Memoirs of Museum Victoria 80: 1-41 (2021) Published March 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/).mmv.2021.80.01

Indo-West Pacific and Australian species of Eucalliacidae with descriptions of four

new species (Crustacea: Axiidea)

(http://zoobank.org/urn:lsid:zoobank.org:pub:EA21667A-77A5-A11 D-9C1A-23ECFFF3D505)

Gary C. B. POORE (http:;//zoobank.org/urn:lsid:zoobank.org:author:c004d784-e842-42b3-bfd3-3174d359f8975)

(https://orcid.org/0000-0002-7414-183X )

Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia. Email: gpoore(?museum.vic.gov.au

Poore, G.C.B. (2021). Indo-West Pacific and Australian species of Eucalliacidae with descriptions of four new species

surveys of coral reefs and sandy shallow environments in the Indo-West Pacific and Australia have discovered 14

species of Eucalliacidae, of which four are new. All species are diagnosed, and Andamancalliax arafura sp. nov.,

Eucalliaxiopsis dworschaki sp. nov., Eucalliaxiopsis paradoxa sp. nov. and Eucalliaxiopsis patio sp. nov. are described as

new. The collection is an opportunity to re-diagnose other species and provide colour photographs of some. Calliaxina

xishaensis Liu and Liang, 2016, is synonymised with Calliaxina novaebritanniae (Borradaile, 1900).

Abstract

(Crustacea: Axiidea). Memoirs of Museum Victoria 80: 1—41.

Keywords Crustacea, Axiidea Eucalliacidae, taxonomy, new species

Introduction

Of eight families of callianassid-like Axiidea, Eucalliacidae

have the most complicated taxonomic history that was most

recently reviewed by Poore et al. (2019). Their revision and the

molecular phylogeny on which it was based (Robles et al. 2020)

depended in part on extensive collections made in the Indo-

West Pacific over the last couple of decades. These collections

also made it possible to re-diagnose known species and

describe others revealed as new during the molecular study.

Eucalliacidae comprise seven genera, diagnosed by Poore et

al. (2019), and 21 species listed by these authors. Six species are

from the Atlantic or Mediterranean and two are deep-sea. This

contribution deals only with those from coral reefs and shallow

environments in the Indo-West Pacific and Australia. Fourteen

species are covered here, of which four are described as new.

Methods

Much of the material comes from the Muséum nationale

d'Histoire naturelle, Paris (MNHN), including expeditions to

Papua New Guinea, Madang Province (PAPUA NIUGINI

stations), and New Ireland Province (KAVIENG 2014

stations). Another sizable Indo-West Pacific collection has

been accumulated by the Florida Museum of Natural History,

University of Florida (UF). Specimens were also examined

from Museums Victoria, Melbourne (NMV); the Australian

Museum, Sydney (AM); Western Australian Museum, Perth

(WAM); and Naturhistorisches Museum, Vienna (NHMW).

Unless otherwise stated, station prefixes and numbers belong

to systems initiated by the museum holding the material.

Size 1S expressed as carapace length (cl.), including

rostrum, in mm. Individuals marked with an asterisk (*) were

sequenced and contributed to molecular analysis (Robles et

al., 2020). Diagnoses have been prepared for all species and

coded into a DELTA database (Dallwitz, 2010). This database

was used to generate the diagnoses presented here; character

states in italics diagnose each species in at least one respect

from every other species. Colour photographs of fresh

specimens were taken in the laboratory shortly after collection

by Arthur Anker and Zdenek Duri&. Photographs 1n figs 12

and 25 were taken using a Leica 205C microscope and the

Leica Application Suite multifocus routine.

Eucalliacidae Manning and Felder, 1991

Eucalliinae Manning and Felder, 1991: 781 (misspelling).

Eucalliacinae Sakai, 1999: 108—109.— Sakai, 2018: 734—738 (partim).

Eucalliacidae.— Sakai, 2011: 491.— Sakai and Türkay, 2014: 190

(outdated key to genera).— Sakai, 2018: 734—738.— Poore et al., 2019:

122 (for complete synonymy).

Remarks. The family was diagnosed by Poore et al. (2019). It 1s

recognised most easily by the truncate setose apex of the

dactylus of maxilliped 3. Callianopsidae, which also share this

feature, differ in that the uropodal exopod lacks a dorsal plate,

indicated in eucalliacids by a secondary dense row of stout

setae diverging on the upper surface from the anterior margin.

Andamancalliax Sakai, 2011

Andamancalliax Sakai, 2011: 494—495.— Sakai, 2018: 738.—

Poore et al., 2019: 126.

Remarks. The genus differs from other eucalliacids in the

prominent sharp rostrum and the unequal dissimilar chelipeds, the

minor one having elongate fingers. The crista dentata of the type

species was not illustrated, but in the new species described here,

it comprises few sharp long spines in lieu of a row of similar even

teeth seen in other genera. Sakai (2011, 2018) stated that the

maxilliped 3 lacks an exopod but in the newly discovered female

of what appears to be a different species, a small exopod is present.

The genus was diagnosed by Sakai (2011) as having the male

pleopod 1 biarticulate, but this was changed by Sakai (2018) to

pleopod 1 absent. I believe itis biarticulate, as in other eucalliacids,

for the reasons given under remarks on the type species.

Andamancalliax andamanica (Sakai, 2002)

Calliax andamanica Sakai, 2002: 463—467, figs 1, 2.— Sakai,

2005: 201.

Andamancalliax andamanica Sakai, 2011: 495.— Sakai, 2018: 738.

Distribution. Thailand, Andaman Sea. Shelf, 31—61 m.

Remarks. Sakai (2002) referred to the holotype of Calliax

andamanica as female in the description and male in the figure

legends. His figure 2B of the "female Plp 1” from the holotype,

described as "uniramous, two-articled, distal article bilobed

distally”, is most probably from an immature male (cf. fig. 16q

of male pleopod | of Eucalliaxiopsis dworschaki sp. nov.).

Andamancalliax arafura sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: | E5B7D96-E101-

45B8-A D22-7156C8B29B30

Figures 2, 3

Material examined. Holotype. Australia, NT, Arafura Sea, 9.2055° S,

133.6483° E, 155-158 m, oozy mud (RV Southern Surveyor stn

032BS006), AM P.74526 (female, 3.7 mm, without major cheliped).

Description of holotype. Carapace smooth; gastric-abdominal

regions together 4.3 times as long as width of these regions

together; branchiostegite fully calcified; cervical groove at 0.62

carapace length, scarcely obvious on branchiostegite.

Rostrum acute, strongly tapering, two-thirds length of

eyestalk. Anterolateral carapace lobe absent. ‘Thoracic sternite

7 1.6 times as long as wide, with well-defined median sulcus

over posterior half, smooth over medial half, weak lateral ridge

anterior to coxal articulation crossing pit at quarter of width.

Pleonite | without anterior transverse groove, without

sternal plates.

Eyestalk 1.5 times as long as greatest width, swollen laterally,

tapering distally around cornea, with defined apical lobe; cornea

wider than long. Antennular peduncle 1.8 times as long as width

of both eyestalks; article 2 twice as long as wide; article 3 as long

as article 2; article 3 with ventrolateral row of few setae,

continued onto flagellum. Antennal peduncle 2.2 times as long as

width of both eyestalks, overreaching antennule peduncle by

most of article 5; scaphocerite ovoid, longer than wide; article 4

G.C.B. Poore

c. 5 times as long as wide; article 5 0.7 length of article 4.

Maxilliped 3 basis with 4 mesial teeth along lower margin,

ischium with strongly expanded proximo-lower lobe, tapering to

two-thirds of its greatest proximal width; crista dentata of 3 short

basal spines, plus 4 long sharp spines, first 2 recurved, last 2

overlapping merus; merus upper margin 1.5 times as long as

ereatest width; carpus-dactylus together reaching back to mid-

ischium; dactylus as wide as long, with setose truncate apex;

exopod third length of upper margin of ischium.

Major cheliped missing. Minor cheliped (right) ischium c.

2.8 times as long as distal width, lower margin with sharp

subdistal spine; merus twice as long as broad, lower margin

convex, unarmed; carpus 1.1 times as wide as upper length,

margins carinate; propodus palm tapering, 0.9 times as wide

as upper palm length, upper margin rounded, distomesial

margin of palm oblique, distolateral margin of palm oblique,

with triangular tooth at base of finger; fixed finger twice

length of upper margin of palm, lateral cutting edge with

proximal sharp oblique tooth, obtuse distal tooth; dactylus as

long as fixed finger, 5 times as long as wide at base, cutting

edge with obtuse rounded tooth at about two-thirds.

Pereopod 2 merus 2.2 times as long as maximum width;

dactylus c. 5 times as long as upper margin of propodus.

Pereopod 3 merus 2.3 times as long as maximum width;

carpus 1.7 times as long as wide; propodus with lower margin

concave, 1.3 times as long as mid-length; dactylus less than

half as long as axial length of propodus. Pereopod 4 merus 3

times as long as maximum width; carpus 3 times as long as

wide; propodus setose, with long distal spiniform seta among

setose margin, as long as dactylus, with 2 long distal setae

overlapping dactylus. Pereopod 5 chelate.

Pleopod | uniarticulate, tapering. Pleopod 2 endopod 2.6

times as long as wide; appendix interna at midpoint of endopod,

reaching two-thirds remaining length of endopod. Pleopod 3

with appendix interna embedded in endopod.

Uropodal endopod ovoid, anterior margin more convex

than posterior margin, widest at midpoint, 1.7 times as long as

wide, without facial setae; exopod ovate, greatest dimension 1.6

times anterior margin, anterior margin straight, curving into

distal margin, sharper curve between distal and posterior

margins, with row of blade-like setae on distal posterior

margin; dorsal plate defined by weak longitudinal ridge,

extending half of exopod width, distal plate margin with a

dozen scattered spiniform setae of various sizes. Telson 1.6

times as wide as long, broadest over anterior half, lateral margin

evenly curving to posterior margin; posterior margin convex,

with posterolateral clusters of long setae; dorsal surface smooth.

Etymology. For the Arafura Sea (noun in apposition).

Distribution. Australia, Arafura Sea. Shelf, 151—158 m.

Remarks. The truncate setose dactylus of maxilliped 3,

presence of a dorsal plate, spike-like rostrum and long fingers

on the minor cheliped place the new species in Andamancalliax.

The sole specimen differs from the illustrations of A.

andamanica in the presence of a short exopod on maxilliped 3,

a distal spine on the lower margin of the ischium of the minor

cheliped, a tooth on the fixed finger of the minor cheliped, a

Indo-West Pacific and Australian Eucalliacidae 3

Figure 1. Live colour photographs: a, Calliaxina bulimba, Papua New Guinea. MNHN-IU-2013-7097; b, c, C. kensleyi, Saudi Arabia, UF 36699;

d, e, C. novaebritanniae, Papua New Guinea, MNHN-IU-2013-7062; f, Eucalliaxiopsis dworschaki, French Polynesia, UF 16286; g, E. dworschaki,

Papua New Guinea, MNHN-IU-2013-7081; h, E. inaequimana, French Polynesia, UF 29162; 1, E. inaequimana, French Polynesia, UF 29208.

Photos a, d, e, g: Z. Duri&; b, c, f, h, i, A. Anker. Various scales.

more oval uropodal endopod and a less pronounced dorsal ^ surface carinate and the anterior half elevated.

plate on the uropodal exopod. The few sharp spines that

constitute the crista dentata are unique within Eucalliacidae; Calliaxina Ngoc-Ho, 2003

the situation in A. andamanica was not reported. The dorsal

plate of the uropodal exopod of A. arafura is less well developed Calliaxina Ngoc-Ho, 2005: 495—494.— Sakai, 2011: 497 (part).—

than in A. andamanica, which was said to have the dorsal Poore etal., 2019: 126—127.

| G.C.B. Poore

J j / i

WE IT -

—

——

K WX

Figure 2. Andamancalliax arafura sp. nov. Holotype, Australia, NT, AM P.74526 (female, 3.7 mm): a, b, anterior carapace, eyestalks, antennule,

antenna, pleonite 1, dorsal, lateral views; c, pleonite 6, telson, uropod (c', detail of uropod exopod spiniform setae); d, thoracic sternite 7; e, f,

minor cheliped (right), mesial, lateral views; g, pereopod 2; h, pereopod 3; 1, J, pereopod 4, with detail of propodus, dactylus; k, pereopod 5. Scale

bars = | mm.

Indo-West Pacific and Australian Eucalliacidae

Figure 3. Andamancalliax arafura sp. nov. Holotype, Australia, NT, AM P.74526 (female, 3.7 mm): a, b, left maxilliped 3, lateral, mesial views; c,

left maxilliped 3, basis, merus, posterior view; d, pleonal sternite 1, pleopod 1; e, f, pleopods 2, 3. Scale bar = 1 mm.

Remarks. Calliaxina differs from Eucalliaxiopsis in lacking a

transverse ridge on the telson, lacking differentiation between

male and female major chelipeds, and always possessing an

exopod on maxilliped 3, sometimes vestigial (Poore et al.,

2019; Robles et al., 2020).

Calliaxina includes the type species, C. punica (de Saint

Laurent and Manning, 1982) from the western Mediterranean

plus four species from the Indo-West Pacific. Separation of

the species is difficult because characters defining each have

historically been incompletely documented. Juveniles of all

species tend to have larger eyes, more acute rostrums and

similar telsons, making differentiation on the basis of these

characters impossible. Adult males of four species are known;

juvenile males of one of these and of the fifth species have

presumably immature pleopods 1 (see Poore et al. [2019] and

discussions below under C. bulimba and C. novaebritanniae).

Liu and Liang’s (2016) key to species included one species

here placed in Eucalliaxiopsis and relied on the shape of the

rostrum and telson, both size-dependent features.

Key to species of Calliaxina

l. Maxilliped 3 exopod vestigial 2

— Maxilliped 3 exopod reaching at least third way along

ischium 3

2. Major cheliped propodus distomesial margin with setae

aligned in 2 groups of c. 10 and c. 6 setae; minor cheliped

distomesial margin with setae aligned in 2 scarcely separate

eroups of c. 20 and c. 8 setae; cheliped merus lower margin

with few denticles; Red Sea, Jordan, Saudi Arabia;

Madagascar C. kensleyi (Dworschak, 2005)

Major and minor chelipeds propodi distomesial margins

with setae aligned in 2 groups of c. 9 setae each; cheliped

merus lower margin with 2 or 3 proximal truncated

curved teeth and diminishing denticles more distally;

Papua New Guinea, SE Australia... s sh hh

C. bulimba (Poore and Griffin, 1979)

Maxilliped 3 exopod overlapping merus (except in

Juveniles) 4

Maxilliped 3 exopod reaching half-way along ischium;

male pleopod | article 2 3 times as long as wide; Japan,

East China Sea C. sakaii (de Saint Laurent, 1979)

Antennule peduncle reaching base of antenna peduncle

article 5; male pleopod 1 with broad subapical notch,

prominent appendix interna, triangular apex; Mediterranean

C. punica (de Saint Laurent and Manning, 1982)

Antennule peduncle reaching midpoint of antenna peduncle

article 5; pleopod | article 2 with narrow apical notch, short

appendix interna, apex prominently curved; Papua New

Guinea, Indonesia, NE Australia, Philippines, South China

Sea Pen em Polynesia: ». eene endete nde tat

C. novaebritanniae (Borradaile, 1900)

Calliaxina bulimba (Poore and Griffin, 1979)

Figures la, 4, 5, 12a, h

Callianassa bulimba Poore and Griffin, 1979: 257, fig. 21.

Calliax bulimba.—de Saint Laurent and Manning, 1982: 222.—

Sakai, 1999: 119, fig. 32a—-c.— Sakai, 2005: 202.

Calliaxina bulimba.—Sakai, 2011: 499.— Poore et al., 2019: 127,

146, fig. 19g, h.

Eucalliax bulimba.— D worschak, 2005: fig. 5c, d.

Eucalliax kensleyi.— Sakai, 2017: 1122-1124 (partim), fig. 2D.

Material examined. Australia. Qld, Britomart Reef front, 18° 17' S,

146° 38' E (NMV stn AIMS 57), NMV 712184 (female, 8.7 mm). Rib

Reef, 18.48° S, 146.86" E, October 1985, shallow lagoon, 8 m, (M.

Riddle stn R/10/1), NMV J71676 (male, 4.6 mm).

Papua New Guinea. Madang Province, lagoon, PAPUA NIUGINI

stations: 05° 10.2' S, 145° 50.4' E, 1-3 m (stn PR243), MNHN-

[U-2013-7097* (male, 4.5 mm); 05? 10.3' S, 145° 48.5' E, 1-18 m (stn

PR213), MNHN-IU-2013-7124 (female, 3.5 mm).

Diagnosis. Sternite 7 with transverse groove visible only

laterally, anteromedial lobe rounded. Antennule peduncle

reaching to midpoint of antenna peduncle article 5. Maxilliped

3 exopod vestigial, about twice as long as wide. Major cheliped

merus, lower margin with 2 or 3 proximal truncate teeth;

propodus distomesial margin with setae aligned in 2 similar

groups of c. 9 setae each. Male pleopod 1 article I linear, with

distal setae; article 2 short, oblique (juvenile form).

Colour. Exoskeleton translucent white (fig. 1a).

Distribution. Indonesia, Ambon; Papua New Guinea, Madang

and New Ireland provinces; Australia, Qld (type locality).

Coral reef lagoon, 1—18 m.

Remarks. Poore and Griffin’s (1979) simple figures are

supplemented here by illustrations of a newly discovered male

and female from Australia. The telson 1s widest proximally,

the anterior margins each side of the rostrum are concave, and

the distomesial margins of the propodi of the chelipeds bear

two widely separate small groups of c. 9 setae each. As is

typical of callianassid-like families, the rostrum of small

individuals 1s more acute than 1n adults and the cornea more

swollen. Separation of the anterior and posterior sections of

thoracic sternite 7 is weak, with only a slight inclination

laterally terminating in a sublateral pit; the posterolateral

margin carries groups of c. 6, 7 and 9 long setae (fig. 12h). As

figured by Dworschak (2005), pleonite 1 lacks obvious ventral

plates but bears two short rows of three or four long setae

anterior to the pleopods.

The 4.5-mm male from Papua New Guinea has a pleopod |

with long distal setae on article 1 and a short simple oblique

article 2 (fig. 5d). This is consistent with Poore and Griffin’s

(1979: fig. 21g) and Dworschak's (2005: fig. 5c, d) illustrations

of the much larger holotype (cl. 10 mm) and resembles those of

Eucalliaxiopsis inaequimana (see Dworschak, 2014: fig. 4j-l),

E. madagassa (see Ngoc-Ho, 2014: fig. 2S, as C. thomassini)

and E. patio sp. nov. Pleopod 1 of the 4.6-mm male from

Australia differs in having a bilobed article 2 with a longitudinal

crease and a separate appendix interna (fig. 5b). As argued by

G.C.B. Poore

Poore et al. (2019), this would appear to be an intermediate

form tending towards the more typical eucalliacid male

pleopod (e.g., fig. 7h, J). Undeveloped male pleopods 1 are

also seen in small juveniles of C. novaebritanniae (see below).

The small specimens from Papua New Guinea can be

distinguished from the holotype in having an apparently

broader uropodal endopod (1.25 times as long as wide vs 1.4

times). They did not possess the fine granular lateral surface

in the gape of the cheliped illustrated by Poore and Griffin

(1979). Sakai (1999: fig. 32) showed the cobblestone pattern

on the lower branchiostegite to be a little more extensive in

the holotype from Queensland than on the new material from

Papua New Guinea.

This 1s one of two species with a rudimentary exopod on

maxilliped 3, overlooked by Poore and Griffin (1979) but noted

on the holotype by Sakai (1999). The exopod bears a few short

setae. The other is C. kensleyi (Dworschak, 2005); the two

were differentiated by Dworschak (2005) but see further

comments under C. kensleyi below.

Sakai (2017) listed and figured an ovigerous female from

Ambon, Indonesia, as Eucalliax kensleyi but on biogeographical

erounds this is certainly C. bulimba.

Calliaxina kensleyi (Dworschak, 2005)

Figures 1b, c, 6, 7, 12b, c, 1

Eucalliax kensleyi Dworschak, 2005: 205, figs 1—4, 5a, b.— Sakai,

2017: 1122-1124 (partim), fig. 2A-C, E-G.

Calliaxina kensleyi.—Poore et al., 2019: 125, 127, 146, fig. 191.

Material examined. Saudi Arabia, Farasan Banks, Marka Island, sandy

cay, 18.22055° N, 41.32438° E (stn SAFA-010), UF 36699 (male, 5.9 mm).

Madagascar, Tuléar (now Toliara), Thomassin stn 648, MNHN-

IU-2014-10476 (female, 7.5 mm); Thomassin stn 650, MNHN-

IU-2016-8086 (female, 6.6 mm). Nosy Bé, MNHN-IU-2016-8084

(male, 7.4 mm).

Diagnosis. Sternite 7 with transverse groove visible only

laterally, anteromedial lobe rounded. Antennule peduncle

reaching to midpoint of antenna peduncle article 5. Maxilliped 3

exopod vestigial, about twice as long as wide. Major cheliped

merus, lower margin with 2 or 3 proximal truncate teeth;

propodus distomesial margin with setae aligned in 2 groups of

c. 10 and c. 6 setae. Male pleopod | article 2 about 2.5 times as

long as wide; appendix interna obsolete.

Colour. Exoskeleton mostly translucent white, gastric region,

epistome, upper parts of chelipeds pink (fig. Ib, c).

Distribution. Red Sea (Jordan, Saudi Arabia); western Indian

Ocean (Madagascar). Intertidal, shallow sediments.

Remarks. Calliaxina kensleyi shares a vestigial maxilliped

3 exopod with C. bulimba. The two species were compared by

Dworschak (2005) but the differences he listed are subtle and

deserve reinterpretation and addition. The more pronounced

rostrum of C. kensleyi is subtly so, but juveniles appear to have

sharper rostrum in both species. The telson of C. kensleyi was

Indo-West Pacific and Australian Eucalliacidae T

jl X» A) á

jj A AA 24 Wa

3 f KA y f

NS SN

Figure 4. Calliaxina bulimba. Australia, Qld, NMV J12184 (female, 8.7 mm): a, anterior carapace, eyestalks, antennule, antenna; b, telson,

uropod; c, thoracic sternite 7; d, pleonal sternite 1; e, maxilliped 3; f, g, major cheliped (right), mesial, lateral views; h, minor cheliped (left),

mesial view; uropod. Scale bars = 1 mm.

b-e

G.C.B. Poore

Figure 5. Calliaxina bulimba. Australia, Qld, NMV J71676 (male, 4.6 mm): a, left coxa 5, pleonite 1, pleopod 1 in situ, lateral view; b, c, pleopods

1, 2. Papua New Guinea, MNHN-IU-2013-7097 (male, cl. 4.5 mm): d, e, pleopods 1, 2. Scale bars = 1 mm.

said to be widest at its midpoint, whereas that of C. bulimba 1s

widest anteriorly; this difference is less obvious in smaller

specimens. The uropodal exopod is oval in C. kensleyi, more

tapering in C. bulimba, but again less clear in smaller

specimens. In both species, the major cheliped (the slightly

wider one) has a tooth on the fixed finger, whereas the minor

cheliped does not, with the apparent exception of the paratype

of C. kensleyi.

Despite their obvious morphological similarity, Calliaxina

kensleyi and C. bulimba were not the two most

phylogenetically related of the four species 1n Robles et al.'s

(2020) study. Calliaxina kensleyi was sister species to C.

novaebritanniae, and these two more related to C. sakait,

which if true suggests that the vestigial maxilliped 3 exopod

has evolved twice. Ovoid plates are visible on the sternite of

pleonite 1 (fig. 6c) but absent in E. bulimba (fig. 4d).

Dworschak (2005) did not describe the dentition of the

lower margin of the merus of the chelipeds of C. kensleyi; in

the new material there are few proximal denticles (fig. 6e, f).

All specimens of C. bulimba possess two or three proximal

truncated curved teeth and diminishing denticles more distally

(fig. 4f). The distomesial face of the propodus of the chelipeds

has two rows of submarginal setae in both species. On the

major cheliped of C. kensleyi, these setae align in two separate

groups of c. 10 and c. 6 setae (figs 6d, 12c); on the minor

cheliped, the setae align 1n two scarcely separate groups of c.

20 and c. 8 setae (fig. 6f; Dworschak, 2005: fig. 3a, c). On both

chelipeds of C. bulimba, distomesial setae align in two widely

separate groups of c. 9 setae each (fig. 4h). The distolateral

propodus of the chelipeds is granulated in a concavity at the

base of the fingers in some individuals of both species but 1s

absent on others, especially smaller specimens.

The two males identified here as C. kensleyi possess a

male pleopod 1 typical of several species of Calliaxina and

Eucalliaxiopsis: C. novaebritanniae, C. paradoxa, C.

punica, C. sakaii, Eucalliaxiopsis jonesi (Heard, 1989), E.

mcilhennyi (Felder and Manning, 1994) and E. panglaoensis

(all with a broad article 2, appendix interna and curved apex;

see fig. 7h, | and earlier papers (Dworschak, 2006, 2014;

Felder and Manning, 1994; Heard, 1989; de Saint Laurent

and Manning, 1982; ). As noted above, the male pleopod | of

C. bulimba differs from all of these, its second article lacking

an appendix interna and distal curved apex. The appendix

masculina on pleopod 2 of C. kensleyi 1s narrower than that

of C. bulimba.

sakai (2017) 1ncluded C. kensleyi 1n Eucalliax although

Indo-West Pacific and Australian Eucalliacidae 9

SSS uw. "m moo aa n o A sih T ji) f

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Figure 6. Calliaxina kensleyi. Madagascar, MNHN-IU-2014-10476 (female): a, anterior carapace, eyestalks; b, thoracic sternite 7; c, pleonal

sternite | (anterior uppermost); d, e, major cheliped (right), mesial, lateral views, with detail of mesial setae on propodus; f, g, minor cheliped

(left), mesial, lateral views, with detail of mesial setae on propodus. Scale bars = 1 mm.

^ j - Ap p 7 oJ 9, //

77 WZ Z 4 E y A

cg 77 A A;

G.C.B. Poore

Figure 7. Calliaxina kensleyi. Saudi Arabia, UF 36699 (male): a, anterior carapace, eyestalks, antennule, antenna; b, maxilliped 3, exopod, basis,

ischium; c, telson, uropod; d, major cheliped (left), mesial view; e, major cheliped fingers (left), lateral view; f, minor cheliped (right), mesial view;

e, minor cheliped fingers (right), lateral view; h, 1, pleopods 1, 2. Madagascar, MNHN-IU-2014-8084 (male): J, pleopod 1. Scale bars = 1 mm.

the species has all the features of Calliaxina, however

diagnosed. His figure 2A-C, E-G of material from the Red

sea resembles the types from the nearby type locality, Aqaba.

His figure 2D of the cheliped of a female from Ambon has a

curiously triangular carpus. The location suggests C. bulimba,

a species he did not mention. He referred to E. kensleyi as

being from Aqaba and Moreton Bay (Calliaxina bulimba 1s

from Moreton Bay, Qld) but listed the type locality as Alona

Beach, the Philippines (the type locality of E. panglaoensis

Dworschak, 2006). The species was not listed by Sakai (2011).

Calliaxina novaebritanniae (Borradaile, 1900)

Figures Id, e, 8—10, 12d, e, j

Callianassa novae-britanniae Borradaile, 1900: 419, pl. 39 figs

]4a-d.

Callianassa (Callichirus) novae-britanniae.—Borradaile, 1903:

Indo-West Pacific and Australian Eucalliacidae 11

P g

m, details

Figure 8. Calliaxina novaebritanniae. Australia, NMV J71677 (female, 11.5 mm): a, carapace, pleon, dorsal; b, carapace, lateral; c, d, anterior

carapace, eyestalks, antennule, antenna, dorsal and lateral views; e, thoracic sternite 7; f, pleonal sternite 1; g, telson, uropod; h, maxilliped 3; 1,

maxilliped 3, crista dentata, basis, ischium; J, major cheliped (right), mesial with detail of merus margin, distal propodus; k, major cheliped

(right), distal propodus, dactylus, lateral; 1, minor cheliped (left), mesial view, with detail of merus margin, distal propodus; m, minor cheliped

(left), fixed finger, lateral. Scale bars = 1 mm.

12 G.C.B. Poore

LoL "

BE LBL 4

Eur rr y

2d eg igi” :

a AEE Da

f E

(- (Pc 6-6 c.

COC.

renr rer ae

(One

oF fe

tw e

Figure 9. Calliaxina novaebritanniae. Australia, NMV J71677 (female, 11.5 mm): a—c, pereopods 2—4; d, e, pleopods 1, 2. NMV J71678 (male,

8.7 mm): f, anterior carapace, eyestalks; g, telson; h, i, major cheliped (right), propodus, dactylus, mesial and lateral; j, k, minor cheliped (left),

propodus, dactylus, mesial and lateral; l, m, pleopods 1, 2. Scale bars = 1 mm.

Indo-West Pacific and Australian Eucalliacidae

Figure 10. Calliaxina novaebritanniae. Papua New Guinea, MNHN-IU-2014-8837 (male, 4.0 mm): a, anterior carapace, eyestalks, antennule,

antenna; b, pleonal sternite 1, right pleopod l(anterior uppermost); c, telson, uropod; d, maxilliped 3; e—g, pereopods 2—4; h, 1, pleopods 1, 2.

French Polynesia, UF 23954 (female, 5.1 mm): j, anterior carapace, eyestalks, antennule, antenna. Scale bars = 1 mm.

547.— Borradaile, 1904: 753. — De Man, 1928a: 48.—De Man, 192S8b:

29, 92, 93, 114 (part).

Calliax novaebritanniae.— de Saint Laurent and Manning, 1982:

211—224, figs 1c, 2b, 6c.— Sakai, 2005: 202.

Paraglypturus novaebritanniae.— Sakai, 1999: 123, fig. 32d-f.

Calliaxina novaebritanniae.—Ngoc-Ho, 2003: 493.— Sakai,

2011: 500—501.—Dworschak, 2018: 17-19, fig. 1b, c (synonymy,

notes). — Poore et al., 2019: 126, 127, 146.— Robles et al., 2020.

Calliaxina xishaensis Liu and Liang, 2016: 83—87, figs 1, 2. Syn.

nov.

not Callianassa (Callichirus) novaebritanniae.— Sakai, 1966: 161—

171 (record from Japan = C. sakaii [de Saint Laurent, 1979]).

not Callianassa (Callichirus) novaebritanniae var.—De Man,

1928b: fig. 20 (= C. punica [de Saint Laurent and Manning, 1982] fide

de Saint Laurent and Manning [1982]).

Material examined. Papua New Guinea. Madang Province, lagoon,

05° 10.2' S, 145? 50.4' E, 1-3 m (PAPUA NIUGINI stn PR243), MNHN-

IU-2013-7062* (ovigerous female, cl. 8.8 mm). New Ireland Province,

Nago Island, 02° 36.3' S, 150° 46.2' E, 3-12 m (KAVIENG 2014 stn

KRO6), MNHN-IU-2014-8837* (male, 4.0 mm); MNHN-IU-2013-7941*

(female, c. 4.0 mm).

French Polynesia, Moorea Island, NW motus, in channel between

islands and beach, 17.488393° S, 149.91342° W (stn BIZ-148), UF

23954 (female, 5.1 mm)

Australia, Qld, Great Barrier Reef. Myrmidon Reef, 18.27° S,

147.38* E, October 1985: lagoon with “Callianassa” mounds, 8 m, (M.

Riddle stn M/10/1), NMV J71677 (female, 11.6 mm); NMV J71678

(male, 8.7 mm); NMV J71669 (2 males, 8.4 mm; 6 ovigerous females,

6.9-10.3 mm; 1 female, 9.4 mm). Myrmidon Reef, 18.27? S, 147.38° E,

October 1985: shallow lagoon, 4 m, (M. Riddle stn M/10/2), NMV

J71670 (2 males, 11.1 mm; 3 ovigerous females, 10.4 mm); outer reef

flat, 2 m, (M. Riddle stn M/10/3), NMV J71673 (3 males). Rib Reef,

18.48" S, 146.56" E, October 1985: reef flat, 4 m, (M. Riddle stn

R/10/2), NMV J71671 (4 males; 16 females). reef flat, 8 m, (M. Riddle

stn R/10/1), NMV J71675 (6 males, smallest 5.5 mm; 4 females).

Lizard Island, 50 m W of Loomis beach, 14.6816? S, 145.4515? E, (stn

AUST-S 1-036), UF 16947 (ovigerous female, 9.0 mm). Lord Howe

Island, Old Settlement Beach, 31.5? S, 159.1? E (stn LHI 2017 059),

AM P.100430 (2 males, 11.9 mm, 14.6 mm).

Diagnosis. Sternite 7 with transverse groove visible only laterally,

anteromedial lobe rounded. Antennule peduncle reaching to

midpoint of antenna peduncle article 5. Maxilliped 3 exopod

reaching third to half-way along ischium (juveniles only) or

reaching to mid-merus. Major cheliped merus, lower margin with

2 or 3 proximal truncate teeth; propodus distomesial margin with

setae aligned in 2 similar groups of c. 9 setae each. Male pleopod

l article 2 twice as long as wide; appendix interna prominent.

Description of female. Carapace scattered with small shallow

depressions; gastric-abdominal regions together 4.6 times as long

as wide; branchiostegite calcified over dorsal half, separate from

wide ventral uncalcified anterior region and narrower triangular

uncalcified posterior section, the two separated from each other

by squarish calcified plate; posteroventral area without small

calcified plates; cervical groove at 0.57 carapace length, scarcely

obvious on branchiostegite; cardiac sulcus at 0.73 carapace length,

not meeting mid-dorsally, extending obliquely anteroventrally to

lower limit of calcified branchiostegite.

Thoracic sternite 7 1.35 times as long as wide, with well-

defined median sulcus over posterior third, smooth over medial

half, lateral ridge anterior to coxal articulation ending in pit at

quarter of width, with 5 posterolateral clusters of long setae.

Rostrum broadly triangular, length about 0.25 width of

eyestalks. Anterolateral carapace lobe half as long as rostrum,

with rounded apex, dorsal anterolateral margin oblique in

dorsal view as far back as anterodorsal angle of

branchiostegite, with c. 10 submarginal dorsal setae. Pleonites

1—6 lengths relative to cl: 0.42 : 0.47 : 0.30 : 0.30 : 0.32 : 0.47;

pleonite 1 with strong dorsal transverse groove, with pair of

oval sternal plates supporting pleopods, each with lateral row

of c. 20 setae; pleonites 3—5 with lateral tufts of setae; pleonite

6 with 8 lateral rows of setae.

Eyestalk 1.85 times as long as wide at base, tapering slightly

to cornea, then more rapidly to subacute apices, sometimes

with minute mesiodorsal tubercle; cornea round, slightly

domed, 0.6 width of maximum eyestalk width. Antennular

peduncle 2.5 times as long as width of both eyestalks; article 2

3 times as long as wide; article 3 0.6 length of article 2; articles

2 and 3 with ventrolateral row of long setae, continued onto

flagellum. Antennal peduncle 3.2 times as long as width of both

eyestalks, overreaching antennule peduncle by all of article 5;

G.C.B. Poore

scaphocerite ovoid, longer than wide; article 4 c. 7 times as long

as wide; article 5 half as long as article 4. Maxilliped 3 exopod

reaching to midpoint of merus; ischium with convex mesial

margin, expanding from narrow proximomesial corner;

ischium-merus 1.8 times as long as greatest width; crista

dentata of c. 25 even denticles on basis and ischium; carpus-

dactylus together as long as ischium—merus.

Chelipeds subequal, major 1.1 length of minor. Major

cheliped ischium c. 2.2 times as long as distal width, lower

margin with truncate denticles becoming more distinct distally;

merus 2.6 times as long as broad, lower margin with 4—5

truncate proximal denticles, diminishing distally; carpus 1.5

times as wide as upper length, margins carinate; propodus

ereatest width in middle of palm, as wide as upper palm length,

distomesial margin of palm with 2 submarginal groups each of

c. 8 setae plus nearby granules, distolateral margin of palm with

submarginal group of c. 10 setae between fingers; fixed finger

half as long as upper margin of palm, cutting edge with

microdenticles over proximal half, blunt triangular tooth third

way along; dactylus as long as fixed finger, twice as long as wide

at base, cutting edge concave over proximal half; submarginal

tufts of long setae on upper and lower mesial margins of carpus

and propodus, opposing mesial margins of fingers.

Minor cheliped ischium c. 2.3 times as long as distal width,

lower margin with truncate denticles becoming more distinct

distally; merus 1.7 times as long as broad, lower margin with

4—5 truncate proximal denticles, diminishing distally; carpus

1.5 times as wide as upper length, margins carinate; propodus

palm more evenly wide than major, 1.1 times as wide as upper

palm length, distomesial margin of palm with 2 submarginal

groups of c. 10 and c. 12 setae, distolateral margin of palm with

submarginal group of c. 10 setae, plus 2 smaller groups further

from margin between fingers; fixed finger 0.4 times as long as

upper margin of palm, mesial cutting edge with microdenticles,

blunt proximal tooth, lateral cutting edge microdenticulate

defining edge of lateral concavity at base of fixed finger, filled

with granules; dactylus overreaching fixed finger, 3 times as

long as wide at base, cutting edge curved; submarginal tufts of

long setae on upper and lower mesial margins of carpus and

propodus, opposing mesial margins of fingers.

Pereopod 2 merus 2.2 times as long as maximum width;

dactylus c. 3 times as long as upper margin of propodus.

Pereopod 3 merus twice as long as maximum width; carpus as

long as wide; propodus with strongly concave lower margin,

twice as long as mid-length; dactylus 2.5 times axial length of

propodus. Pereopod 4 merus 2.3 times as long as maximum

width; carpus 2.5 times as long as wide; propodus typically

setose, with 2 long distal setae overlapping dactylus.

Pleopod 1 article 1 without distal projection beyond article

2; article 2 longer than article 1. Pleopod 2 endopod 2.5 times as

long as wide; appendix interna about quarter length of endopod.

Uropodal endopod ovoid, anterior margin more convex

than posterior margin, widest about third way along, 1.5 times

as long as wide, with subdistal facial group of c. 8 long setae;

exopod 2.4 times as wide as long (length measured from

peduncle to anterior point of dorsal plate), distal and posterior

margins typically setose, with c. 20 blade-like setae near end of

posterior margin; dorsal plate extending almost over half of

Indo-West Pacific and Australian Eucalliacidae

exopod width, posterior margin with spiniform setae. Telson

1.7 times as wide as long, broadest at mid-length, posterolateral

corner more sharply rounded than lateral and posterior margins;

posterior margin convex, with lateral lines of long setae; dorsal

surface with median row of long setae at anterior quarter.

Adult male. As in female except as follows. Major cheliped

propodus, distomesial margin of palm with 2 submarginal groups

each of c. 8 setae, fixed finger with proximal mesial and lateral

eranulate ridges; dactylus cutting edge with basal blunt tooth.

Minor cheliped propodus, distomesial margin of palm with

2 submarginal groups of c. 12 and c. 15 setae, distolateral margin

of palm with submarginal groups of c. 30 and c. 15 setae; lateral

concavity at base of fixed finger, filled with numerous granules.

Pleopod 1 article 1 twice as long as greatest width; article

2 2.3 times as long as wide, appendix interna acute, without

hooks, subdistal lobe small, distal curved lobe acute,

occupying about third of whole. Pleopod 2 endopod 2.3 times

as long as wide; appendix masculina attached at c. 0.4 of

length, 3.5 times as long as distal width, distal margin convex-

truncate reaching 0.9 length of endopod, with c. 20 facial

setae near midpoint, c. 15 subdistal setae; appendix interna

rod-like, with hooks, about fifth length of appendix interna.

Variation. Branchiostegite always divided by oblique

longitudinal edge between calcified dorsal region and variously

uncalcified ventral region, always with midventral rectangular

plate, sometimes with posteroventral tessellated pattern (Sakai,

1999: fig. 32e), sometimes more or less calcified, often not

calcified. Rostrum more acute in smaller specimens (figs 3a, J).

Cornea larger in smaller specimens (fig. 3J). Maxilliped 3 exopod

reaching mid-ischium in some small specimens. Minor cheliped

propodus fixed finger proximal concavity variously granulate.

Colour. Exoskeleton mostly translucent white, middle of

branchiostegites pink (fig. Id, e).

Distribution. Papua New Guinea, Madang, New Britain (type

locality) and New Ireland provinces; Indonesia, Sulawesi, Bali;

Australia, N Qld; Philippines, Pulawan; French Polynesia.

Lagoon sediments, to 12 m.

Remarks. Calliaxina novaebritanniae 1s unique in the Indo-West

Pacific in having the exopod of maxilliped 3 reaching to the

middle of the merus in adults. Small genetic differences were

found between the pair from Nago Island (New Britain), one

from Madang, one from Palawan, Philippines (NHMW 25366),

and one from Indonesia (NHMW 25783) (Robles et al., 2020).

