484E-43A4-4383-AlA6-AE493Q87BCAl

" um:lsid:zoobank.org:author:68E417C5-lAlD-4FBC-B832-2AQEBEE79DFl

^ um:lsid:zoobank.org:author:DD9CQF3A-8512-4AC8-B395-7687CE3FC565

Abstract. For the first time Amphipoda have been diseovered living in Bryozoa. A new genus and

speeies of the amphipod family Chevaliidae, Bryoconversor tutus gen. et sp. nov. is deseribed from New

Zealand at depths of 53Q-15QQ m. The speeies lives in an inquiline relationship with the eheilostome

bryozoan Onchoporoides moseleyi (Calwelliidae), inhabiting an abfrontal basal eoelom of the bryozoan

beneath the membranous eetoeyst (eutieularized epithelium) that eoneeals and proteets the amphipods.

The eolony is strengthened along all edges by a unique intraeoelomie rod of ealeium earbonate that is

formed within the marginal kenozooids of the eolony. The potential benefits and eosts to the bryozoan

are diseussed.

Key words. Amphipoda, Chevaliidae, Bryozoa, eommensalism. New Zealand.

Lorz A-N., Myers A. & Gordon D. 2Q14. An inquiline deep-water bryozoan/amphipod association from New

Zealand, including the description of a new genus and species of Chevaliidae. European Journal of Taxonomy 72:

1-17. http://dx.doi.org/lQ.5852/eit.2Q14.72

Introduction

Amphipoda are known to be assoeiated with many different invertebrates, ineluding deeapod erustaeeans

(Myers 1974; Vader & Myers 1996; Vader & Krapp 2QQ5), molluses (Vader 1972), aetiniarians (Vader

1984), aseidians (White & Reimer 2Q 12a) and sponges (Eorz & De Broyer 2QQ4; White & Reimer 2Q 12b).

Bryozoans are also known to be assoeiated symbiotieally with a variety of invertebrates, ineluding

sponges (Harmelin et al. 1994), eorals (MeKinney et al. 199Q), atheeate hydroids (Puee et al. 2QQ7),

polyehaetes (Ehlers 1876), hermit erabs (Taylor 1994) and other deeapod erustaeeans (e.g., Eggleston

European Journal of Taxonomy 72 : 1-17 ( 2014 )

1971; Gordon & Wear 1999). D’Hondt (1983), Hayward (1985) and Cook (1985) documented a range

of additional associations between bryozoans and other organisms, from mutualistic to parasitic.

Until now, the Chevaliidae was considered a monotypic family and the single genus Chevalia Walker,

1904 has been reported from amongst algae, coral rubble, ‘various invertebrates’ and dead coral.

However, the type species (C. aviculae Walker, 1904) was found in washings from pearl oysters and at

least one species (C. setosa Sousa Filho, Souza & Valerio-Berardo, 2010) is known to be associated with

sponges, although the exact nature of the relationship is unknown.

Here we report the first known association between an amphipod and a bryozoan. A new genus and

species of chevaliid amphipod was discovered beneath the membranous ectocyst (cuticularized

epithelium) covering a colony-wide abfrontal basal coelom in the bryozoan Onchoporoides moseleyi

(Busk, 1884), in New Zealand deep waters. It is described and illustrated here as Bryoconversor tutus

gen. et sp. nov. The bryozoan is an erect, planar form with fiattened fronds that bifurcate at intervals. It

is relatively weakly calcified but its unique mode of strengthening its margins is described herein. The

nature of the relationship and whether one or both parties benefit is discussed.

Material and Methods

Recent and historic collections of the bryozoan Onchoporoides moseleyi from New Zealand waters held

at the Marine Invertebrate Collection, National Institute of Water & Atmospheric research (NIWA),

Sc*OI^ IB.

Vanvoiu

New Cblfidonia

TuD^kjjgtclau

SarflOB

Narfbh I

AUElTBMa

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Fig. 1. Sampling localities of the calwelliid bryozoan Onchoporoides moseleyi harboring Bryoconversor

tutus gen. et sp. nov. The type locality is indicated by a star.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

were investigated. Almost 100 eolonies were examined, -95% of them hosting amphipods, from six

loealities. The distribution map (Fig. 1) was ereated in AreGIS.

