Memoirs of Museum Victoria 62(2): 103-146 (2005)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

The millipede genus Lissodesmus Chamberlin, 1920 (Diplopoda: Polydesmida:

Dalodesmidae) from Tasmania and Victoria, with descriptions of a new genus

and 24 new species

Robert Mesibov

Queen Victoria Museum and Art Gallery, Launceston, Tasmania 7250, Australia (mesibov@southcom.com.au)

Abstract Mesibov, R. 2005. The millipede genus Lissodesmus Chamberlin, 1920 (Diplopoda: Polydesmida: Dalodesmidae) from

Tasmania and Victoria, with descriptions of a new genus and 24 new species. Memoirs of Museum Victoria 62(2):

103-146.

Lissodesmus includes L. adrianae Jeekel, 1984, L. alisonae Jeekel, 1984, L. modestus Chamberlin, 1920 (type species)

and L. perporosus Jeekel, 1984 from Tasmania, L. martini (Carl, 1902) from Victoria, and 23 new species: L. anas,

L. bashfordi, L. clivulus, L. cognatus, L. cornutus, L. devexus, L. hamatus, L. horridomontis, L. inopinatus, L. latus,

L. montanus, L. orarius, L. peninsulensis and L. plomleyi from Tasmania, and L. blackwoodensis, L. catrionae,

L. dignomontis, L. gippslandicus, L. johnsi, L. macedonensis, L. milledgei, L. otwayensis and L. tarrabulga from Victoria.

The new genus Tasmanopeltis, close to Lissodesmus , is erected for T. grandis sp. nov. from Tasmania.

Keywords Diplopoda, Polydesmida, Dalodesmidae, millipede, Australia, Tasmania, Victoria, spiracles, sphaerotrichomes

Introduction

Tasmania and Victoria are home to a diverse group of

dalodesmid Polydesmida with well-developed paranota, a long

dorsal seta near the posterior corner of each paranotum (Fig. 1)

and fairly uniform gonopod structure. Currently all such

forms are referable to either Lissodesmus Chamberlin, 1920

or Dasy stigma Mesibov, 2003. In this paper I place 14

new Tasmanian and nine new Victorian species in

Lissodesmus. The expanded genus is far from homogeneous,

but it is not yet clear how it should be divided into mono-

phyletic subgroups. An unusual form from north-eastern

Tasmania is placed in Tasmanopeltis gen. nov., close to

Lissodesmus.

Characters. Tasmanian and Victorian dalodesmids with a

head + 20 segments, well-developed paranota and a long pos-

terior corner seta (Fig. 1) are here called the “ Lissodesmus

group”. All species have four main branches or processes on the

gonopod telopodite arranged as shown in Fig. 2. In naming

these processes I follow Jeekel (1983, 1984), but Jeekel’s

“solenomerite” is here “solenomere”. This is the mesal or

anteromesal process in which the prostatic groove terminates.

In L. martini (Carl, 1902) it was called the “principal branch”

( Hauptast ) by Carl (1902) and the “seminal groove branch”

( Samenrinnenast ) by Attems (1940), while Chamberlin (1920)

called it the dorsal branch of the mesal spur in L. modestus

Chamberlin, 1920.

The telopodite continues distally from the solenomere origin

as the prefemoral process (pf) (Carl: “neighbouring branch”

( Nebenast) ; Attems: “tibiotarsal branch” ( Tibiotarsalast )).

Arising on the lateral or anterolateral side of the prefemoral

process is the femoral process (f) (Carl: “secondary branch”

(sekunddrer Ast) of the “neighbouring branch”; Attems: “short

side branch” ( kurzer Seitenast )). Finally, arising posterior to the

solenomere is the tibiotarsus (tt) (Chamberlin: ventral branch

of the mesal spur; Attems: “spine” {Dorn)).

The process names as used here are only conventional

labels. Because the development of the telopodite in male

Polydesmida is so abrupt and so cryptic, it is not certain that the

processes are homologous with the podomeres suggested by

the process names, i.e. prefemur, femur, tibia and tarsus.

In addition to the main telopodite processes, most species in

the Lissodesmus group have at least one mesolaterally flattened

projection on the posterior surface of the prefemoral process. In

accord with Jeekel (1984) this projection is here called an

“uncus” (u in Fig. 2), although it is not always hook-shaped.

As background to a discussion of classification in the

Lissodesmus group, I review here a number of characters,

beginning with the telopodite processes. Apart from L. adri-

anae Jeekel, 1984, L. alisonae Jeekel, 1982, L. martini (Carl,

1902), L. modestus Chamberlin, 1920 and L. perporosus Jeekel,

1984, all Lissodesmus species mentioned in this section are

new species described below.

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Robert Mesibov

Solenomere. The solenomere is the least variable of the

telopodite processes. It typically arises at one-third to half the

length of the telopodite on the anteromesal surface, bending

posteriorly and often lying at the bend on a broad indentation in

the telopodite body. The prostatic groove typically enters the

solenomere on its anterior side near its base, then abruptly

curves to run on the posteromesal side, terminating at the

solenomere’s fine, generally truncated tip. The solenomere

varies very little in size relative to the telopodite as a whole

from species to species. In most species the solenomere is

directed distally or posterodistally, but in L. devexus (Fig. 32) it

points posteriorly. A small structure like a toothed half-collar is

found in some species on the anterior or anteromesal side of the

solenomere close to its tip (e.g. in L. martini. Fig. 50). This

structure is reduced to a small, pointed projection in some

species (e.g. L. modestus. Fig. 53) and is absent in L. adrianae,

L. devexus, L. horridomontis, L. peninsulensis, T. grandis and

all four Dasy stigma species (Mesibov, 2003b). The most

unusual solenomeres are found in L. peninsulensis, in which the

solenomere curves helically (Fig. 60), and in L. tarrabulga,

which in place of a subapical projection has a large, stout

process arising about midway along the solenomere on its

anterior side (Fig. 66).

Prefemoral process. The tip of the prefemoral process

generally bends posteriorly. The amount of bending varies from

a slight curvature in L. peninsulensis (Fig. 61) to a 180° turn in

L. catrionae (Fig. 22). InL. adrianae (Fig. 12) andL. modestus

(Fig. 53), the tip is erect. In L. dignomontis (Fig. 35) and

L. johnsi (Fig. 45) the tip is completely unarmed but in other

species it bears teeth or tooth-like projections in a wide range

of sizes, shapes, positions and orientations, and the process in

many species ends in an elaborate comb (e.g. in L. macedonen-

sis, Fig. 48). In most species the tip of the prefemoral process

is undivided. The tip is broadly divided in L. orarius (Fig. 57)

and forked in L. bashfordi (Fig. 17). In L. hamatus (Fig. 39),

L. horridomontis (Fig. 41), L. inopinatus (Fig. 43) and

L. otwayensis (Fig. 59) the distal portion of the process is

abruptly shifted laterally, with a low “shoulder” projection

marking the mesal end of the bend. In T. grandis (Fig. 68) this

“shoulder” is larger and projects distally as a separate sub-

process. The prefemoral process varies in length and armature

in L. devexus (Figs 32, 33), but in other species these two

features are more stable.

Femoral process. The most complex femoral processes are

found in Dasystigma species, in which the process is variously

divided near its tip and armed (Mesibov, 2003b). In other

species the process is typically a smooth, bluntly pointed bar or

blade, often with a posterior branch (e.g. in L. alisonae. Fig.

14). In T. grandis (Fig. 68) the tip of the femoral process is

expanded and carries a few blunt projections. Lissodesmus

group species also vary in where the femoral process originates

on the telopodite, i.e. proximal or distal to the solenomere

origin. In four species the femoral process extends distally as

far as the tip of the prefemoral process: L. bashfordi (Fig. 18),

L. dignomontis (Fig. 35), L. johnsi (Fig. 45) and L. modestus

(Fig. 52). Whether the femoral process in these species has

become a functional adjunct of the prefemoral process is

unknown.

Tibiotarsus. In most species the tibiotarsus is a simple

rod directed posterodistally, but in L. devexus the process is

directed posterobasally (Fig. 32). The tibiotarsus originates just

posterior to the solenomere in all but the four Dasystigma

species, in which the origin is much further lateral, roughly

halfway around the telopodite body towards the origin of the

femoral process. The tip of the tibiotarsus is expanded and

turned distally in a number of species, and in L. catrionae

(Fig. 22), L. comutus (Fig. 30), L. horridomontis (Fig. 41) and

L. montanus (Fig. 55) the tip is armed with blunt, tooth-like,

marginal projections. A curious feature of the tibiotarsus in

L. devexus is a series of annular “wrinkles” visible at high mag-

nification (Fig. 31). The “wrinkles” give the impression that the

tibiotarsus has been compressed along its long axis before the

cuticle had thoroughly hardened. In L. johnsi (Figs 44, 45)

the tibiotarsus is fan-shaped and marginally toothed.

Uncus. Many species have a single small uncus similar to

the one shown schematically in Fig. 2, and some have several

mesolaterally flattened projections in the same region, i.e.

between the solenomere origin and the tip of the prefemoral

process. Because these processes vary in shape, number and

position, it is not clear whether they should be considered

homologous, e.g. whether either or both of the small, ridgelike

projections in L. perporosus (Figs 62, 63), partway along the

prefemoral process on its posterior surface, are development-

ally equivalent to the large, arcuate structure in L. peninsulen-

sis (Fig. 61). Assuming homology, the most variable and

unusual uncus is found in L. devexus (Figs 32, 33). In speci-

mens from the type locality the uncus is a long, blade-like

structure parallel to the tip of the prefemoral process, while in

specimens from a nearby locality the uncus is merely a low

ridge with a slightly hook-shaped tip.

Telopodite setae. All species have sparse, long setae on the

posterior surface of the proximal portion of the telopodite. The

most distal setae are typically at the level of the solenomere

origin or just beyond. In L. otwayensis a row of large setae con-

tinues distally almost to the level of the uncus (Fig. 58), and in

L. adrianae the most distal setae are close to the apex of the

prefemoral process (Fig. 11).

Body size. Overall length ranges from c. 11 to c. 35 mm, but

length measurements of preserved specimens are affected by

the degree to which each prozonite is telescoped into the next

metazonite headwards. The size measure used here, H, is the

height of segment 12 as viewed from the rear (see Fig. 3), i.e.

in the plane of the posterior edge of the metazonite. H was

measured ±0.1 mm on a male of each species judged to be

typical; see remarks on individual species for comments on

within-species variability. Between species H varies from 1.2 to

3.3 mm.

Antennomeres. Chamberlin (1920: 135) observed that the

antenna in L. modestus had “the second and sixth articles

longest, the third a little shorter, the fourth and fifth much

shorter”. In redescribing L. modestus from topotypical mater-

ial, Jeekel (1984) measured the relative lengths of anten-

nomeres 2 to 6 as 1.00, 0.90, 0.65, 0.65 and 0.85 and

commented: “In his description Chamberlin stated that the 2nd

and 6th antennomeres are the longest with which present

observations do not agree. It seems possible that Chamberlin

New genus of millipedes

105

actually meant the length of the 2nd and 6th to 8th anten-

nomeres” (p. 94). I have not seen Jeekel’s material, but I have

examined the L. modestus holotype and Chamberlin’s descrip-

tion is correct. The difference between the two descriptions is

almost certainly due to differences in the degree to which the

antennomeres are telescoped in the preserved specimens, and in

how antennomere length was measured. Antennomere propor-

tions vary little in the Lissodesmus group (Fig. 69; see also Fig.

2 in Mesibov, 2003b), with antennomere 2 slightly longer than

or about equal to 6, followed by 3, then by the more or less

equal 4 and 5; antennomere 6 is invariably the widest.

Paranota. The size, shape and setation of paranota vary

from anterior to posterior within individual specimens in the

Lissodesmus group, as elsewhere in the Polydesmida.

Differences can also be seen when comparing the same seg-

ment from individual to individual: in the shape of the para-

notal margin as viewed from above; in the degree of “inflation”

of the paranota in a transverse section of the segment; in the

number, position and degree of definition of marginal notches;

and in the presence/absence of setae associated with the

notches. This variation devalues some paranotal characters for

separating species. However, the ratio R of paranotal width to

prozonite width (Fig. 3) is useful. In the 29 species considered

here, R of segment 12 ranged from 1.3 (“reduced paranota”) to

1.8 (“wide paranota”). Maximum prozonite width is tightly cor-

related (r 2 = 0.96; n = 29) with H in individual specimens, but

R is apparently not correlated with H between species. A

second useful paranotal character is the degree to which the

posterior corner is turned up (Figs 4, 28). Tumed-up comers are

associated with reduced relative width, occurring in all five

species with R = 1.3, two of the eight species with R = 1.4 and

two of the 13 species with R = 1.5.

Ozopores. The pore formula is normal for head + 20 seg-

ment Polydesmida, i.e. ozopores on segments 5, 7, 9, 10, 12,

13, 15-19, except for L. perporosus, which has the formula 5,

7, 9-19. Ozopores open dorsolaterally near the paranotal

margin, more or less close to the posterior corner (Fig. 70).

Spiracles. In most Lissodesmus species the spiracles are as

shown in Fig. 5A for L. modestus. They have thick and

rounded rims, small apertures, and no projecting stmctures. On

each diplosegment the anterior spiracle opens just above the

coxa of the anterior leg, with a thin shelf-like projection of the

body wall separating the two. The posterior spiracle opens

about midway between the anterior and posterior legs. In both

sexes in Dasy stigma species and in Tasmanopeltis grandis (Fig.

5B) the spiracle rims are very thin and the aperture relatively

wide, and a dense mass of hair-like stmctures projects from the

opening. Further, the posterior spiracle lies immediately adja-

cent to the anterior spiracle in T. grandis and in D. bonhami

Mesibov, 2003 (Mesibov, 2003b, Fig. 5A) and D. margaretae

Mesibov, 2003 (Mesibov, 2003b, Fig. 5C, D). Stadium 5

T. grandis have spiracles of normal appearance and at normal

spacing; the spiracles become progressively “hairier” and

closer together in stadia 6, 7 and 8. In both sexes of L. anas

(Fig. 5C), a dense mass of hair-like structures obscures the

spiracle openings. The size, shape and position of the L. anas

spiracles are seen in cleared preparations to be the same as in

L. modestus. A small clump of hair-like structures arises from

the body wall just posterior and ventral to the posterior spiracle

(just visible in Fig. 5C). Just anterior and ventral to the anter-

ior spiracle, a long, rigid “fox tail” of hair-like stmctures arises

and extends posteriorly almost to the posterior spiracle. Many

of the hair-like structures associated with both spiracles are

forked near their base. The “fox tail” stmcture has not, to my

knowledge, been noted elsewhere in the Diplopoda. I have not

seen stadium 6 or younger L. anas , but a less well-developed

anterior “fox tail” and posterior clump are present in stadium 7.

Limbus. Under relatively low magnification (200x), the

limbus appears to be composed of simple, straight elements.

Under higher magnification the elements in L. latus are seen to

be strongly tapering structures (Fig. 6A), while in L. hamatus

each tapering element has a row of sharp, posteromesally

directed teeth on the mesal surface (Fig. 6B). The limbus in

other species has not been examined at high magnification.

Epiproct. In most Lissodesmus group species the epiproct

ends in two short, well-separated, rounded bumps (Fig. 7A).

The projections are longer in some Lissodesmus species

(Fig. 7B), giving the epiproct a forked appearance even at low

magnification.

Podomeres. As usual for Polydesmida, legs close to the

gonopods in Lissodesmus group males are greatly swollen com-

pared to the corresponding legs in females. The prefemur has a

rounded dorsal projection, and some dorsal expansion is

usually also seen on the femur (Fig. 71). Species differ in the

degree of swelling, e.g. L. anas (Fig. 71ana) has slender legs

and L. macedonensis (Fig. 71mac) has thick, robust ones. In

some species the tarsus is shorter than the femur (e.g.

L. horridomontis, Fig. 71hor), in others the same length (e.g.

L. perporosus , Fig. 71per), and in still others distinctly longer

(e.g. L. peninsulensis , Fig. 71pen). Tibiae are also variable,

with a ventral, distal swelling apparent in some species (e.g.

L. modestus. Fig. 71mod).

Sphaerotrichomes. I presented some preliminary observa-

tions on male leg setae (Mesibov, 2004a) and reported that

beginning with leg 3, most legs in male L. modestus have

sphaerotrichomes on the prefemur, femur, postfemur, tibia and

tarsus. An erratum noted that sphaerotrichomes were present

only on the tibia and tarsus in L. modestus. However, my orig-

inal observations were correct (Fig. 8A), and sphaerotrichomes

can, in fact, be seen on the L. modestus femur in Fig. 2G of

Mesibov (2004a). The same error was made by Jeekel (1984) in

his redescription of L. modestus (“globular setae on tibia and

tarsus”, p. 93), and by Johns (1964) in his revision of

Pseudoprionopeltis, including what is now L. martini. In my

own case the error arose because I habitually viewed legs from

the side, and in this orientation the dense “brush” setae on the

proximal podomeres can hide sphaerotrichomes lying mainly

on the midventral line. The true distribution of sphaerotri-

chomes can be revealed using scanning electron microscopy, or

by clearing the leg (e.g., with 60% lactic acid) and viewing it

from beneath with a light microscope and substage illumin-

ation. Sphaerotrichome numbers and distributions vary from

leg to leg (Mesibov, 2004a). For the present study I cleared and

examined leg 6 from one male of each of the 29 species under

consideration. Sphaerotrichomes were found on the prefemur,

femur, postfemur, tibia and tarsus of 24 species, but in some

106

Robert Mesibov

cases “presence” on the prefemur amounted to only one or two

sphaerotrichomes. Sphaerotrichomes were seen on the tarsus

and tibia only in L. devexus and L. milled gei] on the tarsus, tibia

and postfemur in L. cornutus and L. horridomontis\ and on the

tarsus, tibia, postfemur and femur in L. orarius (Fig. 8B).

Absence of sphaerotrichomes from podomeres was clearly cor-

related with a lower total number of sphaerotrichomes on the

leg, indicating that absence from some leg 6 podomeres in the

five named species results from variation in a quantitative char-

acter. A similar quantitative variation is seen in individual spec-

imens: the total number of sphaerotrichomes per leg decreases

from leg 3 rearwards, and the last legs typically have

sphaerotrichomes on tibia and tarsus only. In most Lissodesmus

species the sphaerotrichome shafts are gradually tapered, as

shown for L. modestus in Fig. 2D in Mesibov (2004a), but in

some species, e.g. L. martini, the tips of the shafts are slightly

expanded, as in Bromodesmus species (Fig. 2B in Mesibov,

2004a).

“Furriness” . The new species L. cognatus and L. latus, are

“furry” in portions of their respective ranges. The “fur” (Fig.

9B) consists of long, thin setae sparsely distributed on the col-

lum and metatergites of all males and females collected in these

areas (for locations, see remarks on the respective species).

Furry individuals have the usual complement of longer setae on

the head, the anterior portion of the collum and the posterior

portion of the preanal segment. Differences between the

gonopods of furry and typical forms are minor. The functions

of the fur are unknown; furry specimens are found in the same

microhabitats as non-furry forms.

Juvenile stadia. The typical pattern of paranotal develop-

ment in the Lissodesmus group is shown in Fig. 10 for L. per-

porosus. Stadium 5 and younger individuals have prominent

teeth armed with setae on the lateral and posterior margins. As

the individual matures, the lateral teeth are progressively

reduced to marginal sections delimited by small notches. All

or most of the lateral setae are lost; the most anterior of the

lateral setae is the one most frequently retained (Fig. 70). The

posterior marginal teeth disappear in most species but are

retained in stadium 8 of Tasmanian L. latus and Victorian

L. blackwoodensis, L. dignomontis, L. gippslandicus, L. johnsi,

L. macedonensis, L. martini and L. tarrabulga (Fig. 70). The

lateral margin of the paranotum is typically straight and

parallel to the longitudinal body axis in early stadia, curving as

the individual matures (Fig. 10).

Classification

Jeekel (1984) reviewed the six species then kn own in the

Lisodesmus group: L. adrianae, L. alisonae, L. margaretae,

L. martini, L. modestus and L. perporosus. The gonopod struc-

ture generalised in Fig. 2 clearly set these six species apart from

the other dalodesmids with a head + 20 segments known at the

time from Tasmania and Victoria, namely Gasterogramma psi

Jeekel, 1982, Gephyrodesmus cineraceus Jeekel, 1983 and

Tasmanodesmus hardyi. Jeekel (1984) applied to that concept

the oldest available name, Lissodesmus.

There are now 33 species in the group, and some subgroup-

ing is apparent. It is less clear how to translate that structure

into genera (or subgenera, which in myriapodological practice

are often “holding bays” for future genera). There are few use-

ful non-sexual characters, and gonopod character states seem to

be almost randomly mixed through the group. I have therefore

looked for discontinuities: multi-character morphological gaps

between species.

Four species of Dasystigma were distinguished by an

unusually wide separation between solenomere and tibiotarsus,

by the presence of hair-like structures in the spiracles, and by a

close similarity in body size and colour, overall gonopod struc-

ture and paranotal form (Mesibov, 2003b). Here a new genus is

erected for Tasmanopeltis grandis, remarkable for its excep-

tionally large size, spiracles with hair-like structures, process

origins unusually far distal along the telopodite, and a unique,

large, distal projection on the prefemoral process.

The other species in the Lissodesmus group include many

which look to be closely related (i.e., have similar gonopods)

and occur in the same region, e.g. L. dignomontis, L. johnsi and

L. tarrabulga in Gippsland, and L. bashfordi and L. modestus in

south-east Tasmania. However, I here leave Lissodesmus as a

heterogeneous assemblage of 28 species. It thus joins

Icosidesmus in New Zealand (Johns, 1964) and Gnomeskelus in

southern Africa (Lawrence, 1953, 1958) as an unresolved knot

of relationships in Dalodesmidae. If Shelley (1990a) is correct,

it adds to a long list of large, “difficult” genera in many other

millipede families. Lissodesmus may not be the largest such

group in the south-east Australian Polydesmida; Tasmaniosoma

Verhoeff, 1936 contains a similarly puzzling diversity of forms

(Mesibov, in preparation).

Ecology and life history

In Tasmania, species in the Lissodesmus group are found in

moist leaf litter, in and under rotting wood, under stones, in the

upper layers of richly organic soil and in the moist skirt of

rotted bark, twigs and leaves around the base of larger trees.

They occur in cool temperate rainforest, wet eucalypt forest,

dry eucalypt forest and subalpine woodland. They are particu-

larly abundant in forest growing on fertile soils (e.g., those

derived from Tertiary basalt). Within the Tasmanian annual

rainfall range of c. 550-3500 mm they appear to be most abun-

dant at intermediate values, c. 1000-2000 mm. Lissodesmus

group species are occasionally found in alpine areas, grassy

woodland and coastal scrub, but are only rarely seen in moor-

land, heathland and grassland. A few species are accidental

cave inhabitants. No species have yet been found in long-

established gardens of exotic plants, but L. alisonae and

L. perporosus can be abundant in plantations of Pinus radiata

(Bonham et al., 2002; Mesibov, 2005).

Victorian Lissodesmus species are so far known from cool

temperate rainforest, wet eucalypt forest and dry eucalypt

forest in areas with an annual rainfall greater than c. 800 mm.

They are mainly found in well-rotted eucalypt logs and in accu-

mulations of moist peaty material in surface depressions on

such logs. In my own, limited experience, Victorian

Lissodesmus are very rarely seen away from logs at lower

elevations, but are sometimes found in leaf litter in the high

country (>900 m).

New genus of millipedes

107

Species in the Lissodesmus group are typically found shel-

tering in mixed- age aggregations. In Tasmania, life histories of

Dasy stigma, Lissodesmus and Tasmanopeltis appear to be only

weakly seasonal. Throughout the year, a collector can find

adults (including pairs in copula) and most juvenile stadia,

even in subalpine habitats. Victoria has hotter and drier

summers than Tasmania, and Victorian Lissodesmus may be

more seasonal than their Tasmanian congeners.

All species in the Lissodesmus group are cryptic in their

habits, but some Tasmanian species, notably L. perporosus, can

be found wandering at night. Judging from gut contents,

Lissodesmus group species feed on well-rotted wood and

richly organic soil particles. Nothing is known of their parasites

or predators, but small mites (unidentified) are often found on

older individuals of both sexes.

Biogeography

Mesibov (2003b) reported that Dasy stigma in Tasmania formed

a mosaic complex of the kind first clearly described by Shelley

(1990a, 1990b). The biogeographical situation is more compli-

cated in Tasmanian Lissodesmus. While mosaics can some-

times be found (e.g. in the north-east with L. adrianae,

L. alisonae and L. hamatus ; see Mesibov, 1997), in most parts

of the island species are sympatric. Three Lissodesmus species

are commonly found at a single site. Four species (L. anas,

L. clivulus, L. latus and L. perporosus ) have been collected in

forest adjoining Balfour in the north-west, and five

Lissodesmus species can occur in the same patch of forest in

parapatric overlap zones, such as the Weavers Creek parapatric

zone in the north-east (Mesibov, 1997; L. adrianae, L. alis-

onae, L. cognatus, L. devexus and L. hamatus), which is

also home to Tasmanodesmus hardyi and Tasmanopeltis

grandis. Two species, L. cognatus and L. devexus, seem to have

naturally disjunct areas of occurrence.

In Victoria, Lissodesmus distributions are mainly allopatric,

but L. blackwoodensis and L. macedonensis have been found

together near Blakeville, L. gippslandicus and L. dignomontis

near Allambee, L. gippslandicus and L. johnsi near Yarragon,

and L. gippslandicus and L. tarrabulga near Balook. L. martini

and L. milledgei probably co-occur near the Acheron Gap in the

Yarra Ranges and at The Beeches near Marysville. Collections

to date in Gippsland indicate that L. dignomontis and L. johnsi

are parapatric south-west of Trafalgar, and that L. gippsland-

icus and L. martini are parapatric near the Narracan Creek

valley.

Lissodesmus has so far been sampled at only c. 60 sites in

Victoria, compared to c. 1200 sites in Tasmania for

Dasy stigma, Lissodesmus and Tasmanopeltis, but the area over

which Lissodesmus could be expected to occur in Victoria is

substantial. Including only the higher-rainfall, southern por-

tions of the Western and Eastern Uplands, and both of the

Southern Uplands blocks, I estimate this area to be 80 000 km 2 ,

which is larger than Tasmania. Although much of the pre-

European cover of wet forest has been lost from this area, it is

likely that new species of Lissodesmus remain to be found in

the Grampians, East Gippsland and physiographically isolated

portions of the eastern highlands.

New, narrow-range species of Tasmanian Lissodesmus are

also likely to be recognised in future, as there are female (and

a few aberrant male) specimens in collections which I cannot

confidently assign to any of the species described below.

Several of these come from Flinders Island, which has not yet

been surveyed carefully for millipedes.

Conservation

The Tasmanian species L. clivulus, L. cognatus, L. hor-

ridomontis, L. orarius and L. peninsulensis have extents of

occurrence (range envelopes) of 300 km 2 or less. Nevertheless,

local populations of these species are often large and well-

distributed through extensive patches of native vegetation

which are unlikely to be cleared in the foreseeable future.

L. montanus, L. peninsulensis and L. plomleyi have larger

ranges but seem to be naturally rare. None of these three

species appears to be under threat. The range of L. montanus

includes parts of several large, formal reserves. L. peninsul-

ensis and L. plomleyi are both known mainly from areas of

public land within which there are sizeable informal reserves of

riparian forest, old-growth eucalypt forest, and steep and rocky

ground. My field experience over the past 25 years is that

Tasmanian Lissodesmus species tolerate the clearfelling, burn-

ing and regeneration of wet eucalypt forest habitat (Taylor,

1990; Mesibov, unpublished results).

Field studies aimed at establishing range boundaries and

habitat preferences have so far been carried out for two

Tasmanian Lissodesmus species. Originally thought to be rare

(Mesibov, 1992), L. orarius was later found to be the dominant

Lissodesmus species in coastal habitats between the Pedder and

Pieman Rivers on the west coast (Mesibov, unpublished report

to the Tasmanian Conservation Trust, 1993). Distribution map-

ping of L. alisonae increased the number of known localities

from nine to 93, and the range envelope (minimum convex

polygon) from 1551 to 4965 km 2 (Mesibov et al., 2002). Within

its range, centred on the densely settled Tamar Valley,

L. alisonae occurs in all native forest types as well as in

native/exotic vegetation mixtures in city parks, and has

recently been found to be abundant in a second-rotation Pinus

radiata plantation (Mesibov, 2005)

Tasmania is a largely forested island, and about 40% of that

forest is currently in formal reserves. The distribution of those

reserves is the result, in part, of efforts to secure large, region-

ally representative blocks of little-disturbed native forest com-

munities. The situation in Victoria is very different. Much of

the wet native forest present when Europeans arrived in the

19th century has been cleared for agriculture or forestry planta-

tions, and much of the surviving forest, reserved or not, has

been degraded by frequent burning and weed invasion. In my

field experience in Victoria, populations of Lissodesmus

species other than L. gippslandicus, L. martini and L. otway-

ensis seem to be small and restricted to little-disturbed native

forest remnants.

L. johnsi was first collected in 1890 near Trafalgar in the

Latrobe Valley, probably in the area’s tall, dense eucalypt

forests (Adams, 1978). The area is now almost entirely covered

with pasture and forestry plantations. Recent searches for

108

Robert Mesibov

L. johnsi have so far yielded specimens from forest remnants at

only two localities, both near Trafalgar and about 5 km apart.

Methods

Specimens were usually killed and preserved in 70-80%

ethanol. For some species, vouchers preserved for molecular

analysis in 95% ethanol have been deposited in QVM.

Preliminary drawings on graph paper were made using an eye-

piece grid; the grid was calibrated for measurements of body

parts. Gonopods and male sixth legs were cleared and tem-

porarily mounted in 60% lactic acid; other body parts were

temporarily mounted in a glycerine-water mixture. A Philips

Electroscan ESEM 2020 operated in high-vacuum mode was

used to examine material which had been air-dried before

sputter-coating with gold. SEM images were acquired digitally.

This review is based on more than 2000 museum samples

containing more than 6500 specimens. Details of specimens

other than holotypes and paratypes have been omitted from the

“Material Examined” sections and are given in a separate doc-

ument (“ Lissodesmus supplement”) available on the Memoirs

of Museum Victoria website, or from the author, or from the

Curator of Zoology at the Queen Victoria Museum and Art

Gallery.

In the text, Tasmanian localities are given with a UTM grid

reference (Grid Zone Designation 55G) and the (calculated)

equivalent latitude/longitude, in both cases with respect to the

AGD66 datum. Victorian localities are given with latitude/

longitude with respect to the WGS84 datum.

Abbreviations are as follows: AM, Australian Museum,

Sydney, NSW; MCZ, Museum of Comparative Zoology,

Cambridge, Mass., USA; NMV, Museum Victoria, Melbourne,

Vic.; QVM, Queen Victoria Museum and Art Gallery,

Launceston, Tas.; TM, Tasmanian Museum and Art Gallery,

Hobart, Tas.; WAM, Western Australian Museum, Perth, WA.

Taxonomy

In the case of millipedes distinguished primarily by gonopod

differences, keys and species diagnoses are necessarily verbose

and often very hard to understand, even for a specialist. The

most taxonomically useful part of a millipede species descrip-

tion is the gonopod illustration, and readers are encouraged

to look first at the scanning electron micrographs and line

illustrations provided.

Note that in the telopodite descriptions and drawings a

“mesal” view is one centred on the solenomere, “lateral” on the

femoral process, “anterior” on a line roughly midway between

the origins of these two processes on the anterior surface, and

“posterior” on a line roughly 180° around the telopodite from

“anterior”. This convention has been adopted for convenience;

as seen in the SEM views, the in situ orientation of the

telopodite varies considerably from species to species.

The five species L. adrianae, L. alisonae, L. martini,

L. modestus and L. perporosus have already been carefully

described by Jeekel (1983, 1984); brief, partial redescriptions

and new illustrations are included here only for the sake of

consistency.

Order Polydesmida Leach, 1815

Suborder Dalodesmidea Hoffman, 1980

Dalodesmidae Cook, 1896

Lissodesmus Chamberlin, 1920

Lissodesmus Chamberlin, 1920: 135. — Attems, 1940: 490. — Jeekel,

1970: 336.— Jeekel, 1983: 150.— Jeekel, 1984: 89.— Mesibov, 2003b:

198.

Pseudoprionopeltis ( Australopeltis ) Johns, 1964: 47.

Australopeltis Hoffman, 1980: 184. — Shelley et aL, 2000: 86.

Type species. Lissodesmus modestus Chamberlin, 1920, by

original designation.

Diagnosis. Small to medium-sized dalodesmids (11-23 mm

long, 1.2-2. 2 mm vertical diameter) with head + 20 segments,

normal pore formula (except L. perporosus ), well-developed

paranota with long posterior corner seta, spiracles well-

separated on diplosegments and without emergent hair-like

structures. Telopodite with small mesal or anteromesal

solenomere and tibiotarsus, small to large lateral or antero-

lateral femoral process and large central prefemoral process;

solenomere arising at one-third to one-half the telopodite

height. Prefemoral process tip typically undivided; in

L. orarius divided apically into two posteriorly directed

branches.

Remarks. Chamberlin (1920) erected the monotypic genera

Lissodesmus and Tasmanodesmus for the Tasmanian species

L. modestus and T. hardyi. The two genera were ignored by

Verhoeff (1932, 1936) on the grounds that Chamberlin had pub-

lished no illustrations, and were regarded as genera of uncertain

family placement by Attems (1940). Chamberlin believed the

two genera were closely related, but as shown in a recent

redescription (Mesibov, 2004b) T. hardyi is substantially differ-

ent in gonopod structure from all other Tasmanian dalodes-

mids, and the posterior comer seta in T. hardyi is short and

inconspicuous.

Pseudoprionopeltis martini Carl, 1902 from Melbourne was

included by Johns (1964) in his revision of New Zealand

Pseudoprionopeltis. He erected the subgenus Australopeltis for

P. martini and illustrated the anterior segments and gonopod of

a specimen from Cockatoo Creek in the Museum Victoria

collection. One of the characters used by Johns to diagnose

the new subgenus was the posterior comer seta, which he

described as “a long seta inserted just anterior and dorsal of the

[posterior paranotal] tooth” (Johns, 1964: 47).

In his reclassification of the Diplopoda, Hoffman (1980)

raised Australopeltis to a genus. Although Chamberlin (1920)

had provided a verbal description of the L. modestus gonopod,

Hoffman (1980: 150) regarded the gonopod structure of

Lissodesmus as “still unknown”. Nevertheless, he placed both

Australopeltis and Lissodesmus in Dalodesmidae Cook, 1896.

In doing so, Hoffman formalised the observation by Brolemann

(1916) that the circular gonopod aperture in P martini indi-

cated an affinity between that species and Semnosomidae

Brolemann, 1916, which Hoffman (1980) considered a

synonym of Dalodesmidae.

New genus of millipedes

109

Jeekel (1983) made Australopeltis a synonym of

Lissodesmus after examining fresh material of Victorian

L. martini and Tasmanian L. modestus. Both species had been

collected by Jeekel during a 1980 field trip to Australia. Jeekel

did not redefine Lissodesmus in the 1983 paper, instead refer-

ring the reader to “a previous paper”, then in press, which

actually appeared the following year (see below). However,

Jeekel (1983) gave a detailed description and clear illustrations

of a male of L. martini from Femtree Gully National Park.

In 1984, Jeekel redescribed L. modestus from topotypical

material and added four new Tasmanian species to the genus:

L. adrianae, L. alisonae, L. margaretae and L. perporosus. In

his key to the genera of Tasmanian Polydesmida, he noted that

Lissodesmus could be separated from Tasmanodesmus by the

presence in the former of “a long hair mesad of caudal edge of

paranota” (Jeekel, 1984: 89), i.e. the posterior comer seta.

L. margaretae Jeekel, 1984 was recently removed from

Lissodesmus and placed with three new Tasmanian species in

the new genus Dasystigma (Mesibov, 2003b).

As delimited here, Lissodesmus is what remains of the group

of south-eastern Australian dalodesmids with a head + 20 seg-

ments and a long posterior corner seta after exclusion of

Dasystigma (spiracles with hair-like structures, wide separation

between solenomere and tibiotarsus origins, complex femoral

process) and Tasmanopeltis gen. nov. (spiracles with hair-like

stmctures, relatively long unbranched portion of telopodite,

mesal “shoulder” process on prefemoral process). Lissodesmus

is thus a product of “taxonomic erosion” of a group of similar

taxa and is not defined by a set of unique character states.

However, it is readily distinguished from the other regional

H+20 dalodesmids Atalopharetra Mesibov, 2005, Bromodes-

mus Mesibov, 2004, Gastero gramma, Gephyrodesmus,

Tasmanodesmus and Victoriombrus Mesibov, 2004 by its pos-

session of a long seta at the posterior corner of each paranotum

and by the gonopod stmcture shown schematically in Fig. 2.

To avoid repetition in the species descriptions below, I have

omitted mention of the nearly constant non-sexual features

which can be seen in the detailed accounts given by Jeekel

(1983; 1984) ofL. adrianae, L. alisonae, L. martini, L. modestus

and L. perporosus. Note that the male antennae are separated by

about twice a socket diameter unless otherwise specified.

Distribution. Throughout Tasmania, and in parts of Victoria

with annual rainfall over c. 800 mm.

Lissodesmus adrianae Jeekel, 1984

Figures 7B, 11, 12, 69adr, 70adr, 71adr, 72 (map)

Lissodesmus adrianae Jeekel, 1984: 94.

Material examined. Holotype (not seen). Male, Australia, Tasmania.

Ben Lomond National Park, 35 km ENE of Evandale, along the road

to the top of Ben Lomond, near the park ranger office (approx.

EQ530040, 41°30'49"S 147°38”06"E), 23.xi.1980, C.A.W. Jeekel &

A. Jeekel-Rijvers. The type is said to be deposited in TM (Jeekel, 1984:

86), but has not yet been received there.

Paratypes. 9 males, 19 females, 10 stadium 7 males, 9 stadium 7

females, 2 stadium 6 females, details as for holotype; female, 10 km

NE of Blessington, 23.xi.1980, C.A.W. Jeekel & A. Jeekel-Rijvers.

These specimens are listed as paratypes by Jeekel (1984: 94) but their

present locations are u nk nown and they have not been examined.

Other material. 456 males, 449 females and 357 juveniles from 174

localities (see “ Lissodesmus supplement” for details).

Description. Male c. 18 mm long, H = 1.7 mm. In alcohol,

well-coloured specimens under low magnification with

light brown body colour dorsally, a transverse band of darker

brown near posterior metatergal margins and a small darker

brown patch medially, paranota near- white. Antenna moder-

ately long (Fig. 69adr). Paranota fairly wide, R = 1.5 (Fig.

70adr); posterior corners not turned up. Legs robust, tarsus

slightly shorter than femur (Fig. 71adr). Epiproct with promi-

nent paired, bluntly rounded projections (Fig. 7B). Telopodite

(Figs 11, 12) sparsely setose almost to apex, reaching almost

to leg 5 when retracted. Solenomere arising at just under half

the telopodite height, directed at c. 45° to telopodite axis

basally but smoothly curving distally, terminating at one-third

to half the prefemoral process length without subapical projec-

tion. Tibiotarsus arising slightly distal to solenomere origin,

rod-like, bluntly tipped, one-third as long as solenomere and

much narrower, directed at c. 45° to telopodite axis. Femoral

process arising at level of solenomere origin, straight, blade-

like, bluntly pointed, pressed close to prefemoral process, par-

alleling and reaching halfway to tip of latter, just past apex of

solenomere, with a short, bluntly pointed posterior branch at

one-third its length. Prefemoral process about two-thirds the

width of telopodite base at origin, straight, mesal edge with

a short series of small teeth midway to apex, the latter

finely-toothed, anteriorly concave. No uncus.

Distribution and habitat. In wet eucalypt forest, cool temperate

rainforest, subalpine woodland and wetter microhabitats in dry

eucalypt forest over c. 3500 km 2 in north-east Tasmania (Fig.

72), from 60 m to at least 1050 m. The western and eastern

range limits of L. adrianae correspond to the biogeographical

divides known as the East Tamar Break and Goulds Country

Break, respectively (Mesibov, 1994, 1997). The most southerly

known occurrence is near Castle Cary, north of Avoca. L. adri-

anae is abundant over most of its range and is by far the com-

monest dalodesmid in the wetter, forested parts of north-east

Tasmania.

Remarks. L. adrianae varies little in size, coloration and

gonopod structure.

Lissodesmus alisonae Jeekel, 1984

Figures 13, 14, 69ali, 70ali, 7 1 ali, 73 (map)

Lissodesmus alisonae Jeekel, 1984: 96.

Material examined. Holotype (not seen). Male, Australia, Tasmania. 8

km NW of Frankford, 15 km SW of Beaconsfield, 24.xi.1980, C.A.W.

Jeekel & A. Jeekel-Rijvers. The type is said to be deposited in TM

(Jeekel, 1984: 86), but has not yet been received there.

Paratypes. 12 males, 21 females, 1 stadium 7 male, 1 stadium 6

female, details as for holotype. These specimens are listed as paratypes

by Jeekel (1984: 96) but their present locations are u nk nown and they

have not been examined.

Other material. 286 males, 308 females and 129 juveniles from 181

localities (see ‘Lissodesmus supplement” for details).

110

Robert Mesibov

Description. Male c. 18 mm long, H = 1.7 mm. In alcohol,

well-coloured specimens under low magnification with pale

brown body colour, pale red posterior metatergal margins.

Antenna moderately long (Fig. 69ali). Paranota reduced, R =

l. 4 (Fig. 70ali); posterior corners slightly turned up. Legs

robust, tarsus about as long as femur (Fig. 7 lali). Telopodite

(Figs 13, 14) broad at base with abrupt narrowing at start of

prefemoral process, reaching almost to leg 5 when retracted.

Solenomere arising at about half the telopodite height, directed

distally at a small angle to the telopodite axis, terminating with

toothed subapical collar at one-third to half the prefemoral

process height. Tibiotarsus pointed and mesolaterally flattened,

parallel to and almost as wide as solenomere but somewhat

shorter and bending slightly laterally. Femoral process arising

just proximal to solenomere origin, not pressed close to

prefemoral process, forked at about one-quarter its length,

branches more or less equal, blade-like, pointed; anterior

branch directed distally and slightly concave posteriorly,

terminating just distal to solenomere tip at half to two-

thirds the length of prefemoral process, posterior branch

gradually curving anteriorly, its tip sometimes lying

between anterior branch and prefemoral process.

Prefemoral process about half the width of telopodite base,

slightly tapered, the tip curved posteriorly with small,

blunt, apical and subapical teeth. Uncus prominent, arising

centrally on prefemoral process at just under half the process

length (at level of solenomere tip).

Distribution and habitat. Common in dry eucalypt forest, wet

eucalypt forest and subalpine woodland over c. 6000 km 2 in

north central Tasmania (Fig. 73), from sea level to at least 1000

m. The principal eastern range limit for L. alisonae is the bio-

geographical divide known as the East Tamar Break (Mesibov,

1994, 1997), although the species extends eastward along the

north coast into the lower Brid River catchment, and a possibly

isolated population has been sampled at Cuckoo, near

Scottsdale. L. alisonae reaches the Don River in the west and

Projection Bluff on the north-east comer of the Central Plateau.

East of the Plateau L. alisonae is parapatric with L. hamatus

(Mesibov, 1997) along a more or less SW-NE line across the

Northern Midlands. L. alisonae has been found to be abundant

in a second-rotation Pinus radiata plantation at Stoodley

(Mesibov, 2005) and sometimes occurs in part-native gardens

in the Launceston area. Throughout its range, populations are

largest in the richly organic soil under dense stands of the

understorey tree Pomaderris apetala.

Remarks. L. alisonae varies somewhat in size and col-

oration, with the smallest and palest forms found in dry forest

on relatively infertile soils. In the north-eastern portion of the

L. alisonae range, the femoral process on the gonopod is longer

and the posterior branch less curved (Fig. 14).

Lissodesmus anas sp. nov.

Figures 4C, 5C, 15, 16, 69ana, 70ana, 71ana, 74 (map)

Lissodesmus sp. Wl. — Mesibov, 1993: 31.

Material examined. Holotype. Male, Australia, Tasmania. Duck Creek,

CP408763 (41°44'56"S 145°05'06"E), 160 m, 6.iii.l999, R. Mesibov,

QVM 23:45823 (ex QVM 23:40749).

Paratypes. 2 males, north of Pieman Head, CP261860 (41°39'31"S

144°54 , 41"E), 10 m, l.vi.1993, R. Mesibov, QVM 23:17656; 2 males,

details as for holotype, AM KS91166 (ex QVM 23:40749); 2 males,

details as for holotype, QVM 23:40749, dissected; 5 females, details as

for holotype, QVM 23:40748.

Other material. 5 males, 9 females and 15 juveniles from Balfour,

Heemskirk Road, Mt Frankland, Newdegate Creek, Pieman Head,

Piney Creek, Roger River West, Savage River and Wild Wave River

(see “ Lissodesmus supplement” for details).

Description. Male c. 16 mm long, 77 = 1.5 mm. In alcohol,

well-coloured specimens under low magnification with pale

brown body colour, slightly darker brown near posterior

metatergal margins. Antenna long, slender (Fig. 69ana).

Paranota reduced with markedly oblique anterior shoulders,

R = 1.3 (Fig. 70ana); posterior corners strongly turned up (Fig.

4C), no marginal setae. Legs fairly slender, tarsus about as long

as femur, tibia with slight ventral distal swelling (Fig. 71 ana).

Spiracles apparently typical for genus but with a “foxtail” of

hair-like structures arising just anterior and ventral to anterior

spiracle, curving dorsally and posteriorly to obscure anterior

spiracle and terminating near posterior spiracle; a small clump

of hair-like structures arising just posterior and ventral to pos-

terior spiracle (Fig. 5C). Telopodite (Figs 15, 16) more or less

uniform in width, slightly curved posteriorly, reaching leg 5

when retracted, with unusually long, sparse setae. Solenomere

arising at about half the telopodite height, directed distally

before curving slightly posteriorly and laterally at about

half its length, terminating with small subapical projection at

about half the prefemoral process height. Tibiotarsus more or

less cylindrical, about half the solenomere diameter and

directed at almost a right angle to telopodite axis, the tip

turned distally and pointed with a small, blunt, subapical

projection on the anteromesal surface. Femoral process

arising proximal to level of solenomere origin, directed

distally, closely pressed to prefemoral process, blade-like with

a blunt, wide posterior projection at about half the process

length, terminating at about half the solenomere length.

Prefemoral process about two-thirds the width of telopodite

base at origin, narrowing and bending sharply laterally at

about half its length before expanding to very wide, flat-

tened tip curving postero-mesally and aligned parallel to

the telopodite axis, terminating in a comb with 20-30 long,

posteriorly directed teeth. Uncus prominent, arising at about

half the length of prefemoral process (just proximal to

solenomere tip), with a few small, blunt teeth subapically on

proximal edge.

Distribution and habitat. Known from c. 1300 km 2 in north-

west Tasmania, from Roger River West south to Piney Creek

(north of Zeehan) (Fig. 74), and from sea level to 450 m. L.

anas is an uncommon species and is likely to have been over-

looked outside its known range. It has mainly been found in wet

eucalypt forest and cool temperate rainforest, but two of the

paratypes are from partly wooded heathland near the coast at

Pieman Head. One specimen was taken from a cave near

Savage River.

Etymology. Latin anas (“duck”), noun in apposition, referring

to the type locality, Duck Creek.

New genus of millipedes

111

Lissodesmus bashfordi sp. nov.

Figures 17, 18, 69bas, 70bas, 71bas, 72 (map)

Lissodesmus sp. SE1. — Mesibov, 1996: 18.

Material examined. Holotype. Male, Australia, Tasmania. Huon River

(Manuka Road), DN764287 (43°05'38”S 146°42’36”E), 140 m,

15.V.1997, R. Mesibov, QVM 23:45945 (ex 23:40746).

Paratypes. Male, Dromedary Creek, EN098692 (42°43'46"S

147°07'H"E), 420 m, 19.iii.1992, R. Mesibov, QVM 23:16178;

female, same details, QVM 23:16179; male, Bracken Ridge,

DN897308 (43°04”31"S 146°52 ,, 25"E), 360 m, 17.U995,

R. Bashford, QVM 23:40745; male, Huon River (Manuka Road),

DN764287 (43°05’38"S 146°42'36"E), 140 m, 15.V.1997, R. Mesibov,

QVM 23:40746; 7 males, Huon River (Edwards Road), DN792288

(43°05'35"S 146°44'40"E), 100 m, 9.i.2001, R. Mesibov, QVM

23:45946, 2 dissected, in 95% ethanol; male, Huon River (Manuka

Road), DN765285 (43°05'45"S 146°42'40"E), 110 m, 3.ii.2001,

R. Mesibov, QVM 23:45947, in 95% ethanol; male, Judds Creek,

DN976476 (42°55'24"S 146°58T5"E), 390 m, 29.xi.2003, R. Mesibov

& K. Bonham, AM KS91167; female, same details, QVM 23:25601;

male, same details but DN970463 (42°56T0"S 146°57'49"E),

220 m, QVM 23:25603; male, Peak Rivulet, DN914020

(43°19’59"S 146°53’44"E), 140 m, 12.ix.2005, W. & L. Clarkson,

QVM 23:46135.

Description. Male c. 15 mm long, H = 1.4 mm. In alcohol,

well-coloured specimens under low magnification with very

pale brown body colour and red-brown speckling transversely

on metatergites, concentrated near posterior margins. Antenna

relatively short (Fig. 69bas). Paranota fairly wide, R = 1.5

(Fig. 70bas); posterior comers not turned up. Legs moderately

robust, tarsus about as long as femur, tibia with slight ventral

distal swelling (Fig. 71bas). Telopodite (Figs 17, 18) reaching

leg 5 when retracted. Solenomere arising at one-third the

telopodite height, directed posterodistally at about 45° to

telopodite axis, curving smoothly laterally and distally, the tip

bending abmptly mesally and armed subapically with a small,

bluntly pointed projection, terminating at about one-third the

prefemoral process height. Tibiotarsus more or less cylindrical,

narrower than solenomere, more or less parallel to solenomere

but about half its length, curving slightly laterally. Femoral

process arising far distal to solenomere origin at half to two-

thirds the telopodite length, slightly flattened near base and

curving outwards, distally expanding to a large, leaf-shaped

structure bent towards prefemoral process at nearly a right

angle and reaching as far distally as that process. Pre-

femoral process somewhat more than two-thirds the width of

telopodite base, bending sharply posteriorly about midway and

curving slightly mesally, terminating in large, flattened,

hand-shaped tip with long lateral “thumb” and several short,

marginal teeth as “fingers”. Uncus small, finger-like, arising

just distal to solenomere tip close to mesal edge of prefemoral

process.

Distribution and habitat. An uncommon species known only

from wet eucalypt forest over c. 500 km 2 in south-east

Tasmania (Fig. 72), from 100 to 420 m.

Etymology. Named for Richard Bashford, who has collected

this and many other invertebrate species in his long and

productive career in forest entomology in Tasmania.

Lissodesmus blackwoodensis sp. nov.

Figures 19, 20, 69bla, 70bla, 71bla, 79 (map)

Material examined. Holotype. Male, Australia, Victoria. 3 km NW of

Blackwood, 37°27'25'S 144°16'09"E, 560 m, 9.ix.2004, R. Mesibov &

T. Moule, NMV K-8933.

Paratypes. 3 males, details as for holotype, AM KS91168; 10 males,

details as for holotype, NMV K-8923 to K-8932; 13 females, details as

for holotype, NMV K-8934 to K-8946; 13 males, 1 km NE of

Blakeville, 37°29’45"S 144°13'07"E, 680 m, 10.ix.2004, R. Mesibov &

T. Moule, NMV K-8947 to K-8959; 8 females, same details, NMV

K-8960 to K-8967.

Description. Male c. 16 mm long, H = 1.4 mm. In alcohol,

well-coloured specimens under low magnification with very

pale brown body colour, red speckling dorsally on both pro-

zonites and on metatergites, a well-defined transverse band of

red speckling near posterior metatergal margins. Antenna mod-

erately long (Fig. 69bla). Paranota reduced but prominent,

R = 1.4 (Fig. 70bla); posterior comers not turned up, two incon-

spicuous posterior marginal teeth on each side. Legs robust,

tarsus about as long as femur (Fig. 71bla). Telopodite (Figs 19,

20) with sparse, long, setae extending posterolaterally to just

distal of tibiotarsus origin, reaching leg 4 when retracted.

Solenomere arising at one-third the telopodite height, directed

posterodistally at about 45° to telopodite axis, gradually curv-

ing distally, terminating with very small subapical projection at

one-quarter to one-third the prefemoral process height.

Tibiotarsus more or less parallel to solenomere but shorter,

about as wide as solenomere at base and widening distally,

forking near tip and terminating in two blunt points. Femoral

process arising proximal to solenomere origin, mesolaterally

flattened and pressed close at base to prefemoral process,

directed distally and widening before terminating in bluntly

pointed apex just distal to solenomere tip, anterior to a slightly

projecting “shoulder”. Prefemoral process very long, about

half as wide as telopodite base, bending laterally just distal

to solenomere tip, then bending mesally and curving

anterodistally before flexing at nearly 180° to point basally,

the lateral edge a comb of c. 40 long, mainly posteriorly

directed teeth for nearly its entire length, a few small teeth

on mesal edge close to apex. Uncus small, arising near base of

prefemoral process on a ridge formed by the mesal edge of the

process.

Distribution and habitat. Known from wet eucalypt forest at

two localities 6 km apart in the southern portion of Wombat

State Forest (Fig. 78).

Etymology. Named for Blackwood, the town closest to the type

locality, 65 km north-west of Melbourne.

Lissodesmus catrionae sp. nov.

Figures 21, 22, 69cat, 70cat, 71cat, 78 (map)

Material examined. Holotype. Male, Australia, Victoria. Mt Cole,

37°16'44"S 143°14'23"E, 900 m, 8.ix.2004, R. Mesibov & T. Moule,

NMV K-8968.

Paratypes. 3 males, details as for holotype, NMV K-8969 to

K-8971, 2 dissected; 7 females, details as for holotype, NMV K-8972

to K-8978; 2 males, Mt Cole, 37°17'09”S 143°14T1"E, 850 m,

112

Robert Mesibov

8.ix.2004, R. Mesibov & T. Moule, AM KS91169; 3 males, same

details, NMV K-8979 to K-8981; female, same details, NMV K-8982.

Description. Male c. 16 mm long, H = 1.3 mm. In alcohol,

well-coloured specimens under low magnification with very

pale brown body colour, sparse red speckling on metatergites,

paranota near-white. Antennae moderately long (Fig. 69cat),

about 1.75x a socket diameter apart. Paranota fairly wide,

R = 1.5, anterior shoulders projecting forward (Fig. 70cat); pos-

terior corners not turned up. Legs with high, rounded prefemur

(Fig. 71cat), tarsus somewhat longer than femur. Telopodite

(Figs 21, 22) with abrupt narrowing at prefemoral process

origin, with sparse, long setae extending posterolaterally to just

distal of tibiotarsus origin, reaching leg 4 when retracted.

Solenomere arising at one-third the telopodite height, directed

posterodistally at about 45° to telopodite axis and gradually

curving distally, terminating with a small subapical projection

at one-third to half the prefemoral process height. Tibiotarsus

more or less parallel to solenomere, slightly flattened and wider

than solenomere and about as long, bending slightly poster-

iorly at about half its length, flattening and widening apically

with a large, rounded notch opening anteromesally and a small,

bluntly pointed projection on anterior (distal) surface at about

two-thirds the process length. Femoral process a very nar-

row, tapering rod arising at about half the telopodite length,

directed distally and terminating just distal to solenomere tip.

Prefemoral process arising from lateral half of telopodite

base, a little less than half the width of the base, narrow,

very long, bending slightly posterodistally at about half its

length and flexing 180° at three-quarters its length to ter-

minate in a bluntly pointed apex not far distal to

solenomere tip, the posterolateral edge in the distal half of

the process a comb of c. 25-30 long, mainly proximally

directed teeth. Uncus large, arising at about one-quarter the

length of prefemoral process (just proximal to solenomere tip).

Distribution and habitat. Known from wet eucalypt forest on

Mt Cole, 50 km north-west of Ballarat (Fig. 78).

Etymology. Named for my wife, Catriona (Trina) Moule, in

gratitude for her help in the field in Tasmania and Victoria.

Lissodesmus clivulus sp. nov.

Figures 23, 24, 69cli, 70cli, 7 lcli, 73 (map)

Material examined. Holotype. Male, Australia, Tasmania. The Clump,

CQ213361 (41°12'23"S 144°52 , 06"E), 190 m, 6.H.1992, R. Mesibov,

QVM 23:45821 (ex QVM 23:17667).

Paratypes. 2 males, details as for holotype, AM KS91170 (ex QVM

23:17667); 11 males, details as for holotype, QVM 23:17667, 3 dis-

sected; 16 females, details as for holotype, QVM 23:17675; male,

south of Italian River, CQ180045 (41°29’25"S 144°49'11"E), 10 m,

30.iv.1993, R. Mesibov, QVM 23:17664.

Other material. 12 males, 26 females and 1 juvenile from Balfour,

Brooks Creek, Dawson River, Little Eel Creek, Mt Frankland, Possum

Creek, Sardine Creek, Sawards Creek, Sundown Creek, Sundown

Point, Temma and The Clump (see “ Lissodesmus supplement” for

details).

Description. Male c. 17 mm long, H - 1.6 mm. In alcohol,

well-coloured specimens under low magnification with light

brown body colour (unusually, extending to legs and antennae),

slightly darker brown transversely on metatergites near poster-

ior margins and distally on antennae. Antenna long, slender

(Fig. 69cli). Paranota reduced but prominent, R = 1.4 (Fig.

70cli); posterior corners not turned up. Legs long, slender,

tarsus about as long as femur (Fig. 7 lcli). Telopodite (Figs 23,

24) almost reaching leg 5 when retracted. Solenomere arising at

half the telopodite height, directed posteriorly at a large angle

(c. 60°) to telopodite axis, curving slightly laterally, terminating

with toothed subapical collar at about one-quarter the

prefemoral process height. Tibiotarsus about as wide as

solenomere, slightly flattened and tapered near tip, directed

posteriorly at almost a right angle to telopodite, curving

slightly distally near tip. Femoral process arising at level of

solenomere origin, a straight, distally directed blade with a

pointed tip, terminating well distal to solenomere tip; at one-

quarter to one-third its length with a short, narrow, bluntly

tipped branch directed posteriorly and curving distally near its

tip. Prefemoral process about half the width of the telopodite

base at its origin, expanded past this point, narrowing near apex

and bending posteriorly, a few short, large, rounded, proxim-

ally directed teeth subapically, the apex flattened and slightly

crenulate. Two inconspicuous unci at about half the length of

the prefemoral process, not obvious in some specimens.

Distribution and habitat. Known from c. 300 km 2 on the west

coast of Tasmania (Fig. 73), from just north of the Arthur River

to just south of the Italian River, inland to Mt Frankland, and

from sea level to 400 m. Despite its small range, L. clivulus is

sometimes locally abundant. It occurs in wet forest, coastal

woodland, tea-tree forest and tea-tree scrub, and (unusually for

Lissodesmus species) has also been found in low heath (on Mt

Frankland, near Balfour).

Etymology. Latin clivulus (“small hill”), noun in apposition.

Named for the type locality, The Clump, a low hill covered with

a clump of tall eucalypt trees. Because the surrounding country

is flat and heathy, The Clump has long been a landmark on the

northern part of the Tasmanian west coast.

Remarks. L. clivulus is similar to L. latus in gonopod structure

but the prefemoral process is shorter, the prefemoral process tip

less expanded and the tibiotarsus bent posteriorly at nearly a

right angle to the telopodite. The two species co-occur along

the coast between the Arthur and Pieman Rivers, and can be

distinguished in the field by coloration and by the smoother

paranotal margin in L. clivulus.

Lissodesmus cognatus sp. nov.

Figures 25, 26, 27, 69cog, 70cog, 71cog, 76 (map)

Lissodesmus sp. NE3. — Mesibov, 1996: 17. — Mesibov, 1997:

567.— Mesibov, 2003a: 209.

Material examined. Holotype. Male, Australia, Tasmania. Weavers

Creek, EQ330091 (41°28’07"S 147°23’42"E), 380 m, 31.vii.1994, R.

Mesibov & T. Moule, QVM 23:15287.

Paratypes. 2 males, North Esk River, EQ336079 (41°28'46"S

147°24'08"E), 490 m, 23.ii.1992, R. Mesibov, QVM 23:15284; 2

males, “Elkington” property, EP3 12979 (41°34’11"S 147°22'27"E),

350 m, 1 8.iii. 1992, R. Mesibov, AM KS91171 (ex QVM 23:15283); 4

New genus of millipedes

113

males, same details, QVM 23:15283, 2 dissected; 2 females, Weavers

Creek, EQ307122 (41°26’27"S 147°22'02"E), 680 m, 19.vii.1994,

R. Mesibov, QVM 23: 15289; male, Tower Hill, EP708983 (41°33'49"S

147°50'56"E), 720 m, 31.xii.1998, R. Mesibov & K. Bonham, QVM

23:40758; 10 males, Coxs Creek, EQ767040 (41°30'42"S

147°55'08"E), 480 m, ii.2001, R. Bashford, pitfall sample, QVM

23:24738, 2 dissected.

Other material. 14 males, 2 females and 5 juveniles from “Aplico”

property, Coxs Creek, Grants Creek, Long Gully Creek, Musselboro,

Rabbity Creek, Tower Hill and Weavers Creek (see u Lissodesmus

supplement” for details).

Description. Male c. 14 mm long, H = 1.3 mm. In alcohol,

well-coloured specimens under low magnification near-white

in body colour with red tinge transversely near posterior

margins of metatergites and dorsa of prozonites. Antennae

long, slender (Fig. 69cog), about 2.5x a socket diameter apart.

Paranota reduced, R = 1.3 (Fig. 70cog); posterior corners

turned up slightly. Legs fairly slender, tarsus about as long as

femur, tibia with slight ventral distal swelling (Fig. 71cog).

Telopodite (Figs 25, 26, 27) reaching leg 5 when retracted.

Solenomere arising at half the telopodite height, directed pos-

terodistally at a small angle to telopodite axis, terminating with

toothed subapical collar at about half the prefemoral process

height. Tibiotarsus a slightly flattened, bluntly pointed rod

about as wide as solenomere but shorter, more or less parallel

to solenomere but gradually curving laterally. Femoral process

arising proximal to solenomere origin (at about one-third

the telopodite length), pressed close to prefemoral process

proximally, forked near base; branches more or less equal,

blade-like, pointed, the anterior branch directed distally

and terminating at level of solenomere tip, the posterior

branch curving anteriorly, its tip sometimes lying between

anterior branch and prefemoral process. Prefemoral

process about half the width of telopodite base, bending

posteriorly at about two-thirds its length and tapering, the

flexed distal section bearing a comb of c. 15-20 large, irreg-

ular, proximally directed teeth. Uncus prominent, arising at

about half the prefemoral process length (about the level of the

solenomere tip) on well-defined longitudinal ridge near lateral

edge of process.

Distribution and habitat. An uncommon species known from

wet eucalypt forest in two disjunct areas c. 35 km apart in

north-east Tasmania (Fig. 76): south and west of Mt Barrow

(c. 30 km 2 ), and north and east of Tower Hill (c. 15 km 2 ).