Nago Island and Madang, Papua New Guinea, are not far distant,

west and east respectively, from the type locality. Morphological

differences between them, the female from French Polynesia and

the large collection from the Great Barrier Reef, Australia, are

small. The telson and uropod of adults are the same, all have a

sulcus on the carapace. The maxillipedal 3 exopod reaches to

mid-merus except in two small specimens where it 1s shorter

than 1n adults, a phenomenon noted by Liu and Liang (2016) in

C. xishaensis. The rostrum is sharper and the eyestalks more

rounded in smaller specimens. Article 2 of pleopod 1 of the

holotype has two similar distal lobes on its 1nner margin (one the

appendix interna) and an apex strongly curving mesially (de

saint Laurent and Manning, 1982: fig. 6). The appendix interna

and second subdistal lobe are less developed in all the Australian

males in this collection (fig. 91). A juvenile male from Papua New

Guinea has a simple pleopod | with three obsolete distal lobes

(fig. 10hb), indicating that this limb metamorphoses with maturity.

The male pleopod 2 has a broad appendix masculina, with distal

and subdistal clusters of long setae, and appendix interna midway

on the endopod (fig. 101).

Calliaxina novaebritanniae co-occurs in one sample from

the Great Barrier Reef with individuals of C. bulimba trom

which it 1s difficult to distinguish at first glance. However, the

exopod of maxilliped 3 reaches to the middle of the merus in

most C. novaebritanniae but 1s vestigial 1n all C. bulimba.

The eyestalks of C. novaebritanniae are narrower than in C.

bulimba (almost twice as long as wide vs 0.8 times as long as

wide in adults) the antennal peduncles are narrower

(antennular peduncle article 2 3.5 times as long as wide vs

twice; antennal Ipeduncle article 4 6 times as long as wide vs

3 times). The telson of C. novaebritanniae is widest at its

midpoint whereas in C. bulimba it tapers from the base,

although the taper is less obvious in juveniles. Pereopods of

C. novaebritanniae are relatively broader than in C. bulimba

(meri of pereopods 2 and 3, 2.5 and 2.0 times as long as wide

vs 2.2 and 2.4 times). The rostrum is more acute in larger C.

novaebritanniae than in similarly sized C. bulimba but is

more acute in small specimens of both species.

Borradaile (1900) based his description on a single male of

total length 37 mm from New Britain (Papua New Guinea).

The holotype was illustrated by Borradaile (1900), de Saint

Laurent and Manning (1982) and Sakai (1999).

Notwithstanding the species’ long and complicated synonymy,

the material listed here and that from the Philippines and

Indonesia (Dworschak, 2018) are the first reported since. De

Man (1928a) described and illustrated a specimen he identified

as a variety of C. novaebritanniae; this was described as

Calliax (now Calliaxina) punica by de Saint Laurent and

Manning (1982), type locality, Salammbo, Gulf of Tunis. De

Man (1928b) identified material from the Maldives as C.

novaebritanniae; this was identified by Sakai (1999) as Calliax

(now Eucalliaxiopsis) aequimana (Baker, 1907) but is almost

certainly E. inaequimana (Dworschak, 2014).

In as far as can be decided from descriptions, C.

novaebritanniae 1s indistinguishable from C. xishaensis Liu

and Liang, 2016. The latter was described from two females

from the South China Sea, not far from Palawan, Philippines. A

specimen from this locality (NHMW 25360) was found to be

genetically similar to those from Papua New Guinea (the type

locality) and Indonesia, suggesting that the species has a wide

distribution. Calliaxina xishaensis is here synonymised with C.

novaebritanniae. Calliaxina sakaii 1s close by in Japan but is

morphologically and genetically quite remote (see below and

Robles et al., 2020).

Sakai’s (1999) designation of the holotype as a lectotype

was unnecessary.

Calliaxina sakaii de Saint Laurent in de Saint Laurent and

Le Loeuff, 19779

Figures 11, 12f, g

Callianassa (Callichirus) novaebritanniae.—Sakai, 1966): 161-

168, figs 1—4.

Calliax sakaii de Saint Laurent in de Saint Laurent and Le Loeuff,

1979: 95.— de Saint Laurent and Manning, 1982: 212, 222—223, figs

1g, 2c.— Sakai, 1987: 305—306.— Sakai, 2005: 203.

Paraglypturus sakaii.— Sakai, 1999: 124, fig. 33d, e.

Calliaxina sakaii.—Ngoc-Ho, 2003: 489, 493, 496, fig. 20J.—

Sakai, 2011: 502.— Sakai, 2018: 739.

Material examined. Japan, Tomioka, MNHN Th312 (holotype male,

10mm), MNHN Th313 (female), NMV J59763 (male, 6.7 mm; female,

7.0 mm; ex ULLZ-8894*).

Diagnosis. Sternite 7 with complete transverse V-shaped

groove, anteromedial lobe acute. Antennule peduncle reaching

to base of antenna peduncle article 5. Maxilliped 3 exopod

reaching third to half-way along ischium. Major cheliped

merus, lower margin with several proximal teeth; propodus

distomesial margin with setae aligned in 4 groups of 10, 5, 7

and 4 setae. Male pleopod | article 2 3 times as long as wide;

appendix interna prominent.

Remarks. Differences between C. sakaii and the more widespread

C. novaebritanniae were confirmed by re-examination of the

holotype and more recently collected material. The male pleopod

l has a bilobed proximal lobe and pleonal sternite | has a pair of

oval plates supporting the pleopods, both similar to C

novaebritanniae. |Calliaxina | sakaii differs from C.

novaebritanniae in the exopod of maxilliped 3 reaching half the

length of the ischium (vs overlapping the merus), a more acute

rostrum, fully calcified branchiostegites (vs weakly chitinised

over the lower half), having a deep sharp transverse groove on

thoracic sternite 7 (vs an obsolete ridge) and the posterior margin

of the telson concave (convex). The distomesial margin of the

propodus of the major cheliped has four groups of 4-10

submarginal setae, two groups on the minor cheliped (vs two

eroups on both chelipeds). A lateral concavity at the base of the

fixed finger is defined by a lower ridge on both chelipeds.

The species was described briefly in footnote number 14

by M. de Saint Laurent alone in de Saint Laurent and Le

Loeuff (1979).

Eucalliaxiopsis Sakai, 2011

Eucalliaxiopsis Sakai, 2011: 503—504.— Poore et al., 2019: 127—

128 (synonymy).

Remarks. Eucalliaxiopsis differs from Calliaxina in having a

strong ridge across the telson, lacking an exopod on maxilliped

3 (with one exception), the appendix masculina overlapping the

endopod of pleopod 2, and in the major cheliped of the male

usually being more robust, the palm longer and wider, than that

of the female. In Calliaxina, the telsonic ridge is absent or

obsolete, maxilliped 3 always possesses an exopod, the appendix

masculina does not reach the apex of the endopod of pleopod 2,

and chelipeds of males and females are not differentiated from

each other nor scarcely from the minor chelipeds. In as much as

G.C.B. Poore

can be deduced from published illustrations, the lower margin

of the merus of the chelipeds is smooth or with small tubercles

at its midpoint in Eucalliaxiopsis, whereas this margin bears

small proximal truncate teeth in Calliaxina. The post-rostral

dorsal area of some species of Eucalliaxiopsis (E. aequimana,

E.madagassa) has a broad median longitudinal ridge, seen also

in Eucalliax quadracuta (Biffar, 1970).

Eucalliaxiopsis includes seven species from the Indo-

West Pacific, of which three are newly described here, plus

three species from the Western Atlantic: E. cearaensis

(Rodrigues and Manning, 1992) from Brazil, E. jonesi from

the Bahamas and E. mcilhennyi from Florida, USA. The

species identified as EF. HWI from Hawaii by Robles et al.

(2020), which is genetically quite separate from six other

species, was not available for study.

Eucalliaxiopsis panglaoensis, E. dworschaki and E.

paradoxa, the last two being new species described here,

differ from the others. All possess a transverse telson ridge

beset with a row of short spiniform setae, a distally excavate

uropodal endopod, a cheliped carpus with two distal spines

on the upper margin, two mesiodistal lobes near the apex of

the lower lateral margin plus a submarginal distal ridge on the

mesial face. The carpus is similar to that of Eucalliax

quadracuta, sole member of its genus. The first two of this

eroup were found to be sister taxa by Robles et al. (2020) but

the others were not monophyletic.

Calliaxiopsis Sakai and Türkay, 2014, was synonymised

with Calliaxina by Poore and Dworschak (2017). Bakercalliax

Sakai, 2018, Heardcalliax Sakai, 2018, and Manningcalliax

Sakai, 2018, were all differentiated from Eucalliax, Calliaxina

and from each other largely on the basis of the male pleopod 1

but also on the relative sizes of the appendices interna and

masculina of pleopod 2. All were synonymised with

Eucalliaxiopsis by Poore et al. (2019), who argued that similar

forms of the male pleopod 1 are found in Eucalliaxiopsis and

Calliaxina and that some of the forms used as generic

characters by Sakai (2018) are characteristic of juveniles.

Key to Indo-West Pacific species of Eucalliaxiopsis

|. Telson with row of spiniform setae on transverse ridge;

cheliped carpus with 2 short grooves perpendicular to

distolateral margin near lower apex; uropodal endopod

with apical concavity 2

— Telson without spiniform setae on transverse ridge;

cheliped carpus distolateral margin smooth; uropodal

endopod ovate 4

2. Cheliped carpus upper margin with proximal hook and

subproximal submesial pit; cheliped propodus lateral face

with longitudinal ridge extending from near carpus to

base of finger; uropodal endopod with 8 short setae

between apical clusters; widespread in Indo-West Pacific

E. dworschaki sp. nov.

— A Cheliped carpus upper margin smooth; cheliped propodus

lateral face with longitudinal ridge extending proximally

slightly from base of finger; uropodal endopod with 10—

1] short setae between apical clusters 3

Indo-West Pacific and Australian Eucalliacidae

u g W 1 i NNNM N AAN

Figure 11. Calliaxina sakaii. Japan, NMV J59763 (male, 6.7 mm): a, telson, uropod; b, thoracic sternite 7; c, pleonal sternite | (anterior uppermost);

d, pleopod 1; e, f, major cheliped propodus, dactylus (right), mesial, lateral views; g, h, minor cheliped propodus, dactylus (left), mesial, lateral

views. Scale bars = | mm.

Chelipeds asymmetrical in male, major cheliped

propodus longer than wide and longer than in female;

distolateral lobe above lower margin of cheliped carpus

with acute apex, ridged, oblique; Philippines, Japan

E. panglaoensis (Dworschak, 2006)

9 9$890998989899296098909948900892849828696899539a490290894092969€809658928929€498

Chelipeds similar in both sexes, major cheliped palm little

longer than wide; distolateral lobe above lower margin of

cheliped carpus with rounded apex, longitudinally

grooved, perpendicular; NE Australia |...

E. paradoxa sp. nov.

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Maxilliped 3 with exopod; cheliped merus lower margin

with 2 denticles at midpoint, fixed finger with row of

equally spaced setae along lateral ridge; telson with

rounded posterior margin; female pleopod 2 with free

extension beyond rami; Madagascar s

E. madagassa (Sakai and Türkay, 2014)

Maxilliped 3 without exopod; cheliped merus lower

margin smooth, fixed finger with clusters of setae along

lateral ridge; telson with truncate or excavate posterior

margin; female pleopod 2 without extension beyond rami .

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Cheliped propodus upper mesial face with row of clusters

of setae without associated transverse ridges; major

cheliped of male upper margin as long as wide, similar to

female; male pleopod | article 1 without distal setae,

article 2 parallel-sided, apically rounded, with marginal

setae, without appendix interna; 5 Australia...

E. aequimana (Baker, 1907)

Cheliped propodus upper mesial face with row of short

transverse ridges associated with clusters of setae; major

cheliped of male upper margin longer than wide,

dissimilar to female; male pleopod 1 not as above 6

G.C.B. Poore

pr =

Figure 12. Calliaxina spp. C. bulimba: a, major cheliped, mesial. C. kensleyi: b, c, major cheliped lateral, mesial. C. novaebritanniae: d, major

cheliped, mesial; e, minor cheliped, mesial. C. sakaii: f, g, minor cheliped mesial, lateral. Thoracic sterna: h, C. bulimba; 1, C. kensleyi; j,

C. novaebritanniae. Various scales.

6. Antenna peduncle 3 times as long as base of both eyes,

most of article 5 exceeding antennule; telsonic ridge

sharply elevated, with median notch; uropodal endopod

1.8 times as long as wide; female pleopod 2 without

appendix interna; widespread in Indo-West Pacific ...........

E. inaequimana (Dworschak, 2014)

— Antenna peduncle 2.4 times as long as base of both eyes,

half of article 5 exceeding antennule; telsonic ridge

slightly elevated, complete; uropodal endopod 1.5 times

as long as wide; female pleopod 2 with appendix interna;

Papua New Guinea E. patio sp. nov.

Peo eee heehee HHO HOO H ETE H HOHE H OHH HERE HH ESO HERE H EET

Eucalliaxiopsis aequimana (Baker, 1907)

Figures 13, 14, 25a, b

Callianassa aequimana Baker, 1907: 182-185, pl. 24 figs 1-8.—

Hale, 1927: 87, fig. 83.—Poore and Griffin, 1979: 245, figs 12, 13.

Callianassa (Callichirus) aequimana.— De Man, 1928b: 28, 93, 114.

Calliax aequimana.—de Saint Laurent and Manning, 1982:

222.— Sakai, 1988: 53, 61.— Sakai, 1999: 118—119 (not fig. 31).—Tudge

et al., 2000: 145.—Davie, 2002: 459.— Sakai, 2005: 202.

Eucalliax aequimanus.— Poore, 2004: 184, f1g.50e, f.

Calliaxina aequimana.—Sakai, 2011: 498—499.

Eucalliax aequimana.— Dworschak, 2014: 236, 244, figs 8, Oh, 1.

Bakercalliax aequimana.—Sakai, 2018: 738.

Material examined. Australia. NSW, Balmoral Beach, 33.8? S, 151.3? E,

AM P3636 (male, 20 mm).

Victoria. San Remo, NW from rock outcrop 1.1 km E of Back

Beach Rd (MRG transect 4), 38.53" S, 145.38? E, NMV J59759

(female, 9.6 mm). San Remo, channel bank end, 38.5233* S, 145.363? E,

NMV J59762 (female, 6.6 mm). Port Phillip Bay, Geelong arm, 3 m,

(NMV stn PPBES 953), 38.155? S, 144.545? E, NMV 716782 (4

females, 3.4—4.7 mm).

Tasmania. Waterhouse Island, 3—5 m, 40° 48' S, 147° 38.7' E

(Moverley stn 52), NMV J71685 (juv., 2.9 mm). Georges Bay, Steiglitz

Beach, 3—4 m, 41° 19.3'S, 148° 19.1' E (Moverley stn 42) NMV J71686

(juv., 2.9 mm).

South Australia. Port MacDonnell, 38.05? S, 140.7? E, NMV

Indo-West Pacific and Australian Eucalliacidae 19

WH TTT BN FA 3

Figure 13. Eucalliaxiopsis aequimana. Australia, NMV J71687 (male, 11.0 mm): a, carapace, dorsal view; b, carapace, lateral view; c, d, anterior

carapace, eyestalks, antennule, antenna, dorsal and lateral views; e, eyestalk, lateral view; fh, telson, dorsal, postero-oblique, lateral views; 1, right

uropod; j, maxilliped 3; k, 1, major cheliped (left), mesial and lateral views; m, n, minor cheliped (right), mesial and lateral views. Scale bars = 1 mm.

G.C.B. Poore

Figure 14. Eucalliaxiopsis aequimana. Australia, NMV J71687 (male, 11.0 mm): a, b, d, pereopods 2, 3, 5; e, pleonal sternite 1, right pleopod

l(anterior uppermost); g, h, left pleopods 1, 2, posterior views. NMV J759651 (female, 11.2 mm): c, pereopod 4; f, pleonal sternite 1, right pleopod

1; 1, left pleopod 1, mesial view; J, k, anterior carapace, antennule, antenna, lateral and dorsal views. Scale bars = 1 mm.

J16783 (male, 12.8 mm). Dutton Bay, eastern shore, 34.0489° S,

136.415° E, NMV J71688 (male, 12.2 mm; 3 females, 8.9-11.9 MM),

NMV J59651* (female, 11.2 mm). NMV J71687 (male, 11.0 mm).

Western Australia. Houtmans Abrolhos, WAM C9113 (male, 14.8

mm). Dalkeith, Swan River, WAM C1410 (male, 16.2 mm). Colpoy

Point, Garden Island, 32° 12'S, 115° 40' E, WAM C11975 (female, 14.4

mm). Rockingham, Point Peron, Cockburn Sound, 32° 11'S, 115° 43' E,

WAM C11973 (male, 12.9 mm). Fremantle Harbour, 32° 03' S,

115° 44' E, WAM C12030 (male, 20.3 mm). Mandurah, 32° 32' S,

115° 43' E, WAM C11978 (female, 15.8 mm). Bunbury, 33° 20' S,

115° 38' E, WAM C11990 (female, 10.1 mm). Harbour Board, Bunbury,

WAM C5056 (female, 21.3 mm). Koombana Beach, Bunbury,

33° 20' S, 115° 38' E, WAM C12029 (male, 14.9 mm). Leschenault

Estuary, Bunbury, 33° 20' S, 115° 38' E, WAM C11986 (male, 11.2

mm). Peel Inlet, 32° 36' S, 115° 43' E, WAM C11968 (male, 8.9 mm;

female, 17.6 mm). Albany, 32° 00'S, 117° 52'E, WAM C11994 (female,

13.0 mm). Albany, WAM C6792 (female, 15.9 mm). Cheyne Beach,

WAM C11987 (female, 19.8 mm). King George Sound, MNHN Th578

(female).

Diagnosis. Pleonite | with pleopods attached to pair of sternal

plates. Eyestalk about twice as long as wide. Antennule peduncle

reaching to base of antenna peduncle article 5. Maxilliped 3

exopod absent. Cheliped carpi distolateral margin square at

lower angle, propodi lateral face with oblique longitudinal ridge

extending from beyond mid-length of palm on to base of finger,

upper mesial face with row of clusters of long setae. Male

pleopod 1 article I without distal setae, article 2 parallel-sided,

apically rounded, with marginal setae, without appendix

interna. Female pleopod 2, appendix interna present. Uropod

endopod ovate. Telson without transverse row of spiniform setae.

Description of male. Carapace smooth, with pair of shallow

longitudinal groove posterior to rostrum; gastric—abdominal

regions together 4.5 times as long as wide; branchiostegite

calcified, with tessellated pattern of sulci; cervical groove at

0.6 carapace length, scarcely obvious on branchiostegite;

cardiac sulcus at 0.76 carapace length, not meeting mid-

dorsally, extending transversely across half of branchiostegite,

with complex branches dorsally.

Rostrum blunt or obsolete. Anterolateral carapace lobe

longer than rostrum, with rounded apex, anteriorly directed,

with numerous submarginal dorsal setae.

Thoracic sternite 7 1.15 times as long as wide, with well-

Indo-West Pacific and Australian Eucalliacidae

defined median sulcus over posterior half, lateral groove anterior

to coxal articulation poorly defined, oblique, not meeting medially,

with 3 posterolateral clusters of long setae. Pleonal sternite with

pair of swollen well marked areas anterior to ridge bearing

pleopods; without setae. Pleonite 6 lateral margin smooth.

Eyestalks 1.6 times as long as wide at base, tapering around

cornea, acute apical tubercle sometimes present; cornea

subcircular. Antennular peduncle 2.5 times as long as width of

both eyestalks; article 2 2.5 times as long as wide; article 3 0.6

times as long as article 2; articles 2 and 3 with ventrolateral row

of long setae, continued onto flagellum. Antennal peduncle 3

times as long as width of both eyestalks, overreaching antennule

peduncle by third of article 5; scaphocerite wider than long,

subcircular; article 4 c. 5 times as long as wide; article 5 0.6

length of article 4. Maxilliped 3 ischium with slightly convex

mesial margin, expanding from narrow proximomesial corner;

ischium—merus upper margin 1.5 times as long as greatest

width; crista dentata of c. 25 small teeth; carpus—dactylus

together almost as long as ischium—merus.

Chelipeds subequal, propodus of major cheliped about 1.2

times that of minor, of similar widths, major dactylus stouter

than that of minor. Major cheliped ischium twice as long as

distal width, lower margin with short distal tooth; merus 1.8

times as long as broad, lower margin barely convex, unarmed;

carpus 1.4 times as wide as upper length, upper margin carinate,

distomesial and distolateral margins simple; propodus greatest

width proximally, upper palm length 1.1 width, distomesial

margin of palm with 2 submarginal groups each of c. 10 setae

plus small submarginal bicuspid tooth, distolateral margin of

palm with submarginal group of c. 10 setae between fingers plus

small submarginal tooth; fixed finger half as long as upper

margin of palm, cutting edge with microdenticles over proximal

half, obsolete tooth at midpoint, with obsolete lateral ridge

extending on to palm; dactylus overreaching fixed finger, 2.2

times as long as width at base, cutting edge sinuous; submarginal

tufts of long setae on upper and lower mesial margins of carpus

and propodus, opposing mesial margins of fingers.

Minor cheliped ischium twice as long as distal width,

lower margin with distal tooth; merus twice as long as broad,

lower margin barely convex, unarmed; carpus 1.2 times as

wide as upper length, upper margin carinate, distomesial and

distolateral margins simple; propodus palm as wide as upper

palm length, upper margin carinate, with blunt distal tooth,

distomesial margin of palm with 2 submarginal groups of c. 10

and c. 12 setae, distolateral margin of palm with 2 submarginal

groups of c. 8 and 10 setae; fixed finger half as long as upper

margin of palm, cutting edge smooth, lateral concavity at base

of fixed finger sharply defined by ridge on distal palm; dactylus

barely overreaching fixed finger, 3.8 times as long as wide at

base, cutting edge sinuous; submarginal tufts of long setae on

upper and lower mesial margins of carpus and propodus,

opposing mesial margins of fingers.

Pereopod 2 merus twice as long as maximum width;

dactylus 3 times as long as upper margin of propodus.

Pereopod 3 merus twice as long as maximum width; carpus

1.6 times as long as wide; propodus with lower margin

concave, 1.6 times as long overall as mid-length; dactylus half

axial length of propodus. Pereopod 4 merus length 2.7 times

maximum width; carpus 2.7 times as long as wide; propodus

1.5 times as long as wide, typically setose, with | long distal

seta overlapping dactylus.

Pleopod | article | without distal setae; article 2 parallel-

sided, apically rounded, 1.8 times as long as article |, 4 times as

long as wide, apex with c. 10 marginal setae, without appendix

interna. Pleopod 2 endopod 2.4 times as long as wide; appendix

masculina overreaching endopod by about fifth its length, setose

along posterior face and distally; lobe-like appendix interna near

midpoint of appendix masculina. Pleopod 3 with appendix interna

submerged in endopod margin.

Uropodal endopod anterior margin convex, posterior margin

almost straight, widest about third way along, 1.7 times as long

as wide, apex broadly rounded, with 2 groups of long setae at

ends of anterior and posterior margins; exopod ovate, all

margins continuous, greatest dimension 1.5 times greatest width,

with row of blade-like distal setae on distal margin; dorsal plate

extending beyond half of exopod width, distal margin with

spiniform setae. Telson 1.5 times as wide as long, broadest at

midpoint, posterolateral corner rounded; posterior margin

deeply excavate, with posterolateral cluster of long setae plus

short spiniform seta; dorsal surface with sharp transverse ridge

at anterior third, notched and with few long setae at midpoint.

Female. Essentially as male except: pleonal sternite 1 with pair of

oval plates supporting pleopods, each with longitudinal row of 6—8

single setae emerging from pore. Pleopod | peduncle curved, with

dense group of plumose setae; article 2 twice as long as article 1.

Distribution. Australia, NSW (as far north as 33? S), Tas., Vic.,

SA, WA (as far north as 25? S). Intertidal to subtidal sediments.

Remarks. Eucalliaxiopsis aequimana is recognised by the

subequal chelipeds that are not sexually dimorphic. In a sample

of 14 large specimens, the ratio between the upper propodus

margins of major and minor chelipeds ranged from 1.1 to 1.3,

with insignificant difference between males and females. The

species 1s notable for the simple setose article 2 of the male

pleopod 1, longer than article 1, lacking an appendix interna and

acute apex.

The collection from Western Australia includes specimens

with carapace lengths ranging 8.9—20.5 mm, most are larger

than any from South Australia or Victoria, maximum cl. 12.8

mm. They differ most obviously in the near absence of a

prominent rostrum (fig. 13)).

sakars (2005) synonymy includes references to other

species from places outside southern Australia.

Eucalliaxiopsis dworschaki sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: ESBF2226-256F-

4EF7-9DEB-DFD6955BB744

Figures If, g, 15—17, 25c-f

Eucalliax panglaoensis.—Dworschak, 2006: 356 (partim), fig.

5.— Kneer et al., 2013: 265.—Dworschak, 2018: 17 (partim; not fig. 1 =

E. panglaoensis).

Calliaxina panglaoensis.—Sakai and Türkay, 2014: 191—192, fig.

12.

Eucalliaxiopsis aff. panglaoensis.— Robles et al., 2020.

22 G.C.B. Poore

Figure 15. Eucalliaxiopsis dworschaki sp. nov. Holotype, MNHN-IU-2014-10003 (male, 8.9 mm): a, b, carapace, pleonite 1, dorsal and lateral

views; c, d, anterior carapace, eyestalks, antennule, antenna, dorsal and lateral views; e, thoracic sternite 7; f, pleonite 1, sternite (anterior

uppermost); g, pleonite 6, right lateral margin, oblique view; h, 1, maxilliped 3; J, maxilliped 3, crista dentata; k, left uropod; |, m, telson, dorsal

and lateral views. Scale bars = 1 mm.

Indo-West Pacific and Australian Eucalliacidae 23

dr P |o tht

T e T

—————————— |

a-l, p

m, n

Figure 16. Eucalliaxiopsis dworschaki sp. nov. Holotype, MNHN-IU-2014-10003 (male, 8.9 mm): a, b, major cheliped (left), mesial and lateral views;

c, d, major cheliped (left), carpus, propodus, dactylus, upper and lower views; e, f, minor cheliped (right), mesial and lateral views; g, h, minor

cheliped (right), carpus, propodus, dactylus, upper and lower views; 1-1, pereopods 2—5; m, right pleopod 1, mesial view; n, right pleopod 2, anterior

view; o, appendices masculina, interna, posterior view; p, right pleopod 3, posterior view. UF 28877 (male, 6.0 mm): q, pleopod 1. Scale bars = 1 mm.

24 G.C.B. Poore

Figure 17. Eucalliaxiopsis dworschaki sp. nov. Paratype, MNHN-IU-2017-1351 (female, 8.2 mm): a, b, major cheliped (right), mesial and lateral

views; c, carpus, lower distolateral margin, detail; d, e, minor cheliped (left), mesial and lateral views; f, carpus, lower distolateral margin, detail;

g, pleonite 1; h, pleonal sternite 1, left pleopod 1 in situ (anterior uppermost); 1, left pleopod 1, anterior view; J, k, left pleopod 2, posterior view,

lateral view of peduncle. Scale bars = 1 mm.

Indo-West Pacific and Australian Eucalliacidae

Material examined. Holotype. Papua New Guinea, New Ireland, NW

corner of Little Nusa Island, sand, 0-1 m, 02° 34.9' S, 150° 46.8' E

(KAVIENG 2014 stn KM13), MNHN-IU-2014-10003* (male, 8.9 mm).

Paratypes. Papua New Guinea. Collected with holotype, MNHN-

[U-2017-1351 (female, 8.2 mm). Madang Province, Riwo, mangrove,

05* 09' S, 145° 48.2' E, 1-2 m (PAPUA NIUGINI stn PR235), MNHN-

[U-2013-7081 (female, 7.3 mm).

French Polynesia, Moorea, Papetoai, near 17.49" S, 149.88° W: stn

BIZ-463, UF 28782* (female, 8.4 mm); stn BIZ-493, UF 28930 (female,

4.4 mm); stn BIZ-539, UF 28878* (male, 7.9 mm), UF 28877* (male,

6.0 mm); stn MIB-208, UF 16416 (female, 6.7 mm); stn MIB-195, UF

16286 (male, 8.8 mm); stn MIB-227, UF 16531 (male, 5.4 mm). Off

Nihimaru River estuary, 17.533° S, 149.9045° W, stn MIB-098, UF

15946 (male, 2.9 mm).

Other material. Australia, Qld, Great Barrier Reef, Myrmidon

Reef, 18.27? S, 147.38° E, October 1986: lagoon with “Callianassa”

mounds, 4 m, (M. Riddle stn M/10/2), NMV J71681 (male, 8.4 mm); 8

m, (M. Riddle stn M/10), NMV J71682 (female, 9.2 mm).

Marianas Island, W coast of Guam, near Fisheye Marine Park,

seagrass bed, 0.2-0.5 m (stn AA05), UF GUOK 10-0378, (male, 3.7

mm).

Madagascar, Toliara (as Tuléar), Thomassin stn 678, MNHN-

[U-2016-8083 (female, 8.9 mm).

Diagnosis. Pleonite 1 with pleopods attached to pair of sternal

plates. Eyestalk about twice as long as wide. Antennule peduncle

reaching to midpoint of antenna peduncle article 5. Maxilliped 3

exopod absent. Cheliped carpi distolateral margin with 5 rounded

lobes near lower margin separated by submarginal grooves

parallel to lower margin, second lobe with slight median

depression, propodi lateral face with horizontal longitudinal

ridge extending from near carpus to base of finger, upper mesial

face with row of clusters of long setae. Male pleopod | article |

linear, with distal setae; article 2 blade-like with subapical medial

notch. Female pleopod 2, appendix interna absent. Uropod

endopod ovate with excavate apex. Telson with row of spiniform

setae on ridge.

Description of holotype male. Carapace length 8.9 mm,

smooth; gastric-abdominal regions together 4.7 times as long

as wide; branchiostegite fully calcified; cervical groove at 0.60

carapace length, scarcely obvious on branchiostegite; cardiac

sulcus at 0.77 carapace length, not meeting mud-dorsally,

extending transversely across third of branchiostegite.

Rostrum acute, tapering to narrow tip, length about 0.4

width of eyestalks. Anterolateral carapace lobe almost as long

as rostrum, with rounded apex, anteriorly directed, with

numerous submarginal dorsal setae.

Thoracic sternite 7 1.2 times as long as wide, with well-

defined median sulcus over posterior half, lateral groove

anterior to coxal articulation well defined, oblique, meeting

medially, with 3 posterolateral clusters of long setae. Pleonal

sternite 1 with pair of swollen relatively unchitinised areas

anterior to ridge bearing pleopods; without setae. Pleonite 6

with small lateral blunt hook-like process at midpoint.

Eyestalks twice as long as wide at base, tapering to cornea

and acute apical tubercle; cornea subcircular. Antennular

peduncle 3 times as long as width of both eyestalks; article 2

3.5 times as long as wide; article 3 half as long as article 2;

articles 2 and 3 with ventrolateral row of long setae, continued

onto flagellum. Antennal peduncle 3.4 times as long as width

of both eyestalks, overreaching antennule peduncle by most

of article 5; scaphocerite wider than long, with acute apex;

article 4 c. 5 times as long as wide; article 5 0.65 length of

article 4. Maxilliped 3 ischium with slightly convex mesial

margin, expanding from narrow proximomesial corner;

ischium—merus upper margin 1.6 times as long as greatest

width; crista dentata of 12 spines, the most proximal reflexed;

carpus—dactylus together almost as long as ischium-merus.

Chelipeds unequal, of similar lengths, propodi of similar

widths, major dactylus stouter than that of minor. Major cheliped

(left in holotype) ischium twice as long as distal width, lower

margin with short spines becoming more distinct distally; merus

1.8 times as long as broad, lower margin mostly straight,

unarmed; carpus 1.4 times as wide as upper length, upper margin

carinate, with proximal hooked tooth, subproximal mesial pit,

distal tooth, distomesial margin with prominent lobe near lower

margin, distomesial face with lower submarginal ridge,

distolateral margin with rounded notch near upper margin, with

3 rounded lobes near lower margin separated by submarginal

erooves parallel to lower margin, second lobe with slight median

depression, 2 short setae in lower groove; propodus greatest

width in middle of palm, upper palm length equal to greatest

width, upper margin carinate, with rounded lobe projecting over

dactylus, lower margin carinate, strongly curved mesially, mesial

face with short distal uneven setose ridge below upper margin,

distomesial margin of palm almost straight, lateral face of palm

with prominent ridge running from tubercle near proximal

margin on to finger, oblique row of tubercles at proximal end of

ridge, each perforated for cluster of setae; fixed finger half length

of upper margin of palm, cutting edge with denticles over

proximal half, blunt triangular tooth at about midpoint; dactylus

as long as fixed finger, twice as long as wide at base, cutting edge

with blunt molar, with deflected tip; submarginal tufts of long

setae on upper and lower mesial margins of carpus and propodus,

opposing mesial margins of fingers.

Minor cheliped (right in holotype) ischium twice as long as

distal width, lower margin with distal spine; merus 1.7 times as

long as broad, lower margin mostly straight, unarmed; carpus

1.4 times as wide as upper length, upper margin carinate, with

proximal hooked tooth, subproximal mesial pit, distal tooth,

distomesial face with lower submarginal ridge, distolateral

margin with notch near upper margin, with 3 lobes near lower

margin separated by perpendicular submarginal grooves,

middle lobe apex rounded, with shallow median groove, 3

short setae in lower groove; propodus greatest width in middle

of palm, upper palm length equal to greatest width, as wide

distally as proximally, upper margin carinate, overhanging

mesial face, with rounded lobe projecting over dactylus, lower

margin carinate, strongly curved mesially, distomesial margin

of palm almost straight, lateral face of palm with prominent

ridge running from tubercle near proximal margin on to finger,

oblique row of tubercles at proximal end of ridge, each

perforated for cluster of setae; fixed finger 0.7 length of upper

margin of palm, width at base half width of palm, cutting edge

smooth; dactylus as long as fixed finger, 3.5 times as long as

wide at base, cutting edge straight; submarginal tufts of long

setae on upper and lower mesial margins of carpus and

propodus, opposing mesial margins of fingers.

Pereopod 2 merus twice as long as maximum width;

dactylus c. 2.5 times as long as upper margin of propodus.

Pereopod 3 merus 2.2 times as long as maximum width; carpus

1.8 times as long as wide; propodus with lower margin concave,

1.5 times as long overall as mid-length; dactylus 0.7 times

length of axial length of propodus. Pereopod 4 merus length 2.7

times maximum width; carpus 2.7 times as long as wide;

propodus 1.5 times as long as wide, typically setose, with 1

long distal seta overlapping dactylus.

Pleopod 1 article | distally setose; article 2 as long as article

l, 4 times as long as wide, with apex curved over subdistal

notch, appendix interna freely produced, unarmed. Pleopod 2

endopod 2.1 times as long as wide; appendix masculina

overreaching endopod by about half its length, setose along

posterior face and distally; appendix interna near base of

appendix masculina with 5 small hooks. Pleopod 3 with

appendix interna submerged in endopod margin.

Uropodal endopod anterior margin convex, posterior

margin convex, widest about third way along, 1.4 times as

long as wide, apex excavate, with rows of long setae at ends of

anterior and posterior margins, with 11 short setae between in

excavation, with subdistal group of facial setae; exopod ovate,

anterior margin straight, distal and posterior margins

continuous, greatest dimension 2.3 times anterior margin,

with row of blade-like distal setae on distal margin, c. 6 not

overlapping with densely setose distal margin; dorsal plate

extending almost half of exopod width, distal margin with

spiniform setae. Telson 1.8 times as wide as long, broadest at

midpoint, posterolateral corner squarish; posterior margin

sinuous, lateral regions most prominent, with posterolateral

cluster of long setae plus 2 short spiniform setae; dorsal

surface with sharp transverse ridge at anterior third, with 30

short spiniform setae, few long setae at midpoint.

Female. Essentially as male except: major cheliped palm more

tapering than in male, widest proximally. Pleonal sternite |with

pair of oval plates supporting pleopods, each with longitudinal

row of c. 20 setae in clusters of 4 or 5. Pleopod | peduncle with

mesial setose lobe longer than base. Pleopod 2 peduncle with

similar lobe directed mesially.

Colour. Exoskeleton mostly translucent white, upper parts of

chelipeds pink (fig. If, g).

Etymology. For Peter C. Dworschak, good friend and colleague,

and in recognition of his career devoted to the biology of

"thalassinideans .

Distribution. Madagascar, Philippines, Mariana Islands, Papua

New Guinea, Indonesia, French Polynesia. Intertidal to 8 m.

Remarks. | Eucalliaxiopsis | dworschaki 1s similar to

E. panglaoensis and E. paradoxa (see remarks under

Eucalliaxiopsis above). Eucalliaxiopsis dworschaki differs

from both species in possession of a proximal hooked spine

and subproximal-mesial pit on the upper margin of the carpus

of both chelipeds (figs 16g, 25d, e) and a lateral ridge extending

almost the entire length of the propodus of both chelipeds (figs

16b, 25b). The palm of the major cheliped of the male is a little

longer than wide, while it is 1.25 times as long as wide in

G.C.B. Poore

E. panglaoensis. The telson is relatively shorter (length: width

ratio 1.8 vs 1.6) and with a more sinuous posterior margin, and

the uropodal endopod is broader (length: width ratio 1.4 vs

1.6). Dworschak (2006; pers. comm. 29 September 2019)

reported a small spine on the merus of maxilliped 3 of the

allotype of E. panglaoensis, and of seven from Sulawesi, here

reidentified as E. dworschaki. None of the material examined

for this study, nor Dworschak's two individuals from Bali,

possess this spine. Eucalliaxiopsis paradoxa is compared

below.

Dworschak (2006: 356, fig. 5) remarked on and illustrated

two females from Panglao that differed from typical

E. panglaoensis. He noted the hook and pit on the upper margin

of the cheliped carpus and the longitudinal ridge on the

propodus, characters used here to differentiate E. dworschaki.

The genetic difference from E. panglaoensis from the type

locality in the Philippines (Robles et al., 2020) and consistent

morphology of E. dworschaki over a wide geographic range in

the Indo-West Pacific support recognition of two species.

Dworschak (2018) listed material of E. panglaoensis from

Sulawesi and Bali, Indonesia. He has now confirmed (pers.

comm. 30 October 2018) that this collection of five males and

four females also belongs to E. dworschaki. The two species

occur sympatrically at Panglao and a single individual of

E. dworschaki was recorded from Guam, not far from the

Philippines. A large sample of ghost shrimps from the Great

Barrier Reef, a sample that included specimens of Corallianassa

sp. (Callichiridae) and Thomassinia aimsae Poore, 1997

(Callianideidae), was dominated by E. paradoxa but also

included one specimen of E. dworschaki.

There is no genetic difference between the individual

from Papua New Guinea and the three sequenced from French

Polynesia.

Sakai and Türkay (2014) illustrated a single somewhat

distorted individual from Papua New Guinea that was

identified as Calliaxina panglaoensis. On the basis of its

recorded locality, the specimen is probably E. dworschaki.

Eucalliaxiopsis inaequimana Dworschak, 2014

Figures Ih, 1, 18, 25h

Callianassa aequimana.—Poore and Griffin, 1979: 245 (partim

from Qld).

Calliax aequimana.—Sakai,

Maldives), fig. 31.

Eucalliax inaequimana Dworschak, 2014: 236—244, figs 3—7, 9f,

g, 10e, f. — Dworschak, 2018: 17.

Eucalliaxiopsis inaequimana.—Poore et al., 2019: 125, 127, fig.

190, p.— Robles et al., 2020.

1999: 118-119 (partim from

Material examined. Papua New Guinea, New Ireland, NW corner of

Little Nusa Island, seagrass, 0-1 m, 02° 34.9' S, 150° 46.4' E

(KAVIENG 2014 stn KM11), MNHN-IU-2013-10006* (female, 5.8

mm), MNHN-IU-2013-10008* (male, 7.2 mm).

Australia. Qld. Great Barrier Reef, Myrmidon Reef, 18.27° S,

147.38° E, October 1986, 4 m (M. Riddle stn M/10/2), NMV J71689 (2

males, 7.1 mm; 5 females, 5.3—8.3 mm); 2 m (M. Riddle stn M/10/3),

NMV J71690 (female, 5.6 mm). Norwest Islet, Capricorn Group,

23.3° S, 151.7? E, AM P.10356 (male, 8.1 mm)

Mariana Islands, Guam, W coast, near Fish Eye Marine Park,

Indo-West Pacific and Australian Eucalliacidae

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Figure 18. Eucalliaxiopsis inaequimana. French Polynesia, UF 16512 (male, 5.0 mm): a, anterior carapace, eyestalks, antennule, antenna; b,

pleopod 1. French Polynesia, UF 28903 (male, 5.4 mm): c, pleopod 1. Papua New Guinea, MNHN-IU-2014-10008 (male, 7.2 mm): d-f, telson,

dorsal, lateral and anterodorsal views. Scale bars a, d-f = 1 mm; scale bars b, c = 0.1 mm.

sand/rubble flat, 0.2-0.5 m, 13.471? N, 144.704° E (stn AA05), UF

27330 (female, 3.5 mm).

French Polynesia. Moorea Island, Motu Tiahura/Fareone channel,

17.4888° S, 149.9134° W (stn MIB-225), UF 16512 (male, 5.0 mm).