Amphipoda were removed after being photographed in their host (Fig. 2A) and transferred to 70%

ethanol. Speeimens were examined and disseeted under a Leiea MZ9.5 stereoseope and an Axioseope 2

Plus Zeiss mieroseope, and disseeted parts were mounted in glyeerin and drawn using a camera lucida.

Fig. 2. A. Photograph of the abfrontal side of a eolony of the bryozoan Onchoporoides moseleyi

showing adults (middle left arrow) and juveniles (upper right arrow) of Bryoconversor tutus gen. et sp.

nov. beneath the semitransparent abfrontal membrane. Note the paired kenozooidal rootlets and their

branehes at lower left; these anehor the eolony to sediment grains in the soft-sediment habitat. The blue

eolour of the eolony is optieal, not staining. B. Optieal photograph of the proximal end of a small eolony

of Onchoporoides moseleyi showing the aneestrular zooid (a) in abfrontal view, with ealeified interiors

(*) of lateral kenozooids that eonverge proximally. Seale bars: A = 3 mm; B = 100 pm.

European Journal of Taxonomy 72 : 1-17 ( 2014 )

Pencil drawings were scanned, digitally inked and arranged as plates using the methods described in

Coleman (2003, 2009).

The body lengths of amphipod specimens examined were measured by tracing the mid-trunk length

from the tip of the rostrum to the end of the telson. For scanning electron microscope (SEM) imaging

the specimens and appendages were dehydrated through a graduated ethanol series, acetone dried,

mounted on stubs, coated with gold-palladium and investigated via a SEM EE01525. SEM imaging of

the bryozoan host skeleton was carried out using a Hitachi TM3000 Tabletop Microscope.

All material is held in the Marine Invertebrate Collection, NIWA, Wellington.

Results

Suborder Senticaudata Eowry & Myers, 2013

Infraorder Corophiida Eeach, 1814

Parvorder Corophiidira Eowry & Myers, 2013

Superfamily Chevalioidea Myers & Eowry, 2003

Family Chevaliidae Myers & Eowry, 2003

Genus Bryoconversor gen. nov.

um:lsid:zoobank.org:act:761AFFll-lBB8-4DFD-9126-E15D7C0B8ECB

Type species

Bryoconversor tutus sp. nov., by monotypy

Etymology

From prefix bryo- (abbreviation of bryozoan) and Eatin conversor, to dwell with.

Diagnosis

Head lateral cephalic lobes weakly produced forward, subtriangular; anteroventral excavation weak.

Eye present. Labrum lacking epistome. Antenna 1 & 2 subequal in length. Gnathopod 2 larger than

gnathopod 1, subchelate; Pereopod 5-7 coxae very shallow. Epimera rounded; Urosome segments 1

and 2 coalesced. Uropods 1-2 biramous, inner ramus longer than peduncle, weakly curved with deep

longitudinal ridges and completely lacking setae. Uropod 2 posterior margin with a strongly developed

fiange with a pitted edge. No sexual dimorphism.

Remarks

The new genus is very close to Chevalia Walker, 1904, which also lacks sexual dimorphism, but it

differs from the latter in the presence, on the posterior margin of the uropod 2 peduncle, of a greatly

expanded fiange with a pitted edge (whose function is unknown). In Chevalia, the uropod 2 peduncle is

unexpanded. In addition, the uropod 1 inner ramus inner margin is smooth in Bryoconversor, but with a

row of large serrate spines in Chevalia. It also differs by the deep longitudinal ridges on uropod 1 and 2

inner ramus that are clearly visible under the light microscope in Bryoconversor. These ridges cannot be

seen in Chevalia under the light microscope, although confirmation of their absence with SEM would

be advisable.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

Bryoconversor tutus sp. nov.

um:lsid:zoobank.org:aet:93156279-10B8-4435-A6EQ-FF53BFD5F252

Figs 3-8

Diagnosis

As for genus.

Etymology

Eatin: ‘tutus’ = safe.

Material examined

Holotype

NEW ZEAEAND: NIWA 88918, ovigerous 1.72 mm long, station G675, 19 Jan. 1970, 45.4500° S,

171.4000° E, 792 m.

Paratypes

NEW ZEAEAND: Same eolleetion data as for holotype. NIWA 52919, juvenile S, 0.64 mm long;

NIWA 89534, 4 juveniles, about 0.4 mm long.