Etymology. Latin cognatus (“kindred”), adjective. At first

glance, L. cognatus seems to be closely related to both

L. alisonae and L. hamatus.

Remarks. Furry individuals of L. cognatus have been found

near the North Esk River, Weavers Creek and Musselboro.

Specimens from the “Elkington” property, a few kilometres to

the south in the South Esk River catchment, are non-furry, as

are all specimens from the Tower Hill portion of the range.

I have previously speculated (Mesibov, 1997, 2003a) that

L. cognatus is a stabilised hybrid of L. alisonae and L. hama-

tus, i.e. a product of reticulate evolution. The two putative par-

ents meet with minimal overlap in the western block of the

L. cognatus range, and L. hamatus occurs in the eastern block.

Lissodesmus cornutus sp. nov.

Figures 28, 29, 30, 69cor, 70cor, 71cor, 74 (map)

Lissodesmus sp. SW1. — Mesibov, 1996: 18.

Material examined. Holotype. Male, Australia, Tasmania. Scotts Peak

Dam, DN425343 (43°02'29"S 146°17'38"E), 280 m, pitfall samples

WY and WZ collected 12-16.xi.2001, D. Driscoll, QVM 23:45827 (ex

QVM 23:24824).

Paratypes. Male, Birchs Inlet, approx. CN753875 (42°33'16"S

145°28'51"E), 18.X.1993, J. Griffith, QVM 23:12036; 2 males, details

as for holotype, AM KS91172 (ex QVM 23:24824); 5 males, details as

for holotype, QVM 23:24824, 2 dissected; female, W of Strathgordon,

DN175655 (42°45'30"S 145°59'30"E), 410 m, l.x.2003, R. Mesibov,

QVM 23:25476.

Other material. 63 males and at least 9 females and juveniles from

Acheron Cave area, Darwin, Deadmans Bay, Denison River valley,

Edgar Dam, Franklin River valley, Gelignite Creek, Gordon River

valley, Hibbs Lagoon, Kutikina Cave area, Mt McCutcheon, Olga

River valley, Orange River valley, Scotts Peak Dam and Wedge Inlet

(see “ Lissodesmus supplement” for details).

Description. Male c. 17 mm long, #=1.6 mm. In alcohol, most

specimens under low magnification near-white, the only pig-

mentation a red tinge distally on antennae. Antennae long and

slender (Fig. 69cor), about 2.25x a socket diameter apart.

Paranota wide, highly modified, R = 1.5 (Fig. 70cor), poster-

ior corners extended as large, upwardly curved, conical

structures with ozopore opening midway to tip at anterior

end of lateral groove (Fig. 28). Legs slender, tarsus about as

long as femur, tibia with slight ventral distal swelling (Fig.

71cor), sphaerotrichomes lacking on leg 6 femur and prefemur.

Telopodite (Figs 29, 30) somewhat sinuous, widening just

proximal to prefemoral process origin, with sparse, long setae

extending posterolaterally to just distal of tibiotarsus origin,

reaching leg 6 when retracted. Solenomere arising at one-third

the telopodite height, directed posteriorly at nearly a right angle

to telopodite axis, curving distally and laterally, terminating

with a small, pointed subapical projection at about one-third the

prefemoral process height. Tibiotarsus much wider and some-

what longer than solenomere, directed more or less parallel to

solenomere in mesal view but strongly curved laterally,

flattened, widening at the apex and terminating in two large,

widely spaced, bluntly pointed teeth with 1 or 2 smaller teeth

between. Femoral process arising well distal to solenomere

origin at about one-quarter the prefemoral process height,

blade-like, pressed close at its base to the prefemoral process,

directed posterodistally and curving distally and slightly mes-

ally to terminate at two-thirds the prefemoral process height

with two small apical teeth, with a large, posterobasally

directed branch arising at nearly half the process length.

Prefemoral process nearly as wide as telopodite base below

solenomere origin, curving mesally, then laterally, then again

mesally, flexing slightly posteriorly at about two-thirds its length

and tapering slightly to a broad, rounded tip, with a comb of c.

15 long, posterobasally directed teeth on mesal edge of flexed

distal section. Uncus large, arising centrally at about half the

prefemoral process height (well distal of solenomere tip).

Distribution and habitat. Known from wet eucalypt forest and

cool temperate rainforest over c. 5000 km 2 in south-west

114

Robert Mesibov

Tasmania (Fig. 74), from Darwin in the north to Deadmans Bay

in the south, east to the Scotts Peak Dam Road, and from sea

level to at least 600 m.

Etymology. Latin cornutus (“homed”), adjective, for the horned

shape of the posterior projections of the paranota.

Lissodesmus devexus sp. nov.

Figures 31, 32, 33, 69dev, 70dev, 71 dev, 74 (map)

Lissodesmus sp. NE2. — Mesibov, 1998: 155. — Mesibov, 1999: 252.

Material examined. Holotype. Male, Australia, Tasmania. Lebrina,

EQ 168424 (41°10'09"S 147°12'00"E), 240 m, 5.i.l993, R. Mesibov,

QVM 23:45948 (ex QVM 23:15515).

Paratypes. Male, Rattler Hill, EQ744353 (41°13’48"S 147°53T5"E),

650 m, 29.viii.1990, R. Mesibov, QVM 23:15512, dissected; 2 males,

details as for holotype, AM KS91173 (ex QVM 23:15515); 2 males,

details as for holotype, QVM 23:15515, 1 dissected; 8 stadium

7 males, 2 stadium 7 females, details as for holotype, QVM 23:15538;

3 females, Retreat, EQ153423 (41°10T3"S 147°10’56 M E), 320 m,

2.vii.l993, T. Kingston, QVM 23:21554; 2 males, same details but

EQ137463 (41°08 , 03”S 147°09 , 47”E), 300 m, QVM 23:21534, 1 dis-

sected; 4 males, Mt Roland, DQ402119 (41 0 26'31”S 146°17'02"E),

310 m, 5.vi.l994, R. Mesibov, QVM 23:15506, 1 dissected; 8 males,

Shepherds Rivulet, EQ217388 (41°12'06"S 147°15'31"E), 150 m,

6.vii.l994, R. Mesibov, QVM 23:15509, 1 dissected; male, Lowes

Mount, EP331953 (41°35’35”S 147°23'49"E), 430 m, 5.vii.l995,

R. Mesibov, QVM 23:21542, dissected.

Other material. 115 males, 71 females and 45 juveniles from 77

localities (see “ Lissodesmus supplement” for details).

Description. Male c. 12 mm long, 77= 1.3 mm. In alcohol, most

specimens under low magnification a uniform very pale brown

in body colour. Antenna slender (Fig. 69dev). Paranota slightly

reduced, R = 1.5 (Fig. 70dev), posterior corners turned up. Legs

slender, tarsus about as long as femur (Fig. 71 dev), sphaerotri-

chomes on tibia and tarsus only of leg 6. Telopodite (Figs 31,

32, 33) widest at prefemoral process origin, almost reaching leg

5 when retracted. Solenomere arising at slightly more than

half the telopodite height, directed posterodistally at a large

angle (c. 60°) to the telopodite axis, terminating without a

subapical process at about two-thirds the telopodite height

in a sharp distal bend. Tibiotarsus a somewhat flattened rod

slightly larger than solenomere and about as long, straight

and directed posterobasally at a large angle (greater than

90°) to the telopodite axis, widening and flattening near the

tip and terminating in a sharp mesal bend, the tip some-

times broadly notched, the midsection with numerous

annular “wrinkles” (Fig. 31). Femoral process arising at about

the level of solenomere origin, small and blade-like with a

somewhat sinuous outline, directed distally, not closely pressed

to prefemoral process, terminating at about three-quarters the

prefemoral process height (well distal to solenomere tip).

Prefemoral process about as wide at its base as the telopodite

base, tapering and sharply flexed posteriorly at its midpoint, the

tip variably armed with teeth on posterior (proximally facing)

surface (Figs 32, 33). Uncus arising on mesal edge of

prefemoral process just proximal to flexure in latter, variably

large (Figs 32, 33) but typically directed posterodistally and in

some forms (see left gonopod in Fig. 31) paired with

prefemoral process tip as a major apical feature of telopodite.

Distribution and habitat. In wet eucalypt forest and cool tem-

perate rainforest over c. 3000 km 2 in the higher-rainfall parts of

northern and north-east Tasmania (Fig. 74), from Browns

Creek near Port Sorell east to Weldborough, south to Lowes

Mount near Deddington, and from near sea level to at least 900

m. Populations around Mt Roland and Liena are apparently dis-

junct (Fig. 74), being separated by about 40 km from known

L. devexus sites in the West Tamar region. L. devexus is

gregarious and often locally abundant.

Etymology. Latin devexus (“sloping down”), adjective, for the

orientation of the tibiotarsus on the gonopod.

Lissodesmus dignomontis sp. nov.

Figures 34, 35, 69dig, 70dig, 71dig, 78 (map)

Material examined. Holotype. Male, Australia, Victoria. “Lapoinya”

property, Yarragon South, 38°14'40"S, 146°05’30"E, 360 m,

17.iv.2005, R. Mesibov, NMV K-9663.

Paratypes. Male, west of Allambee, 38°16'55"S 146°00T2"E, 430

m, 30.ix.2004, R. Mesibov & T. Moule, NMV K-9507, dissected;

2 males, details as for holotype, AM KS91428; 8 males, details as for

holotype, NMV K-9664 to K-9671, 1 dissected; male, Allambee,

38°16'27"S 146°03'03"E, 450 m, 22.iv.2005, R. Mesibov, NMV

K-9682; 2 males, Mt Worth, 38°16T4"S 145°58’54"E, 440 m,

22.iv.2005, R. Mesibov, NMV K-9673 and K-9674.

Description. Male c. 15 mm long, H = 1.4 mm. In alcohol,

well-coloured specimens under low magnification very pale

brown in body colour with two discrete, narrow, transverse

bands of red-brown posteriorly on metatergites. Antenna slen-

der (Fig. 69dig). Paranota fairly wide, R = 1.5, with about 4

posterior marginal teeth (Fig. 70dig), posterior corners not

turned up. Legs robust, tarsus slightly longer than femur, tibia

with prominent ventral distal swelling (Fig. 71dig). Telopodite

(Figs 34, 35) reaching leg 5 when retracted. Solenomere arising

at half the telopodite height, directed posterodistally at a small

angle to telopodite axis, terminating with small subapical pro-

jection at just over half the prefemoral process height.

Tibiotarsus rod-like, directed posterodistally and between

one-third and half as long as solenomere. Femoral process

arising distal to solenomere origin at about one-third the

prefemoral process height, blade-like, curving slightly antero-

laterally and tapering to a blunt point, terminating at about

the level of the prefemoral process tip. Prefemoral process

about half as wide at origin as telopodite base, strongly

tapering distally, curving posteriorly near tip and termin-

ating in flattened, blunt point. Uncus prominent, arising at

about half the prefemoral process height (just basal to

solenomere tip).

Distribution and habitat. Known only from wet eucalypt

forest at four sites within a c. 25 km 2 area near Mt Worth

in West Gippsland (Fig. 78). At three of the four sites

L. dignomontis co-occurs with L. gippslandicus, and at

Yarragon South it co-occurs with L. johnsi.

Etymology. Latin dignus (“worthy”) + montis (“mountain”),

noun in apposition. Named for Mt Worth, one of the known

localities for this rare species.

New genus of millipedes

115

Lissodesmus gippslandicus sp. nov.

Figures 36, 37, 69gip, 70gip, 71gip, 79 (map)

Material examined. Holotype. Male, Australia, Victoria. Mt Fatigue,

38°34’13”S 146°18’25”E, 570 m, 28.ix.2004, R. Mesibov & T. Moule,

NMV K-8990.

Paratypes. 2 males, details as for holotype, AM KS91174; male,

details as for holotype, NMV K-8991; 6 females, details as for holo-

type, NMV K-8992 to K-8997; 9 males, 4 females, Loop Track, E of

Allambee, 38°15'52"S 146°04'27"E, 440 m, 21 .xii.2004, R. Mesibov,

NMV K-9483 to K-9495, 2 males dissected.

Other material. 46 males and 12 females from Allambee, Allambee

South, Balook, Darlimurla, Mirboo North, Mt Worth, Narracan, Tarra-

Bulga National Park, Thorpdale and Yarragon South (see

“ Lissodesmus supplement” for details).

Description. Male c. 18 mm long, H = 1.6 mm. In alcohol,

well-coloured specimens under low magnification with pale

brown body colour, red-purple speckling on metatergites and

intense red-purple transverse banding along posterior margins

of prozonites. Antenna with relatively large antennomere 6

(Fig. 69gip). Paranota fairly wide with strongly produced anter-

ior “shoulders” and two inconspicuous posterior marginal

teeth, R= 1.5 (Fig. 70gip), posterior corners not turned up. Legs

robust, tarsus longer than femur, tibia with prominent ventral

distal swelling (Fig. 71gip). Telopodite (Figs 36, 37) almost

reaching leg 4 when retracted. Solenomere arising at about half

the telopodite height, directed distally with a slight posterior

curvature, terminating with prominent subapical collar at about

half the prefemoral process height. Tibiotarsus origin on pos-

teromesal surface of telopodite, not close to solenomere origin;

tibiotarsus a thin, pointed rod directed posterodistally at a small

angle to telopodite axis, about one-quarter the length of the

solenomere. Femoral process arising well proximal to

solenomere origin, blade-like with a deeply notched tip,

curved anteriorly and pressed close basally to prefemoral

process, terminating at less than one-quarter the

prefemoral process height (well proximal to solenomere

tip). Prefemoral process about half as wide at origin as

telopodite base, tapering slightly to mid-length but distally

widening and flexing posteromesally, the tip pointed postero-

mesally, the lateral edge of the process a comb of c. 20 long,

mainly posterobasally directed teeth from about two-thirds the

process length. Uncus prominent, arising near base of prefemoral

process on mesal side, with a widened, deeply notched tip.

Distribution and habitat. In wet eucalypt forest in West and

South Gippsland (Fig. 79), often locally abundant. The

Thorpdale specimens were collected in 1899, before the

densely forested Thorpdale area was cleared for farming.

The pre-European range of this species may have been as much

as 1000 km 2 , but is now possibly only a third of that figure.

Etymology. Named for the Gippsland district, the southern and

western parts of which are home to this species.

Lissodesmus hamatus sp. nov.

Figures 4B, 6B, 38, 39, 69ham, 70ham, 71ham, 77 (map)

Lissodesmus sp. El. — Mesibov, 1994: 134. — Mesibov, 1996: 17. —

Mesibov, 1997: 567.— Mesibov, 1999: 252.

Material examined. Holotype. Male, Australia, Tasmania. Mt Hobbs,

EN476926 (42°31'02"S 147°34’46”E), 580 m, 21.iii.1992, R. Mesibov,

QVM 23:45944 (ex QVM 23:16073).

Paratypes. 7 males, details as for holotype, QVM 23:16073;

2 males, details as for holotype, AM KS91175 (ex QVM 23:16073);

4 stadium 7 males, 1 stadium 6 male, 7 females, 1 stadium 7 female,

details as for holotype, QVM 23:15966; 4 males, Ringarooma Tier,

EQ829796 (40°49'49"S 147°58'59"E), 80 m, 19.ix.1992, R. Mesibov

& T. Moule, QVM 23:16076, 2 dissected; 3 pairs in copula , same

details, QVM 23:16157; 4 females, same details, QVM 23:15845;

9 males, Lagoon of Islands, DP947376 (42°06'48"S 146°56'09"E), 750

m, 5.iii.l995, R. Mesibov, QVM 23:21349, 1 dissected; female, same

details, QVM 23:21505.

Other material. 371 males, 382 females and 238 juveniles from 234

localities (see “ Lissodesmus supplement” for details).

Description. Male c. 16 mm long, H = 1.7 mm (see Remarks).

In alcohol, well-coloured specimens under low magnification

with pale brown body colour and light red speckling dorsally

on both prozonites and metazonites, more intense medially near

posterior margins. Antenna fairly long and slender (Fig.

69ham). Paranota reduced, R = 1.3 (Fig. 70ham), posterior cor-

ners slightly turned up (Fig. 4B). Limbus with toothed elements

(Fig. 6B). Legs fairly slender, tarsus slightly longer than femur,

tibia with slight ventral distal swelling (Fig. 71 ham). Epiproct

with prominent paired, bluntly rounded projections. Telopodite

(Figs 38, 39) reaching leg 5 when retracted. Solenomere arising

at about half the telopodite height, directed posterodistally at

c. 45° to telopodite axis, terminating with toothed subapical

collar at one-third to half the prefemoral process height.

Tibiotarsus rod-like, pointed, about two-thirds the solenomere

length and apically almost touching solenomere. Femoral

process arising at level of solenomere origin, bladelike and

anteroposteriorly flattened with a short, shelf-like posterior

branch at one-third the process length, bluntly tipped and ter-

minating at half the prefemoral process height, just distal to

solenomere tip. Prefemoral process about two-thirds the

width of telopodite base, bending laterally at about two-

thirds its height with a prominent “shoulder” process on

mesal side of bend, curving posteriorly distal to bend with a

variable number of long, mainly posterobasally directed

teeth forming a comb on lateral edge of tip. Uncus promi-

nent, arising centrally at about half the prefemoral process

height, with a coarsely toothed outer edge.

Distribution and habitat. Common in dry and wet eucalypt for-

est over c. 16 000 km 2 in eastern Tasmania (Fig. 77), from

Ringarooma Tier in the far north-east to Dunalley in the south-

east, from the eastern coast (including Maria Island) across the

Eastern Tiers and the Midlands to the eastern portion of the

Central Plateau, and from sea level to at least 1070 m. In places

along the northern edge of its range, L. hamatus is parapatric

with L. adrianae and L. alisonae (Mesibov, 1997).

Etymology. Latin hamatus (“hooked”), adjective, for the

upturned comers of the posterior projections of the paranota.

The name was suggested by Peter Johns (in litt.), who collect-

ed L. hamatus near Triabunna in 1972.

Remarks. Gonopod form varies over the range of this species.

The number of subterminal teeth on the prefemoral process

116

Robert Mesibov

ranges from 6 to 10, and the width of the flexed tip of the

process can be proportionately greater than shown for speci-

mens from the type locality. In the eastern portion of the range

the short, posterior branch of the femoral process often has a

bluntly forked tip. H ranges widely, from c. 1.2 mm in drier

and more southerly areas to c. 1.8 mm in wetter and more

northerly areas; larger males have markedly more swollen legs.

Lissodesmus horridomontis sp. nov.

Figures 40, 41, 69hor, 70hor, 71hor, 73 (map)

Lissodesmus sp. NE5. — Mesibov, 1994: 134.

Material examined. Holotype. Male, Australia, Tasmania. Mt Horror,

EQ588513 (41°05T4"S 147°42’00"E), 200 m, 24.iv.1993, R. Mesibov,

QVM 23:45829 (ex QYM 23:15491);

Paratypes. Male, Friend Creek, EQ862540 (41°03'37"S

148 o 01’33"E), 150 m, 25.xi.1992, QYM 23:15492; 4 males, Mt Horror,

EQ604547 (41°03'23"S 147°43'07"E), 220 m, 18.iii.1993, R. Mesibov,

QVM 23:15490; 7 females, same details, QVM 23:15484; 7 males,

Speck Creek, EQ566549 (41°03T8"S 147°40'24"E), 290 m,

1 8.iii. 1993, QVM 23:15497; 2 males, details as for holotype, AM

KS91176 (ex QVM 23:15491); 2 males, details as for holotype, QVM

23:15491.

Other material. 21 males, 17 females and 5 juveniles from Connors

Road, Friend Creek, Martins Hill, Mt Horror, Oxberry Creek, Pearly

Brook, Speck Creek and Tomahawk River (see “ Lissodesmus

supplement” for details).

Description. Male c. 13 mm long, H= 1.2 mm. In alcohol, most

specimens under low magnification uniformly near-white.

Antenna slender, not strongly clavate (Fig. 69hor). Paranota

reduced, R = 1.3 (Fig. 70hor), posterior comers slightly turned

up. Legs robust, tarsus shorter than femur (Fig. 71hor),

sphaerotrichomes lacking on leg 6 femur and prefemur.

Epiproct with prominent paired, bluntly rounded projections.

Telopodite (Figs 40, 41) reaching leg 5 when retracted.

Solenomere arising at one-third to half the telopodite height,

directed posterodistally at somewhat less than 45° to telopodite

axis, terminating without subapical projection at slightly less

than half the prefemoral process height. Tibiotarsus about as

large as solenomere but slightly shorter, directed more or less

parallel to solenomere, flattened posterobasally and terminating

in a variably dentate edge with a small, bluntly pointed sub-

apical projection on anterolateral surface. Femoral process aris-

ing at level of solenomere origin, straight, pointed, blade-like,

directed distally and terminating at level of solenomere tip.

Prefemoral process about as wide at origin as telopodite

base, tapering slightly and bending laterally at about mid-

height with a coarsely toothed “shoulder” on the mesal side

of bend, curving posteromesally with the tip flexed later-

ally at almost 180°, the tip bearing a comb of c. 10-15 long,

basally directed teeth. Uncus prominent, arising just proximal

to lateral bend in prefemoral process (just distal to solenomere

tip), outer edge coarsely dentate.

Distribution and habitat. Known from wet eucalypt forest and

cool temperate rainforest over c. 150 km 2 in north-east

Tasmania (Fig. 73), from Williams Hill east to Old Chum Dam,

and from 80 to 650 m. L. horridomontis is abundant in the

western portion of its small range.

Etymology. Latin horridus (“frightful”) + montis (“mountain”),

noun in apposition. Named for the type locality, Mt Horror.

Lissodesmus inopinatus sp. nov.

Figures 42, 43, 69ino, 70ino, 7 lino, 72 (map)

Lissodesmus sp. E2. — Mesibov, 1996: 17.

Material examined. Holotype. Male, Australia, Tasmania. Halls Creek,

EP660382 (42°06'19"S 147°47'53"E), 540 m, 22.iii.1992, R. Mesibov,

QVM 23:45828 (ex QVM 23:16164).

Paratypes. 2 males, details as for holotype, AM KS91177 (ex QVM

23:16164); 7 males, details as for holotype, QVM 23:16164, 2 dissect-

ed; 5 females, 1 stadium VII female, details as for holotype, QVM

23:16168; male, Pinnacles Creek, EP528531 (41°58T9"S

147 0 38'14"E), 520 m, 22.iii.1992, R. Mesibov, QVM 23:16165; male,

Mt Ponsonby, EN441966 (42°28'53"S 147°32T1"E), 610 m, 3.ix.2000,

R. Mesibov, QVM 23:41984; male, Anglers Creek, EP697230

(42°14'30"S 147°50'41"E), 570 m, 7.X.2001, R. Mesibov & T. Moule,

QVM 23:24858.

Other material. 3 males, 16 females and 8 juveniles from Anglers

Creek, Mt Mismanagement, Parramores Tier, Pinnacles Creek and

Rocka Rivulet (see “ Lissodesmus supplement” for details).

Description. Male c. 17 mm long, H = 1.7 mm. In alcohol,

well-coloured specimens under low magnification with very

pale brown body colour, faintly reddish and more intense medi-

ally and posteriorly on metatergites. Antenna long and slender

(Fig. 69ino). Paranota reduced, R = 1.3 (Fig. 70ino), posterior

comers turned up. Legs fairly slender, tarsus shorter than

femur, tibia with slight ventral distal swelling (Fig. 7 lino).

Telopodite (Figs 42, 43) widest at prefemoral process origin,

reaching leg 5 when retracted. Solenomere arising at one-third

the telopodite length, directed posterodistally at a small angle

(c. 30°) to telopodite axis, terminating with toothed subapical

collar at just under half the prefemoral process height.

Tibiotarsus nearly as thick as solenomere but slightly shorter,

directed posterodistally and slightly laterally at acute angle to

solenomere, flattened and bluntly pointed. Femoral process

arising at level of solenomere origin, forked near base, both

branches blade-like, pointed; anterior branch directed distally

and pressed close to prefemoral process, terminating at about

one-third the prefemoral process height (proximal to

solenomere tip); posterior branch smaller than anterior branch,

curved anterodistally. Prefemoral process at base about two-

thirds the width of telopodite base proximal to solenomere

origin, bending sharply posterolaterally at about two-thirds

its height with a large, mesolaterally flattened, broadly

toothed projection on mesal edge of prefemoral process at

bend, the distal portion of process with c. 10-20 teeth lying

at a small angle to process axis, the tip also toothed. Uncus

prominent, arising on mesal edge of prefemoral process at

about half the process height (just distal to solenomere tip).

Distribution and habitat. An uncommon species in dry and wet

eucalypt forest over c. 1000 km 2 in the Eastern Tiers of Tas-

mania (Fig. 72), from Pinnacles Creek (c. 15 km E of Campbell

Town) south to Mt Ponsonby and east to Anglers Creek.

Etymology. Latin inopinatus (“unexpected”), adjective. While

carrying out a Tasmania- wide millipede survey in the 1990s,

New genus of millipedes

117

the only Lissodesmus species I expected to find in the L.

inopinatus range was L. hamatus.

Remarks. L. inopinatus varies considerably in overall size

across its range.

Lissodesmus johnsi sp. nov.

Figures 44, 45, 69joh, 70joh, 71joh, 78 (map)

Material examined. Holotype. Male, Australia, Victoria. Uralla Nature

Reserve, Trafalgar, 38°13’36"S 146°08'53"E, 140 m, 19.iv.2005,

R. Mesibov, NMV K-9682.

Paratypes. Male, near Trafalgar, Gippsland, viii.1890, W. Kershaw,

NMV K-9506, dissected; male, “Lapoinya” property, Yarragon South,

38°14'30"S 146°05'34 M E, 370 m, 17.iv.2005, R. Mesibov, NMV

K-9681; 2 males, details as for holotype, AM KS91429; 10 males,

details as for holotype, NMV K-9683 to K-9692, 1 dissected;

9 females, details as for holotype, NMV K-9693 to K-9701.

Description. Male c. 14 mm long, H= 1.2 mm. In alcohol, well-

coloured specimens under low magnification very pale brown

in body colour with red-brown speckling transversely on

posterior half of metatergites. Antenna short, moderately slen-

der (Fig. 69joh). Paranota prominent but reduced, R= 1.4, with

4-5 posterior marginal teeth (Fig. 71joh), posterior corners not

turned up. Legs short, robust, tarsus about as long as femur,

tibia with prominent ventral distal swelling (Fig. 70joh).

Telopodite (Figs 44, 45) posteriorly excavated at base, reaching

leg 5 when retracted. Solenomere arising at just over half the

telopodite height, directed posterodistally at a small angle to

telopodite axis, terminating at somewhat more than half the

prefemoral process height with flattened, posterolaterally

flexed tip. Tibiotarsus broad and flattened, a marginally

toothed fan in posterior view, directed distally and about

half as long as solenomere. Femoral process arising well

distal to solenomere origin at about half the prefemoral process

height, blade-like, slightly swollen distally but tapering to a

blunt point, terminating distal to prefemoral process tip.

Prefemoral process about half as wide at origin as

telopodite base, curving slightly mesally, sharply tapering

distally and terminating in a simple blunt point. Uncus

prominent, arising at about two-thirds the prefemoral process

height (at about the level of solenomere tip).

Distribution and habitat. Known only from wet eucalypt forest

in a small area near Trafalgar in West Gippsland (Fig. 78); the

two contemporary sites are c. 5 km apart. At the Yarragon

South locality, L. johnsi co-occurs with L. dignomontis and

L. gippslandicus .

Etymology. Named for Peter Johns, formerly of the University

of Canterbury (Christchurch, New Zealand), who recognised

the distinctiveness of the 1890 specimen and illustrated its

gonopod, assigning the species to Pseudoprionopeltis

( Australopeltis ) without naming it (Johns, 1964).

Lissodesmus latus sp. nov.

Figures 6A, 9, 46, 691at, 701at, 7 llat, 75 (map)

Lissodesmus sp. NW1. — Mesibov, 1993: 31. — Mesibov, 1996:

18.— Mesibov, 1998: 155.— Bonham et al., 2002: 240.

Material examined. Holotype. Male, Australia, Tasmania. Wombat

Hill, W of Waratah, CQ703065 (41°28’56"S 145°26'47"E), 670 m,

24.ix.1990, R. Mesibov, QVM 23:17495.

Paratypes. 2 males, Wombat Hill, W of Waratah, CQ702064

(41°29'00"S 145°26'42"E), 680 m, 19.ix.1990, R. Mesibov, AM

KS91178 (ex QVM 23: 17490); 2 males, same details, QVM 23:17490,

1 dissected; 2 males, same details but 22.ix.1990, QVM 23:17491;

2 males, details as for holotype but 23 .ix. 1990, QVM 23:17492;

2 males, Wombat Hill, W of Waratah, CQ704066 (41°28'53"S

145°26'51"E), 690 m, 28.ix.1990, R. Mesibov, QVM 23:17496, 1 dis-

sected; male and female in copula, same details, QVM 23:17497;

7 females, same details, QVM 23:17547; 2 males, same details but

29.ix.1990, QVM 23:17493; 2 males, same details but 30.ix.1990,

QVM 23:17494; 9 males, Montagu Swamp, CQ265620 (40°58’28"S

144°56'16"E), 30 m, 22.vi.1991, R. Mesibov, QVM 23:17499, 1 dis-

sected; 5 males, 2 females, Deep Creek Bay, CQ447787 (40°49'40"S

145°09'29"E), <10 m, 5.ix.2000, K. Bonham, QVM 23:41955, 2 males

dissected.

Other material. 186 males, 116 females and 20 juveniles from 105

localities (see il Lissodesmus supplement” for details).

Description. Male c. 20 mm long, H = 1.8 mm. In alcohol,

well-coloured specimens of typical form (see Remarks ) under

low magnification with near- white body colour and wide, trans-

verse, purple banding, interrupted medially, on posterior

margin of prozonites. Antenna long (Fig. 691at). Paranota

fairly wide, R = 1.5, sometimes with two prominent posterior

marginal teeth (Fig. 701at), posterior corners not turned up (Fig.

9). Limbus elements simple, tapering to point (Fig. 6A). Legs

robust, tarsus longer than femur, tibia with slight ventral distal

swelling (Fig. 7 llat). Telopodite (Fig. 46) reaching leg 5 when

retracted. Solenomere arising between one-quarter and one-

third the telopodite height, directed posterodistally at c. 30° to

telopodite axis, curving laterally, terminating with toothed sub-

apical collar at about one-third the prefemoral process height.

Tibiotarsus wider than solenomere and slightly longer, curving

more sharply laterally, diverging from solenomere at a small

angle, terminating in a thin, dentate edge sometimes turned

anterodistally at level of solenomere tip. Femoral process

arising slightly distal to solenomere origin, forked at about one-

third its length, both branches blade-like, bluntly pointed;

anterior branch closely pressed to prefemoral process at its

base, terminating at level of solenomere tip; posterior branch

typically as long as or slightly longer than anterior branch,

curved anterodistally. Prefemoral process at its origin about

two-thirds as wide as telopodite base, very slightly bent later-

ally, bending posteriorly at just over half its length, the mesal

edge and expanded, mesolaterally flattened tip both crenulate

or shortly toothed. Two small unci arising at just under half the

prefemoral process height (just distal to solenomere tip), one

central and one on mesal edge of process.

Distribution and habitat. In cool temperate rainforest, wet

eucalypt forest and coastal scrub and blackwood/tea-tree forest

over c. 13 000 km 2 in north-west Tasmania (Fig. 75), from the

Hunter Group of islands to the Franklin River, from the west

coast inland to the Leven River and the Cradle Mountain area,

and from sea level to at least 950 m. L. latus has been found in

caves at Gunns Plains, Loongana, Mt Cripps and the Wilson

River. It co-occurs with L. perporosus over most of its range

118

Robert Mesibov

and is typically less abundant than L. perporosus when the two

are syntopic.

Etymology. Latin latus (“broad, wide”), for the wide and

relatively flat paranota.

Remarks. Besides the inland form of L. latus , represented by

the holotype, there is a distinctive coastal form with almost uni-

formly pale brown to chestnut-brown coloration and with

stronger dorsal curvature of the paranota. Despite the striking

difference in overall appearance of the two L. latus forms, the

gonopods of coastal and inland forms in the far north-west are

almost indistinguishable. The coastal form occurs close to the

western and Bass Strait coasts and in the Hunter Group of

islands. 1 have not yet found a site in the far north-west

of Tasmania where the coastal and inland form s co-occur.

Coastal L. latus closely resembles L. perporosus, but females

and juveniles of the two species can be separated by inspection

of the ozopores (i.e., the pore formula in L. latus is normal).

Inland L. latus vary in the height of the paranotal margins

relative to the body axis, in the sharpness of notches on the

lateral edge of the paranota, and in the clear presence or near-

absence of a pair of rounded teeth on the posterior margin of

each paranotum. On the gonopod telopodite, the anterior

branch of the femoral process varies from nearly straight to

arc-like (concave posteriorly), and the tip of the tibiotarsus can

be straight or upturned. Furry specimens of L. latus have been

found in the Mt Cripps area and at Lake Lea and the nearby

Vale of Belvoir (Fig. 9).

Lissodesmus macedonensis sp. nov.

Figures 47, 48, 69mac, 70mac, 71mac, 79 (map)

Material examined. Holotype. Male, Australia, Victoria. Mt Macedon,

37°22'42"S 144°36'35"E, 880 m, 10.ix.2004, R. Mesibov & T. Moule,

NMV K-9527.

Paratypes. 2 males, Mt Macedon, 22.x. 1963, A. Neboiss, NMV

K-9508, K-9509; 2 males, details as for holotype, AM KS91179; 15

males, 1 female, details as for holotype, includes male and female in

copula, NMV K-9511 to K-9526, 2 males dissected; 9 females, details

as for holotype, NMV K-9528 to K-9536; male, 1 km NE of

Blakeville, 37°29'45"S 144°13'07"E, 680 m, 10.ix.2004, R. Mesibov &

T. Moule, NMV K-9510.

Description. Male c. 17 mm long, H = 1.5 mm. In alcohol,

well-coloured specimens under low magnification with very

pale brown body colour and red speckling on metatergites,

more intense medially and in a transverse band close to poste-

rior metatergal margin. Antenna short, stout (Fig. 69mac).

Paranota fairly wide, R = 1.5, with several indistinct posterior

marginal teeth (Fig. 70mac), posterior corners not turned up.

Legs short, robust, tarsus about as long as femur, tibia with

slight ventral distal swelling (Fig. 71mac). Telopodite (Figs 47,

48) reaching leg 5 when retracted. Solenomere arising at one-

third to half the telopodite height, directed posterodistally at

c. 45° to telopodite axis, terminating with small, pointed sub-

apical projection at one-quarter to one-third the prefemoral

process height. Tibiotarsus strongly flattened anteroposteriorly,

about as long as solenomere but diverging from it slightly, the

expanded tip curling back anterodistally. Femoral process aris-

ing distal to solenomere origin at about one-quarter the

prefemoral process height, blade-like, wide, pressed close

to prefemoral process at base, deeply notched distally with

rounded tips, terminating at just under half the height of

prefemoral process. Prefemoral process at its origin about

half as wide as telopodite base, slightly tapered, curving

posteromesally at about two-thirds its length, the lateral

edge of the distal half a comb of c. 25 long, mainly basally

directed teeth, the process tip turned back nearly 180°.

Uncus large, arising from mesal edge of prefemoral process at

about one-third the process height (just distal to solenomere

tip).

Distribution and habitat. Known from wet eucalypt forest at

two localities c. 30 km apart: the upper slopes of Mt Macedon,

where it is abundant, and near Blakeville in the southern

portion of the Wombat State Forest (Fig. 79).

Etymology. Named for the type locality, 55 km north-west of

Melbourne.

Lissodesmus martini (Carl, 1902)

Figures 49, 50, 69mar, 70mar, 71 mar, 79 (map)

Pseudoprionopeltis martini Carl, 1902: 599. — Brolemann, 1916:

541.— Chamberlin, 1920: 133. — Verhoeff, 1932: 1982.— Attems,

1940: 453.

Pseudoprionopeltis (Australopeltis) martini Johns, 1964: 47.

Lissodesmus martini Jeekel, 1983: 150. — Jeekel, 1984: 86.

Material examined. Holotype (not seen). Male, Australia, Victoria.

Melbourne, date of collection unknown, collected by “Consul Martin”,

deposited in the Museum d’Histoire Naturelle in Geneva. According to

Johns (1964: 47), who re-illustrated the gonopod of the type, the holo-

type body has been lost and only a slide mount of the gonopods

remains.

Paratypes. None designated.

Other material. 50 males, 19 females and 8 juveniles from Acheron

Gap, “The Beeches” near Marysville, Belgrave West, the Bennison

Plains area NE of Licola, Coalville, Cockatoo Creek, Emerald, Erica,

Gembrook, the Latrobe River near Powelltown, Mt Observation,

Narracan, Narracan East, Neerim, Noojee, Sherbrooke Forest, Slippery

Hill N of Dargo, Starvation Creek NE of Warburton, Toolangi,

Yalloum North and Yinnar (see “ Lissodesmus supplement” for details).

I have not examined the male from Ferntree Gully National Park

described by Jeekel (1983).

Description. Male c. 18 mm long, H = 1.6 mm. In alcohol,

well-coloured specimens under low magnification with pale

brown body colour, red speckling anteriorly on metatergites

and a red transverse band posteriorly on prozonites, paranota

near-white. Antenna moderately long (Fig. 69mar). Paranota

fairly wide, R = 1.5, with about 3 posterior marginal teeth (Fig.

70mar), posterior comers not turned up. Legs robust, tarsus

longer than femur, tibia with prominent ventral distal swelling

(Fig. 71mar). Telopodite (Figs 49, 50) almost reaching leg 5

when retracted. Solenomere arising at about half the telopodite

height, directed posterodistally at a small angle (c. 30°) to

telopodite axis, terminating with toothed subapical collar at just

under half the prefemoral process height. Tibiotarsus origin on

posteromesal surface of telopodite, not close to solenomere

origin; tibiotarsus a thin, pointed rod almost parallel to

solenomere but curving anterodistally, about half the length of

New genus of millipedes

119

the solenomere. Femoral process arising at level of

solenomere origin, a massive, wide, bluntly pointed blade

directed distally, close to prefemoral process, terminating at

just over half the prefemoral process height (distal to

solenomere tip). Prefemoral process at its base about half as

wide as telopodite base, curving slightly mesally at half

its height, bent posteromesally at about three-quarters its

length, the tip expanded and dentate on its posterior-facing

edge. Uncus prominent, arising near mesal edge of prefemoral

process at just under half the process height (at same level as

solenomere tip), with inconspicuously dentate outer edge.

Distribution and habitat. In dry and wet eucalypt forest and

cool temperate rainforest in the Central Highlands and parts of

Gippsland. To judge from earlier records, L. martini once

ranged east and north-east from Melbourne for ca. 200 km, and

it is still Victoria’s most widely distributed Lissodesmus species

(Fig. 79). It is abundant in the Yarra Ranges and occurs up to at

least 1120 m elevation.

Remarks. Carl (1902) characterised Pseudoprionopeltis as

having 21 segments behind the head, rather than the expected

19 or 20. The correct number, 20, was first given by Johns

(1964) for L. martini. Enghoff et al. (1993: 148) suggested that

Carl’s “21” may have been a typographical error.

Chamberlin (1920) gave Polydesmus ( Oxyurus ) serratus

Hutton, 1877 as a synonym of P. martini , and listed the type

locality of serratus, Dunedin in New Zealand, as the only

P. martini locality. This was clearly an error, as Chamberlin

(1920: 133) also claimed that both the identity and generic

placement of P. serratus were impossible to determine.

This is the fourth time L. martini has been described and

illustrated, and I am contributing little new other than a scan-

ning electron micrograph of the gonopods. L. martini varies

only slightly over most of its range, but specimens from the east

of that range are smaller and paler than those from the west.

Lissodesmus milledgei sp. nov.

Figures 51, 69mil, 70mil, 71 mil, 78 (map)

Material examined. Holotype. Male, Australia, Victoria. Myrtle Gully

Reserve, 3.4 km WSW of Mt Donna Buang, 37°43’S 145°38'30"E,

pitfall emptied 20.L1995, G. Milledge, ex sample NOH-1840, NMV

K-9604.

Paratypes. Male, details as for holotype but 7.iv.l995, ex sample

NOH- 1841, NMV K-9605 ; male, Acheron Gap, 6 km NE of Mt Donna

Buang, 37°40'43"S 145°44'20"E, pitfall emptied 28.xii.1995,

G. Milledge, sample NOH- 1851, NMV K-9606, dissected.

Other material. Male, The Beeches, 37°28’S, 145°49’E, 25.iii.1991,

M.S. Harvey & M.E. Blosfeld, Nothofagus litter, WAM T66223.

Description. Male c. 14 mm long, H = 1.3 mm. In alcohol,

under low magnification with uniformly very pale brown body

colour. Antenna short (Fig. 69mil). Paranota reduced, R = 1.4,

anterior shoulders squared rather than rounded (Fig. 70mil),

posterior corners slightly turned up. Legs moderately robust,

tarsus shorter than femur, tibia with slight ventral distal

swelling, sphaerotrichomes on tibia and tarsus only of leg 6

(Fig. 71mil). Telopodite (Fig. 51) reaching leg 5 when

retracted. Solenomere arising at half the telopodite height,

directed posterobasally at large angle to telopodite axis but

bending at about one-third the process length distally and

slightly mesally, terminating with small subapical projection at

between one-t hi rd and half the prefemoral process height.

Tibiotarsus origin on posteromesal surface of telopodite, not

close to solenomere origin; tibiotarsus a posterobasally flat-

tened triangular plate with a bluntly pointed tip, directed at

c. 45° to telopodite axis, about one-third the solenomere length.

Femoral process arising at level of solenomere origin, directed

distally and pressed close to prefemoral process, blade-like

with a slight anterior shoulder distally, tip rounded, terminating

at one-third to half the prefemoral process height (at level of

solenomere tip). Prefemoral process somewhat narrower

than telopodite base, tapering, the distal one-third curving

posterobasally and slightly mesally and bearing a comb of

c. 15-20 long, mainly basally directed teeth; a cluster of

three small tooth-like projections on mesal edge of

prefemoral process at about two-thirds the process height.

Uncus prominent, arising at level of solenomere and femoral

process tips towards lateral edge of prefemoral process.

Distribution and habitat. Known from three localities in cool

temperate rainforest in the Yarra Ranges east of Melbourne

(Fig. 78).

Etymology. Named for Graham Milledge, who collected this

and many other interesting invertebrate species in the mid-

1990s from Nothofagus forest in Victoria.

Lissodesmus modestus Chamberlin, 1920

Figures 4A, 5A, 7A, 8A, 52, 53, 69mod, 70mod, 71mod, 73

(map)

Lissodesmus modestus Chamberlin, 1920: 135. — Attems, 1940:

490.— Jeekel, 1970: 336.— Jeekel, 1984: 91.

Material examined. Holotype. Male, Australia, Tasmania. Russell

Falls, G.H. Hardy, MCZ 4644. The type is in three pieces but is other-

wise in excellent condition. An accompanying label reads “Jan 1915

Russell Falls”.

Paratypes. Female, details as for holotype, MCZ 4645.

Other material. 57 males, 82 females and 32 juveniles from Arve

Road, Bracken Ridge, Collins Cap, Coopers Creek, Dromedary Creek,

Edwards Road, Espies Craig, Fortescue Bay, Gold Creek, Hastings

Caves, Ida Bay cave IB-51, Judds Creek, Kallista Creek, Little

Florentine River, Mt Clark, Mt Field National Park, Mt Mangana

(Bruny Island), Mt Misery, Mt Wellington, Myrtle Forest Creek,

Needles Picnic Area, Pelham Tier, Pendulum Palace Cave (Precipitous

Bluff), Picton River, Plenty River and Tobys Hill (see “ Lissodesmus

supplement” for details).

Description. Male c. 16 mm long, 77 = 1.4 mm. In alcohol,

well-coloured specimens under low magnification very light

brown in body colour, with fairly uniform red speckling across

metatergites and in a transverse band posteriorly on prozonites.

Antenna long, fairly slender (Fig. 69mod). Paranota fairly

wide, R = 1.5 (Fig. 70mod), posterior corners not turned up

(Fig. 4A) but projecting slightly outwards in most specimens

(see also Jeekel, 1984: 93, Fig. 1). Legs robust, tarsus slightly

shorter than femur, tibia with prominent ventral distal swelling

(Fig. 71mod). Telopodite (Figs 52, 53) reaching leg 4 when

retracted. [Note: the convention adopted here of labelling as

120

Robert Mesibov

“mesal” a telopodite view centred on the solenomere is partic-

ularly misleading for this species. As seen in Fig. 52, the

solenomere origin is on the anterior surface of the telopodite in

situ.] Solenomere arising at about half the telopodite height,

directed basally at about 45° to telopodite axis but gradually

curving distally, terminating with a small subapical projection

at just under half the prefemoral process height. Tibiotarsus

rod-like, pointed, directed more or less parallel to solenomere,

about one-third the length of solenomere and much thinner.

Femoral process arising distal to solenomere origin, large,

blade-like with expanded, leaf-shaped tip, curving pos-

teromesally and terminating just proximal to tip of

prefemoral process. Prefemoral process at origin about

two-thirds as wide as telopodite base, at about half-length

tapering rapidly and bending posteromesally, then bending

distally, the last one-third straight, with a row of short,

mainly posterodistally and mesodistally directed teeth on

mesal edge. Uncus identification uncertain; possibly repre-

sented by low, pointed ridge on prefemoral process at about the

level of solenomere tip.

Distribution and habitat. In wet eucalypt forest and cool tem-

perate rainforest over c. 4500 km 2 in southern and south-east

Tasmania (Fig. 73), from Pelham to the south coast, from the

Little Florentine River east to the Channel district, and from

near sea level to at least 940 m. L. modestus also occurs on

South Bruny Island and the Tasman Peninsula, but has not yet

been recorded on either North Bruny Island or Forestier

Peninsula. It occurs in caves at Ida Bay and Precipitous Bluff.

L. modestus is hard to find in many parts of its range.

Remarks. As noted in the Introduction, there are minor errors

with the antennomeres and sphaerotrichomes in the description

given by Jeekel (1984). The gonopod coxae are also said

(Jeekel, 1984: 93) to be “solidly connected”, which suggests

“fused”; they are in fact separate and lightly joined at only one

point, as in all Lissodesmus species. Jeekel’s description is

otherwise accurate and very detailed.

Lissodesmus montanus sp. nov.

Figures 54, 55, 69mon, 70mon, 71mon, 73 (map)

Material examined. Holotype. Male, Australia, Tasmania. Lake Lea,

DP083989 (41°33'22"S 145°54'01'E), 950 m, 18.xii.1993, R. Mesibov

& T. Moule, QVM 23:17717.

Paratypes. Male, Algonkian Mountain, approx. DP220059

(42°23'42"S 146°03'08"E), 1020 m, 26.ii.1987, S.J. Smith, ex WHA

sample ML011, QVM 23:17715, dissected; female, same details,

QVM 23:17714; male, Algonkian Mountain, approx. DP229067

(42°23'16"S 146°03'48'E), 1020 m, 26.ii.1987, S.J. Smith, ex WHA

sample ML005, QVM 23:17716, dissected; male, same details, AM

KS91180; female, Squires Creek, DP240255 (42°13'07"S

146°04'44'E), 650 m, 16.ii.1994, R. Mesibov, QVM 23:40813; male,

Mt Murchison summit, CP850703 (41°48'38"S 145°36'55'E), 1250 m,

ll.iv.1998, R. van Riet & B. Dudman, QVM 23:40744.

Description. Male c. 23 mm long, H = 2.4 mm. In alcohol,

well-coloured specimens under low magnification an almost

uniform light red-brown in body colour (unusually, extending

to legs and antennae) with small pale brown patches medially

and anteriorly on metatergites, and laterally on paranota.

Antenna fairly short, fairly slender (Fig. 69mon). Paranota

fairly wide, R = 1.5 (Fig. 70mon), posterior corners not turned

up. Legs robust, tarsus longer than femur (Fig. 71mon).

Telopodite (Figs 54, 55) narrowing sharply at prefemoral

process origin, reaching leg 5 when retracted. Solenomere aris-

ing at one-third to half the telopodite height, directed basally at

c. 45° to telopodite axis, curving gradually distally, terminating

with a toothed subapical collar at about one-third the

prefemoral process height. Tibiotarsus as large as solenomere

but diverging from it and curving laterally, greatly expand-

ing apically and deeply and broadly notched, thus ter-

minating in two large spine-like structures. Femoral process

arising just distal to solenomere origin, blade-like with a slight

posterior shoulder apically, pressed close to prefemoral

process, terminating at about half the prefemoral process height

(distal to solenomere tip). Prefemoral process at origin about

half as wide as telopodite base, bending first mesally then

distally, the tip curving posteriorly with a comb of c. 20

long, mainly posterobasally teeth on mesal edge, the comb

continued proximally as a row of more irregular “saw-

teeth” to about the level of solenomere tip. Uncus small,

arising centrally just proximal to solenomere tip.

Distribution and habitat. Known from four localities over a

range of c. 500 km 2 in central western Tasmania (Fig. 73), from

650 to 1250 m. At Algonkian Mountain, Lake Lea and Squires

Creek the habitat was cool temperate rainforest, while on Mt

Murchison L. montanus was found walking by day among bare

rocks (R. van Riet, pers. comm.).

Etymology. Latin montanus (“of mountains”), adjective.

Lissodesmus orarius sp. nov.

Figures 8B, 56, 57, 69ora, 70ora, 71ora, 76 (map)

Material examined. Holotype. Male, Australia, Tasmania. Foam Creek,

CP255877 (41°38'35"S 144°54T6"E), <10 m, l.vi.1993, R. Mesibov,

QVM 23:45822 (ex QVM 23:17693).

Paratypes. 10 males, Pedder River, CQ150149 (41°23’45”S

144°47T3"E), <10 m, 23.U991, R. Mesibov, QVM 23:17683, 2 dis-

sected; 2 males, details as for holotype, AM KS91181 (ex QVM

23:17693); 9 males, details as for holotype, QVM 23:1769, 2 dis-

sected 3; 18 females, 2 stadium VI juvenile females, details as for

holotype, QVM 23: 17704.

Other material. 28 males, 53 females and 20 juveniles from

Chimney Creek, Foam Creek, Hunters Creek, Interview River, Lanes

Tor, Monster Creek, Pedder River, Pieman Head, Rocky Creek, Rupert

Point, Sandy Cape and Sea Devil Rivulet (see “ Lissodesmus

supplement” for details).

Description. Male c. 17 mm long, H = 1.5 mm. In alcohol,

well-coloured specimens under low magnification with almost

uniform light brown body colour, somewhat darker posteriorly

on prozonites and lighter on paranotal edges. Antenna short,

fairly stout (Fig. 69ora). Paranota reduced, R = 1.4 (Fig. 70ora),

posterior comers not turned up. Legs moderately robust, tarsus

much longer than femur, tibia with slight ventral distal swelling

(Fig. 71 ora), sphaerotrichomes lacking on leg 6 prefemur (Fig.

8B). Telopodite (Figs 56, 57) reaching just past leg 6 when

retracted. Solenomere arising at about one-third the telopodite

height, directed posterodistally at c. 45° to telopodite axis,

New genus of millipedes

121

sharply curving distally and laterally at midpoint, terminating

with small subapical projection at about one-third the

prefemoral process height. Tibiotarsus about as large as

solenomere and paralleling its course, the tip turned distally

and bluntly pointed, terminating just proximal to solenomere

tip. Femoral process arising distal to solenomere origin, blade-

like and slightly expanded in middle, directed distally and

pressed close to prefemoral process, bluntly pointed and ter-

minating at about two-thirds the prefemoral process height

(distal to solenomere tip). Prefemoral process as wide as

telopodite base, slightly curved laterally, the tip wide,

greatly flattened and concave posteriorly, few small teeth on

posterior edge of lateral rim, the mesal rim bearing (a) at

the process tip a long, more or less cylindrical branch

curving posterolaterally, (b) a smaller, finger-like branch

directed posterodistally more proximally, and (c) a sharp

triangular point at about the level of the femoral process

tip. Uncus small, arising near mesal edge of prefemoral process

at level of solenomere tip.

Distribution and habitat. Known from a c. 30 km strip along

the west coast of Tasmania (Fig. 76), from the Pedder River

(just north of Sandy Cape) to the mouth of the Pieman River.

Along this strip, L. orarius has been found just above sea level

in dune scrubs, tea-tree forest and tea-tree scrub. Remarkably,

the L. orarius distribution extends down to the high tide line,

and at the type locality this species was found together with

intertidal crabs under driftwood surrounded by sedge and grass.

At most of its known localities L. orarius is abundant.

Etymology. Latin orarius (“of the coast”), adjective.

Remarks. The number and size of teeth on the lateral branch of

the prefemoral process vary a little across the L. orarius range.

Lissodesmus otwayensis sp. nov.

Figures 58, 59, 69otw, 70otw, 71otw, 79 (map)

Material examined. Holotype. Male, Australia, Victoria. Calder Ridge,

38°42'41"S 143°34'03"E, 380 m, 13.xii.2003, R. Mesibov & T. Moule,

NMV K-9614.

Paratypes. Male, 2 miles W of Apollo Bay, 8.iii. 1975, M. Campbell,

NMV K-9607; 1 stadium 7 male, 1 female, 2 stadium 7 females, Aire

Crossing Track, 0.5 km N of Aire R crossing, 38°40'S 143°29'E,

31.L1995, G. Milledge, direct search, sample NOH-1086, NMV K-

9608 to K-9611; 2 males, Turtons Pass, 38°38’43"S 143°40'36"E, 420

m, 12.xii.2003, R. Mesibov & T. Moule, NMV K-9612, K-9613, 1 dis-

sected; 2 males, same details, AM KS91182; 2 males, details as for

holotype, NMV K-9618, K-9619, 1 dissected; 3 females, details as for

holotype, NMV K-9615 to K-9617; 7 males, Cape Horn, 38 0 44'13"S

143°34'30"E, 280 m, 13.xii.2003, R. Mesibov & T. Moule, NMV K-

9620 to K-9626, 2 dissected; 2 males, 1 female, Diamond Hill,

38°27'05"S 143°56T0"E, 360 m, 14.xii.2003, R. Mesibov & T. Moule,

NMV K-9627 to K-9629, 1 male dissected; male, same details but

38°27'00"S 143°56'14"E, NMVK-9630.

Description. Male c. 17 mm long, H = 1.6 mm. In alcohol,

well-coloured specimens under low magnification pale brown

in body colour with red speckling dorsally on prozonites and

metatergites, and a darker, transverse red band posteriorly on

metatergites. Antennae short, clavate (Fig. 69otw), about 1.75x

a socket diameter apart. Paranota reduced, R = 1.4 (Fig. 70otw),

posterior comers not turned up. Legs robust, tarsus longer than

femur, tibia with prominent ventral distal swelling (Fig.

71otw). Telopodite (Figs 58, 59) reaching leg 5 when retracted,

with posterolateral row of sparse long setae mnning well distal

to process origins, the base posteriorly somewhat excavate just

below process origins. Solenomere arising at one-third the

telopodite height, directed posterodistally at c. 45° to telopodite

axis, curving very slightly laterally, terminating with small sub-

apical projection at just under one-third the prefemoral process

height. Tibiotarsus rod-like, bluntly pointed, more or less par-

allel to solenomere but somewhat narrower and about half its

length. Femoral process arising distal to solenomere origin,

blade-like, forked at about half its length, both branches

wide and apically pointed; anterior branch curving anteri-

orly and terminating at about one-third the prefemoral

process height (just distal to solenomere tip); posterior

branch directed posterodistally and curving slightly mesal-

ly. Prefemoral process curving slightly laterally, narrowing

and curving posteriorly and distally at just over half the

process height, with a sharply pointed “shoulder” on the

mesal edge marking the bend, distally bending sharply

mesally and tapering to blunt point, almost the whole of the

lateral edge distal to the bend a comb of c. 30-40 long,

straight teeth. Uncus arising at just under half the prefemoral

process height (distal to solenomere tip) on mesal edge of

process.

Distribution and habitat. In wet eucalypt forest and cool tem-

perate rainforest in the Otway Ranges (Fig. 79).

Etymology. Named for the Otway Ranges, where this species

appears to be abundant.

Lissodesmus peninsulensis sp. nov.

Figures 60, 61, 69pen, 70pen, 71pen, 72 (map)

Lissodesmus sp. PI. — Mesibov, 1996: 18.

Material examined . Holotype. Male, Australia, Tasmania. Fortescue

Bay, EN762228 (43°08'36"S 147°56T3"E), 120 m, ll.iii.1992, R.

Mesibov, QVM 23:45825 (ex QVM 23:16173).

Paratypes. Male, Griffiths Road, Koonya State Forest, approx.

EN650270 (43°06'S 147°47'E), 19.xi.1977, J.L. Hickman, QVM

23:16170, dissected; male, details as for holotype, AM KS91183 (ex

QVM 23:16173); male, details as for holotype, QVM 23:16173, dis-

sected; male, Browns Creek, EN736463 (42°55’56"S 147°54'07"E),

190 m, 20.iii.1992, R. Mesibov, QVM 23:16172, dissected; stadium 7

male, stadium 6 male, 5 stadium 7 females, 2 stadium 6 females, same

details, QVM 23:16175; male, Wellard Rivulet, EN730456

(42°56T8"S 147°53'40"E), 140 m, 24.ii.1993, R. Mesibov, QVM

23:16171, dissected; stadium 6 male, female, stadium 7 female, stadi-

um 6 female, same details, QVM 23:16174; male, Mt Clark, approx.

EN638267 (43°06'34"S 147°47'02"E), 250 m, 4.V.2002, K. Bonham,

QVM 23:25021.

Description. Male c. 16 mm long, H = 1.6 mm. In alcohol,

well-coloured specimens under low magnification pale to light

brown in body colour, somewhat darker posteriorly on pro-

zonites . Antennae long (Fig. 69pen), about 2.5x a socket diam-

eter apart. Paranota wide, R = 1.6 (Fig. 70pen), anterior “shoul-

ders” projecting forward, posterior comers not turned up. Legs

long, moderately robust, tarsus slightly longer than femur, tibia

122

Robert Mesibov

with slight ventral distal swelling (Fig. 71pen). Telopodite

(Figs 60, 61) almost reaching leg 5 when retracted.

Solenomere arising at just under half the telopodite height,

directed posterodistally at c. 45° to telopodite axis, twisted

slightly helically, the tip bending distally, terminating with

no subapical projection at half the prefemoral process

height. Tibiotarsus unusually wide at base, initially parallel to

solenomere but curving posterolaterally, then mesally, then dis-

tally; somewhat shorter than solenomere and terminating prox-

imal to solenomere tip. Femoral process arising just proxi-

mal to solenomere origin, blade-like, forked at about one-

quarter its length and not pressed close to prefemoral

process, the two branches apically pointed and about equal

in size; anterior branch curved anteriorly, then distally,

approaching prefemoral process apically; posterior branch

arcuate, concave anteriorly, bent more strongly anteriorly

near tip; the two branches terminating at the level of

solenomere tip. Prefemoral process at origin about two-thirds

as wide as telopodite base, sharply narrowing at about one-third

its length, the distal two-thirds gradually curving posteriorly,

the tip mesolaterally flattened and not armed with teeth. Uncus

large, arising laterally at level of solenomere tip, flattened pos-

terobasally, the outer edge slightly crenulate.

Distribution and habitat. Known from wet eucalypt forest to

250 m elevation on Forestier and Tasman Peninsulas in south-

east Tasmania (Fig. 72). L. peninsulensis appears to be uncom-

mon and may have a total range of less than 200 km 2 .

Etymology. Latin peninsula + -ensis, adjective, referring to the

Forestier and Tasman Peninsulas.

Lissodesmus perporosus Jeekel, 1984

Figures 1, 10, 62, 63, 69per, 70per, 71 per, 72 (map)

Lissodesmus perporosus Jeekel, 1984: 98.

Material examined. Holotype (not seen). Male, Australia, Tasmania.

Hellyer Gorge, 32 km SSW of Somerset, 25.xi.1980, C.A.W. Jeekel

and A. Jeekel-Rijvers. The type is said to be deposited in TM (Jeekel,

1984: 86), but has not yet been received there.

Paratypes. 6 males, 13 females, 3 stadium 7 males, 2 stadium

7 females, details as for holotype; male, 12 km SW of Derwent Bridge,

26.xi.1980, C.A.W. Jeekel and A. Jeekel-Rijvers; 2 males, 4 females,

Lake St Clair National Park, near Cynthia Bay, 5 km WNW of Derwent

Bridge, 26.xi.1980, C.A.W. Jeekel and A. Jeekel-Rijvers. These

specimens are listed as paratypes by Jeekel (1984: 98) but their

present locations are unknown and they have not been examined.

Other material. 550 males, 616 females and 328 juveniles from 333

localities (see “ Lissodesmus supplement” for details).

Description. Male c. 18 mm long, H = 1.7 mm. In alcohol,

under low magnification with near-white body colour and three

transverse bands dorsally: reddish purple posteriorly on

prozonites, red speckling anteriorly and light brown (mainly

medially) posteriorly on metatergites. Antenna fairly short,

stout (Fig. 69per). Paranota fairly wide, R = 1.5 (Figs 10D,

70per), posterior corners not turned up (Fig. 1). Pore formula

5, 7, 9-19 in males and females. Legs robust, tarsus about as

long as femur, tibia with slight ventral distal swelling (Fig.

71per). Telopodite (Figs 62, 63) reaching leg 5 when retracted.

Solenomere arising at about one-third the telopodite height,

directed posterodistally at a small angle (c. 30°) to telopodite

axis, curving slightly laterally, terminating with toothed sub-

apical collar at about one-third the prefemoral process height.

Tibiotarsus somewhat larger than solenomere, directed at

larger angle to telopodite axis, straight with tip bent anterodis-

tally at a right angle. Femoral process arising just proximal to

solenomere origin, blade-like, forked at about one-third its

length; anterior branch directed distally, pointed, pressed close

to prefemoral process, terminating at just over half the

prefemoral process height (distal to solenomere tip); posterior

branch about one-quarter as long as anterior branch and much

narrower, directed posteriorly and curving slightly distally.

Prefemoral process at origin about half the width of telopodite

base, curving gradually posteriorly from about half the process

height, the mesal edge distally with a few low, tooth-like pro-

jections. Small paired unci arising at about half the prefem-

oral process height (distal to solenomere tip) from middle of

posterior surface and from mesal edge.

Distribution and habitat. In cool temperate rainforest, wet

eucalypt forest, blackwood/tea-tree swamp forest and subalpine

woodland over at least 20 000 km 2 in north-west Tasmania

(Fig. 72), from sea level to at least 1150 m. The L. perporosus

range extends from the west coast near Pieman Head to the

eastern edge of the Central Plateau and to Notley Gorge in

the West Tamar region, and from the Bass Strait coast near the

Black River mouth to Tarraleah in the upper Derwent Valley.

L. perporosus is the most abundant dalodesmid over most of its

range. It occurs in Pinus radiata plantations (Bonham et al.,

2002; Mesibov, 2005) as well as native vegetation and has been

found in caves at Bubs Hill and Mole Creek.

Remarks. Size and coloration vary little in this species, and

females and juveniles are easily recognised by the unusual pore

formula. A single male L. perporosus from Trackham Creek

near Waratah (QVM 23:17460) has a normal pore formula.