Papetoai, 17.4898°S, 149.884°W, (stn BIZ-463), UF 16479 (female, 3.9

mm), UF 28784* (male, 5.1 mm); stn BIZ-493 UF 28903 (male, 5.4

mm), UF 28929 (female, 5.1 mm). S of Nihimaru, 17.535° S, 149.904° W

(stn BIZ-616), UF 29162 (ovigerous female, 6.6 mm). Off Opunohu

public beach, 17.49143° S, 149.85138° W (stn BIZ-636), UF 29208 (male,

6.0 mm). Off Nihimaru river estuary, 17.533° S, 149.9045° W (stn MIB-

098), UF 15915 (male, 5.0 mm), UF 15945 (female, 4.0 mm). Gump reef,

in front of waterfront bungalows, 17.4902° S, 149.826° W (stn MIB-

224), UF 16498 (ovigerous female, 6.2 mm). Between Papetoai and

Hotel, 17.4908° S, 149.8871° W (MIB-227), UF 16535 (male, 2.9 mm),

UF 16542 (female, 4.6 mm; male, 4.1 mm).

Diagnosis. Pleonite | without pair of sternal plates. Eyestalk

about twice as long as wide. Antennule peduncle reaching to

base of antenna peduncle article 5. Maxilliped 3 exopod absent.

Cheliped carpi distolateral margin square at lower angle,

propodi lateral face without longitudinal ridge, upper mesial

face with row of short transverse ridges associated with more

distal clusters of setae. Male pleopod 1 article | linear, with

distal setae; article 2 short, oblique, or of single article with

narrower distal lobe. Female pleopod 2, appendix interna

absent. Uropod endopod ovate. Telson without transverse row

of spiniform setae.

Colour. Exoskeleton mostly translucent white, gastric region,

distal propodus of chelipeds with few pink chromatophores

(fig. Ih, 1).

Distribution. Australia, Cocos (Keeling) Island (type locality),

Qld, Great Barrier Reef; Papua New Guinea, New Ireland;

Indonesia, Sulawesi; Philippines, Panglao; French Polynesia.

Intertidal to 4 m.

Remarks. Dworschak (2014) distinguished this species, as

Eucalliax inaequimana, on several features including

inequality of the chelipeds in both sexes from E. aequimana.

Usually, only males of Eucalliaxiopsis have a significantly

larger major cheliped although not all descriptions are clear on

this point. The same is true of E. patio (described and compared

below) and both species differ from E. aequimana in the ways

listed by Dworschak (2014).

Robles et al.’s (2020) phylogram detected no genetic

difference between two individuals from the type locality,

two from Papua New Guinea and one from French Polynesia.

Dworschak (2014) recorded the maximum carapace length as

9.1 mm; the largest in this collection were a male of 7.2 mm

and ovigerous female of 6.2 mm. His illustrations and

description were comprehensive, and many specific details

appear on the new material, such as the transverse ridges on

the mesial faces of the chelipeds and distal tubercles on the

cheliped propodus. A broad postrostral obsolete median ridge

defined by shallow longitudinal grooves is typical of the new

material (fig. 18a) but was not mentioned by Dworschak

(2014). The feature appears to be more conspicuous in some

individuals than others (Dworschak, pers. comm. 29

September 2019). The male pleopod 1 is manifest in two

forms — a long article and shorter second article (fig. 18b; as

figured by Dworschak) — and a form where these appear fused

(fig. 18c).

Poore and Griffin’s (1979) specimen of Callianassa

aequimana from Queensland was reidentified as

E. inaequimana. Sakai's (1999) record of Calliax aequimana

from the Maldives is almost certainly referable to

E. inaequimana. Sakai (2018) omitted E. inaequimana from

his revision of Eucalliacinae.

Eucalliaxiopsis madagassa (Sakai and Türkay, 2014)

Figure 19

Calliaxiopsis madagassa Sakai and Türkay, 2014: 193, fig. 13.

Calliaxina thomassini Ngoc-Ho, 2014: 549, fig. 2.

Calliaxina madagassa.—Poore and Dworschak, 2017: 120, fig. 1.

Material examined. Madagascar Nosy Bé, W of Hell-Ville,

13.41562° S, 48.246485" E (stn MGNW-50), UF 14090 (ovigerous

female, 5.8 mm). Toliara (as Tuléar), Thomassin stn 676, MNHN-

[U-2016-8085 (ovigerous female, 5.8 mm).

Diagnosis. Eyestalk about twice as long as wide. Antennule

peduncle reaching to midpoint of antenna peduncle article 5.

Maxilliped 3 exopod present. Cheliped carpi distolateral margin

square at lower angle, propodi lateral face without longitudinal

ridge, upper mesial face with row of clusters of long setae. Male

pleopod 1 article 1 without distal setae, article 2 parallel-sided,

apically rounded, with marginal setae, without appendix interna.

Female pleopod 2, appendix interna absent. Uropod endopod

ovate. Telson without transverse row of spiniform setae.

Distribution. Madagascar.

Remarks. Poore and Dworschak (2017) discussed the synonymy

and priority of the species names before Calliaxina was divided

into two by Poore et al. (2019). The species is widespread in

Madagascar (Ngoc-Ho, 2014) and is recognised by rounded

posterolateral corners and convex posterior margin of the telson,

the continuous dense row of setae on the lateral margin of the

fixed finger of the minor cheliped, and the presence (usually) of

two or three tubercles at the midpoint of the lower margin of the

cheliped merus. The ovigerous female differs from that of the

male figured by Ngoc-Ho (2014) in having a slightly narrower

uropodal exopod and in having the chelipeds subequal — they

are unequal in males of most species of this genus.

Eucalliaxiopsis panglaoensis (Dworschak, 2006)

Figure 25g

Eucalliax panglaoensis Dworschak, 2006: 349—358, figs 1—4, 6, 7

(not fig. 5 = E. dworschaki sp. nov.).— Osawa and Fujita, 2016: 40—41,

fig. 3c.—Dworschak, 2018: 17, fig. 1 (partim).

Calliaxina panglaoensis.—Sakai, 2011: 501.

Eucalliaxiopsis panglaoensis.—Poore et al., 2019: 125, 127, 146,

fig. I9m.— Robles et al., 2020.

Material examined. Paratypes. Philippines, Panglao Island, 09° 38.3' S,

123? 49.6' E, MNHN Th15093 (female, 7.1 mm), MNHN Th1504 (male,

4.3 mm).

Diagnosis. Pleonite | with pleopods attached to pair of sternal

plates. Eyestalk about twice as long as wide. Antennule peduncle

G.C.B. Poore

reaching to midpoint of antenna peduncle article 5. Maxilliped 3

exopod absent. Cheliped carpi distolateral margin with 5 acute

teeth near lower margin separated by wide grooves parallel to

lower margin, second tooth with acute ridge, propodi lateral

face without longitudinal ridge, upper mesial face with row of

clusters of long setae. Male pleopod 1 article 1 linear, with distal

setae; article 2 blade-like with subapical medial notch. Female

pleopod 2, appendix interna absent. Uropod endopod ovate with

excavate apex. Telson with row of spiniform setae on ridge.

Distribution. Philippines, Panglao; Japan, Ryukus, Miyako

Group. Sandy beaches, intertidal to 4 m.

Remarks. Eucalliaxiopsis panglaoensis differs from E. paradoxa

and E. dworschaki (see remarks under Eucalliaxiopsis above

and these two species) in armature of the cheliped. The marginal

teeth at the lower distal angle of the carpus are more widely

spaced, and the second more sharply ridged than in the other

two. The longitudinal lateral ridge on the propodus is shorter

than in E. dworschaki but more prominent than in E. paradoxa

(fig. 25g; Dworschak, 2000).

The record of this species, rather than E. dworschaki,

from Japan is based on the illustration of the cheliped with a

short lateral propodal ridge (Osawa and Fujita, 2016: fig. 3c).

Eucalliaxiopsis paradoxa sp. nov.

http://zoobank.org/urn:|sid:zoobank.org:act:2 EEAC4EC-C3EA-

4D17-9102-4090CA6DF02E

Figures 20, 21, 25h, 1

Material examined. Holotype. Australia, Qld, Great Barrier Reef, Rib

Reef, 185.48? S, 146.86" E, October 1985, shallow lagoon, 4 m, (M.

Riddle stn R/10/2), NMV J71683 (male, 9.4 mm).

Paratypes. Collected with holotype. NMV J71679 (ovigerous

female, 9.4 mm), NMV J71684 (3 females, 9.8—10.9 mm). Australia,

Old, Great Barrier Reef, Myrmidon Reef, 18.27? S, 147.38" E, October

1985, lagoon with “Callianassa” mounds, 8 m, (M. Riddle stn M/10/1),

NMV J71680 (7 males, 3.9-10.3 mm; 7 females, 4.4—10.4 mm).

Diagnosis. Pleonite 1 with pleopods attached to pair of sternal

plates. Eyestalk about twice as long as wide. Antennule peduncle

reaching to midpoint of antenna peduncle article 5. Maxilliped 3

exopod absent. Cheliped carpi distolateral margin with 5 acute

teeth near lower margin separated by submarginal oblique

grooves, second tooth with prominent rounded ridge, propodi

lateral face with short horizontal longitudinal ridge extending

from beyond mid-length of palm to base of finger, upper mesial

face with row of clusters of long setae. Male pleopod | article |

linear, with distal setae; article 2 blade-like with subapical

medial notch. Female pleopod 2, appendix interna absent.

Uropod endopod ovate with excavate apex. Telson with row of

spiniform setae on ridge.

Description of holotype male. Carapace length 9.4 mm,

smooth; gastric-abdominal regions together 4.7 times as long

as wide; branchiostegite fully calcified; cervical groove at 0.63

carapace length, scarcely obvious on branchiostegite; cardiac

sulcus at 0.8 carapace length, not meeting mid-dorsally,

extending transversely across quarter of branchiostegite.

Rostrum acute, tapering evenly over most of length, length

0.5 width of eyestalks. Anterolateral carapace lobe 0.6 length

Indo-West Pacific and Australian Eucalliacidae 29

|: 5. PE

— RII

^ M

\ LAS : (us $E

Figure 19. Eucalliaxiopsis madagassa. Madagascar, UF 14090 (ovigerous female, 5.8 mm): a, b, anterior carapace, eyestalks, antennule, antenna,

dorsal and lateral views; c—e, telson, dorsal, posterior and lateral views; f, major (left) cheliped, mesial view; g, major (left) cheliped, propodus,

dactylus, lateral view; h, minor (right) cheliped, mesial view; 1, minor (right) cheliped, propodus, dactylus, lateral view. Scale bars = 1 mm.

30 G.C.B. Poore

PAS :

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Figure 20. Eucalliaxiopsis paradoxa sp. nov. Holotype, NMV J71683 (male, 9.4 mm): a, b, carapace, dorsal and lateral views; c, d, anterior

carapace, eyestalks, antennule, antenna, dorsal and lateral views; e, maxilliped 3; f, g, major cheliped (left), mesial and lateral views, with detail

of distolateral corner of carpus; h, 1, minor cheliped (right), mesial and lateral views, with detail of distolateral corner of carpus; J, k, m, pereopods

2, 3, 5. Paratype, NMV J71680 (female, 9.3 mm): 1, pereopod 4. Scale bars = 1 mm.

Indo-West Pacific and Australian Eucalliacidae 31

EAR? PINTOR

ITA

p q

Figure 21. Eucalliaxiopsis paradoxa sp. nov. Holotype, NMV J71683 (male, 9.4 mm): a, thoracic sternite 7; b, pleonitel sternite, right pleopod 1

(anterior uppermost); c, pleonite 6, telson, lateral view; d, pleonite 6, left lateral oblique view; e, right uropod; f, telson; g, h, pleopod 1, mesial

and posterior views; 1, pleopod 2, posterior view. Paratype, NMV J71679 (ovigerous female, 9.4 mm): j, k, anterior carapace, eyestalks, antenna,

dorsal and lateral views; l, m, major cheliped (left), mesial and lateral views; n, o, minor cheliped (right), mesial and lateral views; p, q, pleopods

1, 2. Paratype, NMV J71684 (female, 10.4 mm): r, minor cheliped carpus, distolateral margin. Scale bars = 1 mm.

rostrum, with rounded apex, anteriorly directed, with

numerous submarginal dorsal setae.

Thoracic sternite 7 1.3 times as long as wide, with well-

defined median sulcus over posterior half, lateral groove

anterior to coxal articulation shallow, oblique, meeting

medially, with 7 posterolateral clusters of 2 or 3 long setae.

Pleonal sternite 1with pair of swollen relatively unchitinised

areas anterior to ridge bearing pleopods. Pleonite 6 with small

lateral blunt hook-like process at midpoint.

Eyestalk 2.2 times as long as wide at base, tapering to

cornea and acute apical lobe; cornea subcircular; apical lobe

extended dorsally. Antennular peduncle 3 times as long as

width of both eyestalks; article 2 3.5 times as long as wide;

article 3 half as long as article 2; articles 2 and 3 with

ventrolateral row of long setae, continued onto flagellum.

Antennal peduncle 3.4 times as long as width of both

eyestalks, overreaching antennule peduncle by half of article

5; scaphocerite wider than long, with acute apex; article 4c. 5

times as long as wide; article 5 0.6 length of article 4.

Maxilliped 3 ischium with slightly convex mesial margin,

expanding from narrow proximomesial corner; ischium-

merus upper margin 1.7 times as long as greatest width; crista

dentata of 12 spines, the most proximal reflexed; carpus—

dactylus together almost as long as ischium—merus.

Chelipeds unequal, major propodus 1.2 times as long as

that of minor, both of similar widths, major dactylus stouter

than that of minor. Major cheliped (right 1n holotype) ischium

twice as long as distal width, lower margin with short spines

becoming more distinct distally; merus twice as long as

broad, lower margin mostly straight, unarmed, upper margin

with distal tooth; carpus 1.2 times as wide as upper length,

upper margin carinate, unarmed except for distal tooth,

distomesial margin with prominent lobe near lower margin,

distolateral margin with acute tooth near upper margin, with 3

acute teeth near lower margin separated by submarginal

oblique grooves, second tooth with prominent rounded ridge,

3 short setae in lower groove; propodus widest proximally,

tapering, upper palm length barely exceeding greatest width,

upper margin straight, carinate, with rounded lobe not

projecting over dactylus, lower margin carinate, strongly

curved mesially, mesial face with short distal uneven setose

ridge below upper margin, distomesial margin of palm almost

straight, lateral face of palm with short horizontal ridge, about

fifth length of palm, extending on to finger, 4 rimmed

tubercles near proximal margin, | at midpoint of palm, | near

lower margin, each perforated for cluster of setae; fixed finger

half length of upper margin of palm, cutting edge with

denticles over proximal half, blunt triangular tooth near

midpoint; dactylus as long as fixed finger, twice as long as

wide at base, cutting edge with blunt proximal molar, with

straight tip; submarginal tufts of long setae on upper and

lower mesial margins of carpus and propodus, opposing

mesial margins of fingers.

Minor cheliped (left 1n holotype) ischium twice as long as

distal width, lower margin with distal spines; merus twice as

long as broad, lower margin mostly straight, unarmed; carpus

1.2 times as wide as upper length, upper margin carinate,

unarmed except for distal tooth, distomesial face with lower

G.C.B. Poore

submarginal ridge, distolateral margin with acute tooth near

upper margin, with 3 teeth near lower margin separated by

submarginal oblique grooves, second tooth acute with

prominent rounded ridge, 2 short setae in lower groove;

propodus greatest width proximally, tapering, upper palm

length equal to greatest width, palm tapering, upper margin

carinate, overhanging mesial face, with rounded lobe projecting

over dactylus, lower margin carinate, strongly curved mesially,

mesial face with 4 blunt teeth at associated with upper row of

clusters of setae, distomesial margin of palm almost straight,

lateral face of palm with prominent ridge running from two-

thirds length of palm to middle of finger, 5 rimmed tubercles

near proximal margin of which | near midpoint of palm is

more distal, each perforated for cluster of setae; fixed finger 0.7

length of upper margin of palm, width at base half width of

palm, cutting edge smooth; dactylus as long as fixed finger, 3.2

times as long as wide at base, cutting edge straight; submarginal

tufts of long setae on upper and lower mesial margins of carpus

and propodus, opposing mesial margins of fingers.

Pereopod 2 merus twice as long as maximum width;

dactylus c. 3 times as long as upper margin of propodus.

Pereopod 3 merus twice as long as maximum width; carpus

1.8 times as long as wide; propodus with lower margin

concave, twice as long overall as mid-length; dactylus 0.7

times length of axial length of propodus. Pereopod 4 merus

length 2.2 times maximum width; carpus 2.5 times as long as

wide; propodus as long as wide, typically setose, with | long

distal seta overlapping dactylus.

Pleopod 1 article 1 distally setose; article 2 as long as

article 1, 4 times as long as wide, with apex curved, appendix

interna lobe-like, not produced, unarmed. Pleopod 2 endopod

twice as long as wide; appendix masculina overreaching

endopod by about third its length, triangular 1n cross-section,

setose along posterior face and distally; appendix interna near

base of appendix masculina without hooks.

Uropodal endopod anterior margin convex, posterior

margin convex, widest about third way along, 1.4 times as

long as wide, apex excavate, with rows of long setae at ends of

anterior and posterior margins, with 8 short setae between in

excavation, with subdistal group of facial setae; exopod ovate,

anterior margin straight, distal and posterior margins

continuous, greatest dimension 2.2 times anterior margin,

with row of blade-like distal setae on distal margin, c. 6 not

overlapping with densely setose distal margin; dorsal plate

extending almost half of exopod width, distal margin with

spiniform setae. Telson 1.6 times as wide as long, broadest at

midpoint, posterolateral corner rounded; posterior margin

sinuous, lateral regions most prominent, with posterolateral

cluster of long setae plus 2 short spiniform setae; dorsal

surface with sharp transverse ridge at anterior third, with 36

short spiniform setae, few long setae at midpoint

Female. Essentially as male except: major cheliped palm

tapering, upper length 0.9 greatest width, upper margin convex,

depressed distally, lateral ridge longer than in male, fixed finger

broader than in male, crenellate proximal and distal to tooth.

Pleonal sternite |with pair of oval plates supporting pleopods,

each with longitudinal row of setae. Pleopod 1 peduncle with

Indo-West Pacific and Australian Eucalliacidae

mesial setose lobe longer than base. Pleopod 2 peduncle with

similar mesial setose lobe; endopod without appendix interna.

Etymology. Paradoxa (Latin), reflecting the subtle differences

from E. panglaoensis and co-occurring E. dworschaki, and a

tribute to the collector’s name, Martin Riddle.

Distribution. Australia, Qld, central Great Barrier Reef (18° S).

Subtidal coral reef lagoons.

Remarks. Eucalliaxiopsis paradoxa shares several features

with E. panglaoensis and E. dworschaki (see remarks under

Eucalliaxiopsis above).The new species differs from these two

in having eight short setae in the uropodal endopod excavation

(fig. 21e; 11 in the other species), a short longitudinal ridge at

the base of the finger on the lateral face of the chelipeds, the

propodus of the male cheliped tapering (more rectangular in

the others), and the middle distolateral cheliped carpal tooth

being oblique and sharply ridged (figs 20g, 1, 21m, o, r, 25h). In

E. dworschaki, the ridge on the palm extends almost its entire

length, and the middle carpal lobe is perpendicular and with a

median groove. Besides having fewer short setae at the end of

the uropodal endopod, Eucalliaxiopsis paradoxa differs from

E.panglaoensis in the upper margin of the male major cheliped

palm being as long as wide (vs 1.3 times as long as palm width),

the palm of the female major cheliped being relatively longer

(1.1 vs 0.9 times as long as greatest width), the uropodal

endopod being more tapered (more oval in E. panglaoensis)

and the sinuous posterior margin of the telson (vs straight).

Eucalliaxiopsis patio sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: F5884500-4430-

493B-9BCD-CA5E769F846E

Figures 22—24

Eucalliaxiopsis PNG-1153C.— Robles et al., 2020.

Material examined. Holotype. Papua New Guinea, New Ireland

Province, Patio Island, 02? 36.2' S, 150? 31.6' E, 6—8 m, coral rubble

(KAVIENG 2014 stn KB38), MNHN-IU-2014-2536* (female, cl. 4.6

mm).

Paratype. Collected with holotype (MNHN-IU-2014-2539 (male,

cl. 4.3 mm, with male and female bopyrid isopods under left

branchiostegite).

Diagnosis. Pleonite I without pair of sternal plates. Eyestalk 1.5

times as long as wide. Antennule peduncle reaching to midpoint

of antenna peduncle article 5. Maxilliped 3 exopod absent.

Cheliped carpi distolateral margin square at lower angle, propodi

lateral face without longitudinal ridge, upper mesial face with

row of short transverse ridges associated with more distal clusters

of setae. Male pleopod | of single article with narrower distal

lobe. Female pleopod 2, appendix interna present. Uropod

endopod ovate. Telson without transverse row of spiniform setae.

Description of holotype female. Carapace length 4.6 mm,

smooth; gastric-abdominal regions together 4.2 times as long

as wide; branchiostegite fully calcified; cervical groove at 0.67

carapace length, scarcely obvious on branchiostegite; cardiac

sulcus at 0.83 carapace length, not meeting mid-dorsally,

extending transversely across two-thirds of branchiostegite.

Rostrum broadly rounded-triangular, length about 0.2

width of eyestalks. Anterolateral carapace lobe as long as

rostrum, with rounded apex, depressed anteriorly, with c. 5

submarginal dorsal setae. Pleonites 1—6 lengths relative to cl:

0.46 : 0.67 : 0.45 : 0.33 : 0.41: 0.51; pleonite 1 with strong dorsal

transverse groove, without pair of sternal plates; pleonites 3—5

with lateral tufts of setae; pleonite 6 with 8 lateral rows of setae.

Thoracic sternite 7 1.2 times as long as wide, with well-

defined median sulcus over posterior half, smooth over medial

half, lateral ridge anterior to coxal articulation crossing pit at

quarter of width, meeting medially, with 3 posterolateral

clusters of long setae.

Eyestalk 1.7 times as long as wide at base, swollen laterally,

tapering to cornea and acute apical tubercle; cornea wider than

long, depressed anteriorly. Antennular peduncle twice as long

as width of both eyestalks; article 2 twice as long as wide;

article 3 as long as article 2; articles 2 and 3 with ventrolateral

row of long setae, continued onto flagellum. Antennal peduncle

2.4 times as long as width of both eyestalks, overreaching

antennule peduncle by half of article 5; scaphocerite ovoid,

longer than wide; article 4 c. 5 times as long as wide; article 5

0.8 length of article 4. Maxilliped 3 ischium with slightly

convex mesial margin, expanding from narrow proximomesial

corner; ischium—merus upper margin twice as long as greatest

width; crista dentata of 6 spines, the 2 most proximal largest;

carpus—dactylus together almost as long as ischium—merus.

Chelipeds unequal, major carpus-propodus upper margin

1.1 length of minor, propodus 1.2 times as wide as minor.

Major cheliped (left in holotype) ischium twice as long as

distal width, lower margin with needle-like spines becoming

more distinct distally; merus 1.7 times as long as broad, lower

margin convex, unarmed; carpus 1.2 times as wide as upper

length, margins carinate; propodus greatest width in middle of

palm, upper palm length 1.15 times greatest width, distomesial

margin of palm with submarginal group of 2 setae, small

tubercle, distolateral margin of palm with submarginal group

of c. 6 setae between fingers; fixed finger 0.4 length of upper

margin of palm, cutting edge with miucrodenticles over

proximal half, blunt triangular tooth at about midpoint;

dactylus as long as fixed finger, twice as long as wide at base,

cutting edge with deflected tip; submarginal tufts of long setae

on upper and lower mesial margins of carpus and propodus,

Opposing mesial margins of fingers; 3 short transverse ridges

associated with setae on submesial upper margin.

Minor cheliped (right in holotype) ischium c. 2.3 times as

long as distal width, lower margin with needle-like spines

becoming more distinct distally; merus 1.7 times as long as

broad, lower margin convex, unarmed; carpus 1.2 times as wide

as upper length, margins carinate; propodus palm more tapering

than major, 0.95 times as wide as upper palm length, distomesial

margin of palm with submarginal group of 4 12 setae,

distolateral margin of palm with submarginal group of c. 4

setae; fixed finger half length of upper margin of palm, mesial

cutting edge straight, smooth, cutting edge straight, fixed finger

with smooth lateral ridge defining edge of lateral concavity at

base of fixed finger, without granules; dactylus as long as fixed

finger, 3 times as long as wide at base, cutting edge straight;

submarginal tufts of long setae on upper and lower mesial

34 G.C.B. Poore

Figure 22. Eucalliaxiopsis patio sp. nov. Holotype, MNHN-IU-2013-2536 (female, 4.1 mm): a, habitus, dorsal; b, carapace, lateral view; c,

thoracic sternite 7; d, pleonitel sternite, right pleopod 1; e, f, anterior carapace, eyestalks, antennule, antenna, dorsal and lateral views; g, pleonite

6, telson, uropod; h, uropod exopod, margin, dorsal plate; 1, J, telson, lateral and dorso-anterior views; k, maxilliped 3; 1 , maxilliped 3, crista

dentata, basis, ischium. Scale bars = 1 mm.

Indo-West Pacific and Australian Eucalliacidae

a-h

Figure 23. Eucalliaxiopsis patio sp. nov. Holotype, MNHN-IU-2013-2536 (female, 4.1 mm): a, major cheliped (left), mesial; b, major cheliped

(left), distal propodus, dactylus, lateral; c, minor cheliped (right), mesial view; d, minor cheliped (right), distal propodus, dactylus, lateral; e—h,

pereopods 2-5; 1, J, pleopods 1, 2. Scale bars = 1 mm.

margins of carpus and propodus, opposing mesial margins of

fingers; 2 short transverse ridges associated with setae on

submesial upper margin, more prominent distally.

Pereopod 2 merus 1.8 times as long as maximum width;

dactylus c. 3 times as long as upper margin of propodus.

Pereopod 3 merus 2.2 times as long as maximum width;

carpus 1.8 times as long as wide; propodus with lower margin

almost straight, 1.8 times as long as mid-length; dactylus half

as long as axial length of propodus. Pereopod 4 merus twice

as long as maximum width; carpus 3 times as long as wide;

propodus typically setose, with distal spiniform setae among

setose margin, with 2 long distal setae overlapping dactylus.

Pleopod | article 1 without distal projection beyond article

2; article 2 longer than article 1. Pleopod 2 endopod 3 times as

long as wide; appendix interna at midpoint of endopod.

Uropodal endopod ovoid, anterior margin more convex

WEN SS

G.C.B. Poore

Figure 24. Eucalliaxiopsis patio sp. nov. Paratype, MNHN-IU-2013-2539 (male, 4.6 mm): a, b, anterior carapace, eyestalks, dorsal and lateral

views; c, major cheliped (right), mesial; d, major cheliped (right), propodus, dactylus, upper; e, minor cheliped (left), mesial view; f, minor

cheliped (left), propodus, dactylus, upper; g, h, pleopods 1, 2. Scale bars = 1 mm.

than posterior margin, widest about third way along, 1.5 times

as long as wide, with subdistal group of facial setae; exopod

ovate, anterior margin straight, distal and posterior margins

continuous, greatest dimension 2.2 times anterior margin, with

row of blade-like distal setae on distal margin, c. 6 not

overlapping with densely setose distal margin; dorsal plate

extending almost half of exopod width, distal margin with

spiniform setae. Telson 1.6 times as wide as long, broadest over

anterior half, posterolateral corner evenly rounded to posterior

margin; posterior margin barely convex, with posterolateral

clusters of long setae; dorsal surface with obscure rounded

transverse ridge at anterior third, without median notch.

Paratype male. Carapace length 4.3 mm. Eyestalks 1.5 times

as long as wide at base, swollen laterally, tapering to cornea

and acute apical tubercle; cornea wider than long, depressed

anteriorly Postrostral dorsal area with pair of shallow pits.

Chelipeds unequal, major carpus-propodus upper margin

1.1 length of minor, propodus 1.3 times as wide as minor.

Major cheliped (right) ischium twice as long as distal width,

lower margin with denticles, | distal spine; merus 1.7 times as

long as broad, lower margin convex, unarmed; carpus 1.4

times as wide as upper length, margins carinate; propodus

ereatest width in middle of palm, upper palm length 1.15

times greatest width, distomesial margin of palm with

submarginal group of 2 setae plus tubercle; fixed finger 0.35

length of upper margin of palm, cutting edge with

microdenticles over proximal half, blunt triangular tooth near

midpoint; dactylus as long as fixed finger, twice as long as

wide at base, cutting edge irregular, with deflected tip;

submarginal tufts of long setae on upper and lower mesial

margins of carpus and propodus, opposing mesial margins of

fingers; 4 short transverse ridges associated with setae on

submesial upper margin.

Minor cheliped (left) ischium c. 2.3 times as long as distal

width, lower margin with denticles, | distal spine; merus 1.8

times as long as broad, lower margin convex, unarmed; carpus

1.1 times as wide as upper length, margins carinate; propodus

palm more tapering than major, 0.95 times as wide as upper

palm length, distomesial margin of palm with 2 submarginal

Indo-West Pacific and Australian Eucalliacidae 37

Figure 25. Eucalliaxiopsis spp. E. aequimana: a, b, major cheliped, mesial. E. dworschaki, Papua New Guinea, MNHN-IU-2013-10003: c, d,

major cheliped lateral, upper margin. E. dworschaki, Philippines, NHMW 20923: e, f, major cheliped carpus lower lateral angle, upper margin.

E. panglaoensis, NHMW 20922: g, minor cheliped, lateral. E. inaequimana: h, major cheliped, mesial. E. paradoxa: h, 1, major cheliped lateral,

mesial. Photos e-g: P.C. Dworschak. Various scales.

eroups of 2—3 setae; fixed finger 0.4 length of upper margin of

palm, mesial cutting edge straight, smooth; dactylus as long as

fixed finger, 2.8 times as long as wide at base, cutting edge

straight; submarginal tufts of long setae on upper and lower

mesial margins of carpus and propodus, opposing mesial

margins of fingers; 3 short transverse ridges associated with

setae on submesial upper margin, more prominent distally.

Pleopod 1 articles indistinguishable; article 1 with 1 short, 4

long distal setae; article 2 a short triangle, half as wide as article

|. Pleopod 2 endopod 2.7 times as long as wide; appendix

masculina attached at mid-length, 5 times as long as width,

distal margin rounded, exceeding endopod by fifth of its length,

with c. 9 facial setae near midpoint, c. 14 subdistal setae;

appendix interna rod-like, about 0.4 length of appendix interna.

Etymology. From Patio Island, Papua New Guinea, the type

locality (noun in apposition).

Distribution. Papua New Guinea. Coral rubble, 6—8 m.

Remarks. The holotype was sister taxon to five individuals of

E. inaequimana on the phylogenetic tree of Robles et al. (2020).

Both species have submarginal mesial transverse ridges on the

cheliped propodi from which tufts of long setae emerge; in other

species of Eucalliaxiopsis the same tufts are not supported in

this way. The male pleopod 1 of both species has a simple

second article, free or fused, a form seen too in E. madagassa

(see Poore and Dworschak, 2017: fig. IC, D), Eucalliax

quadracuta and Calliaxina bulimba (see above). The typical

article 2 of species of Eucalliaxiopsis 1s flattened with a distal

oblique notch. — Eucalliaxiopsis patio differs from

E. inaequimana in shorter, more swollen eyestalks (1.5 vs 1.8

times as long as wide), slightly shorter rostrum (quarter vs fifth

length of eyestalk), shorter antenna peduncle (2.4 vs 3.0 times as

long as base of both eyes), more compact antennular and

antennal peduncular articles, the telsonic ridge being only

slightly elevated and complete (vs clearly elevated, with median

notch), and a broader uropodal endopod (1.5 vs 1.8 times as long

as wide). The appendix interna on the female pleopod 2 is

present while it 1s absent in E. inaequimana. It 1s possible that

this difference may be related to the female's small size;

Dworschak (2006) noted that the smallest female of

E.panglaoensis possessed an appendix interna while larger ones

did not. Two individuals were collected from the type locality,

with carapace lengths of 4.3 and 4.6 mm. The largest known

specimen of E. inaequimana has a carapace length of 9.1 mm.

Identification of these specimens using Sakai's (2011) key to

species of Calliaxina leads to C. aequimana, a southern

Australian species. This species, now Eucalliaxiopsis

aequimana, lacks mesial ridges on the chelipeds and has a

female-like male pleopod 1.

Pseudocalliax Sakai, 2011

Pseudocalliax Sakai, 2011: 505—506.— Sakai, 2018: 744.— Poore

et al., 2019: 128, 146, figs 18a, k, 19e, r.

Remarks. The type and only species was placed in Eucalliacinae,

now Eucalliacidae, as a species of Calliax, by Sakai (2005).

Sakai (2011) erected the new genus Pseudocalliax for it in the

G.C.B. Poore

same subfamily. No molecular sequences were available to

Robles et al. (2020) to challenge this placement that has good

morphological support (Poore et al., 2019). The genus shares

with the three species of Calliax the characteristic short fixed

finger on the minor cheliped.

Pseudocalliax tooradin (Poore and Griffin, 1979)

Figure 26

Callianassa tooradin Poore and Griffin, 1979: 275—277, fig. 36.

Calliax tooradin.—Sakai, 1988: 61.— Davie, 2002: 459.— Sakai,

2005: 204.

Paraglypturus tooradin.— Sakai, 1999: 124, fig. 33a—c.—Poore,

2004: 184, fig. 50h, 1.

Pseudocalliax tooradin.— Sakai, 2011: 506.

Material examined. Australia, Vic., Western Port, Crib Point,

38° 20.23' S, 145° 13.38' E, 5 m (CPBS stn LIN), NMV J301 (holotype

female, 5.2 mm), NMV J302 (2 paratype females); 38° 21.17’ S,

145° 13.15' E,2 m (CPBS stn 000), NMV J303 (paratype male, 6.5 mm).

San Remo, channel edge, E. of No. 7 beacon, 38° 32'S, 145° 23' E(MRG

Transect 1), NMV J59760 (male, 5.1 mm); N from 200 m E of Back

Beach Rd to channel edge, 38° 32'S, 145° 23' E (MRG transect 2), NMV

J59761 (2 males, 4.2, 6.0 mm). Swan Bay, Edwards Point, 38° 13.3' 5,

144° 41.4' E, 2 m, NMV J16722 (juvenile, 1.5 mm).

Distribution. Australia, Vic., entrances of Western Port and

Port Phillip Bay. Subtidal, c. 2 m.

Remarks. Poore and Griffin (1979) provided only a short

description and simple figures. Here, the antennae, chelipeds,

terminal articles of pereopods 3 and 4, male pleopods, telson

and uropod are figured. The species is notable for the setose

thoracic sternite 7 and ventroposterior surface of the coxa—

ischium of pereopod 5, a feature diagnostic for the genus (fig.

26b). The antenna bears a free scaphocerite, longer than

broad, with a rounded apex (fig. 26a). The exopod of

maxilliped 3 typically reaches the middle of the merus but is

shorter on smaller 1ndividuals. While the fixed finger of the

minor cheliped is shorter than the dactylus in most individuals

it is almost as long 1n one male. The propodus of pereopod 3

is suboval, little longer along its upper margin than its greatest

width, and densely setose on its lateral face. The propodus of

pereopod 4 1s elongate-oval, 1.5 times as long along its upper

margin than its greatest width, with a distal long spiniform

setae on the lower margin, and densely setose on its lateral

face. The male pleopod 1, overlooked by Poore and Griffin

(1979), has a shorter second article with an apical notch (fig.

26g). The peduncle of the male pleopod 2 is longer than

broad; the appendix interna is short and lacks hooks; no

appendix masculina is present (fig. 26h). The appendix interna

on the endopod of pleopod 3 1s triangular, longer than broad,

emerging by about half its length (fig. 26]). The uropodal

endopod is oval, 1.8 times as long as wide, with marginal

setae; the exopod is 1.6 times as long as wide, with short

spiniform setae along its apical margin, with a well-defined

dorsal plate with c. 6 short spiniform setae and many longer

setae (fig. 26f). A dorsal oval, cardiac prominence and cardiac

sulcus are all absent; all known material is in NMV (ct.

contradictory statements and figures in Sakai, 1999, and

Sakai, 2011).

Indo-West Pacific and Australian Eucalliacidae 39

LAE

| 7 TIL X

N M

N S

Pres | N

| i

a-f

Figure 26. Pseudocalliax tooradin Poore and Griffin, 1979. Australia, Vic., Western Port, NMV J303 (paratype male, 6.5 mm): a, anterior

carapace, eyestalks, antennule, antenna; b, posterior carapace, pleonite 1, coxa 4, thoracic sternite 7, coxa-basis 5, pleopod 1; c, maxilliped 3, with

crista dentata; d, e, propodus, dactylus, pereopods 3, 4; f, telson, uropod; g, pleopod 1; h, pleopod 2; j, appendix interna, pleopod 3. NMV J301

(holotype female, 5.2 mm): 1, pleopod 2. Scale bars = 1 mm.

Acknowledgements

I thank Philippe Bouchet for his organisation of the Kavieng

Lagoon Biodiversity Survey (Principal Investigators: Philippe

Bouchet, Jeff Kinch), part of the of La Planéte Revisitée

expeditions organised jointly by Muséum national d'Histoire

naturelle, Pro-Natura International and Institut de Recherche

pour le Développement, with support from Papua New

Guinea's National Fisheries Authority. The organisers

acknowledge supporting funding from the Total Foundation,

the Laboratoire d'Excellence Diversités Biologiques et

Culturelles (LabEx BCDiv), the Laboratoire d'Excellence

Diversités Biologiques et Culturelles (LabEx BCDiv, ANR-10-

LABX-0003-BCDiv), the Programme Investissement d'Avenir

(ANR-II-IDEX-0004-02), the Fonds Pacifique and CNRS

Institut Ecologie et Environnement. The expedition was

endorsed by the New Ireland Provincial Administration and

operated under a Memorandum of Understanding with the

University of Papua New Guinea. I am grateful to Laure

Corbari, Paula Lefevre-Martin and Anouchka Krygelmans at

MNHN and Zdeněk Duri&, University of Ostrava, Czech

Republic for help in making the collections available. I

acknowledge all the divers who collected samples, the

expeditioners who sorted them in the field.

I especially thank Peter Dworschak for checking and

photographing NHMW . material of X Eucalliaxiopsis

panglaoensis and E. dworschaki and for valuable comments

on the manuscript. I thank too Gustav Paulay, Amanda Bevis

and Arthur Anker for the loan of collections from the Florida

Museum of Natural History, University of Florida, so too

stephen Keable, Australian Museum, and Andrew Hosie,

Western Australian Museum. I am grateful to Martin Riddle,

Australian Institute of Marine Sciences, Townsville, for the

gift of eucalliacid specimens from the Great Barrier Reef.

Arthur Anker and Zdeněk Ďuriš kindly allowed me to publish

their colour photographs taken in the field.

This study was supported by the National Taxonomy

Research Grant Program (Grant numbers CN216-14 and

CBG18-06) from the Australian Biological Resources Study

and the MNHN Crosnier Fund in Paris.

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Memoirs of Museum Victoria 80: 43-58 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.03

Initiation and early development of the postcanine deciduous dentition in the

dasyurid marsupial Dasyurus viverrinus

WINTER PATRICK LUCKETT!**, NANCY HONG LUCKETT?**, AND TONY HARPER?*

Abstract

! Center for Functional Anatomy and Evolution, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

? Caribbean Primate Research Center Museum, University of Puerto Rico, Medical Sciences Campus, San Juan, Puerto

Rico, USA

? DeBusk School of Osteopathic Medicine, Lincoln Memorial University, Knoxville, Tennessee, USA.

* To whom correspondence should be addressed. Email: anthony.harper@|munet.edu

** Deceased

Luckett, W.P., Hong Luckett, N., and Harper, T. 2021. Initiation and early development of the postcanine deciduous

dentition in the dasyurid marsupial Dasyurus viverrinus. Memoirs of Museum Victoria 80: 43—58.

There has been disagreement for more than 100 years concerning the presence or absence of deciduous and

successional third premolars in the dentition of the Australian dasyurid marsupial Dasyurus viverrinus as well as 1n other

species of the genus Dasyurus. Most authors during the 19th and 20th centuries have considered the missing premolar in

Dasyurus to be the third premolar family in both jaws, in part because of the reduction in size of the third deciduous

premolar in many other genera of dasyurids. While other authors, found "the deciduous premolar to be constantly present

in the young mammary foetus as a small and precociously calcified vestigial tooth". However, the fate of this deciduous

tooth, and its possible successor in later developmental stages, was never described; and the vestigial dp3 and its possible

successor were assumed to undergo absorption during foetal life. With access to later developmental stages of Dasyurus

viverrinus, via the extensive collections of Professor J.P. Hill, we present new evidence for the later development of dP3

and P3 in both jaws, the probable eruption of the successional P3 in both jaws and evidence that dP2 1s the missing tooth

in both jaws of this dasyurid species.

Keywords

Introduction

During the years 1895-1905, when he was a Demonstrator of

Biology at the University of Sydney, Australia, Professor James

P. Hill collected a large sample of juveniles of the dasyurid

marsupial Dasyurus viverrinus, ranging from newborn young

to individuals about four months old. The species was still

relatively common in New South Wales at the time and the

animals lived relatively well in captivity. Many of these

captive-bred dasyurids produced the dated pouch young that

are the subject of this paper. Although Hill and colleagues

published several papers on the early development, placentation

and reproduction in general of Dasyurus, only one paper dealt

specifically with the development of the pouch young.