Additional material

NEW ZEAEAND: Several dry speeimens. NIWA 89536, st. G705, 23 Jan. 1970, 46.0667° S, 172.4750°

E, 1500 m; NIWA 89533, st. TAN1208/58, 24 Jun. 2012, 42.8107° S, 179.8265° E, 1005 m; NIWA

89535, st. TAN0705/155, 16 Apr. 2007, 42.9989° S, 176.3483° W, 648 m; NIWA 89538, st. C620,

2 May 1961, 43.6667° S, 174.7833° W, 752 m; NIWA 89537, st. E827, 24 Get. 1967, 46.5917° S,

166.7417° E, 530 m.

Description

Female (holotype, 1.72 mm long)

Head. Eateral eephalie lobes weakly produeed forward, subtriangular, anteroventral exeavation weak.

Eye present. Antenna 1 8c 2 subequal in length, about half body length. Antenna 1 slender, posterior

margin with long slender setae; peduneular artieles in the ratios 3:4:3; flagellum less than half length of

pedunele, with 7 artieles, aeeessory flagellum with one long and one tiny artiele. Antenna 2 peduneular

artieles 4 and 5 subequal; flagellum shorter than pedunele, with 6 artieles. Labrum laeking epistome.

Mandible molar triturative; palp three-artieulate; artiele 1 short; artiele 3 a little longer than artiele 2,

elongate sub-ovoid with dense distal setae and a few more sparsely on the posterior distal margin. Labium

with subaeute mandibular projeetions. Maxilla 1 inner plate with marginal setae; palp biartieulate with 5

distal robust setae. Maxilla 2 inner plate with oblique setal row. Maxilliped palp mueh longer than outer

plate; daetylus rounded, short, laeking nail.

Pereon. Gnathopod 1 eoxa weakly produeed forward, sub-aeute; basis slender, anterior margin weakly

eoneave; earpus slender, longer than propodus; propodus slender, palm oblique, evenly eontinuous with

posterior margin; daetylus slender, two thirds length of propodus. Gnathopod 2 larger than gnathopod

1; eoxa subquadrangular, equal in depth with eoxa 1; basis stout, a little over twiee as long as broad,

anterior margin eoneave; earpus short, stout, as broad as long; massive propodus nearly twiee length

of earpus, palm with three proeesses, a triangular distal spine and two rounded mid-palmar humps;

daetylus a little shorter than palm. Pereopods 2-4 slender; eoxae subtriangular, about half depth of

eoxael-2; daetylus short, about one third length of propodus. Pereopod 5-7 eoxae very shallow; earpus

small, lunate; merus large and broad; daetylus with aeeessory spine on anterior margin. Pereopod / basis

of posterodistal margin with strong projeetion.

European Journal of Taxonomy 72 : 1-17 ( 2014 )

Fig. 3. Bryoconversor tutus gen. et sp. nov., holotype, NIWA 88918, 1.72 mm long. A. Habitus. B.

Urosome. C. Antenna 2. D. Mandible. Seale bars = 100 pm.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

Fig. 4. Bryoconversor tutus gen. et sp. nov., holotype, NIWA 88918, 1.72 mm long. A. Hypopharynx.

B. Maxilla 2. C. Maxilla 1. D. Maxilliped. Seale bars = 100 pm.

European Journal of Taxonomy 72 : 1-17 ( 2014 )

Fig. 5. Bryoconversor tutus gen. et sp. nov., holotype, NIWA 88918, 1.72 mm long. A. Gnathopod 1.

B. Gnathopod 2. C. Pereopod 3. D. Pereopod 4. Scale bars =100 pm.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

Fig. 6. Bryoconversor tutus gen. et sp. nov., holotype, NIWA 88918, 1.72 mm long. A. Pereopod 5.