Another 114 specimens from the same locality, including

28 males, have the perporosus formula, indicating that this

Trackham Creek male is unique and not representative of a

pore-formula “race” within the species. Gonopod structure also

varies very little in L. perporosus, the gonopods of males from

sites 200 km apart at Rebecca Creek and Dee Lagoon differing

only slightly in size, number and position of teeth on the

prefemoral process. The greatest deviation from the typical

perporosus gonopod structure is found in the vicinity of Maggs

Mountain in the upper Mersey River catchment. Males in this

small area have additional tooth-like projections on the

prefemoral process, and the tip of the process is thickened and

bent over (Fig. 63).

Lissodesmus plomleyi sp. nov.

Figures 64, 69plo, 70plo, 71plo, 76 (map)

Lissodesmus sp. NE4. — Mesibov, 1996: 16. — Mesibov, 1998: 155.

Material examined. Holotype. Male, Australia, Tasmania. East of

Rattler Hill, EQ743353 (41°13'49"S 147°53T2"E), 650 m, 1812005,

R. Mesibov, QVM 23:45893.

Paratypes. Male, Rattler Hill, EQ744353 (41°13'48”S 147°53T5"E),

650 m, 6.ix.l990, QVM 23:15466, dissected; female, Rattler Hill,

EQ744353 (41°13'48"S 147°53’15”E), 650 m, 7.ix.l990, QVM

New genus of millipedes

123

23:15468; female, Rattler Hill, EQ744353 (41°13'48"S 147°53'15"E),

650 m, 8.ix.l990, QVM 23:15467; male, Ben Nevis, EQ538147

(41°25'02"S 147°38'37"E), 890m, 15.iii.1993, QVM 23:15465, dis-

sected; male, Caldbeck Creek, EQ471205 (41 0 21'55"S 147°33’47"E),

850 m, 26.xii.1994, R. Mesibov & T. Moule, AM KS91184 (formerly

QVM 23:17758).

Description. Male c. 11 mm long, H = 1.2 mm . In alcohol,

under low magnification very pale brown in body colour with

darker brown speckling in narrow transverse band posteriorly

on metatergites . Antennae short, clavate (Fig. 69plo), about

1.75x a socket diameter apart. Paranota reduced, R = 1.4 (Fig.

70plo), posterior corners not turned up. Legs robust, tarsus

longer than femur, tibia with prominent ventral distal swelling

(Fig. 71plo). Epiproct with prominent paired, bluntly rounded

projections. Telopodite (Fig. 64) widest near solenomere origin,

reaching just past leg 6 when retracted. Solenomere arising at

one-third the telopodite height, directed posterodistally at a

small angle to telopodite axis, curving gradually laterally, ter-

minating with small sub apical projection at about one-quarter

the prefemoral process height. Tibiotarsus roughly parallel to

solenomere and about as wide but shorter, rod-like and pointed.

Femoral process arising at level of solenomere origin, blade-

like, forked at just over half its length; anterior branch basally

pressed close to prefemoral process, directed distally, pointed

and terminating at one-t hi rd to half the prefemoral process

height (distal to solenomere tip); posterior branch short, wide,

pointed, arising from anterior branch at right angle.

Prefemoral process at origin nearly as wide as telopodite

base, slightly sinuous in anterior view, curving gradually

posteriorly from about half its height, the distal third with

a few very long, straight teeth on mesal edge directed

basally and posterobasally and on lateral edge postero-

basally. Uncus prominent, deeply notched distally, arising from

mesal side of prefemoral process at one-quarter to one-third the

process height (distal to solenomere tip).

Distribution and habitat. Known from wet forest at three local-

ities above 600 m in north-east Tasmania; the maximum

distance between the localities is c. 30 km (Fig. 76). The

L. plomleyi range has been carefully searched for millipedes

and this species appears to be genuinely rare.

Etymology. In honour of Brian Plomley (1912-1994),

Tasmanian scientist and scholar who encouraged biological

research in north-east Tasmania.

Lissodesmus tarrabulga sp. nov.

Figures 65, 66, 69tar, 70tar, 71 tar, 78 (map)

Material examined. Holotype. Male, Australia, Victoria. North-east of

Balook, 38°24'52"S 146°34'15"E, 620 m, 22.xii.2004, R. Mesibov,

NMV K-9636.

Paratypes. Male, Jeeralang West Road, 0.1 km N of Binns Hill

Junction, 38°26'30"S 146°29'E, pitfall emptied lO.i.1996, G. Milledge,

ex sample NOH-2303, NMV K-9631; male, same details but pitfall

emptied 5.iii.l996, ex sample NOH-2304, NMV K-9632; 3 males, W

of Balook, 38°26'25"S 146°31'00"E, 600 m, 29.ix.2004, R. Mesibov &

T. Moule, NMV K-9633 to K-9635, 2 dissected; 2 males, details as for

holotype, AM KS91185; male, details as for holotype, anterior

segments only, NMV K-9637.

Description. Male c. 17 mm long, H = 1.4 mm. In alcohol,

well-coloured specimens under low magnification pale brown

in body colour with dispersed brown speckling on metater-

gites'. Antennae moderately long (Fig. 69tar), about 1.75x a

socket diameter apart. Paranota wide, R = 1.6, lateral edge

parallel to body axis, 2-3 inconspicuous posterior marginal

teeth (Fig. 70tar), posterior comers not turned up. Legs robust,

tarsus considerably longer than femur, tibia with prominent

ventral distal swelling (Fig. 71tar). Telopodite (Figs 65, 66)

reaching leg 6 when retracted. Solenomere arising at half the

telopodite height, directed distally, terminating at just over

half the prefemoral process height, a rod-like, apically

toothed, anteriorly directed projection arising at half the

solenomere length on its anterior surface. Tibiotarsus rod-

like, pointed, directed posterodistally at small angle to

telopodite axis, narrower than solenomere and about half as

long. Femoral process arising at level of solenomere origin,

blade-like, tapering, bluntly pointed, directed distally, not

pressed close to prefemoral process, terminating at almost

three-quarters the prefemoral process height (just distal to

solenomere tip). Prefemoral process at origin about half as

wide as telopodite base, tapering strongly distally with

simply pointed tip curving posteriorly. Uncus prominent,

arising on mesal side of prefemoral process at just over half the

process height (just proximal to solenomere tip).

Distribution and habitat. Known from wet eucalypt forest at

three sites over a 10 km section of the Strzelecki Ranges near

Balook, in Gippsland (Fig. 78).

Etymology. Named for Tarra-Bulga National Park, the apparent

centre of the small known range of this species.

Tasmanopeltis gen. nov.

Type species. Tasmanopeltis grandis, by present designation.

Diagnosis. Larger dalodesmids (30-35 mm long, 3.3 mm verti-

cal diameter in type species) with head + 20 segments, normal

pore formula, well-developed paranota with long posterior

comer seta, hair-like structures arising from spiracles, spiracles

on diplosegments juxtaposed over anterior leg. Telopodite

straight, with small mesal solenomere and tibiotarsus, small

lateral femoral process and large central prefemoral process.

All processes arising at more than half the telopodite height;

solenomere arising at almost two-thirds the telopodite height.

Prefemoral process tip divided subapically into erect lateral

branch with comb of long, mainly posterobasally directed teeth,

and shorter, erect mesal branch with short, blunt, distally

directed teeth.

Distribution. As for T. grandis sp. nov.

Etymology. A variation of Australopeltis Johns, 1964, which

Johns probably derived from Latin pelta (“shield”); masculine.

Remarks. Tasmanopeltis is distinguished from Lissodesmus by

its unusually large size (nearly two and a half times the calcu-

lated body volume of the next largest species, L. montanus, and

more than 10 times that of the type, L. modestus)\ spiracles

close together and with hair-like structures; process origins

more than halfway along the telopodite; and a large, distal

124

Robert Mesibov

projection on the prefemoral process, mesal to the latter’s

comb-like tip.

Tasmanopeltis grandis sp. nov.

Figures 5B, 67, 68, 69gra, 70gra, 71gra, 75 (map)

Lissodesmus sp. NE1. — Mesibov, 1994: 134. — Mesibov, 1996:

17.— Mesibov, 1997: 567.— Mesibov, 1998: 155.

Material examined. Holotype. Male, Australia, Tasmania. Speck

Creek, EQ557542 (41°03'41"S 147°39'46"E), 330 m, 18.iii.1993,

R. Mesibov, QVM 23:15555.

Paratypes. Male, Tin Hut Creek, EQ930505 (41°05'28"S

148°06'26"E), 140 m, 19.L1989, R. Bashford, QVM 23:40767, pitfall;

male, Eagle Hill, EQ550581 (41 o 01'34"S 147°39’15”E), 170 m,

10.iii.1993, R. Mesibov, AM KS91186 (ex QVM 23:15567); male,

same details, QVM 23:15567; female, Speck Creek, EQ558537

(41°03'57"S 147°39'50"E), 150 m, 10.iii.1993, R. Mesibov, QVM

23:15598; male. Speck Creek, EQ576538 (41°03'53"S 147°41'07"E),

230 m, lO.iii. 1993, R. Mesibov, QVM 23:15554, dissected; male, Mt

Horror, EQ604547 (41°03'23’vS 147°43'07"E), 220 m, 18.iii.1993,

R. Mesibov, QVM 23:15557, dissected; male, Mt Horror, EQ588513

(41°05'14"S 147°42'00"E), 200 m, 24.iv.1993, R. Mesibov, QVM

23:15558; male, Mt Dismal area, EQ072330 (41°15'15"S

147°05'09 M E), 320 m, 20.U995, R. Mesibov, QVM 23:17762; male,

Ben Lomond, EQ483057 (41°29’55”S 147°34'43"E), 720 m,

16.vi.1995, R. Mesibov, QVM 23:21516.

Other material. 41 males, 49 females and 90 juveniles from Ansons

River, “Aplico” property, Amon River, Back Creek, Barrow Creek,

Bell Creek, Ben Lomond, Ben Nevis, Blacksnake Marsh, Brady Kayes

Lookout, Branxholm, Burns Creek, Caldbeck Creek, Carneys Creek,

Chinaman Comer, Constance Creek, Duncraggen Hill, Eagle Hill,

Foons Hill, Friend Creek, Glennons Road, Great Forester River,

Harrys Hill, Hogarth Rivulet, Ikes Creek, Jewells Creek, Joseph Creek,

Kamona, Martins Hill, Monazite Creek, Morris Ridge, Mother Logans

Creek, Mt Arthur, Mt Cameron, Mt Dismal, Mt Horror, Mt Stronach,

Musselboro, North Esk River, Northallerton Valley, Nunamara,

Ockerbys Hills, Oxberry Creek, Patersonia, Pearly Brook, Rattler Hill,

Scottsdale Mine, Sideling Range, Simons Road, Speck Creek, Spurrs

Rivulet, Swanee Creek, Telita, Tin Hut Creek, Traills Point, Weavers

Creek, Wild Pig Hill, Williams Creek, Williams Hill and Wyena (see

“ Lissodesmus supplement” for details).

Description. Male c. 32 mm long, H = 3.3 mm. In alcohol,

well-coloured specimens red- to purple-brown dorsally, pale

yellow-orange laterally, ventrally and on legs; at low magnifi-

cation with fairly uniform red- to purple-brown colouring on

metazonite, pale yellow-orange on paranotal and posterior

metazonal margins, somewhat darker red- to purple brown on

head and prozonites, legs darker dorsally, distal podomeres and

antennae pale red. Antennae (Fig. 69gra) moderately long,

slender, separated by about 2x socket diameter, sockets not

deeply impressed; antennomere 6 widest, antennomere lengths

typically decreasing in order 2, 3, 6, (4,5). Head with clypeus

and frons densely setose, vertex bare; vertigial sulcus extending

to point about 1.5x socket diameter from line joining socket

centres. Collum from above slightly wider than head, with

nearly straight anterior border, steeply sloping shoulders,

rounded posterior comers, slightly convex posterior margin

with emarginate centre; posterior corner setae present and a few

long setae in transverse line behind anterior margin and

centrally; collum comers slightly ventral to paranotal margin of

segment 2. Paranota (Fig. 70gra) increasing in width from

segment 2 to 5, fairly uniform in width posteriorly to segment

16, narrowing thereafter; paranota wide, R = 1.8 on segment

12; lateral margins more convex and posterior corners more

produced posteriorly, on most segments the paranota widest

anteriorly; lateral margins with 3-4 notches, typically each with

short seta, the most anterior notch with longest seta, sometimes

as long as posterior comer seta, the latter arising on most seg-

ments on mesal side of projecting posterior corner, close to

base of latter; the margins straight in lateral view but inclined

upwards posteriorly at c. 15° to horizontal. Segment surfaces

smooth, metatergites bare apart from short, sparse setae in 2-3

transverse rows anteriorly. Pore formula 5, 7, 9, 10, 12, 13,

15-19; ozopore opening dorsolaterally on paranotum at about

three-quarters the distance from anterior to posterior paranotal

margins, close to lateral margin. Spiracles (Fig. 5B) enlarged,

on diplosegments the posterior spiracle almost touching anter-

ior spiracle over anterior leg, rims very thin, projecting, all

spiracles with dense mass of emergent hair-like stmctures.

Sternite tetrads longer than wide, transverse furrow deep,

longitudinal furrow less deep. Legs (Fig. 71gra) variably

swollen from segment 3 posteriorly, mostly with strongly

arched prefemur, slightly arched femur, slight ventral distal

swellings on postfemur and tibia; tarsus slightly curved;

podomere lengths decreasing in order tarsus, femur, prefemur,

(postfemur, tibia); dense “brush” setae on prefemur, femur,

postfemur; sphaerotrichomes on all five podomeres on legs

near gonopods; leg 2 coxa with genital opening on mesodistal

swelling, only slightly raised above coxal surface; paired coxae

of legs 6 and 7 well separated, slightly separated on legs 5, not

separated on legs 4. Gonopod aperture about one-third as wide

as segment 7 prozonite, more or less rectangular (wider than

long) with rounded comers in ventral view, in lateral view with

lateral margin strongly produced posteriorly. Gonopod coxae

entirely contained within aperture, fairly small, more or less

ellipsoid, lightly joined along midline distally, with dense, short

setae on ventromesal surface distally; cannulae prominent.

Telopodites (Figs 67, 68) separate, straight, lightly setose pos-

teriorly from base to level of solenomere origin, reaching leg

5 when retracted. Solenomere arising from anteromesal

surface of telopodite at almost two-thirds the telopodite

height, directed posterodistally at somewhat less than 45° to

telopodite axis, curving very slightly laterally, tapering

gradually, tip curving distally, no subapical projection or

similar structure, terminating at about four-fifths the

telopodite height (about two-thirds the prefemoral process

height). Tibiotarsus arising a little posterior and distal to

solenomere and more or less parallel to it, rod-like, bluntly

pointed, about one-third the solenomere length. Femoral

process arising just proximal to solenomere origin on antero-

lateral surface, mesolaterally flattened, pressed close near

origin to prefemoral process, widening distally with truncate,

sinuous, inconspicuously toothed distal edge, terminating at

just over half the prefemoral process height (proximal to

solenomere tip). Prefemoral process at origin about half as

wide as telopodite base, widening distally and dividing at

about half its height into well-separated lateral and mesal

branches; lateral branch curving posteriorly with a comb of

New genus of millipedes

125

10-15 long, straight, mainly posterobasally directed teeth

on mesal edge; mesal branch one-third to one-half the

height of lateral branch (terminating just distal to

solenomere tip), bluntly pointed with mesal surface armed

with 3-4 short, blunt, posterodistally directed teeth. Uncus

a large posterior extension of mesal edge of prefemoral process

just proximal to solenomere tip, widely notched on outer edge.

Hypoproct paraboloid in outline; preanal ring with numerous

long setae; epiproct projecting well past anal valves, in dorsal

outline triangular with truncated apex. Female slightly larger

than male, c. 35 mm long, legs thinner, otherwise with same

coloration, paranotal form and spiracle features. Posterior rim

of epigynal opening strongly produced, with rounded corners

and elevated central portion; cyphopods not examined.

Distribution and habitat. In dry and wet eucalypt forest and

cool temperate rainforest over c. 3500 knri in north-east

Tasmania (Fig. 75), from Mt Dismal in the Tamar Valley east

to Ansons River, from Gladstone south to Ben Lomond, and

from near sea level to 920 m. The western range boundary of

T. grandis lies within the biogeographical divide known as the

East Tamar Break (Mesibov, 1994), and is particularly sharp

near Weavers Creek (Mesibov, 1997). Adults of T. grandis are

generally found in or under rotting logs and only rarely burrow

into the soil.

Etymology. Latin grandis (“large, magnificent”), adjective.

Remarks. Although T. grandis is the largest polydesmidan in

Tasmania, it is remarkably cryptic in its habits, and the first

known museum specimen was collected less than 20 years ago

(on Mt Arthur in 1987).

Acknowledgements

This study was supported by a grant from the Plomley

Foundation. I thank David Steele (University of Tasmania) for

acquiring the SEM images; Gonzalo Giribet (Harvard

University) for access to the Lissodesmus modestus types; Peter

Lilly white (Museum Victoria) for the loan of specimens;

Genefor Walker-Smith (Tasmanian Museum and Art Gallery)

and Cathy Young (Department of Primary Industries, Water and

Environment, Tasmania) for advice on millipede holdings;

Dennis Black (La Trobe University) for the opportunity to

examine the Lower Gordon River Scientific Survey material

held in the Australian National Insect Collection; and Mark

Harvey (Western Australian Museum) for alerting me to his

collection of L. milledgei. I am grateful to Geoff and Jackie

Tims for inviting me to collect on “Lapoinya”, near Yarragon,

and to other Gippsland landowners who allowed me access to

their properties. I am especially grateful to my wife, Trina

Moule, for her able assistance in the field over many years. A

draft of this paper was greatly improved by comments from

Mark Harvey and Gary Poore.

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Figure l.Lissodesmus perporosus Jeekel, 1984, male. Left lateral view

of midbody paranotum, showing posterior corner seta (arrow). Scale

bar =1.0 mm.

Figure 4. (A) L. modestus Chamberlin, 1920, male from Edwards

Road, Huon River, ex QVM 23:15294; (B) L. hamatus sp. nov., male

paratype ex QVM 23:16073; (C) L. anas sp. nov., male paratype ex

QVM 23:17657. Left lateral SEM views of midbody segment, show-

ing nearly straight paranotal margin (A), slightly upturned posterior

corner (B) and strongly upturned posterior comer (C). Scale bars = 0.5

mm.

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Nomenclator generum etfamiliarum Diplopodorum II. A List of the

Genus and Family-Group Names in the Class Diplopoda from 1958

through 1999. Pensoft: Sofia.

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Figure 2. Diagrammatic posteromesal view of the right gonopod

telopodite of a member of the Lissodesmus group, f = femoral process,

pf = prefemoral process, s = solenomere, tt = tibiotarsus, u = uncus.

Figure 3. Diagrammatic posterior (left) and dorsal (right) views of a

polydesmidan midbody segment, showing measurement limits for

height H, prozonite width wp and paranotal width wm. Relative para-

notal width R - wm/wp.

New genus of millipedes

127

Figure 5. (A) L. modestus Chamberlin, 1920, male from Edwards

Road, Huon River, ex QVM 23: 15294; (B) Tasmanopeltis grandis gen.

et sp. nov., male paratype, QVM 23:15554; (C) L. anas sp. nov., male

paratype ex QVM 23:17657. Left lateral SEM views of midbody

segment, showing spiracles. Scale bars = 0.25 mm (A, C) and 0.35

mm (B).

Figure 6. (A) L. latus sp. nov., male paratype ex QVM 23:17490, (B)

L. hamatus sp. nov., male paratype ex QVM 23:16073. SEM views of

a group of limbus elements on a midbody segment. Scale bars = 0.025

mm; anterior is to the left.

Figure 7. (A) L. modestus Chamberlin, 1920, male from Edwards

Road, Huon River, ex QVM 23:15294; (B) L. adrianae Jeekel, 1984,

male ex QVM 23:21220. Dorsal SEM views of epiproct. Scale bars =

0.1 mm.

128

Robert Mesibov

Figure 8. (A) L. modestus Chamberlin, 1920, male from Edwards

Road, Huon River, ex QVM 23:15294; (B) L. orarius sp. nov., male

paratype ex QVM 23:17683. Ventral SEM views of leg 6 prefemur in

situ. Arrow points to one of seven sphaerotrichomes visible among

“brush” setae on L. modestus prefemur; sphaerotrichomes are lacking

on L. orarius prefemur. Scale bars = 0.1 mm.

Figure 10. L. perporosus Jeekel, 1984, males from Wombat Hill, W of

Waratah. SEM views of paranotal margin of midbody segment.

(A) stadium 5, ex QVM 23:17291; (B) stadium 6, ex QVM 23:17291;

Scale bars = 0.25 mm.

Figure 9. L. latus sp. nov., females. Left dorsolateral SEM views of

midbody segment of typical form (A) from Sterling River, QVM

23: 17525, and furry form (B) from S of Mt Cripps, ex QVM 23: 17516.

Figure 11. L. adrianae Jeekel, 1984, male from Ben Lomond, ex QVM

23:21220. SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 12. L. adrianae Jeekel, 1984, male from Ben Lomond, ex QVM

23:21220. Mesal (left) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 14. L. alisonae Jeekel, 1984, males from Franklin Village near

Launceston (left), ex QVM 23:15440, and “Bowood” property near

Bridport (right), ex QVM 23:15442. Lateral views of right gonopod

telopodite. Scale bar = 0.25 mm.

Figure 13. L. alisonae Jeekel, 1984, male from Franklin Village, ex

QVM 23:15440. Mesal (left) and anterior (right) views of right gono-

pod telopodite. Dashed lines indicate course of prostatic groove; scale

bar = 0.25 mm.

Figure 15. L. anas sp. nov., male paratype ex QVM 23:17657. SEM

view of gonopods in situ. Scale bar = 0.5 mm.

130

Robert Mesibov

Figure 16. L. anas sp. nov., male paratype ex QVM 23:40749. Mesal

(left) and anterior (right) views of right gonopod telopodite. Dashed

lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 17. L. bashfordi sp. nov., male paratype ex QVM 23:45946.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 18. L. bashfordi sp. nov., male paratype ex QVM 23:45946.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 19. L. blackwoodensis sp. nov., male paratype, NMV K-8923.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

New genus of millipedes

131

Figure 20. L. blackxvoodensis sp. nov., male paratype, NMV K-8924.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 21. L. catrionae sp. nov., male paratype, NMV K-8970. SEM

view of right gonopod in situ. Scale bar = 0.5 mm.

Figure 22. L. catrionae sp. nov., male paratype, NMV K-8971. Mesal

(left) and anterior (right) views of left gonopod telopodite. Dashed Figure 23. L. clivulus sp. nov., male paratype ex QVM 23:17667. SEM

lines indicate course of prostatic groove; scale bar = 0.25 mm. view of gonopods in situ. Scale bar = 0.5 mm.

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Robert Mesibov

Figure 24. L. clivulus sp. nov., male paratype ex QVM 23:17667.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 25. L. cognatus sp. nov., male paratype ex QVM 23:15283.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 26. L. cognatus sp. nov., male paratypes from Coxs Creek in the

eastern range patch (far left), ex QVM 23:24738, and Weavers Creek

in the western range patch (centre and right), ex QVM 23:21562.

Mesal (left and centre) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 27. L. cognatus sp. nov., male paratypes from Coxs Creek in the

eastern range patch (left), ex QVM 23:24738, and Weavers Creek in

the western range patch (right), ex QVM 23:21562. Lateral views of

right gonopod telopodite. Scale bar = 0.25 mm.

New genus of millipedes

133

Figure 28. L. cornutus sp. nov., male paratype ex QVM 23:24824. Left

lateral SEM view of midbody segment. Scale bar = 0.5 mm.

Figure 29. L. cornutus sp. nov., male paratype ex QVM 23:24824.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 30. L. cornutus sp. nov., male paratype ex QVM 23:24824.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 31. L. devexus sp. nov., male paratype from Shepherds Rivulet,

6 km SE of the type locality, ex QVM 23:15509. SEM view of

gonopods in situ. Scale bar = 0.5 mm.

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Robert Mesibov

Figure 32. L. devexus sp. nov., male paratype ex QVM 23:15515.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 34. L. dignomontis sp. nov., male paratype, NMV K-9664.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 33. L. devexus sp. nov., male paratypes from (A) Retreat, ex

QVM 23:21534, (B) Weldborough, QVM 23:15512, (C) Mt Roland,

ex QVM 23:15506 and (D) Lowes Mount, QVM 23:21542. Mesal

views of right gonopod telopodite. Scale bar = 0.25 mm.

Figure 35. L. dignomontis sp. nov., male paratype, NMV K-9507.

Posterior (left), mesal (centre) and anterior (right) views of right gono-

pod telopodite. Dashed lines indicate course of prostatic groove; scale

bar = 0.25 mm.

New genus of millipedes

135

Figure 37. L. gippslandicm sp. nov., male paratype, NMV K-9484.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 38. L. hamatus sp. nov., male paratype ex QVM 23:16073.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 39. L. hamatus sp. nov., male paratype ex QVM 23:16073.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

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Robert Mesibov

Figure 40. L. horridomontis sp. nov., male paratype ex QVM

23:15490. SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 42. L. inopinatus sp. nov., male paratype ex QVM 23:16164.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 41. L. horridomontis sp. nov., male paratype ex QVM

23:15490. Mesal (left) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 43. L. inopinatus sp. nov., male paratype ex QVM 23:16164.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

New genus of millipedes

137

Figure 44. L. johnsi sp. nov., male paratype, NMV K-9683. SEM view

of gonopods in situ. Scale bar = 0.5 mm.

Figure 47. L. macedonensis sp. nov., male paratype, NMV K-9511.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 45. L. johnsi sp. nov., male paratype, NMV K-9506. Posterior

(left), mesal (centre) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 46. L. latus sp. nov., male paratype ex QVM 23:17490. Mesal

(left) and anterior (right) views of right gonopod telopodite. Dashed

lines indicate course of prostatic groove; scale bar = 0.25 mm.

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Robert Mesibov

Figure 48. L. macedonensis sp. nov., male paratype, NMV K-9512.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 50. L. martini (Carl, 1902), male from Acheron Gap area, NMV

K-9591. Mesal (left) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 49. L. martini (Carl, 1902), male from Acheron Gap area, NMV

K-9590. SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 51. L. milledgei sp. nov., male paratype, NMV K-9606. Mesal

(left) and anterior (right) views of right gonopod telopodite. Dashed

lines indicate course of prostatic groove; scale bar = 0.25 mm.

New genus of millipedes

139

Figure 52. L. modestus Chamberlin, 1920, male from Edwards Road,

Huon River, ex QVM 23:15294. Posterior SEM view of gonopods in

situ. Scale bar - 0.5 mm.

Figure 54. L. montanus sp. nov., male paratype, QVM 23:17716. SEM

view of gonopods in situ. Scale bar = 0.5 mm.

Figure 53. L. modestus Chamberlin, 1920, male from Edwards Road,

Huon River, ex QVM 23:15294. Mesal view of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 55. L. montanus sp. nov., male paratype, QVM 23:17715.

Mesal view of right gonopod telopodite. Dashed lines indicate course

of prostatic groove; scale bar = 0.25 mm.

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Robert Mesibov

Figure 56. L. orarius sp. nov., male paratype ex QVM 23:17683. SEM

view of gonopods in situ. Scale bar = 0.5 mm.

Figure 57. L. orarius sp. nov., male paratype ex QVM 23:17693.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 58. L. otwayensis sp. nov., male paratype, NMV K-9620. SEM

view of gonopods in situ. Scale bar = 0.5 mm.

Figure 59. L. otwayensis sp. nov., male paratype, NMV K-9619. Mesal

(left) and anterior (right) views of right gonopod telopodite. Dashed

lines indicate course of prostatic groove; scale bar = 0.25 mm.

New genus of millipedes

141

Figure 60. L. peninsulensis sp. nov., male paratype, QVM 23:16173.

SEM view of gonopods in situ. Scale bar = 0.5 mm.

Figure 62. L. perporosus Jeekel, 1984, male from Wombat Hill, W of

Waratah, ex QVM 23: 17419. SEM view of right gonopod in situ. Scale

bar = 0.5 mm.

Figure 61. L. peninsulensis sp. nov., male paratype, QVM 23:16172.

Mesal (left) and anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove; scale bar = 0.25 mm.

Figure 63. L. perporosus Jeekel, 1984, males from Wombat Hill, W of

Waratah (centre and right), ex QVM 23:17419, and Maggs Mountain

(left), QVM 23:17411. Mesal (left and centre) and anterior (right)

views of right gonopod telopodite. Dashed lines indicate course of pro-

static groove; scale bar = 0.25 mm.

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Robert Mesibov

Figure 64. L. plomleyi sp. nov., male paratype, QVM 23:15465. Lateral

(left), mesal (centre) and anterior (right) views of right gonopod

telopodite. Dashed lines indicate course of prostatic groove; scale bar

= 0.25 mm.

Figure 65. L. tarrabulga sp. nov., male paratype, NMV K-9634. SEM

view of gonopods in situ. Scale bar = 0.5 mm.

Figure 66. L. tarrabulga sp. nov., male paratype,

NMV K-9635. Mesal (left) and anterior (right)

views of right gonopod telopodite. Dashed lines

indicate course of prostatic groove; scale bar =

0.25 mm.

Figure 68. Tasmanopeltis grandis gen. et sp. nov.,

male paratype, QVM 23:15557. Mesal (left) and

anterior (right) views of right gonopod telopodite.

Dashed lines indicate course of prostatic groove;

scale bar - 0.25 mm.

Figure 67. Tasmanopeltis grandis gen.

et sp. nov., male paratype, QVM

23:15554. SEM view of gonopods in

situ. Scale bar =1.0 mm.

New genus of millipedes

143

Figure 69. Male antennae of Lissodesmus adrianae Jeekel, 1984 (adr; ex QVM 23:21220), L. alisonae Jeekel, 1984 (ali; ex QVM 23:15440),

L. anas sp. nov. (ana; paratype ex QVM 23:40749), L. bashfordi sp. nov. (bas; paratype ex QVM 23:45946), L. blackwoodensis sp. nov.

(bla; paratype, NMV K-8924), L. catrionae sp. nov. (cat; paratype, NMV K-8971), L. clivulus sp. nov. (cli; paratype ex QVM 23:17667),

L. cognatus sp. nov. (cog; paratype ex QVM 23:24738), L. cornutus sp. nov. (cor; paratype ex QVM 23:24824), L. devexus sp. nov. (dev; paratype

ex QVM 23:15515), L. dignomontis sp. nov. (dig; paratype NMV-9507), L. gippslandicus sp. nov. (gip; paratype NMV K-9484), L. hamatus sp.

nov. (ham; paratype ex QVM 23:16073), L. horridomontis sp. nov. (hor; paratype ex QVM 23:15490), L. inopinatus sp. nov. (ino; paratype ex

QVM 23:16164), L. johnsi sp. nov. (joh; paratype NMV K-9683), L. latus sp. nov. (lat; paratype ex QVM 23:17490), L. macedonensis sp. nov.

(mac; paratype NMV K-9512), L. martini (Carl, 1902) (mar; NMV K-9591), L. milledgei sp. nov. (mil; paratype NMV K-9606), L. modestus

Chamberlin, 1920 (mod; ex QVM 23:15294), L. montanus sp. nov. (mon; paratype QVM 23:17716), L. orarius sp. nov. (ora; paratype ex QVM

23:17693), L. otwayensis sp. nov. (otw; paratype NMV K-9619), L. peninsulensis sp. nov. (pen; paratype QVM 23:16171), L. perporosus Jeekel,

1984 (per; paratype ex QVM 23:17419), L. plomleyi sp. nov. (plo; paratype QVM 23:15466), L. tarrabulga sp. nov. (tar; NMV K-9635) and

Tasmanopeltis grandis gen. et sp. nov. (gra; paratype QVM 23:15558). Scale bar =1.0 mm.

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Robert Mesibov

Figure 70. Left side dorsal profiles of segment 12 of male Lissodesmus spp. and Tasmanopeltis grandis gen. et sp. nov., and of segment 13 of male

L. plomleyi sp. nov. Species codes and specimen details as in Fig. 65. The ozopore opening is shown as a small ellipse near the paranotal margin

in all cases. Scale bar =1.0 mm.

Figure 72. Localities in Tasmania of Lissodesmus adrianae (triangles),

L. bashfordi sp. nov. (open squares), L. inopinatus sp. nov. (crosses),

L. peninsulensis sp. nov. (dots) and L. perporosus (filled squares).

Figure 73. Localities in Tasmania of Lissodesmus alisonae (triangles),

L. clivulus sp. nov. (filled squares), L. horridomontis sp. nov. (cross-

es), L. modestus (dots) and L. montanus sp. nov. (open squares).

New genus of millipedes

145

Figure 71. Male leg 7 of Lissodesmus spp. and Tasmanopeltis grandis gen. et sp. nov. Species codes and specimen details as in Fig. 65, but L.

plomleyi sp. nov. specimen is paratype QVM 23:15465 and T. grandis specimen is paratype QVM 23:15557. Scale bar = 1.0 mm.

Figure 74. Localities in Tasmania of Lissodesmus anas sp. nov. (dots),

L. comutus sp. nov. (squares) and L. devexus sp. nov. (triangles).

Figure 75. Localities in Tasmania of Tasmanopeltis grandis sp. nov.

(squares) andL. latus sp. nov. (dots).

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Robert Mesibov

Figure 76. Localities in Tasmania of Lissodesmus cognatus sp. nov.

(squares), L. orarius sp. nov. (dots) and L. plomleyi sp. nov. (crosses).

Figure 78. Localities in Victoria of Lissodesmus blackwoodensis sp.

nov. (crosses), L. catrionae sp. nov. (filled square), L. dignomontis sp.

nov. (filled triangles), L. johnsi sp. nov. (open triangle), L. milledgei

sp. nov. (dots) and L. tarrabulga sp. nov. (open squares). For the sake

of clarity, some neighbouring localities are marked with single

symbols.

Figure 77. Localities in Tasmania of Lissodesmus hamatus sp. nov.

(squares).

Figure 79. Localities in Victoria of Lissodesmus gippslandicus sp. nov.

(crosses), L. macedonensis sp. nov. (triangles), L. martini sp. nov.

(squares) and L. otwayensis sp. nov. (dots). For the sake of clarity,

some neighbouring localities are marked with single symbols.

Memoirs of Museum Victoria 62(2): 147-179 (2005)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A review of pygal-furrowed Synallactidae (Echinodermata: Holothuroidea), with

new species from the Antarctic, Atlantic and Pacific oceans

P. Mark O'Loughlin 1 and Cynthia Ahearn 2

'Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia

(pmo@bigpond.net.au)

2 Museum Specialist - Echinoderms, Department of Invertebrate Zoology, National Museum of Natural History,

Smithsonian Institution, PO Box 37012, Washington DC, 20013-7012, USA (ahearn.cynthia@nmnh.si.edu)

Abstract O’Loughlin, P.M., and Ahearn, C. 2005. A review of pygal-furrowed Synallactidae (Echinodermata: Holothuroidea),

with new species from the Antarctic, Atlantic and Pacific oceans. Memoirs of Museum Victoria 62(2): 147-179.

We review the genera and species of pygal-furrowed Synallactidae and describe new species from the collections of

the United States National Museum of Natural History (Smithsonian Institution). Meseres Ludwig is reviewed and placed

incertae sedis. Molpadiodemas Heding and Pseudostichopus Theel are rediagnosed. Platystichopus Heding is declared a

junior synonym of Molpadiodemas. Filithuria Koehler and Vaney and Peristichopus Djakonov are declared junior

synonyms of Pseudostichopus. Lectotypes are designated for: Meseres involutus Sluiter, M. macdonaldi Ludwig,

M. peripatus Sluiter, Pseudostichopus globigerinae Herouard, P. mollis Theel, P. pustulosus Sluiter, P. trachus Sluiter,

P. villosus Theel. A neotype is designated for Pseudostichopus nudus Ohshima.

Except for the type species, Meseres macdonaldi Ludwig, and Meseres (?) torvus (Theel), all species previously

referred to Meseres are reassigned to Molpadiodemas or Pseudostichopus. Meseres (?) torvus (Theel) is retained in

its original combination. Eleven new synallactid species are described: Molpadiodemas constrictus, M. crinitus,

M. epibiotus, M. helios, M. morbillus, M. neovillosus, M. pediculus, M. porphyrus, M. translucens, M. ustulatus and

Pseudostichopus tuberosus. Molpadiodemas constrictus is fissiparous. Pseudostichopus villosus var. violaceus Theel is

raised to species status; Pseudostichopus alatus Imaoka, P. trachus Sluiter and P. nudus Ohshima are junior synonyms

of Pseudostichopus mollis Theel; Pseudostichopus globigerinae Herouard and P. dilatorbis Imaoka are junior synonyms

of Meseres (= Molpadiodemas ) involutus Sluiter; Pseudostichopus propinquus Lisher and P. lapidus Herouard are

junior synonyms of Meseres (= Pseudostichopus ) peripatus Sluiter; and Pseudostichopus arenosus Ohshima and

P. molpadioides Ohshima are junior synonyms of Meseres (= Pseudostichopus ) hyalegerus Sluiter.

New combinations are: Molpadiodemas atlanticus (Perrier), M. depressus (Herouard), M. involutus (Sluiter),

M. pustulosus (Sluiter), M. villosus (Theel), Pseudostichopus elegans (Koehler and Vaney), P. hyalegerus (Sluiter),

P. papillatus (Djakonov), P. peripatus (Sluiter) and P. spiculiferus (O'Loughlin). Other species discussed are:

Molpadiodemas violaceus (Theel), Pseudostichopus aemulatus Solfs-Marm and Billett, P echinatus Thandar, P. mollis

Theel, P. occultatus Marenzeller, and P. profundi Djakonov. A key is provided for the species of Molpadiodemas and

Pseudostichopus. Tables are provided of genera and subgenera with current systematic status, and species with original

and current combinations.

Introduction

We report on the United States National Museum of Natural

History (Smithsonian Institution) collections of synallactid

holothurian species from the Atlantic, Pacific and Antarctic

oceans. Only pygal-furrowed genera are dealt with. The pygal-

furrowed synallactids substantially lack body wall ossicles and

have a distinct pygal (posterior) vertical furrow.

Pygal-furrowed synallactids in the USNM collections com-

prise 345 catalogued lots (several thousand specimens) from a

variety of sources. Operation Deep Freeze (USS Edisto ) and

the United States Antarctic Program (under the auspices of the

National Science Foundation, Office of Polar Programs) con-

tributed about 47 % of the lots (research vessels USNS Eltanin,

now the ARA Islas Orcadas\ RV Hew, USCGS Eastwind).

Between the years 1884 and 1911, the United States Fish

Commission mounted expeditions to the North-west Pacific,

Philippines, Hawaii, eastern Pacific, California coast, lower

California and western Atlantic, and provided material (USFCS

Albatross ). When the USFC became the United States Bureau

of Fisheries, cruises were conducted throughout the western

Atlantic that contributed several lots from a 1970 cruise to the

Gulf of Mexico (RV Oregon II). The Fish Commission, and

later the Bureau of Fisheries, contributed about 21% of the lots.

148

P. Mark O’Loughlin and Cynthia Ahearn

About 12% of the lots were provided by the Ocean Minerals

Company from a benthic sampling and photographic survey of

the Clarion-Clipperton Fracture Zone in the eastern central

Pacific during 1978-1980 (RV Governor Ray).

The remaining 20% of lots were donated by various institu-

tions, universities and agencies. A significant contribution of

two new species described herein ( Molpadiodemas epibiotus

and M. constrictus) was made by the University of Miami from

cruises in 1973 and 1975 to the Puerto Rico and Cayman

Trenches (RV Gilliss). The Southampton Oceanographic

Centre (now National Oceanographic Centre) surveys of the

Western European Basin in 1979 (RV Challenger) and

Porcupine Abyssal Plain in 1999 (RV Discovery) added two

lots which included paratypes of the recently described

Pseudostichopus aemulatus. Thirty lots were contributed by

Oregon State University from survey cruises off the coast of

Oregon in the 1960s and 1970s (RV Commando, RV Cobb, RV

Yaquina, RV Cay use). The Smithsonian Oceanographic Center,

supporting the International Indian Ocean Expedition and the

Southeast Pacific Benthic Oceanographic Program, added 8

lots from cruises during the 1960s to Thailand and many south-

eastern Pacific localities (RV Anton Bruun, RV Te Vega). Six

lots were contributed by the Woods Hole Oceanographic

Institute following cruises off Argentina in 1960 (RV Atlantis

II), to the West European Basin in 1972 (RV Chain), and to the

North American Basin in 1973 (RV Knorr). Several lots were

added by the Virginia Institute of Marine Science cruises to the

Bahamas in 1978, 1980, and 1981 (RV Columbus Iselin). Texas

A&M University provided one lot from a 1972 cruise to the

Gulf of Mexico (RV Alaminos). Three lots were provided by

the US Bureau of Land Management surveys off South

Carolina in 1985 (RV Cape Hatteras), and off Louisiana during

the Northern Gulf of Mexico Continental Shelf project.

Fourteen lots were donated by M.D. Richardson from a survey

of the Venezuelan Basin in 1981 (RV Bartlett). One lot was

donated by the Vienna Royal Imperial Natural History Museum

from a survey of the eastern Mediterranean from 1890-1894

(HMS. Pola). Two lots were recently donated by Dr Tohru

Imaoka (ex-Seto Marine Laboratory and Faculty of Science at

the Kyoto University in Shirahama), one lot by Dr Ken Smith

(Scripps Institution of Oceanography) and another two lots by

Dr David Stein (Systematics Laboratory, National

Oceanographic and Atmospheric Administration). Two lots

have insufficient information as to the source of the material.

During April 2002, Mark O’Loughlin visited the Natural

History Museum in London and the Zoologisch Museum at the

University of Amsterdam, and designated most of the lecto-

types selected in this work. The scanning electron microscopy

and most of the digital imaging were done by Cynthia Aheam.

We collaborated on systematic observations and decisions.

Because the ossicles in each pygal-furrowed synallactid

species do not have a consistent form and frequently vary

considerably, the SEM images selected for each species do not

represent the full range of forms. They should be viewed in

conjunction with the text description of forms and sizes.

In the diagnosis of Meseres by O’Loughlin (2002),

reference was made to thin tubular appendages which are

rhizomatous and sometimes branched. It is judged here that

these are epibiotes, and not holothurian appendages and not

species specific.

Abbreviations for institutions are: AM, Australian Museum,

Sydney; BMNH, The Natural History Museum, London;

MNHN, Museum National d’Histoire Naturelle, Paris; MOM,

Musee oceanographique de Monaco; NMV, Museum Victoria,

Australia; OMNH, Osaka Museum of Natural History, Japan;

RAS, Russian Academy of Sciences, St. Petersburg, Russia;

SAM, South Africa Museum, Cape Town; SMBL, Seto Marine

Biological Laboratory, Kyoto University, Japan; TM,

Tasmanian Museum, Hobart; UNAM, Universidad Nacional

Autonoma de Mexico; USNM, US National Museum of

Natural History, Smithsonian Institution, Washington; ZMA ,

Zoologisch Museum, Amsterdam; ZMUC, Zoological

Museum, University of Copenhagen, Denmark.

Historically, three types of registration number have been

used for USNM material, and occur in this report. Prior to

1920, museum catalogue numbers without a prefix were used.

In 1920, A.H. Clark began a dedicated echinoderm numbering

system with the prefix E before the registration number. Since

2001, the EMU on-line system is used, and registrations report-

ed without an E prefix.

Throughout this paper Rowe (in Rowe and Gates, 1995) is

referred to as Rowe (1995). Descriptions and measurements

refer to preserved material.

History of USFCS Albatross 1906 material

Numerous holothurian types and other material collected in the

north-west Pacific during the summer of 1906 by the USFCS

Albatross cannot be located. Correspondence in the National

Museum of Natural History’s Registrar Office indicates that the

final dispository of the material was to be the Smithsonian

Institution. After the death of Dr Kakichi Mitsukuri (1909) the

United States Bureau of Fisheries transferred the holothurians

to his graduate student Mr Hiroshi Ohshima. He judged 46 of

the 95 species and one subspecies to be new, and requested

USNM catalog numbers for the type specimens for inclusion in

his manuscript. The results of this study were published

(Ohshima, 1915), but the specimens were not returned at the

time (possibly due to World War I). In 1938, the Director of the

Zoological Institute of the Tokyo Imperial University, Naohide

Yatsu, discovered this collection stored in a cellar. Yatsu corres-

ponded with Alexander Wetmore, then Assistant Secretary of

the Smithsonian Institution, informing him of the holothurian

collection and seeking advice as to the best way to ship the

material. In April 1938 the holothurian collection in four

wooden crates was transported on the Kwansai Maru via the

Panama Canal to New York. In correspondence dated 10 June,

1938, Alexander Wetmore acknowledged receipt of the four

crates in good condition.

The four crates should have been delivered to Dr Arthur H.

Clark, curator of the Echinoderm Division in the NMNH. Some

material was in fact returned and is now in the echinoderm col-

lection. The fate of the remaining material is unknown.

Archives have been searched for official memoranda, curators

of other departments contacted, and the Natural History

Museum basement and attic searched without success. Dr

A review of pygal-furrowed Synallactidae

149

Frederick M. Bayer, curator and research scientist of the

Coelenterata collection, knew Dr Clark and recalled that he

had many unpacked boxes in his office and that some were

probably discarded.

Key to species of pygal-furrowed Synallactidae

(The generic assignment of Pseudostichopus profundi Djakonov

is unresolved. The species is distinguished by the unique

presence of curved mesh-like narrow plates in the tube feet.)

1. Tube feet not in prominent series on paired radii; longitu-

dinal muscles flat, broadly attached to internal body wall;

gonad tubules arising from common source at end of

gonoduct, branched Molpadiodemas Heding ... 2

— Tube feet or papillae in prominent series on paired radii;

longitudinal muscles cylindrical, narrowly attached to

internal body wall; gonad tubules in series along gonoduct,

unbranched Pseudostichopus Theel ... 17

2. Tube feet hair-like or thin, thread-like 3

— Tube feet cylindrical, prominent or inconspicuous .... 4

3. Body lacking brim; tube feet hair-like, over whole body,

sometimes matted ventrally, anteriorly, pygally, frequently

withdrawn ventrally; tentacle ossicles never plate-like rods

with close fine spines Molpadiodemas villosus

— Body with rounded lateral brim, sometimes with pustulose

bulges; tube feet soft, flaccid, thread-like, sometimes

matted on small specimens, readily lost with outer body

layer; tentacle ossicles include wide plate-like rods with

close fine spines Molpadiodemas porphyrus

4. Body with one, sometimes two, deep transverse

constrictions; no tentacle ossicles

Molpadiodemas constrictus

— Body lacking deep transverse constrictions; tentacle

ossicles present 5

5. Body typically with one or two concave depres-

sions created by epibiotic attachments; body strongly

posteriorly tapered; tentacle ossicles include thin, lumpy

rods Molpadiodemas epibiotus

— Body lacking concavities; body not strongly posteriorly

tapered; tentacle ossicles not including thin lumpy rods ... 6

6. Ventrolateral margin with distinct raised protuberances ... 7

— Ventrolateral margin lacking distinct raised protuberances

7. Margin serrated by transverse creasing of body; ventrolat-

eral protuberances lacking tufts of tube feet; body wall

wrinkled, with small digitate projections on low reticulate

ridges; tentacle ossicles include rods; body typically

encrusted with globigerines or small stones

Molpadiodemas involutus

— Ventrolateral margin with pustules surmounted by nipple-

like protuberances; tube feet in tufts on posterior pustules;

body wall not wrinkled, lacking low reticulate ridges;

tentacle ossicles lace mesh only; some globigerine

attachments Molpadiodemas pustulosus

8. Body wall thin, translucent; gonad ossicles predominantly

thin lace mesh, not developed along primary rods

Molpadiodemas translucens

— Body wall not translucent; gonad ossicles, if present, not

predominantly thin lace mesh, if mesh, developed along

primary rods 9

9. Body wall thick, gelatinous; ventrolateral brim thick,

rounded Molpadiodemas depressus

— Body wall not thick, gelatinous; lacking thick, rounded

brim 10

10. Cylindrical tube feet concentrated in conspicuous ventro-

lateral band; frequently contiguous, matted ventrolaterally

— Tube feet not contiguous or matted ventrolaterally ... 13

11. Tube feet typically 2.0 mm long, 0.4 mm diameter;

frequently tubercles and divided pygal lobes posteriorly .

Molpadiodemas crinitus

— Tube feet typically less than 2.0 mm long, 0.4 mm

diameter; lacking posterior tubercles and divisions of

pygal lobes 12

12. Body short, diameter large (about 2:1); tube feet typically

1.6 mm long, 0.3 mm diameter; longitudinal muscles wide;

gonad ossicles long, thin rods

Molpadiodemas neovillosus

— Body elongate, diameter small (about 5:1), long tapers

distally; tube feet typically 1.0 mm long, 0.25 mm

diameter; longitudinal muscles narrow; lacking gonad

ossicles Molpadiodemas pediculus

13. Tube feet cylindrical, most evident as lateral band, not

matted 14

— - Tube feet around body, not more evident as lateral band .

14. Tube feet dark reddish brown creating a lateral spotted

appearance; lacking gonad ossicles

Molpadiodemas morbillus

— Whole body dark brown, scorched appearance; gonad

ossicles present Molpadiodemas ustulatus

15. Tube feet soft, scattered, lying all over body in larger

specimens, sometimes erect orally and pygally; tentacles

with large, conical, pointed peripheral digits

Molpadiodemas helios

— Tube feet not soft, scattered, lying all over body in larger

specimens; tentacle digits not large, conical, pointed . . 16

16. Even cover of small tube feet, frequently in pits or not

evident, cylindrical and firm if extended; tentacle ossicles

lacking knob-like central swelling; high frequency of

gonad ossicles comprising rods with extensive lateral mesh

Molpadiodemas atlanticus

— - Even cover of small, soft, cylindrical tube feet, evident

only on small specimens, never erect; many tentacle

ossicles with knob-like central swelling; rare presence of

gonad ossicles Molpadiodemas violaceus

17. Body densely encrusted with sponge spicules or globi-

gerines or shells or small stones 18

— - Body not encrusted with attachments 22

18. Body covered predominantly with dense mat of sponge

spicules 19

— - Body covered predominantly with globigerines

(foraminiferans) or shells or stones 20

19. Large dense mesh ossicles in pygal lobes; lacking gonad

ossicles Pseudostichopus hyalegerus

150

P. Mark O’Loughlin and Cynthia Ahearn

— Lacking mesh ossicles in pygal lobes; gonad ossicles

include complex branching rods

Pseudo stichopus spiculiferus

20. Predominantly pointed shells with globigerines and sand

cover; closely knobbed tentacle rod ossicles; lacking

ossicles in gonads Pseudo stichopus echinatus

— Predominantly globigerine or small stone cover; tentacle

ossicles not closely knobbed rods; ossicles frequently in

gonads 21

21. Ossicles in pygal lobes and respiratory trees; pygal lobe

ossicles irregular perforated plates, sometimes double

layered Pseudo stichopus occultatus

— Lacking ossicles in pygal lobes and respiratory trees ....

Pseudostichopus peripatus

22. Body with radial series of tubercles ventrolaterally or on

paired radii 23

— Body lacking radial series of tubercles 24

23. Double series of tubercles on paired radii; ossicles present

in papillae, lacking in gonads .Pseudostichopus papillatus

— - Single series of lateroventral tubercles; ossicles in gonads,

lacking in tube feet Pseudostichopus tuberosus

24. Paired radii with long tapering papillae (frequently more

than 10 mm long) Pseudostichopus elegans

— Paired radii with short tapering papillae (up to 5.0 mm

long) 25

25. Ossicles in tube feet, perianally, lacking in gonads

Pseudostichopus mollis

— Ossicles in gonads, lacking perianally and in tube feet . . .

Pseudostichopus aemulatus

Order Aspidochirotida Grube, 1840

Synallactidae Ludwig, 1894

Table 2, Figure 1

Remarks. Although the Synallactidae (sensu lato) are charac-

terized as deep-water forms that possess peltate tentacles, lack

tentacle ampullae, have gonads in one or two tufts, and usually

possess body wall ossicles (including tables and rods), this

work embraces only those species of Synallactidae that sub-

stantially lack body wall ossicles, and have in addition a

distinct pygal (posterior) vertical furrow. The pygal-furrowed

Synallactidae are characterised by: a generally cylindrical body

form; rounded anterior and posterior ends; ventral mouth and

anus; presence of a pygal furrow; 18-20 peltate tentacles;

absence of tentacle ampullae; a complete cover of small and

frequently inconspicuous tube feet; absence of retractor

muscles; longitudinal muscles never divided; a solid calcareous

ring; radial plates broader than high, anteriorly with a central

notch and two central and two lateral low projections, poster-

iorly with a shallow smooth indentation and two pairs of

"teeth" variably evident; interradial plates broad and low, anter-

iorly with a central spire, posteriorly with a smooth shallow

indentation; madreporite sometimes evident externally on the

anterior dorsal body surface; single, sac-like, elongate or short,

ventral polian vesicle; gonad tubules in two groups, one on

each side of the dorsal mesentery; respiratory trees comprising

two long unequal branches of alveolar-like clusters along a

central strap; ossicles absent from the body wall, but present in

the pygal lobes in some species.

Body wall, pygal lobes, tube feet, tentacles, gonads, and

respiratory trees were examined for ossicles in all species, and

are reported only as found, not as absent. The colour of tenta-

cles and internal anatomical features was noted throughout the

study, and found to be not diagnostically reliable. The form of

the peltate tentacles, which varies greatly with state of preser-

vation, was found to be not diagnostically reliable. The pres-

ence or absence of "teeth" on the posterior indentation of the

radial plates of the calcareous ring has been used diagnostic-

ally by some authors. This feature was found to be variable and

subjective, and depended on the angle from which the plates

were viewed. Descriptions of these features have been omitted.

The genera included in this paper have been shifted between

the Gephyrothuriidae (now restricted after O’Loughlin, 1998)

and Synallactidae (see O’Loughlin, 1998, 2002). In the

Zoological Catalogue of Australia Rowe (1995) referred

species of pygal-furrowed Synallactidae to the genera Meseres

Ludwig, 1894 (incertae sedis below) and Pseudostichopus

Theel, 1886 in the Gephyrothuriidae. Following the rediagno-

sis below of Pseudostichopus Theel, Meseres peripatus Sluiter,

1901 is reassigned to Pseudostichopus. Following examination

of the AM collections by one of us (M.O’L), as far as the

Australian fauna is concerned, Rowe (1995) correctly identi-

fied Pseudostichopus mollis Theel and Pseudostichopus

peripatus (Sluiter) material, but wrongly identified some

material as Pseudostichopus pustulosus Sluiter, 1901 (=

Molpadiodemas pustulosus below). We judge that this material

is Pseudostichopus mollis, and that Molpadiodemas pustulosus

(Sluiter) is not to date represented in the Australian fauna.

Table 1 . Index of genera and subgenera in paper, with current status including senior synonym.

Genera and subgenera

Current status or senior synonym

Family

Filithuria Koehler and Vaney, 1905

Pseudostichopus Theel, 1886

Synallactidae

Meseres Ludwig, 1894

Incertae sedis

Molpadiodemas Heding, 1935

Synallactidae

Peristichopus Djakonov, 1952

Pseudostichopus Theel, 1886

Synallactidae

Platy stichopus Heding, 1940

Molpadiodemas Heding, 1935

Synallactidae

Plicastichopus Heding, 1940

Pseudostichopus Theel, 1886

Synallactidae

Pseudostichopus Theel, 1886

Synallactidae

Trachostichopus Heding, 1940

Pseudostichopus Theel, 1886

Synallactidae

A review of pygal-furrowed Synallactidae

151

Table 2. Index of species in paper in original combination, and with current combination or senior synonym.

Species in original combination Current combination or senior synonym

aemulatus, Pseudostichopus , Solis-Marin and Billett, 2004

acaudum, Molpadiodemas, Heding, 1935

alatus, Pseudostichopus , Imaoka, 1990

aleutianus, Pseudostichopus, Ohshima, 1915

arenosus, Pseudostichopus, Ohshima, 1915

atlanticus, Pseudostichopus, Perrier, 1 898

constrictus, Molpadiodemas, sp. nov.

crinitus, Molpadiodemas, sp. nov.

depressus, Pseudostichopus, Herouard, 1902

dilatorbis, Pseudostichopus ( Pseudostichopus ), Imaoka, 1978

echinatus, Pseudostichopus, Thandar, 1992

elegans, Filithuria, Koehler and Vaney, 1905

epibiotus, Molpadiodemas, sp. nov.

globigerinae, Pseudostichopus, Herouard, 1923

helios, Molpadiodemas, sp. nov.

hyalegerus, Meseres, Sluiter, 1901

ingolfi, Plicastichopus, Heding, 1942

involutus, Meseres, Sluiter, 1901

japonensis, Pseudostichopus, Imaoka, 1978

japonensis, Pseudostichopus ( Trachostichopus ), Imoka, 1978

lapidus, Pseudostichopus, Herouard, 1923

macdonaldi, Meseres, Ludwig, 1894

marenzelleri, Pseudostichopus, Herouard, 1923

mollis, Pseudostichopus, Theel, 1886

molpadioides, Pseudostichopus, Ohshima, 1915

morbillus, Molpadiodemas, sp. nov.

neovillosus, Molpadiodemas, sp. nov.

nudus, Pseudostichopus, Ohshima, 1915

occultatus, Pseudostichopus, Marenzeller, 1893

occultatus var. plicatus, Pseudostichopus, Koehler and Vaney, 1905

papillatus, Peristichopus, Djakonov, 1952

pediculus, Molpadiodemas, sp. nov.

peripatus, Meseres, Sluiter, 1901

porphyrus, Molpadiodemas, sp. nov.

profundi, Pseudostichopus, Djakonov, 1952

propinquus, Pseudostichopus, Fisher, 1907

pustulosus, Pseudostichopus, Sluiter, 1901

spiculiferus, Meseres, O'Loughlin, 2002

tachimaruae, Pseudostichopus ( Trachostichopus ), Imaoka, 1978

torvus, Stichopus ?, Theel, 1886

trachus, Pseudostichopus, Sluiter, 1901

translucens, Molpadiodemas, sp. nov.

tuberculatus, Pseudostichopus ( Trachostichopus ), Imaoka, 1990

tuberosus, Pseudostichopus, sp. nov.

unguiculatus, Pseudostichopus, Ohshima, 1915

ustulatus, Molpadiodemas, sp. nov.

villosus, Pseudostichopus, Theel, 1886

violaceus var, villosus, Pseudostichopus, Theel, 1886

Pseudostichopus aemulatus, Solis-Marin and Billett, 2004

Molpadiodemas atlanticus (Perrier, 1898)

Pseudostichopus mollis Theel, 1886

Pseudostichopus peripatus (Sluiter, 1901)

Pseudostichopus hyalegerus (Sluiter, 1901)

Molpadiodemas atlanticus (Perrier, 1898)

Molpadiodemas constrictus sp. nov.

Molpadiodemas crinitus sp. nov.

Molpadiodemas depressus (Herouard, 1902)

Molpadiodemas involutus (Sluiter, 1901)

Pseudostichopus echinatus Thandar, 1992

Pseudostichopus elegans (Koehler and Vaney, 1905)

Molpadiodemas epibiotus sp. nov.

Molpadiodemas involutus (Sluiter, 1901)

Molpadiodemas helios sp. nov.

Pseudostichopus hyalegerus (Sluiter, 1901)

Pseudostichopus peripatus (Sluiter, 1901)

Molpadiodemas involutus (Sluiter, 1901)

Pseudostichopus hyalegerus (Sluiter, 1901)

Pseudostichopus peripatus (Sluiter, 1901)

Meseres macdonaldi Ludwig, 1 894

Pseudostichopus peripatus (Sluiter, 1901)

Pseudostichopus mollis Theel, 1886

Pseudostichopus hyalegerus (Sluiter, 1901)

Molpadiodemas morbillus sp. nov.

Molpadiodemas neovillosus sp. nov.

Pseudostichopus mollis Theel, 1886

Pseudostichopus occultatus Marenzeller, 1893

Pseudostichopus peripatus (Sluiter, 1901)

Pseudostichopus papillatus (Djakonov, 1952)

Molpadiodemas pediculus sp. nov.

Pseudostichopus peripatus (Sluiter, 1901)

Molpadiodemas porphyrus sp. nov.

Pseudostichopus profundi Djakonov, 1952

Pseudostichopus peripatus (Sluiter, 1901)

Molpadiodemas pustulosus (Sluiter, 1901)

Pseudostichopus spiculiferus (O'Loughlin, 2002)

Pseudostichopus hyalegerus (Sluiter, 1901)

Stichopus ? torvus Theel, 1886

Pseudostichopus mollis Theel, 1886

Molpadiodemas translucens sp. nov.

Pseudostichopus peripatus (Sluiter, 1901)

Pseudostichopus tuberosus sp. nov.

Pseudostichopus peripatus (Sluiter, 1901)

Molpadiodemas ustulatus sp. nov.

Molpadiodemas villosus (Theel, 1886)

Molpadiodemas violaceus (Theel, 1886)

Molpadiodemas Heding, 1935

Table 1, Figure 2

Molpadiodemas Heding, 1935: 77-78. — Heding, 1940: 356-357. —

Deichmann, 1940: 209-211. — O'Loughlin, 1998: 497. — O'Loughlin,

2002: 303, 305, 315.

Platy stichopus Heding, 1940: 358 (new synonym).

Type species. Molpadiodemas acaudum Heding, 1935 (junior

synonym of Pseudostichopus atlanticus Perrier, 1898, by

O'Loughlin, 2002).

Other included species. Molpadiodemas atlanticus (Perrier,

1898); M. constrictus sp. nov.; M. crinitus sp. nov.; M. depres-

sus (Herouard, 1902); M. epibiotus sp. nov.; M. helios sp. nov.;

M. involutus (Sluiter, 1901); M. morbillus sp. nov.; M. neovillo-

sus sp. nov.; M. pediculus sp. nov.; M. porphyrus sp. nov.; M.

pustulosus (Sluiter, 1901); M. translucens sp. nov.; M. ustultus

sp. nov.; M. villosus (Theel, 1886); M. violaceus (Theel, 1886).

Diagnosis. Pygal-furrowed Synallactidae displaying: absence

of prominent appendages (tube feet, papillae) along the paired

152

P. Mark O’Loughlin and Cynthia Ahearn

Figure 1. Characters of pygal-furrowed Synallactidae. a, ventral mouth and anus (M. hellos sp. nov., USNM E31104, paratype, 91 mm long);

b, ventral peltate tentacles, polian vesicle ( P. mollis, TM H2004; anterior end dissected off); c, respiratory tree (P. mollis, TM H2004); d, drawing

of dorsal posterior pygal furrow; e, posterioventral view of pygal furrow and anus (M. violaceus, BMNH 86.10.2.150); f, anterior dorsal

madreporite (arrow) (P. peripatus, USNM 1008159); g, drawing of calcareous ring (radial plate right, as in P. spiculiferus)-, h, calcareous ring

(M. crinitus sp. nov., USNM E48644, paratype); i, calcareous ring (M. violaceus, BMNH 86.10.2.145).

Figure 2. Characters of Molpadiodemas. a, drawing of transverse section of mid-body, hair-like tube foot cover, flat broadly attached longi-

tudinal muscles (as for M. villosus)-, b, flat longitudinal muscles, gonad (upper left), respiratory tree (upper right) (M. porphyrus sp. nov., NMV

F101871, paratype, 90 mm long); c, complete cover of tube feet (M. translucens sp. nov., USNM 1008072, paratype); d, drawing of part of gonad,

branched tubules from common origin; e, gonad, flat longitudinal muscles (arrows) (M. villosus, BMNH 86.10.2.155, paralectotype); f, branched

gonad tubules (M. violaceus, BMNH 86.10.2.145).