An extensive series on the development of the growth stages

in the pouch young of Dasyurus viverrinus was examined in a

monograph by Hill and Osman Hill (1955). The authors

presented 16 figures of the pouch young, and they described in

detail the external morphology of the developing young in a

series of stages labelled A to P. These ranged from unattached

newborn young outside and inside the pouch, measuring 5.5—

Dasyurus, deciduous dentition, dental development, dental homologies, pouch young

6.0 mm greatest length (GL) and 2.5 mm head length (HL;

stage A), up to adolescent young that were four months old

(stage P). These oldest young were freely moving about outside

the pouch and measured about 60 mm HL. However, no

description or figures of the microscopic anatomy of the

developing teeth were presented. Unfortunately, Hill passed

away in 1954 during the preparation of their manuscript. We

suspect that Hill would have published a detailed description of

the developing dentition if he had lived longer. After his death,

at least a portion of his collections of mammalian embryos and

young animals was donated to the Hubrecht Laboratory of

Comparative Embryology, Utrecht, Netherlands. These

included a series of histological sections of some of the

Dasyurus viverrinus pouch young described by Hill and

Osman Hill (1955) and some older juveniles preserved as

alcohol specimens. A few of the alcohol specimens from Hill's

collection were borrowed and sectioned by Professor Louis

Bolk, University of Amsterdam, Netherlands, and then returned

to the Hubrecht Laboratory of Comparative Embryology.

Materials and methods

It 1s rare to be able to histologically examine closely related

stages of early developing pouch young for any marsupial,

and we were able to study three unattached intrauterine

embryos and 22 early developing stages of pouch young

attached to the nipple for Dasyurus viverrinus from the Hill

collection. The pouch young examined range in age from

about 30 hours old, measuring about 6 mm GL, to 40—41 days

old, and measuring about 29 mm GL. Given that this species

appears to be a useful model for the developmental

relationships of the dentition 1n the extinct dasyuromorphian

Thylacinus cynocephalus (see Luckett et al., 2019), along with

other marsupicarnivore taxa, we present this overview of

early dental development in Dasyurus viverrinus. These

studies were carried out over a period of several years at the

Hubrecht Laboratory of Comparative Embryology before the

embryological collections were transferred to their present

home in the Museum für Naturkunde, Berlin, Germany.

It should be noted that Dasyurus viverrinus 1s probably

extinct on mainland Australia, with a last sighting on January

1963 in the suburbs of Sydney (Dickman et al., 2001). The

species persists in Tasmania , although it 1s vulnerable there

for several reasons, including climate change and disease,

among other possibilities (Fancourt, 2016).

A brief overview of our findings on dental development in

Dasyurus was presented previously (Luckett, 1989, 1993). We

have made new drawings of four of the developing pouch

young (fig. 1) examined by us from the Hill collection, ranging

from an early attaching pouch young of about 30 hours old to

an embryo of about 41 days old, measuring about 29 mm GL

and 18 mm HL. Most genera of dasyurids, peramelids and

didelphids have three premolars in both jaws of adults. This is

presumably the primitive condition for marsupials (Cifelli et

al., 1996). Emphasis in our study is given to the evidence for

loss of dP2 in this dasyurid with two premolars, and the

accelerated development of dP3 and P3, when compared with

these features in dasyurids with three premolars. A summary

of the major features of early development for the deciduous

and successional dentition in marsupials is presented ın fig. 2.

Results

Early development of the dental lamina

Hill and Osman Hill (1955) noted that the earliest date of

gestation was 16 days in their sample of Dasyurus viverrinus

embryos. The earliest evidence of a dental lamina that we

found was detected in the maxilla of three intrauterine

embryonic stages of 5.5—6 mm GL. In the youngest of these

embryos, the dental lamina was detected bilaterally in the

upper jaw as a localised thickening of the oral epithelium that

was associated with a very early differentiation of bone (or

prebone) in the distal portion of the developing maxilla. In

contrast, the dental lamina and developing bone were not yet

detected in the premaxillary region of the upper jaw. The

more mature intrauterine young showed dental lamina

thickenings in both the premaxilla and maxilla regions of the

upper jaw. The dental lamina was not yet evident 1n the lower

W.P. Luckett, N. Hong Luckett & T. Harper

Jaw of the youngest intrauterine embryo. As explained by Hill

and Osman Hill (1955), the newborn young of Dasyurus

viverrinus are able to find their way to the mother’s pouch,

unaided by the mother and helped by their own well-developed

forelimbs that have very sharp deciduous claws.

In the youngest pouch young attached to the mother’s

nipple, examined at about 30 hours old and measuring 6 mm

GL, a continuous dental lamina was present bilaterally in both

the upper and lower jaws. This specimen was illustrated as

fig. 5 by Hill and Osman Hill (1955) and we have redrawn this

important specimen in our fig. la. The HL of this pouch

young was 3.25 mm. It should be noted here that all of our

measurements from the Hill collection are taken from

specimens that had been placed in spirit fixation by Hull. In

the upper jaw, three early-bud thickenings are evident on the

dental lamina (see fig. 2a for the relations between the dental

lamina and an early bud). The most rostral of these is

associated with the premaxillary ossification, the next is

adjacent to the rostral end of the developing maxillary bone,

and the distal bud occurs at the caudal end of the dental

lamina and is associated with the distal extent of the maxillary

ossification. Based on their bony relationships and comparison

with later ontogenetic stages, these tooth buds can be

identified as dI2 or dI3 anteriorly, dC and dP3, respectively.

The dP3 bud lies beneath the level of the anterior half of the

eye. An overview of the major features of dental development

for the postcanine dentition in the upper and lower jaws is

summarised in Tables ] and 2.

The bud for the early developing incisors in the upper jaw

does not occur at the rostral-most extent of the premaxilla;

instead, it is positioned somewhat more distolateral. It is unclear

whether this developing tooth germ is dI2 or dI3 at this early

stage. The late or retarded development of dll in the upper jaw

of dasyurids, peramelids and didelphids was noted by Thomas

(1887). Later, Guiler and Heddle (1973) suggested that the late

development and eruption of dll helped the continued suckling

of the young. In the lower jaw, a large early bud 1s evident at the

rostral extent of the dentary ossification. Two small bud-like

thickenings occur distal to the rostral early bud (presumably

dil), but these are not as distinct as the dil. It should be noted

here that adult dasyurids possess four upper and three lower

incisors, in contrast to the presumably primitive condition in

ancestral marsupials and didelphids, which possessed five

upper and four lower incisors. It is unclear to us which incisor

has been lost 1n each jaw of Dasyurus, so we will simply call

them [1-4 above and 11-3 below.

We next examined a three-day-old embryo measuring 7

mm GL and 4 mm HL (see fig. 6 in Hill and Osman Hill,

1955). Two small deciduous incisors were in an early-bud stage

rostrally 1n the upper jaw. The moderately sized deciduous

canine was in a more distinct early-bud stage. It had a more

darkly stained abnormal buccal portion and a more normal

pale-stained lingual portion. Based on later developing stages,

the darkly stained buccal portion represents the primordium of

the rudimentary deciduous canine, and the more normal

lingual portion represents the primordium of the successor

canine. These modified relationships for the developing

deciduous and successional incisors and canines were

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus

Table 1. Development of Dasyurus viverrinus upper postcanine dentition

30 hour PY

6 mm GL

(3.25 mm HL)

5—6 day PY

8 mm GL

(4.5 mm HL)

7 day PY

9 mm GL

(5.5 mm HL)

10 day PY

10.5 mm GL

(6.5 mm HL)

10 day PY

11 mm GL

(7 mm HL)

14 day PY

13 mm GL

(8 mm HL)

14 day PY

13.5 mm GL

(8 mm HL)

15-16 day PY

14mm GL

(9 mm HL)

19 day PY

17 mm GL

(10 mm HL)

~19-20 day

16-17 mm GL

~19 day PY

16-18 mm GL

Early bud

No trace No trace

No trace No trace Early to

middle bud

Slight No trace rly

swelling of

Early cap Tiny bud-like | Middle to late

thickening bell, flat

lamina

Early cap No trace Small, late

bell, thin

dentin, flat

successional

lamina

Early bell No trace Moderately

developed

dentin,

successional

lamina

mesially

Late cap to No trace Moderately

thick dentin,

thin enamel;

successional

lamina

mesially

Very early

bud-like

thickening

Late bud to

early cap

Middle to late

Early cap

cap

Early

bud- like

swelling

Middle bell

middle bell

Early cap

Late bud to

early cap

lamina

No trace Early to

middle bell,

flat

successional

lamina

successional

Early bell No trace Moderately

developed

dentin,

successional

lamina

mesially

early bell

46 W.P. Luckett, N. Hong Luckett & T. Harper

25 day PY

20 mm GL

(12.5 mm HL)

Middle to late | No trace thick dentin, | Early to Late bell, Early cap

thin enamel; | middle cap possible

lingual odontoblasts

successional

lamina

No trace Moderately | Late bell, Moderately | Middle to late

thick dentin, | early developed bell

thin enamel; | odontoblasts | dentin, thin

lingual enamel on

successional metacone

lamina

moderately No trace Moderately | Late bell, thin | Moderately | Middle to late

thick dentin, thick dentin, | dentin thick dentin, | bell

moderately thin enamel; thin enamel

developed mesio- on metacone

enamel lingual

successional

lamina intact

thick dentin, thick dentin, | dentin thick dentin | thin dentin middle bud

moderately thin enamel; and enamel

developed incomplete on metacone

enamel mesio-lingual

successional

lamina

Note: GL, greatest length; HL, head length; PY, pouch young

~30-35 day PY

~23 mm GL

~35 day PY

~23—25 mm GL

13.5 mm HL

~40-41 day PY

~29 mm GL

18 mm HL

Table 2. Development of Dasyurus viverrinus lower postcanine dentition

30 hour PY No trace No trace

6 mm GL bud-like

(3.25 mm HL) thickening

5—6 day PY No trace No trace Middle to late

8 mm GL

(4.5 mm HL)

5—6 day PY No trace No trace Late bud to

5.5 mm GL early cap

(4.5 mm HL)

6-7 day PY No trace No trace Late bud

9 mm GL

(5 mm HL)

10 day PY Flattened No trace

11 mm GL early bud

(7 mm HL)

14 day PY Flattened late | Possible

13 mm GL bud to early | slight

(8 mm HL) cap swelling

Middle bell, Early bud

short flat

successional

lamina

lingually

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus 47

Stage {apt j|dp — dp — fmt — |m | — |m ——

~14 day PY

13.5 mm GL

(8 mm HL)

-15-16 day PY

14 mm GL

(9 mm HL)

19 day PY

17 mm GL

(10 mm HL)

~19-20 day PY

16-18 mm GL

Bolk

25 day PY

20 mm GL

(12.5 mm HL)

~30-35 day PY

~23 mm GL

Bolk

~35 day PY

23—25 mm GL

Bolk

13.5 mm HL

~40-41 day PY

~29 mm GL

Bolk

18 mm HL

Late bud to

early cap

Early cap

Late cap to

early bell

Early bell

slight

swelling of

dental lamina

slight

swelling of

dental lamina

slight

swelling of

dental lamina

Possible

slight

swelling

Middle bell

Late bell, thin

dentin

Late bell,

moderately

developed

dentin, thin

enamel

Moderately

thick dentin,

moderately

developed

enamel

Late bell, no

odontoblasts,

lingual

successional

lamina

small

abnormal

bell,

moderately

developed

dentinal cap

Moderately

developed

dentin;

mesio-lingual

successional

lamina

Small,

moderately

thick dentinal

arc, thin

enamel

Small, thick

dentin, thin

enamel

Small,

abnormal

dentinal mass,

with thin

enamel

Abnormal,

moderately

thick dentinal

arc with thin

enamel

Tiny, solid

dentinal mass

with thin

enamel

Possible early | Middle bud

bud-like

swelling

m i BEN

bud-like

Middle to late

bell

swelling

Middle to late | Late cap

bell

Late bell, thin | Middle to late

bell

to moderately

Late bell, Early bell o

Possible early

bud-like

swelling

Early

bud-like

swelling at

mesial end of

dp3

Early bud

Moderately

sized, middle

to late cap,

mesial to

abnormal dp3

Moderately

thick dentin

and

moderately

developed

enamel on

protoconid

developed

dentin

moderately

developed

dentin, thin

enamel

Moderately

large, early to

middle bell

Moderately

thick dentin,

moderately

Moderately

thick dentin,

thin enamel

Middle to late

bell

developed

enamel on

protoconid

Thick dentin,

moderately

thick enamel

on protoconid

Late bell,

moderately

developed

dentin

Probable

early bud

Moderately

large, late

bell, possible

odontoblasts

Moderately

thick dentin,

moderately

developed

enamel on

protoconid

Note: GL, greatest length; HL, head length; PY, pouch young

48 W.P. Luckett, N. Hong Luckett & T. Harper

Figure 1. Four early pouch young of Dasyurus viverrinus: a, 30 hours old, 3.25 mm head length (HL); b, 5—6 days old, 4.5 mm HL; c, 14 days old,

8 mm HL; d, 25 days old, 12.5 mm HL.

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus 49

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Figure 2. Transverse sections of early developmental stages of the dentition 1n marsupials: a, early tooth bud forming from the dental lamina; b,

early-cap stage; c, middle-bell stage with elongate dental lamina; d, late-bell stage with odontoblasts and early lingual successional lamina; e,

well-developed late-bell stage with dentin and enamel. A successional early-cap stage develops from the lingual successional lamina, showing

early fragmentation of the dental lamina. Abbreviations: AM, ameloblasts; D, dentin; DL, dental lamina; DP, dental papilla; E, enamel; IEE,

inner enamel epithelium; OD, odontoblasts; OEE, outer enamel epithelium; SL, successional lamina; SR, stellate reticulum; TB, tooth bud.

previously described in detail for marsupials (Luckett, 1993).

The dental lamina was reduced distal to the developing canine.

The lamina became thickened again distally at the level of the

rostral end of the developing eye. It then gave rise to a moderate

size early- to middle-bud stage dP3. The dental lamina and the

maxillary bone disappeared distal to the dP3 bud.

In a slightly older embryo, 5—6 days old and measuring 8

mm GL and 4.5 mm HL (see fig. 7, Hill and Osman Hill, 1955,

and our fig. 1b), three early developing incisors are now

evident in the premaxillary region of the upper jaw. The

anterior two incisors are moderately sized and flattened in the

early- to middle-bud stages. The third incisor 1s moderately

large and more spherical in the middle-bud stage, and is

located nearer the distal end of the premaxilla. It has a smaller

darkly stained buccal portion and a larger lingual lobular

portion with lighter, normal-appearing cells. Based on its

appearance in later developmental stages, these represent the

buccal deciduous incisor and the more normal-appearing

lingual successor incisor.

A moderately large late-bud to early-cap stage canine

develops at the rostral extent of the maxilla. This also consists

of a smaller darker buccal portion and a larger lingual paler-

staining portion, representing the future deciduous and successor

canines, respectively. The dental lamina 1s reduced distal to the

developing canine and continues distally towards the level of

the middle of the eye. At this region, the dental lamina enlarges

and gives rise to a moderately large early-cap stage dP3, formed

by invagination of the underlying dental papilla cells (see fig. 2b

for an example). The dental lamina then disappears a short

distance distal to this tooth, near the distal end of the eye.

In the lower jaw, three developing incisors are in early- to

late-cap stages at the rostral extent of the dentary. The dil is

in a late-cap stage, with possible odontoblasts buccally and a

lingual projection for an early successor bud for il. The di2

has an early-cap stage buccally and a distolingual projecting

lamina for a possible successor 12. The smaller di3 has a less

distinct buccal early cap with a relatively indistinct lingual

successional lamina. The moderately sized dc is in a late-bud

stage with a lingual successional ridge. The dental lamina

continues distally as a thin ridge with no suggestion of a dpl

or dp2. The dental lamina then thickens again distally,

beneath the level of the distal portion of the eye, and gives rise

to a moderately large late-bud to early-cap stage dp3. The

dental lamina then ends distal to this developing tooth.

In a 10-day-old pouch young with a 6.5 mm HL and a

10—10.5 mm GL (see fig. 9 in Hill and Osman Hill, 1955),

there is little change in the developing incisor and canine

regions of the upper jaw. The dental lamina immediately

distal to the canine exhibits a somewhat flattened early bud-

like thickening for a possible primordium of dP1, but this is

not very distinct. The dental lamina is then reduced and

extends distally. At the level of the middle portion of the eye,

the dental lamina thickens again and gives rise to a moderately

large early-cap stage dP3. The dental lamina then ends distal

to this tooth. In the lower jaw, there is also a moderately sized

early-cap stage dp3 and no trace of a developing dpl.

In a slightly larger 10-day-old pouch young (7 mm HL

and 1] mm GL), several distinct changes have occurred

compared with the preceding specimen. The developing

incisor region remains similar, whereas the deciduous canine

has a tiny buccal nodule associated with a large distolingual

projecting late-bud to early-cap stage successor canine in the

upper jaw. Immediately following the canine is a tiny,

somewhat flattened, probable early bud for dPl. The dental

lamina is reduced distal to this early bud and extends distally

to underlie the eye, where it enlarges to form a moderately

sized middle-cap stage dP3. There was no distinct swelling of

the dental lamina between the dP3 and the probable early-bud

stage dPl. The dental lamina extends slightly distal to dP3

before it disappears.

Note that in the early developing pouch young, there is

little space in the jaws for the developing anterior dentition,

due in part to the greatly enlarged tongue and its intimate

association with the large nipple (fig. 3). This often results in

ereat compression and flattening of the more anterior

developing teeth, whereas there is less compression distally in

the region of the developing dP3 and molars.

In the lower jaw, there is also a small, flattened early-bud

stage for dpl. The dental lamina is then reduced as it extends

distally to form a moderately sized early- to middle-cap stage

dp3. The dental lamina then ends distal to dp3. As in dasyurids

with three premolars, the sequence of early development for

the premolars in both jaws 1s dP3 > dP1 > dP2. As noted in

this and later stages, it is the late-developing dP2 that is lost 1n

both jaws of dasyurids with only two premolars.

In an older 14-day-old pouch young, with 8 mm HL and

13 mm GL (see fig. 10 in Hill and Osman Hill, 1955, and our

fig. Ic), the developing relationships for dll and dI2 in the

W.P. Luckett, N. Hong Luckett & T. Harper

upper jaw were similar to those of the previous stage. In

contrast, dI3 has an early cap knot enclosing a tiny dentinal

arc. The small distolingual successional bud for I3 1s still

relatively small. A small buccal epithelial knot is evident for

dC, and there is a large middle-bud stage successor for the

canine that is associated with a deep maxillary alveolus. A

small, flattened late-bud to early-cap dP! follows the canine

immediately, and then the dental lamina is reduced distal to

this. There is a possible slight swelling of the dental lamina

beneath the anterior end of the eye, but this is not very distinct.

The dental lamina is then reduced distal to this slight swelling.

The dental lamina enlarges further distally and gives rise to a

moderately sized early- to middle-bell stage dP3 beneath the

middle third of the eye (see fig. 2c for a middle-bell stage

tooth). This tooth is still intact with its primary dental lamina

origin. A flattened lingual successional lamina extends mesial

to dP3. Distal to dP3 is a moderately sized late-bud stage M1

underlying the distal end of the large eye. The dental lamina

ends distal to this tooth.

In the lower jaw, the dil is small and cap like, enclosing

thin distinct dentin. Distolingual to this is the larger early-bud

stage successor 1l. The di2 1s also a small cap with an irregular

dentinal fragment; a slightly larger successor 12 is placed

distolingually in a late-bud to early-cap stage. The tiny di3

cap lacks dentin, and its distolingual successor ridge lacks a

distinct successor tooth. The small, dark cap for dc lacks

dentin and has a distolingual middle-bud stage successor

canine. The small, flattened dpl follows immediately and is in

the late-bud to early-cap stage. There 1s no distinct evidence

for a dp2 swelling. Somewhat more distally there is a small

middle-bell stage dp3 with a short, flattened lingual

successional lamina. Immediately distal to this tooth is a

moderately sized early bud for ml. The dental lamina

disappears distal to this developing tooth.

In a slightly larger 14-day-old pouch young (13.5 mm GL

and 8 mm HL; see fig. 10 in Hill and Osman Hill, 1955), some

notable changes are evident in the canine and postcanine

dentition. The successor canine 1s now in the early- to middle-

bell stage in the upper jaw. The tiny, flattened dPl is in the

early-cap stage. The dental lamina becomes reduced distal to

this tooth and then enlarges slightly to form a possible

primordium for a rudimentary dP2. However, the dental

lamina is reduced distal to this slight swelling and then

enlarges further distally and gives rise to a moderately large

middle- to late-bell stage dP3. This tooth has moderately

developed stellate reticulum, and a short, slender lingual

successional lamina extends mesial to this tooth (see fig. 2d

for a late-bell stage with a lingual successional lamina). The

dP3 lies beneath the middle region of the eye. The primary

dental lamina extends distal to dP3 and gives rise to a

moderately large late-bud M1 beneath the distal region of the

eye. The dental lamina then disappears distal to this bud.

In the lower jaw, the successor canine is now in the late-

bud to early-cap stage. This is followed immediately by the

small, elongate late-bud to early-cap stage dpl. The dental

lamina is slightly swollen distal to this tooth, but this is not

very significant. Further distally, the small dp3 is in the late-

bell stage with moderately developed stellate reticulum but no

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus 51

Figure 3. Transverse section through the head at the level of the developing eyes in a 10-day-old Dasyurus viverrinus embryo ( 7 mm head length).

Note the close compression between the large tongue (T) and the nipple (N). No developing teeth are evident.

dentin. There is a lingual successional lamina that extends

mesial to dp3 and terminates with a slight bud-like swelling.

Distal to dp3 is a large middle-bud stage ml, and then the

dental lamina disappears distal to this.

In a slightly older pouch young (13-15 mm GL and 9 mm

HL) estimated to be 15—16 days old, a significant change has

occurred in the postcanine region of the upper jaw. The small

dPl is in the early-cap stage and the dental lamina 1s greatly

reduced distal to this tooth without any suggestion of a

swelling for a rudimentary dP2. Further distally, the small to

moderate sized dP3 is in the late-bell stage with thin dentin on

its apex. The tooth 1s relatively superficial with a short, intact

primary dental lamina stalk to the oral epithelium. The tooth

lies beneath the anterior 75 of the eye. The slender lingual

successional lamina extends mesial to the tooth and then

enlarges to form a moderately sized early-cap stage successor

P3. The successor P3 extends mesially beneath the anterior

margin of the eye. Distal to this, a moderately large middle- to

late-cap stage M1 develops beneath the level of the middle of

the eye. The dental lamina then disappears distal to this tooth.

In the lower jaw, the dpl is a moderately sized early-cap

stage. The dental lamina 1s reduced distal to this tooth; there

is a slight swelling distally, but this 1s not very distinct.

Further distally, the dental lamina enlarges again and gives

rise to a small to moderately sized, somewhat abnormal dp3

with an irregular, moderately developed dentinal cap. There is

a short, primary dental lamina stalk to the oral epithelium.

The tooth is in a shallow bony alveolus and has a flattened

lingual successional lamina. Distal to this tooth is a large

middle-bell stage ml. The dental lamina disappears distal to

the developing molar. In both jaws, the dP3 appear to be

relatively smaller than 1n earlier stages and they lie closer to

the oral epithelium.

In the later developmental stages, we will only discuss the

postcanine development of the Dasyurus pouch young

because there 1s little controversy concerning the development

of the incisor and canine regions. [n a 19-day-old pouch young

(I7 mm GL and 10 mm HL; see fig. 11 in Hill and Osman

Hill, 1955), the small dPl is somewhat flattened or compressed

and has now attained the early-bell stage in the upper jaw. The

dental lamina is reduced distal to this tooth and becomes

detached from the oral epithelium. The dental lamina

becomes reattached further distally and forms an early bud-

like swelling; this possibly represents the rudiment of a dP2.

The dental lamina is then reduced again and extends further

distally; it then enlarges to form a small late-bell stage dP3.

There is moderately developed dentin but no enamel on this

small tooth. It lies beneath the middle of the eye in a shallow

bony alveolus. A lingual successional lamina extends mesially

from dP3 and forms a slight bud-like thickening. As noted in

the later developmental stages, this thickening is the

primordium for the later developing successor P3. The dental

lamina extends distal to dP3 and then enlarges to form a

moderately large middle-bell stage M1. This tooth lies

beneath the distal half of the eye; the dental lamina disappears

distal to this tooth.

In the lower jaw, dpl 1s a moderately sized, late-cap to early-

bell stage tooth with a shallow bony alveolus. The dental lamina

is slightly swollen distal to this tooth (possibly representing the

site of the lost dp2); then further distally, the dental lamina

thickens again and gives rise to a small late-bell stage dp3 with a

moderately developed layer of dentin. The lingual successional

lamina of this tooth extends mesially, where it is slightly

thickened terminally. The dental lamina extends distal to this

tooth and gives rise to a large middle- to late-bell stage ml. The

dental lamina disappears distal to this tooth.

In a later pouch young of 19—20 days old (16-18 mm GL),

sectioned by Dr Louis Bolk, the dPl is a small tooth in the late-

cap to early-bell stage 1n the upper jaw. The dental lamina distal

to this tooth is detached from the oral epithelium. There is no

trace of a possible rudimentary dP2. Further distally, the dental

lamina reappears and gives rise to a small, late-bell stage dP3

with moderately thick dentin and a thin film of early enamel.

The lingual successional lamina of dP3 extends mesially and

gives rise to a small late-bud to early-cap stage successor P3,

which lies anterior to dP3 (see fig. 2e for a late-bell stage dP3

and its successor early-cap stage P3). Distal to dP3, the dental

lamina gives rise to a moderately large early- to middle-bell

stage M1. The dental lamina then disappears distal to this tooth.

In the lower jaw, the moderately sized dpl is in the early-

bell stage. The tooth 1s somewhat compressed and lies in a

shallow bony alveolus. There is only a slight possible swelling

of the dental lamina distal to this tooth 1n the region of the

missing dp2. Further distally, the dental lamina thickens again

and gives rise to a small late-bell stage dp3 with a moderately

thick arc of dentin. There is also a thin layer of darkly stained

enamel. The slender lingual successional lamina adjacent to

dp3 extends mesially and 1s enlarged as a slight early bud-like

swelling for the future successional p3. The dental lamina is

reduced distal to dp3 and then enlarges again to form a large

middle- to late-bell stage ml. Distal to this tooth is a

moderately large late-cap stage for m2. The dental lamina

disappears distal to this tooth.

In another 19—20-day-old pouch young (10 mm HL and

16-17 mm GL), there are slight advancements for some of the

developing teeth. A moderately sized dPl is 1n the early-bell

W.P. Luckett, N. Hong Luckett & T. Harper

stage in the upper jaw. There is no distinct evidence for a

rudimentary dP2 swelling distal to this tooth. The dental

lamina thickens further distally and gives rise to a small to

moderately sized late-bell stage dP3 with moderately

developed dentin on its crown. There is a shallow bony

alveolus for the tooth, which lies beneath the anterior third of

the eye. The lingual successional lamina of dP3 extends

mesially and thickens to give rise to a small early-cap stage

successor P3. Distal to the dP3, the dental lamina thickens

again and gives rise to a moderately large middle-bell stage

M1. This tooth is positioned beneath the middle of the eye.

The dental lamina thins and then disappears distal to MI.

In the lower jaw, a moderately sized dpl is 1n the late-cap

to early-bell stage. There is no distinct evidence for a

rudimentary dp2 distal to this tooth. Further distally, there 1s

a small, somewhat abnormal dp3 with a moderately developed

dentinal arc but no stellate reticulum. The lingual successional

lamina of dp3 ıs slightly thickened mesially and then gives

rise to a small middle-bud stage successor p3 further

anteriorly. Distal to dp3, the dental lamina enlarges and gives

origin to a large middle- to late-bell stage ml. Distal to this

tooth, the dental lamina gives rise to a moderately sized late-

cap stage m2. The dental lamina then disappears distal to this

tooth. Note that in the 19—20-day-old pouch young, the

dentition in the lower jaw is accelerated in its development

compared with that in the upper jaw; this trend continues to

increase in later developmental stages.

In a later 25-day-old pouch young (20 mm GL and 12.5

mm HL; see fig. 12 in Hill and Osman Hill, 1955, and our

fig. Id) that has been sectioned longitudinally, dPl is a small

to moderately sized tooth in the middle- to late-bell stage in

the upper jaw (fig. 4). As in the previous stage, there is no

suggestion of a dP2 rudiment. Further distally, the small to

moderately sized dP3 has relatively thick dentin and thin

enamel. The tooth is located beneath the anterior 1⁄3 of the eye.

The lingual successional lamina of dP3 extends mesially and

thickens to form a moderately large early to middle-cap stage

successor P3. The deciduous and successor P3 appear to arise

from a common primary dental lamina stalk. This is due to

the partial fragmentation of the primary dental lamina stalk

in later stages of development (compare fig. 4 with 2e). Distal

to the developing dP3/P3 complex, the M1 is now in a large

late-bell stage with possible early odontoblasts but no dentin.

This tooth underlies the middle of the eye. A moderately sized

early-cap stage M2 develops distal to Ml. The dental lamina

then disappears distal to this tooth.

In the lower jaw, dpl 1s now a moderately large middle-bell

stage tooth, developing immediately distal to the large

developing canine. There is no distinct evidence for a rudiment

of dp2. Further distally, the small to moderately sized dp3 is

partially covered with a thick layer of dentin and a thin layer of

enamel. Mesiolingual to the small dp3, the lingual successional

lamina gives rise to a larger early-bud stage p3. Further distally

is a large late-bell stage m1 with moderately developed dentin

on the apex of the tall protoconid. Distal to this tooth, the

dental lamina gives rise to a large middle- to late-bell stage

m2. A possible early bud for m3 lies distal to this tooth, but

problems with the tissue make it difficult to be certain of this.

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus

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Figure 4. Parasagittal section through the dentition 1n a 25-day-old pouch young (12.5 mm head length) showing the developing teeth between

dPl and M2 in the upper jaw. Arrowhead refers to the site of the missing dP2. Abbreviation: D, location of developing dP3 follicle.

In a 30—35-day-old pouch young (23-24 mm GL and 13.5

mm HL; see fig. 13 in Hill and Osman Hill, 1955), the dP1 is

now a moderately sized tooth with moderately thick dentin

and thin to moderately developed enamel on its crown in the

upper jaw. The tooth lies in a shallow bony alveolus, just

below the oral epithelium. This 1s related 1n part due to the

large canine that partially underlies the smaller dPl

throughout its length. There is no trace of a rudimentary dP2.

Further distally, the dental lamina thickens and gives rise to a

small, somewhat elongate dP3 with moderately thick dentin

and a partially disrupted layer of enamel. The tooth lies

anterior to the developing eye. The lingual successional

lamina of dP3 extends mesially and gives rise to a larger

moderately sized late-bell stage successor P3 with

odontoblasts and a thin layer of predentin on its crown. The

P3 extends both anterior and posterior to dP3 because of its

larger size. There is a normally sized developing bony

alveolus for P3, whereas a distinct bony alveolus is lacking for

the small dP3. Instead, the small deciduous tooth lies more

superficially, in the apex of the deeper alveolus for the larger

P3. Distal to the dP3/P3 complex, the large M1 has moderately

thick dentin and thin enamel on its elevated metacone. There

is a flattened residual lingual lamina along the extent of the

tooth. This tooth lies beneath the anterior half of the eye. A

moderately large middle- to late-bell stage M2 lies beneath

the distal half of the eye. The dental lamina disappears distal

to this tooth.

In the lower jaw, dpl is a relatively large tooth with

moderately developed dentin and thin enamel on its apex.

There is no suggestion of a rudimentary dp2. Further distally,

the small dp3 has a relatively thick dentinal arc overlain by a

thin film of enamel. The enamel is covered by inactive cuboidal

ameloblasts. Lingual to the small abnormal dp3 is a thin

successional lamina; this extends further mesially and gives

rise to a moderately large early- to middle-bell stage p3 with a

loosening of its stellate reticulum. Distal to the dp3/p3

complex, the ml is a large tooth with moderately thick dentin

and thin to moderately developed enamel on the apex of its tall

protoconid. The tooth has a short, flat residual lamina. Distal

to this tooth is a large m2, also with moderately developed to

moderately thick dentin and thin enamel on its tall protoconid.

The m2 also has a short, flat residual lamina. A moderately

large m3 lies distal to m2 and is in the middle- to late-bell

stage. The dental lamina disappears distal to this tooth.

Another pouch young 30-35 days old (23 mm GL) was

sectioned by Louis Bolk and showed some small differences

with the previous specimen. The dPl is small to moderately

sized and is in the late-bell stage with early odontoblasts and

thin predentin in the upper jaw. The dpl has a short, flat

lingual successional lamina. Again, there 1s no suggestion of a

rudimentary dP2. The small, somewhat abnormal dP3 has

moderately thick dentin and possibly a thin film of enamel.

Mesio-lingual to dP3 is a moderately sized late-bell stage P3

with very early odontoblasts. The elongate lingual

successional lamina is fragmented in places and it is not

clearly connected to dP3, in contrast to earlier stages. Distal

to dP3 is a large late-bell stage M1 with moderately developed

dentin and thin enamel on the tall metacone. A prominent,

flattened residual lamina is present lingually. Distal to this

tooth is a moderately large middle- to late-bell stage M2. The

dental lamina disappears distal to this tooth.

In the lower jaw, the moderately large late-bell stage dpl

has thin predentin on the apex of its single cusp; it immediately

follows the large canine. There is no suggestion of a

rudimentary dp2. Further distally is a small, abnormal

dentinal mass for the rudimentary dp3 with a thin layer of

enamel on the surface of the dentin. The primary dental

lamina connection to the oral epithelium persists. Anterior to

the abnormal dp3 is a moderately sized middle- to late-cap

stage p3. Only a few fragmented epithelial nodules represent

the earlier remnants of the lingual successional lamina from

dp3. Distal to dp3 is the large m1 with moderately thick dentin

and thin to moderately developed enamel on the apex of its

tall protoconid. A thin, flattened residual lamina is evident

mesially. Distal to this is a large late-bell stage m2 with

moderately developed dentin and thin enamel on the apex of

the tall protoconid. There is also a prominent, short residual

lamina lingually. Distal to this tooth is a moderately sized

early-bell stage m3; the dental lamina then disappears distal

to this tooth.

In a later pouch young of about 41 days old (29 mm GL

and 18 mm HL; see fig. 14 in Hill and Osman Hill, 1955), also

sectioned by Louis Bolk, further changes occurred, especially

with regards to the dP3 and P3 in both jaws. In the upper jaw,

the small dP! now has moderately thick dentin and moderately

developed enamel on the apex of the unicuspid tooth. There is

a short, flattened lingual successional lamina, and again no

trace of a rudimentary dP2. Further distally, there is a small

dP3 with moderately thick dentin and moderately developed

enamel. Ameloblasts are low cuboidal and probably are no

longer functional. Fragmented strands of the lingual

successional lamina extend anteriorly and give rise to a

moderately sized late-bell stage successor P3 with thin early

predentin on its apex. Further distally, the large M1 appears; it

has moderately thick dentin and enamel on its tall disto-

buccal metacone. Distal to this tooth is a large late-bell stage

M2. This tooth also has an elevated disto-buccal cusp that has

odontoblasts and a very thin layer of predentin. Following this

tooth is a moderately sized early to middle-bud stage M3. The

dental lamina then disappears distal to this developing tooth.

In the lower jaw, dpl is a moderately sized tooth with a

relatively thick layer of dentin and a moderately developed

layer of enamel (partially disrupted). Its primary dental lamina

stalk is still attached to the oral epithelium. Only a slight

suggestion of the thin lingual successional lamina is evident

mesially. There is no trace of a rudimentary dp2. Distal to dpl

is a very small, abnormal dp3 with a prominent dentinal mass.

There is also a very thin film of enamel, although there are no

distinct remnants of ameloblasts. The tooth is considerably

reduced in size compared with earlier developmental stages.

The primary dental lamina connection of dp3 with the oral

epithelium and its lingual successional lamina are also broken

in places. Nevertheless, the fragmented lingual successional

lamina extends mesially and gives rise to a moderately large

late-bell stage successor p3. The p3 has well-developed stellate

reticulum and possibly early odontoblasts, although this is not

clear. The tiny dp3 lies more superficial than its larger

successor and lacks a distinct bony alveolus.

Distal to dp3 is a large ml with thick dentin and

moderately thick enamel on its tall, mesio to buccal

protoconid. The tooth lies in a deep bony alveolus. Distal to

ml is a large m2 with moderately thick dentin and moderately

developed enamel on its tall protoconid. Further distally, m3

is a moderately large tooth in the late-bell stage with a

moderately developed layer of dentin on its tall protoconid.

There are also early ameloblasts on the tooth cusp, but no

enamel. There is a well-developed lingual residual lamina

mesially on the tooth. Distal to this tooth is a possible early

W.P. Luckett, N. Hong Luckett & T. Harper

bud for m4 developing on both sides of the jaw, but tissue

disruption makes this difficult to confirm. The dental lamina

disappears distal to this region.

Later development and eruption of the dentition in

Dasyurus viverrinus

We would like to have been able to microscopically examine

later developmental stages of the dentition in Dasyurus

viverrinus, but at the time we were working at the Hubrecht

Laboratory of Comparative Embryology, later stages were not

available. Many older specimens, from 46 days old to about

120 days old, were described by Hill and Osman Hill (1955),

but only a few brief comments were provided by them

concerning the early eruption of a few teeth. Presumably,

these later stages of alcohol specimens were included in the

Hill collection when it was moved with the Hubrecht

collection to the Museum für Naturkunde.

Concerning the later development and eruption of teeth in

Dasyurus viverrinus, Hill and Osman Hill (1955) cited the

earlier studies by Thomas (1887) and especially Woodward

(1896), and accepted their beliefs that P3 is the tooth that has

been suppressed in this species with two premolars. They all

agreed that a vestigial dP3 developed in the jaws and that the

lingual successional lamina of this tooth was much swollen

and probably represented a rudiment for the successor P3,

although it probably does not develop much further. All

agreed that the vestigial dP3 1s present but does not erupt and

that the swollen primordium of P3 does not differentiate much

further or erupt.

It seems clear to us that none of these authors

microscopically examined later developmental stages of

Dasyurus viverrinus beyond the “much swollen" stage of P3.

In our studies, this stage occurred at 15-20 days of

development. Indeed, in some 19—20-day-old pouch young,

the successor P3 was already in the late-bud to early-cap

stage. Between 20 and 41 days of development in our study,

the P3 developed normally in both jaws, attaining the late-bell

stage. In contrast, the dP3 in both jaws became relatively

smaller and more abnormal in these later developmental

stages; this would be consistent with continued reduction and

non-eruption of dP3 in later stages.

The absence of a developing dP2 and the normal

development of the successor P3 in the 25—41-day-old pouch

young in our study provide a strong basis for our belief that

dPl and P3 are the two erupting premolars 1n both jaws of

later developmental stages 1n Dasyurus viverrinus. Hopefully,

future investigators will microscopically examine the later

developmental stages and early eruption of the jaws in

Dasyurus viverrinus to corroborate (or refute) our hypothesis

of dental homologies.

Comparison of Dasyurus viverrinus with the three-

premolared dasyurids Sminthopsis virginiae (Luckett and

Woolley, 1996) and Antechinus stuartii (Luckett and Hong

Luckett, pers obs) shows that there 1s a similar pattern 1n the

early postcanine development of the premolars in these two

eroups, despite the loss of one deciduous premolar in the

Dasyurus species. While the deciduous premolars differentiate

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus OO

and develop in a posterior to anterior series (dP3 > dP2 > dPI)

in both jaws of the didelphids Didelphis and Monodelphis, the

dasyurids show a different developmental sequence of dP3 »

dPl » dP2. In Sminthopsis virginiae, a previous study showed

that dP2 was relatively late in its initiation, not appearing until

about 20 days of pouch young development (Luckett and

Woolley, 1996). The same is true for Antechinus stuartii. As

shown ın our developmental series of Dasyurus viverrinus, it

is this later developing dP2 that has been lost in this species

with only two premolars. In our Dasyurus viverrinus series, it

was during the 15 to 20 days of development that there was a

slight suggestion of thickening of the dental lamina between

dPl and dP3; this probably represented a rudimentary attempt

to develop a dP2. In the later stages of development, from 25 to

41 days, there was no longer any sign of a rudimentary dP2.

Another interesting and unusual condition in Dasyurus

viverrinus, when compared with the dasyurids with three

premolars, 1s the nature of the development of dP3 in both jaws.

The dP3 is the first postcanine tooth to initiate differentiation

in both jaws, as also occurs in all three-premolared dasyurids

examined, as well as in didelphids. In contrast with these

marsupials with three premolars, however, the dP3 in Dasyurus

undergoes an abnormal state of development in later stages in

both jaws, including a relative reduction in size and having a

relative superficial position in the jaw with only minimal

development of a bony alveolus. This abnormal development of

dP3 is even more noticeable in the lower jaw. There is a large

amount of dentin but only minimal amounts of enamel. In

contrast, the successional P3 is developing normally in both

jaws and it 1s quickly larger in size and deeper in the jaw than

its deciduous predecessor. Although we could not follow later

stages histologically, it is the normal developing successor P3

that erupts adjacent to dP1 in both jaws. In contrast, we are

unaware of any evidence for eruption of the small, abnormal

dP3 in Dasyurus viverrinus.