B. Pereopod 6. C. Pereopod 7. D. Pleopod 1. Seale bars =100 pm.

European Journal of Taxonomy 72 : 1-17 ( 2014 )

Pleon. Epimera 1-3 rounded. Urosome segments 1 and 2 eoaleseed. Uropod 1 biramous, inner ramus

longer than pedunele, weakly eurved with deep longitudinal ridges and eompletely laeking setae; outer

ramus a little shorter than pedunele with distal robust setae and one sub-distal seta. Uropod 2 posterior

margin with a strongly developed flange with a pitted edge; inner ramus longer than pedunele, weakly

eurved with deep longitudinal ridges and eompletely laeking setae; outer ramus a little shorter than

pedunele, with distal robust setae. Uropod 2 pedunele short, rami subequal in length with pedunele, with

terminal setae. Telson dorsoventrally thiekened with one long dorsal seta on eaeh side; subdistal margin

with a small group of flat-topped projeetions visible under SEM.

Fig. 7. Bryoconversor tutus gen. et sp. nov., paratype, S, NIWA 52919, 0.64 mm long. A. Habitus. B.

Head. C. Epimeron. D. Urosome. E. Uropod 2 and uropod 1. F. Telson. Seale bars: A = 300 pm; B-D =

100 pm; E = 20 pm; F = 10 pm.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

Male

(Sexually dimorphie eharaeters.) Not different from female, exeept for mueh smaller body size.

The bryozoan host

Onchoporoides Ortmann, 1890 is a nominally monotypie genus of Calwelliidae (Cheilostomata) that is

known only from the southwestern Paeifie. The sole ineluded speeies, O. moseleyi, was first eolleeted by

the R. V. Challenger from a depth of 951 m on the Kermadee Ridge north of Maeauley Island. Additional

Fig. 8. Bryoconversor tutus gen. et sp. nov., paratype, S, NIWA 52919, 0.64 mm long. A. Detail of

serrated surfaee ontelson. B. Gnathopod 1 and larger gnathopod2. C. Daetylus gnathopod 1. D. Daetylus

gnathopod 1. E. Dorsal view telson, uropod 3 and uropod 2. F. Flat-topped projeetions on telson. Seale

bars: A = 1 pm; B = 100 pm; C-D = 10 pm; E = 20 pm; F = 2 pm.

European Journal of Taxonomy 72 : 1-17 ( 2014 )

Fig. 9. A. Optical view of the abfrontal side of a distal marginal autozooid (polypide redueed to a

residual brown body, bb) adjaeent to a lateral kenozooid in whieh there is a distally tapered (i.e., apieally

developing) intraeoelomie ealeareous rod (arrow). B. SEM of a similar view to Abut of skeletal elements

only, showing the distal tip of the ealeareous rod (white arrow) and the eommunieation pores (blue

arrowhead) that allow nutrient eonneetivity between the feeding autozooid and non-feeding kenozooid

in whieh the eoelomie eavity eomes to be filled by the ealeareous rod. C. Fraetured transverse seetion

of a ealeareous rod showing alternating thiek and thin wall-perpendieular prismatie fabrie. D. Part of C

magnified. Seale bars: A = 200 pm; B = 100 pm; C = 25 pm; D = 5 pm.

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

speeimens were subsequently taken by the then New Zealand Oeeanographie Institute (now NIWA)

from 526 m in the Solander Trough, at the western approaehes to Foveaux Strait, and from 2677 m in

the Hikurangi Trough east of Cook Strait (Gordon 1989). This paper adds additional reeords, within

this depth range, mostly from stations east of South Island. Gordon & d’Hondt (1997) also reported the

speeies from the Loyalty Basin.

Onchoporoides moseleyi forms planar ereet eolonies up to 52 mm high. These ean braneh diehotomously

to give an overall flabellate form and the whole eolony is anehored in soft substrata by root-like rhizoids

(non-feeding zooidal polymorphs). Like the zooids in the eolony, eaeh rhizoid has a ealeified body

wall. A tubular kenozooid (also a non-feeding bryozoan polymorph) runs down the lateral margin

of the eolony on eaeh side, eonverging proximally to form a short stem that divides proximally into

numerous individual rhizoids (Fig. 2 A) with small lateral proeesses. The founding individual of a eolony

(aneestrula) is unusually long (2 mm) for bryozoans. It tapers proximally into two rhizoid-like proeesses

that anehor it. The eolony grows by distal budding of new zooids from the aneestrula and its daughter

zooids.