A review of pygal-furrowed Synallactidae

153

radii; longitudinal muscles flat, not cylindrical, broadly

attached to the inner body wall; gonad tubules branched, aris-

ing from a common base on the gonoduct, not in series along

the gonoduct; ossicles never present in tube feet; tentacle ossi-

cles frequently rods with ends intertwining and side branches

fused to create mesh.

Distribution. Cosmopolitan; 103-7086 m.

Remarks. Heding (1935) erected the monotypic genus

Molpadiodemas for his new species acaudum, and subsequent-

ly (Heding, 1940) reassigned Pseudostichopus atlanticus

Perrier, P. occultatus Marenzeller and P. villosus Theel to

Molpadiodemas. O'Loughlin (1998) reassigned these four

species to Meseres Ludwig, 1894, Molpadiodemas becoming a

junior synonym of Meseres. Molpadiodemas is raised here out

of synonymy with Meseres (see below under Meseres).

O'Loughlin (2002) considered M. acaudum to be a junior syn-

onym of P. atlanticus, and that decision is confirmed here.

Heding (1940) erected the new genus Platystichopus for

Pseudostichopus depressus Herouard, 1902, which is reas-

signed below to Molpadiodemas. Platystichopus becomes a

junior synonym of Molpadiodemas.

Molpadiodemas atlanticus (Perrier, 1898) comb. nov.

Figures 3 a, b, 4a-d, 6a, b

Pseudostichopus atlanticus Perrier, 1898: 1665. — Perrier, 1902:

333-338, pi. 17 figs 15-20.— Mortensen, 1927: 386-387.—

Deichmann, 1930: 87-88. — Deichmann, 1940: 209, 211. — Heding,

1942: 5.— O’Loughlin, 2002: 315.

Molpadiodemas acaudum Heding, 1935: 78-80, pi. 6 figs 1-2. —

Heding, 1940: 354-357.— Deichmann, 1940: 209, 211.— Heding,

1942: 4-5.

Molpadiodemas atlanticus . — Heding, 1940: 353-359.

Meseres atlanticus . — O’Loughlin, 1998: 497. — Thandar, 1999:

376-379, fig. 4.

Meseres acaudum . — O’Loughlin, 1998: 497, fig. lg-h.

Material examined. Pseudostichopus atlanticus Perrier, 1898.

Holotype. North Atlantic, off the Azores, 42°19'N, 23°31'W, 4060 m.

Talisman stn 134, 1883, MNHN EcHh 2772. Paratype. 42°19'N,

23°36'W, 4060-1010 m, MNHN EcHh 658.

Molpadiodemas acaudum Heding, 1935. Paratype. North Atlantic,

60°17'N, 54°5'W, 3230 m, Ingolf stn 37, 1885, ZMUC.

Other material. Pacific Ocean, New Zealand, Chatham Is,

2610-2668 m, USNM E49319 (1); Chile, 34°07’S (incorrectly

recorded as N in Theel, 1886), 73°56'W (incorrectly as E), 4069 m,

Challenger stn 298, BMNH 1886.10.2.149 (1); 33°31'S, 74°43'W,

3950 m, Challenger stn 299, BMNH 1886.10.2.185 (1); Peru, 3500 m,

USNM 1020067 (1); California, off Point Conception, 4100 m,

1073654 (1); Oregon, 3021 m, USNM E16475 (1); 3700 m, E16488

(3); Clipperton Fracture Zone, 13°08'-13°53'N, 129°51'-129 0 55’W,

4801-4923M, E31100-E31102 (3); E31107, E31108 (2); E31121-

E31123 (3); E31125 (1); E53278 (1); NMV F101845 (1).

Atlantic Ocean, Bahamas, 4828^1873 m, USNM 1021592 (10); NE

of Bahamas, 4930—4940 m, E49466 (9); West European Basin,

4780-4795 m, E38320 (1); Caribbean Sea, Venezuelan Basin,

13°30'-14°50'N, 64°45'-67°30'W, 3411-5062 m, E38789- E38791

(20); E38798 (1); E39266 (20); 1023352 (1); NMVF101846 (2).

Description. Large, up to 210 mm long; body rounded in

section, sometimes sac-like, sometimes with slight brim,

usually no detrital attachments; body wall moderately thick,

soft leathery or firm gelatinous, frequently smooth surface,

frequently wrinkled ventrally; uniform cover of very small tube

feet, frequently not evident or in pits, most evident ventrally

and py gaily, from 0. 1 mm long, 0.2 mm diameter up to 1.0 mm

long, 0.3 mm diameter, cylindrical and firm if extended;

ossicles frequently abundant in tentacles and gonads, some-

times rare or absent; tentacle ossicles frequently irregular thin

rods slightly to strongly curved with ends intertwined to create

small perforations, less frequently irregular straight to curved,

narrow to wide, smooth to laterally spinous, variably perfor-

ated rods, ossicles variable in length, up to 500 pm long; gonad

ossicles irregular rods, frequently with thin lateral branches

joined to create extensive irregular lace-like mesh, non-

spinous, up to 420 pm long.

Colour. Body and tube feet brown to grey-brown to off-white;

frequently some residual mauve markings.

Distribution. North Atlantic, 3230-4060 m (Perrier, 1902, as

P. atlanticus ; Heding, 1935, as M. acaudum)-, south-east

Atlantic, 3155-3255 m (Thandar, 1999); West European Basin,

4780-4795 m (this paper); Venezuelan Basin, 3411-5062 m

(this paper); Pacific Ocean, off Chile, 3950-4069 m (Theel, 1886;

see below); Pacific Ocean, 2610-4923 m (this paper).

Remarks. In agreement with the opinions of Deichmann (1940)

and Heding (1942), M. acaudum was made a junior synonym

of P. atlanticus by O’Loughlin (2002). That decision is

confirmed here.

Theel (1886) noted that two Challenger specimens, taken in

the Pacific Ocean off Chile at stations 298 and 299, closely

resembled Pseudostichopus villosus but lacked tube feet. Both

specimens were examined in this study, and determined as

P. atlanticus. Theel (1886) reported sizes up to 280 mm long.

The larger specimen is now only 150 mm long.

The very small dark brown attachments on the body which

were noted by O’Loughlin (2002) and Theel (1886) are judged

to be epibiotes. O’Loughlin (2002) judged the “mulberry” bod-

ies in the body wall of P. atlanticus, illustrated by Perrier (1902)

and Thandar (1999), and referred to by Theel (1886) as present

in some specimens of P. villosus, to be detrital accretions and

not holothurian ossicles. That opinion is maintained here.

The presence of P. atlanticus in the Pacific Ocean is

confirmed by the numerous USNM specimens determined in

this study. O’Loughlin (2002) summarised P. atlanticus

distribution records as Atlantic Ocean only. This work extends

the distribution to the Pacific Ocean, and to greater depths (at

least 5062 m).

Amongst Molpadiodemas species the distinguishing char-

acters of M. atlanticus are: large, sac-like form; even cover of

small tube feet, frequently inconspicuous; frequent presence

of large, irregular, open mesh-like gonad ossicles.

Molpadiodemas constrictus sp. nov.

Figures 3c-e

Material examined. Holotype. North Atlantic Ocean, Puerto Rico

Trench, 20°08'N, 65°27'W, 7086 m, RV Gilliss, 20 Jan 1973, USNM

1022814.

154

P. Mark O’Loughlin and Cynthia Ahearn

Figure 3. a-b, M. atlanticus (NMV F101845, 210 mm long); a, ventral view; b, small epibiotes on body wall, c-e, M. constrictus sp. nov. (USNM

1022815, paratypes); c, dorsal view (51 mm long); d, ventral view (51 mm long); e, tube feet, f-g, M. crinitus sp. nov. (USNM 1023615, holo-

type, 96 mm long); f, ventral view; g, ventrolateral tube feet, h, M. depressus, pygal furrow, gelatinous body, lateral brim (NMV F101849).

i-k, M. epibiotus sp. nov. (USNM 1008183, holotype, 110 mm long); i, ventral view, posterior taper; j, ventral mouth, epibiote concavities

(arrows); k, ventral view, epibiote concavity (left arrow), few tube feet (central arrow). 1, M. helios sp. nov., tentacle digits, tube feet (lower right)

(USNM E31104, paratype). m-o, M. involutus (NMV F101850, 40 mm long); m, ventral view, serrated margin; n, dorsal anterior view,

globigerines attached; o, posterior ventrolateral tube feet.

A review of pygal-furrowed Synallactidae

155

Figure 4. Tentacle ossicles (SEM). a-d, Meseres atlanticus, a-c (USNM E49466), d (USNM 1020067). e-h, M. crinitus sp. nov. (USNM E48662).

i-1, M. depressus (USNM E53279). m-p, M. epibiotus sp. nov. (USNM 1008310, paratype). q-t, M. helios sp. nov., q-r (USNM E31104,

paratype), s-t (USNM E31117, paratype). u-x, M. involutus (USNM E49256).

156

P. Mark O’Loughlin and Cynthia Ahearn

Figure 5. a-d, Meseres morbillus sp. nov., a, lateral view (USNM 1008093, paratype, 91 mm long); b, tentacles, ventrolateral tube feet “spotting” (NMV

F101865); c, tentacles, digits (USNM E49243); d, calcareous ring (radial plate right half) (USNM 1008093, paratype). e-g, M. neovillosus sp.

nov.; e, dorsal view (USNM 1008458, paratype, 71 mm long); f, pygal end, tube feet (USNM 1008458); g, ventrolateral tube feet (BMNH

86.10.2.151, holotype). h-k, M. pediculus sp. nov.; h, dorsal view, elongate form (USNM 1008318, holotype, 56 mm long); i, ventrolateral tube

feet (NMV FI 04796, paratype); j, tentacles (holotype); k, broadly attached, relatively narrow, slightly thickened, longitudinal muscles

(arrows) (paratype). 1-m, M. porphyrus sp. nov. (USNM E38795, holotype, 114 mm long); 1, dorsal view, thick rounded brim; m, gonad,

n-o, M. translucens sp. nov. (USNM E48652, holotype, 78 mm long); n, dorsal view, longitudinal muscles visible through translucent body wall

(arrows); o, tentacles.

A review of pygal-furrowed Synallactidae

157

Figure 6. Gonad ossicles (SEM). a-b, Meseres atlanticus (USNM E53278). c-d, M. helios sp. nov. (USNM E31106, holotype). e-h, M. neovillosus

sp. nov. (USNM 1008458, paratype). i-1, M. porphyrus sp. nov. (USNM E38795, holotype). m-p, M. ustulatus sp. nov. (USNM 1022493, holotype).

Paratypes. Type locality and date, USNM 1022815 (20); NMV

F101857 (5).

Other material. Type locality and date, USNM 1022816 (90); NMV

F101856 (10).

Description. Up to 62 mm long; body rounded anteriorly,

frequently strongly tapered and narrow posteriorly, dorsoven-

trally depressed, low convex dorsally, flat ventrally fre-

quently with longitudinal median depression, body typically

with one rarely two deep transverse constrictions most

frequently posterior to calcareous ring, body rounded ventro-

laterally sometimes subacute; body wall thick, firm, soft,

semi-gelatinous; even cover of very small tube feet all over

body, some very thin erect, about 0.2 mm long, some lying on

body, cylindrical, thin, typically 0.8 mm long, 0.15 mm

diameter, up to 1.5 mm long, rarely contiguous; longitudinal

muscles flat, broadly attached, narrow, slightly thickened; no

ossicles found.

Colour. Body dark mauve-brown (largest), pale grey-brown to

translucent (smallest), largest sometimes with dark red-brown

markings dorsally, sometimes with even cover of mauve to

brown spots ventrally creating a mottled appearance; largest

tube feet body colour, smallest off-white.

Etymology. From the Latin constrictus (drawn together,

contracted), referring to the characteristic deep transverse

constrictions on specimens.

Distribution. North Atlantic Ocean, Puerto Rico Trench,

7086 m.

Remarks. The transverse constrictions of the body are growth

constrictions. They closely resemble similar growth con-

strictions observed by one of us (M.O’L) in the fissiparous

cucumariid species Squamocnus aureoruber O’Loughlin

and O’Hara, 1992. Emson and Wilkie (1980) listed six fissi-

parous holothurian species, four of them aspidochirotids.

M. constrictus is judged to be fissiparous.

The material was at some stage preserved in formalin solu-

tion, which might account for the complete absence of ossicles.

Amongst Molpadiodemas species the distinguishing characters

of M. constrictus are: body form typically with a transverse

constriction; distinct posterior taper; body colour frequently

with dark markings dorsally and mottled ventrally; minute thin

erect tube feet; absence of ossicles.

Molpadiodemas crinitus sp. nov.

Figures lh, 3f, g, 4e-h, 7h

Pseudostichopus sp. MoV 2033. — O’Loughlin et al., 1994:

253-254.

158

P. Mark O’Loughlin and Cynthia Ahearn

Meseres villosus . — O’Loughlin, 2002: 313, figs 3a, b, tables 1, 2

(non Pseudostichopus villosus Theel, 1886).

Material examined. Holotype. Eastern Antarctica, Ross Sea, 72°57'S,

171°35'E, 573-576 m, USARP, Eltanin stn 1878, USNM 1023615.

Paratypes. Type locality and date, USNM E48644 (8); NMV

F101872 (3).

Other material. Eastern Antarctica, Prydz Bay, 66°46'-68°50'S,

72°14'-77 0 19'E, 333-765 m, NMV F68152 (9); F68158 (1); F68162

(5); F72534 (2); F76583 (2); F76597 (1); F76606 (1); F81816- F81817

(3); Ross Sea, 67°24'-78°23'S, 166°15’E-163°02’W, 223-923 m,

USNM E48578- E48580 (8); E48597 (4); E48621, E48623 (4);

E48642 (6); E48646 (9); NMV F101874 (3); USNM E48662,

E48663 (32); NMV F101873 (4); E49244- E49245 (12); E49270

(1); E49283 (1); E49286 (2); E49299- E49300 (3); E49368 (1);

1005125 (1); 1005129 (2); New Zealand, Antipodes Is, Campbell

Plateau, 2010-2100 m, E48655 (2); W of Antipodes Is, 103 m,

E49357 (4).

Description. Up to 150 mm long; body rounded in transverse

section, lacking brim and distinctive ventrolateral margin,

sometimes transverse grooves, frequently tubercles and pygal

sublobes posteriorly; body wall firm, thick to thin; lacking

significant globigerine or sponge spicule attachments; body

covered with thick, cylindrical tube feet, typically extended,

longest and often matted pygally and in a conspicuous lateral to

ventrolateral band, feet typically 2.0 mm long, 0.4 mm diame-

ter, up to 3.0 mm long, smaller and more spaced dorsally and

ventrally; multiple-branching gonad tubules; ossicles in

tentacles only, abundant to sparse to absent, thick irregular non-

spinous rods, frequently short thick rods with bluntly rounded

ends, some with swelling mid-rod, some as irregular tuberous

lumps, some with short branching, some with blunt spines,

some ends branched, some branches intertwined to create per-

forations, lace-like mesh development on rods rare, rods up to

270 pm long.

Colour. Body and tube feet similar in colour, dark to pale

brown, contracted tube feet rare and sometimes with dark discs.

Etymology. From the Latin crinitus (long-haired, fringed),

referring to the lateral band of long, fringing tube feet.

Distribution. Eastern Antarctica, Prydz Bay, 333-765 m

(O'Loughlin, 2002, as Meseres villosus)-, Ross Sea, 223-923 m;

Antipodes Is, 103-2100 m (this paper).

Remarks. O’Loughlin (2002) considered Antarctic material to

be Pseudostichopus villosus, and although it is similar in form

it is not conspecific with any of the syntypes assigned to

P. villosus by Theel (1886). Amongst Molpadiodemas species

the distinguishing characters of M. crinitus are: wide ventrolat-

eral band of thick, cylindrical, extended, sometimes matted

tube feet, similar in colour to the body wall, typically 2.0 mm,

long, 0.4 mm diameter; tentacle ossicles usually abundant,

frequently short thick rods rounded distally; absence of

gonad ossicles.

Molpadiodemas depressus (Herouard, 1902) comb. nov.

Figures 3h, 4i— 1

Pseudostichopus depressus Herouard, 1902: 15-16, pi. 2 figs 15-18.

Platystichopus depressus . — Heding, 1940: 353-358.

Holotype. North Atlantic Ocean, between Portugal and the Azores,

4360 m, Monaco stn 753, 1896, MOM (cannot be located).

Material examined. North Atlantic Ocean, West European Basin,

4426-4435 m, USNM 1005343 (2); NMV F101849 (1); NE of

Bahamas, 4383-4558 m, USNM 1008242 (1); E53279 (21); NMV

F101868 (4); Sargasso Sea, 5690 m, USNM E49454 (1); N of Puerto

Rico, 5248-5278 m, USNM 1008241 (2).

Provisional (very damaged): Gulf of Mexico, 1353-1399 m, USNM

E48661 (1).

Description. Up to 100 mm long, not tapered; body wall gelat-

inous, thick, firm to soft; sometimes with globigerine attach-

ments; flat ventrally, low convex dorsally, dorso ventrally

depressed; thick rounded ventrolateral brim; mouth anterio-

ventral; dorsally smooth to wrinkled, ventrally wrinkled,

pustulose; small soft tube feet all over body, thick cylindrical

not thread-like, most concentrated and largest orally and pyg-

ally, up to 1.0 mm long, 0.3 mm diameter, inconspicuous and

dome-like over rest of body, 0.2 mm diameter, closer ventrally

than laterally and dorsally; longitudinal muscles flat, slightly

thickened, narrow; ossicles in tentacles, large, thick to thin,

curved to straight rods, smooth or with few blunt to sharp spines,

some with mid-rod swelling, some with branched intertwined

ends creating perforations, some irregularly tuberous and rugose,

some thickly bifurcate at ends, some with one or two large

perforations, no lateral perforations or mesh, up to 390 pm long.

Colour. Body grey to off-white; tube feet off-white.

Distribution. North and South Atlantic Ocean, West Euro-

pean Basin, Sargasso Sea, off Bahamas and West Indies;

1353-5690 m.

Remarks. The holotype of P. depressus cannot be located

(M. Bruni, MOM, pers. comm.), and has been missing for a

long time. Belloc (1950) noted the absence when creating a

catalogue of types, and there is no slide in Cherbonnier’s

comprehensive collection (F. Solis-Marin, pers. comm.).

Based on the description and figures of Herouard (1902), the

characteristics of P. depressus are: up to 74 mm long; off-white

colour; pygal furrow; dorsoventrally depressed; thickened

lateral brim extending around oral end; distinct dorsal

madreporite plate; numerous inconspicuous tube feet all over

body; narrow longitudinal muscles; two tufts of gonad tubules;

lacking ossicles in body wall and gonads.

In this work numerous North Atlantic specimens are judged

to be conspecific. Three were taken from the West European

Basin at 4426-4435 m, a locality close to the type locality for

P. depressus (between Portugal and the Azores, 4360 m). The

conspecific material is characterized by an off-white colour,

close cover of very small tube feet, absence of gonad ossicles,

dorso ventral depression, and frequently a thick brim. No dorsal

madreporite similar to that observed and illustrated by

Herouard (1902) for P. depressus has been seen. But the

external evidence of a dorsal madreporite has been rare in this

work, and not observing it is considered to be not significant

diagnostically. Herouard (1902) illustrated tufts but not series

of gonad tubules (it is not possible to judge branching or not),

and illustrated cylindrical longitudinal muscles. On the basis of

the consistency of the diagnostic characters of Molpadiodemas

A review of pygal-furrowed Synallactidae

159

Figure 7. a-c, Meseres ustulatus sp. nov.; a, ventral view (USNM 1022566, paratype, 55 mm long); b, ventrolateral tube feet (USNM 1022493,

holotype); c, tube feet (NMV F101858, paratype). d-f, M. villosm ; d, ventral view (BMNH 86.10.2.147, lectotype, 90 mm long); e, ventrolateral

tube feet (lectotype); f, posterior tube feet (NMV F98745). g-i, M. violaceus (BMNH 86.10.2.156, holotype, 142 mm long); g, ventrolateral view;

h, tube feet of M. crinitus (left, NMV F101872, paratype) contrasted with M. violaceus (right, NMV F101848); i, tentacles (USNM 1022608).

(above), it is anticipated that the longitudinal muscles were

broadly attached to the body wall, and that the gonad tubules

were branched.

Heding (1940) erected the monotypic genus Platystichopus

for the species depressus, and referred Platystichopus with

Benthothuria Perrier to the Benthostichopodinae (subfamily of

the Gephyrothuriidae). P. depressus has a distinct pygal furrow,

has a close cover of minute tube feet, and lacks body wall

ossicles. It shares more significant diagnostic characters with

Molpadiodemas than Benthothuria, and is referred here to

Molpadiodemas. Platystichopus is a junior synonym of

Molpadiodemas (see above).

On specimens of M. depressus there are ventral minute red

to violet conical attachments similar to those found on

M. atlanticus, and considered here to be epibiotes. Amongst

Molpadiodemas species the distinguishing characters of

M. depressus are: thick gelatinous grey to off-white body wall;

dorsoventrally depressed body form; ventrolateral brim; thick

cylindrical small tube feet; absence of gonad ossicles.

Molpadiodemas epibiotus sp. nov.

Figures 3i-k, 4m-p

Material examined. Holotype. North Atlantic Ocean, Caribbean Sea,

Cuba, Cayman Trench, 19°39’N, 76°17’W, 6195-6320 m, RV Gilliss,

stn GS-121, 21 Jul 1975, USNM 1008183.

Paratypes. Type locality and date, USNM 1008310 (4); NMV

F101862 (1).

Description. Up to 125 mm long; body strongly tapered poster-

iorly, dorsoventrally depressed, low convex dorsally, flat ven-

trally; thin brim anteriorly around mouth, variably developed

laterally, posteriorly; all specimens with anterior large concave

depression suggesting epibiote attachment, some smaller

depressions mid-body; body wall firm, thin, soft, semi-

gelatinous, embedded with white granules, surface wrinkled,

additionally pustulose ventrally; complete cover of small tube

feet, translucent and merged into body wall dorsally; latero-

ventral tube feet in widely spaced band, sparse, subglobular, up

to 0.5 mm long, 0.25 mm diameter, some with dark red mark-

ings; ventral tube feet closer, smaller, some with red markings,

not extended, 0.2 mm diameter as domes; ossicles in tentacles

only, abundant, variable, rods, thick to thin, lumpy to smooth to

bluntly spinous, with or lacking swelling in mid-rod, limited

branching at ends, some branches fused to create perforations,

ossicles up to 300 pm long.

Colour. Dorsally off-white to greyish- white; ventrally reddish-

brown around mouth, dark red flecking on surface and on some

tube feet.

Etymology. From the Greek epi (upon) and bios (life), referring

to the evidence for epibiotic attachments on all specimens.

160

P. Mark O’Loughlin and Cynthia Ahearn

Distribution. North Atlantic Ocean, Caribbean Sea, Cayman

Trench, 6195-6320 m,

Remarks. Amongst Molpadiodemas species, the distinguishing

characters of M. epibiotus are: distinctly posteriorly tapered

body; dorsoventral depression of the body; presence of a thin

brim anteriorly; presence of anterior concavities from

epibiotes; fine dark red markings ventrally on the body surface

and tube feet.

Molpadiodemas helios sp. nov.

Figures la, 31, 4q-t, 6c, d

Material examined. Holotype. Central Pacific Ocean, Clipperton

Fracture Zone, 13°13'N, 129°53'W, 4877 m, RY Governor Ray, Ocean

Minerals Co., 15 Feb 1980, USNM E31106.

Paratypes. Type locality, 13°13’N, 129°52-53’W, 4724-4892 m,

1980, USNM E31104 (1), E31117 (1), E31134 (1), E31137 (1),

E31141 (1), NMV F101852 (1), F101854 (1).

Other material. Type locality, 13°13'N, 129°52-55'W, 4755-4938

m, 1980, USNM E31097 (1), E31103 (1), E31105 (1), E31110-

E31114 (5), E31118 (1), E31129 (2), E31133 (1), E31138-E31140 (3),

E31144 (1), E48515 (1), NMV F101853 (1).

Description. Up to 170 mm long; body wall soft-leathery to

semi-gelatinous; no body attachments (encrustations); body

rounded in transverse section, lacking brim; tentacles with

about 10 large conical pointed digits peripherally, some digits

bifid; tube feet soft, white to semi-translucent, scattered and

lying all over body, most dense around oral and pygal areas, up

to 2.0 mm long, 0.15 mm diameter, frequently cylindrical

digitate erect orally and pygally, typically 0.8 mm long, 0.2 mm

diameter; ossicles in tentacles and gonad; tentacle ossicles

sparse to rare, thick to thin rods up to 150 pm long, rare

branches, rare perforations at ends or branches joined to form

perforations, lacking spines, sometimes swollen mid-rod;

gonad tubules with abundant irregular thin rods, some

irregularly branched, branches sometimes joined to create

perforations, lacking spines, swellings rare, up to 280 pm long.

Colour. Body off-white to grey, frequently semi-translucent,

smaller specimens sometimes with pale brown.

Etymology. From the Greek helios (sun), referring to the sun-

flower appearance of the radiating tentacle digits.

Distribution. Central Pacific Ocean, Clipperton Fracture Zone,

13°13'N, 129°53'W, 4724-4938 m.

Remarks. Amongst Molpadiodemas species, the distinguishing

characters of M. helios are: relatively prominent soft tube feet

scattered over the body, sometimes cylindrical and erect

posteriorly and anteriorly; prominent conical tentacle digits;

predominantly smooth rod ossicles in gonad.

Molpadiodemas involutus (Sluiter, 1901) comb. nov.

Figures 3m-o, 4u-x

Meseres involutus Sluiter, 1901a: 11-12. — Sluiter, 1901b: 49-50,

pi. 8 fig. 6. — Perrier, 1902: 359. ( Meseres convolutus) [lapsus]. —

O'Loughlin, 2002: 306, fig. 2e, tables 1, 3, 4.

Pseudostichopus globigerinae Herouard, 1923: 23-25, pi. 4 fig.

6. — Mortensen, 1927: 386-388. — Deichmann, 1930: 87, 90.

Pseudostichopus ( Pseudostichopus ) globigerinae. — Heding, 1940:

353-360.— Imaoka, 1978: table 1-1. — Thandar, 1992: 167.

Pseudostichopus villosus . — Hansen, 1956: 47-48 (part; non

Pseudostichopus villosus Theel, 1886).

Pseudostichopus ( Pseudostichopus ) dilatorbis Imaoka, 1978:

378-380, 384, fig. 1 B-E, table 1-1 (new synonym).

Meseres globigerinae. — O’Loughlin, 2002: 305, tables 1, 3 (new

synonym).

Material examined. Meseres involutus Sluiter. Lectotype (designated

here). Indonesia, Sawu Sea, 10°49'S, 123°23'E, 918 m, Siboga stn 300,

ZMA V.Ech.H1052. Paralectotype. Seram Sea, 3°27'S, 131°01'E, 567

m, Siboga stn 173, ZMA V.Ech.H 1051 (1).

Pseudostichopus globigerinae Herouard, 1923. Lectotype (larger

syntype, with pygal furrow, designated here). North Atlantic, off

Newfoundland, 46°18'N, 50°42'W, 4380 m, Monaco stn 2964, 20 Jul

1910, MOM (examination via pers. comm, by M. Bruni at MOM).

Second syntype, type locality and date, MOM.

Pseudostichopus ( Pseudostichopus ) dilatorbis Imaoka, 1978.

Holotype. Japan, W of Kyushu, near Shimo-Koshiki I., 400-450 m.,

SMBL 309.

Other material. Pacific Ocean, eastern Australia, Tasman Sea, Lord

Howe Rise, 1423 m, AM J23326 (1); off Newcastle, 2984-3058 m,

J16833 (1); off Nowra, 1650-1750 m, NMV F80451 (1); Japan, East

China Sea, off Kagoshima, 420 m, USNM 1071585 (3), NMV

FI 04793 (2); 300-330 m, USNM 1071802 (1); Galapagos, 3667 m,

USNM E950 (1).

Atlantic Ocean, West European Basin, 4426-4435 m, USNM

1005340 (8); NMV F101850 (2); off South Carolina, 2100 m, USNM

E41392 (3); NMV F101851 (1); Gulf of Mexico, 1668 m, USNM

E46796 (3); off Louisiana, 2063-2085 m, USNM 1008165 (2); NMV

F101869 (1); 1829 m, USNM E46798 (2); 1646 m, E46799 (1).

South Atlantic Ocean, Argentina, Buenos Aires, SE of Mar Del

Plata, 5208-5223 m, USNM 1005317 (1); 1008207 (1); off Falkland

Is, 5687-5801 m, E49256 (5); E49359 (1); Scotia Sea, 2250-2404 m,

E49295 (2).

Description. Up to 155 mm long; typically encrusted with

globigerine or small stone attachments; body usually dorso-

ventrally depressed; flat ventrally, low convex dorsally; margin

subacute, partly serrated because of transverse creasing of body

wall, sometimes narrow rounded brim; body wall thin to thick,

firm, wrinkled, with small digitate projections on low reticulate

ridges; raised protuberances ventrolaterally, pygally; close

cover of thin cylindrical tube feet, sometimes larger and more

evident and reddish-brown in ventrolateral band, sometimes

matted, typically 1.0 mm long, 0.2 mm diameter, up to 3.0 mm

long, 0.5 mm diameter; multiple branching gonad tubules;

ossicles in tentacles only, sometimes not detected; ossicles

unbranched rods with thick central swelling and branched rods,

branches frequently intertwining at ends or joining laterally to

create irregular perforated mesh, ossicles up to 330 pm long.

Colour. Body grey to off-white; tube feet sometimes reddish-

brown ventrolaterally.

Distribution. Indonesia, Seram and Sawu Seas, 567-918 m

(Sluiter, 1901a); eastern Australia, Tasman Sea, 1423-3058 m

(O’Loughlin, 2002); Japan, W of Kyushu, 400-450 m (Imaoka,

1978; as P. dilatorbis)-. East China Sea, off Kagoshima,

300-420 m (this paper); Galapagos Islands, 3667 m (this

paper); northern Atlantic Ocean, off Newfoundland, 4380 m

(Herouard, 1923, as P. globigerinae ); West European Basin,

A review of pygal-furrowed Synallactidae

161

4426-4435 m; western Atlantic Ocean, off South Carolina,

Gulf of Mexico, 1646-2100 m; southern Atlantic Ocean, off

Argentina, 5208-5223 m; Falkland Is, 5687-5801 m; Scotia

Sea, 2250-2404 m (this paper).

Remarks. Sluiter (1901a) described two syntypes, and both

were examined during a visit to the University of Amsterdam

(Apr 2002). One is designated above as the lectotype.

Hansen (1956) considered P. globigerinae to be a junior

synonym of P. villosus. O'Loughlin (2002) rejected the

synonymy on the evidence that P. villosus never has tufts of

tube feet on tubercles along the posterior margin, or a globi-

gerine cover, or translucent body wall. A further synonymy was

not suggested. Based on the description by Herouard (1923), on

observations communicated by Michele Bruni (MOM), and on

photographs by Francisco Solis-Marin (UNAM) of a tentacle

ossicles slide prepared by Gustav Cherbonnier (MNHN box

108 slide 44), the characters of the larger syntype are: up to 30

mm long; body completely covered by globigerines; body wall

thin, soft, translucent; pygal furrow present; villous-like cover

of unequal and unevenly distributed small tube feet, rare mid-

ventrally, in tufts on posterior ventrolateral tubercles creating

serrated appearance; flat longitudinal muscles; lacking gonad;

and tentacle ossicles branched closed mesh fragments.

O’Loughlin (2002) noted that there was inadequate descriptive

information to confirm any synonymy, but in the light of

further data P. globigerinae is considered here to be a junior

synonym of M. involutus.

Herouard noted that there was no pygal furrow on the 15

mm long smaller syntype of P. globigerinae , and M. Bruni

observed a cylindrical longitudinal muscle. The smaller of the

two syntypes is not conspecific with P. globigerinae, but

cannot be identified. The larger syntype is designated above as

lectotype.

The description of P. dilatorbis by Imaoka (1978) refers to

the external characters of the large paratypes and the internal

characters of the holotype. The holotype has the diagnostic

characters of Molpadiodemas', the paratypes those of

Pseudostichopus. Although the holotype lacks tentacles

ossicles, several specimens of Pseudostichopus dilatorbis

recently donated by Tohru Imaoka yielded abundant tentacle

ossicles and possess external characters similar to those of

M. involutus. Pseudostichopus dilatorbis is considered here to

be a junior synonym of Molpadiodemas involutus. The two

paratypes of P. dilatorbis are much larger (up to 180 mm long)

than the holotype, and are very close in appearance and diag-

nostic characters to the types of Pseudostichopus trachus

Sluiter, 1901, which is considered below to be a junior

synonym of Pseudostichopus mollis. The paratypes are referred

below to P. mollis.

Material determined here extends the distribution of

M. involutus from the western Pacific Ocean into the eastern

Pacific Ocean, northern and southern Atlantic Ocean and Scotia

Sea.

Amongst Molpadiodemas species, the distinguishing char-

acters of M. involutus are: body wall with small digitate pro-

jections on low reticulate ridges; margin partly serrated because

of transverse creasing of body wall; cylindrical tube feet most

evident in ventrolateral band; tentacle ossicles rods and mesh

on primary rods; typical dense cover of globigerines.

Molpadiodemas morbillus sp. nov.

Figures 5a-d, 8a-d

Material examined. Holotype. Antarctic Ocean, South Shetland Is,

Bransfield Strait, Livingston I, 62°38'S, 59°37'W, 681-1409 m, RV

Eltanin, USARP Cr 6 stn 430, 7 Jan 1963, USNM E48647.

Paratypes. Type locality and date, USNM 1008093 (31); NMV

F101866 (5).

Other material. Scotia Sea, 55 o 01'-59°37'S, 26°00’-45°05'W,

1071-3040 m, USNM E48601 (4); E48653 (3); E49243 (10); NMV

F101865 (3); USNM E49251 (1); E49253 (12); E49254 (49); NMV

F101867 (5); USNM 1008094-1008096 (7); 1022457 (9); 1022461

(1); South Sandwich Trench, 5350 m, USNM 1071583 (8); South

Shetland Is, 884-935 m, USNM 1005127 (2); 2119-2562 m, E48569

(1); W of Elephant I., 2672-3020 m, E48625 (1); N of Amundsen Sea,

4682 m, USNM 1022460 (1); 4866-4881 m, E48649 (2); 4709 m,

E48650 (1); NW of Amundsen Sea, 4575^1813 m, E48631 (1).

Description. Up to 143 mm long; body tapered anteriorly and

posteriorly, frequently rounded in transverse section, some-

times with transverse folds and ridges; ventrolateral margin

commonly rounded, sometimes with distinct serrated

margin created by transverse folds; body wall firm, soft, thin

(small to medium specimens) to thick (largest specimens),

surface wrinkled and pustulose; some specimens with grit or

globigerine attachments; complete cover of scattered small tube

feet, predominantly not extended, frequently evident as a broad

ventrolateral band, frequently extended and close pygally,

frequently lost with loss of outer body wall, cylindrical not

thread-like when extended, not villous or matted, sometimes

contiguous pygally, never ventrolaterally, typically 0.5 mm

long, 0.2 mm diameter pygally and when extended ventro-

laterally, 0.2 mm diameter when contracted; ossicles in tenta-

cles only, abundant to sparse to absent, irregular rods, thick to

thin to wide, some perforated, frequently narrow rods with

large central knob and pointed ends, some long thin rods

with closely intertwined branches at ends, some with short

branches, some with blunt spines, up to 300 pm long.

Colour. Body reddish-brown to brown, smallest grey, largest

off-white; tube feet typically dark reddish-brown to chocolate,

creating a spotted appearance.

Etymology. From the Latin morbillus (measles), referring to the

typically spotted body appearance created by the dark-coloured

tube feet.

Distribution. Amundsen Sea, Scotia Sea, 681-4881 m.

Remarks. One specimen was covered by a rhizomatous epibiote

with vertical tongue-like stems, that could be mistaken as a

character of the species. M. morbillus is similar to M. crinitus

(above), from which it is distinguished by: smaller, usually con-

tracted, tube feet (thick and extended in M. crinitus, typically 2.0

mm long, 0.3 mm diameter, up to 3.0 mm long, 0.5 mm diam-

eter); tube feet which are scattered and never contiguous ven-

trolaterally (often matted ventrolaterally in M. crinitus)', and

tube feet which are darker in colour than the body wall (same

colour as body wall in M. crinitus).

162

P. Mark O’Loughlin and Cynthia Ahearn

Table 3. Revised systematic status of the ten syntypes of Pseudostichopus villosus Theel, 1886 and variety P. villosus var. violaceus Theel, 1886.

Challenger

BMNH Registration

Type status

Revised systematic status

stn 244

86.10.2.147

Lectotype

Molpadiodemas villosus (Theel, 1886)

stn 157

86.10.2.155

Paralectotype

Molpadiodemas villosus (Theel, 1886)

stn 156

86.10.2.156

Holotype

Molpadiodemas violaceus (Theel, 1886)

stn 61

86.10.2.145 (2)

Syntypes

Molpadiodemas violaceus (Theel, 1886)

stn 147

86.10.2.154

Syntype

Molpadiodemas violaceus (Theel, 1886)

stn 325

86.10.2.150

Syntype

Molpadiodemas violaceus (Theel, 1886)

stn 216

86.10.2.151

Holotype

Molpadiodemas neovillosus sp. nov.

stn 302

86.10.2.153

Syntype

Pseudostichopus ? peripatus (Sluiter, 1901)

Unconfirmed (small specimens):

stn 146

86.10.2.148

Paralectotype

Molpadiodemas villosus (Theel, 1886)

stn 156

86.10.2.146

Paralectotype

Molpadiodemas villosus (Theel, 1886)

stn 296

86.10.2.152

Paralectotype

Molpadiodemas villosus (Theel, 1886)

Amongst Molpadiodemas species, the distinguishing char-

acters are: conspicuous ventrolateral band of small dark

cylindrical tube feet which create a spotted appearance;

tentacle ossicles irregularly present, frequently narrow rods,

tapered at ends, with central knob; absence of gonad ossicles.

Molpadiodemas neovillosus sp. nov.

Table 3, Figures 5e-g, 6e-h, 8e, f

Pseudostichopus villosus Theel, 1886: 170-171 (part; syntype from

Challenger stn 216 only; non Pseudostichopus villosus Theel, 1886,

lectotype designated below).

Material examined. Holotype. Syntype, P. villosus Theel, 1886.

Northern Pacific Ocean, Caroline Is, 2°56'N, 134°H'E, 3658 m,

Challenger stn 216, 16 Feb 1875, BMNH 86.10.2.151.

Paratypes. Galapagos Is, 2°34'N, 92°6’W, 2487 m, USNM 18275

(1); NMV F101847 (1); 4°33'S, 87°42’W, 3667 m, USNM 1008458 (1).

Other material. North Pacific Ocean, Mexico, S of Punta

Maldonado, 3436 m, USNM 18276 (1).

Description. Up to 70 mm long; body rounded in transverse

section, narrower anteriorly and posteriorly, lacking tubercles

on pygal lobes, lacking brim; body wall thin, smooth, thicker

and wrinkled if contracted, lacking globigerine or sponge

spicule attachments; tube feet in conspicuous ventrolateral

wide band, continuous anteriorly and pygally, feet cylindrical,

frequently overlapping and matted, typically 1.6 mm long, 0.3

mm diameter; tube feet smaller, more spread, dorsally, smallest

ventrally; broad, flat longitudinal muscles about 4.0 mm wide;

multiple branching gonad tubules; ossicles in tentacles and

gonads; tentacle ossicles abundant rods, variable form, pre-

dominantly thin, curved, smooth to rugose, not spinous, blunt

to tapered, frequently intertwined at ends, rarely branched or

with slight mid-rod swelling, some with lateral fused branches

creating narrow mesh, rods up to 320 pm long; gonad ossicles

abundant rods, predominantly thin, long, rarely branched,

straight to slightly bent to irregular, rarely with slight swelling

mid-rod, up to 350 pm long.

Colour. Body and tube feet pale brown to off-white.

Etymology. From the Latin neo (new, recent), referring to this

recently recognized new species with villous ventrolateral tube

feet.

Distribution. Central Pacific Ocean, Caroline and Galapagos Is,

Mexico, 2487-3667 m.

Remarks. Theel (1886) commented specificially on this speci-

men, noting that “the individual obtained at Station 216 is

remarkable in that the pedicels of the dorsal surface and the

sides of the body are slightly thicker and larger than those of the

ventral surface which are thread-like and very minute”. Further

to this comment is the observation that the tube feet are largest

and most dense and frequently matted in a wide ventrolateral

band which extends around the anterior and posterior ends of

the body.

Amongst Molpadiodemas species, the distinguishing char-

acters are: ventrolateral band of villous cylindrical tube feet

that are distinctly smaller than those of the similar M. crinitus

(above); tentacle ossicles predominantly thin curved rods with

intertwined ends, some with slight swelling centrally; gonad

ossicles abundant thin, short to very long, rods.

Molpadiodemas pediculus sp. nov.

Figures 5h-k, 8g-h

Material examined. Holotype. South Atlantic Ocean, NE of South

Sandwich Is, 55°07'S, 25°59'W, 5435-5453 m, RV Eltanin, Cr 8 stn

591, 29 Apr 1963, USNM 1008318.

Paratypes. Type locality and date, NMV F104796 (1).

Other material. Scotia Sea, N of South Orkney I„ 2800 m, USNM

E48635 (1).

Description. Up to 60 mm long; body small, rounded in

section, elongate, tapered anteriorly and posteriorly, lacking

brim; body wall firm, semi-gelatinous, surface smooth, not

covered with attachments; tube feet all over body, overlapping

ventrolaterally, distinctly cylindrical, frequently erect, close

cover, anteriorly and pygally, typically 1.0 mm long, 0.25 mm

diameter, up to 1.5 mm long; tube feet typically shorter, more

thread-like, dorsally and ventrally than ventrolaterally; longi-

tudinal muscles broadly attached, relatively narrow, slightly

thickened; some unbranched gonad tubules; ossicles in ten-

tacles only, predominantly irregular, thick, rough to smooth,

sometimes bluntly spinous rods, swollen mid-rod, tapering at

ends, rare branches joined to create thick intertwining mesh,

sometimes irregularly tuberous, up to 280 pm long.

A review of pygal-furrowed Synallactidae

163

Colour. Body, tube feet off-white to pale brown.

Etymology. From the Latin pediculus (little foot), referring to

the prominent ventrolateral band of small tube feet.

Distribution. South Atlantic Ocean, off South Sandwich and

South Orkney Is, 2800-5453 m.

Remarks. Amongst Molpadiodemas species, the distinguish-

ing characters are: body tapering orally and anally; close

cylindrical tube feet overlapping ventrolaterally; longi-

tudinal muscles broadly attached, relatively narrow, slightly

thickened.

Molpadiodemas porphyrus sp. nov.

Figures 2b, 51, m, 61 — 1, 8i-l

Material examined. Holotype. North-west Atlantic Ocean, Carribean

Sea, Venezuelan Basin, 13°30'N, 64°45’W, 3459-3503 m, Norda Thru

IRCZM, RV Bartlett 1301-82, 27 Nov 1981, USNM E38795.

Paratypes. Type locality, date, depth, USNM 1008206 (1); NMV

F101870 (1); type locality, 3428-3476 m, USNM E38792 (1); NMV

F101871 (1); type locality, 3422-3464 m, USNM E38797 (2).

Other material. Type locality, date, 3476-3518 m, USNM E38793

(2); 3967-4009 m, USNM E38799 (1); Caribbean Sea, off Cuba, 2997

m, E2589 (1); South-west Atlantic Ocean, off Brazil, 1227 m, E2584

(14).

Description. Up to 115 mm long; body wall thick, firmly gelat-

inous; some globigerines attached, no overall cover; to varying

degrees flattened dorsoventrally, distinct rounded brim lateral-

ly, sometimes with slightly pustulose bulges; body covered

with small soft flaccid thread-like tube feet, usually spread,

sometimes matted in smaller specimens, readily lost with loss

of outer body layer, typically 0.7 mm long, 0.15 mm diameter,

pygally up to 1.0 mm long, 0.2 mm diameter; ossicles in tenta-

cles and gonad; tentacle ossicles numerous, variable, curved

thin spinous rods (predominant in small specimens), closely

finely spinous and smooth unbranched thick to broad plate-like

rods, sometimes perforated, sometimes branched and inter-

twined at ends, spinous or smooth mesh, up to 270 pm long;

gonad ossicles abundant to sparse, small thin rods, bluntly

spinous or smooth or knobbed, branched or unbranched, some

branching creating narrow mesh, up to 140 pm long.

Colour. Very soft outer body layer brown, thick semi-gelati-

nous layer grey with distinct overall violet mauve hue, some

sparse violet mauve colour spots; tube feet off-white.

Etymology. From the Greek porphyro (purple), referring to the

violet mauve colour hue and colour spots.

Distribution. North-west and South-west Atlantic Ocean,

Caribbean Sea, and off Brazil; 1227-4009 m.

Remarks. Amongst Molpadiodemas species, the distinguishing

characters are: firm thick gelatinous body wall; lateral brim;

grey-mauve colouration; thread-like flaccid tube feet; tentacle

ossicles frequently closely bluntly spinous.

Molpadiodemas pustulosus (Sluiter, 1901) comb. nov.

Pseudostichopus pustulosus Sluiter, 1901a: 16-17. — Sluiter, 1901b:

53-55, pi. 4 fig. 6, pi. 9 fig. 1.

Material examined. Lectotype (designated here). Indonesia,

Halmahera Sea, 0°45'S, 128°40'E, 827 m, Siboga, stn 145, ZMA

V.Ech.H1012. Paralectotypes. Lectotype locality and date, ZMA

V.Ech.HlOll (1); Ceram Sea, 3°38’S, 131°26'E, 924 m, Siboga stn

170, ZMA V.Ech.H1006 (1).

Description. Up to 115 mm long; some globigerines attached;

distinct ventrolateral margin with series of nipple-like pustules,

most conspicuous posteriorly and around anus on lobes of

pygal furrow, some pustules lobed; body covered with elongate

tube feet, present but not extended mid-ventrally; tube feet

clustered in tufts on posterior pustules; gonad tubules short, fat,

multiple-branching; ossicles in tentacles only; tentacle ossicles

non-spinous lace mesh, comprising thin branched joined rods,

not perforated plates.

Distribution. Indonesia, Halmahera and Ceram Seas, 827-924 m.

Remarks. During an examination (Apr 2002) of the holothurian

types in the University of Amsterdam, four specimens were

found to be registered as syntypes of R pustulosus. This was in

accord with the catalogue published by Jangoux (1991). But

only three syntypes were recorded by Sluiter (1901a). One of

the four, registered and recorded by Jangoux (1991) as ZMA

1303 from Station 170, was not conspecific with the other syn-

types. Sluiter (1901) reported, and Jangoux (1991) listed, two

syntypes of P. trachus, but the syntype registered as ZMA

E2496/2 was missing from the collection. The extraneous syn-

type of P. pustulosus (ZMA 1303) was conspecific with the

syntype of P. trachus, and is now registered as a P. trachus type

(ZMA V.Ech.H2496/2, following the catalogue registration

number of Jangoux, 1991). A lectotype of P. pustulosus is

designated above.

The designated lectotype is the specimen figured by Sluiter

(1901b), and does not show all of the characters referred to here

in the description. The syntype showing the diagnostic charac-

ters most completely, such as the tube feet, is the smallest

(ZMA V.Ech.H1006; 63 mm long).

Rowe (1995) judged that Pseudostichopus propinquus

Fisher, 1907 and Pseudostichopus nudus Ohshima, 1915 were

junior synonyms of P. pustulosus. M. pustulosus has the diag-

nostic characters of Molpadiodemas (above), not Pseudo-

stichopus (below). Both synony mi es are rejected on the grounds

of the relevant generic diagnostic characters. P. propinquus and

P nudus are discussed below. Rowe (1995) identified material

from off Newcastle on the eastern Australian slope as Pseudo-

stichopus pustulosus. All Australian slope material has been

examined and none determined as P. pustulosus. Material from

off Newcastle (AM J16749) was determined as P. mollis (below).

Amongst Molpadiodemas species, the distinguishing char-

acters of M. pustulosus are: conspicuous nipple-like pustules on

the ventrolateral margin; tufts of tube feet on the pustules pos-

teriorly; lace-mesh tentacle ossicles.

Molpadiodemas translucens sp. nov.

Figures 2c, 5n, o, 8m-p, 12e, f

Material examined. Holotype. Weddell Sea, Antarctic Peninsula,

64°07'S, 40°48W, 4465^1557 m, RV Eltanin, USARP Cr 12 stn 1018,

21 Mar 1964, USNM E48652.

164

P. Mark O’Loughlin and Cynthia Ahearn

Paratypes. Type locality and date, USNM 1008072 (5); NMY

F101863 (2).

Other material. South-east Pacific Basin, 4575-4813 m, E49487 (2).

Description. Up to 79 mm long; body wall thin, soft, trans-

lucent; no body attachments; body typically rounded in

transverse section, not elongate, tapered anteriorly and poste-

riorly, lateroventral margin sometimes subacute or slightly

bulbous; complete body cover of small, cylindrical, soft, scat-

tered tube feet, erect or lying on body, typically 0.3 mm long,

0.1 mm diameter, rarely contiguous over most of body, slightly

more concentrated ventrolaterally, most concentrated and

frequently contiguous pygally, typically 0.6 mm long, 0.2 mm

diameter; longitudinal muscles flat, slightly narrow; ossicles in

tentacles and gonads; tentacle ossicles abundant, variable in

form, thin to thick tapered smooth distally bluntly pointed rods

sometimes with central rounded thickening, rods with inter-

twined branched ends creating small perforations, rods with

fused side branches creating small lateral perforations, fused

branches sometimes creating large perforated irregular mesh,

ossicles up to at least 520 pm long; gonad ossicles abundant,

predominantly thin lace mesh with irregular large perforations,

not developed on primary thick rod, some irregular finely

spinous thin rods with irregular branching closing to create per-

forations, ossicles up to 320 pm long.

Colour. Pale greyish brown, with traces of mauve colouration;

tube feet off-white to semi-translucent.

Etymology. From the Latin lux (light) and trans (through),

referring to the translucent body wall.

Distribution. South-east Pacific Basin, Weddell Sea, Antarctic

Peninsula, 4465-4813 m.

Remarks. Amongst Molpadiodemas species, the distinguish-

ing characters are: translucent body wall; lace mesh gonad

ossicles.

Molpadiodemas ustulatus sp. nov.

Figures 6m-p, 7a-c, 12q-t

Material examined. Holotype. South-east Pacific Ocean, Peru, Peru-

Chile Trench, 8°48'S, 80°40'W, 5069-5173 m, RV Anton Bruun , Cr 11

stn 178, 3 Nov 1965, USNM 1022493.

Paratypes. Type locality and date, USNM 1022566 (3); NMV

F101858 (1).

Description. Up to 73 mm long; form cylindrical, narrowly

elongate, slightly dorsoventrally depressed (possible artefact),

lacking brim; outer body wall soft, pustulose, wrinkled; inner

body wall soft, thin, semi-gelatinous; close cover of small

cylindrical tube feet all over body, extended but not erect,

longest and most evident in lateral band, not matted, up to 1.0

mm long, typically 0.6 mm long, 0.2 mm diameter; ossicles in

tentacles and gonad; tentacle ossicles abundant smooth rods,

thick to thin, straight to curved, some with central swellings,

some with short branches, some with intertwined joined ends

creating large and small perforations, some irregularly tuber-

ous, up to at least 270 pm long; gonad ossicles numerous thick

to thin smooth rods, some lateral branches fused to form mesh

with large perforations, fragments up to 280 pm long.

Colour. Body and tube feet very dark brown.

Etymology. From the Latin ustulatus (scorched), referring to

the dark brown blistered appearance of the specimens.

Distribution. South-east Pacific Ocean, Pern-Chile Trench,

5069-5173 m.

Remarks. The specimens are in poor condition, but adequate for

establishing morphological diagnostic characters. Amongst

Molpadiodemas species the distinguishing characters of

M. ustulatus are: very dark brown colour; narrowly elongate

body; tube feet most evident in lateral band; gonad rod ossicles

with open lateral mesh.

Molpadiodemas villosus (Theel, 1886) comb. nov.

Table 3, Figures 2a, e, 7d-f, 8q-t, 12a-d

Pseudostichopus villosus Theel, 1886: 170-171 (part; syntypes

from Challenger stns 157, 244 only; not Pseudostichopus villosus var.

violaceus Theel, 1886).

Material examined. Lectotype (designated here). North-west Pacific

Ocean, off Japan, 35°22’N, 169°53'E, 5304 m. Challenger stn 244,

BMNH 86.10.2.147.

Other material. Paralectotype, Southern Indian Ocean, 53°55'S,

108°35E, 3566 m, Challenger stn 157, BMNH 86.10.2.155 (1).

Northern Pacific Ocean, Clarion-Clipperton Fracture Zone, no

depth, USNM E48492 (1); Galapagos Is, 3667 m, E9929 (1); South-

east Pacific Basin, N of Amundsen Sea, 5042-5045 m, USNM 48659

(3); 4709 m, E49280 (1); 4676 m, E48584 (1); 4740-4742 m, E49333

(5); N of Marie Byrd Land, 4682 m, E48658 (1); Ross Sea, 3495-3514

m, E48583 (4); 2005-2010 m, E48620 (7); NMVF101859 (2).

North Atlantic Ocean, North American Basin, 3264-3356 m, USNM

1006292 (1).

Scotia Sea, S of South Georgia I., 3413-3446 m, USNM E48634

(1); E48633 (8); NMV F98745 (2); 2742-2758 m, USNM E48640 (2);

NMV F101860 (2); 2384-2416 m, USNM E49257 (1, with commen-

sal gastropods); South Georgia I., 3166-3255 m, USNM E48609 (1);

South Shetland Is, 681-1409 m, E48571 (2); N of Bellingshausen Sea,

4941 m, E48589 (1).

Description. Up to 110 mm long; body variably elongate and

tapered anteriorly and posteriorly; subcylindrical, lacking brim,

sometimes sac-like; lacking globigerine or sponge spicule or

grit attachments; body wall thick semi-gelatinous to thin firm,

surface smooth to wrinkled and pustulose; tube feet small, soft,

hair-like, never erect, close to sparse cover over whole body,

dorsally typically 1.5 to 2.0 mm long, up to 3.0 mm long, 0.05

mm diameter; tube feet sometimes matted ventrolaterally, ante-

riorly and pygally, typically 1.0 to 1.3 mm long, 0.2 mm dia-

meter; ventral tube feet frequently withdrawn, if extended

typically 0.5 mm long, 0.2 mm diameter; gonad tubules

unbranched and multibranched; ossicles in tentacles and gonad;

tentacle ossicles abundant, variable form, large, up to 530 pm

long, frequently rods perforated entire length, rods variably

with mid-rod swelling, thick rods with branches at ends inter-

twined to create mesh, thick rods with lateral joined branches

creating mesh, large mesh ossicles (intertwined, joined rods,

not perforated plates), club-shaped rods with minute perfora-

tions, smooth thick to thin rods tapered at ends, sometimes

blunt spines, some knots of thick irregularly branching rods;

gonad ossicles abundant, long thin branched and unbranched

A review of pygal-furrowed Synallactidae

165

irregular rods, many with lateral mesh with large perforations,

many thick smooth rods with central knobbed swelling, many

central knobs elongating into short thick branches which fuse to

create perforations, up to 270 pm long.

Colour. Greyish-white, sometimes with reddish-brown hue;

tube feet off-white, sometimes pale brown ventrolaterally.

Distribution. Atlantic, Indian, Pacific and Southern Oceans,

Ross Sea, N of Amundsen Sea, N of Bellingshausen Sea, Scotia

Sea; 681-5304 m.

Remarks. Theel (1886) listed ten syntypes of P. villosus

(nominal subspecies villosus ), plus a type of his variety

P. villosus var. violaceus. All have been examined in this study.

Theel noted the “villous” appearance of the specimen from stn

244 and it is this one we have chosen as lectotype. One other,

from stn 157, is a paralectotype. Of the others, the one from stn

216 is not conspecific and is described above as the new

species, M. neovillosus.

Four of the remaining types are also not P. villosus. The type

of the variety and three of the syntypes are referred below to the

species Molpadiodemas violaceus (Theel, 1886). The seventh

syntype (Southern Pacific Ocean, off Chile, 42°43'S, 82 0 ll'W,

2652 m, Challenger stn 302, BMNH 86.10.2.153) is not

P. villosus. It was 80 mm long, and had a close cover of glo-

bigerines, strong transverse ridges ventrally, an acute serrated

ventrolateral margin, and cylindrical longitudinal muscles. No

gonad was present. It is probably Pseudostichopus peripatus

(Sluiter), but there is insufficient evidence to confirm a deter-

mination. The three remaining syntypes are small, and remain

unconfirmed paralectotype s: stn 146, BMNH 86.10.2.148

(1); stn 156 (of two lots from this Station the lot with the

P. villosus syntype), BMNH 86.10.2.146 (1); stn 296, BMNH

86.10.1.152 (1).

Dr Frank Rowe (pers. comm.) observed eggs or embryos

amongst the tentacles of the paralectotype (stn 157), and con-

sidered the possibility of brood-protection. The observation

was confirmed during this study.

The distribution of P. villosus given by O’Loughlin (2002)

is not valid, since only one of the syntypes is conspecific with

the designated lectotype, and most previous determinations of

P. villosus are in doubt.

Amongst Molpadiodemas species, the distinguishing char-

acters are: close to sparse body cover of small, hair-like, off-

white tube feet, sometimes matted ventrolaterally and py gaily;

gonad ossicles.

Molpadiodemas violaceus (Theel, 1886) comb. nov.

Table 3, Figures le, i, 2f, 7g-i, 8u-x

Pseudostichopus villosus var. violaceus Theel, 1886: 172, pi. 10 fig

6b (raised to species status here).

Pseudostichopus villosus Theel, 1886: 170-171 (part; syntypes

from Challenger stns 61, 147, 325; non Pseudostichopus villosus

Theel, 1886).

Material examined. Holotype. Pseudostichopus villosus var. violaceus

Theel, 1886. Southern Ocean, off Shackleton Ice Shelf, 62°26'S,

95°44'E, 3612 m, Challenger stn 156, BMNH 86.10.2.156.

Other material. Pseudostichopus villosus Theel, 1886. Syntypes.

North-western Atlantic Ocean, 34°54'N, 56°38'W, 5212 m. Challenger

stn 61, BMNH 86.10.2.145 (2); southern Indian Ocean, off Crozet I.,

46°16'S, 48°27'E, 2926 m, Challenger stn 147, BMNH 86.10.2.154

(1); south Atlantic Ocean, off Argentina, 36°44'S, 46°16'W, 4846 m,

Challenger stn 325, BMNH 86.10.2.150 (1).

Weddell Sea, S of South Orkney Is, 3587-3660 m, USNM E48600

(1) ; 1008106 (4); NMV F101844 (2).

South Atlantic Ocean, Scotia Sea, South Orkney Is, 2196 m, USNM

E49337 (2); 3250-3285 m, E48651 (11); NMV F101843 (2);

Burdwood Bank, 3514-3642 m, USNM E48627 (1); 4026-4063 m,

E48654 (2); South Sandwich Is, 5435-5453 m, E48637 (6);

NMV F101842 (2); 4758 m, E48626 (1); 1190-1469 m, 1022456 (3);

Drake Passage, 3788-3944 m, USNM 1008117 (1); 2324-3020 m,

E48591 (1).

South Pacific Ocean, Chile, depth unknown, USNM E48636 (1);

Pern-Chile Trench, 6006 m, E48611 (1); 6146-6354 m, 1022608

(2) ; NMV F101848 (1); 5069-5173 m, USNM 1022609 (1); SE

Pacific Basin, 4731 m, E48630 (3); 4773 m, E49334 (1); 3859 m,

E48657 (3); 4572-4848 m, E49366 (1); 3694 m, E48605 (2); 4682 m,

1022455 (1).

Description. Up to 220 mm long; body wall leathery, pliable,

surface folded and wrinkled, sometimes mucilaginous; sparse

small grit and globigerine attachments; body rounded in trans-

verse section, tapered anteriorly, posteriorly, frequently sac-

like; ventrolateral margin rounded, not distinctive, lacking

brim, mouth terminal ventral; even cover of small soft

cylindrical tube feet all over body, never erect, evident in small

specimens only, slightly larger anteriorly and pygally, may be

matted, typically 1.0 mm, long, 0.3 mm diameter, frequently

withdrawn in small pits, not evident on smooth leathery body

wall of largest specimens; ossicles in tentacles abundant, rare in

gonad; tentacle ossicles variable, predominantly bluntly

tapered rarely curved unbranched smooth thick rods with dis-

tinct knob-like central swelling, some rods lacking central

swelling, some rods with bifurcate or intertwined ends, some

with lateral and terminal perforations, rare blunt spines and

knobs, rods frequently 250 pm long, up to 350 pm long; gonad

ossicles rarely present, straight and irregular sometimes

bluntly spinous short thin rods, some with central swelling,

some branching creating perforations, never mesh ossicles, up

to 150 pm long.

Colour. Brown to grey -brown to off-white; tube feet brown to

pale brown to off-white.

Distribution. Atlantic, southern Indian and Pacific, and

Southern Oceans, Weddell Sea, Scotia Sea, SE Pacific Basin,

2 196-63 54m.

Remarks. Theel (1886) established ten syntypes for

Pseudostichopus villosus, and a holotype for P. villosus var.

violaceus. Three of the ten syntypes of P villosus are con-

specific with the type of P. villosus var violaceus.

Amongst Molpadiodemas species, the distinguishing char-

acters are: leathery body wall; tube feet inconspicuous in large

specimens, when small even cover of soft cylindrical tube feet

with some anterior and pygal matting; tentacle ossicles fre-

quently thick smooth bluntly-tapered rods, many with large

knob-like central swelling; rare presence in gonad of short, thin

rod ossicles.

166

P. Mark O’Loughlin and Cynthia Ahearn

Figure 8. Tentacle ossicles (SEM). a-d, Meseres morbillus sp. nov. (USNM E49251). e-f, M. neovillosus sp. nov. (USNM 1008458, paratype).

g-h, M. pediculus sp. nov. (USNM 1008318, holotype). i— 1, M. porphyrus sp. nov. (USNM E38795, holotype). m-p, M. translucens sp. nov.

(USNM E48652, holotype). q-t, M. villosus, q-r (USNM E48640), s-t (USNM E9929). u-x, M. violaceus, u-v (USNM 1022608), w-x (USNM

E49366).

A review of pygal-furrowed Synallactidae

167

Figure 9. Characters of Pseudostichopus. a, drawing of transverse section of mid-body (as in P. mollis)-, b, cylindrical longitudinal muscles

detached from calcareous ring (P. tuberosus sp. nov., NMV F101864, paratype); c, dorsolateral radial papillae (P elegans, NMV F97447,

papillae up to 8 mm long); d, drawing of part of gonad, unbranched tubules, arising separately along gonoduct; e, two series of unbranched gonad

tubules along gonoduct ( P. mollis , TM H2004); f, gonad (P. tuberosus sp. nov., paratype, USNM 1008333).