Another significant difference between Dasyurus

viverrinus and the dasyurids with three premolars is the

accelerated rate of development and eruption of P3 in Dasyurus,

when compared with these other dasyurids and with didelphids.

In the 97-day-old Sminthopsis virginiae examined by us (see

Luckett and Woolley, 1996), M1-2 are erupted in both jaws and

M3 ıs partially (75) erupted in the upper jaw. In the lower jaw,

m3 1s almost completely erupted and the protoconid of m4 is in

an early stage of eruption. The dP! to dP3 are erupted in both

jaws, whereas the moderately large and well-developed

successor P3 is still unerupted in both jaws.

In contrast, there 1s no sign of an erupted or unerupted

dP3 in either jaw of Dasyurus viverrinus at about 95 days old,

and dPl is in early eruption. The successor P3 is in early

eruption at about 98 days old; this 1s about the same time as

the early eruption of M1. These data on Dasyurus viverrinus

eruption are from the study by Merchant et al. (1984),

although we have changed the names of their premolars ("first

and second upper premolars") to correspond to our more

precise and correct terminology because they considered dP2

to be present and P3 to be lost. As with most other studies on

Dasyurus viverrinus, Merchant et al. (1984) did not

microscopically examine any of their extensive series of

developing pouch young. We also note that these authors

found no trace of an erupted dP3 ın earlier or later stages of

development; this 1s consistent with our findings of abnormal

development of dP3 in later stages of development.

Variation in the fate of dP3 within Dasyurus

Neither Hill and Osman Hill (1955) nor Merchant et al. (1984)

could detect any evidence for the eruption of a rudimentary

dP3 in Dasyurus viverrinus, and we have not seen such a

tooth in any of the juvenile or subadult skulls that we have

examined. However, in another Dasyurus species that we

have studied, Dasyurus albopunctatus, we have noted similar

developmental stages to those of Dasyurus viverrinus, as well

as an unusual condition during later developmental stages and

early eruption (Luckett pers obs). Our specimens of Dasyurus

albopunctatus consist of microscopic sections of two early

pouch young and a larger number (26) of skulls of juvenile

and subadult specimens with erupting postcanine dentitions.

In the youngest specimen (fig. 5) that we examined

microscopically (AMNH 195149; 12 mm HL), the postcanine

region contained a moderately large late-bell stage dP1 with no

Figure 5. Dasyurus albopunctatus pouch young (AMNH 195149, 12 mm

head length). Scale bar is 5 mm.

odontoblasts in the upper jaw. There was a diastema for the

missing dP2 with no swelling of the dental lamina. Further

distally was a moderately sized dP3 with moderately thick dentin

and moderately developed enamel. Extending mesially from the

dP3 was a lingual successional lamina that gave rise to an early-

to middle-bud stage successional P3. Further distally was a large

M1 with moderately developed dentin but no enamel. Distal to

this was an early- to middle-cap stage for M2; the dental lamina

ended distal to this tooth. This specimen is very similar in its

developmental features to that of our Dasyurus viverrinus

25-day-old pouch young (12.5 mm HL). A later developmental

stage of Dasyurus albopunctatus (AMNH 193967; 32 mm HL)

was considerably advanced. The dPl had thick dentin and

enamel, and again, there was no trace of a rudimentary dP2 in

the upper jaw. The now tiny calcified dP3 was elongate and

slender, considerably reduced in relative size compared with the

previous specimen; it was very close to the overlying alveolar

margins. Deeper in the jaw and lingual to this was a moderately

large successor P3, with thick dentin and enamel. This specimen

was considerably more advanced in its development compared

with our 41-day-old Dasyurus viverrinus.

In a slightly later juvenile skull of Dasyurus albopunctatus

(AM M24313; skull damaged; not measured), the second to

fourth incisors were in early eruption in the upper jaw and the

canine was in an earlier stage of eruption. The dPl was 4-7

erupted and there was no trace of a dP2. Immediately distal to

this, the moderately sized successor P3 was in very early

eruption, less than that of dPl. Lying on the apex of the P3

crown was a tiny, elongate and pearl-shaped dP3 on both sides

of the jaw. The tiny dP3 had a crown height of 1.3 mm on the

right side and 1.4 mm on the left side. The M1 was almost

completely erupted and M2 was in early eruption. In comparison

with the data on Dasyurus viverrinus from Merchant et al.

(1984), this specimen would be 95—98 days old. In the lower jaw,

the incisors are almost completely erupted, as is the canine. The

dpl is also almost completely erupted. The successor p3 1s about

^5 erupted and there is no trace of a tiny dp3. The ml is almost

completely erupted and m2 1s about 2 erupted.

In nine juvenile skulls of Dasyurus albopunctatus from

later developmental stages that we examined, we could see the

tiny dP3 lying adjacent to the erupting P3 in the upper jaw in

skulls where the P3 was not completely erupted (see fig 6a—d).

It would be incorrect to say that the tiny dP3 were erupted,

and we prefer to consider them to be displaced during the

eruption of P3. The oldest specimen that we examined with

the tiny dP3 was a partially damaged juvenile skull (AMNH

151980; 43.51 mm Skull Length). The dPl was erupted and

P3 was almost completely erupted on the right side of the

upper jaw. The MI] was erupted and M2 was almost

completely erupted. In the lower jaw, dpl, p3 ml and m2 were

erupted and m3 was about ?4 erupted. As in all the juvenile

specimens examined by us for this species, there was no trace

of a dp3 1n the lower jaw. This specimen was briefly noted by

late (1947, p. 142) who stated that the "upper and lower fourth

premolars are absent. The milk P4, however, can still be

detected". As we have shown in this and earlier developmental

stages, it is dP2 that is the missing tooth in both jaws, and P3

(his fourth premolars) 1s developed in both jaws. It was the

W.P. Luckett, N. Hong Luckett & T. Harper

tiny dP3 (his “milk P4") that he detected in the upper jaw.

In later developmental stages of Dasyurus albopunctatus,

in which P3 was completely or almost completely erupted in

the upper Jaw, the tiny dP3 was absent in most cases. However,

in a few instances of some specimens with P3 erupted, tiny

tooth-like fragments of a probable apical or basal remnant of

dP3 were found unilaterally (AMNH 151973, 151971). The

oldest juvenile that we found with a slender root remnant of

dP3 unilaterally (fig. 6d) was in a male (BBM-NG 97868;

47.25 mm Skull Length) with dP1, P3, MI and M2 erupted in

the upper jaw, and M3 was about 4 erupted.

We suggest that the pattern of dental development and

eruption in Dasyurus albopunctatus is an intermediate

condition between that of the dasyurids with three premolars

that we have examined and Dasyurus viverrinus. Both of our

species of Dasyurus have a similar accelerated rate of

development for the eruption of P3 at about the same time as

MI and M2, compared with that for the dasyurid species

Sminthopsis and Antechinus with three premolars. The

ereatly reduced dP3 adjacent to the erupting P3 1n the upper

Jaw of Dasyurus albopunctatus 1s an 1ntermediate condition

between the presence of an erupted dP3 in the dasyurids with

three premolars and its absence in Dasyurus viverrinus.

Although we have not made an extensive search for

rudimentary dP3 remnants in other species of Dasyurus, we

suspect that they can and will be found in some of these

species. We have seen one such probable tiny dP3 in Dasyurus

geoffroii and Dasyurus hallucatus, and we have seen a similar

condition of accelerated eruption of P3 in these two species.

We also note that Archer (1975, p. 255) reported the presence

of a rudimentary spicule-like tooth in the position of the

“normally absent" dP3 in Dasyurus geoffroii (WAM M6370);

we suspect this was also a dP3 fragment.

A final thought. Although Dasyurus viverrinus 1s considered

to be extinct on mainland Australia, we hope that investigators

in Tasmania will help complete our studies on the later

development and eruption of the dentition in this species, which

still exists there but is threatened. Investigators there have

studied many other aspects of the biology of this interesting

species, and we hope that this paper will stimulate some to

further support or challenge our findings on dental development.

Acknowledgements

We are grateful to all the personnel from the Hubrecht

Laboratory of Comparative Embryology, from the Director to

the kitchen staff, for making our numerous stays in Utrecht

and at their laboratory a special experience for us, both

scientifically and personally, during our studies in their

collections. Our colleague and friend Patricia Woolley in

Australia helped considerably with the preparation and

examination of the specimens of Dasyurus albopunctatus in

her care. Colleagues at the American Museum of Natural

History in New York, and at the Bernice Bishop Museum in

Honolulu, Hawaii, helped considerably with specimens in

their care. Our colleague Stephanie Canington at Johns

Hopkins helped greatly in the organisation of the manuscript.

Initiation and early development of the postcanine deciduous dentition in the dasyurid marsupial Dasyurus viverrinus 57

Figure 6. Dasyurus albopunctatus pouch young: a, (AMNH 190927; 36.64 mm skull length). Erupting teeth in the upper jaw with canine about 1⁄2

erupted. The dP1 is almost completely erupted and P3 is in early eruption. A tiny, slender dP3 is lying on the distal margins of the erupting P3,

pointing posteriorly; its apex is not evident. M1 is erupted and the M2 is about 74 erupting (not shown); b, (AMNH 221650; 37.53 mm skull length)

slightly older than the preceding. The dP1 is erupted in the upper jaw and P3 is erupting (2-7). A tiny, elongate dP3 is evident along the posterior

margins of the erupting P3; c, (BBM — NG 28247; 37.89 mm skull length). The dP1 is erupted, and the P3 is almost completely erupted in the upper

Jaw. A tiny, slender and elongate dP3 is evident along the distal margin of P3 and its root extends distally beneath the anterior opening of the

infraorbital canal; d, (BBM — NG 97868; 47.25 mm skull length). In this older young, the P3 is almost completely erupted in the upper jaw. A tiny,

slender root remnant of dP3 is evident, distal to the buccal end of P3.

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Memoirs of Museum Victoria 80: 59-99 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/|.mmv.2021.80.04

A review of the Australasian genus Pseudophycis (Gadiformes: Moridae),

redescribing its four species and resurrecting the name Physiculus palmatus

Klunzinger, 1872, for the Australian Red Cod

MARTIN F. Gomon! 4t, CARL D. STRUTHERS? and JoDIE KEMP”

' Museums Victoria, Melbourne Museum, 11 Nicholson Street, Carlton, Victoria, 3053, Australia.

Email: mgomon@museum.vic.gov.au

^ Museum of New Zealand Te Papa Tongarewa, 467 Cable Street, Wellington, New Zealand. Email: CarlS@tepapa.govt.nz

* 25 Grigg Court, Wallington, Victoria, 3222, Australia. Email: jodie@hindson.com.au

* Corresponding author

Gomon, M.F., Struthers, C.D. and Kemp, J. 2021. A review of the Australasian genus Pseudophycis (Gadiformes:

Moridae), redescribing its four species and resurrecting the name Physiculus palmatus Klunzinger, 1872, for the

The commercially fished Red Cod Pseudophycis bachus Forster in Bloch and Schneider, 1801, 1s shown on the basis

of both morphological and genetic evidence to be confined to New Zealand waters. ‘The separation of the closely related but

distinct Australian cognate brings the number of species in the Australasian genus Pseudophycis to four. The name

Physiculus palmatus Klunzinger, 1872, which was long thought to be a junior synonym of Pseudophycis barbata Günther,

1862, 1s demonstrated to have been originally proposed for the Australian endemic. Pseudophycis palmata (new combination)

differs from P. bachus 1n having a prominent black blotch or spot basally on the pectoral fin that does not extend dorsally

onto the body, in contrast to distinctly extending onto body, the second dorsal fin with 47—56 rather than 40—45 rays, the anal

fin with 50—57 rather than 42—48 rays and the vertebral column with 47—50 rather than 44—46 total vertebrae. Diagnoses,

colour descriptions, images and distributions for all four species and a key to members of the genus are provided.

Abstract

Australian Red Cod. Memoirs of Museum Victoria 80: 59—99.

Keywords Teleostei, Actinopterygii, morphology, re-description, comparison, distribution, COI

Introduction

In an annotated catalogue of gadiform fishes of the world,

Cohen et al. (1990) treated the morid genus Pseudophycis as

comprising three species: Pseudophycis bachus Forster in

Bloch and Schneider, 1801, P. barbata Günther, 1863, and P.

breviuscula Richardson, 1846, all confined to the temperate

waters of New Zealand and Australia. The taxonomy followed

Paulin's (1983) review of New Zealand species of Moridae

that concluded that only three are present in New Zealand and

Australian waters, despite the historical confusion over which

names apply to the various Pseudophycis species. Paulin did,

however, observe meristic variations in fin ray and scale

counts between New Zealand specimens and the limited

number of Australian specimens of P. bachus at his disposal,

saying "examination of a larger sample of Amustralian

specimens might show some differences but, in view of the

variability within species of Moridae, there would probably

be little justification for separating the populations at the

species level" (Paulin, 1983, p. 92). Subsequent authors (e.g.

Gomon in Gomon et al., 1994; Kuiter, 1993; Last et al., 1983;

Paxton and Hanley in Paxton et al., 1989; Paxton et al., 2006;

Yearsley et al., 1999) dealing with temperate Australian fish

biodiversity followed Paulin (1983) and Cohen et al. (1990) in

treating the Australian Red Cod as conspecific with the New

Zealand species.

smith et al. (2008) detected genetic differences between

Australasian samples of Pseudophycis using the COI locus.

Their work included eight samples of P. bachus, four each

from Australia and New Zealand, six samples of P. barbata

from Australia, but no samples of the third species, P.

breviuscula. They found differences between the P. bachus

samples from the two countries that were equivalent to

differences between P. barbata and each of the two P.

bachus morphs, suggesting species-level differences

between all three. These genetic discrepancies were

supported by work of Kemp (2010), who observed that there

are a number of differences between individuals from

Australian and New Zealand waters and that morphometric

and meristic characteristics of Pseudophycis species in

Australian waters have been poorly documented. She

compared 50 Australian specimens with 160 from New

Zealand, both series identified as P. bachus, and found

significant differences in mean values for the second dorsal

(50.2 vs. 42.8, respectively) and anal (52.1 vs. 45.0) fin ray

counts, as well as pyloric caeca (9.6 vs. 6.0). These

observations provided the incentive to further compare the

Australian and New Zealand populations with each other

and with the other two long-recognised members of the

genus, P. barbata and P. breviuscula. The outcome was a

recognition of four easily distinguished species in the genus

and the initiation of research on the scientific name for the

Australian Red Cod.

The Australian Red Cod is a relatively common demersal

species, distributed in the shelf waters of south-eastern

Australia (Gomon in Gomon et al., 2008). It 1s an important

prey species for some high trophic level predators, including

little penguins, Eudyptula minor (Chiaradia et al., 2003;

Cullen et al., 1992; Montague and Cullen, 1988), Australian

fur seals, Arctocephalus pusillus doriferus (Gales and

Pemberton, 1994; Hume et al., 2004; Kirkwood et al., 2008;

Litnan et al, 2007) and New Zealand fur seals,

Arctocephalus forsteri (Page et al., 2005). The trophic role of

Australian Red Cod as prey, coupled with its relatively

common occurrence, suggests that this species 1s particularly

important in Australian waters. Humans include these fishes

as part of the non-targeted bycatch consumed annually,

although the species is more frequently discarded than

retained (pers. comm. M. Tudman, Bycatch Reduction

Program Manager, Australian Fisheries Management

Authority, 2008).

This contribution provides a name for the endemic

Australian Red Cod, a detailed description that distinguishes

it from the New Zealand P. bachus, descriptions for the four

Species in the genus and a dichotomous key to the species.

Methods

Morphometric methodology mostly followed Paulin (1983),

Trunov (1990) and Markle and Melendez (1988);

measurements were taken directly (point-to-point) except

that measurements between fins were horizontal distances

between vertical projections at the origins of each fin,

caudal peduncle length was taken from the posterior end of

the anal fin base to the hypural joint, caudal-fin length was

from the hypural joint to the fin tip, and lengths of gill

rakers and filaments were taken at or adjacent to the angle

of the gill arch. In lists of specimens examined, numbers of

specimens and their standard lengths in mm are placed in

parentheses after registration numbers, except for lots

containing only a single specimen where only the length is

presented. Meristic methods are those of Hubbs and Lagler

(1947) where counts of unpaired fins and vertebrae were

taken from radiographs, the number of trunk centra was

counted from the cranium to the last centrum with pleural

ribs and counts of tail centra exclude the urostyle. Selected

meristic and relative morphometric ranges for the four

congeners are presented in Tables 2—4. Other materials

examined are in the ichthyological collections of the

Australian Museum (AMS), Natural History Museum

(formerly British Museum [Natural History]; BMNH),

M.F. Gomon, C.D. Struthers & J. Kemp

Australian National Fish Collection (CSIRO), Museums

Victoria (NMV), Museum of New Zealand Te Papa

Tongarewa (NMNZ), South Australian Museum (SAMA),

Staatliches Museum für Naturkunde, Stuttgart (SMNS) and

Western Australian Museum (WAM).

Genetic material and sequences. A genetic analysis of

mitochondrial cytochrome oxidase subunit I (COI) gene was

carried out on 72 sequences from muscle tissues, representing

5 terminal taxa of the family Moridae.

The dataset included 61 sequences from the BOLD

database (www.boldsystems.org) and 11 sequences newly

generated for this study (see Appendix | for list of material

and BOLD and GenBank accession numbers; vouchers for

tissues yielding sequences in the material examined lists are

indicated by a dagger ^?) including 18 specimens of

Pseudophycis bachus from New Zealand, 23 P. barbata from

Australia and New Zealand, 12 P. breviuscula from Australia

and New Zealand, I2 P. palmata from Australia, and 7

Auchenoceros punctatus from New Zealand as a morid

outgroup.

Genomic DNA was extracted from the 11 (7 P. bachus

[including neotype], 2 P. breviuscula and 2 P. barbata)

additional ethanol-preserved muscle tissues (Appendix 1)

using DNeasy blood and tissue kit (Qiagen). A fragment of

the COI gene was PCR amplified and sequenced using

thermocycling conditions and primers (FishF2 and FishR1)

from Ward et al., 2005). PCR products were purified by

digestion with the ExoProStar (Illustra) or with the Zymo

DNA Clean and Concentrator (Zymo Research, USA),

following the manufacturer’s instructions. PCR products

were sequenced in both directions on an ABI-3730 (Massey

University Genome Service, Palmerston North, New

Zealand or Macrogen, Seoul, South Korea). Newly

generated sequences were deposited in GenBank (see

Appendix | for a list of voucher specimens and their

accession numbers).

Phylogenetic analyses. All 72 sequences were individually

aligned in ClustalW using the software Geneious Prime

v10.0.5. The COI alignment contained no insertions or

deletions. Maximum likelihood, neighbour joining and

maximum parsimony trees were used to visualise distance

relationships within and between species; node support was

assessed with 1,000 bootstrap replications of each tree. For

the maximum likelihood analyses, the most appropriate

model of sequence evolution for each dataset was determined

using the Akaike information criterion in ModelTest; the

HKY + G model was selected (Hasegawa et al., 1985) because

it was the best performing of 24 different nucleotide

substitution models tested in the model selection option of

MEGA version X (Kumar et al., 2018). A discrete gamma

distribution was used to model evolutionary rate differences

between sites (5 categories, +G, parameter = 0.3772). In

addition, a neighbour joining tree was estimated using

p-distances (uncorrected distances). The p-distance is the

proportion of nucleotide sites at which two sequences being

compared are different. It is obtained by dividing the number

of nucleotide differences by the number of nucleotides

Australasian red cods

compared. It does not make any correction for multiple

substitutions at the same site, substitution rate biases (e.g.

differences in the transitional and transversional rates), or

differences in evolutionary rates between sites. For the

maximum parsimony analyses, a heuristic search algorithm

with 1000 random addition sequence replicates and tree

bisection and reconnection branch swapping methodology

was used. Nodal support was tested with 500 bootstrap

pseudo-replicates, with ten random addition replicates for

each pseudo-replicate.

Multivariate analysis of morphological data

To investigate the a priori grouping by species, a canonical

analysis of principal coordinates (CAP) was performed using

normalised Euclidean distances between specimens

(Anderson and Willis, 2003). CAP is equivalent to a classical

discriminant analysis. Leave-one-out mis-classification error

(Lachenbruch and Mickey, 1967) was used to obtain a direct

measure of the ability of the CAP discriminant model to

identify species on the basis of the 20 characters. This cross-

validation provided a rigorous assessment of the

distinctiveness of the four species of Pseudophycis. The

following 20 characters, measured in 124 Pseudophycis

specimens of the four species (see material examined), were

used in the analysis: anal fin base length, barbel length, caudal

peduncle depth, caudal peduncle length, caudal fin length,

first dorsal fin base length, first dorsal fin longest ray length,

head length, orbit diameter, pectoral fin base width, pectoral

fin length, pelvic fin length, postorbital length, pre-dorsal fin

length, pre-anal fin length, pre-pectoral fin length, pre-pelvic

fin length, pelvic fin origin to anal fin origin length, second

dorsal fin base length, and snout length.

Genus Pseudophycis Gunther, 1862

Pseudophycis Günther, 1862: 350, type species: Lota breviuscula

Richardson, 1846, by monotypy.

Austrophycis Ogilby, 1897: 90, type species: Austrophycis

megalops Ogilby, 1897 (= P. breviuscula) by monotypy.

Physiculus (Pseudophycis) Whitley, 1956: 403,

decision.

taxonomic

Diagnosis. Morid fishes with an elongate body. Luminescent

organs absent. Snout broadly rounded in dorsal view, not

projecting greatly beyond mouth, length about equal to

interorbital width. Eye diameter much less than postorbital

length. Each jaw with broad band of small, depressible,

caniniform teeth of uniform size. Vomerine teeth absent. Chin

with well-developed, often small, barbel. Branchiostegal rays

seven. Gill rakers of outer branchial arch short or of moderate

length. Otolith with ostium approximately equal to the cauda

(1:1 to 1:1.4), the crista superior as long as or longer than the

crista inferior; otolith thickness 20—22% of its length (Paulin,

1983). Scales small, covering body and head; extending onto

loose membranes enveloping the bases of dorsal and anal fins;

lateral line pores at ends of short tubes from naked strip not

associated with individual scales. Two dorsal fins and one anal

fin, first dorsal slightly taller than, immediately adjacent to and

Joined to second by low membrane, with 8—14 rays, first ray not

prolonged; second dorsal and anal fins with elongate bases, of

nearly uniform height; caudal fin separate, rounded or truncate.

Ventral fins composed of five or six rays, the two outermost

longest and projecting beyond fin membrane but not greatly

prolonged.

Etymology. The feminine name Pseudophycis is an

amalgamation of the Greek words pseudos, meaning “lie” or

"false", and phykis, "a fish living among seaweeds” (Brown,

1954) and the source word of the scientific name for the North

Atlantic gadid genus Phycis Walbaum, 1792, in reference to the

overall resemblance of species of the two genera to one another.

Distribution. Species of the genus are endemic to coastal

waters of temperate Australia and New Zealand.

Remarks. Species referable to the genus Pseudophycis were

described or treated as members of seven morid genera based

on the superficial resemblances of species in the genera to

one another. These include: Gadus Linnaeus, 1758, type

species Gadus morhua Linnaeus, 1758, by subsequent

designation of Jordan and Gilbert (1883: 802); Enchelyopus

Bloch and Schneider, 1801, type species Gadus cimbricus

Bloch and Schneider, 1801, by subsequent designation of

Jordan and Evermann (1898: 2560); Lota Oken, 1817, type

species Gadus lota Linnaeus, 1758, by subsequent tautonomy

(Fricke et al., 2019); Lotella Kaup, 1858, type species Lota

phycis Temminck and Schlegel, 1846, by monotypy and

Physiculus Kaup, 1858, type species Physiculus dalwigki

Kaup, 1858, by monotypy (see species treatments for details).

Austrophycis Ogilby, 1897, caused particular confusion until

sazanov (2001: 343) identified the type specimen of A.

megalops as P. breviuscula (Richardson, 1846). The genus

Pseudophycis comprises four species.

Genetic analysis. Concerted efforts to document genetic

sequences for the COI gene across Australasian fish species as

part of the Fish Barcode of Life project provided an additional

means of testing suspect species concepts. A preliminary

comparison of sequences of New Zealand and Australian

specimens identified as P. bachus and P. barbata in BOLD had

already revealed potentially cryptic sister species on either side

of the Tasman Sea. Sequences for specimens identified as P.

bachus had low divergence among specimens within Australia

(0.002) and within New Zealand (0.000) but high sequence

divergence in the pooled data (0.086) typical of species pairs

(Smith et al., 2008: 8, fig. 5).

A comparison of a larger set of sequences for Australian

and New Zealand specimens identified as species of

Pseudophycis, plus sequences of specimens of Auchenoceros

punctatus as an outgroup, generated a tree with five inferred

species branches, four having a common basal node for

vouchers identifiable as P. barbata, P. breviuscula, P. bachus

and P. palmata (Ward pers. comm.; fig. 1). Importantly, the

branches of P. bachus and P. palmata, previously thought to

be conspecific, are of about the same lengths as those of

P. barbata and P. breviuscula, despite the morphological

differences distinguishing the latter two (Table 1).

Key to species of Pseudophycis

l. Caudal margin truncate with angular corners in specimens

larger than about 150 mm SL, central caudal fin rays

distinctly shorter than those extending to the corners; basal

portion of pectoral fin with distinct dark blotch or spot 2

— Caudal margin rounded with rounded corners, central

caudal fin rays equal in length or longer than those

extending to the corners; no distinct dark blotch or spot

basally on pectoral fin 3

2. Dark blotch or spot basally on pectoral fin extending

dorsally onto body; second dorsal fin with 40—45 rays;

anal fin with 42—48 rays; total vertebrae 44—46

— Dark blotch or spot basally on pectoral fin not extending

dorsally onto body; second dorsal fin with 47—56 rays;

anal fin with 50—57 rays; total vertebrae 47—50

3. Scales above lateral line 16—22; second dorsal fin with

54-59 rays; anal fin with 54—63 rays; pyloric caeca 14—20;

total vertebrae 48—51 Pseudophycis barbata

— Scales above lateral line 7—9; second dorsal fin with 44—52

rays; anal fin with 49—56 rays; pyloric caeca 6-9; total

vertebrae 42—45 Pseudophycis breviuscula

Pseudophycis palmata (Klunzinger, 1872)

Proposed name: Australian Red Cod (Australian Standard)

Figures 1, 2, 3A, 4, 5; Tables 1—4

Physiculus palmatus Klunzinger, 1872: 38. Type locality:

Hobsons Bay, Victoria, Australia. Types: SMNS 1589.

Pseudophysis barbatus (nec Günther, 1862). McCoy, 1878: 29, pl.

20 (in part; description).

Pseudophycis bacchus (nec Forster in Bloch and Schneider, 1801).

Günther, 1880a: 28 (Twofold Bay, NSW ); Waite, 1899: 119 (distribution).

Pseudophycis bachus (nec Forster in Bloch and Schneider, 1801).

Ogilby, 1886: 48 (list, in part); Edgar etal., 1982: 32, fig. 17 (description,

in part); Last et al., 1983: 234, fig. 21.7 (description, in part); Paulin,

1983: 93 (distribution, in part); Hutchins and Swainston, 1986: 34, 124,

fig. 113 (description); May and Maxwell, 1986: 194 (description);

Paxton and Hanley in Paxton et al., 1989: 302 (list, 1n part); Grant,

1991: 75, fig. 164 (list, in part); Gomon in Gomon et al., 1994: 333, fig.

M.F. Gomon, C.D. Struthers & J. Kemp

297 (description, in part); Yearsley et al., 1999: 87, fig. (description);

Paxton et al., 2006: 616 (taxonomy, in part); Gomon in Gomon et al.,

2008: 313, fig. (description, 1n part).

Physiculus bachus (nec Forster in Bloch and Schneider, 1801).

Waite, 1904: 24; Stead, 1906: 86 (description, in part); Waite, 1907: 18,

fig. (South Australia); Stead, 1908: 48, pl. 16; Waite, 1921: 67;

McCulloch, 1921: 42; McCulloch, 1922: XVII, 32; Waite, 1923: 92,

fig. (description, in part); Lord and Scott, 1924: 8, 43 (description, in

part); Lord, 1927: 13; McCulloch, 1927: 32, pl xi, fig. 112a (list); Waite,

1928: 6 (listed); McCulloch, 1929: 129; McCulloch, 1930: 129 (in

part, listed); Norman, 1935: 3; Norman, 1937: 54, 55 (in part, listed);

Munro, 1961: 62, fig. 441, (description; in part); Scott, 1962: 84, fig.

(description, in part); Whitley, 1962: 58; Whitley, 1964: 40 (list, in

part); Walker, 1972: 2; Suda, 1973: 2150—2152 (distribution); Scott et

al., 1974: 95, 96, fig. (description, in part).

Pseudophycis barbata (nec Günther, 1862). Kuiter, 1993: 59, fig.

(description, in part).

Diagnosis. First dorsal fin with. 10—12 rays, second dorsal fin

with 47—56 rays; anal fin with 50—57 rays; total vertebrae 47—

50; nostrils located about two-thirds to three-quarters of the

way from snout tip to eye; gill rakers of outer arch of moderate

length, almost as long near angle as opposing gill filaments;

chin barbel short, 6-17% HL; scales above lateral line in

oblique series from base of first dorsal fin ray 11—15; oblique

rows of scales intersecting with lateral line 96—116; pyloric

caeca 8-10; caudal fin truncate with angular corners in

specimens larger than about 150 mm SL, middle rays shorter

than rays extending to corners; dark blotch basally on pectoral

fin not extending onto side above fleshy pectoral fin base. A

large species reaching at least 598 mm TL or approximately

532 mm SL (Kemp, 2010: 26).

Description. (Values for non-type specimens when different

from type in parentheses; see Tables 2—4 for summary of

selected meristic and comparative morphometric values.) First

dorsal fin 11 (10—12, rarely 10, first ray usually minute); second

dorsal fin 56 (47—56, rarely 47 or 54—55); anal fin 55 (50—57,

rarely less than 51); pectoral fin 22 (22-27); pelvic fin 5 (5 or 6,

rarely 6); caudal fin 37 (35—41) rays; gill rakers 4 + 10 (3—4 +

9—11 = 12-15); lateral line pores not associated with individual

scales; oblique scale rows intersecting with lateral line

unknown (96-116), scales 1n oblique series above lateral line

15 (11—15), scales in oblique series below lateral line unknown

(28—40); vertebrae 16 + 34 (15-18 + 31-34) = 50 (47—50);

pyloric caeca 10 (8—10).

Table 1. Estimates of evolutionary divergence over sequence pairs within (left) and between (right) species. Kimura-2-parameter genetic distances

as averages over all sequence pairs within species are in the left column, with standard error estimates on the right (left table). Genetic distances

as averages over all sequence pairs between species are below the diagonal with standard error estimates above it (right table).

Between species

Distance below and Standard Error above diagonal

Species Within species

standard

N Distance Error A. punctatus

A. punctatus ji 0.002 0.001

P. bachus 18 0.005 0.001 0.160

P. breviuscula 12 0.001 0.001 0.170

P. palmata 12 0.005 0.002 0.158

P. barbata 23 0.003 0.001 0.171

P. bachus P. breviuscula | P. palmata P. barbata

0.017 0.017 0.017 0.017

0.015 0.013 0.015

0.143 0.016 0.011

0.092 0.139 0.014

0.127 0.085 0.128

Australasian red cods

Table 2. Selected counts for types and representative specimens of the four species of Pseudophycis.

AIP PISFUBIU guv

DINISNIAIAG “J

əd) N A9p pis F ue&oui

0c - VI

9S - 8t

cc 7 9I

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əƏdÁ}| N A9p pis F ueoui

yc - 9c

cl - OL

9€] -TOI

6t -tt

01 -9

Vc - 6l

6-9

9] - VI

e-v

ISUBI

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ISUB.I

0043} N Aop pis F ue&oui

nud ‘J

92992 OIIO[Ád

OUT] [&19]e| ^O[9q so[eog

OUT] TEINE] 9A0qe so[eog

SMOI 9[09S 9SIOASUUI[,

SABI UY [epned eon,

sÁKvI ur [epneo JUSLINIOId IOMO'T]

SÁLI UY [epneo (pououeiq) [edr»uuq

SÁLI ur [epneo juaumooud 1odd()

SIDYPI [[I8 [RIOT

SJo*e1 [fis ədd N

SIDYCI [IIo I20'T

IVIGOLIOA [VIOL

3UJQ33I9A [Epn€

IBIQOLIOA [epneoodq

sABI UY [eJO0]2od

SAEI uj euy

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SABI UY [BSIOP 3SILT

J3]9€.IEQ2 JI)SLI9 [AI

64 M.F. Gomon, C.D. Struthers & J. Kemp

Table 3. Frequencies of selected meristic characters in Pseudophycis species. Shaded values are those of types.

Second dorsal fin rays 40 41 42 43 44 45 46 47 48 49 50 51 52 33 54 #55 56 57 58 39 N

P. palmata 2 8 4 10 6 7 4 3 2 1] 47

P. bachus 1 4 13 10 4 | 33

P. barbata 7 90 F 5 5 D 36

P. breviuscula | 29M 13 5 141 1 2 2 48

Anal fin rays 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 N

P. palmata l 5$ 4 Il 7 Big 3s 7 48

P. bachus 1] 4 5 8 8 6 | 33

P. barbata 2 4 7 8 6 5 ]| 1 1 1 36

P. breviuscula 7 Nu > 58 & 5S 4 | 46

Total caudal fin rays 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 4l N

P. palmata 3 10 6 19 2 |] | 42

P. bachus ] 1 3 EOS 9 1 2 23

P. barbata T m 5 1] 18

P. breviuscula fm 2 4 7 12 7 4 37

Caudal vertebrae 27 28 29 30 31 32 33 34 35 36 37 N

P. palmata 1 13 26 6 46

P. bachus 1 9 18 5 33

P. barbata 4 11 12 NEM 1 35

P. breviuscula 4 $239. 18 3 48

Total vertebrae 42 43 44 45 46 47 48 49 50 5I N

P. palmata 2 10 29 6 47

P. bachus 6 20 7 33

P. barbata 2 8 19 6 35

P. breviuscula 3 18 21 6 48

Scales above lateral line 7 8 9 10 Il 12 13 14 #15 16 17 18 19 20 21 22 N

P. palmata 4 10 4 4 5 27

P. bachus 6 12 16 34

P. barbata 4 8 10 2 3 1 2 30

P. breviuscula 7 13 2 22

Pyloric caeca 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

P. palmata | 2 95 8

P. bachus 29 29

P. barbata 2 T 58 3 2 l 20

P, breviuscula 4 4 8

Table 4. Selected proportional measurements for types and representative specimens of the four species of Pseudophycis.

Australasian red cods

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N Aeppjscueour oSgu&e1 əd | N Aeppjscueoui osuvrl odd) | N AS3pDpIScUPQ9UI — ssuBl əd} | N Asppiscxueour oSue1 — odÁ JU3UIQ3JnSe2][AI

DINISNIAIAG d DIDGADG d snunq d nud “J

66 M.F. Gomon, C.D. Struthers & J. Kemp

NMNZ P.045835| NZ. ECNI| FNZC252-09

NMNZ P.053551| NZ. Bay of Islands

NMNZ P.044103| NZ. White Island | FNZC280-09

NMNZ P.037124] NZ. White Island | FNZC189-09

NIWA no voucher SER6] NZ. Ngawi] FNZAG08-08

NIWA. no voucher SERS| NZ. | FNZA387-06

NIWA no voucher SERAH NZ. | FNZA386-08

NIWA no voucher SER3| MZ. | FNZAXES-06

NIWA no voucher SBR7|NZ. Ngawil FNZAGQ9-08

NIWA no voucher SER2| NZ. | FNZA384-08

NIWA no voucher SER1[ NZ, | FNZA383-06

NMNZ P.052745|NZ. Capre Brett

NMNZ P.037109| NZ. White Istand| FNZC188-09

| CSIRO H 7949-13 AUS, SA, GABIBW-A13730

99955 |L CSIRO no voucher] BW-1698] AUS. NSW. S of Diaster Bay| FOAD138-05

CSIRO H 3791-01] AUS. VIC. $ of Gabo Eland| FOADI35-05

CSIRO H 4500-01|AUS. VEC. 5 of Cape Everard| FOAD141-05

| CSIRO no voucher|BW-1699| AUS. NSW. 5 of Diaster Bay| FGADI39-05 -

CSIRO H 7539-03| AUS. TAS. E of Orford| FOACUSO-14

| CSIRO H 7904-03| AUS. SA. GAB|BW-A13489

CSIROno voucher|&W-1697| AUS. NSW. N of Cape Howe] FOADI37-05

CSIRO no voucher|EW-1700| AUS. NSW. S of Diaster Bay] FOADI40-05.

CSIRO no voucher jT 10004| AUS. SA. GAB|BW-AT4L20

CSIRO H 7698-09| AUS. TAS. Huon Commonwealth Marine Reserve| BW-A13043

| MMNZ P. 046285 | HZ. Bay of Iskandis| FNZC272-09

MMNZ P.046285-1|NZ. Bay of Iands|FNZC271-09 ——

CSIRO H 6845-01/3|AUS. NSW. E of Broken Bay] FOAKG10-10 ———

CSIRO H 6838-08) AUS. NSW. E of Broken Bay] FOAKÉ32-10 b.

NMAN P.052719| MZ. Ranfurly Bank :

| CSIRO H 6836-08/2] AUS. NSW. E of Broken Bay| FOAKG04-

CSIRO H 6845-011] AUS. NSW. E of Broken Bay| FOAKB26-10.

CSIRO H 6845-01/2] AUS. NSW. E of Broken Bay| FOAKBL;

CSIRO H 6980-04) AUS. TAS. E of Tasmania | BW-ABD56

CSIRO H 6991-01|AUS, TAS. E of Tasmaría|EW-ABO67

CSIRO no voucher] GT 7975] AUS. TAS. The Friars] BW-A13029-

NMW A 31132-001 AUS. TAS. off Taroona| BW-A12965

CSIRO no voucher! GT 7974] AUS. TAS. The Friars] BW-A:

CSIRO no voucher| BW-1693|AUS. VIC. Bass Strait| FOAD133-05 -

J| CSIRO H 7715-01]AUS. TAS. Munro Bight] BW-A13031 ——

CSIRO H 4225-01] AUS. TAS. S of Port Arthur] FOAD136-05 _

Wut.

CSIRO urreg GT 7976|AUS. TAS. The Friars| BW-A13030

CSIRO H 7717-01|AUS. TAS. Hinsby Beach|BW-A13032

CSIRO no voucher| BW-1692| AUS. VIC. Bass Strait| FOAD132-05.

SS NMNZ P.044349] NZ. Chatham Rise] FNZA558-08

NIWA no voucher RCOS| N2. S of Auckland Island| FNZA351-07-

NIMNZ P.044350|N2. Chatham Rise| FN2A559-08

NMNZ P.053749| NZ. Stewart Island

asses || NIWA no voucher RCO1| NZ. ME of Campbell Island|FNZ949-07

NIWA no voucher RECHI] NZ. NE of Campbe! EHand|FNZ950-07

NIWA no voucher RCO6| NZ. S of Auckland Esland| FNZA352-07

NIWA no voucher RCO?7| NZ. |FNZASEB-08

NIWA no voucher ROOS] NZ.|FNZA389-08

NIWA no voucher ROOSI NZ. |FNZA390-08

NMNZ P.044338| MZ. ECSTIFNZASS5-08

NMNZ P.044339| NZ. ECSI]FNZAS56-08

NMNZ P.044348| NZ. Chatham Rise

NMMHZ P.053748| NZ. Stewart Island

NMNZ P.053750| NZ. Stewart Island

NMNZ P.053751|NZ. Stewart Island

NMNZ P.053752| NZ, Stewart Island

NMNZ P.054&828| NZ. Whanganui Bar| Neotype |

NIWA no voucher PCO2| NZ. WESI] FNZA334-07

NIMNZ P.045575|NZ. WCSI| FN2C107-09

39/9/99 |. NMINZ P.045S76|NZ. WCSI] FNZCC108-09

NMNZ P.045577]NZ. WCST| FNZC109-09 A. punctatus

NIWA na voucher PCO1| NZ. WESH FNZA333-07

NIMNZ P.045573|NZ. WCSI| FNZC- 105-09

NMNZ P.045574| NZ. WCSI| FNZC 106-09

0.05

Figure 1. Hasegawa-Kishino-Yano maximum likelihood tree of 65 COI sequences from four purported species of Pseudophycis, along with seven

sequences for an outgroup species, Auchenoceros punctatus. Percentage bootstrap values followed by bootstrap values in p-distance neighbour

joining and maximum parsimony trees are given for all well supported nodes (2 of 3 exceeding 80%). Specimen numbers are those of registered

museum vouchers or sequence numbers in the Barcode of Life Database (BOLD, http://www.barcodinglife.org/). o

Australasian red cods 6/

20mm

E d

=a IN

^ $

a bh

z — -

g UV -

jd 7 7 -

20 mm

Figure 2. Pseudophycis palmata. A, B, CSIRO H 7366-01, 265 mm SL, Storm Bay, east of Variety Bay on North Bruny Island, Tasmania,

Australia, fresh specimen, lateral view of body and lateral view of head and anterior body with black pectoral blotch, respectively, (photographs

C. Devine, CSIRO); C, SMNS 1589, holotype, 172 mm SL, Port Phillip, Hobsons Bay, Victoria, Australia (photograph C. Struthers, NMNZ).