The zooids of O. moseleyi open only on one faee, with resident amphipods dwelling on the abfrontal side

(Fig. 2A) where they apparently do not interfere with the life proeesses of the bryozoan. Enelosing the

amphipods is a semitransparent membrane that is attaehed to eaeh outer margin, eonforming to the shape

of the eolony. The spaee enelosed by the membrane is interpreted to be a eolony-wide extrazooidal basal

eoelom and its membrane an eetoeyst (eutieularized epithelium). Gordon (1989) noted the apparent

universality of the membrane in the eolonies he examined, with the possible exeeption of very small

eolonies in whieh he failed to deteet a membrane, perhaps beeause of its transpareney. Busk (1884)

deseribed the speeies on the basis of a single speeimen 19 mm high and did not mention an abfrontal

membrane or any assoeiation with amphipods.

Examination of growing margins of O. moseleyi by light mieroseopy and SEM indieates that the basal

eoelom and eetoeyst are laterally eontiguous with the eontinuous marginal kenozooid (‘extrazooidal

marginal eoelom’ of Cook & Chimonides 1981) on eaeh side of the braneh. The basal eoelom probably

originates from the vieinity of the aneestrula (Fig. 2B) when it buds the first daughter zooids distally,

eaeh with a lateral kenozooid and the presumed ineeption of the basal eoelom and eetoeyst between

them. Later in eolony ontogeny, the first settler amphipod (possibly a hermaphrodite or a female with

an attaehed male eompanion) is inferred to eut an entranee slit in the eetoeyst, presumably using its

gnathopods, in order to insert itself beneath, but not so destruetively as to wholly interfere with the

integrity of the basal eoelom (though there must be some leakage or loss of eoelomie fiuid). One dried

eolony shows a longitudinal slit in the eetoeyst where it would otherwise attaeh to the lateral kenozooid,

with the tips of amphipod appendages protruding, but it is not elear if the tear is a eonsequenee of drying

or made by the amphipod. Self-repair is well doeumented in bryozoans (e.g., Ryland 1970) and a slit

made by an amphipod to enter the basal eoelom would quiekly repair. We believe it mueh more likely

that resident amphipods would remain within the basal eoelom, surviving on nutrient transfer from the

bryozoan, rather than eonstantly exiting and returning or sustaining an opening in the basal membrane

for feeding. Continual loss of eoelomie fiuid would eompromise the integrity of the extrazooidal basal

eoelom, whereas it is evident from the eontinued differentiation of the distal braneh tips of the eolony

by intussuseeption that this must not be the ease.

There are no eommunieation pores in autozooidal basal walls (whieh are, however, unealeified, allowing

diffusion aeross the thinly eutieularised epithelium). There are sueh pores in the lateral walls, ineluding

those abutting the marginal kenozooidal tube as it differentiates on eaeh side of a braneh. At their distal

tips, the marginal kenozooids are hollow and filled with eoelomie fiuid. Proximal to eaeh tip the tapered

distal end of a developing intraeoelomie ealeified rod is visible (Fig. 9A-B). This rod oeeupies the

remainder of the kenozooid, fully oeeupying the eoelomie eavity. A transverse fr aeture of the rod shows

European Journal of Taxonomy 72 : 1-17 ( 2014 )

it to have a wall-perpendicular prismatic fabric (cf Weedon & Taylor 2000) made up of thick and thin

layers (Fig. 9C-D).

Discussion

Commensalism describes a relationship between two living organisms in which one benefits and the

other is not significantly harmed or helped and that appears to be the case here - the bryozoan provides

shelter or housing for the amphipod, hence the association is a case of inquilinism.

The existence of the extrazooidal basal coelom that accommodates the amphipod residents is puzzling,

as it has not been reported in any other calwelliid species. The precise conformity of its enclosing

membrane to the shape of the bryozoan, from the proximal-most zooids to the branch tips, supports

the inference that it is produced by the bryozoan. All colonies in the collection have a basal coelom

and ectocyst. In addition, it appears improbable that it could have been produced by the amphipod,

as there is no known mechanism whereby it could secrete (or induce) such a membrane and secure it

precisely to the lateral margins of the bryozoan. Extrazooidal basal coeloms have been found in several

cheilostome families. Examples include species of Cupuladriidae (Hakansson 1973; Tavener-Smith

1973), Euthyrisellidae and Didymosellidae (Cook & Chimonides 1981), Selenariidae (Chimonides &

Cook 1981) and Urceoliporidae (Gordon 1988). In their discussion. Cook & Chimonides (1981, p. 81)

also briefiy mention the presence of “an extensive” extrazooidal basal coelom in Petraliidae and in

their text (p. 62) they note that “extrazooidal marginal coelomic systems ... resemble those formed by

lateral kenozooids in ... Onchoporella buskiE (see Florence et al. 2007: fig. 201). However, they did

not mention an extrazooidal basal coelom in this calwelliid species, in which it thus seems to be absent.