Figure 10. a, Pseudostichopus hyalegerus, sponge spicule cover (P japonensis, paratype, SMBL 314, junior synonym, 29 mm long), b-c, P. mollis ;

b, rugose ventrum (TM H3111, 145 mm long); c, calcareous ring (TM H2004). d-e, P. papillatus (RAS, syntype, 27 mm long); d, dorsal view of

paired radial tubercles; e, ventrolateral view, tube feet, f-h, P. peripatus ; f, ventral view, marginal tubercles, grit attachment, transverse creases

(NMV F101840, 60 mm long); g, ventral view, marginal tubercles, grit attachment, transverse creases (NMV F101839, 45 mm long); h, cylindrical

muscles (arrows), gonad tubules ( P. tuberculatus , holotype, OMNH Ivll90, junior synonym), i, P. tuberosus sp. nov. (USNME16721, holotype,

115 mm long), ventral view, lateroventral tubercles (arrow).

168

P. Mark O’Loughlin and Cynthia Ahearn

Figure 11. Tentacle ossicles (SEM). a-b, Pseudo shichopus aemulatus (USNM 1025525, paratype). c-d, P. elegans (USNM E16505). e-f, P. mollis

(USNM E48656). g-h, P. tuberosus sp. nov. (USNM E16721, holotype). i-1, P. peripatus , i (USNM 12198), j (USNM E38794), k (USNM

E38796), 1 (USNM E49349).

Pseudostichopus Theel, 1886

Table 1, Figure 9

Pseudostichopus Theel, 1886: 169. — Ludwig, 1894: 38. — Perrier,

1902: 337-338.— Herouard, 1902: 11.— Fisher, 1907: 691.—

Mitsukuri, 1912: 3. — Herouard, 1923: 21-23. — Ekman, 1925:

32-36.— Ekman, 1926: 451-470, fig. I.— Mortensen, 1927:

386-388.— Deichmann, 1930: 86-87.— Heding, 1940: 356, 358-

360.— Djakonov, 1952: 125.— Imaoka, 1978: 377-378.— Thandar,

1992: 163-164.— Rowe, 1995: 285.— O’Loughlin, 1998: 497.—

O’Loughlin, 2002: 304.

Filithuria Koehler and Vaney, 1905: 81-81. — Heding, 1940:

356-357 (new synonym).

Pseudostichopus ( Pseudostichopus ). — Heding, 1940: 357, 360. —

Imaoka, 1978: 378.— Thandar, 1992: 164.

Pseudostichopus (Trachostichopus) Heding, 1940: 357, 361. —

Imaoka, 1978: 380 (new synonym).

Plicastichopus Heding, 1940: 357 (nomen nudum). — Heding, 1942:

5-6 (new synonym).

Peristichopus Djakonov, 1952: 125 (new synonym).

Type species. Pseudostichopus mollis Theel, 1886 (subsequent

designation by Fisher, 1907).

Other included species. Pseudostichopus aemulatus Solfs-

Marin and Billett, 2004; P. echinatus Thandar, 1992; P. elegans

(Koehler and Vaney, 1905); P. hyalegerus (Sluiter, 1901);

P. mollis Theel, 1886; P. occultatus Marenzeller, 1893;

P. papillatus (Djakonov, 1952); P. peripatus (Sluiter, 1901);

P. profundi Djakonov, 1952; P. spiculiferus (O'Loughlin, 2002);

P. tuberosus sp. nov..

Diagnosis. Characters of pygal-furrowed Synallactidae

(above); prominent appendages (tube feet, papillae) along the

paired radii only; longitudinal muscles cylindrical, not flat,

narrowly attached to the body wall; gonad tubules not

branched, arising in series along the gonoduct, not from a com-

mon base; ossicles sometimes present in tube feet and papillae;

tentacle ossicles predominantly unbranched rods, rarely rods

with ends intertwining and side branches fused to create mesh.

Distribution. Cosmopolitan; 91-5453 m.

Remarks. The larger radial appendages of the species of

Pseudostichopus are frequently tapered and variably elongate,

extremely so in P. elegans (below), and when distinctly tapered

are described in this work as papillae rather than tube feet.

The monotypic genus Filithuria Koehler and Vaney, 1905

was established for F. elegans Koehler and Vaney, 1905 which

is assigned here to Pseudostichopus and described in detail

below. F. elegans has the diagnostic characters of

Pseudostichopus (above), and the dorsolateral radial papillae

are distinctively elongate. The dorsolateral appendages in the

type species for Pseudostichopus, P. mollis, are moderately

elongate and distinctly tapered, and have the appearance of

papillae more than tube feet. The length of the dorsolateral

papillae is an inadequate single character on which to maintain

the monotypic genus Filithuria, which becomes a junior

synonym of Pseudostichopus.

Heding (1940) erected the genus Plicastichopus citing as

type species the manuscript name Plicastichopus ingolfi. Until

he formalised the name in 1942, the genus remained a nomen

nudum. O'Loughlin (2002) synonymised P. ingolfi Heding,

1942 with Meseres peripatus (Sluiter), Plicastichopus Heding,

1942 becoming a junior synonym of Meseres. Meseres peri-

patus is reassigned below to Pseudostichopus, Plicastichopus

now becoming a junior synonym of Pseudostichopus.

A review of pygal-furrowed Synallactidae

169

Figure 12. Gonad ossicles (SEM). a-d, Meseres villosus (USNM E9929). e-f, M. translucens sp. nov. (USNM E48652, holotype).

g-h, Pseudostichopus peripatus, g (USNM 12198), h (USNM E48638). i-1, P. tuberosus sp. nov. (USNM E16721, holotype). m, P. aemulatus

(USNM 1025525, paratype).

Posterior lobe ossicles (compound microscope), n-p, P. hyalegerus, n-o (NMV F80181), p (NMV F80178).

Tentacle ossicles (compound microscope), q-t, M. ustulatus sp. nov. (NMV FI 01 858, paratype).

Heding (1940) erected the subgenus Trachostichopus (of

Pseudostichopus ), with type species Pseudostichopus trachus

Sluiter. Rowe (1995) referred trachus to Meseres Ludwig,

Pseudostichopus ( Trachostichopus ) becoming a junior syn-

onym of Meseres. P. trachus is considered below to be a junior

synonym of Pseudostichopus mollis, Trachostichopus now

becoming a junior synonym of Pseudostichopus.

The monotypic genus Peristichopus was established for

Peristichopus papillatus Djakonov, 1952. Djakonov (1952)

characterised the genus by: cylindrical body; ventral mouth,

anus; pygal furrow; small tube feet on paired radii; cylindrical

longitudinal muscles; unbranched gonad tubules in two bun-

dles; tentacles with rod ossicles, some rod ossicles perianally

and in tube feet; and body wall, genital tubules, respiratory

trees lacking ossicles. Djakonov (1952) noted that

Peristichopus was close to Pseudostichopus, but gave no

diagnostic distinctions. Peristichopus papillatus is very close to

Pseudostichopus mollis, and no justification is found here for

maintaining Peristichopus Djakonov which is a junior synonym

of Pseudostichopus Theel.

Pseudostichopus aemulatus Solfs-Marin and Billett, 2004

Figures 11a, b, 12m

Pseudostichopus sp. — Billett, 1988: 196-197. — Billet et al., 2001:

325-348.

Pseudostichopus aemulatus Solfs-Marin and Billett (in Solfs-Marin

et al.), 2004: 1079-1084, figs 1, 2, tables 5, 6.

170

P. Mark O’Loughlin and Cynthia Ahearn

Material examined. Paratypes. Northeast Atlantic Ocean, Porcupine

Abyssal Plain, 48°45’-48°48'N, 16°36'-16°41'W, 4835-4838 m, RRS

Discovery , stn 54901#5, 28 Apr 1999, USNM 1025525 (3).

Description. Up to 146 mm long; body wall thick, gelatinous,

opaque, sometimes encrusted with foraminiferans and sand;

distinct brim, flat ventrally, convex dorsally; body surface

smooth, lacking close wrinkling with ridges and digitate pro-

jections; tapering papillae in double irregular series on dorso-

lateral radii, series not widely separated, papillae typically up to

1.5 mm long, 1.0 mm diameter at base; papillae in widely sep-

arated double series ventrolaterally, dorsal to or ventral to or on

brim; ossicles in tentacles and gonad; tentacle ossicles abun-

dant, tapering rods, thick to thin, finely to bluntly spinous or

smooth, with or lacking mid-rod swelling, rarely with short

branches, up to 260 pm long; gonad ossicles predominantly

small, thin, pointed, unbranched rods with mid-rod swelling,

some with sparse blunt spines, rods up to 140 pm long.

Colour. Body off-white.

Distribution. NE Atlantic Ocean, Porcupine Abyssal Plain,

4350M-850 m (Solfs-Marfn et al., 2004).

Remarks. R aemulatus is similar to P. peripatus (below), but is

distinguished by: body wall off-white (not greyish), opaque

(not semi-translucent), smooth (lacking ridges with digitate

projections); dorsolateral double series of papillae not widely

separated; ventrolateral double series of papillae widely

separated; papillae lack ossicles; gonad ossicles predominantly

unbranched pointed rods with mid-rod swelling. These charac-

ters also distinguish P. aemulatus from other Pseudostichopus

species.

Pseudostichopus echinatus Thandar, 1992

Pseudostichopus ( Pseudostichopus ) sp. — Heding, 1940: 360-361,

fig. 16.

Pseudostichopus ( Pseudostichopus ) echinatus Thandar, 1992:

164-167, fig. 2.

Description (based on the descriptions by Heding, 1940, as

P (Pseudostichopus) sp., and Thandar, 1992, and on additional

observations by A. Thandar, pers. comm.). Up to 65 mm long;

dense dorsal and lateral encrustation of shells, globigerines,

sand; subcylindrical body; 19 tentacles; scattered cover of very

small tube feet, some more prominent dorsolateral tube feet

evident amongst encrustations, series of ventrolateral tube feet

in prominent series; longitudinal muscles cylindrical;

unbranched gonad tubules in 2 series along gonoduct; ossicles

in tentacles and tube feet only; tentacle ossicles irregular

closely knobbed rods with some branching and perforations

created by joined branches, not mesh-like, up to 300 pm long.

Distribution. Indian Ocean, off eastern Africa, 1°4T-28 0 22'S,

32°35'^U°47’E, 200-825 m.

Remarks. P. echinatus is known from the holotype (SAM

A23435), a specimen discussed by Heding (1940), and a recent

specimen from KwaZulu-Natal, 200 m (A. Thandar, pers.

comm.). It has the diagnostic characters of Pseudostichopus

above. Amongst Pseudostichopus species the distinguishing

characters of P. echinatus are: encrusting cover of shells,

globigerines and sand; very irregular closely knobbed tentacle

rod ossicles; absence of gonad ossicles.

Pseudostichopus elegans (Koehler and Vaney, 1905) comb. nov.

Figures 9c, 11c, d

Filithuria elegans Koehler and Vaney, 1905: 81, pi. 6 figs 1, 2, pi.

1 2 figs 29-3 1 .—Heding, 1 940: 357, 362-363, fig. 18.

Material examined. NE Pacific Ocean, off Oregon, 3021 m, USNM

E16815 (1); 1008097 (3); 2510 m, USNM E16505 (3); NMV F97447

(1); 2884 m, USNM E16497 (1); 2710 m, E16490 (4); NMV F97446

0 ).

Description. Up to 145 mm long; elongate, subcylindrical,

tapering anteriorly and posteriorly, flat ventrally, high convex

dorsally, sometimes brim weakly developed; body wall thick,

leathery, firm thin outer layer, soft thin semi-gelatinous inner

layer; surface smooth dorsally and laterally, rugose ventrally;

paired radii with double series of long tapering papillae, very

thin distally, longest in upper dorsolateral series, up to 22 mm

long, smaller in lower dorsolateral and double ventrolateral

series; small tube feet all over body, up to 0.5 mm long, 0.3 mm

diameter; ossicles in tentacles, papillae, tube feet; tentacle

ossicles abundant tapering rods, long to short, thick to thin,

with or lacking central thickening, rare short branching,

spinous or smooth, up to 490 pm long; large dorsal papillae

with rods similar to tentacles, some more sharply and abun-

dantly spinous, shorter, up to 260 pm long; small papillae and

tube feet with thick to thin rods, many with central swelling,

bluntly spinous or smooth, short, up to 160 pm long.

Colour. Body violet-brown to reddish-brown to off-white;

gonad tubules and respiratory trees pale brown; longitudinal

muscles chocolate to reddish-brown.

Distribution. Bay of Bengal, off Andaman Is., 741 m (Koehler

and Vaney, 1905); Indian Ocean, off Sumatra, 750 m (Heding,

1940); North-east Pacific Ocean, off Oregon, 2510-3021 m

(this paper).

Remarks. Koehler and Vaney (1905) and Heding (1940)

reported single specimens of Filithuria elegans. Filithuria

Koehler and Vaney is considered above to be a junior synonym

of Pseudostichopus Theel. The distribution of P. elegans is

extended here from the Indian Ocean to the NE Pacific Ocean,

to greater depths. Amongst Pseudostichopus species the distin-

guishing characters of P. elegans are: distinctly long papillae on

the paired radii; numerous tapered rod ossicles in the papillae.

Pseudostichopus hyalegerus (Sluiter, 1901) comb. nov.

Figures 10a, 12n-p

Meseres hyalegerus Sluiter, 1901a: 12. — Sluiter, 1901b: 50-51, pi.

5 figs 2-4.— Perrier, 1902: 359.

Pseudostichopus trachus. — Mitsukuri, 1912: 3-9, pi. 1 figs 1-5. —

Ohshima, 1915: 227-228. — Mortensen, 1918: 80-81, fig. 16 (non

Pseudostichopus trachus Sluiter, 1901).

Pseudostichopus ( Trachostichopus ) tachimaruae Imaoka, 1978:

380-382, fig. 2A-E, table 1-2 (synonymy by O'Loughlin, 2002).

Pseudostichopus ( Trachostichopus ) japonensis Imaoka, 1978:

382-384, fig. 3A-D, table 1-2. — Imaoka, 1990: 148 (synonymy by

O'Loughlin, 2002).

A review of pygal-furrowed Synallactidae

171

Pseudostichopus molpadioides Ohshima, 1915: 228-229, pi. 8 figs

6a-c.— Heding, 1940: 353-359.— Imaoka, 1978: 384, tables 1, 2.—

Imaoka, 1990: 152. — O’Loughlin, 2002: 304-305 (new synonym).

Pseudostichopus arenosus Ohshima, 1915: 229. — Heding, 1940:

353-359.— Imaoka, 1978: table 1-2.— O’Loughlin, 2002: 304-305

(new synonym).

Material examined. Meseres hyalegerus Sluiter. Syntypes. Indonesia,

Banda Sea, 5°28'S, 132°00'E, 204 m, Siboga stn 251, ZMA

V.Ech.H9499 (1); ZMA V.Ech.H2 177/1 (6); ZMA V.Ech.H2 1772/2 (1,

mounted on glass in alcohol).

Pseudostichopus ( Trachostichopus ) tachimaruae Imaoka. Holotype.

Japan, W of Kyushu, near Shimo-Koshiki I., 400-450 m, T. Imaoka,

27 Oct 1976, SMBL 311. Paratypes. Type locality, date, SMBL 312

( 2 ).

Pseudostichopus ( Trachostichopus ) japonensis Imaoka. Holotype.

Japan Sea, off the Akita Prefecture, 200-300 m, S. Nishimura,

1972-1973, SMBL 313. Paratypes. Type locality and date, SMBL

314 (3).

Other material. North-west Pacific Ocean, Japan, 350-400 m,

USNM 1025085 (3); 300-150 m, 1025086 (17); NMV F101861 (5);

420 m, 1071801 (1); Sagami Bay, 128-553 m. Northwest Pacific

Expedition, Albatross stn 5092, USNM E17147 (42); NMV F101855

(5); stn 4968, USNM 1001645 (1); stn 5069, 1001646 (1); stn 5093,

1001647 (1); stn 5055, 1001648 (5); stn 5094, 1001649 (7).

Eastern Australian continental slope, Tasman Sea and Bass Strait,

293-1100 m, NMV F80171- F80175 (21), F80178 (3), F80181 (19),

F80448 (1); 3300-3350 m, AM J23009 (1).

Description. Up to 120 mm long; typically encrusted with

dense mat of predominantly sponge spicules, with shells and

globigerines; frequently densely covered with tubular, thin,

branched, tangled, epibiote; body dorsoventrally depressed,

subacute margin; body wall semi-gelatinous, firm, translucent

to opaque, wrinkled with pits and raised protuberances all over

body; short papillae on paired radii, sparse irregular double

series dorsolaterally, more numerous in band ventrolaterally;

papillae cylindrical at base, tapering distally, typically 2.0 mm

long, 0.3 mm diameter (at base); gonad tubules predominantly

unbranched, rarely singly branched, rarely bifid distally;

ossicles in tentacles, papillae, pygal lobes; tentacle ossicles

irregular, frequently spinous, elongate rods, branching rare,

up to 340 pm long; papillae with rare irregular rods up to 100

pm long; papillae sometimes with “endplates”, very irregular

perforated plates or irregular mesh of branched rods, up

to 70 pm wide; pygal lobe ossicles single, large, elongate, up

to 800 pm long, comprising dense perforated mesh of

branched rods connected with open mesh, bluntly spinous

marginally.

Colour. Body grey to off-white; papillae pale brown to off-

white.

Distribution. West Pacific, Indonesia, Banda Sea, 204 m

(Sluiter, 1901a); Japan, Sagami Sea, 141-564 m (Mitsukuri,

1912, as P. trachus ); Japan Sea, south of Honshu, 128-553 m

(Ohshima, 1915, as P. trachus)-, 200-300 m (Imaoka, 1978, as

P. japonensis ); Hokkaido, Shiribeshi, 714 m (Ohshima, 1915,

as P. molpadioides ), Kyushu, Koshiki Is, 781 m (Ohshima,

1915, as P. arenosus ); 400-450 m (Imaoka, 1978, as P. tachi-

maruae)’, eastern Australia, Tasman Sea and Bass Strait,

293-1100 m (this work).

Remarks. Eight of the eleven syntypes were present (Apr 2002)

in the ZMA and examined. All were taken from the same

locality, and the establishment of a lectotype is not necessary.

The characters of Meseres hyalegerus are consistent with the

diagnosis of Pseudostichopus, and M. hyalegerus is reassigned

to Pseudostichopus.

The types of Pseudostichopus molpadioides Ohshima, 1915

and P. arenosus Ohshima, 1915 cannot be located (see

Introduction). The principal characters of Pseudostichopus

molpadioides given by Ohshima (1915) are: up to 52 mm long;

cylindrical form; pygal furrow; covered by sand, foraminifer-

ans, some sponge spicules; 20 tentacles; tube feet in double

rows on paired radii, most numerous ventrolaterally, up to 2.0

mm long; unbranched gonad tubules; irregular spiny ossicles

perianally, up to 200 pm long; tube feet support rods up to 120

pm long, endplates up to 140 pm wide; lacking gonad ossicles.

Ohshima (1915) considered that Pseudostichopus arenosus

closely resembled P. molpadioides, the two species differing

only in minor differences in the form of tentacle ossicles and

absence of tube feet support rods in P. arenosus. In this study

the detection of tube foot support rods in P. hyalegerus was not

consistent, and the differing details of tentacle ossicle form in

P. hyalegerus fell within the variations reported by Ohshima

(1915). P. arenosus is considered here to be a junior synonym

of P. molpadioides. The only characters distinguishing

P. molpadioides from P. hyalegerus are the typical cover of

predominantly sponge spicules in the latter, and possible

differing form of perianal ossicles. The drawing of a perianal

ossicle for P. molpadioides by Ohshima (1915) is identical with

a fragment of the large perianal ossicles found in

M. hyalegerus. A variation in body cover from predominantly

to only some sponge spicules is considered here to be not

diagnostically adequate for distinguishing the species.

P, molpadioides and P. arenosus are thus junior synonyms of

P. hyalegerus.

O'Loughlin (2002) considered P. tachimaruae and

P. japonensis to be junior synonyms of M. hyalegerus, based on

the descriptions by Imaoka (1978). The types were examined in

this work, and the pygal lobe mesh ossicles unique to

P hyalegerus were found in paratypes of both P. tachimaruae

and P. japonensis. The synonymies are confirmed here.

Amongst Pseudostichopus species, the distinguishing char-

acters of P. hyalegerus are: encrusting mat of sponge spicules

and other attachments; ossicles in the papillae and pygal lobes.

Pseudostichopus mollis Theel, 1886

Figures lb, c, 9a, e, 10b, c, lie, f

Pseudostichopus mollis Theel, 1886: 169-170, pi. 10 figs 5, 6. —

Ludwig, 1898: 7.— Perrier, 1902: 337-338.— Fisher, 1907: 691.—

Ekman, 1925: 5, 28-36, figs 4, 5.— Ekman, 1926: 451-470, fig. Id.—

Heding, 1940: 353-360.— Imaoka, 1978: table 1-1.— Imaoka, 1990:

148.— Gutt, 1991a: 147, 152, figs 3, 6, table 2.— Gutt, 1991b: 321,

324.— Thandar, 1992: 167.— Rowe, 1995: 285.— O'Loughlin, 2002:

304.

Pseudostichopus trachus Sluiter, 1901a: 15-16. — Sluiter, 1901b:

52-53, pi. 5 fig. 1, pi. 8 fig. 8.— Perrier, 1902: 337-338.— Fisher,

1907: 693. — Ekman, 1925: 32-36. — Savel'eva, 1941: 74. — Djakonov,

1952: 127, 129.— Baranova, 1957: 239.— Djakonov et al., 1958:

172

P. Mark O’Loughlin and Cynthia Ahearn

366.— Imaoka, 1978: 384.— Cherbonnier and Feral, 1981: 383, 385,

fig. 16.

Pseudostichopus nudus Ohshima, 1915: 230. — Ekman, 1925:

32-36.— Heding, 1940: 353-359.— Djakonov et al., 1958: 367.—

Rowe, 1995: 285 (as junior synonym of Pseudostichopus pustulosus ;

non Pseudostichopus pustulosus Sluiter, 1901; new synonym).

Pseudostichopus ( Trachostichopus ) trachus . — Heding, 1940:

353-362, fig. 17.— Imaoka, 1978: table 1-2.— Thandar, 1992: 166.

Pseudostichopus ( Pseudostichopus ) dilatorbis Imaoka, 1978:

378-380, 384, fig. 1 A-E, table 1-1 (part; paratypes non P. dilatorbis-,

new synonym).

Pseudostichopus ( Pseudostichopus ) alatus Imaoka, 1990: 146-148,

fig. 1A-E (new synonym).

Meseres trachus . — Rowe, 1995: 285. — O'Loughlin, 1998: 497. —

O'Loughlin, 2002: 300, 312, table 3 (new synonym).

Pseudostichopus pustulosus . — Rowe, 1995: 285 (non

Pseudostichopus pustulosus Sluiter, 1901).

Material examined. Pseudostichopus mollis Theel, 1886. Lectotype

(designated here). Southern Pacific Ocean, off Chile, 52°45'S,

73°46'W, 448 m, Challenger stn 311, BMNH 2002.293.

Paralectotypes, type locality and date, 86.10.2.143 (3), 1956.10.3.6

(2); off Chile, 50°56'S, 74°14'W, 256 m, Challenger stn 309A,

86.10.2.142 (3); Southern Indian Ocean, off Marion I., 46°48'S,

37°49'E, 91-137 m, Challenger stn 144A, 86.10.2.144 (2).

Pseudostichopus trachus Sluiter, 1901. Lectotype (designated here).

Indonesia, Ceram Sea, l 0 ll'S, 130°09'E, 798 m, Siboga stn 161, ZMA

V.Ech.H2496.2 (1). Paralectotype, Timor Sea, 8°50’S, 127°02'E, 883

m, Siboga stn 286, V.Ech.H2496.1 (1).

Pseudostichopus nudus Ohshima, 1915. Neotype (designated here).

North Pacific Ocean, Bering Sea, USA, Alaska, Aleutian Is, Bowers

Bank, 54°33'N, 178°45'E, 1019-1068 m, RV Albatross, North-west

Pacific Expedition, stn 4774, 4 Jun 1906, USNM E10787. Same local-

ity and date, 1008140 (4).

Pseudostichopus ( Pseudostichopus ) alatus Imaoka, 1990.

Holotype. Japan, Tosa Bay, 350 m, H. Horikawa, 30 Jan 1989, OMNH

Ivll88. Paratype. Tosa Bay, 390^410 m, Ivll89 (1).

Other material. North Pacific Ocean, Bering Sea, Alaska, Bowers

Bank, 1068 m, USNM E10697 (1); off southern California, W of San

Nicolas I., 825 m, 32416 (19); off Oregon, 44°36'-46°01'N,

1 24°40'-l 24°45'W, 320-466 m, USNM E16503 (13); E16734-

E 16742 (20); E16745, E167446 (6); E16749-E16752 (9); E16886 (3);

E16888 (16); E53270 (2); off San Diego, 572 m, E2042 (2); 762 m,

E02035 (4); 379-636 m, E17014 (1); 914-969 m, E17016 (1); 1587 m,

32410 (3); Costa Rica, 245 m, 18269 (2).

South Pacific Ocean, south-eastern Australian continental slope,

460-1200 m, AM J16749 (1); J16836 (1); J20028 (2); J22248 (1);

J22938, J22939 (3); J22943 (2); J22955 (5); J22957 (1); J22960 (7);

J22964 (1); J22967- J22969 (9); J22972 (1); J22974, J22975 (6);

J23218 (1); J23220 (1); J23268 (1); NMV F80176, F80177 (2),

F80179, F80180 (5); TM H3111 (1); H3114 (2); W of Cape Sorrell,

972 m, H2004 (2); Tasman Sea, Lord Howe Rise, 1920 m, NMV

F97692- F97694 (4); New Zealand, Campbell I., 589-594 m, USNM

E48604 (2); Chatham Is, 964-1067 m, E48641 (1); Antipodes Is,

384-397 m, E48665 (7); off Chile, 960 m, 1002911 (2); Macquarie I.,

NMV F89907 (1); F85036 (4); 900 m, F89908 (1); Strait of Magellan,

485 m, USNM E48639 (3); 256-320 m, E48666 (25); 769-869 m,

E48656 (7).

Antarctica, Palmer Archipelago, 460-500 m, USNM 1005855 (1);

460-500 m, 1008441 (2); Antarctic Peninsula, 500-670 m, 1005856

(1); 630-650 m, 1022611 (2).

Description. Up to 225 mm long; body rounded in transverse

section, frequently flat ventrally, high convex dorsally; some-

times thick to thin, rounded, ventrolateral brim, especially

anteriorly; pygal furrow sometimes barely evident in largest

specimens; body wall leathery, firm, thick to thin; anterior mid-

dorsal soft inconspicuous madreporite; ventrally frequently

rugose, wrinkled with pockets and fine transverse ridges, folds

and protuberances sometimes evident along ventrolateral

margin, sometimes with raised multilobed tubercles pygally or

midventrally or in lateroventral series, tubercles sometimes sur-

mounted by tube feet, ventrum frequently encrusted with

globigerines; papillae most evident dorsolaterally in sparse

radial band, frequently 2 up to 4 wide, strongly tapered, some-

what whip-like, up to 5 mm long (170 mm long specimen),

frequently withdrawn or lost leaving pits; double series of ven-

trolateral tube feet rarely externally evident; minute tube feet

ventrally, rarely evident; ossicles in tentacles, tube feet,

perianally; tentacle ossicles numerous to sparse, irregular rods,

typically 120-240 pm long, frequently swollen mid-rod, some

thin branches, rare blunt spines, branches rarely connecting to

create perforations, lacking mesh ossicles, rare large irregular

tuber-like ossicles, up to 380 pm long; tube feet ossicles pre-

dominantly irregular, smooth, rarely branched rods, frequently

with swelling mid-rod, typically 100-130 pm long; pygal lobes

with small globular mesh ossicles, difficult to detect.

Colour. Body, tube feet pale brown to off-white; tentacles,

longitudinal muscles dark brown to brown; gonad tubules,

respiratory trees pale brown.

Distribution. Western Antarctica, Weddell Sea, 65°19'S,

56°48'W, 400 m (Ekman, 1925); Weddell Sea, 340-470 m

(Gutt, 1991); North and South Pacific Ocean, southern Indian

Ocean, eastern Australian continental slope, Antarctic Ocean in

Palmer Archipelago and off Antarctic Peninsula, 91-1587 m

(this work).

Remarks. Theel (1886) listed “numerous” syntypes from

Challenger stn 311. Six of these specimens (in three lots) were

examined in the BMNH (Apr 2002), and one disintegrated

specimen examined (Apr 2002) in ZMU (ZMA V.Ech.H2999).

One of these specimens is designated as lectotype above. Theel

(1886) also listed six specimens from stn 309A of which three

were examined in the BMNH. Theel (1886) further listed two

specimens from stn 144A, and both were present in the BMNH.

During an examination (Apr 2002) of holothurian types in

the University of Amsterdam, a type specimen of Pseudo-

stichopus trachus Sluiter was found to be registered as

Pseudostichopus pustulosus (discussed above under M. pustul-

osus). This specimen is designated the lectotype for P. trachus.

O'Loughlin (2002) judged material referred to P. trachus by

Mitsukuri (1912) and Ohshima (1915) to be M. hyalegerus, and

raised doubts about determination of material as P. trachus by

Heding (1940), Savel'eva (1941), Baranova (1957), Djakonov

et al. (1958), and Cherbonnier and Feral (1981).

Significant features of the types of P trachus are: large size,

up to 180 mm long (Sluiter, 1901b); slightly dorsoventrally

depressed, flat ventrally, convex dorsally, pygal furrow, leath-

ery wri nkl ed body wall, subacute pustulose margin with dis-

tinct small irregular protuberances; densely covered with

globigerine attachments; tube feet all withdrawn, paired radial

A review of pygal-furrowed Synallactidae

173

series ventrolaterally, small ones ventrally (Sluiter 1901b), dor-

sal tube feet sparsely evident dorsally (Sluiter, 1901b);

cylindrical longitudinal muscles ("not round in cross section"

according to Sluiter, 1901b), but seen here to be round but

sometimes slightly flattened); long thin unbranched gonad

tubules; "teeth" on posterior margin of radial plates of ring; ten-

tacle ossicles irregular thick rods, sometimes branched, branch-

es sometimes fused to create perforations, smooth or with large

thorny spines, sometimes with central swellings, up to 200 pm

long; tube foot ossicles short smooth tapered rods, frequently

with central swelling; body wall, gonads, respiratory trees lack-

ing ossicles. The types were taken in Indonesia at 798-883 m.

In all respects, except the dorsal cover of globigerines and

uncertain distribution of dorsolateral tube feet, P. trachus is

indistinguishable from P. mollis. Many specimens of P. mollis

have a dense ventral encrustation of globigerines, and quite

wrinkled and pitted and ridged and pustulose ventrum and

lateroventral edge. The presence of encrusting globigerines

dorsally is considered to be not grounds for separating the

species diagnostically. The various forms and sizes of the

tentacle ossicles are noticeably similar for the P. mollis and

P. trachus types, as are the teeth on the radial plates of the

ring. The longitudinal muscles are cylindrical, and sometimes

slightly flattened, in both species. The gonad tubules are

uniquely long and thin and unbranched in both species.

P. mollis is found on the Australian continentlal slope at depths

similar to those of the P. trachus types. This study has shown

that the degree to which the ventrolateral margin develops pro-

tuberances and serrations varies from strongly developed to

non-existent within the same species such as in

Pseudostichopus peripatus below. Similarly the ventrum may

be relatively smooth or strongly pustulose with raised protu-

berances as observed here for P. mollis. P. trachus is thus

judged here to be a junior synonym of P. mollis.

The “Type” for/! nudus nominated by Ohshima (1915), and

given USNM Cat. No. 34150, cannot be traced (see

Introduction). A neotype, selected from Ohshima’ s original

material, is designated above. P. nudus has all but one of the

distinguishing characters of P. mollis : firm, leathery body wall;

potentially large sac-like form; small tapered tube feet on the

paired radii; cylindrical longitudinal muscles; very long

unbranched gonad tubules, in series along gonoduct; tentacles

with irregular rod ossicles; tube feet with small rod ossicles

with central swellings; absence of gonad, respiratory tree, pos-

terior lobe ossicles. Ohshima (1915) distinguished P. nudus by

a midventral band of tube feet which were each surrounded by

pustular protuberances. A prominent midventral band of tube

feet arising from protuberances was seen in a number of speci-

mens in this study, but other specimens showed a range of ven-

tral conditions intergrading with P. mollis. Small tube feet are

present ventrally in P. mollis, and in some specimens become

evident and in some arise from irregular swollen protuberances.

P. nudus is also considered here to be a junior synonym of

P. mollis.

Rowe (1995) considered P. nudus to be a junior synonym of

P. pustulosus. This synonymy is rejected above (see M. pustu-

losus ). Rowe (1995) identified material from off Newcastle on

the eastern Australian slope as Pseudostichopus pustulosus. All

Australian slope material has been examined and none deter-

mined as P. pustulosus. Material from off Newcastle (AM

J 16749) was determined as P. mollis. Ludwig (1894) referred

material to P. mollis, but illustrated (Ludwig, 1894, pi. 9 figs

5-9) ossicles from the genital tubules that are similar to those

in the gonads of P. peripatus (see below). At least some of

the material was not P. mollis, which consistently lacks gonad

ossicles.

The two paratypes of P. dilatorbis have all the diagnostic

characters of P. mollis, and are so assigned (see above).

The distinctive features of the two type specimens of

P. alatus are: up to 115 mm long; pale brown to white, opaque

body encrusted with sponge spicules, foraminiferans, shell,

sand; 19 pale brown tentacles; tube feet on paired radii, more

numerous ventrolaterally; very small tube feet ventrally; long

thin unbranched gonad tubules in series along gonoduct; ossi-

cles in tentacles only, not in gonads or tube feet; tentacle ossi-

cles irregular rods about 150-400 pm long, frequently with

fairly large spines, rare branching. The types of P. alatus

exhibit all of the characteristics of P. mollis and P. trachus, and

these observations are confirmed by the description of Imaoka

(1990). P. alatus is considered here to be a junior synonym of

P. mollis.

Amongst Pseudostichopus species, the distinguishing char-

acters of P. mollis are: leathery brown body wall; typical

absence of a dense cover of globigerines or sponge spicules;

distinct papillae on the paired radii; absence of gonad ossicles.

Pseudostichopus occultatus Marenzeller, 1893

Pseudostichopus occultatus Marenzeller, 1893a: 15-17, pi. 4 fig.

9. — Marenzeller, 1893b: 10, pi. 2 fig. 3a-c. — Perrier, 1902:

337-338.— Mortensen, 1918: 81.— Mortensen, 1927: 387-388.—

Deichmann, 1930: 89-90.

Molpadiodemas occultatus. — Heding, 1940: 353-359.

Meseres occultatus . — O’Loughlin, 1998: 497. — O’Loughlin, 2002:

307, tables 1,3.

Material examined. Syntypes. Mediterranean Sea, Crete, 35°4'N,

24°17'E, 1445 m, 31 Aug 1891, MNHN EcHh 3658 (2); USNM 18294

( 2 ).

Description. Up to 40 mm long; encrusted with shell, sand,

sponge spicule attachments; body dorso ventrally depressed,

flat ventrally, low convex dorsally, acute ventrolateral margin

or thin brim; body wall parchment-like, wrinkled with low

reticulate ridges; appendages most evident as papillae dorsolat-

erally and in ventrolateral band, typically 1 mm long; ossicles

in tentacles, pygal lobes, tube feet, respiratory trees; tentacle

ossicles thick to thin irregular rods, some with central

swellings, rare short branches, rare blunt spines, lacking mesh

developments, up to 280 pm long; pygal lobe ossicles irregular

perforated plates, partly double-layered, up to 400 pm long;

tube foot ossicles smooth tapered rods with central swelling,

typically 80 pm long, “endplates” clusters of very irregular

twisted branched rods or irregular perforated plates, typically

80 pm wide; respiratory tree ossicles abundant thin, smooth,

symmetrically branched, pointed rods, typically 80 pm long.

174

P. Mark O’Loughlin and Cynthia Ahearn

Colour. Body surface appearance brown, body wall texture

grey.

Distribution. Mediterranean Sea, 415-1445 m (Marenzeller,

1893a); North Atlantic, Spain, off Cape Finisterre, 363-510 m

(Marenzeller, 1893b).

Remarks. P. occultatus has the diagnostic characters of

Pseudostichopus, and is returned to its original combination.

O’Loughlin (2002) described thin, tubular appendages cover-

ing the whole body. These are considered here to be epibiotes.

Amongst Pseudostichopus species, the distinguishing charac-

ters of P. occultatus are: short, thin, smooth, distally pointed,

symmetrically branched, respiratory tree ossicles; “endplate”

ossicles; irregular, partly double-layered, perforated plate

ossicles in the pygal lobes.

Pseudostichopus papillatus (Djakonov, 1952) comb. nov.

Figures lOd, e

Peristichopus papillatus Djakonov, 1952: 125-127, figs 11-14.

Material examined. Syntypes. Russia, SE of Kamtchatka, 52°45'N,

161°41'E, 4100^1200 m, 25 Jul 1946, RAS (6).

Description. Up to 65 mm long; torpedo-shaped body, some

gravel attachments; translucent body wall; prominent small

tapered papillae, arising singly on wart-like tubercles in double

rows on paired radii, typically 1.5 mm long; ossicles in

tentacles, papillae; tentacle ossicles unbranched thick to thin,

irregular rods, some swollen centrally, rugose at ends, some

bluntly spinous, up to 280 pm long; papillae ossicles irregular

branching rosette-like endplates, irregular rods, some swollen

centrally, some spinous, up to 260 pm long.

Distribution. Russia, SE of Kamtchatka, 4100-4200 m.

Remarks. Djakonov (1952) referred this species to his new

genus Peristichopus, which is considered above to be a junior

synonym of Pseudostichopus. Amongst Pseudostichopus

species, the distinguishing characters of P. papillatus are: the

regular paired series of low tubercles and papillae on each of

the paired radii; rosette-like “endplate” ossicles in papillae.

Pseudostichopus peripatus (Sluiter, 1901) comb. nov.

Figures If, lOf-h, 1 11— t, 12g, h

Meseres peripatus Sluiter, 1901a: 10-11. — Sluiter, 1901b: 48-49,

pi. 5 fig. 5, pi. 8 fig.7. — Perrier, 1902: 359. — Rowe, 1995: 285.

Pseudostichopus occultatus . — Herouard, 1902: 14-15, pi. 2 figs

4-14 (part, illustrated; non Pseudostichopus occultatus Marenzeller,

1893).

Pseudostichopus occultatus var. plicatus Koehler and Vaney, 1905:

9-10, pi. 3 fig. 8, pi. 9 figs 1-3.— Heding, 1940: 353 (non

Pseudostichopus occultatus Marenzeller, 1893).

Pseudostichopus propinquus Fisher, 1907: 691-693, pi. 71 fig. 3, pi.

72 fig. 2, pi. 73 fig. 3, pi. 74 fig. 1, pi. 76 fig. 3.— Imaoka, 1978: 382

(new synonym).

Pseudostichopus aleutianus Ohshima, 1915: 228, pi. 8 figs 5a-c. —

Imaoka, 1978: 380.

Pseudostichopus unguiculatus Ohshima, 1915: 230-231, pi. 8 figs

7a-c.— Imaoka, 1978: 384.— Rowe, 1995: 285.

Pseudostichopus marenzelleri Herouard, 1923: 25. — Mortensen,

1927: 287-288.— Deichmann, 1930: 90.

Pseudostichopus lapidus Herouard, 1923: 26-28, pi. 4 fig. 5.

Pseudostichopus ( Pseudostichopus ) lapidus. — Heding, 1940:

353-360 (new synonym).

Pseudostichopus ( Pseudostichopus ) marenzelleri. — Heding, 1940:

353-359. — Imaoka, 1978: table 1-1. — Thandar, 1992: 167 (synonymy

by O’Loughlin, 2002).

Pseudostichopus ( Pseudostichopus ) unguiculatus. — Heding, 1940:

353-360. — Imaoka, 1978: table 1-1. — Imaoka, 1990: 152. — Thandar,

1992: 167 (synonymy by Rowe, 1995).

Pseudostichopus ( Trachostichopus ) aleutianus. — Heding, 1940:

353-359. — Imaoka, 1978, table 1-2 (synonymy by O’Loughlin 2002).

Plicastichopus plicatus . — Heding, 1940: 354-359. — Heding, 1942:

6. (synonymy by O’Loughlin, 2002).

Pseudostichopus ( Trachostichopus ) propinquus. — Heding, 1940:

357.— Imaoka, 1978: table 1-1.— Imaoka, 1990: 148, 152.

Plicastichopus ingolfi Heding, 1942: 5-6, figs 4-5, pi. 1 figs 4-5.

Pseudostichopus ( Trachostichopus ) tuberculatus Imaoka, 1990:

149-152, pi. p. 149, fig. P. 15. (synonymy by O’Loughlin, 2002).

Meseres ingolfi . — Rowe, 1995: 285 (synonymy by O’Loughlin,

2002 ).

Meseres propinquus. — O'Loughlin, 2002: 309.

Material examined. Meseres peripatus Sluiter, 1901. Lectotype (desig-

nated here). Indonesia, Flores Sea, 7°24'S, 118°15E, 794 m, Siboga stn

45, ZMA V.Ech.H9500. Paralectotypes. Type locality and date, ZMA

V.Ech.H1048 (2); Maluka Sea, 1°59’N, 125°01E, 1200 m, Siboga stn

122, ZMA V.Ech.H 1049 (1); Banda Sea, 5°41'S, 120°46E, 1158 m,

Siboga stn 211, ZMA V.Ech.H1050 (2).

Pseudostichopus propinquus Fisher, 1907. Holotype. Hawaiian Is,

21°11'N, 156°35'W, 518-519 m, USNM 21217.

Pseudostichopus lapidus Herouard, 1923. Syntypes. North Atlantic

Ocean, off the Azores, 4020 m, Monaco stn 527, 1895, MOM (3)

(examination by M. Bruni, pers. comm., MOM).

Pseudostichopus ( Trachostichopus ) tuberculatus Imaoka, 1990.

Holotype. Japan, Tosa Bay, 660-700 m, H. Horikawa, 23 May 1989,

OMNH Ivll90.

Pseudostichopus unguiculatus Ohshima, 1915. Syntypes. North

Pacific Ocean, off Japan, 1058-1680 m, Albatross stns 4960, 5083,

5084, USNM 34151 (2), E24543 (1), E24544 (1).

Other material. Pacific Ocean, Tasman Sea, eastern Australia conti-

nental slope, 823-1750 m, AM J20026, J20027 (3); J23219 (1); J23267

(2); J22980 (9); NMV F80449, F80450 (3); F90070 (2); Philippines,

878 m, USNM E48764 (2); est. 1300 m, E48770 (2); off Thailand, 370

m, 1017465 (7); off Oregon, 1646 m, E48586 (3); 411 m, E17035 (2);

1946 m, E1972 (29); Mexico, 1608 m, 32389 (1); 2014 m, 32391 (2);

Costa Rica, 1789 m, 18267 (1); Peru-Chile Trench, 3500 m, E53272

(1) ; NMV F101841 (1); Galapagos Is, 2418 m, USNM 18272 (4); 1158

m, E949 (1); 3667 m, 1008450 (1); 1008457 (2); South-west Pacific

Basin, 3386-3422 m, E49306 (1); South-east Pacific Basin, N of

Amundsen Sea, 4978-5043 m, E48660 (5); 4709 m, E48590 (1); 4682

m, 1022459 (1); 4575-4813 m, E48629 (1); W Balleny Is, 2836-2864

m, E48632 (2); off Victoria Land, 3459-3492 m, E48624 (2); 566-569

m, 1022604 (2); Ross Sea, 3495-3514 m, 1008176 (1).

North Atlantic Ocean, off Florida, 931m, USNM El 990 (1); off

Massachusetts, 3235 m, 12198 (1); 3682 m, E53743 (1); off South

Carolina, 1337 m, E2581 (2); Bahamas, 4763^1803, 1021900 (1);

4578-4778 m, 1021902 (1); 4783-4823 m, 1021901 (1); Caribbean

Sea, Venezuelan Basin, 3428-3476 m, E38794 (2); 3411-3459 m,

E38796 (1).

Weddell Sea, 1025-1153 m, USNM E48573 (1).

South Atlantic Ocean, Scotia Sea, 52°00'-62 o 30'S, 14 o 54'-60°40'W,

267-5453 m, USNM E48572 (1); E48574- E48577 (15); E48581 (1);

E48585 (9); E48587, E48588 (2); E48592 (1); E48596 (6); E48599

(2) ; E48602, E48603 (5); E48607, E48608 (5); E48628 (2); NMV

A review of pygal-furrowed Synallactidae

175

F101840 (1); USNM E48638 (45); E48648 (23); NMV F101839 (5);

USNM E49241 (5); E49255 (10); E49325, E49326 (7); E49348,

E49349 (2); E49351 (1); E49444, E49445 (3); 1008141 (6); NMV

F101838 (3); USNM 1008159, 1008160 (7); 1008166 (5); 1008297

(4); 1022445 (2); 1022463 (3).

Antarctic Ocean, South Orkney Is, 3250-3285 m, USNM 1008177

(3); 1228-1400 m, E49393 (1); South Shetland Is, 662-1120 m,

E48570 (1); 884-935 m, E48610 (7); 213-311 m, E49350 (1); South

Sandwich Trench, 5350 m, USNM 1071584 (2); Palmer Peninsula, 134

m, E49259 (1).

Description. Up to 140 mm long; form of body variable, typic-

ally with encrusting globigerine or grit attachments; body elon-

gate, slightly tapered anteriorly and posteriorly, depressed to

varying degrees dorsoventrally, flat ventrally, typically low

convex dorsally; body wall firm, leathery, thick to thin, semi-

gelatinous, frequently with deep transverse grooves and ridges,

frequently wrinkled with low reticulate ridges surmounted by

small digitate projections, frequently pitted with withdrawn

very small interradial tube feet; ventrolateral margin acute to

subacute, rounded, variably serrated by the transverse body

folds and irregular transverse creases; ventrolateral margin

variable from rounded, to mammiform, to rounded protuber-

ances surmounted by elongate to multiple-knobbed extensions,

to kn obbed domes further surmounted by smaller knobs or

small tube feet or larger radial tube feet; larger tube feet sur-

mount double radial series of mammiform bases, not present

midventrally, mammiform base and papilla typically 2.0 mm

long; ossicles in tentacles and gonad, larger papillae; tentacle

ossicles abundant large irregular rods, thick to thin, terminally

tapered, frequently with central swelling, sometimes swollen

distally, sometimes with small ter mi nal knobs, sometimes

bluntly spinous, sometimes branched along rod with

branches closed to create perforations, rods up to 360 qm

long; papillae with spinous to smooth rods, frequently with

central swelling, up to 160 pm long; “endplates” sometimes

detected in papillae as perforated plates with mesh collar or

tangled mesh of joined irregular rods, up to 340 pm diameter;

gonad ossicles frequently present, abundant, typically small

predominantly slender tapering rods with small central hub,

spinous to smooth, unbranched or Y- or X-shaped, sometimes

irregular thick variably spinous branched rods, up to 200 pm

long.

Colour. Body grey to off-white, to residual pale reddish-brown

on some small specimens, sometimes semi-translucent; radial

tube feet brown, smaller tube feet off-white.

Distribution. Indo-Pacific Region, North and South Pacific

Ocean, North and South Atlantic Ocean, Scotia Sea, Antarctic

Ocean, Ross Sea, Weddell Sea; 134-5453 m.

Remarks. Seven syntypes of Meseres peripatus Sluiter, 1901

are listed (ZMA E1050) by Jangoux (1991). Only two were

present when the syntypes were examined (April 2002).

M. peripatus has the diagnostic characters of Pseudostichopus

detailed above, and is reassigned.

The description of P. propinquus Fisher, 1907 referred to:

thin translucent body wall, with some foraminiferan attach-

ment; distinct thickened lateroventral margin, with mammi-

form tubercles surmounted by tube feet; paired radii with tube

feet; round longitudinal muscles; unbranched gonad tubules;

gonad with fine branched sometimes spinous rod ossicles, up to

about 200 pm long; respiratory tree ossicles similar to gonad.

All of these characters accord with the diagnosis of

Pseudostichopus above, and propinquus is returned to its orig-

inal combination. All of the features of P. propinquus , except

the presence of respiratory tree ossicles, are diagnostic features

of Pseudostichopus peripatus. Fisher (1907) referred to two

specimens. The holotype is in very poor condition, and respir-

atory tree ossicles could not be confirmed. The second

specimen was not located. If gonad-type ossicles were found in

the respiratory trees of material otherwise conspecific with

P. peripatus, this would be judged to be an individual variation

rather than a basis for a separate species. P propinquus is con-

sidered here to be a junior synonym of P. peripatus. Rowe

(1995) made P. propinquus a junior synonym of P. pustulosus

Sluiter, 1901. This synonymy is rejected. P. propinquus has the

diagnostic characters of Pseudostichopus, and P. pustulosus

those of Molpadiodemas (above).

Based on the description by Herouard (1923), on observa-

tions communicated by M. Bruni (MOM), and on photographs

by Francisco Solis-Marin (UNAM) of a tentacle ossicles slide

prepared by Gustav Cherbonnier (MNHN box 108 slide 45),

the characters of P. lapidus are: up to 15 mm long; pygal fur-

row; encrusting cover of foraminiferans, sand, stones; tube feet

on paired radii, small to absent on midventral radius; longitudi-

nal muscles narrow; gonad tubules unbranched; ossicles in ten-

tacles only, lacking in body wall and internal organs; tentacle

ossicles predominantly rods without branches and associated

mesh. Although gonad ossicles have not been detected,

P. lapidus has the distinctive characters of Pseudostichopus

peripatus and it is judged here to be a junior synonym.

O’Loughlin (2002) considered P. marenzelleri to be a junior

synonym of P. peripatus, based on the description and figures

by Herouard (1923). Based on the additional description by M.

Bruni (MOM, pers. comm.) of cylindrical longitudinal muscles

and unbranched gonad tubules, the synonymy is confirmed

here. O'Loughlin (2002) considered P. tuberculatus to be a

junior synonym of P. peripatus, based on the photograph and

description in Imaoka (1990). Following an examination of the

holotype the synonymy is confirmed.

The considerable variety in ossicle and body form, and cos-

mopolitan distribution and depth range, suggested to us that

there are probably more than one species involved. We were

unable to recognize discrete diagnostic characters on which to

distinguish further species.

The distribution summary by O’Loughlin (2002) indicated

an Indo-Pacific and North Atlantic distribution for P. peripatus.

Data above extend the distribution to the eastern Pacific,

western and southern Atlantic, and Antarctic Oceans, and to

significantly shallower and greater depths.

Amongst Pseudostichopus species, the distinguishing char-

acters of P. peripatus are: dorsoventrally depressed body, with

strong wrinkling and transverse creases creating a serrated

ventrolateral margin; typical encrusting cover of globigerines

or rounded grit attachments, not sponge spicules; rods of

various forms in gonads, no mesh ossicles.

176

P. Mark O’Loughlin and Cynthia Ahearn

Pseudostichopus profundi Djakonov, 1952

Pseudostichopus profundi Djakonov, 1952: 127-129, figs 15-18.

Material examined. Syntype (RAS). Russia, SE of Kamtchatka,

52°45'N, 161°41E, 4100-4200 m, 25 Jul 1946.

Description. Up to 56 mm long; body cylindrical; body wall

thick, opaque, soft; few sand grain attachments; very small tube

feet unevenly distributed all over body, lengths variable, largest

anteriorly and posteriorly, fewest mid-dorsally and mid-

ventrally; 20 tentacles; radial plates of calcareous ring lacking

"teeth"; longitudinal muscles cylindrical; gonad tubules

unbranched; tentacle ossicles thick, broad, bent, irregular,

variably perforated, somewhat mesh-like, typically 200 pm

long; tube feet ossicles variable from irregular rods, to knots, to

irregular curved mesh-like narrow plates, up to 180 pm long;

posterior lobes with rare unbranched rods, irregular ends, up to

O. 17 mm long; gonad and respiratory trees with small, smooth,

sparse rod fragments.

Colour. Body and tube feet brown to grey.

Distribution. Russia, SE of Kamtchatka, 4100-4200 m.

Remarks. The description above is based on Djakonov (1952).

The syntype that was examined here was 27 mm long, and very

soft. The longitudinal muscles were rounded laterally, but

broadly attached to the body wall and not cylindrical, and are

considered here to be flat. No gonad was found. The pygal tube

feet were soft, cylindrical, and typically 0.3 mm long, 0.15 mm

diameter. No ossicles were found in the tube feet. The absence

of ossicles in the tube feet, and flat longitudinal muscles, sug-

gest that the syntype examined here is not conspecific with

P profundi as described. Djakonov (1952) described very

small, smooth rods in genital tubules and respiratory trees. We

judge these to be probably artefacts and not ossicles, since such

very small fragmentary rods are frequently present in gonad

and respiratory tree preparations.

Reference by Djakonov (1952) to cylindrical longitudinal

muscles and unbranched gonad tubules indicate an appropriate

assignment of the species to Pseudostichopus. But the flat

longitudinal muscles, and absence of prominent tube feet on the

paired radii, indicate that the syntype examined here belongs in

Molpadiodemas. The characters of the species, and generic

assignment, remain unresolved.

Amongst Pseudostichopus species the unique character of

P. profundi (as described by Djakonov) is the presence of

curved mesh-like narrow plates in the tube feet.

Pseudostichopus spiculiferus (O’Loughlin, 2002) comb. nov.

Figure lg

Pseudostichopus sp. MoV 2068. — O’Loughlin et al., 1994:

253-255.

Meseres spiculiferus O’Loughlin, 2002: 309-312, figs la-f, 2a-d,

tables 1-3.

Material examined. (See O’Loughlin, 2002). Antarctic Ocean, Weddell

Sea, 1554 m, USNM 49279 (1); Antarctic Peninsula, 370-375 m,

E48582 (1); 246-270 m, E49242 (2); 326 m, 49258 (2); 460-500 m,

E49472 (4); Ross Sea, 73°58'-78°23'S, 168°50’E-161°57'W,

364-728 m, E48598 (2); E48643 (2); E49268, E49269 (7);

1005124 (1); E49287 (2); E49354 (1); E49358 (2); 485^190 m,

1005121 (4); 1005123 (3); New Zealand, Antipodes Is, 2010-2100 m,

1005113 (1).

Description. Up to 105 mm long; body typically encrusted with

sponge spicules, sometimes globigerines; body flat ventrally,

convex dorsally; lateroventral margin subacute, some reticulate

ridges with small digitate projections, lacking prominent ven-

trolateral projections; irregular double rows of tube feet dor-

solaterally, larger along ventrolateral margin in band up to 5

irregular rows wide and continuous around anterior body,

sometimes contiguous, up to 3.0 mm long, 0.8 mm diameter;

ossicles in tentacles, tube feet and gonad; tentacle ossicles thick

to thin, curved to straight irregular rods, frequently swollen

mid-rod, rarely swollen terminally, rarely branched, rarely

bluntly spinous, up to 320 pm long; tube feet with rods

distally, same form as tentacle rods, up to 200 pm long; “end-

plates” of close cluster of irregular rods, up to 130 pm wide;

gonad ossicles abundant smooth rods, tapered to points at ends,

variable shape, frequently with central swelling, X- or Y-

shaped or complex multiple branching, branches sometimes

joined to create large perforations, ossicles frequently 120 pm

long, up to 300 pm long.

Colour. Body off-white or grey or grey-brown; tube feet off-

white to reddish-brown.

Distribution. Eastern Antarctica, off Wilkes, MacRobertson,

Enderby Lands, 177-695 m (O’Loughlin, 2002); Ross Sea,

364-728 m; Antarctic Peninsula, 246-500 m; Weddell Sea,

1554 m; New Zealand, Antipodes Is, 2100 m.

Remarks. Meseres spiculiferus has the diagnostic characters of

Pseudostichopus above, and is reassigned. The above data

extend the distribution of O’Loughlin (2002) from eastern

Antarctica to the Weddell Sea and New Zealand Antipodes

Is, and depth to 2100 m. Amongst Pseudostichopus species,

the distinguishing characteristics of P. spiculiferus are:

dense cover of sponge spicules; absence of large pygal lobe

ossicles; lateral band of tube feet; complex branching gonad

ossicles.

Pseudostichopus tuberosus sp. nov.

Figures 9b, f, lOi, 1 lg, h, 12i— 1

Material examined. Holotype. North Pacific Ocean, off Oregon,

45°43'N, 125°13W, 1920 m, Commando , 28 May 1964, USNM

E16721.

Paratypes. Holotype locality and date, USNM 1008333 (1); NMV

F101864 (1).

Other material. North Pacific Ocean, off southern California,

32°33'N, 118°04'W, 1937 m, USNME17015 (1).

Description. Up to 140 mm long; body wall leathery, thin, firm;

body flat ventrally, convex dorsally; prominent single series of

ventrolateral protuberances (“warts”), irregularly rounded, typ-

ically 2.0 mm wide, domed, soft and thin-walled, or firm and

flat, not surmounted by papillae, ventral to series of ventrolat-

eral papillae; body transversely creased and wrinkled ventrally,

sometimes dorsally, abundant grit attachments, ventrolateral

margin with indentations and lobes created by transverse

A review of pygal-furrowed Synallactidae

177

creases; irregular double rows of tapering papillae on both

paired radii, up to 2.0 mm long; minute tube feet all over body,

up to 0.2 mm long; ossicles in tentacles and gonad; tentacle

ossicles abundant, predominantly small thick to thin curved

rods, bluntly to finely spinous or smooth, swelling mid-rod

sometimes, rare short branches, rods up to 340 pm long; gonad

ossicles abundant, small unbranched rods, tapering to pointed

ends, slightly spinous to smooth, typically with central

swelling, up to 140 pm long.

Colour. Body, tube feet and protuberances off-white.

Etymology. From the Latin tuberosus (“full of lumps or pro-

tuberances”), referring to the distinct large protuberances along

each ventrolateral margin.

Distribution. North-eastern Pacific Ocean, off Oregon and

southern California, 1920-1937 m.

Remarks. The ventrolateral protuberances are wart-like ("fungi-

form papillae"), and resemble similar features in Bathyplotes

moseleyi. Amongst Pseudostichopus species the distinguishing

characters of P. tuberosus are: ventrolateral series of wart-like

protuberances; presence of small unbranched tapered rods in

the gonad.

Incertae Sedis

Meseres Ludwig, 1894

Meseres Ludwig, 1894: 34, 36, 38. — Perrier, 1902: 359. —

Rowe, 1995: 284-285.— O’Loughlin, 1998: 497.— O’Loughlin, 2002:

303-305.

Type species. Meseres macdonaldi Ludwig, 1894 (monotypy).

Diagnosis (emended from O'Loughlin, 2002). Up to 30 mm

long; completely lacking ossicles; thin translucent body wall;

body rounded anteriorly, posteriorly; mouth, anus ventral; lack-

ing pygal furrow; ventrolateral margin with series of pyramidal

protuberances surmounted by typically 3 digitate projections /

(? tube feet); close body cover of small papillae / (? tube feet)

and thin tubular appendages (probably epibiotic); 15 small

shield-shaped tentacles; lacking tentacle ampullae; five small

radial and five interradial plates in ring; longitudinal muscles

undivided; unbranched gonad tubules on each side of dorsal

mesentery; respiratory tree comprising two thin-walled tubes

with sacs.

Material examined. Meseres macdonaldi Ludwig, 1894. Lectotype

(designated here). Pacific Ocean, off Costa Rica, Cocos I., 5°56'N,

85°10'W, 2149 m, RV Albatross stn 3362, 26 Feb 1891, USNM 30501

(1). Paralectotypes. Pacific Ocean, off Colombia, Malpelo I., 4°3'N,

81°31'W, 1644 m, RV Albatross stn 3380, 5 Mar 1891, USNM 18190

(5 now unrecognisable specimens).

Remarks. The diagnosis of Meseres by O'Loughlin (2002) was

based on the species assigned at that time to Meseres , and not

exclusively on the type species M. macdonaldi. The emended

diagnosis is based solely on the description of the type materi-

al by Ludwig (1894), and some observations on the remaining

type material. The syntypes of M. macdonaldi were taken off

Costa Rica at 2149 m (USNM 30501 (1)), and off Colombia at

1644 m (USNM 18190 (5)). All are in very poor condition, but

a single syntype (USNM 30501) retains some recognisable tis-

sue and body parts. For the purpose of having a precise type

locality for this type species of Meseres, and some specific tis-

sue for possible molecular analysis, the syntype registered

alone is designated here as lectotype.

The characters of M. macdonaldi were not illustrated by

Ludwig (1894), and all of the type material is in an advanced

state of decomposition. Whether the body was covered with

tube feet or papillae remains subjective. The upper limit of 15

tentacles (in this study Synallactidae species 30 mm long have

more than 15 tentacles), and complete absence of ossicles, indi-

cate that Meseres does not belong in the Synallactidae. Meseres

has most of the characters of the Gephyrothuriidae as re-diag-

nosed by O'Loughlin (1998), with the exception of digitate ten-

tacles. Ludwig (1894) described small shield-shaped tentacles,

but there is no evidence as to what precise form they took. It is

relevant to note that one of the two species of the

Gephyrothuriidae reviewed by O'Loughlin (1998),

Hadalothuria wolfii Hansen, had ventrolateral protuberances

surmounted by rudimentary (?) tube feet. This feature appears

to be shared with M. macdonaldi. The description of shield-

shaped tentacles is an obstacle to referring the genus to the

Gephyrothuriidae and Meseres, as exemplified by its type

species, is treated here as incertae sedis.

Sluiter (1901) and O'Loughlin (1998, 2002) referred species

to Meseres principally on the basis of a distinctive ventrolater-

al margin, and Rowe (1995) on the basis of “gonads in two dis-

crete bunches of divided or undivided tubules, one on each side

of the dorsal mesentery”. All of these species referred to Meseres

had 18-20 tentacles, a distinct pygal furrow, and tentacle ossi-

cles, and are hence not congeneric with M. macdonaldi. Except

for the type species M. macdonaldi, and Meseres (?) torvus

(Theel, 1886), all species previously referred to Meseres are reas-

signed above to Molpadiodemas Heding or Pseudostichopus

Theel. O’Loughlin (2002) listed Molpadiodemas Heding, 1935,

Trachostichopus Heding, 1940 and Plicastichopus Heding

1940 as junior synonyms of Meseres Ludwig, 1894. These

three genera are removed from the synonymy, and are

discussed under Molpadiodemas and Pseudostichopus above.

O’Loughlin (2002) referred Stichopus (?) torvus Theel,

1886 to Meseres, based on the original description and the com-

parison with Meseres macdonaldi by Ludwig (1894). The

single type specimen was collected in the South Pacific Ocean

off Chile at 2516 m. A subsequent examination (Apr 2002) of

the type revealed the following: strongly contracted, 160 mm

long (210 mm in Theel 1886); cylindrical; irregular double

rows of large tube feet on paired radii; dorsal and lateral sur-

faces covered with long conical papillae of varying sizes,

largest ventrolaterally, rare branching; small pygal furrow

present as a split above anus (possible artefact); longitudinal

muscles round, deeply split (probable artefact); gonad tubules

short, sac-like, branched; ossicles not detected in body wall,

tentacles, papillae, tube feet, gonad, or respiratory trees. The

calcareous ring was not Pseudostichopus- like. These characters

have little affinity with Molpadiodemas Heding or

Pseudostichopus Theel, and torvus is referred back to the

original combination Stichopus (?) torvus.