M.F. Gomon, C.D. Struthers & J. Kemp

- Gr.

-A0

P. palmata F. bachus

(S) Holotype ^ (9) Nectype

> othermaterial 9 other material

150

P. barbata

@) Holotype

@ data

^ ether material y

120 l 180

P. breviuscula

(9) Holotype

@ data

^? ether material

120 150 ` 180

Figure 3. Collection localities for specimens of Pseudophycis spp. in museums. A, P. bachus, green and P. palmata, orange; B, P. barbata, blue;

C, P. breviuscula, red.

Australasian red cods 69

Caudal Peduncle Length vs SL

Q P. palmata holotype

& P, palmata

e P. barbata

Q P. palmata holotype

9 P. palmata

e P. barbata

Ə P. palmata

e F. barbata

Q P. palmata holotype

e P. palmatu

@ P. barbata

Figure 4. Plots of proportional measurements (76 SL) against standard lengths for non-type specimens of Pseudophycis palmata, P. barbata and

holotype of P. palmata. A, caudal peduncle length; B, interorbital width; C, barbel length; D, pelvic fin length.

M.F. Gomon, C.D. Struthers & J. Kemp

Resemblance: D1 Euclidean distance

0.2

0.1

CAP2

-0.1 -

-0.2

-0.2 -0.1

CAP 1

Key

3€ P palmata Holotype

A P palmata

MX P. bachus Neotype

O P bachus

$- P barbata Holotype

O P barbata

A P. breviuscula Holotype

1 0.2

Figure 5. Scatter plot of canonical analysis of principal coordinates scores on the first and second canonical axes for 124 specimens of

Pseudophycis, including types.

Body moderately slender, compressed laterally (fig. 2),

greatest depth at anal fin origin 23.6 (18.7-26.5)% SL,

tapering uniformly from second dorsal fin origin to shallow

caudal peduncle; caudal peduncle moderately short, 8.9 (6.7—

10.8)% SL, strongly compressed, depth subequal to orbital

diameter. Distance between middle of anus and base of anal

fin slightly less than half suborbital depth. Body cavity

extending to above origin of anal fin.

Head acute, of moderate size, length 26.1 (25.9—-30.0)%

SL, width 14.5 (15.3—21.6)% SL and depth 12.8 (10.2—18.4)%

SL; snout of moderate length 7.6 (6.7—9.1)96 SL, not projecting

in advance of upper jaw, rounded in dorsal view. Nostrils

small, located about two-thirds to three-quarters of the way

from snout tip to eye, distance from nostril to orbital margin

less than (equal to or less than) diameter of combined nostrils;

nostrils positioned above horizontal through centre of eye;

posterior opening larger than (about half size to larger than)

anterior opening, separated from it by raised skin flap; most of

margin (most or just posterior opening) encircled by tubular

flap. Interorbital broad, slightly convex. Eye of moderate size,

orbital diameter 28.0 (18.9—31.5)96 HL, 1.33 (0.66—1.87) times

in interorbital space, 2.41 (1.65—7.06) times suborbital

distance, circular, upper edge of eye adjacent to dorsal margin

of head in lateral view, transparent skin covering eye faintly

pigmented near orbital dorsal margin. Postorbital moderately

long, 0.91 (0.55—1.01) times length of upper jaw. Mouth large,

terminal, upper jaw terminating posterior to (at or posterior

to) vertical through posterior margin of eye. Jaw teeth

caniniform, slightly curved, depressible, band of up to four or

five irregular rows in upper jaw with distinct hiatus at

symphysis; teeth in outer row more regularly arranged than in

inner rows; band tapering near rear of Jaw. Teeth of lower Jaw

similar in form to those of upper jaw; band of two or three

rows with broader patch on either side of symphysis, tapering

to single, widely spaced row posteriorly; rows almost

contiguous across symphysis. Vomerine teeth absent.

Australasian red cods

Opercular bones strong; upper extremity of gill opening at

horizontal through middle of eye; gill membranes continuous

across isthmus. Gill rakers on outer arch slender, of moderate

length, about 0.8 to 1.3 times length of opposing gill filaments,

8.4—12.2 times in head, their inner margin denticulate. Chin

barbel short, subconical 11.3 (6.0—17.4)96 HL.

Small cycloid scales covering all of head, including gular

region and isthmus, and body except for branchiostegal

membranes, surface of maxilla and premaxilla, lower lip and

distal parts of fins; thick mucus covering obscuring scales and

pores in freshly preserved material. Most head pores tiny,

following main cephalic sensory canals; row of slightly raised

pores from nostrils to tip of snout and then posteriorly just

above lower edge of suborbital; row of enlarged mandibular

pores on underside of lower jaw. Lateral line comprising

widely spaced pores on short tubes arising from narrow scale-

less gap, anterior end curved upwards slightly, then gradually

descending to lateral midline below posterior 25% of second

dorsal fin, remaining on lateral midline posteriorly.

Membranes of first and second dorsal fins continuous at

base; first dorsal originating distinctly behind vertical through

pectoral fin base; anterior two-thirds of second dorsal of

uniform height, lOth to 12th ray from posterior end of fin

longest, its height about 1.4 times length of 2nd ray at anterior

end of fin, last ten or so rays progressively shorter; rays of both

dorsal fins mostly unbranched, only last 13 to 16 rays, apart

from last one to three, branched. Fleshy, fine scale-covered

basal sheath on third or more of first dorsal and anterior

portion of second dorsal fins; fin rays interconnected by

membranes to tips, sheath gradually decreasing slightly in

coverage and thickness posteriorly, encompassing about half

of second dorsal fin near its insertion; sheath extending onto

body for anterior third of combined dorsal fin base, broadest

anteriorly, narrower posteriorly. Profile of anal fin like second

dorsal fin with comparable unbranched and branched rays.

Anal fin enclosed in broad fleshy sheath like dorsal fins; sheath

also extending onto body for anterior third or less of fin.

Caudal fin truncate with distinct dorsoposterior and

ventroposterior corners in adults, posterior margin with slight

convex curve; longest rays to corners, middle rays about

85—90% length of longest rays; fin more rounded in juveniles;

base of fin covered by indistinct sheath sharply demarcated

from scales of caudal peduncle. Pectoral fin tip reaching (not

quite to or to) vertical through anal fin origin, sixth or seventh

ray longest. Pelvic fin inserted anterior to vertical through

posterior edge of preopercle (more posteriorly in some

Specimens); outer two rays longer than inner rays; second ray

longest, 15.1 (11.7-17.3)76 SL, nearly twice length of

subsequent ray, reaching vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 2A,

B.) Medium brown above extending ventrally to about ventral

portion of pectoral fin base, white below, suffused with pinkish

hue, especially above anal fin base; lateral line slightly paler at

least anteriorly. Underside of head, jaws and barbel white,

sometimes tinged with pink. Dorsal, caudal and distal half of

anal fins medium brown; proximal half of anal fin white,

especially anteriorly, with pink hue; dorsal and anal fins with

‘1

fine black edge; posterior edge of caudal fin with broad black

margin. Pectoral fin medium brown with semi-circular black

basal spot covering dorsal 80% of proximal edge of fin,

extending little 1f at all onto side dorsal to fin base. Pelvic fin

rays white with pink hue.

Preserved colour. Mostly pale (upper half of head and body

pale dusky to dusky, lower half very pale, frequently pearly

white. Dorsal, caudal and distal half of anal fins pale dusky;

distal edges of dorsal and anal fins with fine dark margin;

posterior edge of caudal fin broadly dark; pelvic fin, pectoral

fin and basal half of anal fin very pale; pectoral fin with

prominent dark spot (faded in type) covering dorsal half or

more of basal edge, not extending onto side of body dorsally.

Etymology. The specific epithet palmata appears to be Latin

for “embroidered with palm branches”, although the reason for

the name is unknown.

Distribution. Endemic to coastal temperate waters of south-

east Australia at least from Port Lincoln, Spencer Gulf, South

Australia (34° 44' S, 135° 52' E, SAMA F2766), to Port

Stephens, New South Wales (32° 49' S, 152° 05' E, AMS

1.25865-003), including all of Tasmania (fig. 3a). An unverified

record at Coles Point, South Australia, 34° 22' 06” S, 135° 21'

09" E (SAMA F11864), may extend the distribution slightly

farther to the west (fig. 3A). Occurs on soft bottom habitat

at 2-115 m.

Remarks. Pseudophycis palmata 1s closely related to the New

Zealand P. bachus, with which it was confused (Günther, 1880a:

28; McCulloch, 1921: 42; Ogilby, 1886: 48; Waite, 1904: 24).

Both differ from the remaining two congeners by the possession

of a black spot or blotch at the base of the pectoral fin and a

truncated caudal fin margin in adults. The two also differ from

the others in having the lower half of the body and the basal half

or more of the anal fin mostly white, rather than tan to brown,

although these areas are occasionally suffused with orange to

pink in specimens of all four species. Morphometrically, the

pair appear to have a deeper caudal peduncle (3.8—5.9, mean

4.7% SL vs. 3.3—5.2, mean 4.4% SL), shorter barbel (1.7—4.9,

mean 3.5% SL vs. 4.7—9.6, mean 6.5% SL) and shorter or longer

head (25.9—30.0, mean 27.4% SL vs. 23.3—31.6, mean 28.8 in P.

barbata and 23.4—26.0, mean 24.8% SL in P. breviuscula), as

well as other relative lengths of features that correspond with

the general body form. All four species in the genus are

separable by vertical fin ray, scale and vertebral counts, as

identified 1n the above key and in Table 2. Both P. palmata and

P. bachus attain a large size, as does Pseudophycis barbata

(well over 600 mm SL), while P. breviuscula 1s the smallest

species, reaching only about 150 mm SL.

The two P. bachus-like species are very similar to each

other proportionally, although the eye of the Australian

species is slightly larger than that of P. bachus, the orbital

diameter 5.3—9.7, mean 7.4% SL vs. 4.6—8.1, mean 6.2% SL,

the paired fins proportionally shorter, pectoral fin 13.5—17.7,

mean 15.8% SL vs. 14.6—19.4, mean 17.1% SL and pelvic fin

11.7-17.3, mean 14.3% SL vs. 11.5—21.1, mean 16.7% SL, and

caudal fin similarly shorter 9.9—15.3, mean 12.2% SL vs.

11.2-15.6, 13.7% SL. Pseudophycis palmata is readily

separable from P. bachus by the smaller pectoral fin blotch

that fails to extend onto the body above the pectoral fin base.

It also differs subtly from P. bachus in having the distal half

of the anal fin brownish with a fine black margin rather than

whitish, like the basal half, and the black margin confined to

the posterior lobe of the fin, if it is present at all. The

Australian cognate has fewer transverse scale rows (96-116

vs. 102-136 in P. bachus) but more pyloric caeca (8 — 10 vs.

6), anal fin rays (50—57 vs. 42—48) and second dorsal fin rays

(47-56 vs. 40—45).

In the course of exploring the taxonomic identity of the

Australian Red Cod, the species to which the name Physiculus

palmata Klunzinger, 1872, 1s referable became less and less

clear. The name was regarded as a junior synonym of P.

barbata for more than 80 years (McCulloch, 1929: 129).

Klunzinger’s (1872) original description is inadequate for

separating the three species of Pseudophycis occurring at or

in the vicinity of the type locality. A specimen (SMNS 1589)

in the Stuttgart Museum, the repository of Klunzinger's type

material, registered the year the name was published,

apparently collected at the type locality and clearly identified

by Klunzinger as that species, was regarded as a syntype by

Fricke (1992: 13; 2005: 48). It 1s much smaller (173 mm SL)

than the 50 cm length given in the description, implying

Klunzinger was aware of or had other material. The dark spot

on the base of the pectoral fin that 1s diagnostic for the species,

as well as dark pigment on the distal edge of the caudal fin,

has completely faded. The only other specimen of

Pseudophycis (SMNS 2242) dating from that approximate

time currently in the Stuttgart collection was identified by

Klunzinger in 1877 and is unlikely to have been available

when the description was published. We therefore follow

Fricke in regarding SMNS 1589 as the only known type.

Although the three species occurring near the type

locality are clearly separable by morphological characters, the

colours and markings on the type specimen that would have

been diagnostic have faded. Meristic characters including

vertebral numbers, fin ray and scale counts have ranges that

overlap slightly in the two most likely candidate species, P.

barbata and the Australian P. bachus cognate. Unfortunately,

the type specimen of P. palmata has meristic values that fall

in the overlap zone for all but three characters, the number of

scales above the lateral line 15 (11—15 in P. bachus-like vs.

16—22 in P. barbata), total caudal fin rays 37 (35—41 in P.

bachus-like vs. 32—35 in P. barbata) and pyloric caeca 10

(8—10 in P. bachus-like vs. 14—20 1n P. barbata) that favour its

identity as the P. bachus cognate (Table 1). A comparison of

relative morphometric values for the type with those of these

two species, however, revealed clear support for the identity of

the type as the P. bachus-like species with eight of the type's

relative measurements (head length, caudal peduncle length,

caudal peduncle depth, post-orbital length, interorbital width,

barbel length, pelvic fin length and caudal fin length)

positioned closer to the P. bachus-like species proportional

curves relative to standard length (e.g. fig. 4) and curves for

the same measurements (not head length) equally close to the

proportional curve relative to its head length. Only predorsal

fin length has clearly different proportional curves for the P.

M.F. Gomon, C.D. Struthers & J. Kemp

bachus-like species and P. barbata, with the predorsal value

for the type of P. palmata falling equidistant between the

curves of the two species for that measurement.

The diagnostics of a CAP analysis revealed that 96.8% of

124 specimens of Pseudophycis were correctly classified as

their respective species based on the 20 characters examined.

By species, the percentage of individuals correctly classified

by the CAP discriminant model was 96.6% for P. palmata

(with one misclassified as P. bachus), 97.3% for P. bachus

(with one misclassified as P. palmata), 94.3% for P. barbata

(with one misclassified as P. palmata) and 100% for P.

breviuscula. Separation of the four species of Pseudophycis 1s

shown in fig. 5, with some overlap of P. palmata and P.

bachus. SMNS 1589 (holotype of P. palmata) 1s grouped

within the other specimens of P. palmata and 1s well separated

from P. barbata. Consequently, the CAP analysis and fig. 5

support the taxonomic decision to resurrect Pseudophycis

palmata as the appropriate name for the Australian endemic

previously thought to be conspecific with P. bachus. Overlap

of P. palmata and P. bachus 1s consistent with the historical

confusion involving the two species.

Despite its close relationship with P. bachus, the first

known detailed description of this species appears to be that

of McCoy (1878) who based his description of P. "barbatus"

almost entirely on specimens of the Australian P. bachus-like

species. That species was probably the common representative

of the genus in the Melbourne markets at the time. Meristic

and morphometric data presented by McCoy appear to be

entirely attributable to specimens of P. palmata, with only the

illustration (McCoy, 1878: pl. 20) based on a specimen of P.

barbata. Although several of McCoy's specimens were either

lost or obscured by early collection practices, NMV A23366-

001 is likely to be the specimen featured in the illustration,

while counts and measurements were probably taken from

NMV 43104, 43105, A841 and A23366-002. Other old NMV

collection specimens of the new species that lack documented

provenance probably comprised the remaining three.

Considering the frequent occurrence of this species at the

type locality of P. palmata, together with the strong

morphological support discussed above, we consider the name

to be applicable to the Australian P. bachus-like cognate.

Paulin (1983) failed to deal with the name in the synonymies

of the three species he recognised.

This species has the most restricted range of Australian

Pseudophycis species, with no records of it west of Spencer

Gulf, South Australia. Its latitudinal limits approach those of

P. bachus in New Zealand, as do those of the two species

occurring in both Australia and New Zealand where

bathymetrically feasible.

Material examined. Type. Physiculus palmatus SMNS 1589 (172,

holotype) Port Phillip, Hobsons Bay (northernmost section of Port

Phillip Bay immediately south of Melbourne), Victoria.

Other material. (51 non-type specimens examined for meristic or

morphometric values, 90.1—380 mm SL; see Appendix 2 for additional

material in Australasian collections.) Australia, New South Wales:

AMS 1.34462-004 (2, 230—233) north-east of Lookout Point, Twofold

Bay, 37° 4.2'S, 149° 56.1' E, 100 m, J.K. Lowry and S.J. Keable, 26—27

November 1988; AMS 1.34567-001 (146) middle of Long Beach,

Australasian red cods

Batemans Bay, 35° 42'S, 150° 13' E, 50 m, J.K. Lowry and S.J. Keable,

23—24 November 1988; AMS 1I.34569-003 (112) east of Lookout Point,

Twofold Bay, 37° 46'S, 149° 56' E, 50 m, J.K. Lowry and S.J. Keable,

26—27 November 1988; AMS 1.34570-001 (3, 117—190) east of

Lookout Point, Twofold Bay, 37° 46' S, 149° 55' E, 50 m, 26-27

November 1988. Victoria: NMV A840-001 (2, 95.5211) old

collection, no data; NMV A840-002 (2, 198-202) old collection, no

data; NMV A3859 (187) eastern Bass Strait, 6 km west-south-west of

Cape Conran, 37° 49.8' S, 148° 40' E, 26 m, BSS 208 T, M.F. Gomon

and R.S. Wilson, 30 July 1983; NMV AS$8870-001 (4, 95.7—113) and

NMV A$870-002 (121) Bass Strait, 1 km off Lake Tyers, 37° 51.1' S,

148° 9' E, 15 m, Marine Science Laboratories, 5 June 1984. Tasmania:

AMS I.23880-004 (192) north of George Town, 40 57.90' S, 146 46.20'

E, 50 m, K. Graham and FRV Kapala, 13 July 1980; AMS I.34952-003

(258) mouth of Fortescue Bay, 43° 7.77' S, 147° 59.47' E, 50 m, J.K.

Lowry and K. Dempsey, trawl, 9-10 April 1994; CSIRO H 4229-01'

(380) south of Port Arthur, 43° 17.3' S, 147? 47.1' E-43? 16.5' S, 147°

49.6' E, 115—119 m, B. Evans, 27 May 1996; CSIRO H 6205-02 (136)

Battery Point, CSIRO wharf, 42° 53' S, 147° 20' E, H. Motomura, 20

February 2005; CSIRO H 7366-01' (265) Storm Bay, east of Variety

Bay on North Bruny Island, 43° 12.17' S, 147° 27.46' E, 40—45 m, hook

and line, A. Pender, 6 May 2012; CSIRO H 7716-01' (350) Munro

Bight, 43° 11'S, 147° 59' E, 22 m, A. Pender , January 2012; CSIRO H

7T117-01* (362) Hinsby Beach, Taroona, 42? 57.22' S, 147? 20.82' E, 2

m, A. Pender, February 2012; CSIRO T 1186-02" (8, 110-262) Nutgrove

Beach, Sandy Bay, Derwent River, P.R. Last, 19 May 1980; CSIRO T

1417 (113 mm SL) Derwent Estuary, 7 m, P.R. Last; NMNZ P.024340

(4: 142—197) Nakyrare Beach, Derwent Estuary, 43° 3.000' S, 147°

22.000' E, FV Ophelia, 29 March 1988; NMV A 1218-001 (316),

NMV A 1218-002 (318) central Bass Strait, 20 km north-north-east of

North Point, 40? 31.8' S, 145° 22.8' E, 44 m, BSS 116 T, M.F. Gomon,

G.C.B. Poore and P. Forsyth, 4 November 1980; NMV A1275 (378)

central Bass Strait, 23 km east of Cape Rochon, Three Hummock

Island, 40° 22.8' S, 145° 16.998' E, 40 m, BSS 112 T, M.F. Gomon,

G.C.B. Poore and P. Forsyth, 3 November 1980; NMV A1528-001 (6,

73.5—161) and NMV A1528-002 (3, 93.6—151) central Bass Strait, 30

km north of Wynyard, 40° 33.07' S, 145° 44.69' E, 67.7 m, 4 February

1981; NMV A 31132-001* (318) off Taroona between Taroona and

Alum Cliffs, 42? 57.470' S, 147? 20.797' E, 6 m, hook and line, B.

Barlow, 10 May 2014.

Pseudophycis bachus (Forster in Bloch and Schneider,

1801)

Common name: New Zealand Red Cod (new Australian name);

red cod, hoka (New Zealand)

Figures 1, 3A, 5, 6; Tables 1—4

Enchelyopus bachus Forster 1n Bloch and Schneider, 1801: 53.

Type locality: Queen Charlotte Sound, New Zealand. No types known.

Based on manuscript description of Gadus bacchus by Forster.

Gadus bacchus. Cuvier, 1817: 486; Forster in Lichtenstein, 1844:

120, 420; Whitehead, 1978: 40; Hoare, 1982: 269.

Lota bacchus Cuvier, 1829: 334; Richardson, 1846: 61.

Brosmius venustus Richardson and Gray, 1843: 222 (generic

referral of Parkinson's unpublished manuscript name Blennius

venustus, written on the painting used by Forster for his description of

Gadus bacchus). Richardson, 1843: 27; Taylor, 1855: 413.

Enchelyopus bacchus. Lichtenstein, 1844: 419 (1ndex only).

Lotella bacchus. Günther, 1862; 347; Hutton and Hector, 1872: 46,

115; Hector, 1875: 239; Hector, 1884: 55; Hector, 1886: 28; Hutton,

1875: 134; Thomson, 1877: 485; Thomson, 1878: 326; Thomson, 1879:

382; Dambeck, 1879: 536, 547, 555; Parker, 1882: 263; Parker, 1883:

[à

234, 235, pl. 33; Sherrin, 1886: 16, 17, 93, 304; Thomson, 1890: 370, pl.

28; Beattie, 1891: 71,81, 82, pl. 12, pl. 13, pl. 14, pl. 15; Thomson, 1892:

212; Ayson, 1900: 14; Mair, 1903: 319; Ayson, 1907: 22; Johnson,

1921: 473; Carter and Malcolm, 1926: 647; Malcolm, 1926: 658;

Svetovidov, 1937: 1285; Svetovidov, 1948: 17, 60 (anatomy).

Pseudophycis bacchus. Günther, 1880a: 26 (Port Hardy, D'Urville

Island); Murray, 1895: 599 Gill, 1893: 94, 95 100, 120 (list); Karrer,

1971: 153, 179, 180, 185, 195; Habib, 1975: 1; Ayling and Cox, 1982:

142, pl 9. (description).

Pseudophycis bachus. Günther, 1880b: 542—543, fig. 248; Ogilby,

1886: 48 (list, in part); Hutton, 1890: 282; Fitch, 1972: 570, 573;

Marshall and Cohen, 1973: 490; Edgar et al., 1982: 32, fig. 17

(description, in part); Paulin, 1983: 91 (description, taxonomy, in part);

Paulin and Stewart, 1985: 22; Paul, 1986: 57, fig. (1n part); Francis

1988: 21, pl. 18; Paulin, 1988: 450, 451, 453; Paxton and Hanley in

Paxton et al., 1989: 302 (list, in part); Paulin et al., 1989: 119 (keyed),

255 (list); Cohen in Cohen et al., 1990: 373 (taxonomy, in part); Paulin

in Amaoka et al., 1990: 155, fig. 103 (description, in part); Paulin and

Roberts, 1992: 130, fig. 62a (description, 1n part); Kuiter, 1993: 59

(description, in part); Gomon in Gomon et al., 1994: 333 (description,

in part); Francis, 1996: 20, pl. 18; Horn, 1996: 151, 158; Kuiter, 1997:

50 (description, in part); Paulin, 1998: 52, fig. (description); Paul,

2000: 57 (description); Francis, 2001: 25, pl. 19; Beentjes and Renwick,

2001: 315, 316; Paxton et al., 2006: 616 (taxonomy, in part); Hirt-

Chabbert, 2006: 39; Gomon in Gomon et al., 2008: 313 (description, in

part); Roberts et al., in Gordon et al., 2009: 532 (listed); McMillan et

al., 2011a: 161, 162; McMillan et al., 2011b: 94; Francis, 2012: 5; Horn

et al., 2012: 624, 625, 627—629, 631—633; Roberts et al., 2014: 18

(listed); Struthers et al., in Roberts et al., 2015: 864, fig. 107.20

(description); Roberts et al., 2015: S164 (listed); Roberts et al., 2017:

8] (listed), Roberts et al., 2019: 90 (listed).

Physiculus bacchus. Günther, 1887: 87; Goode and Bean, 1895:

365, 549; Hutton, 1896: 316; Hutton, 1904: 48 (listed); Thomson, 1906:

551; Thomson and Anderton, 1921: 74; Thomson and Thomson, 1923:

111; Frost, 1924: 609; Frost, 1926: 488, 490; Young, 1925: 370; Archey

in Speight, Wall and Laing, 1927: 203; Anonymous, 1931: 32; Frost,

1933: 140; Benham, 1934: 31; Benham, 1935: 22; Benham, 1938: 56;

Graham, 1938: 405; Graham, 1939: 364; Doogue and Moreland, 1960:

197, 288; Doogue and Moreland, 1961: 208, 316; Moreland, 1963: 20;

Webb, 1966: 52, fig. 2.8, 70, 128, table 3.4, 164, 209, 230, 231, 234,

238—240, 257, 263, 266, 280, 294b, 294c; Webb, 1972b: 43; Webb,

1973: 307—309; Heath and Moreland, 1967: 37, 56; Whitehead, 1969:

pl. 11; Anonymous, 1971: 17; Anonymous, 1972c: 47; Watkinson and

Smith, 1972: 31; Knox and Kilner, 1973: 354; Vooren, 1974: 43, 44.

Physiculus bachus. Stead, 1906: 86 (in part); Waite, 1907: 18

(listed); Zietz, 1909: 266; Waite, 1909: 51, 52, 57, 134; Waite, 1911:

162, 183, 259, 265, 2770, pl. 31; Thomson, 1913: 233; Phillipps, 1918:

271; Phillipps, 1921: 121, 125; Phillipps and Hodgkinson, 1922: 95;

Waite, 1923: 92 (1n part); Ayson, 1924: 7; Lord and Scott, 1924: 8, 43;

Phillipps, 1926: 528; Lord, 1927: 13; Phillipps, 1927a: 128; Phillipps,

1927b: 23, 60; Phillipps, 1927c: 12; Waite, 1928: 6 (listed); McCulloch,

1929: 129 (in part); Young, 1929: 141; Anonymous, 1930: 28; Finlay,

1930: 47; Anonymous, 1934: 43; Anonymous, 1935: 34; Norman,

1935: 3; Benham, 1936: 26; Hefford, 1936: 71, 74; Cunningham, 1937:

898, Norman, 1937: 54, 55 (in part, listed); Shorland, 1937: 223;

Wilson, 1937: 31; Johnston, 1938: 47; Munro, 1938: 62; Graham,

1939b: 364; Fowler, 1940: 758; Rapson, 1940: 35; Phillipps, 1947: 42;

Phillipps, 1948: 129; Shorland, 1948: 109; Laird, 1949: 14, 19, 36, 37,

39, 53, 56, 60, 6l, 137, 145, 146; Phillipps, 1949: 24, 59; Shorland,

1950: 35; Laird, 1951: 287, 298, 306, 308; Laird, 1952: 589, 590, 595,

596, 600; Graham, 1953: 166, 173, 399; Manter, 1954. 498, 545, 547,

549, 559; Robinson, 1955: 10, 71, 105, pls 9 and 14; Anonymous, 1957:

69; Kaberry, 1957: 90; Moreland, 1957: 34, 36; Parrott, 1957: 47, 175;

Anonymous, 1958: 73; Parrott, 1958: 117; Anonymous, 1959: 70;

74 M.F. Gomon, C.D. Struthers & J. Kemp

-> * - e - -

m. "u^ |

il » oa € Ys “a

— ` è - ` ^ " T i

| Qi +. a p

= 9 ~~ E

= ww. Da <

Figure 6. Pseudophycis bachus. A, B, NMNZ P.054828, neotype, 351 mm SL, 2.5 km off coast from Whanganui River bar, Whanganui, New

Zealand, fresh specimen, lateral view of body and anterior body showing black pectoral blotch respectively (photographs C. Struthers, NMNZ);

C, unpublished line drawing by S. Parkinson of Enchelyopus bachus Forster in Bloch and Schneider, 1801 (reproduced from Whitehead, 1969: pl. 31).

Australasian red cods

Robinson, 1959: 152; Anonymous, 1960: 70; Parrott, 1960: 67, 164;

Meglitsch, 1960: 321—323; Scott, 1962: 84 (in part); Gorman, 1963: 29;

Graham, 1963: 167; Doogue and Moreland, 1964: 205, 311; Street,

1964: 18; Anonymous, 1965: 15, 22; Churchman, 1965: 56; Doogue

and Moreland, 1966: 205, 311; Elder, 1966: 96, 97; Howell, 1966: 33;

Paul, 1966: 372, 373; Svetovidov, 1967: 1685, 1686, 1689, 1690;

Sorensen, 1968: 148; Tong and Elder, 1968: 64; Doogue and Moreland,

1969: 205, 311; Russell, 1969: 108; Cowper, 1970: 45; Godfriaux,

1970: 257; Iwai et al., 1970: 21; Shuntov, 1970: 373, 374, 376; Sorensen,

1970: 4, 17; Coakley, 1971: 24; Russell, 1971a: 9, 19, 21, fig. 3, 38, 41,

94, 173, table 10, fig. 13, 191, 197; Russell, 1971b: 83; Anonymous,

1972a: 229; Anonymous, 1972b: 187—189, 278; Doak, 1972: 19, 101,

131; Hewitt and Hine, 1972: 92; Iwai et al., 1972: 29, 36; Shuntov,

1972: 339; Walker, 1972: 2; Webb, 1972a: 16; Suda, 1973: 2150-2152;

Waugh in Williams, 1973: 257, 274; Williams, 1973: 427; Godfriaux,

1974: 502; Ryan, 1974: 133, 135; Scott et al., 1974: 95, 96 (in part).

Lotella bachus. Ayson, 1908: 28; Rendahl, 1926: 2.

Physiculus backus. Fowler, 1940: 758 (near Loba, New Zealand).

Physiculus (Pseudophycis) bachus. Graham, 1956: 166; Whitley,

1956: 403; Whitley, 1968: 40 (list).

Pscudophycis bachus. Doogue and Moreland, 1982: 218, fig.

(misspelling).

Diagnosis. First dorsal fin 11—13 rays, second dorsal fin 40—45

rays; anal fin 42—48 rays; total vertebrae 44—46; scales above

lateral line 1n oblique series from base of first dorsal fin ray

10-12; oblique rows of scales intersecting with lateral line

102-136; pyloric caeca 6; caudal peduncle moderately short

28.0—38.9% HL; barbel short, 8.6—16.9% HL; caudal fin

truncate with angular corners in specimens larger than about

150 mm SL, middle rays shorter than rays extending to corners;

dark blotch basally on pectoral fin extending well onto side

above fleshy pectoral fin base; underside of head and body

whitish; anal fin mostly whitish with faint narrow black distal

margin only posteriorly, if at all. A large species reaching at

least 650 mm SL.

Description. (Values for neotype with those for non-type

specimens when different in parentheses; see Tables 2—4 for

summary of selected meristic and comparative morphometric

values.) First dorsal fin 11 (11—13, rarely 13, first ray usually

minute); second dorsal fin 42 (40—45, usually 42 or 43); anal fin

45 (42—48, usually 45—47); pectoral fin 24 (22-25, rarely 22);

pelvic fin 5; caudal fin 36 (33-39); gill rakers 4 + 10 (4—5 +

10-12 = 14—16); lateral line pores not associated with individual

scales; oblique scale rows intersecting with lateral line 112

(102—136); scales 1n oblique series above lateral line 11 (10—

12), scales in oblique series below lateral line 30 (26—34, rarely

33 or 34); vertebrae 16 + 29 (16—17 + 27—30 = 44—46); pyloric

caeca 6.

Body moderately slender, compressed laterally (fig. 6),

greatest depth at anal fin origin 19.5 (18.4—23.7)% SL,

tapering uniformly from second dorsal fin origin to shallow

caudal peduncle; caudal peduncle short, 8.5 (7.7—10.5)% SL,

strongly compressed, depth less than orbital diameter.

Distance between middle of anus and base of anal fin less

than suborbital depth. Body cavity extending posteriorly to

above origin of anal fin.

Head acute, of moderate size, length 26.5 (25.9—29.6)96

SL, its width 14.4 (13.4-17.7)% SL and depth 11.7 (10.2—

15.6)% SL; snout of moderate length 7.5 (7.1-9.0)% SL, not

projecting in advance of upper jaw, rounded in dorsal view.

Nostrils small, located about two-thirds to three-quarters of

the way from snout tip to eye, distance from nostril to orbital

margin equal to or greater than diameter of combined nostrils;

nostrils positioned above horizontal through centre of eye;

posterior opening larger than anterior, separated from it by

raised skin flap; posterior opening with tubular flap encircling

margin. Interorbital of moderate width, very slightly convex.

Eye of moderate size, orbital diameter 21.1 (17.0—30.4)% HL,

1.58 (0.98—2.00) times in interorbital space, 2.20 (1.50—4.25)

times suborbital distance, circular, upper edge of eye adjacent

to dorsal margin of head in lateral view, transparent skin

covering eye faintly pigmented near orbital dorsal margin.

Postorbital moderately long, 1.01 (0.89—1.22) times length of

upper jaw. Mouth large, terminal, upper jaw terminating at

vertical through posterior margin of eye. Jaw teeth caniniform,

slightly curved, depressible, band of up to four or five irregular

rows in upper jaw with distinct hiatus at symphysis; teeth in

outer row more regularly arranged than in inner rows; band

tapering near rear of jaw. Teeth of lower jaw similar in form

to those of upper jaw; band of two or three rows with broader

patch on either side of symphysis, tapering to single, widely

spaced row posteriorly; rows almost contiguous across

symphysis. Vomerine teeth absent. Opercular bones strong;

upper extremity of gill opening at horizontal through middle

of eye; gill membranes continuous across isthmus. Gill rakers

on outer arch slender, of moderate length, almost as long near

angle as opposing gill filaments, 12.8 (9.5—12.8) times in

head, their inner margin denticulate. Chin barbel short,

subconical 12.0 (8.6—16.9)96 HL.

small cycloid scales covering all of head, including

isthmus and gular region, and body except for branchiostegal

membranes, surface of maxilla and premaxilla, lower lip and

distal parts of fins; thick mucus covering obscuring scales and

pores in freshly preserved material. Most head pores tiny,

following main cephalic sensory canals; row of slightly raised

pores from nostrils to tip of snout and then posteriorly just

above lower edge of suborbital; row of enlarged mandibular

pores on underside of lower jaw. Lateral line comprising

widely spaced pores on short tubes arising from narrow scale-

less gap, anterior end curved upwards slightly, then gradually

descending to lateral midline below posterior 25% of second

dorsal fin, remaining on lateral midline posteriorly.

Membranes of first and second dorsal fins continuous at

base; first dorsal originating vertically above pectoral fin

base; anterior two-thirds of second dorsal of uniform height,

7th to 9th ray from posterior end of fin longest, its height

about 1.4 times length of 2nd ray at anterior end of fin,

subsequent rays progressively shorter; rays of both dorsal fins

mostly unbranched, only last 13 to 16 rays of second dorsal,

apart from last one to three, branched. Fleshy, fine scale-

covered basal sheath on third or more of first dorsal and

anterior portion of second dorsal fins; fin rays interconnected

by membranes to tips, sheath gradually decreasing 1n coverage

and thickness posteriorly, encompassing about half of second

dorsal fin near its insertion; sheath extending onto body for

anterior third of combined dorsal fin base, broadest anteriorly,

narrowing posteriorly. Profile of anal fin like that of second

dorsal fin, although shallower posteriorly, with comparable

unbranched and branched rays. Likewise, anal fin enclosed in

broad fleshy sheath that 1s like that of dorsal fins; sheath also

extending onto body for anterior third or less of fin. Caudal fin

truncate with distinct dorsoposterior and ventroposterior

corners in adults, posterior margin with little if any convex

curve; longest rays to corners, middle rays about 85—90%

length of longest rays; fin more rounded in juveniles; base of

fin covered by indistinct sheath sharply demarcated from

scales of caudal peduncle. Pectoral fin tip variably reaching

almost to, to or past vertical through anal fin origin, sixth or

seventh ray longest. Pelvic fin inserted anterior to vertical

through posterior edge of preopercle (more posteriorly in

some specimens); outer two rays longer than inner rays;

second ray longest, 16.0 (11.5—21.1)% SL, nearly twice length

of subsequent ray, reaching past a vertical through first dorsal

fin origin.

Fresh colour. (Based on images of neotype; fig. 6A, B.) Medium

brown above extending ventrally to level of ventral edge of

pectoral fin base, white below, suffused with pinkish hue,

especially above anal fin base; lateral line slightly paler at least

anteriorly. Underside of head, jaws and barbel white, tinged

with pink. Dorsal and caudal fins medium brown, caudal paler

with reddish tinge ventrally; anal fin white with pink hue;

dorsal fins with fine black distal margin; caudal fin margin with

broader, more diffuse black margin; anal fin with faint black

distal margin, darkest posteriorly. Pectoral fin medium brown

with semi-circular black basal spot covering dorsal half of fin

edge proximally, extending well onto side of body above fin

base; ventroposterior margin of fin pale to translucent. Pelvic

fin rays white with pink hue.

Preserved colour. Upper half of head and body pale dusky

to dusky, lower half very pale, frequently pearly white.

Dorsal and caudal fins pale dusky; distal edge of dorsal fin with

fine dark margin; posterior edge of caudal fin broadly dark;

anal fin very pale, distal edge of fin posteriorly with fine dark

margin in some; pelvic fin, pectoral fin and basal half of anal

fin very pale; pectoral fin with prominent dark spot covering

dorsal half or more of basal edge that extends onto side above

pectoral fin base.

Etymology. The name bachus is an altered spelling of bacchus,

the Greek god of wine, adopted for the wine-red colouration

assumed shortly after capture but lost soon after death.

Distribution. Endemic to New Zealand, occurring around both

the North and South Islands, and reaching the Chatham Islands

in the east (fig. 3A). A demersal species on soft bottom at 2—570

m depth.

Remarks. Pseudophycis bachus was originally described as

Enchelyopus bachus Forster in Bloch and Schneider, 1801,

based on Forster’s manuscript description of “Gadus bachus",

subsequently published by Lichenstein (1844: 120). Forster’s

drawing No. 180, on the back of which is pencilled “ex Queen

Charlotte Sound”, is evidently an illustration of the fish on

which the description was based. Although both Bloch and

Schneider (1801: 53) and Lichtenstein gave the origin of the

M.F. Gomon, C.D. Struthers & J. Kemp

Specimen simply as “in the seas around New Zealand”,

Whitehead (1969: pl. 11), Paulin (1983: 91) and Fricke et al.

(2019) regarded Queen Charlotte Sound, the presumed

collection locality of the specimen implied in the note on

Forster’s illustration, as the type locality. However, Hoare’s

(1982: 269) reproduction of Forster’s Resolution Journal has a

footnote stating “Gadus bacchus of Descr. Anim., p. 120 and

Forster drawing 180 (undated). The description is from a

specimen of 22 inches (559 mm), dated 13 April 1773”. That

date puts the type locality as Dusky Sound from a larger fish.

Paulin (1983: 92) provided a detailed discussion of the origin

and various iterations of Bloch and Schneider’s name.

An unfinished drawing by Parkinson (vol. 2, no. 5, ex

Totarra ‘nue, 404 mm tot. 1.) reproduced by Whitehead (1969:

pl. 11; here as fig. 6C), has “18, Blennius venustus” pencilled

on the reverse side and 1s the basis for Richardson and Gray’s

(1843: 222) Brosmius venustus. The fish figured is clearly an

example of P. bachus, despite the incomplete nature of the

illustration. Totaranui 1s the Maori name for Queen Charlotte

sound. The scientific name is regarded as unavailable because

no distinguishing features were provided by the authors

(Fricke et al., 2019). The illustrated characters agree with

those of specimens from New Zealand identified as P. bachus.

Because no type specimen of P. bachus 1s known (Fricke

et al., 2019), the designation of a neotype for P. bachus 1s

considered justified. A specimen (NMNZ P.054828, 351 mm

SL; fig. 6a) collected from 2.5 km off the coast from

Whanganui River bar, Whanganui, New Zealand, is proposed

as the neotype for P. bachus. Accordingly, Whanganui, New

Zealand, becomes the type locality of P. bachus under Article

76.3 (ICZN, 1999), replacing the original type locality Queen

Charlotte Sound, New Zealand.

Material examined. Neotype. NMNZ P.054828' (351) 2.5 km off the

coast from Whanganui River bar, Whanganui, New Zealand, 39° 53'S,

174° 49' E, 18 m, C. Papple, 10 October 2012.