The kenozooidal intracoelomic calcareous rod has not been reported before and the alternation of dense

layers with thinner layers (Fig. 9C-D) is likewise novel. The thin layers appear as though they may

have a higher organic content. The layered nature of the rod would render it less prone to fracture from

stresses on the colony, especially if semi-organic layers allowed for minute internal “slippage” of layer

against layer. Preserved colonies can bend slightly which implies that this is possible in nature.

Although the bryozoan host does not appear to benefit from or be harmed by the association, the question

arises concerning the selection pressure that resulted in the unique internal calcification of the marginal

kenozooids. As indicated above, other members of the family Calwelliidae, especially Onchoporella

buskii and species of Ichthyaria that also form planar fronds, either lack marginal kenozooids (four of

the five species of Ichthyaria) or the kind of calcification that results in a vertical intracoelomic rod.

{Ichthyaria simplex Gordon & d’Hondt, 1997 from New Caledonia has marginal kenozooids and lacks

a basal uncalcified window in each zooid and should be transferred to Onchoporoides. It is not known

if the marginal kenozooids have an intracoelomic rod.) We have no evidence that the presence of the

amphipod induces the formation of either the extrabasal coelom or the kenozooidal intracoelomic rod.

Both would impart strengthening of the bryozoan colony, preventing excessive intracolony Hexing or

bending although, arguably, given the fact that the colony is rooted in soft sediment, bending of the

entire colony from the base is likely to be possible. An extrazooidal basal coelom would also strengthen

the colony and reduce intracolony bending and Hexing, especially in combination with intracoelomic

calcification of the marginal kenozooids.

Our interim conclusions and hypotheses are: 1) the extrazooidal basal coelom is produced by the

bryozoan, not induced by the amphipod; 2) a hermaphrodite or a female, accompanied by a smaller,

attached male gains entrance to the extrabasal coelomic cavity by making a slit that subsequently repairs

itself; 3) the amphipods live and reproduce within the coelom, relying on nutrients provided by the

bryozoan; 4) when the bryozoan colony is damaged or dies the amphipods exit the coelom and colonize

LORZ A-N., MYERS A. & GORDON D., A new genus and speeies of Chevaliidae

other available bryozoans in the same habitat. The relationship appears to be one-sided, benefiting the

amphipod by providing nutrients and shelter (proteetion from predation). Potentially, an entire bryozoan

eolony and its resident amphipods eould be eaten by a relatively large predator (see Lidgard 2008

for examples of predation on bryozoans) but many bryozoans have bioehemieal defenses that deter

predation. The bryozoan host would seem not to benefit from the assoeiation, unless the amphipod

seeretes or exeretes organie moleeules that the host ean uptake. It has been established, for example, that

bryozoans are able to uptake dissolved organie earbon (examples in Gordon et al. 1987).

Acknowledgements

Speeimens studied here were eolleeted under numerous researeh projeets funded by the New Zealand

Ministry for Primary Industries; the historie samples were eolleeted by the former New Zealand

Oeeanographie Institute (now the National Institute of Water & Atmospherie Researeh, NIWA).

We thank the NIWA Marine Invertebrate eolleetion team for registration and euration of the speeimens,

Erika MaeKay (NIWA) for digitally inking several peneil drawings and Peter Gerring (NIWA) for

ereating the map. Renate Walter (Zoologieal Museum Hamburg) kindly assisted with the seanning

eleetron mieroseope. Two anonymous reviewers vastly improved an earlier version of this paper.

This work has been supported by NIWA through Coasts and Oeeans Researeh Programme 2 (Marine

Biologieal Resourees) taxonomy projeet COBRI402.

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Manuscript received: 24 October 2013

Manuscript accepted: 1 January 2014

Published on: 29 January 2014

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium; National History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.