178

P. Mark O’Loughlin and Cynthia Ahearn

Acknowledgments

We are grateful for the commitment of resources by successive

United States Administrations, which has resulted in the invalu-

able collections now held by the Smithsonian Institution and

which were the principal subjects of this work. We are appre-

ciative of the assistance with materials, resources and advice

given by: Nadia Ameziane (MNHN); Joke Bleeker (ZMA ); Ben

Boonen (manipulation of images and format of figures); Susann

Braden, Scott Whittaker (USNM; assistance with SEM);

Michele Bruni (MOM); Andrew Cabrinovic, Sheila Halsey,

Miranda Lowe (BMNH); Tohru Imaoka (OMNH); David

Pawson (USNM); Gary Poore (NMV); Frank Rowe (AM);

Alexei Smirnov (RAS); Francisco Solis-Marin (UNAM; recog-

nised additional pygal-furrowed synallactid material in the

unidentified collections of the USNM which yielded two of the

new species); Ahmed Thandar (University of KwaZulu-Natal);

Linda Ward (USNM; assistance with EMU); and Shigeyuki

Yamato (SMBL). We are particularly grateful for corrections

and suggestions proposed by the editor, Gary Poore, and three

reviewers David Pawson, Frank Rowe and Ah med Thandar.

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Memoirs of Museum Victoria 62(2): 181-189 (2005)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A new asterinid genus from the Indo-West Pacific region, including five new

species (Echinodermata: Asteroidea: Asterinidae)

P. Mark O’Loughlin 1 and Francis W.E. Rowe 2

‘Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia

(pmo @ bigpond.net.au)

2 Research Associate, Australian Museum, Sydney, New South Wales, Australia (Fwer.Goldbrookenn@btintemet.com).

Private address: Goldbrook Kennels, Hoxne, Suffolk, IP21 5BB, U.K.

Abstract O’Loughlin, P. M., and Rowe, F.W.E. 2005. A new asterinid genus from the Indo-West Pacific region, including five new

species (Echinodermata: Asteroidea: Asterinidae). Memoirs of Museum Victoria 62(2): 181-189.

A new genus of asterinid asteroid, Ailsastra gen. nov., and five new species, Ailsastra achituvi, sp. nov., A. amezianeae,

sp. nov., A. booneni sp. nov., A. eleaumei sp. nov., A. paulayi sp. nov. (type species), are described from the Indo-West

Pacific. Asterina heteractis H.L. Clark, 1938 is reassigned from Aquilonastra O’Loughlin, 2004 to Ailsastra gen. nov.

The external morphological characters distinguishing Ailsastra from other genera of Asterinidae are the distinctively

small oral plates and the variably sunken regular carinal series of plates. Internally, superambulacral and superactinal

plates, where investigated, are known to be lacking in three species, including the type species. A key to the six species

and a distribution map of type localities are provided.

Keywords Echinodermata, Asteroidea, Asterinidae, Ailsastra, new genus, new species, taxonomy

Introduction

The Asterinidae have undergone three major revisions (Verrill,

1913; A.M. Clark, 1983; O’Loughlin and Waters, 2004). The

most recent included for the first time a combination of mor-

phological and (limited) molecular analyses. This resulted in

the reassessment and rationalisation of some established

genera, the description of four new genera (bringing the total to

21), and the inclusion of 103 valid species in the family.

Rowe and Richmond (2004) found, in the course of identi-

fying specimens of two species of ‘ Asterina ’ (= Aquilonastra

O’Loughlin, 2004) from Rodrigues Island in the south-western

Indian Ocean, that a problem lay in determining the correct

identification of Aquilonastra burtoni (Gray, 1 840) and the dif-

ferentiation of supposedly conspecific species included in its

synonymy. We have widened their work and undertaken a com-

parative and systematic study of Indo-West Pacific asterinid

taxa.

In the process of solving this problem, additional new

material became available from Drs Chantal Conand (La

Reunion Island), Gustav Paulay (Indo-West Pacific), Matthew

Richmond (western Indian Ocean), and Yves Samyn (south-

eastern Africa). Loans of material (including type specimens)

have been made by Museum National d’Histoire Naturelle,

Paris, The Natural History Museum, London, South African

Museum, Cape Town, Australian Museum, Sydney, Museum of

Comparative Zoology, Harvard University, Massachusetts, The

Hebrew University of Jerusalem, and Tel-Aviv University.

The availability of this abundant material for comparative use

has led to the recognition of yet further, numerous, new aster-

inid taxa that require description, and has allowed a review of

some past, published determinations.

Amongst the new taxa it became apparent that both mor-

phological and molecular (Jon Waters, pers. comm.; paper in

preparation) evidence supported the recognition of a new,

wide-spread, Indo-West Pacific genus of small, stellate aster-

inids with regular carinal series and small oral plates. Included

in this genus are the eastern Australian species Asterina heter-

actis H.L. Clark, 1938 and five new species, extending in range

from the Red Sea and western Indian Ocean to Indonesian

waters. This paper includes descriptions of these taxa, and fore-

shadows an anticipated series on further Indo-West Pacific

Asterinidae. The number of taxa included in the family now

rises to 22 genera and 108 species.

Terminology follows O’Loughlin and Waters (2004). R/r

values are given with the recognition that they vary within a

species on the same specimen, amongst specimens, and with

size of specimen. Abbreviations for institutions are: AM,

Australian Museum, Sydney; MNHN, Museum National

d’Histoire Naturelle, Paris; NHM, The Natural History

182

P. Mark O’Loughlin and Francis W.E. Rowe

Museum, London; NMV, Museum Victoria, Melbourne,

Australia; UF, University of Florida, USA; MCZ, Museum of

Comparative Zoology, Harvard University, Massachusetts,

USA. Photographs were made with a Leica MZ16 stereomicro-

scope, DC300 Leica digital camera, and “Auto-Montage” soft-

ware for composition of images.

Asterinidae Gray, 1840

Ailsastra gen. nov.

Figure 1

Type species. Ailsastra paulayi sp. nov.

Other included species. Ailsastra achituvi sp. nov.; A.

amezianeae sp. nov.; A. booneni sp. nov.; A. eleaumei sp. nov.;

Asterina heteractis H.L. Clark, 1938.

Diagnosis. Small, stellate asterinids, up to R = 16 mm; rays pre-

dominantly 5; one species fissiparous (5-7 rays); interradial

margin incurved, rays discrete, broad to narrow at base, short to

long, rounded distally; flat actinally, convex abactinally, infero-

marginal plates project to form acute ventrolateral margin; disc

distinctly to irregularly defined; carinal series of plates present

for most of ray, doubly papulate with 2 longitudinal series of

large papulae for most of ray, each plate with rounded anterior

projection; carinal plates imbricated by adcarinal plates on both

sides, forming characteristic ‘sunken’ series; other abactinal

plates closely imbricate, not notched, arched over papular space

more than indented proximally for papula; lacking secondary

plates; papulae on rays single per papular space, large, in

longitudinal series; spinelets glassy, thick to thin, conical or

sacciform or subsacciform, pointed or splay-pointed, in clusters

or single or double series across proximal plates, sometimes in

round or oval bowl-shaped series on plate; lacking pedicel-

lariae with differentiated valves; plates with prominent glassy

convexities; superomarginal and inferomarginal plates in

regular series; actinal interradial plates in predominantly

oblique series; oral plates distinctively small, constricted dis-

tally (pair with form of short-handled, smooth-headed mace or

club); oral spines up to 6, series incurved distally; suboral

spines variably present, if present in continuous series with

distal oral spines; lacking superambulacral and superactinal

plates in type species and two other investigated species.

Distribution. Lord Howe I., NE Australia, Sulawesi I.,

Sudanese Red Sea, Mauritius, NW Madagascar; rock and coral;

0-16 m.

Etymology. From Ailsa with the Latin astrum (star), in recog-

nition of the substantial contribution by Ailsa Clark to seastar

systematics (feminine).

Remarks. Superambulacral and superactinal plates are absent

from the type species A. paulayi, and A. amezianeae and

A. eleaumei. The remaining three species (which have not been

dissected) have the other generic characters of Ailsastra, and it

is anticipated that they too lack these internal plates. Ailsastra

shares the lack of superambulacral and superactinal plates with

seven other genera currently included in the Asterinidae:

Asterina, Asterinides, Kampylaster, Meridiastra, Pseuda-

sterina, Pseudopatiria and Tremaster (see O’Loughlin and

Waters, 2004). However, the form of the oral plates and

arrangement of the carinal series of plates distinguish Ailsastra,

morphologically, from these and all other genera in the family.

The morphological justification for recognising the new genus

is supported by limited molecular data that places the two

specimens of the type-species A. paulayi sp. nov. (see below) in

a highly divergent but strongly monophyletic clade within a

molecular phylogeny of Asterinidae (Jon Waters, pers. comm.;

paper in preparation).

Key to species of Ailsastra

1. Rays 5-7; more than 1 madreporite; fissiparous

A. heteractis

— Rays predominantly 5; single madreporite; not fissiparous

2. Integument obscures abactinal and actinal plates; lacking

superomarginal spinelets; proximal actinal large pyramidal

spines present A. booneni

— Abactinal and actinal plates not obscured by integument;

superomarginal spinelets present; lacking proximal actinal

large pyramidal spines 3

3. Rays narrow at base, R/r > 2.5, digitiform; spinelets up to

0.25 mm long A. paulayi

— Rays broad at base, R/r < 2.0, not digitiform; spinelets not

longer than 0.20 mm 4

4. Form subpentagonal; up to 8 longitudinal series of papulae

across mid-rays; spinelets up to 0.10 mm long

A. amezianeae

— Rays discrete, form not subpentagonal; 6 longitudinal

series of papulae across mid-rays; some spinelets more

than 0. 10 mm long 5

5. Proximal abactinal spinelets frequently in bowl-shaped

arrangement A. eleaumei

— Proximal abactinal spinelets never in bowl-shaped

arrangement A. achituvi

Ailsastra achituvi sp. nov.

Figures 1, 2a, 3a, 4a-c

Asterina burtoni . — Clark and Rowe, 1971: 70, table 1 (part: speci-

men 1951.5.7.12) (non Asterina burtoni Gray, 1840).

Material examined. Holotype (in alcohol). Sudanese Red Sea, Khor

Inkeifail, on coral, Manihine Collections, 29 Dec 1950, NHM

1951.5.7.12.

Diagnosis. Ailsastra with, at R = 10 mm, discrete rays, medium

length, wide at base, strongly tapered, rounded distally,

R/r about 1.9; carinal series of plates narrowly visible,

below adcarinal plates; 6 longitudinal series of papulae across

mid-ray; spinelets splay-pointed, subsacciform to conical, up to

0.15 mm long; inferomarginal spinelets twice the length of

superomarginal spinelets; up to 6 interradial actinal spines

per plate.

Description. 5 rays, R = 10 mm; integument evident; rays discrete,

wide at base, tapered, rounded distally, R/r about 1.9; single

A new asterinid genus

183

madreporite, not fissiparous. Carinal series of plates narrowly visible,

below adcarinal plates, doubly papulate, other ray plates slightly

incurved proximally for 1 papula, 2 longitudinal series of papulae on

each upper side of ray, 6 longitudinal series across mid-ray; disc close

to regularly defined by 5 radial, 5 interradial plates; spinelets glassy,

splay-pointed, subsacciform to conical, up to 14 spinelets in single or

double series across proximal edge of plate, spinelets up to 0.15 mm

long, 3-6 on distal interradial plates, proximal and distal spinelets

subequal; some spinelets may act as pedicellariae, but not differen-

tiated as valves; up to about 6 superomarginal spinelets per plate, up to

about 14 inferomarginal spinelets per plate, longest distally, about

twice length of superomarginal spinelets. Actinal spines per plate: oral

5; suboral 0-1; furrow (proximal) 3^1; subambulacral 3; adradial

actinal 4-6; interradial 4-6; interradial spines glassy, conical, splay-

pointed, in webbed tufts.

Distribution. Sudanese Red Sea.

Etymology. Named in recognition of the research on the aster-

inids of the eastern Mediterranean Sea and Red Sea by Dr Yair

Achituv of the Bar Ilan University in Israel.

Remarks. The carinal plate arrangement and oral structure are

consistent with this unique asterinid combination in the

Ailsastra type species A. paulayi. The single specimen was not

dissected. An absence of superambulacral and superactinal

plates was not confirmed. Molecular data are not available for

A. achituvi.

Ailsastra amezianeae sp. nov.

Figures 1, 2b, 3b, 4d-f

Material examined. Holotype (dry). NW Madagascar, Nossi Be,

G. Cherbonnier, 6 Oct 1959, MNHN EcAs 11845.

Paratype (dry, part dissected). Type locality, G. Cherbonnier, 2 Dec

1959, MNHN Ec As 11846.

Diagnosis. Ailsastra with, at R = 10 mm, discrete, short, broad

rays, R/r about 1.5; 8 longitudinal series of papulae across mid-

ray; abactinal spinelets stout, short, up to about 0.10 mm long;

inferomarginal spinelets twice as long as superomarginal

spinelets; up to 4 actinal interradial spines per plate.

Description. Integument evident; 5 rays, R up to 10 mm; rays discrete,

short, wide at base, broad, tapered to narrow rounded end, R/r about

1.5; single madreporite, not fissiparous; abactinal gonopores. Carinal

series of plates broadly visible, not below adcarinal plates, doubly

papulate, other abactinal ray plates sometimes slightly incurved proxi-

mally for single papula, 3 longitudinal series of papulae on each upper

side of rays, 8 longitudinal series of papulae at mid-ray; disc close to

regularly defined by 5 radial, 5 interradial plates; spinelets glassy,

stout, splay -pointed subsacciform, up to about 16 spinelets in double or

single series across proximal edge of plates, up to about 0.10 mm long,

up to 6 in subpaxilliform tuft on distal interradial plates, proximal

and distal spinelets subequal; up to about 6 short stout sacciform super-

omarginal spinelets per plate, up to about 16 longer sacciform

184

P. Mark O’Loughlin and Francis W.E. Rowe

Figure 2. Abactinal view of holotypes of species of Ailsastra gen. nov. (photographed at same magnification), a, Ailsastra achituvi sp. nov.,

R = 10 mm, NHM 1951.5.7.12; b, Ailsastra amezianeae sp. nov., R =10 mm, MNHN EcAs 11845; c, Ailsastra booneni sp. nov., R = 9 mm,

UF 1938; d, Ailsastra eleaumei sp. nov., R = 7.5 mm, MNHN EcAs 11843; e, Ailsastra heteractis (H.L. Clark, 1938), R = 7 mm, MCZ 3258;

f, Ailsastra paulayi sp. nov., R = 16 mm, UF 1815.

A new asterinid genus

185

Figure 3. Actinal view of holotypes of species of Ailsastra gen. nov. a, Ailsastra achituvi sp. nov., R =10 mm, NHM 1951.5.7.12; b, Ailsastra

amezianeae sp. nov., R = 10 mm, MNHN EcAs 11845; c, Ailsastra booneni sp. nov., R = 9 mm, UF 1938; d, Ailsastra eleaumei sp. nov., R = 7.5

mm, MNHN EcAs 11843; e, Ailsastra heteractis (H.L. Clark, 1938), R = 7 mm, MCZ 3258; f, Ailsastra paulayi sp. nov., R = 16 mm, UF 1815.

186

P. Mark O’Loughlin and Francis W.E. Rowe

Figure 4. a-c, Ailsastra achituvi sp. nov., NHM 1951.5.7.12: a, low carinal series of plates; b, abactinal spinelets, glassy convexities on plates; c,

small oral plates, d-f, Ailsastra amezianeae sp. nov.: d, abactinal interradial plates removed to show absence of superambulacral (arrow) and

superactinal plates (paratype, MNHN EcAs 11846); e, abactinal spinelets (holotype, MNHN EcAs 11845); f, small oral plates (paratype, MNHN

EcAs 11846). g-i, Ailsastra booneni sp. nov., UF 1938: g, two proximal carinal plates with longitudinal series of spinelets (arrows), breaks in

enveloping integument which obscures plates, abactinal spinelets; h, distal interradius, inferomarginal spinelets, absence of superomarginal

spinelets; i, small oral plates, proximal interradial single pyramidal spines.

inferomarginal spinelets per plate, up to twice as long as supero-

marginal spinelets. Actinal spines per plate: oral 2-5; suboral 0; furrow

(proximal) 3; subambulacral 2-3; adradial actinal 2^1; interradial 2-4;

interradial spines stout, subsacciform, pointed, in webbed tufts.

Distribution. NW Madagascar, Nossi Be.

Etymology. Named in recognition of Dr Nadia Ameziane of

the Museum National d’Histoire Naturelle, Paris, who has

graciously assisted the authors with information and the loan of

materials for echinoderm research.

Remarks. The carinal plate arrangement, oral structure, and

absence of superambulacral and superactinal plates, are con-

sistent with this unique asterinid combination in the Ailsastra

type species A. paulayi. Molecular data are not available for

A. amezianeae.

Ailsastra booneni sp. nov.

Figures 1, 2c, 3c, 4g-i

Material examined. Holotype (alcohol). Indonesia, N Sulawesi I., Bay

of Tomini, off Salongon, Mulibagu, 0 o 21'31"S, 124°3'14"E, 2-15 m,

G. Paulay, 18 Sep 1999, UF 1938.

Diagnosis. Ailsastra with, at R = 9 mm, discrete, narrow rays,

R/r up to 1.9; 6 longitudinal series of papulae across mid-ray;

integument obscuring abactinal and actinal plates; abactinal

spinelets granuliform to sacciform conical, up to about 0.20

mm long; lacking superomarginal spinelets; proximal actinal

large pyramidal spines present on some interradii; up to 2

actinal interradial spines per plate.

Description. Integument obscures abactinal and actinal plates; 5 rays,

asymmetrical form, R up to 9 mm; rays discrete, narrow, tapered to

A new asterinid genus

187

Figure 5. a-c, Ailsastra eleaumei sp. nov., MNHN EcAs 11843: a, low carinal series of plates with bowl-shaped spinelet arrangements;

b, bowl-shaped arrangements of spinelets on disc plates; c, interradial abactinal gonopores (arrows), d-f, Ailsastra heteractis (H.L. Clark, 1938),

MCZ 3258: d, lateral ray abactinal spinelets; e, inferomarginal spinelets; f, small oral plates, proximal actinal pores (arrows), g-i , Ailsastra paulayi

sp. nov., UF 1815: g, abactinal spinelets; h, abactinal interradial plates removed to show absence of superambulacral plates (arrow); i, distal

abactinal interradius.

narrow rounded end, R/r up to 1.9; single large madreporite, not fissi-

parous; abactinal paired or single interradial gonopores. Carinal series

of plates extend to end of ray, below adcarinal plates creating furrow,

each plate doubly papulate, other abactinal ray plates obscured by

integument, crescentiform series of spinelets pointing through integu-

ment, single papula per plate, 2 longitudinal series of papulae on each

upper side of rays, 6 longitudinal series of papulae across mid-ray; disc

distinctly bordered by 5 radial, 5 interradial plates; spinelets glassy,

stout, granuliform on upper ray, sacciform conical in interradius, up to

about 0.20 mm long; up to 5 spinelets in longitudinal series on

proximal carinal plates, 1-2 spinelets on carinal plates in mid ray, up

to about 8 spinelets in predominantly single or double series across

raised proximal edge of plates; single spinelets in mid- interradius;

superomarginal plates lacking spinelets; up to about 8 long, thin, glassy

spinelets per inferomarginal plate. Actinal plates obscured by integu-

ment, spines in oblique series, variably webbed by integument. Actinal

spines per plate: oral 3; suboral predominantly 1 (0-2), some longer

than proximal oral spines, continuous as series with oral spines; furrow

3-2; subambulacral 2-1; adradial actinal 1; interradial 2-1; interradial

spines digitiform, partly obscured by integument.

Colour (live). Mottled dark mauve and white, disc reddish-

brown (holotype photo by G. Paulay: GP Sula-664).

Distribution. Indonesia, N Sulawesi, 2-15 m.

Etymology. Named in appreciation of the assistance of Ben

Boonen, who has graciously and skilfully prepared the images

for this and other echinoderm papers.

Remarks. The carinal plate arrangement and oral structure are

consistent with the patterns shown in the type species, Ailsastra

paulayi. The holotype was not dissected and the absence of

188

P. Mark O’Loughlin and Francis W.E. Rowe

superambulacral and superactinal plates was not confirmed.

Molecular phylogenetic analysis by Jon Waters (pers. comm.;

paper in preparation) places this species in a clade with the type

species, Ailsastra paulayi (below).

Ailsastra eleaumei sp. nov.

Figures 1, 2d, 3d, 5a-c

Material examined. Holotype (dry). Mauritius (tie Maurice), MAU 74-

12, Peyrot-Clausade, 1974, MNHN EcAs 11843.

Paratype (dry, part dissected). Type locality, MAU 74-13, MNHN

EcAs 11844.

Diagnosis. Ailsastra with, at R = 7.5 mm, discrete, short,

strongly tapered rays, R/r about 1.6; frequent bowl-shaped

arrangement of numerous, small, thin, webbed spinelets on abac-

tinal plates, up to 0.17 mm long; 4 longitudinal series of papu-

lae across mid-ray; up to 5 interradial actinal spines per plate;

longitudinal alignment of spines on some distal actinal plates.

Description. Integument not evident; rays 6 (holotype), 5 (paratype),

R up to 7.5 mm; rays discrete, short, strongly tapered, rounded dis-

tally, R/r about 1.6; single madreporite, not fissiparous; abactinal

gonopores. Carinal series of plates variably narrowly evident below

adcarinal plates, doubly papulate, other abactinal ray plates lacking

papular indentation, single longitudinal series of non-carinal papulae

on each upper side of rays, 4 longitudinal series of papulae across mid-

ray; disc not delineated; spinelets glassy, thin, subsacciform to conical,

finely splay-pointed, up to about 20 spinelets in 2 webbed series across

proximal edge of plate, frequently in bowl-shaped series around

periphery of projecting plate, up to about 0.17 mm long, up to 8 in sub-

paxilliform tuft on distal interradial plates; proximal and distal

spinelets subequal; up to about 10 webbed spinelets around periphery

of each superomarginal plate, up to about 16 longer spinelets on each

inferomarginal plate, most in peripheral webbed series on abactinal

surface of plate, few on actinal surface. Some non-plated areas

actinally. Actinal spines per plate: oral 4; suboral 0; furrow 3; sub-

ambulacral 3; adradial actinal 1-3; interradial 1-3, up to 5 distally,

sometimes in longitudinal alignment; interradial spines sacciform,

thin, pointed.

Distribution. Mauritius (lie Maurice).

Etymology. Named in recognition of Marc Eleaume of

the Museum National d’Histoire Naturelle, Paris, who has

graciously assisted the authors with information and the loan of

materials for echinoderm research.

Remarks. The carinal plate arrangement, oral structure, and

absence of superambulacral and superactinal plates are consis-

tent with the type species. Variable ray number is common

amongst asterinid species, and the predominant ray number for

A. eleaumei is not evident. Molecular data are not available for

A. eleaumei.

Ailsastra heteractis (H.L. Clark)

Figures 1, 2e, 3e, 5d-f

Asterina heteractis H.L. Clark, 1938: 152-153, pi. 22 fig. 5. — H.L.

Clark, 1946: 130, 133.— Rowe (in Rowe and Gates), 1995: 34-35.

Aquilonastra heteractis. — O’Loughlin and Waters, 2004: 11, 13, 15.

Material examined. Holotype (dry). Australia, Lord Howe I., Neds

Beach, under rock fragment, littoral, K. Birmingham, Apr 1932, MCZ

3258.

Paratypes (dry). Lord Howe, Neds Beach, MCZ 3259 (1); dredged

from lagoon near Goat I., MCZ 3260 (2).

Other material (dry). Queensland, Townsville, AM J9541 (2); reef

flat, Heron I., AM J19449 (1).

Diagnosis. Ailsastra with 6-7 discrete high rays with narrow

bases, R/r about 2.0; multiple inconspicuous madreporites; fis-

siparous habit; abactinal gonopores; spinelets up to 0.15 mm

long; large proximal actinal interradial pores; up to 4 spines per

actinal interradial plate.

Description. 6-7 rays, R up to 7 mm; integument evident; rays dis-

crete, narrow base, long, blunt distally, high convex abactinally, R/r

about 2.0; multiple inconspicuous madreporites; fissiparous, ray

lengths frequently unequal; abactinal paired interradial proximal gono-

pores. Carinal series of plates broadly visible, slightly below adcarinal

plates, doubly papulate for most of ray, not proximally, other ray plates

slightly indented for single large papula, single longitudinal series of

papulae on each upper side of rays, 4 longitudinal series across mid-

ray; papular spaces large; disc not delineated; spinelets glassy, stout,

conical to subsacciform, splay-pointed, up to about 8 per plate, in sin-

gle or double series across plates or in apical tufts, up to 0.15 mm long;

superomarginal plates with up to 4 spinelets; inferomarginal plates

with up to 10 longer, splay-pointed webbed spinelets. Single large

proximal actinal interradial pores. Actinal spines per plate: oral 4-6;

suboral 0-3; furrow (proximal) 3-4; subambulacral 3-4; adradial acti-

nal 2-4; interradial 1-3; interradial spines subsacciform, elongate,

pointed.

Colour (live). Salmon pink to light orange to nearly white

abactinally, whitish actinally (H.L. Clark, 1938).

Distribution. Lord Howe I., NE Australia, littoral to shallow

sublittoral.

Remarks. The presence of large single proximal actinal pores is

unique amongst the asterinids. There was inadequate material

to confirm by dissection the nature of the pores. They were

observed only on the holotype. The distal oral spines are close

to the suboral spines on the characteristic small oral plates, and

sometimes appear to be in continuous series. O’Loughlin

and Waters (2004) assigned Asterina heteractis to Aquilonastra

O’Loughlin, 2004, but noted some exceptional characters.

A. heteractis has the carinal plate arrangement and small oral

plates of Ailsastra paulayi, the type species of Ailsastra, and is

reassigned to this new genus. No specimen was dissected to

confirm an absence of superambulacral and superactinal plates.

Ailsastra paulayi sp. nov.

Figures 1, 2f, 3f, 5g-i

Material examined. Holotype (in alcohol, part dissected). Indonesia,

Sulawesi I., Pulau Talatakoh, 0°28'2"S, 122°8’22"E, 1-10 m,

G. Paulay, 19 Sep 1999, UF 1815.

Paratype. Sulawesi I., 1-2°S, 121° E, 1-16 m, G. Paulay, 26 Sep

1999, UF 910.

Diagnosis. Ailsastra with, at R = 16 mm, discrete, elongate,

narrow, slightly tapered rays, R/r about 2.7; 8 longitudinal

series of papulae across mid-ray; disc regularly bordered; long,

thin, spinelets, up to about 0.25 mm long; proximal and distal

interradial spinelets subequal; superomarginal and infero-

marginal spinelets subequal; up to 7 actinal interradial spines

per plate.

A new asterinid genus

189

Description (holotype). Morphological characters of Ailsastra\ integu-

ment evident; 5 rays, R up to 16 mm; rays discrete, narrow at base,

elongate, low, slightly tapered, rounded distally, R/r about 2.7; single

madreporite, not fissiparous; abactinal gonopores. Carinal series of

plates broadly visible, plates not noticeably below adcarinal plates,

doubly papulate, other ray plates not indented for single papula, 3 lon-

gitudinal series of papulae on each upper side of rays, 8 longitudinal

series across mid-ray; disc regular, defined by 5 radial, 5 interradial

plates; spinelets glassy, thin, subsacciform to conical, each with few

small points distally, up to about 20 spinelets in single or double

webbed series across proximal edge of plate, or in splayed tufts on

plates, spinelets up to about 0.25 mm long on proximal ray, up to about

10 in subpaxilliform splayed tuft on distal interradial plates; proximal

and distal interradial spinelets subequal in length; superomarginal and

inferomarginal spinelets subequal in length, up to about 6 per supero-

marginal plate, about 14 per inferomarginal plate. Actinal spines per

plate: oral 5-6; suboral 0-1; furrow (proximal) 5; subambulacral 4;

adradial actinal 3; interradial 3-7; interradial spines glassy, thin,

subsacciform to conical, splay-pointed, in webbed tufts.

Colour (live). Mottled red and orange (holotype photo by

G. Paulay: GP 670:24).

Distribution. Indonesia, Sulawesi; 1-16 m.

Etymology. Named in recognition of Dr Gustav Paulay of the

University of Florida, who collected this material and who has

facilitated extensive collecting of asterinid material throughout

the Indo-Pacific region.

Remarks. The paratype is small (R = 7 mm), and the rays are

not as narrow as those of the holotype. Molecular phylogenetic

data from Jon Waters (pers. comm.) places the two specimens

of A. paulayi in a highly divergent but strongly monophyletic

clade within a molecular phylogeny of Asterinidae (Jon Waters,

pers. comm.; paper in preparation).

Acknowledgments

We are grateful for the assistance provided by Nadia Ameziane

and Marc Eleaume (MNHN, loans, data), Penny Berents (AM,

loan), Ben Boonen (figure preparation), Andrew Cabrinovic

and Sheila Halsey (NHM, loans), Fred Collier and Bob

Woollacott (MCZ, loans), Ruth Doran (NMV, map), Gustav

Paulay (UF, loans, photo), Chris Rowley (NMV, photography),

and Jon Waters (University of Otago, molecular phylogeny

data). We are appreciative of the comments offered by the

referees, Alan Dartnall and Tim O’Hara.

References

Clark, A.M. 1983. Notes on Atlantic Asteroidea. 3. Families

Ganeriidae and Asterinidae. Bulletin of the British Museum of

Natural History (Zoology) 47: 19-51.

Clark, A.M., and Rowe, F.W.E. 1971. Monograph of shallow-water

Indo-west Pacific Echinoderms. British Museum (Natural History):

London. 234 pp.

Clark, H.L. 1938. Echinoderms from Australia. An account of collec-

tions made in 1929 and 1932. Memoirs of the Museum of

Comparative Zoology at Harvard College 55: 1-596, 28 pis, 63

figs.

Clark, H.L. 1946. The echinoderm fauna of Australia. Its composition

and its origin. Carnegie Institution of Washington Publication 566:

1-567.

O’Loughlin, P.M., and Waters, J.M. 2004. A molecular and morpho-

logical revision of genera of Asterinidae (Echinodermata:

Asteroidea). Memoirs of Museum Victoria 61(1): 1^40.

Rowe, F.W.E., and Gates, J. 1995. Echinodermata. In: Wells, A. (ed.),

Zoological Catalogue of Australia 33: i-xiii, 1-510. CSIRO:

Melbourne.

Rowe, F.W. E., and Richmond, M.D. 2004. A preliminary account of

the shallow- water echinoderms of Rodrigues, Mauritius, western

Indian Ocean. Journal of Natural History 38: 3273-3314.

Verrill, A.E. 1913. Revision of the genera of starfishes of the family

Asterininae. American Journal of Science 35 (4): 477-485.

Memoirs of Museum Victoria 62(2): 191-212 (2005)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A review of soles of the genus Aseraggodes from the South Pacific, with

descriptions of seven new species and a diagnosis of Synclidopus.

John E. Randall

Bishop Museum, 1525 Bernice St. Honolulu, Hawai’i 96817-2704, USA (jackr@hawaii.rr.com)

Abstract Randall, J.E. 2005. A review of soles of the genus Aseraggodes from the South Pacific, with descriptions of seven new

species and a diagnosis of Synclidopus. Memoirs of Museum Victoria 62(2): 191-212

The soleid genus Parachirus Matsubara and Ochiai is referred to the synonymy of Aseraggodes Kaup. Aseraggodes

persimilis (Gunther) and A. ocellatus Weed are reclassified in the genus Pardachirus Gunther. Synclidopus macleayanus

(Ramsay) from Queensland and NSW is redescribed. A diagnosis is given for Aseraggodes, and a key and species

accounts provided for the following 12 species of the genus from islands of Oceania in the South Pacific and the east

coast of Australia: A. auroculus, sp. nov. from the Society Islands; A. bahamondei from Easter Island and Lord Howe

Island; A. cyclurus, sp. nov. from the Society Islands; A. lateralis, sp. nov. from the Marquesas Islands; A. lenisquamis,

sp. nov. from NSW; A. magnoculus sp. nov. from New Caledonia; A. melanostictus (Peters) from 73 m off Bougainville

and a first record for the Great Barrier Reef from 115 m; A. nigrocirratus, sp. nov. from NSW; A. normani Chabanaud

from southern Queensland and NSW; A. pelvicus, sp. nov. from the Great Barrier Reef; A. ramsaii (Ogilby) from Lord

Howe Island, with a first record for New Caledonia; and A. whitakeri Woods as first records from the Caroline Islands,

Coral Sea, New Caledonia, Fiji, American Samoa, Phoenix Islands, and Society Islands. Presumed hybrids of

A. lenisquamis and A. nigrocirratus were found in two Australian Museum lots of specimens from NSW.

Keywords Taxonomy, soleid fishes, Aseraggodes, Synclidopus macleayanus

Introduction

With the advent of scuba diving and the use of the ichthyocide

rotenone, ichthyologists have made major collections of shore

fishes, particularly in the tropical and subtropical regions of the

world. The coral reefs and adjacent habitats have yielded a

multitude of new species of fishes. Taxonomic research tended

to focus on the colourful species in well-known genera with

ample material. Collections rarely resulted in more than one or

two specimens of soles of any species of Aseraggodes Kaup.

Small and usually not distinctly coloured, they have not received

the scientific attention they require. As noted by Randall (1996),

some specimens of Aseraggodes still remain on museum

shelves identified only to genus or the family Soleidae.

Nevertheless, the genus Aseraggodes is second only to

Solea in the number of species of the family Soleidae.

Eschmeyer’s Catalog of Fishes (updated to 14 January 2005

at web site www.calacademy.org/research/ichthyology/catalog/

fishcatsearch.html) lists 26 Indo-Pacific species as valid for the

genus and one, A. herrei Seale, from the Galapagos Islands. Of

the 26 species of Eschmeyer’s list, A. filiger Weber, with its

slender body and filamentous first dorsal ray, should remain

in the monotypic genus Coryphillus Chabanaud, 1932.

Aseraggodes persimilis (Gunther) from New Britain and

A. ocellatus Weed from Sri Lanka, both described as having a

pore at the base of most of the dorsal and anal rays, are here

reclassified as synonyms of Pardachirus pavoninus

(Lacepede). The species of Aseraggodes lack these distinctive

externally visible pores.

Clark and George (1979) showed that basal dorsal- and

anal-ray pores in species of Pardachirus are the release sites for

a powerful toxin when these soles are threatened. Randall and

Melendez (1987) reported what seems to be a comparable skin

toxin in Aseraggodes bahamondei at Easter Island. They found

small pores beneath occasional scales toward the periphery of

the body on the ocular side, which they believe elicit the toxin.

Randall (2002) discovered that A. therese in the Hawaiian

Islands is unpalatable to the jack Caranx melampygus. He did

not examine specimens of therese for pores at that time, but

after long searching and with the help of staining, these were

later detected beneath a few scales on a large non-type

specimen. Finding the pores on small specimens or old

museum specimens is difficult. These pores were detected in

this study only in A. melanostictus and one of the new species,

but they are probably present in some other species as well.

Solea macleayana was described by Ramsay (1881) from

NSW. Ogilby (1916) illustrated the species, described it more

192

John E. Randall

fully, and reclassified it in Aseraggodes. He has been followed

by subsequent authors. Chabanaud (1943), however, selected

Solea macleayana as the type species of a new genus,

Synclidopus, one of eight new soleid genera briefly described

in the paper. The type species of Synclidopus is more fully

described here.

The classification of the species Aseraggodes has been dif-

ficult because of the paucity of specimens, ontogenetic

changes, variability in colour pattern (see Figs 3-5), and the

broad range in the counts of dorsal rays, anal rays, and lateral-

line scales. For example, Randall and Melendez (1987) found a

range of seven dorsal rays, eight anal rays, and 12 lateral-line

scales for 27 specimens of A. bahamondei, and Randall, 2002

reported 29 specimens of A. therese with a range of eight

dorsal rays, eight anal rays, and seven lateral-line scales.

Nevertheless, the combination of these three counts will often

be of diagnostic value to the species of the genus. The number

of vertebrae also may vary within a species, though usually

not more than two or three. Ochiai (1963) reported 34-37 verte-

brae for 67 specimens of the Japanese species A. kobensis

(Steindachner).

The species of Aseraggodes generally have a thin mem-

branous ridge along the dorsal and anal rays, often disappear-

ing on posterior rays, and many species have cirri along the

edges of these ridges, especially on anterior rays. In old

museum specimens or poorly preserved specimens, the ridges

may not be apparent, and the cirri often cannot be detected.

This limits the usefulness of these features as diagnostic

characters.

Twelve species of Aseraggodes have been reported from

eastern Australia and the islands of the Oceania. Peters (1877)

described A. melanostictus, from a specimen collected in 73 m

off Bougainville. Ogilby (1889) named A. ramsaii from one

specimen from Lord Howe Island. Norman (1926) reported

three specimens, 130-142 mm total length, from Queensland as

A. melanostictus. Chabanaud (1930a) realized that these were

not correctly identified and described a 135-mm one as a new

species, A. normani. Schultz (1943) reported a small speci-

men of the genus from Hull Island in the Phoenix Islands. He

wrote, “It may be a specimen of Aseraggodes melanostictus

(Peters).” Woods in Schultz and collaborators (1966) described

A. whitakeri and A. smith i as new species from the Marshall

Islands. He identified a specimen from Kwajalein Atoll,

Marshall Islands as Aseraggodes melanostictus “with uncer-

tainty.” Randall and Melendez (1987) named A. bahamondei

from Easter Island and Lord Howe Island. Randall (1996)

described A. borehami and A. therese, and Randall (2002)

added A. holcomi, all three from the Hawaiian Islands. Randall

and Bartsch (2005) determined that the Marshall Islands speci-

men identified as A. melanostictus by Woods is a new species,

A. heraldi, described A. firmisquamis from Palau, and reported

A. smithi from Palau. The present paper provides the descrip-

tions of seven new species of Aseraggodes from the South

Pacific, as well as range extensions for A. melanostictus,

A. ramsaii, and A. whitakeri.

Materials and methods. Type specimens of Aseraggodes have been

variously deposited at the Australian Museum, Sydney (AMS); Natural

History Museum, London (BMNH); Bishop Museum, Honolulu

(BPBM); California Academy of Sciences, San Francisco (CAS);

Museum Victoria, Melbourne (NMV); National Science Museum,

Tokyo (NSMT); Royal Ontario Museum, Toronto (ROM); and US

National Museum of Natural History, Washington, DC (USNM).

Standard length (SL) is measured horizontally from the front of the

upper lip to the base of the caudal fin (end of hypural plate). Body

depth is the maximum distance between the bases of the dorsal and

anal fins. Body width is the maximum thickness midlaterally between

the ocular and blind surfaces. Head length (HL) is measured from the

front of the upper lip to a vertical at the posterior end of the operculum.

Preorbital length is the distance from the front edge of the upper eye

directly forward to the most anterior edge of the head. Snout length is

taken from the front of the upper Up to the nearest edge of the upper

eye. Eye diameter is the greatest diameter of the lower eye (the dark

eyeball, not the surrounding cutaneous part). The interorbital width is

the vertical distance between horizontal lines at the lower edge of the

upper eye and upper edge of the lower eye (i.e. between the dark edges

of the two eyes). Upper jaw length is measured on the blind side from

the front of the upper lip to the rear edge of the maxilla (often too

difficult to determine the posterior end of the maxilla on the ocular

side). Caudal-peduncle depth is the least depth, or if the caudal

peduncle is absent, the depth is measured at the base of the caudal fin.

Caudal-peduncle length is the horizontal distance between verticals at

the rear base of the anal fin and the base of the caudal fin at its ventral

edge. Lengths of fin rays are measured from the ray base in a straight

line to the tip. Caudal-fin and pelvic-fin measurements are the length

of the longest ray.

Tables 1-3 provide the counts of the dorsal rays, anal rays, and

lateral-line scales, respectively. Proportional measurements of the

new species are given in Tables 4-11 as percentages of the standard

length. Measurements (ratios related to SL, head length, or body depth)

in the text are rounded to the nearest 0.05. Data in parentheses in the

descriptions refer to paratypes.

Lateral-line scales are counted from the base of the caudal fin to the

front of the straight part of the lateral line on the head (hence 5-15

scales anterior to the upper end of the gill opening). Scale counts above

and below the lateral line are the maximum number of scales in an

oblique row between the lateral line and the outer edge of the scaly

sheath at the base of the dorsal and anal fins, respectively.

Vertebral counts for soles are often given in two parts, the abdomi-

nal vertebrae, followed by the caudal vertebrae. There are ten abdom-

inal vertebrae in all the species of Aseraggodes examined (the count

includes the very small first vertebra overlooked by some authors), so

only the total count, which includes the urostyle, is given here, which

includes the urostyle.

Ochiai (1963) used the count of the number of dorsal pterygio-

phores (he called these interneural spines) associated with the first

three vertebrae (actually, four as he did not include the first very small

vertebra) as a taxonomic character. He is followed in the use of this

count.

Synclidopus Chabanaud, 1943

Synclidopus Chabanaud, 1943: 291.

Type species. Solea macleayana Ramsay, 1881.

Diagnosis. Dorsal rays 62-66; anal rays 49-53; caudal rays

18-20; pelvic rays 5, the fifth ray of ocular-side fin joined by

membrane to base of first anal ray; lateral-line scales 96-113;

lateral line extending forward on head to within an eye dia-

meter of upper eye; a second lateral line branching off dorsally

on head, about two eye diameters behind upper eye, angling

sharply posteriorly about 8 scale rows beneath naked part of

A review of soles of the genus Aseraggodes

193

dorsal fin, and continuing onto anterior body; body deep, the

depth 2.2-2.25 in SL; head short and obtuse, its length 5.25-5.5

in SL; eyes small, 8. 1-9.0 in HL; tubular anterior nostril short,

not reaching lower eye when laid back; rays of median fins

short; anus and genital papilla on blind side, adjacent to base of

first anal ray; no pore at base of dorsal and anal rays, and no

small pores detected beneath scales of ocular side; vertebrae

36-38; dorsal pterygiophores anterior to fourth neural spine 7;

unique colour pattern of many narrow dark bars on ocular side

of head and body.

Remarks. The type species of this monotypic genus was first

described in Solea Cuvier. Other authors such as Ogilby (1916),

Norman (1926), McCulloch (1929), Allen et al. (1976), and

Grant (1987) classified it in Aseraggodes Kaup. It is clearly

distinct at the generic level from Aseraggodes by having a sec-

ond lateral line on the ocular side of the head that continues

dorsoanteriorly on the body, the deepest body, shortest and

most obtuse head, smallest eyes, anus and genital papilla on the

blind side, highest number of lateral-line scales (96-113, com-

pared to 53-96 for species of Aseraggodes), 7 dorsal pterygio-

phores anterior to fourth neural spine (species of Aseraggodes

with 7-16; only one with 7 or 8), and the colour pattern

of narrow dark bars. Ogilby (1916) reported the maximum

size as 280 mm total length (largest species of Aseraggodes,

192 mm TL).

Synclidopus macleayanus (Ramsay, 1881)

Figure 1

Solea Macleayana Ramsay, 1881: 462.

Solea fluviatilis Ramsay, 1882: 111 (type locality, Hunter River,

NSW).

Material examined. NSW: Port Jackson, AMS 1.16278-01, 135 mm,

syntype of Solea macleayana. Eight miles from North Head,

Richmond River, R/V “Endeavour”, BMNH 1925.7.22.72, 112 mm.

Sydney, Hawkesbury River, Gentleman’s Halt, AMS 1.19951-003, 6:

72-105 mm. Off Sydney, 33°51'S, 151°18’E, 40-15 m, BPBM 39454,

149 mm. Lord Howe Island: AMS 1. 12664, 103 mm.

Type locality. Manly Beach, Port Jackson, NSW.

Remarks. Ramsay (1881) briefly described this species. He

wrote, “A number were taken in the net at Manly Beach,

September 11th, 1880, with Solea microcephala .” Only one

type specimen of Solea macleayana has been found, labeled as

a syntype at the Australian Museum. A more detailed descrip-

tion of the species was provided by Ogilby (1916: 127, pi. 15)

who had three specimens, 154-192 mm in total length. He

described the colour as “Lavender grey, with from 32-36 nar-

row brown cross-bars, which are usually rather wider than the

interspaces, and of which 6 or 7 are on the head and 1 or 2 on

the base of the caudal fin; . . .” He placed Solea fluviatilis

Ramsay, described from one 76-mm specimen from freshwater

in Hunter River, in synonymy, adding that Ramsay was “pos-

sibly misled by the different character of the element in which

it was found.” Ogilby summarized the reproductive cycle.

Adults spend the winter months in moderately deep water,

gradually move to shallower water in spring. On reaching

Figure 1. Synclidopus macleayanus , BPBM 39454, 149 mm SL, off

Sydney, NSW.

maturity during summer months, they collect in the vicinity of

river mouths, where they spawn. “The young fishes, as soon as

the yolk-sac is absorbed, make their way into the estuaries and

gradually work up these even to far beyond the limit of the tide,

as we know from the Hunter River example . . .” Under the

heading Uses, Ogilby wrote, “A delicious pan-fish, fully equal

in flavor to its famous European relative, Solea solea.”

Norman (1926) gave the distribution of this species as

“Coasts of New South Wales and southern Queensland (from

southern NSW to Caloundra).” Allen et al. (1976: 437)

reported a specimen from Lord Howe Island.

Grant (1987) noted that this sole is adept at burrowing

beneath the sand and “actually swims beneath the sand.” He

summarized the food habits as “shellfish and worms that live

on and in the substrate.”

Aseraggodes Kaup, 1858

Aseraggodes Kaup, 1858: 103.

Parachirus Matsubara and Ochiai 1963: 93 (type species,

Parachirus xenichus Matsubara and Ochiai, 1963 opening by original

designation and monotypy).

Type species. Aseraggodes guttulatus Kaup, 1858, by subsequent

designation of Jordan and Evermann, 1898.

Diagnosis. Dorsal rays 58-79; anal rays 39-61; caudal rays

typically 18 (usually 14-16 branched in adults); no pectoral

fins; pelvic rays normally 5; lateral-line scales 39-96 (includ-

ing those extending onto head); no gill rakers; abdominal

vertebrae 10 (including the first vertebra, not counted by some

authors, very narrow, the neural spine slender and short, not

extending above cranium); total vertebrae 33-40; first two

dorsal pterygiophores joined to a thicker bone (termed the

erisma and counted as the first pterygiophore, though branched

distally to support the first two dorsal rays), its origin between

second neural spine and cranium, 7-16 dorsal pterygiophores

anterior to fourth neural spine; body an elongate oval, the depth

2.0-2.8 in SL, and very thin; eyes on right side, elevated, sep-

arated by a narrow scaled space; upper eye in advance of lower

eye (rarely directly above); caudal peduncle, if present, very

short; scales small, ctenoid (except cycloid lateral-line scales);

a straight lateral line midlaterally on both sides, with a short

anterodorsal branch on blind side; no prominent pore at base of

dorsal and anal rays; gill membranes united, free from isthmus,

194

John E. Randall

the lower part of head scaled over from ocular to blind side;

mouth ventral and small; jaws strongly curved; a band of villi-

form teeth on blind side of jaws; two nostrils on each side, the

anterior nostril of ocular side tubular, but not longer than eye

diameter; posterior nostril of ocular side a narrow opening in

labial groove before lower eye; dorsal fin originating anterior-

ly on snout, the first ray not prolonged; caudal fin rounded to

slightly pointed, not broadly connected by membrane to

dorsal and anal fins; pelvic fins on ventral edge of body, close

together anteriorly, adjacent or with ocular- side fin slightly

anterior; anus anterior or ventroanterior to first anal ray. Sciatic

part of urohyal forming an angle of about 60-85° to horizontal

main part of bone.

Remarks. Kaup (1858: 103) briefly described Aseraggodes

guttulatus as a new genus and species, but gave no locality for

the holotype, as noted by Gunther (1862: 477). Chabanaud

(1930b) revised the 15 species of the genus then known. He

mistakenly placed A. kaianus (Gunther) in the synonymy of

A. guttulatus and gave two localities, Kei Islands (Gunther’s

type locality of kaianus) and the Maidive Islands. Desoutter et

al. (2001) resolved the locality problem by finding the holotype

in the Museum National d’Histoire Naturelle (MNHN 1246,

79.0 mm SL). Kaup’s original label indicated the specimen as

the type and the collection locality as Bourbon (= Reunion).

Kaup wrote in his description of Aseraggodes guttulatus that

the height of the body is half the total length. Gunther (1862:

477) questioned this in a footnote. Martine Desoutter (pers.

comm.) measured the height of the body of the holotype as 3.1

in total length. She confirmed Kaup’s counts of the dorsal and

anal rays as 64 and 42, respectively. She also provided the

lateral-line scale count of 84 and an x-ray, which indicates a

vertebral count of 34, and 14 dorsal pterygiophores before the

fourth neural spine.

Matsubara and Ochiai (1963) described Parachirus xenicus

as a new genus and species of sole from Japan. In a review of

the Soleidae and Cynoglossidae of Japanese waters, Ochiai

(1963) separated Parachirus from Aseraggodes by having the

dorsal, anal, and pelvic fins slightly branched (as opposed to

not branched in Aseraggodes ), the tubular anterior nostril

reaching the edge of the lower eye (not reaching in

Aseraggodes ), vertebrae 32-33, revised in this paper to 33-34

because the tiny first vertebra is now included in the vertebral

count (vs 37-39 vertebrae in Aseraggodes ), about 15 inter-

neural spines (= dorsal pterygiophores) associated with the

anterior 4 neural spines, and the pelvic fins attached by mem-

brane to the genital papilla. In a generally favorable review of

Ochiai’s publication, Hubbs (1967) pointed out its limitation

from dealing mainly with Japanese species. Chapleau (1989)

made a study of the anterior dorsal pterygiophores, erisma, and

neural spines of 41 species of 26 genera of soleid fishes. He

recognized Parachirus as a valid genus; however, he included

only four species of Aseraggodes in his study.

This study of Aseraggodes has shown that the dorsal, anal

and pelvic fins may be simple or branched (the young of

those with branched rays have unbranched rays); the tubular

anterior nostril often reaches the edge of the lower eye; the

vertebrae vary from 33 to 40; the dorsal pterygiophores anter-

ior to the fourth neural spine vary from 7 to 15; and the

pelvic fins may be attached by membrane to the genital papilla

(as in A. normani). Therefore, Parachirus is a synonym of

Aseraggodes.

Key to species of Aseraggodes of the South Pacific

1. Caudal peduncle present, though very short (7.2-10.8 in

HL) 2

— No caudal peduncle (rear base of anal fin below or

posterior to base of lowermost caudal ray) 5

2. Dorsal and anal rays short, the longest dorsal ray 1.9-2.15

in HL; dorsal and anal rays unbranched (young to adults);

vertebrae 39-40; dorsal pterygiophores (including erisma)

anterior to fourth neural spine 7-8; maximum size 156 mm

SL (Easter Island and Lord Howe Island) A. bahamondei

— Dorsal and anal rays not short, the longest dorsal ray

1.25-1.8 in HL; dorsal and anal rays of adults branched;

vertebrae 36-38; dorsal pterygiophores anterior to fourth

neural spine 13-15; largest specimen, 67.5 mm 3

3. Head large, its length 4.1-4.35 in SL; body slender, the

depth 2.55-2.75 in SL; largest specimen, 43 mm SL

(Micronesia and New Caledonia to Society Islands) ....

A. whitakeri

— Head not large, its length 4.45-4.75 in SL; body not

slender, the depth 2. 4-2.6 in SL; attains at least 63 mm

SL 4

4. Longest dorsal ray 1.25 in HL; length of caudal fin 3.4 in

SL; HL 4.75 in SL; pelvic fins long, 1.6 in HL; lateral-line

scales 81; edge of membranous ridge of anterior dorsal

rays with a row of small tubercle-like papillae, many

ending in a tiny cirrus (one specimen, 67.5 mm SL, Swain

Reefs, Great Barrier Reef) A. pelvicus

— Longest dorsal ray 1.65-1.7 in HL; length of caudal fin

3.85-3.95 in SL; HL 4.45-4.5 in SL; pelvic fins not long,

2.3-2.35 in HL; lateral-line scales 86-88; edge of

membranous ridge of anterior dorsal rays without a row of

papillae or cirri (Lord Howe Island and New Caledonia)

A. ramsaii

5. Lateral line of ocular side with 3 branches on head; mem-

brane from last rays of pelvic fins joined to genital papilla

(NSW and southern Queensland) A. normani

— Lateral line of ocular side without branches on head;

no membrane linking last ray of pelvic fins to genital

papilla 6

6. Dorsal, anal, and pelvic rays unbranched; lateral-line

scales 78-79 (2 specimens, 74-86.5 mm SL, Bougainville

and Great Barrier Reef, 73-115 m) .... A. melanostictus

— Dorsal, anal, and pelvic rays of adults branched; lateral-

line scales 61-73 7

7. Lateral-line scales 61-68; dorsal rays 62-70; anal rays

46-52; dorsal pterygiophores anterior to fourth neural

spine 8-9 8

— Lateral-line scales 69-73; dorsal rays 67-77; anal rays

49-57; dorsal pterygiophores before fourth neural spine

10-12 9

8. Surface of scales smooth, the posterior edge somewhat

pointed, with only the tips of cteni visible; snout length

2.45-2.5 in HL; caudal-peduncle depth 1.25-1.45 in HL;

A review of soles of the genus Aseraggodes

195

caudal-fin length 4.6-5.05 in SL (NSW) . A. lenisquamis

— Surface of scales with texture, the posterior edge rounded,

the cteni strongly projecting; snout length 2. 8-3.0 in HL;

caudal-peduncle depth 1.45-1.75 in HL; caudal-fin length

3.95—4.8 in SL (NSW) A. nigrocirratus

9. Eye large, 3.95-4.2 in HL; caudal fin long, 3.9-3.95 in SL;

longest dorsal ray 1.4 in HL (New Caledonia)

A. magnoculus

— Eye not as large, 4.55-7.0 in HL; caudal fin not long,

44-5.3 in SL; longest dorsal ray 1.45-1.8 in HL .... 10

10. Anal rays 58-59; dorsal rays 78-83; longest dorsal ray 1.8

in HL; caudal fin short, 4.7-5. 3 in SL (3 specimens,

Marquesas Islands) A. lateralis

— Anal rays 53-57; dorsal rays 68-77; longest dorsal ray

1.45-1.7 in HL; caudal fin not short, 44-4.8 in SL . . 11

1 1 . Dorsal rays 74-77 ; anal rays 56-57 ; head large, the length

3.85-3.95 in SL; body depth 2.45-2.55 in SL; caudal fin

4.4-4.45 in SL (Society Islands) A. auroculus

— Dorsal rays 68-71; anal rays 53; head not large, the length

4.65-4.95 in SL; body depth 2.25-2.4 in SL; caudal fin

4.7-4. 8 in SL (Society Islands) A. cyclurus

Aseraggodes auroculus sp. nov.

Figure 2, Tables 1-4

Holotype. ROM 61358, 35.5 mm, Society Islands, Moorea, off NW

coast, 17 o 31'0"S, 149°55'30"E, reef slope of coral rubble, with some

live coral (including a few large heads of Pontes), 18-24 m, rotenone,

Figure 2. Holotype of Aseraggodes auroculus, ROM 61358, 35.5 mm,

Moorea, Society Islands (R. Winterbottom).

R. Winterbottom and R. Mooi, 10 Dec 1989.

Paratypes. ROM 61357, 34.0 mm, Society Islands, Moorea, W side

of pass off Maharepa about middle of its length, 17°29'24"S,

149°48'0"W, 15-18 m, steep slope with coral rubble, sand, and a 3-m

wall, rotenone, R. Winterbottom and R. Mooi, 5 Dec 1989; BPBM

39690, 30.4 mm and USNM 381623, 29.8 mm, same data as holotype.

Diagnosis. Dorsal rays 74-77; anal rays 56-57; most dorsal

and anal rays double branched; lateral-line scales 69-73,

including 6-7 anterior to a vertical at upper end of gill opening;

vertebrae 37-38; dorsal pterygiophores anterior to fourth

neural spine 12; body depth 2.45-2.6 in SL; HL 3.85-3.95 in

SL; eye diameter 4.55-4.9 in HL; upper eye overlapping a

anterior two-thirds to three-fourths of lower eye; interorbital

Table 1. Dorsal Rays of South Pacific Species of Aseraggodes

A. auroculus

A. bahamondei

A. cyclurus

A. lateralis

A. lenisquamis 1

A. magnoculus

A. melanostictus

A. nigrocirratus

A. normani

A. pelvicus

A. ramsaii

A. whitakeri

Table 2. Anal rays of South Pacific species of Aseraggodes

48 49

A. auroculus

A. bahamondei

A. cyclurus

A. lateralis

A. lenisquamis

A. magnoculus

A. melanostictus

A. nigrocirratus

2 1

A. normani

A. pelvicus

A. ramsaii

A. whitakeri

1 1

196

John E. Randall

Table 3. Lateral-line scales of South Pacific species of Aseraggodes

59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88

A. auroculus

A. bahamondei

1 2 1

2 1 2 3 4 3

5 14 3 11

A. cyclurus

A. lateralis

1 1 1

1 1

A. lenisquamis

2 2 12 13 2

A. magnoculus

1 1

A. melanostictus

A. nigrocirratus

12 12 12

1 1

A. normani

A. pelvicus

1 2 3 1

A. ramsaii

1 1 1

A. whitakeri

112 3

112 1 1

Table 4. Proportional measurements of type specimens of Aseraggodes

auroculus as percentages of standard length

Holotype

Paratypes

ROM

61358

USNM

381623

BPBM

39690

ROM

61357

Standard Length (mm)

35.5

29.8

30.4

34.0

Body depth

38.1

40.7

39.6

39.3

Body width

7.1

8.4

7.3

Head length

25.3

25.4

26.0

25.8

Snout length

8.6

9.2

8.2

8.8

Preorbital length

7.1

7.6

7.5

7.8

Eye diameter

5.5

5.6

5.3

5.7

Interorbital width

1.4

1.5

1.0

1.1

Upper-jaw length

8.7

8.8

8.9

8.8

Caudal-base depth

13.4

14.1

13.2

13.8

Predorsal length

6.2

6.1

6.3

5.9

Preanal length

30.6

29.4

29.6

30.0

Prepelvic length

23.8

23.2

24.1

First dorsal ray

6.7

6.9

6.8

6.5

Longest dorsal ray

15.2

16.4

15.8

15.0

First anal ray

7.0

6.4

6.7

6.6

Longest anal ray

15.4

16.7

broken

15.1

Caudal-fin length

22.6

broken

22.7

22.6

Pelvic -fin length

10.2

10.6

9.7

9.5

space narrow, the vertical distance separating eyes about one-

fourth to one-sixth eye diameter; no caudal peduncle; short

fleshy cirri on ventral edge of head; lateral line aligned with

ventral part of upper eye; longest dorsal ray 1.55-1.7 in HL;

caudal fin rounded, its length 4.4-4.55 in SL; pelvic fins

2.4-2.7 in HL, the tip of longest ray reaching base of second or

third anal ray; colour of ocular side in alcohol pale yellowish

brown with 3 rows of large irregular blackish blotches, one

dorsal, one ventral, and one midlateral; a few small dark spots

on fin rays; blind side of body pale yellowish, the dark spots on

rays faint.

Description. Dorsal rays 75 (74-77); anal rays 56 (56-57); dorsal rays

branched except first 9 dorsal rays of holotype and first 19 of smallest

paratype; anal rays branched; caudal rays 18, the middle 16 of holotype

double-branched (middle 12-14 of paratypes double-branched); pelvic

rays 5, branched except first; lateral-line scales 69 (71-73), including

6-7 anterior to a vertical at upper end of gill opening; scales above

lateral line on ocular side to dorsal-fin base about 22; scales below

lateral line to anal-fin base about 24; vertebrae 37 (37-38); erisma

(counted as the first dorsal pterygiophore) about twice as thick as

remaining pterygiophores, its inner half narrowly branched; next

2 pterygiophores before tip of second neural spine; space between

second and third neural spines with 6 pterygiophores; space between

third and fourth neural spines with 3 pterygiophores; total of 12 dorsal

pterygiophores anterior to fourth neural spine; ventroanterior margin of

urohyal forming an angle of about 80°, the comer broadly rounded.

Body depth 2.6 (2.45-2.55) in SL; body width (thickness) 5.3

(4.7-5.75) in body depth; ventral profile of head posterior to mouth

slightly convex; HL 3.95 (3.85-3.95) in SL; snout length 2.7 (2.6)

in HL; eye diameter 4.6 (4.55-4.9) in HL; upper eye overlapping

anterior two-thirds to three-fourths of lower eye; least vertical

interorbital width 3.55 (2.75-3.35) in HL; upper end of gill opening on

a horizontal passing about one-half eye diameter ventral to lower eye;

no caudal peduncle (base of last two or three anal rays posterior to base

of lowermost caudal ray); depth at base of caudal fin 1.9 (1.8-1.95)

in HL.

Maxilla extending to below front edge of pupil, the upper-jaw length

(measured on blind side) 2.9 (2.9-2.95) in HL; blind side of upper and

lower jaws with a dense band of villiform teeth (difficult to see because

just medial to a labial fold); no teeth on ocular side of jaws; tubular

anterior nostril of ocular side membranous, just above upper lip, anter-

ior to upper edge of lower eye, slightly tapering, reaching a little

posterior to front edge of eye when laid back, its length about three-

fourths eye diameter; posterior nostril an oblique slit in labial groove

directly in front of dorsal half of lower eye; anterior nostril of blind

side a more slender, slightly tapering, membranous tube above about

middle of upper lip; posterior nostril of blind side a shorter, broader

membranous tube posterior and slightly dorsal to anterior nostril

(internarial distance about three-fourths eye diameter).

Scales ctenoid on both sides (except those of lateral line partially

embedded); scales of ocular side of body with 6-9 cteni; about 2 rows

of scales in interorbital space, with about another 5 rows extending

onto medial and anterior part of each eye; scales on ocular side of head

progressively smaller anteriorly and ventrally with fewer cteni,

replaced on snout by fleshy papillae; scales on blind side of head anter-

ior to a demarcation just posterior to end of jaws replaced by a dense

zone of fleshy papillae that are progressively longer anteriorly, about

15 visible on ventral edge of head posterior to mouth (long for

papillae, but too stout and short to call cirri). Lateral line straight on

both sides along middle of body, projecting on ocular side toward

ventral edge of upper eye; lateral line of blind side replaced by a row

of sensory papillae on head (differentiated from surrounding papillae

by a narrow papilla-free zone on each side), which curves ventrally at

front of head; supratemporal branch of lateral line on blind side of head

clearly visible as a similar row of low sensory papillae just below basal

A review of soles of the genus Aseraggodes

197

sheath of scales, becoming faint at end of about anterior third of body.

Each dorsal and anal ray with a thin lengthwise membranous ridge,

narrowing distally; ridges progressively less developed posteriorly;

small scales and papillae extending out on ridges of both sides of about

first 20 dorsal rays, making edges of membranous ridges jagged; about

basal fourth of caudal fin with progressively smaller scales on both

sides to at least three-fourths length of fin.