Other material. (38 non-type specimens examined for meristic or

morphometric values, 110—539 mm SL; see Appendix 2 for additional

material in Australasian collections.) New Zealand, North Island:

NMNZ P.001718 (539) off Castlepoint, Wairarapa, 40° 54' S, 176° 13'

E, 73-110 m, F. Abernethy, 8 July 1955; NMNZ P.033899 (5, 121—202)

off Raglan, Waikato, 37° 48.33' S, 174° 34.00' E, RV Kaharoa, 61—63

m, 26 October 1996. South Island: NMNZ P.025037 (2, 479—512)

Blueskin Bay, Otago Peninsula, Otago, 45° 43.28' S, 170° 40.33' E,

20-22 m, C.D. Roberts and C.D. Paulin, 9 May 1990; NMNZ P.025129

(4, 137—192) Entrance mole, Otago Harbour, Otago, 45° 46.45' S, 170°

43.23' E, 6—9 m, C.D Roberts, 9 May 1990; NMNZ P.027580 (239)

Harrold's Bay, Halfmoon Bay, Stewart Island, 46° 53.75' S, 168° 9.25'

E, 2-5 m, NMNZ Stewart Island field team, 4 March 1992; NMNZ

P.032385 (2, 328—395) Mooring, head of Gold Arm, Charles Sound,

Fiordland, 45° 8.55' S, 167° 8.78' E, 5-10 m, NMNZ Fiordland 1995

field team, 24 March 1995; NMNZ P.032393 (3, 292—309) Mooring at

Toe Cove, head of Nancy Sound, Fiordland, 45° 10.55' S, 167° 8.85' E,

25 m, NMNZ Fiordland 1995 field team, 26 March 1995; NMNZ

P.03599] (388) Flowerpot Rock, Jackson's Bay, West Coast, 43° 58.03'

S, 168° 37.30' E, 5-8 m, NMNZ Jackson-Haast 1999 field team, 7

February 1999; NMNZ P.044338' (136) south of Timaru, Canterbury,

44° 45.04' S, 171° 18.40' E, 26-31 m, RV Kaharoa, 3 June 2007;

NMNZ P.044339' (176) south off Timaru, Canterbury, 44° 45.04' S,

171° 18.40' E, 26-31 m, RV Kaharoa, 3 June 2007; NMNZ P. 047709

(229) Lyttelton Port, Canterbury, 43° 36.33' S, 172° 43.10' E, NIWA

Australasian red cods

Port Survey, 4 November 2004; NMNZ P.049392 (343) c. 7 km east-

north-east of Kaikoura Peninsula, 42° 24.77' S, 173° 48.22' E, 93 m,

C.D. Struthers, 16 November 2010; NMNZ P.049401 (370) c. 6 km

north-east of Kaikoura Peninsula, 42° 23.60' S, 173° 47.59' E, 49 m,

C.D. Struthers, 16 November 2010; NMNZ P.049402 (545) c. 5 km

east-south-east of Kaikoura Peninsula, 42° 26.77' S, 173° 46.39' E, 94

m, C.D. Struthers, 17 November 2010; NMNZ P.049403 (519) c. 5 km

south off Kaikoura Peninsula, 42 28.09' S, 173 43.02' E, 91 m, C.D.

Struthers, 19 November 2010; NMNZ P.049406 (485) and NMNZ

P.049407 (467) c. 3 km south off Kaikoura Peninsula, 42° 24.72' S,

173° 45.22' E, 47 m, C.D. Struthers, 17 November 2010; NMNZ

P.049678 (358) Bay View, end of Franklin Road, Hawke's Bay, 10—15 m,

C.D. Struthers and D.H. Struthers, 27 December 2010; NMNZ

P.052508 (297) and NMNZ P.052509 (290) south of Pegasus Canyon

mouth, Canterbury Bight, 43° 33.99' S, 173° 33.44' E, 86 m, RV

Tangaroa, 13 May 2011; NMNZ P.053748' (383), NMNZ P.053749'

(405), NMNZ P.053750' (368), NMNZ P.053751' (373) and NMNZ

P.053752' (337) Port Pegasus; Stewart Island, 47? 13.20' S, 167? 41.38'

E, 64 m, C.D. Struthers, 20 February 2012. Chatham Rise: NMNZ

P.044348' (273) central Chatham Rise, 43? 47.42' S, 179? 30.01' W,

327—328 m, RV Tangaroa, 31 December 2007; NMNZ P.044349'

(382) central Chatham Rise, 43° 47.42' S, 179° 30.01' W, 327—328 m,

RV Tangaroa, 31 December 2007; NMNZ P.044350' (297) central

Chatham Rise, 43° 43.78' S, 179° 24.00' W, 367—375 m, RV Tangaroa,

3] December 2007.

Pseudophycis barbata Günther 1862

Common name: Bearded Rock Cod (Australian Standard);

southern bastard cod (New Zealand)

Figures 1, 3B, 4, 5, 7; Tables 1—4

Pseudophycis barbatus Günther, 1863: 116. Type locality:

Victoria, South Australia. Holotype: BMNH 1863.1.15.38 (skin in

alcohol).

Pseudophycis barbatus. McCoy, 1878: 29, pl. 20 (in part,

description); Johnston, 1883: 126 (description); Ayling and Cox, 1982:

144, fig. (description).

Lotella grandis Ramsay, 1881: 462. Type locality: Wollongong,

New South Wales. Holotype: AMS I.696 (decision by Paulin, 1983: 94).

Lotella grandis. Steindachner, 1901: 509 (description).

Physiculus barbatus. Zietz 1909: 266; Waite, 1923: 91, fig.

(description); Lord and Scott, 1924: 8, 43 (description); McCulloch,

1927: 32 (listed); Waite, 1928: 6 (listed); McCulloch, 1929: 128 (list);

Norman, 1937: 55 (listed); Munro, 1938: 62, fig. 440 (description);

Whitley, 1955: 119 (taxonomy); Scott, 1962: 84, fig. (description);

Whitley, 1964: 40 (list); Svetovidov, 1967: 1686; Scott et al., 1974: 95,

fig. (description).

?Pseudophycis breviusculus (nec Richardson, 1846) Graham,

1939: 405 (after Paulin, 1983: 93).

Pseudophycis breviusculus (nec Richardson 1846). Habib, 1975:

32—57.

Physiculus (Pseudophycis) breviculus (nec Richardson, 1846).

Graham, 1956: 173 (in part, locality implies multiple species;

misspelling).

Physiculus (Pseudophycis) breviusculus (nec Richardson, 1846)

Whitely, 1956: 403 (in part?).

Pseudophycis barbata. Paulin, 1983: 94, fig. 10 (description, new

record for NZ); Paulin and Stewart, 1985: 22; Hutchins and Swainston,

1986: 34 124, fig. 114 (description); May and Maxwell, 1986: 195

(description); Paul, 1986: 58, fig; Francis, 1988: 21, pl. 19; Paulin,

1988: 453; Paxton and Hanley in Paxton et al., 1989: 302 (list); Paulin

et al., 1989: 119 (key), 255 (listed); Cohen in Cohen et al., 1990: 374

(taxonomy); Paulin in Amaoka et al., 1990: 156 (description); Kuiter,

1993: 59 (description); Gomon in Gomon et al., 1994: 334 (description);

Francis, 1996: 21, pl. 19; Horn, 1996: 157; Kuiter, 1997: 50 (description);

Paulin, 1998: 67, fig. (description); Paul, 2000: 58; Francis, 2001: 25,

pl. 18; Hutchins, 2001: 23 (description); Paxton et al., 2006: 616 (list);

Hirt-Chabbert, 2006: 40; Gomon in Gomon et al., 2008: 314

(description); Roberts et al. in Gordon et al., 2009: 532 (listed);

McMillan et al., 201 1a: 161, 162; McMillan et al., 2011b: 94; Francis,

2012: 52; Roberts et al., 2014: 18 (list); Struthers et al. 1n Roberts et al.,

2015: 863, fig. 107.21 (description); Roberts et al., 2015: S164; Roberts

et al., 2017: 81 (list), Roberts et al., 2019: 90 (listed).

Diagnosis. First dorsal fin with. 10—11 rays, second dorsal fin

with 54—59 rays; anal fin with 54—63 rays; total vertebrae 48—

51; scales above lateral line in oblique series from base of first

dorsal fin ray 16—22; oblique rows of scales intersecting with

lateral line 123—162; pyloric caeca 14—20; caudal peduncle

short 13.5-18.6% HL; chin barbel of moderate length, 16.5—

35.1% HL; caudal fin rounded without angular corners, the

middle rays equal to or longer than rays above and below; no

distinct, dark blotch basally on pectoral fin although fin base

sometimes overall dark; underside of head and body tan to

brown; anal fin tan to brown with distinct black distal margin.

A large species reaching at least 620 mm SL.

Description. (Values for non-type specimens when different

from type in parentheses; see Tables 2—4 for summary of

selected meristic and comparative morphometric values.) First

dorsal fin 10 (10—11), first ray small to minute; second dorsal fin

59 (54-59, rarely 59); anal fin 59 (54—63); pectoral fin 25 (24—

27, rarely 24); pelvic fin 5; caudal fin 33 (32—35 rays); gill rakers

unknown (3—4 + 8-11 = 12-14); lateral line pores not associated

with individual scales; oblique scale rows intersecting with

lateral line unknown (123-162); scales in oblique series above

lateral line unknown (16—22), scales in oblique series below

lateral line unknown (38—56, rarely less than 45); vertebrae

unknown (14—16 + 33-37, rarely 37 = 48—51); pyloric caeca

unknown (14—20).

Body of moderate depth, moderately compressed laterally

(fig. 7b), greatest depth at anal fin origin unknown (19.7—

29.3)% SL, tapering uniformly from second dorsal fin origin

to shallow caudal peduncle; caudal peduncle short, 5.4 (5.2—

9.2)70 SL, strongly compressed, depth subequal to orbital

diameter. Distance between middle of anus and base of anal

fin slightly less than suborbital depth. Body cavity extending

to above origin of anal fin.

Head acute, of moderate size, length 23.3 (23.3—31.6)% SL,

width unknown (14.9—19.8)76 SL and depth unknown (11.0—

17.9)% SL; snout of moderate length 7.5 (6.5—9.6)96 SL, not

projecting in advance of upper jaw, rounded in dorsal view.

Nostrils small, located about two-thirds to three-quarters of the

way from snout tip to eye, distance from nostril to orbital

margin about equal to or greater than diameter of combined

nostrils; nostrils positioned above horizontal through centre of

eye, both with low tubular rim; posterior opening about half

size of anterior, separated from it by raised skin flap. Interorbital

of moderate width, very slightly convex. Eye of moderate size,

orbital diameter 20.8 (16.8—28.5)% HL, unknown (0.91—1.68)

times in interorbital space, unknown (1.46—4.71) times

suborbital distance, circular, upper edge of eye adjacent to

M.F. Gomon, C.D. Struthers & J. Kemp

50mm

50 mm

Figure 7. Pseudophycis barbata. A, NMNZ P.037124, 450 mm SL, Bay of Plenty, east of north-east point, North Island, New Zealand (photograph

C. Struthers, NMNZ); B, BMNH 1863.1.15.38, holotype of Pseudophycis barbata, 394 mm SL, Victoria, South Australia (photograph © The

Trustees of the Natural History Museum, London).

dorsal margin of head in lateral view. Postorbital moderately

long, 1.00 (0.52-1.21) times length of upper jaw. Mouth large,

terminal, upper jaw terminating at vertical through posterior

margin of eye. Jaw teeth caniniform, slightly curved,

depressible, band of up to four or five irregular rows in upper

jaw with distinct hiatus at symphysis; teeth in outer row more

regularly arranged than in inner rows; band tapering near rear

of jaw. Teeth of lower jaw similar in form to those of upper jaw;

band of two or three rows with broader patch on either side of

symphysis, tapering to single, widely spaced row posteriorly;

rows almost contiguous across symphysis. Vomerine teeth

absent. Opercular bones strong; upper extremity of gill opening

at horizontal through middle of eye; gill membranes continuous

across isthmus. Gill rakers on outer arch slender, moderately

short, unknown (about 0.6—0.8) times length of opposing gill

filaments near angle, unknown (19—22) times in head length,

slightly denticulate. Chin barbel of moderate length, subconical

21.1 (16.5—25.3)96 HL.

Very small cycloid scales covering all of head and body

except for branchiostegal membranes, surface of maxilla and

premaxilla, lower lip and distal parts of fins; thick mucus

covering obscuring scales and pores in freshly preserved

material. Most head pores tiny, following main cephalic

sensory canals; row of slightly raised pores from nostrils to

tip of snout and then posteriorly just above lower edge of

suborbital; row of enlarged mandibular pores on underside of

lower jaw. Lateral line comprising widely spaced pores on

short tubes arising from narrow scale-less gap, anterior end

Australasian red cods

curved upwards slightly, then gradually descending to lateral

midline below posterior 25% of second dorsal fin, remaining

on lateral midline posteriorly.

Membranes of first and second dorsal fins continuous at

base; first dorsal originating distinctly behind vertical through

pectoral fin base; anterior two-thirds of second dorsal of

uniform height, 7th to 9th ray from posterior end of fin

longest, its height about 1.4 times length of 2nd ray at anterior

end of fin, last ten or so rays progressively shorter; rays of

both dorsal fins mostly unbranched, only last 13 to 16 rays,

apart from last one to three, branched. Fleshy, fine scale-

covered basal sheath on third or more of first dorsal and

anterior portion of second dorsal fins; fin rays interconnected

by membranes to tips, sheath gradually decreasing slightly in

coverage and thickness posteriorly, encompassing about half

of second dorsal fin near its insertion; sheath extending onto

body for anterior third of combined dorsal fin base, broadest

anteriorly, narrowing posteriorly. Profile of anal fin like that

of second dorsal fin, although less deep posteriorly, with

comparable unbranched and branched rays. Likewise, anal fin

enclosed in broad fleshy sheath that is like that of dorsal fins;

sheath also extending onto body for anterior third or less of

fin. Caudal fin distinctly rounded without obvious

dorsoposterior and ventroposterior corners in adults; all but

anterior-most rays of similar length; base of fin covered by

indistinct sheath with much smaller scales (sheath sharply

demarcated from scales of caudal peduncle). Pectoral fin tip

variably reaching not quite to, or past vertical through anal fin

origin, sixth or seventh ray longest. Pelvic fin inserted anterior

to vertical through posterior edge of preopercle; outer two

rays longer than inner rays; second ray longest, 13.3 (13.3—

24.9)% SL, nearly twice length of subsequent ray, reaching

vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 7A.)

Medium to dark reddish brown above, extending ventrally to

level of ventral portion of pectoral fin base, white below in

smaller individuals, large individuals becoming brown

ventrally and often much darker; lateral line not distinctively

coloured. Underside of head, jaws and barbel white, tinged with

pink to orange, becoming more orange to brown in larger

individuals; ventral margin of suborbital dusky to dark. Dorsal

and caudal fins of similar colour to brown of sides, anal fin pink

or orange in smaller individuals, becoming brown like other

median fins in large individuals, dorsal, caudal and anal fins

with distinct broad to narrow black margin distally, margin on

caudal fin broadest. Pectoral fin orange to brown with faintly

darker semi-circular brown blotch basally, covering dorsal

80% of edge of fin proximally. Pelvic fin rays white with pink

to orange hue, becoming brownish in large individuals.

Preserved colour. Head and body dusky to dark dusky; underside

of head, belly and side adjacent to anal fin pale, especially in

small specimens, extent of pale underside less in large individuals.

Dorsal, caudal and anal fins dusky to dark dusky, anal fin often

less dark than dorsal and caudal; distal edges of dorsal, caudal

and anal fins with distinctly dark margins, dark margin on caudal

fin broad; pectoral fin broadly dusky basally, becoming pale

toward outer margin. Pelvic fin pale dusky.

Etymology. The specific epithet barbatus 1s Latin for “bearded”

in reference to the chin barbel, which 1s characteristic of this

species, its congeners and many other members of the family.

Distribution. Endemic to coastal waters of Australia and New

Zealand from Rottnest Island, Western Australia (32° OI' S,

115° 30' E), to Port Stephens, New South Wales (32° 30'S, 152°

35' E), including all of Tasmania and around both the North and

South Islands of New Zealand, reaching the Chatham Islands

in the east (fig. 3B). A demersal species on rocky bottom at | to

at least 520 m. Although found at considerable depths in New

Zealand waters, this is a shallow reef-dwelling species in

southern Australia, where it shelters in caves and beneath

overhangs during daylight hours, venturing out at night to feed.

Remarks. Pseudophycis barbata has an overall darker

colouration than its congeners, often with a reddish hue in fresh

material, and distinct black margins to the dorsal, caudal and

anal fins. [n particularly dark individuals, the basal portion of

the pectoral fin 1s similarly dark but lacks the distinct black

spot present in P. bachus and P. palmata. Pseudophycis

barbata 1s easily separable from P. breviuscula by the more

numerous scales (16—22 vs. 7—9) 1n a transverse row between

the dorsal fin origin and lateral line.

As mentioned above under Remarks for P. palmata,

McCoy's (1878: 29) treatment of P. barbata, the first published

after Günther's original description, was largely based on

specimens of P. palmata, but the accompanying illustration 1s

of a large specimen of P. barbata. Although Ramsay’s (1881)

description of Lotella grandis 1s inadequate for a conclusive

identification of the species, the lone type specimen (AMS

1.696) is clearly P. barbata. Early taxonomic confusion

resulted in Steindachner (1901: 509) reporting P. barbata

from New Zealand as L. grandis, Norman (1937: 55)

speculating that P. barbata (as Physiculus barbatus) may be

identical with P. bachus, and Graham (1938: 399) and Habib

(1975) treating it in New Zealand as P. breviusculus. The last

stems from our assumption that P. breviusculus does not

occur in the waters of the South Island based on specimens

examined and reliable identification records.

Material examined. Types. Pseudophycis barbatus BMNH

1863.1.15.38 (394, holotype) Victoria, South Australia (fig. 7a); Lotella

grandis AMS 1.696 (503, holotype) Wollongong, New South Wales.

Other material. (38 specimens examined for meristic or

morphometric values, 111—531 mm SL; see Appendix 2 for additional

material in Australasian collections.) Australia, Victoria: CSIRO H

3791-01" (340) south of Gabo Island, 37? 43.30' S, 149° 55.20' E-37°

41.10' S, 149° 57.10' E, 116-107 m, A. Graham, 17 September 1994;

CSIRO H 4500-01' (410) south of Cape Everard, 38° 07.81' S, 149?

30.87' E, 156 m, A. Williams and M. Lewis, 13 January 1997; NMV

A2248 (5, 134—185) Portland Harbour, halfway along lee breakwater,

adjacent to oil wharf, 38° 20.916' S, 141° 37.398' E, 7-12 m, R. Kuiter,

R. Wilson and I. Head, 22 October 1981; NMV A2261-001 (4, 149—180)

Portland Harbour, end of lee breakwater, 38? 21' S, 141° 36' E, 12 m, R.

Kuiter, R. Wilson and I. Head, 21 October 1981; NMV A13046 (203)

eastern Bass Strait, 110 km E of Paradise Beach, 38° 18.7' S, 148? 50.7'

E, 201-208 m, M. Gomon, 7 August 1993. Tasmania: AMS I.20079-

009 (90.1) south side of Rocky Cape, in National Park, 40° 51' S, 145°

31'E, 2-8 m, B.C. Russell, 8 December 1977; CSIRO H 7539-03! (185)

south-east of Orford, rock point W of Triabunna woodchip mill, 42°

32.92' S, 147° 54.80' E, 3 m, P.L Last, W. White and J. Pogonoski, 01

April 2014. South Australia: CSIRO CA 3526 (410 mm SL) Great

Australian Bight, 32° 11.8' S, 131° 22.4' E-32° 13.4' S, 131° 22.2' E, 60

m, 7 December 1981; CSIRO H 7949-13! (2, 47—70) Great Australian

Bight, 33° 20.20' S, 130° 15.42' E33? 20.20' S, 130° 16.26' E, 191-188

m, A. Graham, J. Pogonoski, M.F. Gomon and D.J. Bray, 15 December

2015. Western Australia: WAM P.26621.003 (135) Porpoise Bay,

32?00' S, 115°30' E, 1 m. New Zealand, North Island: NMNZ P.01930

(514) Wellington, Ohau Point, Makara Coast, 41 14.00' S, 174 39.00' E,

18 m, J. Moreland, 10 June 1956; NMNZ P.01492] (2, 207—240) Three

Kings Islands, west end of Great Island, 34^ 8.5' S, 172° 9.1' E, 5—7 m,

G.S. Hardy and A.L. Stewart, 28 November 1983; NMNZ P.037109*

(518) Mahina Knoll, NW of White Island, Bay of Plenty, 37° 20.46' S,

177? 5.43' E, 303 m, NMNZ/MARS White Island field team, 22 March

2009; NMNZ P.037124* (450) east of north-east point; White Island,

Bay of Plenty, 37° 30.96' S, 177? 13.54' E, 296 m, NMNZ/MARS

White Island field team, 19 March 2009; NMNZ P.044103' (250)

southeast of Club Rock; White Island, Bay of Plenty, 37? 32.35' S, 177°

11.92' E, 99 m, NMNZ/MARS White Island field team, 16 March

2009; NMNZ P.044272 (376) and NMNZ P.044273 (361) Three Kings

Islands, northern edge of Three Kings Shelf; north off Princess Islands,

114m, NMNZ/MARS Three Kings field team, 10 March 2010; NMNZ

P.045835' (197) east off Parengarenga Harbour entrance, 34? 30.93' S,

173° 18.05' E, 176 m, RV Kaharoa, 4 August 2009; NMNZ P.046563

(339) Three Kings Islands, at anchor, Northwest Bay; Three Kings

Island, 34 9.16' S, 172 8.06' E, 27 m, C.D Struthers, 6 March 2010;

NMNZ P.052585 (339) Northland, off North Cape, 34° 16.47' S, 173°

0.80' E, 130-132 m, RV Tangaroa, 25 March 2011; NMNZ P.052745'

(443) eastern side of Motkokako Island, Cape Brett, 35° 9.90' S, 174°

20.27" E, S. Tindale, 7 September 2011; NMNZ P.053551' (531) Kingfish

Reef, Bay of Islands, 35° 11.20' S, 174° 15.40' E, 35 m, 5. Tindale, 8

April 2012. South Island: NMNZ P.035978 (2, 234—266) West Coast,

Open Bay Island, south end Popotai Islet, 43° 51.82' S, 168° 52.48' E,

14-19 m, NMNZ Jackson-Haast field team, 14 February 1999; NMNZ

P.036535 (3, 186—256) West Coast, Open Bay Island, south end Popotai

Islet, 43? 51.82' S, 168° 52.48' E, 14-19 m, NMNZ Jackson-Haast field

team, 14 February 1999; NMNZ P.037525 (4, 111—148) West Coast,

North of Murphy Beach, Whakapohai Rocks, 43° 42.27' S, 169° |4.33'

E, 12-14 m, NMNZ Haast-Buller field team, 18 February 2000. NMNZ

P.053753 (281) off Snares Islands, 47? 59.99' S, 166? 36.25' E, 160 m,

NMNZ/MARS Auckland Island field team, 21 February 2012; NMNZ

P.053754 (390) Stewart Island, North Arm, Port Pegasus, 47° 10.73' S,

167? 40.95' E, 38 m, NMNZ/MARS Auckland Island field team, 20

February 2012.

Pseudophycis breviuscula (Richardson 1846)

Common name: Bastard Red Cod (Australian Standard);

northern bastard cod (New Zealand)

Figures 1, 3C, 5, 8; Tables 1—4

Lota breviuscula Richardson, 1846: 61, Pl. 38 (figs 1-2). Type

locality: Bay of Islands, New Zealand. Holotype: BMNH

1855.9.19.1182.

Pseudophycis breviusculus. Günther, 1862: 350; Günther, 1863:

116; Hector in Hutton and Hector, 1872: 116, pl. 8; Hutton in Hutton

and Hector, 1872: 47; Dambeck, 1879: 536, 547, 555; Hector, 1884: 55;

Hector, 1886: 28; Sherrin, 1886: 304; Gill, 1893: 120 (list); Hutton,

1904: 48; Thomson, 1906: 551; Young, 1925: 370; Graham, 1938: 405;

Graham, 1939b: 365; Ayling and Cox, 1982: 143, fig. (description).

Austrophycis megalops Ogilby, 1897: 91. Type locality: Maroubra

Bay, New South Wales, Australia. Holotype: AMS 1.3655.

Austrophycis megalops. McCulloch, 1927: 32 (list); McCulloch,

M.F. Gomon, C.D. Struthers & J. Kemp

1929: 129 (list); Munro, 1938: 62, fig. 443 (description); Whitley, 1964:

40 (list); Paxton and Hanley in Paxton et al., 1989: 302 (list).

?Pseudophycis breviusculus. Graham, 1939: 399.

Physiculus (Pseudophycis) breviusculus. Graham 1956: 173, 174

fig.; Whitley, 1956: 403 (list); Svetovidov, 1967: 1686, 1689; Whitley,

1968: 40 (list).

Pseudophycis breviuscula. Paulin, 1983: 93 (description,

taxonomy); Francis, 1988: 21; Paxton and Hanley in Paxton et al.,

1989: 302 (list); Paul, 1986: 58; Paulin, 1988: 450, 451, 453; Paulin et

al., 1989: 119 (key), 255 (list); Cohen in Cohen et al., 1990: 375

(taxonomy); Paulin and Roberts, 1992: 128, fig. 68a (description);

Kuiter, 1993: 60 (description); Gomon in Gomon et al., 1994: 335

(description); Francis, 1996: 21; Paul, 2000: 58; Francis, 2001: 25;

Hutchins, 2001: 23 (distribution); Sazonov, 2001: 293 (taxonomy of A.

megalops), Paxton et al., 2006: 616 (list); Gomon in Gomon et al.,

2008: 314 (description); Roberts et al. in Gordon et al., 2009: 532 (list);

McMillan et al., 2011a: 161, 162; McMillan et al., 2011b: 94; Francis,

2012: 52; Struthers et al., 1n Roberts et al., 2016: 864, fig. 107.22

(description); Roberts et al., 2015: S164; Roberts et al., 2017: 81,

Roberts et al., 2019: 90 (listed).

JAustrophycis marginata (nec Günther, 1878). Cohen in Cohen et

al., 1990: 356 (taxonomy of A. megalops).

Diagnosis. First dorsal fin with 8—10 rays, second dorsal fin

with 44—52 rays; anal fin with 49—56 rays; total vertebrae 42—

45; nostrils adjacent to anterior margin of orbit, interval

between nostril and orbital margin less than diameter of

combined nostrils; posterior nostril with raised anterior border

but without tubular flap posteriorly; gill rakers of outer arch

moderately short, those near angle much shorter than opposing

gill filaments; caudal peduncle short 28.8—33.4% HL; chin

barbel of moderate length, 23.4—30.5% HL; scales above lateral

line in oblique series from base of first dorsal fin ray 7—9;

oblique rows of scales intersecting with lateral line 77—93;

pyloric caeca 7—8; caudal fin rounded without angular corners,

the middle rays equal to or longer than the rays above and

below; no distinct dark blotch at base of pectoral fin; underside

of head, body and anal fin tan to brown, anal fin with fine black

distal margin. A small species reaching at least 151] mm SL.

Description. (Values for non-type specimens when different

from type in parentheses; see Tables 2—4 for summary of

selected meristic and comparative morphometric values.) First

dorsal fin 9 (8—10, rarely 8), first ray often minute; second

dorsal fin 46 (44—52, rarely less than 46); anal fin 50 (49-56);

pectoral fin 21 (20—23); pelvic fin 4 (4—5); caudal fin 26 (26—32,

rarely 26); gill rakers 3 (23) + 8 (7—8) = 9-12; lateral line

pores not associated with individual scales; oblique scale rows

intersecting with lateral line 93 (77-93); scales in oblique

series above lateral line 7 (7—9), scales 1n oblique series below

lateral line 23 (22-28, rarely 28); vertebrae 13 (11—14, rarely 11)

+ 31 (30—33) = 42-45, rarely 42; pyloric caeca 7-8.

Body of moderate depth, moderately compressed laterally

(fig. 8), greatest depth at or just in advance of anal fin origin

22.8 (20.6—23.3)% SL, tapering gradually from second dorsal

fin origin to posterior portion of fin and then more steeply to

shallow caudal peduncle; caudal peduncle short, 4.8 (5.7—

8.5)% SL, strongly compressed, depth subequal to orbital

diameter. Distance between middle of anus and base of anal

fin less than diameter of posterior nostril. Body cavity

extending to above origin of anal fin.

Australasian red cods

10 mm

Figure 8. Pseudophycis breviuscula. A, CSIRO H 4384-01, 110 mm SL, Albany, Western Australia (photograph compliments CSIRO Marine

Research); B, BMNH 1855.9.19.1182, holotype of Lota breviuscula, 152 mm SL, Bay of Islands, New Zealand (photograph © The Trustees of the

Natural History Museum, London).

Head acute, moderately short, length 25.8 (23.4—26.0)%

SL, its width 16.6 (12.4—17.9)% SL and depth 12.1 (10.8—15.2)%

SL; snout moderately short 5.9 (5.1—7.0)% SL, not projecting in

advance of upper jaw, rounded in dorsal view. Nostrils small,

located on posterior half of snout just in front of eye, distance

from nostrils to orbital margin less than diameter of combined

nostrils; nostrils positioned above horizontal through centre of

eye; diameter of posterior opening about 1.5 diameter of

anterior, separated from it by raised skin flap; posterior

opening without tubular flap encircling posterior margin.

Interorbital of moderate width, slightly convex. Eye of

moderate size, orbital diameter 27.4 (24.7—35.7)% HL, 0.91

(0.53—1.10) times 1n interorbital space, 3.72 (2.20—5.78) times

suborbital distance, circular, upper edge of eye just below

dorsal margin of head in lateral view, skin covering eye

unpigmented. Postorbital moderately long, 1.09 (0.97—1.50)

times length of upper jaw. Mouth large, terminal, upper jaw

terminating just posterior to vertical through posterior margin

of eye. Jaw teeth small, caniniform, slightly curved,

depressible, band of up to four or five irregular rows 1n upper

Jaw with narrow hiatus at symphysis; band tapering near rear

of jaw. Teeth of lower jaw similar in form to those of upper

Jaw; band of several rows with broader patch on either side of

symphysis, tapering posteriorly; rows almost contiguous

across symphysis. Vomerine teeth absent. Opercular bones

strong; upper extremity of gill opening at horizontal through

middle of eye; gill membranes continuous across isthmus. Gill

rakers on outer arch club-shaped, moderately short, about

0.3—0.8 times length of opposing gill filaments at angle, 15—23

times in head, slightly denticulate. Chin barbel of moderate

length, subconical 21.0 (19.6—30.5)% HL.

Moderately small cycloid scales covering all of head and

body except for branchiostegal membranes, surface of maxilla

and premaxilla, lower lip and distal parts of fins; very thick

mucus covering obscuring scales and pores in freshly

preserved material. Most head pores tiny, following main

cephalic sensory canals; row of slightly raised pores extending

from tip of snout posteriorly just above lower edge of

suborbital; row of enlarged mandibular pores on underside of

lower jaw. Lateral line comprising pores on tubes arising

above upper end of gill opening, anterior end curved upwards

slightly, then gradually descending to lateral midline below

posterior third of second dorsal fin, remaining on lateral

midline posteriorly.

Membranes of first and second dorsal fins continuous at

base; first dorsal originating distinctly behind vertical through

pectoral fin base; anterior two-thirds of second dorsal of

uniform height, 7th to 9th ray from posterior end of fin longest,

its height about 1.2 times length of 2nd ray at anterior end of

fin, last eight or so rays progressively shorter; rays of both

dorsal fins mostly unbranched, only about last 14 or 15 rays,

except last few, branched. Fleshy basal sheath on base of first

dorsal and anterior portion of second dorsal fins not especially

prominent with scales apparently confined to basal margin of

fin; fin rays interconnected by membranes to tips; sheath more

obvious on body below much of combined dorsal fin base,

broadest anteriorly, narrowing posteriorly. Profile of anal fin

similar to that of second dorsal fin, with comparable unbranched

and branched rays. Fleshy sheath on and below anal fin similar

to that of dorsal fins. Caudal fin rounded without distinct

dorsoposterior and ventroposterior corners, posterior margin

distinctly convex; all but anteriormost rays of similar length;

base of fin covered by indistinct sheath (sheath demarcated

from scales of caudal peduncle). Pectoral fin tip reaching past

vertical through anal fin origin, seventh or eighth ray longest.

Pelvic fin inserted anterior to vertical through posterior edge of

preopercle; outer two rays longer than inner rays; second ray

longest, 17.6 (16.6—21.6)% SL, two to four times length of

subsequent ray, reaching vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 8B.)

Medium brown to pale brown above, paler below with white

chest and belly in smaller individuals, bronze in larger

individuals; lateral line not distinctive. Underside of head and

jaws brownish; ventral edge of suborbital darker brown; barbel

white. Dorsal, anal and caudal fins very pale brown in smaller

individuals, darker in large individuals; fins with distinct broad

to narrow black margin distally, more diffuse in large individuals.

Pectoral fin orange with very faint semi-circular brown blotch

basally. Pelvic fin rays white with pink to orange hue.

Preserved colour. Head, body and fins pale dusky to

dusky, chest and belly very pale, ventral edge of suborbital

darker; distal edges of dorsal, caudal and anal fin with diffuse

darker margins.

Etymology. The name breviuscula is Latin for “rather short”,

probably in reference to the relatively small maximum size of

the species compared with others of the genus, although

Richardson separated it from other New Zealand morids simply

by fin counts.

Distribution. Endemic to coastal waters of Australia and New

Zealand, from Perth, Western Australia (32° Ol' S, 115? 30' E),

to Tweed Heads, New South Wales (28° 14' S, 153° 50' E),

including all of Tasmania and around the North Island of New

Zealand (fig. 3C). A demersal reef-dwelling species at 0—273 m,

although most often encountered at less than 100 m on rocky

and boulder reets (Struthers et al. 1n Roberts et al., 2016: 864).

Remarks. P. breviuscula 1s the smallest of the four species in

the genus, reaching less than half the maximum length of its

M.F. Gomon, C.D. Struthers & J. Kemp

congeners. It also occurs in slightly warmer waters compared

with the other three members of the genus, reaching well onto

the northern New South Wales coast in eastern Australia and

around New Zealand's North Island.

Paulin (1983: 94) provided a thorough synonymy of P.

breviuscula from a New Zealand perspective but failed to

mention Austrophycis megalops Ogilby, 1897, which was

based on a specimen from New South Wales, Australia. This

omission was possibly due to the widespread uncertainty

about the identity of Waite's shrivelled 65 mm type specimen.

some authors regarded Ogilby's species as congeneric with

Günther's (1878: 19) A. marginatus based on material from

the south-eastern Pacific and others thought the two were

likely to be conspecific (Cohen in Cohen et al., 1990: 356).

sazonov (2001: 343) reported that he had examined the type

specimen and found it to be P. breviuscula relegating the

name A. megalops to synonymy with that species.

Material examined. Types. Lota breviuscula BMNH 1855.9.19.1182

(152, holotype) Bay of Islands, New Zealand (fig. 8a); Austrophycis

megalops AMS 1.3655 (65, holotype) Maroubra Bay, New South

Wales, Australia, 33° 57' S, 151° 16' E, T. Whitelegge, 1897 (after

sazonov, 2001: 343, holotype, not re-examined for this study).

Other material. (47 specimens examined for meristic or

morphometric values, 70.2—139 mm SL; see Appendix 2 for additional

material in Australasian collections.) Australia, Victoria: NMV

A2261-002 (2, 106-111) Portland Harbour, end of lee breakwater, 38°

21' S, 141° 36' E, 12 m, R. Kuiter, R. Wilson and I. Head, 21 October

1981; NMV ASS82 (2, 84.0-127) Bass Strait, 80 km south-east of

Loch Sport, 38° 34.3' S, 148° 18.2' E, 86 m, Victorian Marine Sciences

Laboratory, 6 June 1984. Tasmania: CSIRO H 7698-09* (66) Huon

Commonwealth Marine Reserve, 43° 42.72' S, 147° 11.32' E—43?

42.97" S, 147° 10.35' E, 122 m, A. Graham and J. Pogonoski, 8 April

2015. Western Australia: WAM P.25342.004 (112) Cape Naturaliste,

33?32' S, 115*0T' E, J. Scott, 16 February 1964; WAM P.25343.017 (2,

108-119) Fremantle, 32°02' S, 115°40' E, L.M. Marsh et al, 24 June

1975; WAM P.26616.005 (125) Point Clune, 32°00' S, 115°30' E, 8 m,

J.B. Hutchins et al, 29 March 1979; WAM P.28297.006 (5, 59.4—139)

Lucky Bay, 34^08' S, 122°15' E, 8-10 m, J.B. Hutchins, 13 April 1984;

WAM P.28300.005 (5, 63.8—131) Lucky Bay, 34°05' S, 122°15' E, 11—

12 m, J.B. Hutchins et al, 16 April 1984. New Zealand, North Island:

NMNZ P.028044 (9, 532-127) Matatuahu Point, Tawharanui Peninsula,

Hauraki Gulf, 36? 23' S, 174° 49' E, 0—5 m, A.L Stewart and C.D.

Paulin, 8 April 1992; NMNZ P.028118 (7, 62—119) Onepoto Bay, Hicks

Bay, East Cape, 37° 35.25' S, 178° 18.00' E, 0-3 m, NMNZ East Cape

1992 field team, 4 May 1992; NMNZ P.029805 (3, 55-135) Waihau

Bay, Bay of Plenty, 37° 36.8' S, 177° 54.6' E, 4-6 m, NMNZ East Cape

1993 field team, 27 January 1993; NMNZ P.030036 (4, 103-135)

inside of Tuamotu Island, Gisborne Harbour, 38? 42.2' S, 178? 2.4' E,

4—7 m, NMNZ East Cape 1993 field team, 21 January 1993; NMNZ

P.046263 (103) north of Tom Bowling Bay; North Cape, 34^ 22.56' 5,

172? 55.28' E, 61—72 m, RV Tangaroa, 14 July 2009; NMNZ P.046285'

(2, 72.2—81.4) east of Purerua Peninsula, Bay of Islands, 35° 6.36' S,

174° 17.10' E, 119—121 m, RV Tangaroa, 7 July 2009; NMNZ P.048310

(70.4) western end of Omapere wharf, 35° 32.05' S, 173° 23.14 E, 4 m,

NMNZ & AIM Northland 2011 field team, 13 February 2011; NMNZ

P.048311 (70.5) western end of Omapere wharf, 35° 32.05' S, 173°

23.14' E, 4 m, NMNZ & AIM Northland 2011 field team, 13 February

2011; NMNZ P.048312 (82.6) western end of Omapere wharf, 35°

32.05' S, 173° 23.14' E, 4 m, NMNZ & AIM Northland 2011 field

team, 13 February 2011; NMNZ P.048380 (124), NMNZ P.048381

(96.8) and NMNZ P.048382 (3, 59.9-101) reef inside southern

Australasian red cods

headland of Hokianga Harbour, 35° 32.01' S, 173° 22.10' E, 5-8 m,

WCN 11/09, NMNZ & AIM Northland 2011 field team, 15 February

2011; NMNZ P.049600 (84.8) and NMNZ P.049703 (93) Tauroa Point,

Ahipara, 35° 10.48' S, 173° 2.73' E, 21 m, WCN 11/35, NMNZ & AIM

Northland 2011 field team, 22 February 2011; NMNZ P.049708 (100)

Tauroa Point, Ahipara, 35° 9.96' 5, 173° 3.12' E, 14m, NMNZ & AIM

Northland 2011 field team, 22 February 2011; NMNZ P.051786 (119)

Tauroa Point, Ahipara, 35° 10.48' S, 173° 2.73 E, 21 m, NMNZ &

AIM Northland 2011 field team, 22 February 2011; NMNZ P.057218'

(78.5) and NMNZ P.052719' (82.3) Ranfurly Bank, 37? 32.78' S,

178° 53.42' E, 68—70 m, RV Tangaroa, 30 May 2011.

Acknowledgements

We thank P. McMillan for providing insightful comments on

the manuscript. We are indebted to George Habib, who

provided a wealth of information about the morphology of

P. bachus in his unpublished thesis (Habib, 1975), and have

taken the liberty of including the vast majority of his

nomenclatural citations in the above synonymies, as well as

Larry Paul's unpublished bibliography. Amanda Hay and Sally

Reader (AMS), John Pogonoski (CSIRO), Jeremy Barker and

Romain Crechriou (NMNZ), and James Maclaine (BMNH)

provided radiographs of the types of the four species of the

genus. We are particularly grateful to John Pogonoski for

access to sequence data and for arranging the collection of

fresh material and to the angler Andrew Pender for assisting.

We are deeply grateful to Lara Shepard (NMNZ) who

extracted and prepared 11 additional sequences (including the

Neotype of P. bachus) to supplement existing BOLD

sequences. Alastair Graham (CSIRO), Mark McGrouther

(AMS), Glenn Moore and Mark Allen (WAM) and Dianne

Bray (NMV) for access to specimens. Ken Graham for his

collection of specimens and images. Bob Ward (CSIRO) made

available CO] sequences in his care and provided useful

comments on the manuscript. Scott Tindale assisted with the

collection of specimens. Special thanks go to Jeremy Austin

for his failed attempt to amplify adequate lengths of CO1 from

the purported type of P. palmata for comparison with those of

the three Australian species of Pseudophycis. We thank NIWA

skippers, crew and science staff (especially Peter McMillan

and Dan MacGibbon) of both RV Tangaroa and RV Kaharoa

for the important help and ongoing collecting, depositing of

Specimens, tissue samples and photographs to Te Papa. MV

Tranquil Image (Western Workboats), G. Gibbs, and S. Kelly

aided in logistics for Te Papa field collections. Collecting of

fresh material was also aided by Te Papa Collection

Development Programme (AP 3126). This work was supported

(in part) by the NZ National Institute of Water and Atmospheric

Research Ltd Core Funded Coasts and Oceans Programme 2:

Biological Resources subcontract for fundamental knowledge

of marine fish biodiversity with the Museum of New Zealand

Te Papa Tongarewa.