Origin of dorsal fin anterior to lower edge of upper eye, the predor-

sal length 4.1 (4.15—4.4) in HL; first dorsal ray (only the tip free) 3.8

(3.7-3.95) in HL; longest dorsal ray 1.65 (1.55-1.7) in HL; origin of

anal fin below base of 20th dorsal ray, slightly posterior to a vertical at

end of opercular membrane, the preanal length 3.3 (3.35-3.4) in SL;

length of first anal ray 3.6 (3.9-3.95) in HL; longest anal ray 1.65

(1.5-1. 7) in HL; caudal fin rounded, 4.45 (4.4-4.45) in SL; pelvic fin

bases adjacent on ventral edge of body, third and fourth pelvic rays

longest, reaching to base of second or third anal ray, 2.5 (2.4-2. 7) in

HL; anus anterior to first anal ray; genital papilla dorsoposterior to

anus, not connected by membrane to ocular-side pelvic fin.

Colour of ocular side of holotype when fresh: brownish yellow with

numerous whitish blotches about half eye diameter in size, many inter-

connected; three rows of very irregular, large, blackish blotches, the

dark pigment on scale edges, or isolated scales entirely black; a small

squarish white spot behind upper end of gill opening; eyes golden with

faint blackish bands, partly rimmed in black; fins with translucent

membranes and brownish yellow rays, some with 1 or 2 blackish spots;

scaly basal part of caudal fin coloured like body.

Colour of ocular side of holotype in alcohol: pale yellowish brown

with 3 rows of large irregular blackish blotches, one dorsal, one

ventral, and one midlateral; a few small dark spots on fin rays; blind

side of body pale yellowish, the dark spots faint.

Etymology. The species name auroculus is from the Latin

aurum for gold and oculus for eye, in reference to the bright

golden colour of the eyes.

Remarks. The four specimens of this species were collected in

1989 off Moorea in two rotenone stations from steep sloping

bottoms dominated by coral rubble at depths of 15-24 m. They

were deposited in the Royal Ontario Museum with a tentative

identification of Aseraggodes melanostictus (Peters), the name

often given to specimens of soles of this genus with an ocular-

side colour pattern of large blackish blotches. Although sharing

the same number of dorsal rays, vertebrae, and dorsal ptery-

giophores with A. melanostictus, A. auroculus is easily

distinguished by a higher count of anal rays, fewer lateral-line

scales (Tables 2 and 3), and having branched instead of

unbranched dorsal and anal fin rays. It also appears to be a

much smaller species. The two known specimens of

Ligure 3. Paratype of Aseraggodes bahamondei , BPBM 30851, 46.9

mm SL, Easter Island.

A. melanostictus measure 74 and 86.5 mm SL.

Aseraggodes auroculus is more closely related to

A. cyclurus, also collected from the Society Islands. One

rotenone station resulted in a specimen of both species. The two

are separated by dorsal- and anal-ray counts and differences in

body depth, eye size, and length of the caudal fin (see Key).

Also, A. cyclurus seems to be a larger species. The three

Society Islands specimen range from 61.5 to 73.3 mm SL,

compared to 29.8-35.5 mm for the four type specimens of

auroculus. The 34.0 mm paratype of A. auroculus is a fully

mature female.

The photograph taken of the holotype of Aseraggodes

auroculus (Fig. 2), shows a broad interorbital space, about

three-fourths the diameter of the lower eye. The interorbital

width of the preserved specimen is only one-fourth the eye

diameter. The same shrinkage of the interorbital space was

noted for one specimen of A. lenisquamis (see description

below).

Aseraggodes bahamondei Randall and Melendez, 1987

Figures 3-5, Tables 1-3

Aseraggodes bahamondei Randall and Melendez, 1987: 99, figs

1-3.

Material examined. See Randall and Melendez (1987).

Type locality. Easter Island.

Diagnosis. Dorsal rays 65-71; anal rays 50-56; dorsal and anal

rays branched, except in juveniles; lateral-line scales 75-86;

1-4 pores beneath many scales peripherally on ocular side of

Figure 4. Paratype of Aseraggodes bahamondei, BPBM 14790, 68.3

mm SL, Lord Howe Island.

Figure 5. Holotype of Aseraggodes bahamondei, BPBM 6610, 149.3

mm SL, Easter Island.

198

John E. Randall

body; vertebrae 39-40; dorsal pterygiophores anterior to fourth

neural spine 7-8; body depth 2.3-2.5 in SL; HL 3.8-4.55 in SL

(relatively longer in small individuals); upper lip not overlap-

ping lower lip when mouth closed; eye diameter 5. 8-6. 8 in HL;

upper eye varying from slightly anterior to one-half eye

diameter before lower eye; interorbital space 6.5-9.55 in HL;

tubular anterior nostril of ocular side not reaching edge of

lower eye when laid back; prominent lappet-like cirri on ven-

tral edge of head; caudal peduncle present, its length 11.0-15.5

in HL; lateral line aligned with ventral half of upper eye;

dorsal and anal rays short, the longest dorsal ray 1.9-2.15 in

HL; small scales extending out on membranous ridge of dorsal

and anal rays, but no cirri at free edge of ridges; caudal

fin rounded, 4.0-5. 1 in SL; origin of ocular-side pelvic ray

slightly anterior to blind- side fin; third pelvic ray longest,

reaching to or a little beyond base of second anal ray, 2. 8-3. 2

in HL; pale brown with dark-edged white spots and 3 rows

of black spots, these markings relatively smaller and more

irregular, in general, with growth. Largest specimen, 156

mm SL.

Remarks. Currently known only from Easter Island and Lord

Howe Island, but is likely to occur at some intermediate south-

ern subtropical islands such as Pitcairn, Rapa, Kermadec

Islands, or Norfolk Island. Collected from sand at depths of

2-25 m. This species is unique in having the highest vertebral

count and lowest number of anterior dorsal pterygiophores. It

also seems to reach the largest size of species of the genus.

Randall and Melendez demonstrated the toxicity of the milky

secretion exuded by this species when threatened, presumably

from the small pores beneath scales near the edge of the ocular

side of the body.

Aseraggodes cyclurus sp. nov.

Figure 6, Tables 1-3, 5

Holotype. BPBM 8105, 73.3 mm, Society Islands, Tahiti, Papara,

Teavaraa Pass, SE side, sand at entrance to cave, 27.5 m, rotenone, J.E.

Randall, 21 Sep 1967.

Paratypes: USNM 379462, 70.2 mm, same data as holotype; ROM

61359, 61.5 mm SL, Society Islands, Moorea, W side of pass off

Maharepa about middle of its length, 17°29'24"S, 149°48'0"W, 15-18

m, steep slope with coral mbble, sand, and 3-m wall, rotenone,

R. Winterbottom and R. Mooi, 5 Dec 1989.

Diagnosis. Dorsal rays 68-71; anal rays 53; most dorsal and

Figure 6. Holotype of Aseraggodes cyclurus , BPBM 8105, 73.3 mm

SL, Tahiti, Society Islands.

anal rays double branched; lateral-line scales 70-73 (including

6-7 anterior to a vertical at upper end of gill opening); verte-

brae 36-37; dorsal pterygiophores anterior to fourth neural

spine 10-11; body depth 2.25-2.4 in SL; HL 4.65 (4.7^1.95) in

SL; eye diameter 4.65-4.9 in HL; upper eye overlapping about

anterior half of lower eye; interorbital space narrow, the

vertical distance separating eyes about three-fourths eye

diameter; no caudal peduncle; no prominent cirri on ventral

edge of head; lateral line aligned with ventral edge of upper

eye; longest dorsal ray 1.45-1.6 in HL; caudal fin rounded,

its length 4.7-4. 8 in SL; pelvic fins 2.2-2.4 in HL, the tip of

longest ray reaching base of third anal ray; ocular side mottled

brown, the scale edges dark brown to black; large irregular

blackish blotches, the most prominent comprising 4 below base

of dorsal fin, 3 on lateral line, and 2 above posterior half of

anal fin.

Description. Dorsal rays 71 (68-70); anal rays 53; dorsal rays double-

branched except first 19 (first 21 and 25 of paratypes); anal rays

double-branched except first 5 and 9 rays of paratypes; caudal rays 18,

all branched, the middle 16 double-branched; pelvic rays 5, all

branched; lateral-line scales on ocular side 73 (70-72), including 6-7

anterior to a vertical at upper end of gill opening; scales above lateral

line on ocular side to dorsal-fin base about 24; scales below lateral line

to anal- fin base about 27; vertebrae 37 (36-37); erisma (counted as the

first dorsal pterygiophore) about twice as thick as remaining pterygio-

phores, its inner half narrowly branched; next 2 pterygiophores before

tip of second neural spine; space between second and third neural

spines with 5 pterygiophores; space between third and fourth neural

spines with 2 (2-3) pterygiophores; total of 10 (10-11) dorsal ptery-

gio-phores anterior to fourth neural spine; ventroanterioir margin of

urohyal forming an angle of about 80°, the comer well-rounded.

Body depth 2.25 (2.25-2.4) in SL; body width (thickness) 5.65

(4.6-6.0) in body depth; ventral profile of head posterior to mouth

nearly straight; HL 4.65 (4.7-4.95) in SL; snout length 2.7 (2.6) in HL;

eye diameter 4.9 (4.65^1.7) in HL; least vertical interorbital width 8.0

Table 5. Proportional measurements of type specimens of Aseraggodes

cyclurus as percentages of standard length

Holotype

Paratype

BPBM

8105

ROM

61359

USNM

379462

Standard Length (mm)

73.3

61.5

70.2

Body depth

44.7

44.6

42.1

Body width

7.9

9.7

7.0

Head length

21.6

21.4

20.2

Snout length

8.0

8.3

7.8

Preorbital length

7.8

7.5

7.7

Eye diameter

4.4

4.6

4.3

Interorbital width

2.7

2.8

Upper-jaw length

8.2

8.1

Caudal-base depth

13.8

14.6

14.3

Predorsal length

5.3

5.5

5.4

Preanal length

25.3

24.1

25.7

Prepelvic length

19.3

18.2

18.5

First dorsal ray

3.9

4.4

4.2

Longest dorsal ray

14.4

15.4

14.6

First anal ray

6.6

6.4

6.7

Longest anal ray

15.0

15.6

15.7

Caudal-fin length

21.2

22.7

22.6

Pelvic-fin length

9.6

9.5

9.7

A review of soles of the genus Aseraggodes

199

(7.2-7.95) in HL; a vertical at posterior edge of upper eye (edge of

dark eyeball) passing approximately through middle of lower eye;

upper end of gill opening on a horizontal passing about one-half eye

diameter ventral to lower eye; no caudal peduncle (base of lowermost

caudal ray ending above base of last anal ray); depth of body at base of

caudal fin 1 .55 (1 .4-1 .45) in HL.

Snout not overlapping lower jaw when mouth closed; maxilla

extending slightly posterior to a vertical at front edge of lower eye, the

upper jaw length (blind side) 3.65 (2.5-2.65) in HL; blind side of

upper and lower jaws with a dense band of slender, inward-projecting,

slightly curved teeth up to about 7 rows; no teeth on ocular side of

jaws; anterior nostril a tapering membranous tube anterior to upper

edge of lower eye, just reaching anterior edge of eyeball when laid

back, its length nearly equal to eye diameter; posterior nostril an

oblique slit in labial groove directly in front of ventral part of lower

eye; anterior nostril of blind side a short tapering membranous tube just

above anterior third of upper lip; aperture of posterior nostril of blind

side dorsoposterior to anterior nostril (internarial distance about equal

to eye diameter), covered anteriorly with a flattened papilla.

Scales ctenoid on both sides (except those of lateral line partially

embedded); scales of ocular side of body with 10-13 cteni; 3 rows of

scales in interorbital space, with about another 6 rows extending onto

medial half of each eye; scales on ocular side of head progressively

smaller anteriorly and ventrally, the very small scales at front of snout

without cteni; scales on blind side of head replaced anteriorly by small

slender stout papillae on front of snout; a dense zone of small fleshy

papillae ventral and adjacent to lower jaw on blind side and another

adjacent to upper jaw, the latter not much broader than jaw width;

anterior edge of snout and ventral edge of head with very fine cirri,

none along edge of operculum at gill opening on either side. Lateral

line straight on both sides along middle of body, projecting on ocular

side toward ventral edge of lower eye; lateral-line of blind side obscure

on head in zone of papillae where it curves well dorsal to upper jaw to

tip of snout; a supratemporal branch of lateral line on blind side of head

faintly visible, beginning at front of snout, and continuing along base

of dorsal fin to anterior body.

Dorsal and anal fins with a basal sheath of 2 to 3 rows of scales;

small scales continuing out on rays and adjacent membrane on first 25

rays of dorsal fin of ocular side of holotype, those on rays on a thin

membranous ridge basally on each ray; only a few scales basally on

first 7 rays of ocular side of anal fin; scales basally on rays of blind side

except for last 19 rays of dorsal fin and last 17 of anal fin; small cirri

projecting from edge of membranous ridge of anterior dorsal and

anal rays of ROM paratype (but not apparent on Bishop Museum

specimens); about basal third of caudal fin with scales on both sides;

tiny, well-spaced, isolated scales still with cteni, on each side of rays

posteriorly to within outer fourth of fin.

Origin of dorsal fin anterior to lower edge of upper eye, the pre-

dorsal length 4.85 (4.6) in HL; first dorsal ray (only the tip free) 6.55

(4.9) in HL; longest dorsal ray 1.75 (1.7) in HL; origin of anal fin

below base of 20th dorsal ray and in line with posterior end of oper-

cular membrane, the preanal length 3.95 (3.9) in SL; length of first

anal ray 3.9 (3.7) in HL; longest anal ray 1.7 (1.6) in HL; caudal fin

rounded, 4.8 (4.7) in SL; ocular-side pelvic fin on ventral edge of body,

blind-side fin adjacent, both covered anteriorly by a basal band of

small scales; third pelvic ray of each fin longest, reaching to base

of third anal ray, 2.65 (2.55) in HL; anus ventroanterior to first anal

ray; genital papilla on ocular side of base of first anal ray.

Colour of ocular side of holotype in alcohol pale tan, fins with pale

yellowish rays and transparent membranes; no dark markings apparent.

Colour of holotype when fresh light brown, edges of the scales dark

brown to black, with scattered small irregular pale blotches and small-

er dark brown spots; large irregular blackish blotches in 3 rows: upper

row of 4 blackish blotches along dorsal contour of body, middle row of

four blotches on lateral line, the first at origin of lateral line, the 2 mid-

dle blotches clearly largest on body; lower row of 4 blotches below

base of anal fin, less distinct than other blotches; blind side of body

whitish; opercular membrane pale; fin rays mottled brown to dark

brown, the membranes pale.

Etymology. Named cyclurus from the Greek for the nearly cir-

cular caudal fin.

Remarks. This species is described from three specimens, two

from sand at the entrance to a cave in Tahiti at a depth of 27.5

m, and the third on a steep rubble and sand slope in 15-18 m in

a pass at Moorea.

A fourth specimen, BPBM 20864, 56.6 mm, collected at

Budd Reef, Ringgold Isles, Fiji, in 11-26 m with rotenone by

B.A. Carlson and M. Gawel on 14 Apr 1973, is provisionally

identified as Aseraggodes cyclurus. The head and body were

preserved in a curve, so it is difficult to make accurate propor-

tional measurements, no photograph or colour notes were

taken, and the colour has largely faded. The specimen has 70

dorsal rays, those posterior to 18th ray double-branched; 52

anal rays, those posterior to third ray double-branched, 74

lateral-line scales, 36 vertebrae, and 11 dorsal pterygiophores

anterior to the fourth neural spine.

Aseraggodes cyclurus seems most closely related to

A. heraldi Randall and Bartsch, described from two specimens,

38.5 and 47 mm SL, from the Marshall Islands. The two species

share the following characters: no caudal peduncle, same ver-

tebral count, lack of prominent cirri along the ventral edge of

the head, scales extending onto anterior dorsal and anal rays

beyond the basal scaly sheath, and many double-branched rays

of the dorsal and anal fins. They differ slightly in the number of

dorsal rays (70-73 for A. cyclurus vs 75 for A. heraldi ) and anal

rays (51 and 52 for A. cyclurus vs 57 and 58 for A. heraldi ) and

the number of dorsal pterygiophores before the fourth neural

spine, 10 for A. cyclurus (11 for the non- type from Fiji), com-

pared to 12 for A. heraldi. The difference in snout length is the

only difference in proportional measurements that seems great

enough to be beyond individual variation, given the difference

in size of the specimens under comparison. The snout length of

A. heraldi is longer, 8. 1-8.2 in SL, compared to 6.8-6.9 for

A. cyclurus. Also significant is the more extensive area of papil-

lae anteriorly on the blind side of the snout of A. heraldi ; the

entire snout dorsal to the straight anterior part of the upper jaw

is densely covered with small papillae. In A. cyclurus there is

only a zone of papillae adjacent to the jaw that is not much

wider than the jaw. There is also a difference in the basic colour

pattern of the two species. The dark blotches in the three basic

rows on the body are more numerous, and their relative size

smaller in A. heraldi.

Aseraggodes lateralis sp. nov.

Figures 7, 8, Tables 1-3, 6

Holotype. BPBM 10992, 64.2 mm, Marquesas Islands, Ua Huka, W

side, small bay 0.4 miles NE of Motu Takatai, N side of bay in 4.5-9

m, rotenone, J.E. Randall, J.R. Haywood, and R.M. McNair, 7 May 1971.

Paratypes. BPBM 12757, 27.8 mm, Marquesas Islands, Nuku Hiva,

Sentinelle de l’Est, W side, steep rocky slope with no visible sand, 23

m, quinaldine, J.E. Randall and D.B. Cannoy, 17 May 1971; USNM

200

John E. Randall

Figure 7. Holotype of Aseraggodes lateralis, BPBM 10992, 64.2 mm

SL, Ua Huka, Marquesas Islands.

Figure 8. Paratype of Aseraggodes lateralis, BPBM 12757, 27.8 mm

SL, Nuku Hiva, Marquesas Islands.

Table 6. Proportional measurements of the type specimens of

Aseraggodes lateralis as percentages of standard length

Holotype

BPBM

10992

Paratype

BPBM

12757

Paratype

U S NM

382053

Standard Length (mm)

64.2

27.8

37.7

Body depth

50.2

41.6

42.0

Body width

8.1

7.2

7.9

Head length

22.2

22.4

23.8

Snout length

7.5

7.4

Preorbital length

6.5

6.4

6.6

Eye diameter

3.8

4.2

3.9

Interorbital width

2.9

2.1

2.0

Upper-jaw length

7.0

7.1

6.7

Caudal-base depth

15.3

13.9

12.3

Predorsal length

4.5

4.2

4.9

Preanal length

29.7

28.6

29.2

Prepelvic length

25.2

24.1

24.6

First dorsal ray

4.6

4.3

5.3

Longest dorsal ray

13.3

14.0

14.3

First anal ray

7.3

6.8

7.4

Longest anal ray

13.7

14.3

14.4

Caudal-fin length

19.0

21.4

20.9

Pelvic-fin length

7.8

8.0

7.9

392053, 37.7 mm, Marquesas Islands, Fatu Hiva, Hanavave Bay, N

side, 20 m, rotenone, J.L. Earle, L.A. Rocha, and W. Robbins, 23 Aug

2003.

Diagnosis. Dorsal rays 77-78; anal rays 58-59; lateral-line

scales 78-83; vertebrae 37-38; dorsal pterygiophores anterior

to fourth neural spine 12; body depth 2.0-2.4 in SL; HL

3.9-4.15 in SL; eye diameter 6.1-6.35 in HL; upper eye from

directly above lower eye to overlapping anterior two-thirds of

lower eye; interorbital space equal to one-half to three-fourths

eye diameter; no caudal peduncle; upper lip not overhanging

lower jaw when mouth closed; very fine cirri on ventral edge of

head; lateral line aligned with ventral third of upper eye;

longest dorsal ray 1.8 in HL; caudal fin rounded to slightly

pointed, its length 4.7-5.25 in SL; longest pelvic ray 3. 1-3. 2 in

HL, the tip reaching base of second anal ray; ocular side

mottled brown; scale edges dark brown to black; large irregular

blackish blotches, the most prominent comprising 4 below base

of dorsal fin, 3 on lateral line, and 2 above posterior half of anal

fin; lateral line as a broken black line.

Description. Dorsal rays 77, the first 15 branched, remaining rays dou-

ble-branched; anal rays 58 (59), all double branched; caudal rays 18

(uppermost and lowermost rays branched, the middle 16 double

branched; pelvic rays 5, all branched; lateral-line scales 78 (79-83),

including 7-8 anterior to a vertical at upper end of gill opening; scales

above lateral line on ocular side to dorsal-fin base about 28; scales

below lateral line to anal-fin base about 29; vertebrae 37 (38); erisma

(counted as the first dorsal pterygiophore) nearly 3 times thicker than

remaining pterygiophores, its inner three-fourths narrowly branched;

next 2 pterygiophores before tip of second neural spine; space

between second and third neural spines with 6 pterygiophores; space

between third and fourth neural spines with 3 pterygiophores (total of

12 dorsal pterygiophores anterior to fourth neural spine); ventro-

anterior margin of urohyal forming angle of about 80°, the comer

strongly rounded.

Body oval and deep for genus, the depth 2.0 (2.35-2.4) in SL; body

thin, the width 6.2 (5.7-5. 8) in body depth; ventral profile of head pos-

terior to mouth slightly convex; HL 4.5 (4.45-4.6) in SL; snout length

2.95 (2.95-3.0) in HL; eyes small, the eye diameter 5.85 (5.35-5.6) in

HL; upper eye of holotype directly above lower (overlapping about

anterior two-thirds of lower eye in paratypes); least vertical interorbital

space 7.65 (10.7-10.9) in HL; upper end of gill opening at level of

ventral edge of lower eye; no caudal peduncle (base of lowermost

caudal ray ending above base of last anal ray); depth of body at base of

caudal fin 1.45 (1.6-1.75) in HL.

Front of upper lip not overlapping lower jaw when mouth closed;

maxilla extending to a vertical at anterior fourth of lower eye, the

upper jaw length (blind side) 3.2 (3.15-3.25) in HL; blind side of jaws

with a dense band of slender, inward-projecting, slightly curved teeth

nearly the full length of each jaw, up to about 6 rows at broadest place;

anterior nostril a tapering tube before upper edge of lower eye, reach-

ing anterior edge of eye when laid back, its length about three-fourths

eye diameter; posterior nostril an oblique slit in labial groove directly

in front of ventral part of lower eye; anterior nostril of blind side a

short tapering membranous tube just above anterior third of upper jaw;

posterior nostril of blind side dorsoposterior to anterior nostril;

intemarial distance about equal to eye diameter.

Scales of both sides ctenoid (except cycloid on lateral line and

partially embedded), usually with 10 cteni (a few with 11 or 12);

4 rows of scales in interorbital space, with small scales extending

broadly onto median and anterior edge of eyes (apparently lost in

some); scales progressively shorter anteriorly on head and with fewer

cteni (front of snout with very small scales without cteni, but not in the

form of little papillae or tubercles); anterior edge of snout and ventral

edge of head with a row very fine cirri (especially fine anteriorly on

A review of soles of the genus Aseraggodes

201

snout); no cirri along edge of operculum; opercular membrane in form

of a short triangle near upper end of gill opening. Lateral line straight

on both sides along middle of body, on ocular side aligned with ventral

third of upper eye; an indistinct supratemporal branch of lateral line on

blind side of head, continuing faintly aanteriorly on body along base of

dorsal fin.

Dorsal and anal fins with a basal sheath of 2 to 3 rows of scales;

ocular side with small scales continuing a short distance out on rays

and adjacent membrane on first 31 rays of dorsal fin of ocular side of

holotype, and on the first 8 rays of ocular side of anal fin; scales

extending out on rays of dorsal and anal rays of blind side except for

last 21 rays of dorsal fin and last 33 of anal fin; a thin lengthwise mem-

branous scaly ridge basally on anterior dorsal and anal rays of both

sides, progressively shorter posteriorly; about basal third of caudal fin

with scales on both sides, and well-spaced tiny scales, still with cteni,

continuing on each side of rays about half way out in fin.

Origin of dorsal fin anterior to lower edge of upper eye, the pre-

dorsal length 4.95 (4.45^4.65) in HL; first dorsal ray 4.85 (4.15-5.2)

in HL; longest dorsal ray 1.65 (1.5-1.55) in HL; origin of anal fin

below base of 20th dorsal ray and in line with posterior end of oper-

culum, the preanal length 3.35 (3.45-3.5) in SL; anus anterior to first

anal ray; genital papilla dorsoposterior to anus; first anal ray 3.05

(2.95-3.3) in HL; longest anal ray 1.6 (1.5-1.55) in HL; caudal fin

rounded to slightly pointed, 5.25 (4.7-4.8) in SL; pelvic fins close

together anteriorly on ventral edge of body, their origins adjacent; third

pelvic ray of each fin longest, reaching base of third anal ray, 2.85

(2.75-2.8) in HL.

Colour of ocular side in alcohol light yellowish brown with scat-

tered irregular brown blotches of variable size, the most conspicuous

in 3 rows: one below base of dorsal fin, one following lateral line, and

one just above anal fin; blind side uniform light yellowish brown; fins

pale with a few faint small brown spots basally in dorsal and anal fins.

Colour of ocular side of holotype when fresh: light brown with

small irregular pale and dark brown blotches; scale edges dark brown

to black; large irregular blackish blotches containing small pale spots

in 3 rows, dorsal, ventral, and along lateral line; lateral line clearly evi-

dent as a broken black line; posterior edge of operculum broadly white;

eyes pale with radiating dark lines on dorsal half of iris; scaled basal

part of fins coloured like body; remaining part of fins pale with small

dark blotches and a few larger ones along base.

Etymology. Named lateralis from the Latin word with the same

meaning in English, in reference to the distinct pigmented lat-

eral line in life.

Remarks. The holotype was collected from rock and sand bot-

tom of a protected bay, but the habitat of the smallest paratype

was unexpected. It was taken from a steep rocky slope with no

obvious sand during the day. Species of Aseraggodes are

typically found on sedimentary bottoms, at least by day, and

they are usually buried during daylight hours.

The smallest paratype of Aseraggodes lateralis is in poor

condition with the fins badly eroded, a result of long retention

in isopropanol of insufficient concentration. A comparison of

the figures of the holotypes of Aseraggodes lateralis and

A. cyclurus reveals similarity in body and fin morphology and

in colour pattern. Both species have 37 vertebrae and nearly the

same structure of the fins. Aseraggodes cyclurus differs in

having higher dorsal, anal, and lateral-line scale counts (see

Tables 1-3), extremely small cirri on the ventral edge of the

head, and small slender tubercles instead of scales anteriorly on

the blind side of the head.

The 37.7-mm paratype is a mature female. An x-ray

revealed that it had eaten two gastropods with intact shells 1.7

and 1.9 mm long.

Aseraggodes lenisquamis sp. nov.

Figure 9, Tables 1-3, 7

Aseraggodes sp. Kuiter, 1993: 391, upper fig. (central to south-

Table 7. Proportional measurements of type specimens of Aseraggodes lenisquamis as percentages of standard length

Holotype Paratypes

NMV NMV NSMT AMS BMNH CAS BPBM NMV

A 19646 A 25543 P 70086 1.27047 4.10.26.4 221846 39610 A 25543

Standard Length (mm)

Body depth

Body width

Head length

Snout length

Preorbital length

Eye diameter

Interorbital width

Upper-jaw length

Caudal-base depth

Predorsal length

Preanal length

Prepelvic length

First dorsal ray

Longest dorsal ray

First anal ray

Longest anal ray

Caudal-fin length

Pelvic-fin length

102.0

64.5

39.4

38.6

7.2

7.4

20.3

19.9

8.0

8.1

6.4

4.1

4.0

2.0

1.9

5.9

6.2

14.9

15.5

4.5

5.0

26.1

25.2

17.8

18.3

6.7

5.6

11.8

11.9

7.3

7.6

12.1

11.9

21.8

21.9

9.8

10.9

69.0

73.0

40.6

37.8

7.5

7.8

20.6

20.4

8.3

8.2

6.5

6.4

4.5

4.2

1.5

2.8

6.3

5.9

16.2

15.3

4.4

4.5

24.7

26.2

18.3

18.7

6.1

5.8

12.0

12.3

7.0

8.2

11.6

12.6

21.0

21.6

9.9

11.1

77.5

78.0

41.1

41.3

7.8

7.5

20.8

20.3

8.4

8.3

6.7

6.4

3.9

4.1

2.6

2.5

6.4

6.3

15.2

14.9

4.9

5.0

25.7

25.6

18.7

17.7

6.4

6.5

11.7

11.5

8.1

7.8

11.6

11.5

20.8

19.8

10.7

10.2

82.0

82.5

41.2

39.9

7.8

7.3

21.1

20.9

8.6

8.3

6.2

6.1

3.7

3.8

2.4

2.7

6.1

6.5

14.5

15.6

5.1

4.7

26.2

26.4

19.4

19.0

6.7

6.3

11.6

11.8

7.2

7.4

11.7

11.8

19.9

20.9

10.5

202

John E. Randall

Figure 9. Holotype of Aseraggodes lenisquamis , NMV A 19646, 102

mm, NSW.

em NSW).

Holotype. NMV A 19646, 102 mm, Australia, NSW, Sydney Harbor,

Camp Cove (33°50’S, 151°17’E), 4 m, hand net, R.H. Kuiter, 17 Jan

1985.

Paratypes. NMV A 3607, 65 mm, Australia, NSW, Bermagui,

Horseshoe Bay, 36°25’S, 150°4’E, 10 m, hand net, R.H. Kuiter, 24 Jan

1984; NMV A 5827, 88 mm, same locality as preceding, 4-10 m, hand

net, R.H. Kuiter, 30 Jan 1984; NMV A 25543-002, 2: 64.5-82.5 mm,

BPBM 39610, 82 mm, CAS 221846, 78 mm, NSMT P 70086, 69 mm,

BMNH 2004.10.26.4, 77.5 mm, USNM 380210, 68 mm, all with same

data as holotype; AMS 1.27047-001, 73 mm, NSW, Jervis Bay, Hare

Bay, 35°3'S, 150°44'E, 6 m, beam trawl, F.R.I. Jervis Bay study, 28 Oct

1986; AMS 27063-013, 65 mm, NSW, Jervis Bay, Hare Bay, 35°0’S,

150°45'E, 2-7 m, J. Bell (State Fisheries), Dec 1986; AMS 1.28514-

002, 79 mm, NSW, Jervis Bay, Darling Road, 35°3’S, 150°44’E, 5 m,

beam trawl, J. Bell and party, 28 Sep 1988.

Diagnosis. Dorsal rays 62-70; anal rays 46-52; dorsal and anal

rays branched; lateral-line scales 62-68, including 8-9 anterior

to a vertical at upper end of gill opening; vertebrae 36-38;

dorsal pterygiophores anterior to fourth neural spine 8-9; body

depth 2.4-2.65 in SL; head short, its length 4.75-5.05 in SL;

eye diameter 4.6-5 .7 in HL; upper eye overlapping about ante-

rior one-third to one-half of lower eye; interorbital space vari-

able in width, the vertical distance separating eyes 7.3-13.7 in

HL; no caudal peduncle; lappet-like cirri on ventral edge of

head, but not on front of snout; numerous cirri on opercular

edge of gill opening on both sides; dense cirri over much of

ventral part of head; exposed surface of scales overlaid with

soft tissue; only tips of cirri visible at scale margins, capped

with soft tissue; lateral-line scales with fleshy cirri, often one

above and one below pore (cirri better developed on ocular than

blind side); scattered other scales with a slender fleshy cirrus,

often one from each comer of scale; membranous ridges of

both sides of dorsal and anal rays with a conspicuous fringe of

cirri, some of which are bifid; lateral line aligned with upper

eye; longest dorsal ray 1.65-1.8 in HL; caudal fin rounded, its

length 4.6-5.05 in SL; pelvic fins short, 1.8-2.2 in HL, the tip

of longest ray reaching base of second anal ray; ocular side

light brown, with scattered small dark brown blotches; rays

of fins with small dark brown spots. Largest specimen, 102

mm SL.

Description. Dorsal rays 69 (62-70); anal rays 51 (46-52); dorsal and

anal rays branched, most double-branched in large specimens; caudal

rays 18, branched, the middle 16 double-branched; pelvic rays 5,

double-branched; lateral-line scales on ocular side 67 (62-68), includ-

ing 8-9 anterior to a vertical at upper end of gill opening; scales above

lateral line on ocular side to dorsal-fin base about 21; scales below lat-

eral line to anal-fin base about 23; vertebrae 37 (five paratypes with 37,

two with 36, and one with 38); dorsal pterygiophores anterior to fourth

neural spine 9 (8-9); only the erisma (counted as the first dorsal ptery-

giophore before tip of second neural spine) about twice as thick as

remaining pterygiophores, its inner third narrowly branched; next 5

(5-6) pterygiophores in space between second and third neural spines;

space between third and fourth neural spines with 3 (2-3) pterygio-

phores; total pterygiophores before fourth neural spine 9 (five

paratypes with 9, and four with 8); ventroanterior margin of urohyal

forming an angle of about 55°, the comer only slightly rounded.

Body depth 2.55 (2.4-2.65) in SL; body width 5.45 (4.85-5.45) in

body depth; ventral profile of head posterior to mouth nearly straight;

head short, its length 4.85 (4.75-5.05) in SL; snout length 2.55

(2.45-2.5) in HL; eye diameter 4.95 (4.6-5.7) in HL; least vertical

interorbital width 10.2 (7.3-13.7) in HL; upper eye overlapping anter-

ior one-third to one-half of lower eye; upper end of gill opening on a

horizontal passing about an eye diameter ventral to lower eye; no

caudal peduncle (base of lowermost caudal ray ending above base of last

anal ray); depth of body at base of caudal fin 1.35 (1.25-1.45) in HL.

Front of upper lip not overlapping lower lip when mouth closed;

maxilla extending to below anterior margin of pupil, the upper jaw

length (blind side) 3.45 (3.25-3.45) in HL; blind side of upper and

lower jaws with a dense band of slender, inward-projecting, slightly

curved teeth in a maximum of about 8 rows; no teeth on ocular side of

jaws; anterior nostril a short membranous tube, tapering very little,

anterior to upper edge of lower eye, and not reaching edge of eye when

laid back; posterior nostril a slit in labial groove in front of upper half

of lower eye; anterior nostril of blind side a membranous tube above

anterior third of upper lip; posterior nostril of blind side dorso-

posterior to anterior nostril; intemarial distance about two-thirds eye

diameter.

Surface of scales smooth, the ridges of cteni covered with cutaneous

tissue; free margin of scales angular (though posterior end not acutely

pointed); only tips of cteni of scales visible at scale margin (up to 14

on holotype), each covered with soft tissue; 2 (2^1) rows of scales in

interorbital space, with another 2-3 rows extending medially and ante-

riorly onto eyes; no pores detected beneath scales on ocular side of

body; scales on ocular side of head progressively smaller anteriorly,

replaced at front of snout and ventrally on head with band of dense cirri

nearly eye diameter in width; cirri at front edge of snout very small,

those of ventral edge of head longer and more lappet-like, the longest

nearly pupil diameter in length, a few branched at tips; a broader band

of dense cirri anteriorly on blind side of head, around mouth, and

extending in a zone along supratemporal branch of lateral line, which

continues, progressively less distinct, onto about anterior half of body;

band of cirri at edge of operculum on both sides, a dense fringe along

gill opening; lateral-line scales with cirri, usually one above and one

below the pore; scattered other scales with cirri, generally one at each

comer; dorsal end of gill opening on a horizontal line passing about an

eye diameter below lower eye. Lateral line straight midlaterally on

both sides, projecting on ocular side toward upper eye.

Dorsal and anal fins with a basal sheath of 2 rows of scales; dorsal

and anal rays on both sides with a fleshy lengthwise ridge bearing a

fringe of prominent cirri; ridges on rays less developed posteriorly and

with fewer cirri, especially on blind side; scales on basal half of caudal

fin progressively smaller distally.

Origin of dorsal fin anterior to upper eye, the predorsal length 4.5

(4.0-4.65) in HL; first dorsal ray 3.0 (3.1-3.55) in HL; longest dorsal

ray 1.7 (1.65-1.8) in HL; origin of anal fin below base of seventeenth

A review of soles of the genus Aseraggodes

203

dorsal ray and slightly posterior to end of opercular membrane, the

preanal length 3.85 (3.8^1.05) in SL; length of first anal ray 2.8

(2.5-2.95) in HL; longest anal ray 1.7 (1.6-1. 8) in HL; caudal fin

rounded, 4.6 (4.6-5.05) in SL; anus directly before first anal ray, pre-

ceded by fan-like semicircle of plicate tissue; genital papilla on ocular

side of anus, its length nearly three-fourths eye diameter in holotype;

bases of pelvic fins adjacent anteriorly, diverging posteriorly, the

ocular-side fin a little before blind-side fin; pelvic fins not joined by

membrane or to genital papilla; third pelvic ray of each fin longest,

reaching to base of second anal ray, 2.05 (1.85-2.0) in HL.

Colour of ocular side of holotype in alcohol light brown with many

small dark brown blotches, some stellate or cross-shaped; fins pale

yellowish with small dark brown spots along rays, most spots

coinciding with one or a clump of cirri (see below).

Etymology. This species is named lenisquamis from the Latin

lenis for soft or smooth and squama for scale, in reference to

the distinctive scale structure. The cteni of the scales are

nearly covered by soft epidermal tissue, with only the tips

exposed at the scale margin.

Remarks. Kuiter (1993: 391) figured two specimens as

Aseraggodes sp., his upper figure labelled as the estuarine

form, and the lower as coastal form. Upper figure (NMV A

25543-002, 82.5 mm SL) is included here as a paratype. When

fresh, it was orangish brown with diffuse blackish blotches, the

three along lateral line and one above the first blotch the

largest; head and body with numerous, small, very irregular,

dark-edged whitish spots; fin rays grey-brown with small dark-

edged whitish spots, the ray tips white. The colour of the lower

figure is very different, mottled lighter brown, likely to have

been collected from a paler sand bottom.

This species was collected from sand in bays in NSW from

depths of 4-10 m. It is among the most distinctive of the genus

because of its angular and smoother scales and the profusion of

fleshy cirri, in particular those on most of the lateral-line scales.

The covering of the ctenii of the scales by soft tissue except

the small tips is also found in A. normani, but its scales have a

rounded posterior margin instead of an angular one. It is easily

distinguished from lenisquamis in other characters such as the

branching of its lateral line on the ocular side of the head, its

unbranched dorsal and anal rays, and the pelvic fins joined to

the base of the genital papilla.

Aseraggodes lenisquamis is most similar to A. nigrocirratus,

described below (see Remarks for the latter species).

The lower figure of this species in Kuiter (1993) mentioned

above, was not identified initially as A. lenisquamis because of

the broad interorbital space and the upper eye being almost

directly over the interorbital space. However, the specimen was

found as NMV A5827, 88 mm SL, from its shape, details of

colour, and a distinct small tear in the caudal fin. The inter-

orbital of the preserved specimen is much narrower, and the upper

eye is more forward. A similar shrinkage of the interorbital with

preservation was found in the holotype of A. auroculus.

Aseraggodes magnoculus sp. nov.

Figure 10, Tables 1-3, 8

Holotype. ROM 64830, 39.8 mm, New Caledonia, Isle Ua, E side,

22°42'40"S, 166°48'50"E, steep slope of fringing reef, with coral rock,

mbble, and sand at base, 9-18 m, rotenone, R. Winterbottom and R

Tirard, 13 Sep 1991.

Paratypes. ROM 76686, 29.2 mm, New Caledonia, just inside bar-

rier reef (Recif Mbere) NW of Dumbea Pass, 22°20'30"S, 166°14'5"E,

large Porites coral head surrounded by sand, 3-6.5 m, rotenone, R.

Winterbottom, G. Klassen and P. Tirard, 11 Sep 1991; BPBM 39691,

31.3 mm, same data as holotype.

Diagnosis. Dorsal rays 67-72; anal rays 51-53; dorsal rays

branched except anterior 16-24 rays; anal rays branched; later-

al-line scales 71-76, including 7-8 anterior to a vertical at

upper end of gill opening; vertebrae 36; dorsal pterygiophores

anterior to fourth neural spine 10; body depth 2.45-2.5 in SL;

HL 4.15-4.3 in SL; eye diameter 3.95-4.2 in HL; upper eye

overlapping about anterior one-third to one-half of lower eye;

interorbital space narrow, the vertical distance separating eyes

about one-third to one-sixth eye diameter; no caudal peduncle;

very fine cirri on ventral edge of head; lateral line aligned with

ventral edge of upper eye; longest dorsal ray 1.4-1.45 in HL;

caudal fin rounded, its length 3.9-3.95 in SL; pelvic fins

Table 8. Proportional measurements of type specimens of Aseraggodes

magnoculus as percentages of standard length

Holotype

Paratype

ROM

BPBM

64830

76686

39691

Standard Length (mm)

39.8

29.2

31.3

Body depth

40.9

39.7

40.2

Body width

7.9

7.8

7.6

Head length

23.2

24.0

24.1

Snout length

8.9

9.3

9.2

Preorbital length

7.7

7.5

8.8

Eye diameter

5.5

5.8

6.1

Interorbital width

1.0

1.9

Upper-jaw length

7.5

7.4

Caudal-base depth

13.8

14.3

13.3

Predorsal length

6.3

6.2

6.1

Preanal length

26.6

27.5

26.9

Prepelvic length

20.6

21.0

21.8

First dorsal ray

7.3

7.5

7.0

Longest dorsal ray

16.8

16.6

17.2

First anal ray

8.8

8.7

8.9

Longest anal ray

15.8

16.6

Caudal-fin length

25.2

25.7

broken

Pelvic-fin length

12.8

12.6

Figure 10. Holotype of Aseraggodes magnoculus , ROM 64830, 39.8

mm, New Caledonia.

204

John E. Randall

1. 8-2.0 in HL, the tip of longest ray reaching base of second or

third anal ray; colour of ocular side in alcohol light yellowish

brown with 3 rows of dark brown blotches, one row below base

of dorsal fin, one above base of anal and pelvic fins, and one

with two largest blotches well- spaced on lateral line; other

brown markings mainly vertically elongate, some enclosing

small irregular areas of ground colour; fins pale yellowish

except for faint dark blotches along base.

Description. Dorsal rays 72 (67-70); anal rays 53 (51-52); dorsal rays

branched except first 16 (first 23-24 of paratypes); anal rays branched;

caudal rays 18, the middle 16 branched, but not double-branched;

pelvic rays 5, branched; rays of fins slender, the branches not broadly

separated; lateral-line scales 71 (72-76), including 7-8 anterior to a

vertical at upper end of gill opening; scales above lateral line on

ocular side to dorsal-fin base 21-22; scales below lateral line to anal-

fin base 23-24; vertebrae 36; erisma (counted as the first dorsal ptery-

giophore) about twice as thick as remaining pterygiophores, its inner

half narrowly branched; next pterygiophore before tip of second

neural spine; space between second and third neural spines with

5 pterygiophores; space between third and fourth neural spines with

3 pterygiophores (total of 10 dorsal pterygiophores anterior to fourth

neural spine); ventroanterior margin of urohyal strongly curved, the

two limbs, if projected, forming an angle of about 60°.

Body depth 2.45 (2.5) in SL; body width (thickness) 5.2 (5. 1-5.3)

in body depth; ventral profile of head posterior to mouth very slightly

convex; HL 4.3 (4.15) in SL; snout length 2.6 in HL; eyes large, 4.2

(3.95-4.15) in HL; interorbital space very narrow, the least vertical

interorbital width 23.2 (12.7-24.0) in HL; upper eye overlapping anter-

ior one-fourth to one-third of lower eye; upper end of gill opening on

a horizontal passing slightly ventral to lower eye; no caudal peduncle

(base of last anal ray posterior to base of lowermost caudal ray); depth

of body at base of caudal fin 1.7 (1.7— 1.8) in HL.

Snout slightly overhanging lower jaw; maxilla nearly reaching to

below center of eye, the upper jaw length (blind side) 3.1 (3.2-3.25) in

HL; blind side of upper and lower jaws with a dense band of villiform

teeth, obscured laterally because of a labial fold; no teeth on ocular

side of jaws; tapering tubular anterior nostril of ocular side membra-

nous, before upper edge of eye just above upper lip, nearly reaching

anterior edge of pupil when laid back, its length equal to eye diameter;

posterior nostril a slit in labial groove in front of lower eye; anterior

nostril of blind side a slender tube only about twice as long as sur-

rounding papillae, just above middle of upper lip; posterior nostril of

blind side an opening covered by a triangular papilla-like structure,

dorsoposterior to anterior nostril; intemarial distance nearly equal to

eye diameter.

Scales ctenoid on both sides (except partially embedded scales of

lateral line); scales of ocular side of body with up to 10 cteni; only 1 or

2 rows of scales in interorbital space, with about another 5 rows

extending onto medial and anterior part of each eye; scales on ocular

side of head smaller anteriorly, replaced on snout by longitudinal rows

of small fleshy papillae; scales on blind side of head anterior to a trans-

verse demarcation just posterior to end of jaws replaced by a dense

zone of fleshy papillae, largest above posterior half of upper lip; only

fine cirri at front edge of snout and along ventral part of head. Lateral

line straight midlaterally on both sides, projecting on ocular side

toward middle of upper eye; lateral line of blind side altering to a row

of sensory papillae on head (separated from surrounding papillae by a

narrow papilla-free zone on ventral side that curves on head to tip of

snout; supratemporal branch of lateral line on blind side of head

clearly visible as a similar row of low sensory papillae just below basal

sheath of scales on dorsal fin, less obvious on body posterior to head.

Dorsal and anal rays each with a lengthwise thin membranous ridge,

narrowing distally, less developed posteriorly; membranous ridges

without cirri; small scales basally on ocular side of dorsal fin before

about 15th ray, progressively fewer scales approaching 15th ray; anter-

ior part of dorsal fin on blind side with small papillae, progressively

fewer posteriorly, and absent after about the 24th ray.

Origin of dorsal fin anterior to upper eye, the predorsal length 3.7

(3.9-3.95) in HL; first dorsal ray (tip not free; tips of next few rays

visible) 3.8 (3.7-3.95) in HL; longest dorsal ray 1.45 (1.4) in HL;

origin of anal fin below base of seventeeth dorsal ray, slightly poster-

ior to end of opercular membrane, the preanal length 3.75 (3.65-3.7)

in SL; length of first anal ray 3.75 (3.65) in HL; longest anal ray 1.45

(1.45-1.5) in HL; caudal fin rounded, 3.95 (3.9) in SL; pelvic fins

adjacent on ventral edge of body, third and fourth pelvic rays longest,

reaching to base of third anal rays, 1.8 (1.9) in HL; anus anterior to

first anal ray; genital papilla short, dorsoposterior to anus, and not

connected by membrane to ocular-side pelvic fin.

Colour of holotype in alcohol: ocular side light yellowish brown

with 3 rows of dark brown blotches, one below base of dorsal fin, one

above base of anal and pelvic fins, and one midlateral, the two largest

blotches well-spaced on lateral line; other lighter brown markings

irregular, mainly transversely elongate, some enclosing small areas

of lighter ground colour; fins pale yellowish except for faint dark

blotches along base; blind side uniform pale yellowish.

Etymology. This species is named magnoculus from the Latin

magnus for large and oculus for eye, in reference to its having

the largest eyes, relative to the head length, of any species of

the genus examined.

Remarks. Aseraggodes magnoculus is described from three

small specimens collected in two stations from sand and rubble

in coral-reef areas of New Caledonia within the depth range of

3-18 m. No gonad was detected in the largest specimen, so it

may be immature. Aseraggodes auroculus, described above,

appears to be the most similar species to A. magnoculus. The

two share the same body depth, large eyes, and number of ver-

tebrae, dorsal pterygiophores anterior to the fourth neural spine,

dorsal rays, and lateral-line scales. A. auroculus differs in

having a higher count of anal rays (56 or 57, compared to

51-53 for magnoculus ), a larger head (hence the eye size

relative to the head is smaller than that of magnoculus ), and in

having shorter fins.

Aseraggodes melanostictus (Peters, 1877)

Figure 11, Tables 1-3

Solea melanosticta Peters, 1877: 845

Aseraggodes melanostictus . — Randall and Bartsch, 2005: figs. 3-5

Figure 11. Aseraggodes melanostictus, AMS 1.24499-003, 86.5 mm,

North Reef, Great Barrier Reef.

A review of soles of the genus Aseraggodes

205

(of holotype).

Type locality. Bougainville

Diagnosis. Dorsal rays 74-75, unbranched; anal rays 53-54,

unbranched; caudal rays 18 (uppermost and lowermost 2 rays

simple, middle 12 double-branched); pelvic rays 5,

unbranched; lateral-line scales on ocular side 78-79, including

9 anterior to a vertical at upper end of gill opening; scales of

ocular side of body with 12-16 cteni; scales anteriorly on head

progressively shorter, with fewer cteni, modified to small

flattened papillae, each with a small flat brown cirrus in zone

anteriorly on snout; no broad lappet-like cirri at front of snout

or on ventral edge of head, instead cylindrical cirri of variable

size, the longest less than pupil diameter; well-spaced slender

cirri on opercular edge of gill opening on both sides; vertebrae

38; dorsal pterygiophores anterior to fourth neural spine 11-13;

body depth 2.3-2.35 in SL; HL 4.35-4.5 in SL; snout slightly

overhanging lower lip when mouth closed; lateral line directed

anteriorly toward dorsal edge of upper eye; eye diameter

6.4-6. 8 in HL; upper eye overlapping anterior one-half to

three-fourths of lower eye; least vertical interorbital space

8. 3-8. 6 in HL; upper end of gill opening at level of ventral edge

of lower eye; no caudal-peduncle; depth at base of caudal

fin 1.5-1.65 in HL; longest dorsal ray 1.45 in HL; caudal fin

rounded, 5. 1-5.8 in SL; ocular-side pelvic fin on ventral edge

of body, its base slightly anterior to that of fin of blind side;

third pelvic ray longest, 2.0-2. 8 in HL, the tip reaching base of

second anal ray; lengthwise membranous ridge on dorsal rays

of both sides, progressively reduced posteriorly; small scales

on about basal half of membranous ridge of anterior dorsal rays

of ocular side; edge of membranous ridge of about first 15

blind-side dorsal rays with prominent cirri; colour of ocular

side in alcohol light brown, the scale edges darker than centers,

with faint irregular dark-edged pale markings over head and

body, and very irregular dark brown blotches, most much

larger than eye, the two largest on lateral line to either side of

middle of body; fins with dark dots. Largest of two known

specimens, 86.5 mm SL.

Remarks. Randall and Bartsch (2005) described a specimen

from Kwajalein Atoll, Marshall Islands identified as

Aseraggodes melanostictus by Woods in Schultz and collabor-

ators (1966) from Kwajalein Atoll, Marshall Islands as a

new species, A. heraldi. They illustrated an x-ray and two

photographs of the holotype of melanostictus from the Museum

fur Naturkunde in Berlin (ZMB 9814, 74 mm SL) and listed

differences from A. heraldi, notably its having unbranched

dorsal and anal rays.

The holotype of A. melanostictus was collected in 73 m off

the island of Bougainville. A second specimen, AMS 1.24499-

003, 86.5 mm, sent on loan from the Australian Museum, is

provisionally identified as this species. It was dredged in 115 m

NE of North Reef of the Great Barrier Reef (23°8'S, 152°12'E)

by W. Ponder et al. on 14 Dec 1977. It differs in having 13

instead of 11 dorsal pterygiophores anterior to the fourth neural

spine, and in some proportional measurements, but none

clearly beyond expected infraspecific variation.

Aseraggodes nigrocirratus sp nov.

Figure 12, Tables 1-3, 9

Holotype. AMS 1.26535-001, female, 79.0 mm, Australia, NSW, SE of

Evans Head, off Iluka, 29°20'S, 153°29'E, 40-51 m, trawl, FRV

“Kapala”, 24 May 1986.

Paratypes. AMS 1.321, 89.4 mm, Australia, NSW, Sydney, Port

Jackson, 33°51'S, 151°16'E, 1886; AMS 1.636, mature female, 68.8

mm SL, Australia, NSW, Sydney, Port Jackson, 33°51'S, 151°16'E, J.

Table 9. Proportional measurements of type specimens of Aseraggodes nigrocirratus as percentages of standard length

Holotype

Paratypes

AMS

1.26535

BPBM

39610

AMS

IA.5449

AMS

IA.5449

AMS

1.636

AMS

1.321

Standard Length (mm)

79.0

52.3

55.0

56.8

68.8

89.4

Body depth

41.0

38.2

39.7

40.7

42.5

42.7

Body width

8.1

7.6

7.1

7.2

7.6

7.7

Head length

21.7

21.0

21.4

21.3

20.8

21.3

Snout length

7.4

7.5

7.2

7.3

7.2

Preorbital length

6.2

6.1

5.9

6.0

5.9

5.8

Eye diameter

3.7

4.0

3.8

3.7

3.4

Interorbital width

2.0

1.9

2.1

2.2

Upper-jaw length

6.3

6.0

6.4

6.3

6.1

6.4

Caudal-base depth

13.9

12.8

13.2

14.1

14.7

14.6

Predorsal length

5.7

6.1

5.7

5.8

5.7

5.6

Preanal length

26.6

25.7

26.5

26.8

24.8

25.6

Prepelvic length

19.2

20.3

19.8

19.3

19.9

19.4

First dorsal ray

7.8

7.1

7.5

8.1

8.0

7.6

Longest dorsal ray

12.7

13.3

13.5

13.2

12.0

First anal ray

7.7

7.9

8.0

8.5

7.8

7.3

Longest anal ray

12.6

13.3

13.7

13.1

13.2

12.1

Caudal-fin length

21.4

23.0

23.2

broken

23.1

20.8

Pelvic-fin length

10.7

11.2

11.6

11.4

10.9

10.5

206

John E. Randall

Figure 12. Holotype of Aseraggodes nigrocirratus, AMS 1.26535-001,

79.0 mm SL, off Iluka, NSW (KJ. Graham).

Hunt, 1886; AMS IA.5449, 2: 55.0-56.8 mm, NSW, Pittwater,

33°38'S, 151°18'E, dredge, M. Ward, 1932; NMV A 21382, 61 mm,

Sydney, Coogee Beach, 33°55'S, 151°15'E, R.H. Kuiter, 26 Jan 1980;

AMS 1.26535-008, 2: 65.5-82 mm, and ROM 77681, 67 mm, same

data as holotype.

Diagnosis. Dorsal rays 63-68; anal rays 47-53; most dorsal

and anal rays branched; lateral-line scales 61-67, including 8-9

anterior to a vertical at upper end of gill opening; one to three

pores beneath some scales on ocular side of body near base of

dorsal and anal fins; vertebrae 36-38; dorsal pterygiophores

anterior to fourth neural spine 8-9; body depth 2.35-2.65 in

SL; head short, length 4.7-4.85 in SL; upper lip not overlap-

ping lower lip when mouth closed; no prominent cirri on ven-

tral edge of head; lateral line aligned with dorsal third of upper

eye; eye diameter 4.6-6.25 in HL; upper eye overlapping anter-

ior half to two-thirds of lower eye; interorbital space narrow,

the vertical distance separating eyes about half eye diameter; no

caudal peduncle; longest dorsal ray 1.5-1.75 in HL; caudal fin

slightly pointed, its length 3.95-4.8 in SL; pelvic fins 1.8-2.0

in HL, the tip of longest ray reaching base of third anal ray;

ocular side light brown with numerous small variable dark

brown spots of variable size, some in form of a small cross;

cirri on lengthwise membranous ridge of rays of dorsal and

anal fins dark brown (appearing as small dark spots without

magnification).

Description. Dorsal rays 64 (63-68); anal rays 49 (47-53); dorsal and

anal rays unbranched (none double-branched), except unbranched first

9 dorsal and anal rays (first 19 dorsal and anal rays and last few rays

of 43.6-mm paratype); caudal rays 18, branched, middle 16 double-

branched; pelvic rays 5, branched; lateral-line scales on ocular side 65

(61-67), including 8-9 anterior to a vertical at upper end of gill open-

ingl scales above lateral line to dorsal-fin base on ocular side about 20;

scales below lateral line to anal-fin base about 24 (23-24); vertebrae

37 (two paratypes with 36, five with 37, and two with 38); erisma

(counted as the first dorsal pterygiophore) about twice as thick as

remaining pterygiophores, its inner two-thirds branched; only erisma

before the second neural spine; space between second and third neural

spines with 5 pterygiophores; space between third and fourth neural

spines with 3 (2-3) pterygiophores, total 9 (8-9) dorsal pterygiophores

anterior to fourth neural spine; ventroanterior margin of urohyal

forming an angle of about 80° (65-80°), the comer slightly rounded.

Body depth 2.45 (2.35-2.65) in SL; body width 5.05 (5.05-5.85) in

body depth; dorsal profile of head slightly more convex than ventral;

HL 4.6 (4.7-4.85) in SL; snout length 2.95 (2.7-3 .0) in HL; preorbital

length 3.5 (3.3-3.75) in HL; eye diameter 5.85 (4.6-6.25) in HL; least

vertical interorbital width 10.8 (8.9-11.1) in HL; upper eye over-

lapping anterior one-half to four-fifths of lower eye; upper end of gill

opening on a level one-half to one eye diameter below ventral edge of

lower eye; no caudal peduncle (base of lowermost caudal ray above

or slightly anterior to base of last anal ray); depth of body at base of

caudal fin 1.55 (1.4-1.75) in HL.

Front of upper lip not overlapping lower lip when mouth closed;

maxilla extending slightly posterior to a vertical at front edge of lower

eye, the upper jaw length (measured on blind side) 3.45 (3.3-3.6) in

HL; blind side of upper and lower jaws with a dense band (broader

posteriorly) of slender, inward-projecting, slightly curved teeth, up to

about 7 rows in lower jaw and 5 or 6 in upper jaw (teeth on upper jaw

beneath a thin labial fold, hence difficult to see); anterior nostril a

tapering membranous tube anterior to upper edge of lower eye, just

reaching fleshy base of orbit when laid back (reaching to edge of eye-

ball in smaller paratypes), its length in holotype about three-fourths

eye diameter; posterior nostril an oblique slit in labial groove in front

of base of lower eye; anterior nostril of blind side a slender mem-

branous tube just above anterior third of upper lip; posterior nostril of

blind side a short strongly tapering membranous tube dorsoposterior to

anterior nostril; intemarial distance on blind side about three-fourths

eye diameter.

Scales ctenoid on both sides (cycloid on lateral line and partially

embedded); scales of ocular side of body usually with 8-10 cteni (up

to 12 in largest paratype); one to three pores beneath some scales on

ocular side of body near base of dorsal and anal fins; 3 rows of scales

in interorbital space, with about another 3 rows extending onto medial

and anterior part of each eye; scales on ocular side of head progres-

sively smaller anteriorly and ventrally, the most anterior at front of

snout very small and without cteni; fleshy front of snout with small

short cirri on edge; scales on blind side of head gradually replaced

anteriorly by papillae, ending with small slender papillae on edge of

snout; broad zone of small fleshy papillae on blind side dorsal to upper

jaw; ventral edge of head with cirri as long as half eye diameter; no

cirri along edge of operculum at gill opening on either side. Lateral line

straight midlaterally on both sides of body, directed on ocular side

toward dorsal third of upper eye; supratemporal branch of lateral line

on blind side of head faintly visible, continuing dorsally 2 to 3 scale

rows below dorsal fin, persisting to about middle of body.

Dorsal and anal fins with a basal sheath of 2 to 3 rows of scales;

lengthwise thin membranous ridge basally on dorsal and anal rays with

well-spaced dark brown cirri on free edges except posterior rays; small

scales basally on membranous ridges of anterior dorsal rays on both

sides (first 19 rays of holotype), the height of the scale bands pro-

gressively shorter posteriorly; about basal third of caudal fin with

scales on both sides, with small, well-spaced, isolated scales, with a

few prominent cteni on each side of rays to within outer third of fin.

Origin of dorsal fin anterior to lower fourth of upper eye, the pre-

dorsal length 3.8 (3.25-3.8) in HL; first dorsal ray 2.8 (2.6-2.95) in

HL; longest dorsal ray 1.7 (1.5-1.75) in HL; origin of anal fin below

base of fifteenth or sixteenth dorsal ray and about one-half eye

diameter behind posterior end of operculum, the preanal length 3.8

(3.75^1.15) in SL; length of first anal ray 2.8 (2.5-2.9) in HL; longest

anal ray 1.7 (1.5-1.75) in HL; caudal fin slightly pointed, relatively

shorter with growth, 4.7 (3.95 — 4.8) in SL; ocular-side pelvic fin on

ventral edge of body; base of blind- side fin adjacent anteriorly, its

origin slightly posterior to ocular-side fin; third pelvic ray of each fin

longest, reaching to base of third anal ray, 2.0 (1. 8-2.0) in HL; anus

directly before base of first anal ray at end of short semicircular

column of ridged fleshy tissue, an upper anterior triangular part cover-

ing opening; genital papilla on ocular side of base of first anal ray, very

large, its length equal to orbit diameter; membrane from near base of

last pelvic ray of ocular side joined to extreme base of genital papilla.

Colour of holotype in alcohol: ocular side light grey-brown with

numerous small dark brown spots on head, body, and basal scaly part

A review of soles of the genus Aseraggodes

207

Figure 13. Holotype of Aseraggodes normani, BMNH 1925.7.22.73,

109 mm SL, Queensland.

of caudal fin, some as small dark crosses from dark edges of adjacent

scales; dorsal and anal fins with translucent membranes, the rays light

yellowish brown, most with 1-3 small dark brown spots associated

with a fleshy cirrus; naked part of caudal fin like dorsal and anal fins,

but small dark spots on the rays without a cirrus; blind side of body

pale yellowish, slightly dusky in a broad peripheral zone. Figure 12

taken of a fresh specimen, very similar to colour in preservative; pale

spots on figure from missing scales.

Etymology. Named nigrocirratus from the Latin, in reference to

the black cirri on the dorsal and anal rays of the ocular side of

this species.

Remarks. Aseraggodes nigrocirratus is presently known only

from NSW from latitudes 29°20'S to 33°55'S, from inshore to

about 50 m.

Randall and Melendez (1987) mentioned this species in

their description of Aseraggodes bahamondei as probably

undescribed, noting that it also has pores under some scales of

the ocular side near the base of the dorsal and anal fins. They

attributed the presence of these pores in bahamondei to the pro-

duction of a strong skin toxin, probably comparable to that of

the species of Pardachirus (Clark and George, 1979).

Although Aseraggodes bahamondei is similar in its colour-

ation, general morphology, and the low number of dorsal ptery-

giophores anterior to the fourth neural spine to A. nigrocirratus,

it is readily distinguished by having a caudal peduncle (though

this difference is actually very slight), 39 to 40 vertebrae,

75-86 lateral-line scales, and by attaining a significantly larger

maximum size of at least 156 mm SL.

Of the known species of the genus, A. nigrocirratus is most

similar to A. lenisquamis, described above. The two have the

same vertebral count (36-38), the same number of dorsal

pterygiophores anterior to the fourth neural spine (8-9), and

nearly the same number of dorsal and anal rays and lateral-line

scales (see Tables 1-3). They differ mainly in the structure of

the scales. Those of nigrocirratus are typical of most species

of the genus, with prominent cteni that project well beyond the

posterior margin of the scales, and the exposed surface of the

scales with a pattern like miniature scales. The scales of

lenisquamis are generally more pointed posteriorly and are

covered with soft cutaneous tissue, such that only the tips of the

cteni project beyond the scale margins (each capped with soft

tissue). Also the lateral-line scales and scattered other scales of

lenisquamis have small fleshy cirri. Three different propor-

tional measurements of the two species are evident in Tables 7

and 9. A. lenisquamis has a longer snout, broader caudal-fin

base, and is broader, and shorter caudal fin.

Six specimens of AMS 1.24035-003, 42.0-52.3 mm SL, col-

lected in 1 m at Wagstaff Point, Brisbane Waters, NSW,

33°28'S, 151°21'W, in 1982 include four provisionally

identified as A. nigrocirratus and two of about 45 mm SL that

appear to be hybrids of A. lenisquamis and A. nigrocirratus.

The latter two specimens have scales of both types, though

mainly those typical of A. lenisquamis, and only a few short

cirri on lateral-line scales near the middle of the body. Both

specimens are misshapen and difficult to measure. The four

specimens identified as A. nigrocirratus are excluded from the

type series.

Similarly, the two specimens of AMS 1.38279-001,

79.4-83.0 mm, collected by trawl in 48 m off the Clarence

River, NSW at 29°19.5'S, 153°24'E, include one of

A. lenisquamis (the larger) and one that is regarded as a hybrid

of A. nigrocirratus and A. lenisquamis. This lot is also un-

divided, and the specimen of lenisquamis is not designated as a

paratype. The presumed hybrid has scales characteristic of each

of the species, and it is intermediate in the three proportional

measurements that are diagnostic.

Aseraggodes normani Chabanaud

Figure 13, Tables 1-3

Aseraggodes melanostictus . — Norman, 1926: 290, fig. 12 (off

Gladstone, Queensland) (non Peters)

Aseraggodes normani Chabanaud, 1930: 241.

Material examined. Queensland: BMNH 1925.7.22.73, 109 mm, holo-

type of Aseraggodes normani. WSW of Townsend Island, 22°21.5'S,

150°25'E, AMS 1.34377-005, 83 mm. 2 km NE of Cape Clinton,

22°32'S, 150°48'E, AMS 1.34399-032, 93.5 mm. 2 km NE of Cliff

Point, 22°35'S, 150°49’E, AMS 1.34361-024, 3: 74.5-89 mm. Off

Gladstone, AMS IA.2993, 104.5 mm. Off Bustard Head, 24°1’S,

151°46'E, AMS IB. 1 105, 97.5 mm. Moreton Bay, AMS 1.484, 116 mm.

Type locality. Coast of Queensland.

Diagnosis. Dorsal rays 64-71, anal rays 50-52; dorsal and anal

rays unbranched (Chabanaud incorrect in reporting bifid tips);

pelvic rays 5; caudal rays 18, 14 branched; lateral-line scales

68-73 (counted to origin of dorsoanterior branch on head); only

tips of cteni projecting beyond posterior edge of scales, with at

most 8 cteni tips posteriorly on body, fewer anteriorly); eyes

separated by 3 rows of scales at narrowest place, with an

additional row medially and anteriorly on each eye; vertebrae

35-36 (usually 35); dorsal pterygiophores anterior to fourth

neural spine 10-11; body depth 2.3-2.55 in SL; HL 4.4-4.8 in

SL; snout length 2.3-2.5 in HL; scales anteriorly on head

replaced by slender cirri, progressively longer, those at ventral

edge of head and front of snout up to three-fourths eye dia-

meter in length; lateral line aligned with dorsal edge of upper

eye, ending with a dorsoanterior branch of 7-9 pored scales,

straight branch of 4-6 scales, and ventral branch of 7-9 scales;

no pores detected beneath scales on ocular side of body; eye

diameter 5. 0-6. 5 in HL; upper eye overlapping anterior one-

half to three-fourths of lower eye; narrowest vertical interor-

bital space 8.2-9.0 in HL; upper end of gill opening at level of

208

John E. Randall

ventral fleshy edge of lower eye; tubular anterior nostril broad,

not reaching fleshy base of lower eye when laid back; no

caudal peduncle; depth at base of caudal fin 1.55-1.8 in HL;

caudal fin rounded, its length 4. 1^1.5 in SL; longest dorsal ray

1.35-1.5 in HL; blind side of dorsal and anal rays with a length-

wise thin membranous ridge, broad at base and narrowing as it

extends up to three-fourths ray length anteriorly, progressively

shorter and narrower posteriorly; edge of membrane on anter-

ior rays of blind side of dorsal fin with cirri; pelvic fins

1.9-2.05 in HL, the tip of longest ray extending to base of

second or third anal ray; ocular-side pelvic fin distinctly

anterior and larger than fin of blind side; both fins broadly

joined by membrane from their fifth rays and jointly to the large

genital papilla about one-half length above its base; colour in

alcohol light brown, densely dotted with black and short black

scale edges; scattered roundish pale spots smaller than eye var-

iously present, some free of black dots; median fins with black

dots, but fewer than on body; one specimen with a few large

dark blotches on lateral line and on either side of lateral line.

Largest specimen examined, 116 mm SL.

Remarks. Known only from off the coast of Queensland from

22.5° to 27° S, taken by trawls at depths of 15 to 27 m. Norman

(1926) reported three specimens, 130-142 mm total length,

collected off Queensland during the experimental cruises of

R/V “Endeavour”, 1909-1914, as Aseraggodes melanostictus

Peters, previously known from a single specimen taken in 73 m

off New Britain. Chabanaud (1930a) found one of Norman’s

three specimens at the Natural History Museum, London

(BMNH). Recognizing it as misidentified, he named it in

honour of J.R. Norman. An additional (non-type) “Endeavour”

specimen in the Australian Museum was collected off

Gladstone, 23°S 151°E on 9 Jul 1910.

Chabanaud (1943) established the valid genus Synclidopus

for the species Solea macleayanus, but he erred in placing

Aseraggodes normani as a second species of this genus.

Aseraggodes pelvicus sp. nov.

Figure 14, Tables 1-3, 10

Holotype. AMS IB.6134, 67.5 mm, Australia, Queensland, Great

Barrier Reef, Swain Reefs, Gilette Cay, rotenone, Australian Museum

Expedition, October 1962.

Diagnosis. Dorsal rays 71; anal rays 49; lateral-line scales 81;

vertebrae 36; dorsal pterygiophores anterior to fourth neural

spine 14; body depth 2.45 in SL; HL 4.75 in SL; eye diameter

6.4 in HL; upper eye overlapping anterior half of lower eye;

interorbital space narrow, 12.0 in HL; caudal peduncle present,

its depth 1.45 in HL, its length 10.0 in HL; prominent lappet-

like cirri on front of snout and ventral edge of head; lateral line

aligned with dorsal edge of upper eye; longest dorsal ray 1.25

in HL; membranous edges of anterior dorsal rays with a row of

tubercle-like papillae, many with cirri, a few darkly pigmented;

caudal fin rounded and moderately long, 3.4 in SL; pelvic fins

long, 1 .6 in HL, the tip of longest ray reaching base of fifth anal

ray; colour of ocular side in alcohol pale yellowish brown with

numerous irregular pale spots, none as large as eye, and many

smaller brown blotches and dots, the most conspicuous along

Figure 14. Holotype of Aseraggodes pelvicus, AMS IB.6134, 67.5

mm, Swains Reefs, Great Barrier Reef.

Table 10. Proportional measurements of holotype of Aseraggodes

pelvicus as percentages of standard length

Standard Length (mm)

67.5

Body depth

41.2

Body width

7.8

Head length

21.0

Snout length

7.6

Preorbital length

8.1

Eye diameter

3.3

Interorbital width

2.1

Upper-jaw length

6.7

Caudal-peduncle depth

14.2

Caudal-peduncle length

1.7

Predorsal length

6.5

Preanal length

25.2

Prepelvic length

20.3

First dorsal ray

7.5

Longest dorsal ray

16.5

First anal ray

10.3

Longest anal ray

16.6

Caudal-fin length

29.5

Pelvic-fin length

10.2

middle of lateral line.

Description. Dorsal rays 71, branched except first 9, the tips broadly

free; anal rays 49, branched except first; caudal rays 18, the uppermost

and lowermost unbranched, the middle 14 double-branched; pelvic

rays 5, branched except last ray of blind side; lateral- line scales 81,

including 1 1 anterior to a vertical at upper end of gill opening; scales

on ocular side above lateral line to dorsal-fin base 27; scales below

lateral line to anal-fin base about 33; vertebrae 37; erisma (counted as

the first dorsal pterygiophore) about twice as thick as remaining

pterygiophores, its inner three-fourths narrowly branched; next 2

pterygiophores before tip of second neural spine; space between

second and third neural spines with 7 pterygiophores; space

between third and fourth neural spines with 3 pterygiophores (total of

13 dorsal pterygiophores anterior to fourth neural spine); ventro-

anterior margin of urohyal forming an angle of about 80°, the comer

slightly rounded.

Body depth 2.45 in SL; body width 5.3 in body depth; HL 4.75 in

SL; ventral profile of head below mouth nearly straight; eye diameter

6.4 in HL; least vertical interorbital width 10.0 in HL; a vertical at pos-

terior edge of upper eye (edge of dark eyeball) passing through middle

of lower eye; ventral edge of lower eye one-half eye diameter above

level of upper end of gill opening; caudal-peduncle depth 1.45 in HL;

caudal-peduncle length 10.0 in HL.

A review of soles of the genus Aseraggodes

209

Snout not overhanging lower lip when mouth closed; maxilla

extending to below middle of lower eye, the upper jaw length (blind

side) 3.15 in HL; jaws too firmly closed to obtain details of dentition;

anterior nostril a tapering membranous tube anterior to upper edge of

lower eye, reaching anterior edge of eye ball when laid back, its length

equal to eye diameter; posterior nostril an oblique slit in labial groove

directly in front of ventral third of lower eye; anterior nostril of blind

side a very slender membranous tube nearly an eye diameter in length

just above anterior third of upper lip; posterior nostril of blind side an

eye diameter dorsoposterior to anterior nostril, covered by a broadly

curved pointed flap on ventral side.

Scales ctenoid on both sides (except those of lateral line, partially

embedded); scales of ocular side with 11-14 cteni; 3 rows of scales in

interorbital space, with about another 4 or 5 rows extending onto

medial and anterior part of each eye; scales on both sides of head pro-

gressively smaller anteriorly and ventrally, with fewer cteni, replaced

more anteriorly with isolated papillae and at front of snout by cirri; 15

large lappet-like cirri along ventral edge of head below mouth, and

several at front of snout; slender well-spaced cirri along edge of

opercle at the gill opening on both sides. Lateral line straight mid-

laterally on both sides, directed on ocular side toward dorsal edge

of upper eye; supratemporal branch of lateral line curving at front of

head on blind side, continuing posteriorly onto body 2 to 3 scale rows

below dorsal fin, becoming obscure posterior to base of about 23rd

dorsal ray.

Dorsal and anal fins with a basal sheath of 2 to 3 rows of scales; a

thin membranous lengthwise ridge on dorsal and anal rays of both

sides, progressively smaller posteriorly; front of dorsal fin on ocular

side densely covered with papillae to base of about fourth ray; edge of

membranous ridge on rays of about anterior third of dorsal fin with a

row of small tubercle-like papillae, most ending in a slender cirrus;

rows of similar papillae on anterior dorsal rays of blind side, but with-

out cirri; small scales extending out on anterior dorsal rays, progres-

sively reduced posteriorly, and absent beyond 25th dorsal ray; small

tubercle-like papillae on basal part of rays of anal fin on ocular side,

disappearing posterior to 20th ray.

Origin of dorsal fin anterior to lower edge of upper eye, the pre-

dorsal length 3.25 in HL; first dorsal ray 2.8 in HL; first 5 dorsal rays

free of membrane at tips; longest dorsal ray 1.25 in HL; origin of anal

fin below base of 22nd dorsal ray and slightly behind posterior end of

opercular membrane, the preanal length 4.0 in SL; length of first anal

ray 2.05 in HL; longest anal ray 1.25 in HL; caudal fin rounded and

moderately long, 3.4 in SL; pelvic fins adjacent on ventral edge of

body; ocular-side pelvic fin slightly longer, the second and third pelvic

rays longest, reaching to base of fifth anal ray, 2.05 in HL; anus anter-

ior to first anal ray; genital papilla on ocular side of anus, about equal

in length to pupil diameter.

Colour of ocular side in alcohol pale yellowish brown with numer-

ous irregular pale spots (probably near-white in life), some approach-

ing eye size; many brown blotches and brown dots (mainly from dark

posterior edge of scales) scattered on head and body, the middle three

or four of about ten along lateral line the most conspicuous (each

covering 4 or 5 lateral- line scales); dorsal, anal, and pelvic fins with

pale yellowish rays and translucent membranes; a few isolated cirri of

dorsal fin brown; scaled basal part of caudal fin coloured like body, the

rest of fin with pale yellowish rays and transparent membranes, except

for small scales on rays that are brown; blind side of body pale

yellowish brown with no markings.

Etymology. Named pelvicus from the Latin pelvis, in reference

to the long pelvic fins, the longest for species of the genus.

Remarks. This species is known only from the holotype from

the Swain Reefs of the Great Barrier Reef of Australia. It

appears to be most closely related to Aseraggodes ramsaii from

Lord Howe Island and New Caledonia and to A. whitakeri,

wide-ranging in islands of the Pacific. It shares the same counts

of vertebrae, dorsal pterygiophores, dorsal rays, and anal rays

with these two species. It differs from both in its smaller head,

smaller eye, and longer dorsal and pelvic fins. Aseraggodes

ramsaii has a higher number of lateral-line scales, 86-88, com-

pared to 81 for pelvicus, but additional specimens are needed to

confirm this difference. The series of small tubercle-like papil-

lae at the edge of the membranous ridges of the dorsal and anal

rays, many with a small cirrus, and the long pelvic fins appear

to be unique to A. pelvicus.

Aseraggodes ramsaii (Ogilby, 1889)

Figure 15, Tables 1-3

Solea ramsaii Ogilby, 1889: 70, pi. 3 fig. 4.

Aseraggodes haackeanus ramsaii — Munro, 1957: 73.

Aseraggodes haackeanus. — Hoese in Allen et al., 1976: 437.

Aseraggodes ramsaii. — Randall and Melendez, 1987: 105.

Material examined. Lord Howe Island: AMS 1.1951, 57 mm, holotype

of Solea ramsaii', AMS 1.5387, 63.5 mm; BPBM 34265, 41.7 mm. New

Caledonia: lagoon near SE end of St Vincent Pass (22°2'6"S,

165°57'48"E).

Type locality. Lord Howe Island.

Diagnosis. Dorsal rays 69-72, branched except first 14-22

rays; anal rays 47-50, all but branched; caudal rays 18 (upper-

most and lowermost two rays simple, middle 12-14 double-

branched); pelvic rays 5; lateral-line scales on ocular side

86-88, including 12 anterior to a vertical at upper end of gill

opening; scales with 10-14 cteni; scales progressively shorter

anteriorly on head and with fewer cteni; 2 rows of scales in

narrowest interorbital space, with small scales in 4 or 5 rows

extending onto median and anterior edges of eyes; broad

lappet-like cirri on ventral margin of head; vertebrae 36; dorsal

pterygiophores anterior to fourth neural spine 13-14 (14 in

holotype); body depth 2.4-2.6 in SL; HL 4.45-4.5 in SL; upper

lip slightly overhanging lower lip when mouth closed; eye

diameter 4.05-4.65 in HL; posterior edge of upper eye over

middle of lower eye; interorbital space narrow, the least

vertical distance separating eyes 12.3-13.3 in HL; lateral line

angling upward anteriorly, directed toward dorsal edge of upper

eye; ventral edge of lower eye about one-half eye diameter

Figure 15. Aseraggodes ramsaii, BPBM 34265, 41.7 mm, New

Caledonia.

210

John E. Randall

above upper end of gill opening; caudal peduncle present, its

length 8.1-10.8 in HL; longest dorsal ray 1.65-1.7 in HL;

caudal fin rounded, 3.85-3.95 in SL; ocular-side pelvic fin on

ventral edge of body and slightly anterior to fin of blind side,

the second or third ray longest, 2.3-2.35 in HL, the tip reach-

ing to base of second or third anal rays; small scales extending

nearly to dorsal-fin margin anteriorly on both sides, the scale

coverage gradually reduced to fifteenth to 20th ray, and absent

posteriorly; a thin membranous lengthwise ridge basally on

dorsal and anal rays, disappearing posteriorly; a dark brown

cirrus occasional on membranous ridges of dorsal rays on

ocular side. Colour of holotype in alcohol “pale yellow with

many small black spots and short wavy lines on head and body,

which on the lateral line take the form of streaks extending

from two to five scales” (Ogilby, 1889). Largest of 3 specimens

examined, 63.5 mm SL.

Remarks. Munro (1957) regarded Aseraggodes ramsaii as a

subspecies of A. haackeanus (Steindachner). He was followed

by Hoese in Allen et al. (1976), who reported two specimens

from Lord Howe Island collected from sand outside the lagoon

in 5-25 m (specimens not found). Randall and Melendez

(1987) showed that A. ramsaii and A. haackeanus are readily

separated by counts of dorsal rays, anal rays, and lateral-line

scales. There is also a consistent difference in the number of

dorsal pterygiophores before the fourth neural spine, 14-15 for

ramsaii and 7-9 for A. haackeanus.

Aseraggodes ramsaii is otherwise known from the holotype

in the Australian Museum, one other AMS specimen from Lord

Howe Island, and one reported here from New Caledonia col-

lected by Michel Kulbicki and the author in 2-4 m. The New

Caledonia specimen was first believed to be Aseraggodes

holcomi Randall, described from five specimens from the

Hawaiian Islands. It was reidentified as A. ramsaii after com-

parison with the Lord Howe Island specimens. The two species

are very similar, differing in A. holcomi having a broader

interorbital space (6.8-10.9 in HL, compared to 12.2-13.2 for

ramsaii ), longer pelvic fins (1.85-2.1 in HL, compared to

2.3-2.35 for ramsaii ), and a higher average count of lateral-line

scales (87-96, compared to 86-88 for ramsaii ).

Aseraggodes whitakeri Woods

Figures 16, 17, Tables 1-3

Aseraggodes melanostictus. — Schultz, 1943: 60 (Hull Island,

Phoenix Islands) (non Aseraggodes melanostictus Peters).

Aseraggodes whitakeri Woods in Schultz et al., 1966: 71, fig. 150.

Aseraggodes sp. — Wass, 1984: 31 (Taema Bank, Tutuila, American

Samoa).

Material examined. Marshall Islands: Rongelap Atoll, USNM 141765,

38 mm (holotype). Caroline Islands: Kapingamarangi Atoll, CAS

205944, 28 mm. Coral Sea: Chesterfield Bank, BPBM 33731, 22 mm.

New Caledonia, ROM 64828, 42 mm; ROM 64829, 37 mm; ROM

64831, 21.5 mm. Fiji: Ringgold Islands, BPBM 20863, 32 mm.

American Samoa: Tutuila, Taema Bank, BPBM 24113, 43.0 mm;

BPBM 24130, 2: 25-25.5 mm. Phoenix Islands: Hull Island, USNM

115223, 34 mm. Society Islands: Moorea, ROM 61360, 14 mm; ROM

61361, 23 mm.

Type locality. Rongelap Atoll, Marshall Islands.

Diagnosis. Dorsal rays 71-78; anal rays 48-53; pelvic rays 5 (3

in one side on one aberrant specimen); caudal rays 18, 14

branched (17 rays, 13 branched in one specimen); lateral-line

scales 77-86, including 10-12 anterior to a vertical at upper

end of gill opening); most scales of ocular side of body with

8-10 cteni (up to 11 on largest specimens); narrowest inter-

orbital space with 2 scales; eyes with only 1 or 2 scales extend-

ing onto medial edge, and only about 3 rows anteriorly; verte-

brae 36-38; dorsal pterygiophores anterior to fourth neural

spine 14-15; body depth 2.55-2.75 in SL; HL 4.1^1.35 in SL;

an overhanging fleshy snout, the lower edge of upper lip

usually extending ventral to lower lip and jutting anterior to

profile of head below mouth (more evident in smaller speci-

mens); eye diameter 4.7-5.55 in HL; upper eye varying from

one-half to full eye diameter anterior to lower eye; interorbital

space very narrow, the vertical distance separating eyes less

than half eye diameter; caudal peduncle present, its length

7.2-10.0 in HL; ventral edge of head posterior to mouth with

10-19 lappet-like cirri; edge of operculum at gill opening

with slender well-spaced cirri on both sides; edge of lengthwise

membranous ridges of dorsal and anal rays of ocular side with

cirri, reduced and disappearing on about posterior half of fins;

cirri also present on rays of blind side, but fewer and restricted

to more anterior rays; lateral line aligned with dorsal edge of

upper eye; upper end of gill opening in line with ventral fleshy

edge of lower eye; anterior nostril reaching fleshy base of lower

eye when laid back; caudal fin slightly pointed, its length

3.05-3.65 in SL; longest dorsal ray 1.4— 1.8 in HL; anal rays

and all but anterior dorsal rays of larger specimens branched

/ ' > - % ’

v \ . 4 * ? ; *> l } ~'MV' • :-J

. . ■ ’ . .. . ;.. , ■ V ■ ' . ■/' '

t ,

i .;. vs.,;

Figure 16. Aseraggodes whitakeri , BPBM 24113, 43.0 mm SL, Figure 17. Aseraggodes whitakeri , ROM 64828, 42.0 mm SL, New

Tutuila, American Samoa. Caledonia (R. Winterbottom)

A review of soles of the genus Aseraggodes

211

Figure 18. Aseraggodes diringeri, ROM 56917, 43.4 mm SL, Mayotte,

Comoro Islands (R. Winterbottom).

distally; pelvic fins long, 1.65 -1.85 in HL, the tip of longest

ray extending to base of third or fourth anal ray; colour in

alcohol of ocular side of most specimens pale tan without any

dark markings; specimens from American Samoa with faint

dark blotches in 3 rows on the ocular side, one row below base

of dorsal fin, one along lateral line, and one above base of anal

fin. Yellowish tan with many irregular pale markings partially

outlined with dusky brown; fins pale with dusky specks

Remarks. Richard C. Wass recorded the brief colour note

(above) on the label of the largest of three specimens, 43 mm

SL, that he collected in American Samoa. A photograph of his

specimen after 20 years in alcohol is presented as Figure 16.

Figure 17 is a photograph of a fresh 42-mm specimen from

New Caledonia taken by Richard Winterbottom. Most speci-

mens of this species have an overhanging snout, the tip of the

upper lip anterior and ventral to the lower lip, as shown in

somewhat exaggerated form in the drawing of the holotype

(Woods in Schultz et al., 1966: fig. 150). Unfortunately, Woods

placed Aseraggodes whitakeri in the couplet of his key begin-

ning with “snout not overlapping tip of the lower jaw”, and he

wrote in his description, “snout not hooked over lower jaw.” He

evidently confused this part of his description with that of

A. smithi Woods. The dorsal-ray count of Woods is corrected

from 72 to 76, and the anal-ray count from 51 to 52.

The drawing of the holotype is in error in the relative posi-

tion of the eyes. The upper eye should overlap only about

the anterior tenth of the lower eye. Also the dark blotches of the

drawing are in the wrong positions. Those along the dorsal edge

of the body should be 5 or 6 dorsal rays anterior to the position

depicted. The largest dark blotch on the lateral line is also

placed too posteriorly. It should be below the base of the

fiftieth dorsal ray.

Wass (1984) reported three specimens from American

Samoa. The largest, BPBM 24113, collected on the Taema

Bank in 21.5 m, has only 3 rays in the right pelvic fin, a

shorter base, and the left fin is shorter than normal. The other

two specimens, BPBM 24130, from the same bank, but in 35

m, have normal pelvic fins.

The first specimen of this species was reported by Schultz

(1943) from a channel at Hull Island, an atoll in the Phoenix

Islands. He wrote, “it may be a specimen of Aseraggodes

melanostictus (Peters).” The holotype was collected in 6 m

from a lagoon coral head at Rongelap Atoll, Marshall Islands

by V.E. Brock and E.S. Herald in 1946. A second specimen

from Micronesia was taken on a reef flat at Kapingamarangi

Atoll in the Caroline Islands. The remaining specimens from

the South Pacific are from the Chesterfield Bank in the Coral

Sea and New Caledonia to the Society Islands. Depths of

capture ranged from a tidepool at low tide in less than 0.2 m on

the Chesterfield Bank to 37 m on the Taema Bank, American

Samoa.

Aseraggodes diringeri Quero, 1997 from the western Indian

Ocean (type locality Reunion) has the same counts of fin-rays,

lateral-line scales, vertebrae, and dorsal pterygiophores as

A. whitakeri. It differs from whitakeri in its larger size. The

holotype of diringiri measures 104 mm SL, and the one

paratype is 59 mm in SL. Of 23 specimens of diringeri

examined by the author, ten are larger than the largest

whitakeri (43 mm SL). Judging from a single colour

photograph of each species, the white markings of the body of

whitakeri are nearly all vermiculate, as shown in Figure 17,

whereas they are nearly all as discrete spots in diringeri

(Fig. 18).

As noted in the two species accounts above, A. whitakeri is

also similar to A. pelvicus and A. ramsaii.

Acknowledgements

I am grateful to the following for the loan of specimens of the

genus Aseraggodes : Hiromitsu Endo of Kochi University; Jon

Fong of the California Academy of Sciences; Dianne J. Bray

and Martin F. Gomon of Museum Victoria; Ofer Gon and Vusi

Mthombgeni of the South African Institute for Aquatic

Biodiversity; James Maclaine of the Natural History Museum,

London; Mark A. McGrouther and Sally Reader of the

Australian Museum; Kazuo Sakamoto of the University

Museum, University of Tokyo; Jeffrey T. Williams of the

Smithsonian Institution, and Mary Burridge and Richard

Winterbottom of the Royal Ontario Museum. Richard

Winterbottom and Ken J. Graham of NSW Fisheries are also

thanked for their valuable photographs. I am indebted as well

to Martine Desoutter of the Museum National d’Histoire

Naturelle in Paris for information on types and other specimens

of Aseraggodes , Paul Bartsch of the Museum fiir Naturkunde in

Berlin for data on the holotype of A. melanostictus , Loreen R.

O’Hara of the Bishop Museum, Sandra J. Raredon of the

Smithsonian Institution, and Sally Reader for providing x-rays.

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Starck W.A. II, Talbot, F.H., and Whitley, G.P. 1976. Annotated

checklist of the fishes of Lord Howe Island. Records of the

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Chabanaud, P. 1930b. Revision du genre Aseraggodes Kaup.

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dorsale chez les Soleidae (Teleosteens, Pleuronectiformes). Cybium

13(3): 271-279.

Clark, E., and George, A. 1979. Toxic soles, Pardachirus marmoratus

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Grant, E. 1987. Fishes of Australia. Grant: Zillmere, Qld. 480 pp.

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Gunther, A. 1909. Andrew Garrett’s Fische der Stidsee. Journal des

Museum Godeffroy 6(16): 261-388.

Hubbs, C.L. 1967. Review of Fauna Japonica/Soleina (Pisces) by

Akira Ochiai. Copeia 1967(2): 490^191.

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Pleuronectida e. Archiv fur Naturgeschichte 24(1): 94-104.

Kuiter, R.H. 1993. Coastal fishes of south-eastern Australia.

University of Hawaii Press: Honolulu, xxxi + 437 pp.

Jordan, D.S., and Evermann, B.W. 1898. The fishes of North and

Middle America: a descriptive catalogue of the species of fish-like

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Matsubara, K., and Ochiai, A. 1963. Report on the flatfish collected by

the Amami Islands expedition in 1958. Bulletin of the Misaki

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McCulloch, A.R. 1929-1930. A check-list of the fishes recorded from

Australia. Memoirs of the Australian Museum 5: x + 534 pp.

Munro, I.S.R. 1957. Handbook of Australian fishes, no. 17. Fisheries

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Norman, J.R. 1926. Flatfishes. Pp. 219-308 in: Biological Results of

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Ochiai, A. 1963. Fauna Japonica Soleina (Pisces). Biographical

Society of Japan: Tokyo, vi + 114 pp.

Ogilby, J.D. 1889. Reptiles and fishes of Lord Howe Island. Memoirs

of the Australian Museum 2: 49-74.

Ogilby, J.D. 1916. Edible fishes of Queensland. Parts IV-IX. Memoirs

of the Queensland Museum 5: 127-177.

Peters, W.C.H. 1877. Ubersicht der wahrend der von 1874 bis 1876

unter dem Commando des Hm. Capitan z. S. Freiherm von

Schleinitz ausgefiihrten Reise S.M.S. Gazelle gesammelten und

von de Kaiserlichen Admiralitat der Koniglichen Akademie der

Wissenschaften iibersandten Fische. Monatsberichte der Koniglich

preussischen Akademie zu Berlin 1876: 831-854.

Quero, J.-C. 1997. Soleidae et Cynoglossidae (Pleuronectiformes) de

Tile de la Reunion (Ocean Indien): description d’une nouvelle

espece. Cybium 21(3): 319-329.

Ramsay, E.P 1881. Description of two new species of Australian fish-

es. Proceedings of the Linnean Society of New South Wales 5:

462-463.

Randall, J.E. 1996. Two new soles of the genus Aseraggodes

(Pleuronectiformes: Soleidae) from the Hawaiian Islands. Pacific

Science 50(4): 427-440.

Randall, J.E. 2002. Aseraggodes holcomi, a new sole

(Pleuronectiformes: Soleidae) from the Hawaiian Islands. Pacific

Science 56(3): 247-253.

Randall, J.E. and Bartsch, P. 2005. Two new soles of the genus

Aseraggodes (Pleuronectidae: Soleidae) from Micronesia.

Micronesica 38: 125-139.

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Aseraggodes from Easter Island and Lord Howe Island, with com-

ments on the validity of A. ramsaii. Bishop Museum Occasional

Papers 27: 97-105.

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202(3): vii + 176 pp.

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Memoirs of Museum Victoria 62(2): 213-221 (2005)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

The species of Dasycercus Peters, 1875 (Marsupialia: Dasyuridae)

P. A. Woolley

Department of Zoology, La Trobe University, Bundoora Vic. 3086, Australia (p.woolley@latrobe.edu.au)

Abstract Woolley, P.A. 2005. The species of Dasycercus Peters, 1875 (Marsupialia: Dasyuridae). Memoirs of Museum Victoria

62(2): 213-221.

Two species of Dasycercus (mulgaras) are recognised. They can be distinguished by the form of the tail, the number

of upper premolar teeth in each jaw and, in the female, by the number of nipples in the pouch. Dasycercus blythi has a

non-crested tail, two upper premolars (and a diastema between P 2 and M 1 ) and six nipples; D. cristicauda has a crested

tail, three upper premolars (the third very small and sometimes present on one side only) and eight nipples. Both species

have a wide geographic range in the arid zone of Australia, and overlapping distributions. The identity of specimens pre-

viously referred to D. cristicauda in the W. B. Spencer collection of Museum Victoria has been reassessed and both

species found to be represented.

Keywords Marsupialia, Dasyuridae, Dasycercus blythi, Dasycercus cristicauda

Introduction

Four named forms of small carnivorous marsupials have been

assigned to the genus Dasycercus Peters, 1875; three concern

us here. The first was Chaetocercus cristicauda Krefft, 1867.

Krefft’s description was based on a single specimen sent to him

by F. G. Waterhouse of the then Institute Museum of Adelaide

(now the South Australian Museum). A second form,

Phascogale blythi Waite, 1904, was described from specimens

collected by A. C. Blyth in the Pilbara district of Western

Australia. Two of Blyth’s live animals had been left in the care

of B. H. Woodward, Director of the Western Australian

Museum and Art Gallery, and three were taken to Sydney,

where E. R. Waite of the Australian Museum made obser-

vations on the live animals before returning them to Blyth,

following which they escaped. Soon after, Woodward asked

Waite to examine the remaining two specimens, expressing the

wish that “if new, Mr Blyth’s name might be associated with

them”. Woodward himself (1902) had mentioned them as

“ Phascologale blighV without description, thereby erecting a

nomen nudum. The third, Phascogale hillieri Thomas, 1905,

was described from the skin of a specimen from Killalpaninna,

South Australia, presented by H. J. Hillier.

Iredale and Troughton (1934) and Tate (1947) recognised

two species of Dasycercus, namely D. blythi and D. cristi-

cauda and these authors placed D. hillieri in the synonomy of

D. cristicauda. Ride (1970) referred to only a single species,

D. cristicauda and later Mahoney and Ride (1988) formally

placed all three nominal species in the synonymy of

D. cristicauda. Consequently most authors have followed Ride.

Mahoney and Ride included Dasyuroides byrnei Spencer, 1896

in the same genus but there is no consensus on the inclusion of

byrnei in Dasycercus ; byrnei is not considered here.

On the basis of unpublished information (received from

M. Adams of the South Australian Museum) for the existence

of two genetically distinct forms of Dasycercus, referred to as

D. cristicauda and D. hillieri. The 1996 Action Plan for

Australian Marsupials and Monotremes (Maxwell et al., 1996),

listed D. cristicauda as “vulnerable” and D. hillieri as “endan-

gered”. Since then, in an unpublished report, Adams et al.

(2000) have given their reasons for the names they applied to

the two species. Further support for the existence of two genet-

ically distinct forms of Dasycercus, based on gene sequences

obtained from three whole mitochondrial genes (cytb, 12SrRNA

and 16 SrRNA ) has been obtained by M. Westerman and

C. Krajewski (pers. comm.).

I first became aware of the existence of two distinct forms

of Dasycercus when, in 1967, I collected specimens in south-

western Queensland that differed from others that I had col-

lected previously in the Northern Territory. The morphology of

the tail clearly distinguished them as did the number of nipples

in the pouch of females. There were also differences between

the two forms in dentition. I maintained animals from both

areas in captivity, and reported observations on the reproduc-

tion of animals from the Northern Territory (Woolley, 1971)

and on the burrows made by animals from both the Northern

Territory and Queensland (Woolley, 1990). The different sites

reported for the burrows of the two forms suggest a difference

214

P.A. Woolley

in their habitat requirements. Although I was aware of differ-

ences between animals from different localities, I followed

Ride (1970) and referred to them as D. cristicauda.

Now that two forms of Dasycercus are again recognised, the

discrepancy between the names applied by earlier authors

(Iredale and Troughton, 1934; Tate, 1947) and those by Adams

et al. (2000) warrants attention. In the present study, the type

specimens of Chaetocercus cristicauda, Phascogale blythi and

Phascogale hillieri have been examined and early literature

reviewed. This was done to gain an understanding of the mor-

phology of the three previously named forms and the reasons

for them being considered to represent only two species. The

reasons given for the use of the names hillieri and cristicauda

for the two genetically distinct forms are then discussed.

Because Spencer (1896) questioned the accuracy of Krefft’s

description of Chaetocercus cristicauda, the specimens studied

by Spencer were examined to gain an understanding of his

reasons. Specimens collected by me in south-western

Queensland and the Northern Territory and thought to represent

two species were compared with the types to establish their

identities. Specimens in Museum Victoria (NMV), the

Australian Museum (AM), the South Australian Museum

(SAM) and the Western Australian Museum (WAM) have also

been examined and they provide information on the distribution

of the two species.

The type specimens

Chaetocercus cristicauda Krefft, 1867. The holotype (AM

Ml 1342) is a mounted skin and skull (cranium and dentaries)

of indeterminate sex. Krefft (1867) gave the following dimen-

sions: total length 8 inches (203 mm), tail 374 inches (82.5

mm), tarsi and toes 17s inches (28.5 mm).

Krefft founded the genus and species “upon a very singular

animal, approaching in its dentition Dasyurus proper much

more closely that any other known genus” [of the Dasyuridae],

in reference to the reduction in the number of premolar teeth. In

Dasyurus there are two upper and two lower premolar teeth.

Key features of Krefft’s description are: (i) the form of the tail,

described as thick and compressed [i.e., having the two oppo-

site sides nearly plane or flat] with a crest of black hair upon

the apical half [my emphasis]; and (ii) the presence of three

premolars in the upper jaw, the third being “diminutive and

tubercular”, and only two in the lower jaw. The general colour

of the fur was described as leaden grey at the base, tipped with

sandy and rufous, darker towards the tip. A colour plate accom-

panied the description and is reproduced here (Fig. la). The

specimen was said by Krefft to be in not very good condition

when it was received by the Museum, where it was mounted.

Examination of the holotype revealed that the mounted

specimen bears little resemblance to the animal illustrated but

part of the diagnostic crest of black hairs upon the apical half

of the tail can be seen, and the hairs, at least on the head and

back where the pelage has not been patched, are of the colour

described, but are very much less rufous than depicted by the

artist who prepared the colour plate. The compressed nature of

the tail is not apparent because it appears to have been over-

stuffed during preparation, and the left foot appears to be a

mismatched addition. The specimen was originally illustrated

with the left foot partly obscured, perhaps because it was miss-

ing. All teeth (1 4 /a, C 7i, PM di, M 4 A) have erupted in the holo-

type but some upper and all lower incisors have been lost. The

right and left third upper premolars are present, and are small

and tubercular as described by Krefft. Although it is unlabelled,

I have no reason to doubt the association of the skull with the

mount. It complies with Krefft’s description, and in later stud-

ies by Iredale and Troughton (1934), Tate (1947) and Jones

(1949), no doubts have been raised.

The type locality “South Australia, probably the neighbour-

hood of Lake Alexandria” was considered doubtful by Jones

(1949), “since Dasycercus is typically an inhabitant of the arid

and semi-arid Centre and has since been obtained only from

such environment, no specimen having been subsequently

recorded from localities within 200 miles of the Coorong

Lakes”. Calaby (1996) expressed the same opinion, noting that

Lake Alexandrina is “ecologically a most unlikely locality and

almost certainly an erroneous one”. Perhaps, in the absence of

precise information from F.G. Waterhouse, who sent the speci-

men to him, Krefft may have seized on the comment “I hope

soon to be able to send you some good skins of animals and

some skeletons as my man is going to settle down near Lake

Alexandrina . . .”, made by Waterhouse in a letter dated 21 June

1895 (Archives of the Australian Museum C. 20.65. 21). In an

earlier letter, 14 May 1865 (C. 20.65. 19), Waterhouse expressed

disappointment that he had been unable to proceed, because of

drought, on a trip from Port Augusta to Coffin Springs where

he had hoped to trap some small mammals is suggestive of a

more likely source area. Waterhouse said “I am convinced that

the small marsupials and rodents of the northern part of South

Australia are but little known - I know of several that are not

described in Gould’s work and I intend to do my best to procure

them”. Letters written by Waterhouse (held in the State

Archives of South Australia) and by Krefft (held in the Mitchell

Library, Sydney), even after the date of publication of the

description, might yield information on where the specimen

was collected.

Jones (1923) provided a very detailed description of the

external characters of specimens of Dasycercus from Ooldea,

South Australia collected by A.G. Bolam, referring to them as

Krefft’s Pouched Mouse, Chaetocercus cristicauda. Jones

added some details to the description of the form of the tail. He

noted that “The terminal half of the tail is ornamented with a

large dorsal and a small ventral crest of shining black hairs. The

ventral crest is inconspicuous, and consists of short stiff hairs

which do not increase in length as they are followed towards

the tip of the tail. The dorsal crest consists of long hairs which,

starting as a mere ridge of fine black hairs, increases in length

towards the tip of the tail and constitutes a fine fin-like crest.”

The ‘inconspicuous’ ventral crest may not have been obvious to

Krefft (1867) because of the poor condition of the specimen

upon which his description was based, or he may have over-

looked it, as commented on by Spencer (1896: 22). Illustrations

of the skull by Jones (1923, 1949) show three upper, and two

lower, premolar teeth in each jaw. Jones also noted that the

central area of the pads of the pes were striated and those of the

manus, unstriated.

The species of mulgaras

215

Figure 1. a, Chaetocercus cristicauda (= Dasycercus cristicauda) from Krefft (1866). b, tail of Phascogale hillieri Thomas, 1905

(= D . cristicauda ) (photograph of type specimen © The Natural History Museum, London), c, Phascologale cristicauda (-D.

blythi ) from Spencer (1896).

Figure 2. Tails of Dasycercus species, a, D. cristicauda (NMV C5385). b, D. blythi (NMV C5340).

216

P.A. Woolley

Phascogale blythi Waite, 1904. The syntypes, held in the

Western Australian Museum, are mounted skins of a male

M578 (renumbered M41476) and a female M579 (renumbered

M41477), together with the skull (cranium only) of one bearing

the label ‘Type Phascologale blighi (Waite)’, the nomen nudum

introduced by Woodward. The dimensions of the specimens

recorded by Waite (1904) were, for the male (adult); head and

body 150 mm, tail 102 mm and hind foot 27.8 mm, and for the

female (juvenile); head and body 132 mm, tail 95 mm and hind

foot 26.5 mm. They were collected in the Pilbara region of

Western Australia. Waite described the tail as: “. . . of moderate

length, shorter than the head and body, incrassated; the prox-

imal two-fifths above covered with short stiff yellow hairs, the

remainder with gradually lengthening black hairs that do not

however form a crest [my emphasis]. The whole of the lower

surface is black, with the exception of a small proximal portion

which is yellow.” The colour of the pelage was given as “upper

parts sandy, speckled with brown, the basal portion of the fur

being dark grey and the whole of the under parts, together with

the inner side of the limbs and the lining of the pouch is pure

white”. The footpads of both the fore and hind feet were said to

be striated. He recorded various dimensions of the skulls of the

two specimens, giving the ‘basal length’ of the skull of the male

as 35.0 mm, and that of the female as 33.0 mm. Based on my

measurement of the basicranial length (32.5 mm), the single

skull now in the collection is probably that of the female.

Details of the dentition are given. There are only two premolar

teeth in the upper jaw, the first smaller than the second and, if

as it appears that a superscript rather than a subscript has been

used in error, two in the lower jaw. The lower premolars touch

each other, and the first is in contact with the canine and the

second with the first molar tooth. Jones (1949), without making

specific reference to the apparent error, also interpreted this

part of the description as a reference to the lower premolars.

Jones (1949) commented that Waite became acquainted with

Dasycercus cristicauda during the time he was in charge of the

Adelaide Museum (1914-1928), but never suggested that it

was in any way akin to the Western Australian form that he had

described in 1904.

Phascogale hillieri Thomas, 1905. The holotype is an adult

male, skin without right hind foot and part of leg (Natural

History Museum, London, no. 5.3.28.1). It was collected

by H. Hillier at Killalpaninna, east of Lake Eyre, South

Australia. Dimensions of the type (measured in the flesh): head

and body 150 mm, tail 100 mm and hind foot 30 mm. The skull

is lost.

Thomas (1905) considered the specimen to be “near Ph.

cristicauda, but considerably paler in colour” [presumably by

comparison with the colour plate accompanying the description

by Krefft (1867)]. He described the tail as “slightly incrassated

basally; crested above for its terminal two inches [my empha-

sis], the hairs attaining a length of 16 mm, nothing that can be

called a crest below although some of the hairs are slightly

longer than in the proximal part. In colour the main part of the

tail is rather more rufous than the general body colour, though

nothing like the corresponding part in cristicauda ; under

surface indistinctly darker, not black; crest glossy black”.

Photographs of the holotype show the tail to be as described

by Thomas (lateral aspect, Fig. lb), and very like the tail of D.

cristicauda described and illustrated by Krefft (1867) and Jones

(1923). Dorsal fur colour of D. hillieri is similar to that of the

type specimen of Chaetocercus cristicauda Krefft and thus

only paler in reference to the colour plate.

On the morphological evidence, the type of P. hillieri cannot

be distinguished from that of C. cristicauda. Iredale and

Troughton (1934) therefore seem to have been fully justified in

placing P. hillieri in the synonomy of D. cristicauda, a decision

with which Jones (1949) agreed, having pointed out that in the

intensity of coloration “ cristicauda is a very variable animal,

and that specimens that are considerably lighter in colour than

usual are often captured with the more normal coloured

individuals”.

Conclusion. From the foregoing it is clear, based on morpho-

logical features, that there are two species of Dasycercus,

namely D. blythi and D. cristicauda, as recognised by Iredale

and Troughton (1934). The two are distinguishable by the form

of the tail (not crested in blythi, crested in cristicauda ), by the

dentition (PM 2 h in blythi, PM 3 A> in cristicauda ) and to a

lesser extent by their general body coloration {blythi a little less

rufous than cristicauda ). By association of their other morpho-

logical characteristics with those of the type specimens, for

which nipple number was not available, the females of the two

species can be distinguished by nipple number (six in blythi,

eight in cristicauda ).

On the names of the genetically distinct forms

Adams et al. (2000) presented evidence for the existence of two

clades using DNA sequence data for a small part of the mito-

chondrial gene cytb of 33 specimens of Dasycercus in museum

collections. They were able to obtain DNA for sequencing from

the type specimens of Phascogale hillieri and Phascogale

blythi, but not from Chaetocercus cristicauda, and to establish

that one clade includes the holotype of hillieri and the other, the

syntypes of blythi. They presented a subset of the gene

sequences of 171 base pairs from four specimens of each clade

to illustrate the genetic differences between them.

Various morphological attributes of the 23 genetically typed

individuals in the clade containing the type specimen of hillieri

and the 10 in the clade containing the syntypes of blythi were

then examined. They found that the two clades could be distin-

guished morphologically, the most diagnostic features being

“cresting on tail”, distribution of hair on the pes and nipple

number. Specimens in the clade containing the holotype of

hillieri had uniformly long black hairs on the dorsal surface of

the tail [which, by reference to illustrations in their report,

means crested]; hairs from the outer side of the foot covering

33% of the sole, and seven or eight nipples. Those in the clade

containing the syntypes of blythi, on the other hand, had black

hairs covering most of the tail with the hairs increasing in

length on the distal third [which, by reference to illustrations

in their report, means non-crested], hairs from both sides of the

feet covering 75% of the sole and six nipples.

Then, without any specific reference to the characteristic

The species of mulgaras

217

features of the species given in the original descriptions, Adams

et al. attempted to associate the type specimens with the speci-

mens in each of their two clades. They stated, in reference to

the clade containing the holotype of hillieri, “The cabinet skin

of the holotype, Phascogale hillieri, appeared similar to the

bodies (cabinet skins and spirit bodies) of animals referred to

Dasycercus hillieri in Australian Museum collections. This

diagnosis was fully supported by its DNA profile.” Further, in

reference to the clade containing the syntypes of blythi , “The

cabinet skin of the holotype” [of Dasycercus cristicauda ] “was

referable to the bodies of animals assigned to Dasycercus cristi-

cauda in this study.” and “The cabinet skins of the holotypes,

Phascogale blighil blythi, are similar to the bodies of animals

referred to Dasycercus cristicauda, a result consistent with the

DNA profiles for both specimens.”

The association of the name hillieri with one clade is

supported by genetic evidence, “cresting on tail” and by the dis-

tribution of hair on the feet. Nipple number is not useful

because it cannot be established in the holotype. However, the

association of the name cristicauda with the other clade

containing the syntypes of blythi has no support from either

genetic or morphological evidence. Despite repeated attempts,

DNA could not be extracted from the holotype of cristicauda

and the tail of cristicauda is crested whereas that of blythi is

non-crested. Distribution of hair on the pes and nipple number

cannot be used in support of the association because these

attributes cannot be assessed in the mounted type specimens of

cristicauda and blythi. It is my conclusion that Adams et al.

(2000) misplaced cristicauda in the clade containing the

syntypes of blythi. The number of premolar teeth, a character

not considered by them, differs and the tail of cristicauda has a

crest like that of hillieri, which allies cristicauda with the clade

containing hillieri. The correct name for specimens in this clade

is therefore D. cristicauda (with which hillieri has been syn-

onymised in the past) and, for the specimens in the other clade,

D. blythi.

Spencer collection

W. Baldwin Spencer, Professor of Biology at Melbourne

University, was the zoologist on the Horn Expedition to Central

Australia, May- August 1884. No Dasycercus were obtained in

the course of the expedition but soon after Spencer started

receiving specimens from friends. These post-Horn Expedition

‘collectors’ included P.M. Byrne of Charlotte Waters, C.E.

Cowle of Illamurta and FJ. Gillen of Alice Springs. The real

collectors were the Aboriginal people and precise localities

where the specimens were found were not recorded. The exact

number sent to Spencer is not known (Calaby, 1996) but 28

specimens (not including pouch young) of Dasycercus acquired

between November 1894 and September 1895 can be found in

the Spencer collection of Museum Victoria. Nineteen of the 28

were from Charlotte Waters, two from Crown Point and seven

from Illamurta, all in southern Northern Territory.

Spencer (1896) identified them as Phascologale cristicauda

Krefft, the Crest-tailed Phascologale, based on the “peculiarity

of the dentition [a reference to the reduction in the number of

premolar teeth], the crested tail and the general measurements

of the body.” However, variability in form of the tail, dentition,

nipple number and colour among he specimens he studied led

him to give an amended description of the species (Spencer,

1896: 21-23).

Most of Spencer’s specimens are intact (in spirit). The tail

was examined and classed as either C (to denote the crested

form of the tail of D. cristicauda ) or B (the non-crested form of

the tail of D. blythi ). The dentition was examined in prepared

skulls and intact specimens with particular reference to the

upper third premolar tooth position (P = P 3 present, NP = P 3 not

present, dP 3 = deciduous tooth in place). Tooth nomenclature

follows Luckett (1993). Body measurements (total length, tail

length and foot length in mm) of spirit specimens were taken,

if not previously recorded. The reproductive status of individu-

als was assessed by examination of the pouch area and repro-

ductive organs of both females and males. Females were

assessed as mature if they were lactating or showed signs of

having reared young previously (nipples elongated, pouch fur

stained), or, in one case, the appearance of the uteri indicating

that the individual had very recently given birth. Nipple num-

ber was determined in most immature and mature females.

Males were assessed as mature if the scrotum was large and the

prostate gland developed and showing zonation. The size of the

scrotum alone is not a good indicator of maturity because

development of the prostate occurs after the testes have reached

full size.

Two species of Dasycercus can be recognised (Table 1), five

specimens (four from Charlotte Waters and one from Crown

Point) of D. cristicauda (crested tail, P 3 present) and 23 of D.

blythi (non-crested tail, P 3 not present). This agrees with the

Spencer’s (1896) observation that P 3 was absent in most of his

specimens. Females identified as D. cristicauda have eight

nipples and D. blythi have six. None was found to have four

nipples as reported by Spencer; he may have overlooked very

small, unsuckled nipples in the pouch of lactating females. In

this sample, specimens of D. cristicauda are larger than

D. blythi-, two mature males of D. cristicauda having head-

body lengths of 180 and 185 mm and foot length of 33.5 mm,

and two D. blythi with a head-body length of 135 mm and foot

lengths of 26.5 and 27.5 mm. In both species tail length is

less than head-body length. I was unable to distinguish between

the pads of the manus of the two species (said to be unstriated

in D. cristicauda and striated in D. blythi ) but in this long

preserved material it was difficult to see striations on any

pads.

Spencer (1896) failed to recognise that the variability he

saw could be accounted for by two species. The specimen that

he illustrated, as Phascologale cristicauda (Fig. lc), has the tail

of D. blythi. A skull of a specimen from Charlotte Waters

(C.6158, not included in Table 1), is labelled in Spencer’s hand

“Fig. spec.” It has the characteristics of D. blythi (ie P 3 absent,

diastema between P 2 and M 1 ) and may be the missing skull of

C.5388 (see Table 1). If so, specimen C.5388 would be the

specimen illustrated. The tails of the two species are illustrated

in Fig. 2 and dentition in Fig. 3.

In a further 120 specimens, identified as either D. blythi

or D. cristicauda on the form of the tail, P 3 was absent

from all specimens of D. blythi except one (SAM M3 115).

218

P.A. Woolley

Table 1 . Data for 28 specimens of Dasycercus from Charlotte Waters, Crown Point and Illamurta held in the Spencer collection, Museum Victoria.

See text for explanation of assessment of reproductive status, form of the tail (B = D. blythi and C - D. cristicauda) and P 3 .

Reg. no. C

Accession

date

Specimen

Sex

Reproductive

status

Form

of tail

No. of

nipples

P 3

H-B

Tail

Foot

Charlotte Waters (25°55’S, 134°56’E)

234

23.3.1916

spirit, skull

mature

24.5

5336

-.9.1895

spirit

mature

130

24.5

5337

-.12.1894

spirit

mature

125

25.5

5338

-.9.1895

spirit

mature

130

25.0

5339

-.9.1895

spirit

mature

125

24.0

5340

-.9.1895

spirit

mature

135

100

26.5

5359

-.2.1895

spirit

mature

130

23.5

5360

-.2.1895

spirit

weaner

dp 3 *

19.5

5361

-.2.1895

spirit

immature

125

25.5

5369

-.2.1895

spirit

mature

125

24.5

5370

-.2.1895

spirit

mature

130

25.0

5371

-.12.1894

spirit

mature

120

23.5

5372

-.2.1895

spirit

immature

115

25.5

5384

18.12.1895

spirit

mature

130

26.0

5385

18.12.1895

spirit

mature

180

33.5

5387

1895

spirit

weaner

dp 3 *

19.5

5388

1895

spirit (-head)

mature

26.0

5391

1.7.1895

spirit

immature

155

100

30.5

5392

-.7.1895

spirit

mature

185

120

33.5

Crown Point (25°30’S, 134°23’E)

5357

-.8.1895

spirit

mature

150

100

30.5

5358

-.8.1895

spirit

mature

135

27.5

Illamurta (24°18’S, 132°41’E)

4828

-.11.1895

spirit

immature

dp 3 (L)

23.0

4829

-.11.1895

spirit

immature

dp 3 (R)

105

25.0

4830

-.11.1895

spirit

immature

105

24.5

4831

-.11.1895

spirit

immature

dp 3 (L)

100

23.0

5362

4.10.1895

spirit (-gut)

150

27.0

5380

16.11.1894

spirit

immature

dp 3

21.5

5381

16.11.1894

spirit (-tail)

immature

21.5

*not erupted - gum previously cut to expose the tooth

A characteristic feature of D. blythi is the presence of a

diastema between P 2 and M 1 in mature specimens. P3 was pres-

ent in both mature and immature specimens of D. cristicauda

with the exception of two mature specimens (SAM M3 106,

M3 154). These three aberrant individuals formed part of a col-

lection from the Canning Stock Route (Table 2), which raises

the possibility that the skins and skulls of two (M3 106 and

M3 115, both with adult dentition) were mismatched, but pro-

vides no obvious explanation for the absence of P 3 in the mature

specimen (M3 154). However, the teeth of this specimen were

very worn, and the rough appearance of the bone between P 2

and M 1 suggested that P 3 might have been lost. In specimens of

D. blythi the deciduous third premolar was found unerupted

(Table 1) in ‘weaners’, young close to independence in which

all upper incisor teeth except the first pair had erupted. The

eruption of I 1 later than the other incisor teeth is thought to be

an adaptation for suckling (Luckett and Woolley, 1996).

Deciduous premolar teeth were seen in some immature

individuals of D. blythi (Table 1), and D. cristicauda (e.g.,

SAM Ml 9672, M20913 and M20925) and the right dP 3 was

seen in a mature D. blythi (SAM M3 146).

Woolley specimens

Specimens collected on Sandringham Station, south-western

Queensland (Woolley, 1990) have features characteristic of

D. cristicauda (Western Australian Museum, M8207, M9670-

73, M13716; Queensland Museum, JM5532). Specimens

collected in the vicinity of Papunya, near Uluru, on the old

Docker River Road west of Uluru, and near Refrigerator Well,

Tanami Desert, all Northern Territory (see Woolley, 1990 for

details of localities) have features characteristic of D. blythi

(Museum Victoria, Uluru, C35885, C35886; Papunya,

C35887-C35895; Tanami Desert, C35896, C35897;

Queensland Museum, Docker River Road, JM1443; Tanami

Desert, JM2311, JM2312, JM2317.

The species of mulgaras

219

Figure 3. Lateral view of the skulls of Dasycercus species, a, D. cristicauda female (NMV C676), showing P 3 (arrowed), b, D.

blythi female (NMV C234), showing the diastema (arrowed) between P 2 and M 1 .

Distribution

Dasycercus has a wide distribution across the arid zone of

Australia (see maps in Woolley, 1995 and Australian Mammal

Map Updates at http:/www.naturebase.net/science/mupdates.

html). The past and present distribution of each species requires

clarification to assess their conservation status. Of the many

museum specimens available, those from two areas at the

species’ north-western and south-western limits have been

examined, in addition to those above from central Australia in

the Spencer collection.

Canning Stock Route. Otto Lipfert, taxidermist of the Western

Australian Museum, accompanied the Canning Stock Route

Expedition of 1930-1931 through the north of Western

Australia. Twenty-four specimens collected by him and

lodged in the South Australian Museum have been examined

(Table 2). Based on the form of the tail, 13 were identified as

D. cristicauda and 11 as D. blythi. The specimens were col-

lected along the Stock Route, from Well 27 to Well 49, and both

species were found in the vicinity of Wells 27, 35 and 45.

Lipfert reported that he collected specimens up to 2-3 miles

from well sites (McKenzie and Youngson, 1983). An indication

that the two species may occupy different habitats comes from

the collector’s notes on two specimens: D. cristicauda M3 107

- “top of sandhill” and D. blythi M3 144 - “spinifex flat”. The

annotated map of the Stock Route accompanying the article on

Mr Canning’s Expeditions in Western Australia 1906-1907 and

1908-1910 (Anon, 1911) shows the positions of the wells and

220

P.A. Woolley

Table 2. Data for 24 specimens of Dasycercus from the Canning Stock Route held in the South Australian Museum. Explanation as for Table 1.

* aberrant specimens referred to in text

Well no. (coordinates)

Reg.

no. M

Date

Specimen

Sex

Reproductive

status

Form

of tail

No. of

nipples

P 3

27 (22°48'S, 123°39'E)

3140

-.7.1930

spirit, skull

immature

3141

-.7.1930

skin, skull

28 (22°39'S, 123°45'E)

3142

-.8.1930

spirit

mature

29 (22°34'S, 125°53'E)

3143

-.8.1930

skin, skull

„

3144

-.8.1930

skin, skull

„

3154*

-.8.1930

spirit, skull

c 5

mature

33 (22°20'S, 124°44'E)

3145

-.10.1930

skin, skull

34 (22°16'S, 124°54'E)

3146

-.10.1930

spirit, skull

mature

dp 3 R

35 (22°13'S, 125°03'E)

3105

29.10.1930

skin, skull

„

3147

-.10.1930

skin, skull

„

3148

-.10.1930

skin

36 (22°08'S, 125°17'E)

3106*

5.11.1930

skin, skull

3149

-.11.1930

skin, skull

41 (21°33'S, 125°51’E)

3107

4.6.1931

skin, skull

3150

-.6.1931

skin, skull

—

42 (21°19'S, 125°53E)

3108

24.5.1931

skin, skull

43 (21°12'S, 125°54'E)

3109

17.5.1931

skin, skull

P(L)

44 (20°02'S, 126°04'E)

3110

2.5.1931

skin, skull

45 (20°48'S, 126°11’E)

3111

6.5.1931

skin, skull

3112

18.4.1931

skin, skull

„

3113

19.4.1931

skin, skull

3114

18.4.1931

skin, skull

47 (20°26'S, 126°18'E)

3115*

1.4.1931

skin, skull

49 (20°10'S, 126°41’E)

3117

2.3.1931

skin, skull

Table 3. Data for 13 specimens from Ooldea, Fisher and Rawlinna held in the Australian Museum. Explanation as for Table 1. * Too young for

P 3 to have erupted. (I 1 not erupted in M4923; partially erupted in others).

Locality

Reg.

no. M

Date

Specimen

Sex

Reproductive

status

Form

of tail

No. of

nipples

P 3

Ooldea (30°27’S, 131°50’E)

2987

12.8.1921

skin, skull

„

2988

12.8.1921

spirit, skull

mature

3025

29.8.1921

spirit, skull

mature

4923

23.10.1921

spirit

immature

Fisher (30°33’S, 130°58’E)

4862

22.10.1921

skin, skull

„

4863

23.10.1921

skin, skull

mature

4864

23.10.1921

skin

mature

4865

23.10.1921

skin, skull

immature

4866

24.10.1921

skin, skull

immature

„

4924

24.10.1921

spirit

immature

„

31546

20.10.1921

spirit

mature

Rawlinna (31°01’S, 125°20’E)

4355

30.7.1928

spirit, skull

mature

”

4356

30.7.1928

skin, skull

gives details of the country traversed, which includes a mixture

of spinifex flats and sand-ridge country.

Ooldea, Fisher, Rawlinna. These localities are near the

south-western limit of the distribution of Dasycercus. It was

from Ooldea that Jones (1923) obtained his specimens of

D. cristicauda but his material has not been located. Thirteen

specimens (four from Ooldea, seven from Fisher and two from

Rawlinna) held in the Australian Museum have been identified

as D. cristicauda based on the crested tail, presence of P 3 in

mature specimens and eight nipples in females (Table 3).

Remarks by Bolam (1925) suggest that D. blythi may also

occur in the area. He wrote (p. 25) “This animal [the Crested-

tailed phascogale], which bears the scientific name of

Dasycercus cristicauda, is known to the blacks as Mul-gurra.

Two species of the Phascogale family are found on the

Nullarbor Plain and the sandhills of Ooldea. These are the

Crested- tailed and the Brush- tailed; but with the exception of a

small difference in the hair on the tail the species are alike in

appearance and habits ” [my emphasis].

The presence of the two species, D. cristicauda and

D. blythi, in central Australia (the Spencer collection) and

The species of mulgaras

221

north-western Australia (the Canning Stock Route collection)

together with the possibility that both may be found in the

south-western limit of the range (Ooldea/ Nullarbor Plain),

suggests that the two could occur throughout much of the

range.

Common names

Mulgara, the name by which Dasycercus cristicauda was

known to the Aboriginal people at Ooldea, has been used as its

common name. Spencer (1896) knew that the Aborigines at

Charlotte Waters and Illamurta used the name Amperta but it is

impossible to know to which of the two species this might have

applied. Many Aboriginal names for Dasycercus have been

recorded by Finlayson (1961) and Burbidge et al. (1988), one

of which, Ampurta, has been used as the common name for

D. hillieri, here no longer recognised as distinct from D. cristi-

cauda. With his recognition of two species, Troughton (1965)

referred to them as the Crest-tailed Marsupial Mouse (for

D. cristicauda ) and the Western Crest- tail (for D. blythi). The

latter common name is inappropriate because D. blythi is not

restricted to the western part of the range, and it does not have

a crested tail. Because Mulgara was a familiar common name

for Dasycercus when it was thought by some to be a monotyp-

ic genus I propose that it be retained; with D. blythi to be

known as the Brush-tailed Mulgara and D. cristicauda as the

Crest-tailed Mulgara. Both species have black hairs on the dis-

tal half of the tail but the dorsal, fin-like crest of cristicauda

provides a simple means of differentiating between the two

species in the field.

Acknowledgments

I am grateful to the following museum personnel for assistance:

Wayne Longmore, Rory O’Brien, John Augier, Sally Rogers-

Davidson and Sandra Winchester, Museum Victoria; Sandy

Ingleby, Tish Ennis and Jan Brazier, Australian Museum; Cath

Kemper, David Stemmer and Fran Zilio, South Australian

Museum; Norah Cooper and Claire Stephenson, Western

Australian Museum, and Paula Jenkins and Louise Tomsett,

Natural History Museum, London. I thank Mark Adams for

access to the unpublished report on Dasycercus systematics,

and Michael Westerman for helpful discussions while the man-

uscript was in preparation.

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