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Appendix |. COI sequences and source information for Pseudophycis spp featuring in Figure 1.

Species Source Voucher Reg No./ Country Collection Locality BOLD GenBank

Institution Sample ID No. Sequence ID accession

A. punctatus NMNZ P.045573 New Zealand South Island, west coast FNZC105-09 MN200065

A. punctatus NMNZ P.045574 New Zealand South Island, west coast FNZC106-09 MN200089

A. punctatus NMNZ P.045575 New Zealand South Island, west coast FNZC107-09 MN200090

A. punctatus NMNZ P.045576 New Zealand South Island, west coast FNZC108-09 MN200094

A. punctatus NMNZ P.045577 New Zealand South Island, west coast FNZC109-09 MN200092

A. punctatus NIWA no voucher PCOI New Zealand South Island, west coast FNZA333-07 MN200088

A. punctatus NIWA no voucher PCO2 New Zealand South Island, west coast FNZA334-07 MN200093

P. bachus NIWA no voucher RCOI New Zealand Campbell Island FNZ949-07 MN20003 1

P. bachus NIWA no voucher RCO4 New Zealand Campbell Island FNZ950-07 MN200023

P. bachus NIWA no voucher RCO5 New Zealand Auckland Islands FNZA35 1-07 MN200039

P. bachus NIWA no voucher RCO6 New Zealand Auckland Islands FNZA352-07 MN200027

P. bachus NIWA no voucher RCO7 New Zealand no data FNZA388-08 MN200026

P. bachus NIWA no voucher RCO8 New Zealand no data FNZA389-08 MN200030

P. bachus NIWA no voucher RCO9 New Zealand no data FNZA390-08 MN200028

P. bachus NMNZ P.044338 New Zealand South Island, south of Timaru FNZA555-08 MN200037

P. bachus NMNZ P.044339 New Zealand South Island, south of Timaru FNZA 556-08 MN200029

P. bachus NMNZ P.044348 New Zealand Chatham Rise FNZA557-08 MN200062

P. bachus NMNZ P.044349 New Zealand Chatham Rise FNZA558-08 MN200024

P. bachus NMNZ P.044350 New Zealand Chatham Rise FNZA559-08 MN200040

P. bachus NMNZ P.053748 New Zealand Stewart Island, Port Pegasus MN200063

P. bachus NMNZ P.053749 New Zealand Stewart Island, Port Pegasus MN200069

P. bachus NMNZ P.053750 New Zealand Stewart Island, Port Pegasus MN200076

P. bachus NMNZ P.053751 New Zealand stewart Island, Port Pegasus MN200075

P. bachus NMNZ P.053752 New Zealand Stewart Island, Port Pegasus MN200035

P. bachus NMNZ P.054828 (Neotype) New Zealand North Island, Whanganui Bar MN200038

P. barbata CSIRO H 3791-01 Australia VIC, Gabo Island FOAD135-05 MN200087

P. barbata CSIRO H 4500-01 Australia VIC, Cape Everard FOAD 141-05 MN200080

P. barbata CSIRO H 7539-03 Australia TAS, Orford FOAO050-14 MN200085

P. barbata CSIRO H 7904-03 Australia SA, Great Bight FOAPO09- 16 MN200086

P. barbata CSIRO H 7949-13 Australia SA, Great Bight FOAP215-16 MN200079

P. barbata CSIRO no voucher Australia NSW, Cape Howe FOAD137-05 MN200083

P. barbata CSIRO no voucher Australia NSW, Diaster Bay FOAD138-05 MN200078

P. barbata CSIRO no voucher Australia NSW, Diaster Bay FOAD139-05 MN20008 1

P. barbata CSIRO no voucher Australia NSW, Diaster Bay FOAD 140-05 MN200082

P. barbata CSIRO no voucher GT 10004 Australia SA, Great Bight FOAP335-17 MN200084

P. barbata NIWA no voucher SBRI New Zealand no data FNZA383-08 MN200073

P. barbata NIWA no voucher SBR2 New Zealand no data FNZA384-08 MN200072

P. barbata NIWA no voucher SBR3 New Zealand no data FNZA385-08 MN200074

P. barbata NIWA no voucher SBR4 New Zealand no data FNZA386-08 MN200070

P. barbata NIWA no voucher SBR5 New Zealand no data FNZA387-08 MN200066

P. barbata NIWA no voucher SBR6 New Zealand North Island, Ngawi FNZA608-08 MN200036

P. barbata NIWA no voucher SBR7 New Zealand North Island, Ngawi FNZA609-08 MN200068

P. barbata NMNZ P.037109 New Zealand North Island, White Island FNZC188-09 MN200091

P. barbata NMNZ P.037124 New Zealand North Island, White Island FNZC189-09 MN200077

P. barbata NMNZ P.044103 New Zealand North Island, White Island FNZC280-09 MN20007 1

P. barbata NMNZ P.045835 New Zealand North Island, East coast FNZC252-09 MN200067

P. barbata NMNZ P.052745 New Zealand North Island, Cape Brett MN200025

P. barbata NMNZ P.053551 New Zealand North Island, Bay of Islands MN200033

P. breviuscula CSIRO H 6838-08 Australia NSW, Broken Bay FOAK632-10 MN200059

Australasian red cods

Species

P. breviuscula

P. palmata

Source

Institution

CSIRO

NMNZ

NMV

CSIRO

Voucher Reg No./

sample ID No.

H 6838-08/2

H 6845-01/1

H 6845-01/2

H 6845-01/3

H 6980-04

H 6991-01

H 7698-09

P.046285

P.046285-1

P.052718

P.052719

A 26125-002

A 31132-001

H 4229-01

H 7366-01

H 7716-01

H 7717-01

no voucher BW- 1692

no voucher BW-1693

no voucher BW- 1694

no voucher GT 7974

no voucher GT 7975

no voucher GT 7976

Country

Australia

New Zealand

Australia

Collection Locality

NSW, Broken Bay

TAS, east coast

TAS, northeast

North Island, Bay of Islands

North Island, Ranfurly Bank

VIC, Port Phillip Bay

TAS, east coast

TAS, southeast

TAS, North Bruny Island

TAS, Munro Bight

TAS, Hinsby Beach

VIC, Bass Strait

TAS, east coast

TAS, southeast

BOLD

sequence ID

FOAK604- 10

FOAK626-10

FOAK612-10

FOAK610-10

FOAK240-10

FOAK247-10

FOAO011-15

FNZC272-09

FNZC271-09

FMVIC234-08

FOAO533-15

FOAD1306-05

FOAO5909-15

FOAO000-15

FOADI32-05

FOAD133-05

FOAD 134-05

FOAOS596-15

FOAO597-15

FOAO598-15

GenBank

accession

MN200053

MN200054

MN200055

MN200060

MN200057

MN200056

MN200064

MN200058

MN200061

MN200032

MN200034

MN200052

MN200043

MN20004 1

MN200048

MN200047

MN20005 1

MN200050

MN200044

MN200042

MN200045

MN200046

MN200049

M.F. Gomon, C.D. Struthers & J. Kemp

Appendix 2. Specimens registered in Australasian collections not used for compiling morphological data. Not all specimens listed were examined

in the course of the study.

Physiculus palmata

(78 specimens, 30—410 mm SL.) Australia: AMS 1.7534 (310)

no locality, 1905. New South Wales: AMS 1.15024 (420)

Merimbula, 36°53' S, 149°56' E, 1908; AMS I.34458-001 (87)

one third of way to Tollgate Island from Three Islet Reef

Batemans Bay, 35°44' S, 150°15.5' E, 100 m, 23-24 November

1988; AMS 1.34461-001 (190) Lookout Point, Twofold Bay,

37°4.5' S, 149°55' E, 50 m, 26-27 November 1988; AMS

1.34566-001 (225) Batemans Bay west end of Long Beach,

35°42' S, 150°13' E, 50 m, 23-24 November 1988; AMS

1.34568-001 (128) North Head Beach, Batemans Bay, 35°43’

S, 150°16' E, 50 m, 23-24 November 1988; CSIRO H3537-01

(5, 54-97) south of Disaster Bay, 37°24.2' S, 149?58.4

E-37?22.9' S, 149°58.7' E, 42-44 m, 13 August 1993.

Victoria: NMV A639 (221) off Phillip Island, 38?39' S,

145°19.8' E, 07 August 1979; NMV A836 (2, 104—138) Port

Phillip Bay, 38°06' S, 144°52.8' E, 1970; NMV A837 (2, 129—

133) Port Phillip Bay, 38°06' S, 144°52.8' E, 1971; NMV A838

(2, 134—146) Port Phillip Bay, 3.2 km (2 miles) Port Phillip

Bay, west of Sandringham, 37°57' S, 144°57' E, 30 March

1971; NMV A839 (250) Port Phillip Bay, 38°06' S, 144°52.8'

E, 25 July 1927; NMV A841 (2, 245—250) Port Phillip Bay,

Hobsons Bay, 37°52.2' S, 144°55.8' E, September 1867; NMV

A846 (2, 190—207) Port Phillip Bay, Hobsons Bay, 37°52.2' S,

144°55.8' E, September 1867; NMV A848 (315) no data, old

collection; NMV A2285 (51.4) central Bass Strait, 38 km

southwest of Cape Paterson, 38°55.5' S, 145°17' E, 70 m, 12

November 1981; NMV A2667 (51.8) Eastern Bass Strait, 8 km

south of South East Point, Wilsons Promontory, 39°12.9' S,

146°27.3' E, 65 m, 18 November 198]; NMV AIO0580 (242)

Western Port, 5 km north of Cowes, 38°26.1' S, 145?15.3' E, 20

m, 22 October 1986; NMV A20816 (2, 201—210) no data, old

collection, August 1864; NMV A26125-002 (61.0) Port

Phillip Bay, central part of bay, east of St Leonards, 38°12.8'

S. 144*50' E, 24 m, 12 December 1996; NMV A23366-002

(401) no data, old collection; NMV A31156-001 (237) Port

Phillip Bay, Hobsons Bay, 37.87°S, 144.93°E; NMV A31157-

001 (223) same collection data as NMV A31156-001.

Tasmania: AMS IB.1192 (142) d'Entecasteaux Channel,

simpsons Bay, 43°17' S, 147°18' E, 7 July 1942; AMS 1.10287

(140), Oyster Bay, 42°40' S, 148°03' E, 1909; AMS 1.14182

(300) Port Arthur, 43°09' S, 147°51' E, 13 April 1917; AMS

1.14183 (248) Port Arthur, 43?09' S, 147?51' E, 13 April 1917;

AMS 1.6243 (62) NW coast: Ulverston, 41?10' S, 146?11' E,

1903; AMS 1.6275 (340) Tamar River Heads, 41?20' S, 147°02'

E, 1903; AMS 1.6276 (245) Tasmania, Tamar River Heads,

41°20' S, 147?02' E, 1903; AMS 1.9259 (252) Port Arthur,

43°09' S, 147°S1' E, 1908; AMS 1.9989 (177) Bass Strait, east

of Flinders Island, 40?01' S, 148°42' E, 1909; AMS 1.9990

(195) Bass Strait, east of Flinders Island, 40?01' S, 148?42' E,

1909; NMV A1218-003 (2, 297—299) Central Bass Strait, 20

km north-northeast of North Point, 40°31.8' S, 145°22.8' E, 44

m, BSS 116 T, M.F. Gomon, G.C.B. Poore, and P. Forsyth, 4

November 1980; NMV A1289 (294) central Bass Strait, 23

km east of Cape Rochon, Three Hummock Island, 40°22.2' S,

145°17' E, 40 m, 3 November 1980; NMV A1381 (38.5)

central Bass Strait, 6 km northeast of Stanley, 40°48.8' S,

145°22' E, 22 m, 4 November 1980; NMV A1479 (67.2) central

Bass Strait, 32 km west-southwest of Settlement Point,

Flinders Island, 40°9.95' S, 147?31.8' E, 51—52 m, 6 February

1981; NMV A1535 (8, 163—220) central Bass Strait, 32 km

northwest of Devonport, 40°56' S, 146°5.4' E, 68—64 m, 4

February 1981; NMV A9776 (272) near Marlo Reef, 37°48' S,

148°31.8' E, 1991; NMV A9777 (276) Marlo Reef, 37°48' S,

148°31.8' E, 1991; NMV A20556 (202) Bass Strait, east coast

of Flinders Island, 40° S, 148°20' E, 73 m, 16 June 1909;

NMV A21590 (4, 150-182) Bass Strait, east of Flinders

Island, 40°S, 148°33' E, 1909; WAM P.27554.001 (186) Spring

Bay, 42°32' S, 147°55' E, 1-8 m; South Australia: SAMA F

2777 (1) Gulf St Vincent, 34°11' S, 138°9' E, 14 July 1953;

SAMA F4624 (8) Robe, 37°10' S, 139°45' E, 9 September

1979; SAMA F7172 (1) Spencer Gulf, Port Lincoln, 34°44' S,

135°52' E, 1992; SAMA FI10581 (1), Robe, bay near river

mouth, 16 March 2003; SAMA F10855 (1) Investigator Strait,

35°24'28" S, 137°54'40" E, 16 December 2006; SAMA

F10900 (1) Investigator Strait, 35°19' 09" S, 137?46' 05" E, 15

December 2006; SAMA F11864 (1) Great Australian Bight,

Coles Point, 34?22'06" S, 135°21'09" E, 21 March 2003;

SAMA F12755 (1) Victor Harbour, 35?33' S, 138°37' E, 14

september 2003.

Pseudophycis bachus

(416 specimens, 21-573 mm SL.) New Zealand, North

Island: NMNZ P.001129 (373) Wellington, Ngauranga,

Wellington Harbour, 41715' S, 174°50' E, 3 August 1952;

NMNZ P.001809 (174) North Auckland, off Kaipara Bar,

36°24' S, 174°9.5' E, 183 m, August 1955; NMNZ P.002271

(227) Wellington, Wellington Harbour, off Petone, 41°14.0' 5,

174°52.5' E, 13-18 m, 22 August 1957; NMNZ P.002372

(neurocranium) Wellington, York Bay, Wellington Harbour,

41°15.9' S, 174°54.2' E, 10-11 m, 31 May 1953; NMNZ

P.002440 (2, otoliths Wellington, Wellington Harbour,

41°16.5' S, 174°51.0' E, 1953; NMNZ P.003011 (otoliths and

skeleton) Wellington, Paraparaumu Beach, 40*53' S, 174°56'

E, 8 January 1961; NMNZ P.004788 (11, 71—128) Wellington,

Palliser Bay, 41°26.3' S, 175°3.0' E, 64-82 m, 15 February

1968; NMNZ P.006562 (10, 73—99) north of Kapiti Island, off

Foxton, 40°30.5' S, 174°53.0' E, 101 m, 1 March 1976; NMNZ

P.006802 (2, 212—243) Bay of Plenty, 8 km north of Mayor

Island, 37°12' S, 176°15' E, 366 m, 28 September 1962; NMNZ

P.007357 (otoliths) Wellington, Te Mimi, south end of Kapiti

Island, 40°52.35' S, 174°54.60' E, 3 m, 22 August 1977;

NMNZ P.00744] (otoliths) Wellington, 11 January 1978;

NMNZ P.008363 (3, 90—94) South Auckland, east-northeast

of Tolaga Bay, 38°15.2' S, 178°38.6' E, 139 m, 16 January

1979; NMNZ P.009415 (84) Gisborne, Matakaoa, East Cape,

37°34' S, 178°20' E, 15 m, 26 June 1988; NMNZ P.009823 (3,

79-112) Taranaki, west-northwest of Cape Egmont, 38°48.8'

S, 173°29.6' E, 146 m, 9 January 1981; NMNZ P.010543 (315)

Bay of Plenty, north of Mayor Island/east of Slipper Island,

Australasian red cods

37°4.95' S, 176°12.70' E, 315—352 m, 18 April 1981; NMNZ

P.014082 (2, 107-109) Wellington, off Castle Point, 40°56.25'

S, 176°22.80' E, 115-140 m, 17 April 1975; NMNZ P.016708

(10, 32-68) Wellington, east of Cape Campbell, 41°44.15' S,

174°27.65' E, 47 m, 15 December 1978; NMNZ P.017108 (293)

laranaki, Seal Rocks, Sugar Loaf Islands, New Plymouth,

39°3.25' S, 174°0.20' E, 30 m, 24 March 1985; NMNZ

P.017508 (2, 96-100) Wellington, Hikurangi Trench, 40°54.8'

S, 176°25.4' E, 140 m, 17 April 1976; NMNZ P.017509 (102)

Wellington, Hikurangi Trench, 40*57.7' S, 176°20.2' E, 115 m,

17 April 1976; NMNZ P.018714 (2, 52-57) Manawatu,

southwest of Whanganui, 40°22.65' S, 174°22.80' E, 62—80 m,

17 July 1985; NMNZ P.019115 (57) Gisborne, east of Tolaga

Bay, 38°23.05' S, 178°26.65' E, 30 m, 11 January 1980; NMNZ

P.019331 (50) Hawke's Bay, Cape Kidnappers, 39°39' S,

177°10' E, 36 m, 19 October 1969; NMNZ P.023430 (230)

Gisborne, Matakaoa, East Cape, 37°34' S, 178°20' E, 15 m, 26

June 1988; NMNZ P.029685 (5, 310—360) off Mohaka, 39°15'

S. 177°20' E, 17 November 1992. South Island: AMS I.14733

(440) Portobello, Otago Harbour, 45°51' S, 170?39' E, 13

December 1918; AMS 1.14734 (550) Otago, Blueskin Bay,

45?44' S, 170?35' E, 29 November 1918; NMNZ P.001753

(148) Otago, edge of Karitane Canyon, northeast of Tairoa

Head, 45°38.5' S, 171°2.0' E, 220 m, 14 August 1955; NMNZ

P.006548 (2, 90—109) Marlborough, Western Cook Strait; ca

14 km northeast of Stephens Island, 40°33' S, 174°7' E, 130—

132 m, 4 March 1976; NMNZ P.006704 (278) Canterbury,

northeast Mernoo Bank, Chatham Rise, 43?10.5' S, 174?58.5'

E, 298—422 m, 23 June 1975; NMNZ P.007180 (305) Snares

Islands, west side of western chain, Snares Island, 48°3' S,

166°30' E, 120 m, 4 December 1976; NMNZ P.007349 (200)

Marlborough, Kaikoura area, 41°20' S, 174°9' E; NMNZ

P.007442 (otoliths) Marlborough, ‘Run Under Point’; Cook

Strait between Port Underwood and Tory Channel, 41°18.25'

S, 174°14.55' E, 19 January 1978; NMNZ P.007457 (505)

Canterbury, Western Chatham Rise, off Mernoo Bank,

43°16.45' S, 174°55.50' E, 220 m, 11 December 1977; NMNZ

P.007730 (12, 31-123) Marlborough, Cloudy Bay, 41°26.1' S,

174°15.9' E, 59—64 m, 28 January 1979; NMNZ P.008107 (79)

Nelson, 6.4 km northwest of Farewell Spit, 40°27.0' S,

172°48.5' E, 70 m, 10 March 1976; NMNZ P.008355 (185)

Southland, Crooked Arm, Fiordland, 45°25' S, 166°58' E, 16

m, 7 January 1977; NMNZ P.008356 (14, 67-105)

Marlborough, Cloudy Bay, 41°26.35' S, 174°9.70' E, 27-28 m,

28 January 1979; NMNZ P.008365 (18, 60—118) Marlborough,

Cloudy Bay, 41°26.35' S, 174°9.70' E, 27—28 m, 28 January

1979; NMNZ P.009282 (5, 228—360) Canterbury, Western

Chatham Rise, off Mernoo Bank, 43°16.45' S, 174°55.50' E,

220 m, 11 December 1977; NMNZ P.009283 (7, 170—213)

Canterbury, Western Chatham Rise, off Mernoo Bank,

43°16.45' S, 174°55.50' E, 220 m, 11 December 1977; NMNZ

P.010598 (2, 168—170) Otago, off Oamaru, 45*5' S, 170*59' E,

46 m, July 1962; NMNZ P.010647 (115) Otago, Cape

Wanbrow, off Oamaru, 45°7' S, 171°3' E, 100-101 m, June

1963; NMNZ P.010703 (65) Otago, Oamaru, 45°7' S, 174722

E, 73 m, March 1973, NMNZ P012069 (14, 57-75)

Marlborough, Mernoo Bank, 43°6.1' S, 175*20.5' E, 153 m, 12

January 1979; NMNZ P.012088 (2, 65-80) Marlborough,

Cloudy Bay, 41°26.35' S, 174°9.70' E, 27—28 m, 28 January

1979; NMNZ P.013119 (490) Snares Islands, Puysegur Trench,

46°49.15' S, 165°50.50' E, 459—515 m, October 1982; NMNZ

P.016344 (272), Snares Islands, off North Promontory, 48*0'

S, 166°36' E, 110 m, 5 December 1984; NMNZ P.017527 (2,

34—35) Stewart Island, Foveaux Strait, 46°30' S, 167°30' E,

May 1976; NMNZ P.017876 (12, 58—80) Nelson, north of

Cape Farewell, 40*4.65' S, 172*57.09' E, 102 m, 14 December

1978; NMNZ P.0O17879 (2, 36—60) Canterbury, northern

Canterbury Bight, off Waiau River mouth, 42°36.8' S,

173°40.4' E, 81 m, 16 December 1978; NMNZ P.017884 (80)

Nelson, north of Cape Farewell, 40°8.75' S, 173*11.15' E, 90

m, 14 December 1978; NMNZ P.O17885 (5, 38—46)

Canterbury, northern Canterbury Bight, off Kaikoura

Peninsula, 42°39.30' S, 173°35.95' E, 130 m, 16 December

1978; NMNZ P.017891 (12, 28—40) Marlborough, off Cape

Campbell, 41°42.1' S, 174°25.5' E, 60 m, 15 December 1978;

NMNZ BP017892 (15, 33-64) Canterbury, northern

Canterbury Bight, off Kaikoura Peninsula, 42°38.15' S,

173*39.75' E, 91 m, 16 December 1978; NMNZ P.017893 (15,

2]-46) Marlborough, Cook Strait, east of Cape Campbell,

41°46.8' S, 174°28.6' E, I8 m, 15 December 1978; NMNZ

P.018854 (48) Marlborough, east of Flaxbourne Depression,

41°59.9' S, 174°26.9' E, 100 m, 19 January 1982; NMNZ

P.018865 (45) Marlborough, east of Flaxbourne Depression,

4|*59.50' S, 174°30.55' E, 100 m, 19 January 1982; NMNZ

P.018878 (69, 26—75) Canterbury, Canterbury Bight/southeast

of Timaru, 44°43.4' S, 171754.7 E, 95 m, 16 January 1982;

NMNZ P.019322 (2, 45) Westland, northwest of Hokitika,

42?34.18' S, 170°15.19' E, 204 m, 10 December 1978; NMNZ

P.019509 (99, 29—55) Canterbury, northeast of Clarence River

mouth/central Campbell Bank, 42°6.00' S, 174°6.95' E, 84—86

m, 24 November 1982; NMNZ P.019831 (500) Southland,

North Port, Fiordland, 45°59' S, 166°34' E, 27 m, 8 May 1986;

NMNZ P019949 (205) Southland, 30 Fathom Point,

Fiordland, 45°43.3' S, 166°30.3' E, 36—37 m, 13 May 1986;

NMNZ P.020802 (110) East coast South Island, 1982; NMNZ

P.020806 (2, 72—76) East coast South Island, 1982; NMNZ

P.025063 (3, 88-137) Otago, off Tairoa Head, Otago

Peninsula, 45*45' S, 170*45' E, 1990; NMNZ P.025077 (3, 96—

135) Otago, off Otago Peninsula, 45°36.55' S, 170°51.88' E,

41-59 m, 10 May 1990; NMNZ P.025093 (8, 76—116) Otago,

off Otago Peninsula, 45°41.85' S, 170°58.72' E, 97-100 m, 10

May 1990; NMNZ P.025186 (90) Otago, Blueskin Bay, Otago

Peninsula, 45°43.28' S, 170°40.33' E, 20-22 m, 9 May 1990;

NMNZ P.025782 (2, 443—480) Canterbury, Pegasus Canyon,

43°23.78' S, 173°40.50' E, 113-109 m, 1 June 1990; NMNZ

P.030708 (7, 87—120) Snares Islands, Southern Snares Shelf,

48°30.75' S, 166°58.50' E, 134-136 m, 1 April 1993; NMNZ

P.032382 (420) Southland, Waterfall mooring, head of Bligh

Sound, Fiordland, 44°50' S, 166°32' E, 8—10 m, 21 March

1995; NMNZ P.032398 (3, 395—410) Southland, mooring at

Precipice Cove, Bradshaw Sound, Fiordland, 45°15' S, 167°10'

E, 20 m, 28 March 1995; NMNZ P.036007 (2, 266—300)

Westland, outer Frog Rock, 43°58.52' S, 168°33.48' E, 14—20

m, 11 February 1999; NMNZ P.036769 (2, 120-130)

Canterbury, southwest of Timaru, 44°35.71' S, 171°12.55' E,

18—19 m, 20 December 1998; NMNZ P.045793 (104, C&S)

Marlborough, Cloudy Bay, 41°26.1' S, 174°15.9' E, 59-64 m,

28 January 1979. Bounty Islands: NMNZ P.006643 (573)

Bounty Platform, southeast of Islands, 48°7.25' S, 179°16.25'

E, 230—238 m, 18 November 1975. Auckland Islands: NMNZ

P.007136 (264) northwest Auckland Islands/ Campbell

Plateau, 50°10.35' S, 167°43.10' E, 120 m, 21 January 1977.

Chatham Rise: NMNZ P.008357 (2, 78—79), Chatham Rise,

Mernoo Bank, 43°36' S, 175°31' E, 375 m, 12 February 1954;

NMNZ P.008358 (66) Mernoo Bank, Chatham Rise, 42°59.4'

S, 175°30.5' E, 112 m, 23 January 1954; NMNZ P.009414 (2,

94—188) east of Chatham Islands, 43?30.95' S, 176°9.25'W,

176—205 m, 22 May 1987; NMNZ P.020910 (4, 100—125)

Subantarctic Slope, 43°39.95' S, 175°51.40'W, 220—246 m, 28

May 1987; NMNZ P.020922 (270) northeast of Chatham

Islands, 43°33.6' S, 176°0.5'W, 228—234 m, 26 May 1987;

NMNZ P.020952 (273) east of Chatham Islands, 43°30.95' S,

176°9.25'W, 176—205 m, 22 May 1987; NMNZ P.020989 (2,

127-129) east of Chatham Islands, 43°39.15' S, 175°52.10'W,

216—244 m, 24 May 1987; NMNZ P.035283 (142) Urry Bank,

Chatham Rise, 44°9.0' S, 176°6.5' E, 126 m, October 1979.

Campbell Island: NMNZ P.009284 (390) Campbell Island

Rise, northeast of Island, 51°51' S, 169*40' E, 250—271 m, 10

April 1982; NMNZ P.012110 (530), Campbell Island Rise,

northeast of Island, 51?51' S, 169*40' E, 250—271 m, 10 April

1982.

Pseudophycis barbata

(211+ specimens, 26—620 mm SL.) Australia: AMS IB.7565

(465) no locality, 1966. New South Wales: AMS 1.16970-014

(176) Boydtown, Nullica Bay, 37°6' S, 149°53' E, 13 March

1972; AMS I.19833-003 (5, 48—60) east of Ulladulla, 35°20' S,

150*51' E, 128 m, 14 May 1974; AMS 1.25974-003 (91) Eden,

eastern Bass Strait, 37°S' S, 149?55' E, 18—46 m, 26 November

1984; AMS I.27359-008 (270) Quarantine Beach inside North

Head, 33°49' S, 151°18' E, 6 April 1967; AMS I.27359-000 (4,

105-180) Quarantine Beach inside North Head, 33°49' S,

151°18' E, 6 April 1967; AMS 1.28908-001 (305) east of

Shoalhaven Bight, 34°53' S, 151?06' E, 18 April 1985; AMS

1.34570-002 (140) east of Lookout Point Twofold Bay, 37?46'

S, 149°55' E, 26—27 November 1988; AMS I.34854-003 (115)

off Newcastle, 33?2' S, 151?58 E, 117-121 m, 24 March 1993;

AMS IB.8192 (514) Eden, 37°4' S, 149°55' E, 9 June 1968;

CSIRO A 1117 (72 mm TL) between Cronulla and Eden,

October 1948; CSIRO B 3455 (4, 59-96 mm TL) off Moruya

at “The Pines Close", 35°55' S, 150°10' E, 90 m, 28 April 1939.

Victoria: AMS L16987-006 (5, 75-150) Petersborough,

estuary, near boat ramp, 38°37' S, 142°52' E, 6 m, 21 March

1972; CSIRO H 3793-03 (83) south of Gabo Island, 57?44.0'

S. 149°58.0' E-37?*42.3 S, 149°59.5' E, 108-115 m, 17

september 1994; NMV 30055 (502) Western Port, 38°22.2' S,

145°22.2' E, October 1858; NMV 43104 (375) Port Phillip

Bay, Hobsons Bay, 37°52.2' S, 144°55.8' E, October 1878;

NMV 43105 (375) Port Phillip Bay, Hobsons Bay, 37°52.2' S,

144°55.8' E, October 1879; NMV A842 (252) No location

data, | January 1971; NMV A2941 (102) Port Phillip Heads,

38°16.8' S, 144°37.8' E, 1864; NMV AI0567 (225) Western

Port, 5 km north of Cowes, 38°26.1' S, 145°15.3' E, 20 m, 22

M.F. Gomon, C.D. Struthers & J. Kemp

October 1986; NMV A23366-001 (438) no data, old

collection. Tasmania: AMS _ 1.17545-016 (70) Eaglehawk

Neck, 43?2' S, 147°56' E, 29 November 1972; AMS I.17555-

011 (6, 62—95) The Gardens, north of Binalong Bay, 41°13' S,

148°13' E, 1 m, 6 December 1972; AMS L.20086-001 (75)

spring Beach, 42°35' S, 147°54' E, 2-4 m, 16 December 1977;

AMS I.34952-003 (260) Mouth of Fortescue Bay, 43°07'46" S,

147°59'28" E, 9-10 April 1994; CSIRO B 1259 (6, 50—62 mm

TL) NE coast, 43°03' S, 148°03' E, 17 December 1976; CSIRO

C 1405 (165) St Helens, 28 February 1951; CSIRO H 1151-02

(90) 40°55.05' S, 147?21.35' E, 33 m, 17 June 1987; CSIRO T

1105 (26) east of Babel Island, 120 m, 12 April 1984; CSIRO

T 1252 (1) St Helens, 41?25' S, 148°16' E, 30 m; CSIRO T 1523

(1) St Helens Point, east coast of Tasmania, 25 February 1982;

CSIRO T 1625-01 (27) Flinders Island; WAM P.27553.002

(103) Mercury Passage, 42?33' S, 147°57' E, 7-8 m; WAM

P.27554.002 (2, 136-206) Spring Bay, 42?32' S, 147°55' E, 1-8

m; WAM P.27559.002 (2, 74—94) Saint Helens Point, 41?16' S,

148°22' E, 3 m; WAM P.27560.002 (2, 111—296) Bridport,

41°00' S, 147?23' E, 8-9 m; South Australia: AMS 1.18470-

005 (191) Robe, southwest coast, 37°10' S, 139°45' E, 1 m, 3

October 1975; AMS I.20180-021 (5, 69-112) Kangaroo Island,

Penneshaw, 35°44' S, 137°58' E, 0—5 m, 9 March 1978; CSIRO

CA 3516 (281 mm TL) west of Investigator Group, Great

Australian Bight, 33°43.8' S, 132°07.8' E-33°45.1' S, 132°08.9'

E, 180-184 m, 8 December 1981; SAMA AMSTACIO2 (1)

Great Australian Bight, Anxious Bay, 33°20'03" S, 13473847"

E, 25 February 1981; SAMA AMSTACISI (1) 30 miles west

of Robe, 37°09'47" S, 139°12'31" E, 7 April 1981; SAMA

AMSTAC213 (1) 4.5 miles west of Beachport, 37°29' S,

139*55' E, 8 April 1981; SAMA AMSTAC288 (1) Cape

Buffon, 1 April 1981; SAMA AMSTAC289 (1) Cape Buffon,

] April 1981; SAMA AMSTAC292 (1) 2 miles west of Cape

Buffon, 1 April 1981; SAMA AMSTACT7II (1) Great

Australian Bight, South East Isles, 34°20' S, 123°42' E, 26 July

1981; SAMA AMSTACI331 (1) Spencer Gulf, between

Wardang Island and Tipara Reef, 34°12'06" S, 137°18'24" E;

SAMA AMSTACIA90 (1) Kangaroo Island, 2-3 miles off

West Bay, 36°03'40" S, 137°12'39" E, 16 January 1982; SAMA

AMSTAC1602 (1) off Port MacDonnell, 38°03'S, 140°42'E, 1

January 1982; SAMA F505 (1) 1918; SAMA FI1012 (1) Gulf St

Vincent, 35°10' S, 137?55' E, 1928; SAMA F1144 (1) 1928;

SAMA 1145 (1); SAMA 1146 SAMA 1736 (1) St Vincent

Gulf, Port Wakefield, 34°11' S, 138°29' E, 8 July 1932; SAMA

F2011 (1) Spencer Gulf, Moonta, 34°3' S, 137?34' E, 14

October 1937; SAMA F2031 (1) Eyre Peninsula, Venus Bay,

33°12' S, 134°40' E, 29 April 1938; SAMA F2766 (1) Spencer

Gulf, Port Lincoln, 34?44' S, 135°52' E, 28 January 1953;

SAMA F3045 (1) Adelaide, Outer Harbour, 34°47' S, 138°29'

E, 26 January 1962; SAMA F3377 (1) Kangaroo Island,

Stokes Bay, 35°37' S, 137°12' E, 15 August 1966; SAMA

F3392 (1) Kangaroo Island, Stokes Bay, 35°37' S, 137°12' E,

15 August 1966; SAMA F3649 (1) Kangaroo Island, Osmanli

Reef, D’Estrees Bay, 35°59' S, 137?38' E, 13 August 1966;

SAMA F3972 (1) near Coffin Bay, Farm Beach, 34°31' S,

135°23' E, 31 August 1974; SAMA F3980 (1) Encounter Bay,

Port Elliott, Bashams Beach, 35?32' S, 138°41' E, 13 October

1974; SAMA F4093 (1) Encounter Bay, 35?33' S, 138°38' E,

Australasian red cods

20 July 1975; SAMA F4281 (1) Kangaroo Island, Emu Bay,

35°35' S, 137°31' E, 21 January1968; SAMA F5162 (1) Port

MacDonnell, 38°3' S, 140°42' E, 1984; SAMA F5204 (1) Gulf

St. Vincent, 34°26' S, 137?55' E, June 1984; SAMA F5529 (1)

Kangaroo Island, Penneshaw, 35°43' S, 137°56' E, 29 October

1985; SAMA F6416 (1) Spencer Gulf, Cowell, 33°41’ S,

136°55' E, 1988; SAMA F7174 (1) Spencer Gulf, 34°44' S,

135°52' E, 17 February 1993; SAMA F9097 (1) Kangaroo

Island, north of Hog Bay, 35°43' S, 137°57' E, 20—22 m, 6 June

1954; SAMA F9576 (1) Investigator Strait, Althorpe Islands,

35°22'00" S, 136°52'00" E; SAMA F10526 (1) Investigator

Strait, 17 km N of Point Marsden, 35?24'58" S, 137?38'15" E,

28 June 2001; SAMA F10723 (1) Great Australian Bight,

32°15'31" S, 132°39'15" E, 20 April 2002; SAMA F10749 (1)

Great Australian Bight, 34?01'26" S, 134?28'34" E, 18 April

2002; SAMA FI11194 (1) Gulf St Vincent, 35?16'54" S,

138°09'36" E, 17 May 2007; SAMA FI11195 (1) Gulf St

Vincent, 35°16'54" S, 138°09'36" E, 17 May 2007; SAMA

F11196 (1) Gulf St Vincent, 35?16'54" S, 138?09'36" E, 17

May 2007; SAMA F14170 (1) Rivoli Bay, 37°31'44" S,

140°00'04" E, 6 April 2009. Western Australia: AMS 1.12324

(225) Great Australian Bight, southwest of Eucla, 32?00' S,

128?00' E, September 1912; CSIRO H 4383-08 (72) Bunbury,

inner harbour, general berth, 33°19.54' S, 115°39.57' E, 10.5

m, 6 March 1996; CSIRO H 4904-07 (108) Bunbury, outer

harbour (disused jetty groyne), 3.5 m, 6 March 1996; NMV

A9232 (350) Great Australian Bight, 120 km south of Middini

Beach, 33°17.1' S, 127°29.7' E, 162-160 m, 14 February 1990;

SAMA F73 (1) Abrolhos Islands, 28°45' S, 113°47' E, 22 April

1913; WAM P.1794.001 (365) Hopetoun, 33°57' S, 120°07' E;

WAM P.7687.001 (350) Albany, 35?00' S, 117°52' E; WAM

P.14596.001 (lost?) 31°54' S, 110°10' E; WAM P.26119.001

(149) Lookout Point, 34?53' S, 118?25' E; WAM P.29035.009

(270) 34?03' S, 122?00' E; WAM P.29339.001 (160) Mandurah.

Tasman Sea: NMNZ P.009285 (280) southern Lord Howe

Rise, 38? S, 168° E, 260 m, 7 December 1993; NMNZ

P.030825 (460) southern Lord Howe Rise, 38? S, 168? E, 400—

405 m, 31 August 1993; NMNZ P.031243 (540) southern Lord

Howe Rise, 38? S, 168? E, 260 m, 7 December 1993. New

Zealand, North Island: NMNZ P.000256 (135) Auckland,

Hauraki Gulf, 36°30' S, 175*0' E, September 1920; NMNZ

P.001236 (486) Wellington, Makara, 41°13.0' S, 174°42.5' E,

40 m, 28 February 1953; NMNZ P.001462 (528) Wellington,

Cape Terawhiti, 41717' S, 174°37' E, 9 m, 28 February 1954;

NMNZ P.001970 (165) Bay of Plenty, northwest of Mayor

Island, 37°15' S, 176°12' E, 146-219 m, 18 August 1956;

NMNZ P.002233 (379) Wellington, North of Castle Point,

41°37' S, 175°16' E, 65 m, 25 September 1957; NMNZ

P.002795 (500) Wellington, South of Titahi Bay, 41°6' S,

17450 E, 3 m, 12 July 1959; NMNZ P.006783 (308)

Wellington, Breaker Bay, Seatoun, 41°20' S, 174°50' E, 9

October 1973; NMNZ P.007789 (92) Hawke's Bay, 30 km east

of Portland Island, 39°18.25' S, 178°12.00' E, 258—306 m, 27

January 1979; NMNZ P.008360 (116) Bay of Plenty, southeast

of Mayor Island, 37°22.5' S, 176°22.0' E, 207-219 m, 27

February 1957; NMNZ P.008361 (8, 74—120) South Auckland,

east-northeast of Tolaga Bay, 38°15.2' S, 178°38.6' E, 139 m,

l6 January 1979; NMNZ P.009286 (2, 322-331) South

Auckland, Anchor Reef, southeast corner Whale Island,

37*5l' S, 176°59' E, 1 July 1998; NMNZ P.010327 (3, 315—352)

Bay of Plenty, north of Mayor Island/east of Slipper Island,

37*4.95' S, 176°12.70' E, 315—352 m, 18 April 1981; NMNZ

P.013608 (550) Wellington, Mana Island, 41*5' S, 174^52' E, 74

m, 12 March 1983; NMNZ P.014893 (2, 360—410) Three

Kings Islands, NW Bay, Great Island, 34°9.0' S, 172°8.5' E, 14

m, 25 November 1983; NMNZ P.017924 (3, 42—48), off Mayor

Island, Bay of Plenty, 37°14.35' S, 176°21.15' E, 218—225 m, 13

December 1975; NMNZ P.018168 (422) Bay of Plenty, off east

side of Mayor Island, 37°18' S, 176718 E, 37-91 m, I7

February 1986; NMNZ P.023405 (385) Gisborne, Waiaka Bay

near Lottin Point, East Cape, 37°33' S, 178°9' E, 6 m, 24 June

1988; NMNZ P.028461 (288) Gisborne, southeast of Cape

Runaway, East Cape, 37°32.9' S, 178*0.5' E, 8—12 m, 24 April

1992; NMNZ P.030132 (570) Gisborne, Monowai Reef,

38°35.9' S, 178°16.7' E, 15-21 m, 20 January 1993; NMNZ

P.030137 (286) Gisborne, inside of Tuamotu Island, Gisborne

Harbour, 38°42.2' S, 178°2.4' E, 4-7 m, 21 January 1993;

NMNZ P.031312 (279) Hawke’s Bay, “White Cliffs’, ~3 km

south of Cape Kidnappers, 39°42' S, 177°3' E, 40 m, 23

January 1994; NMNZ P.031697 (200) Bay of Plenty, southern

Kermadec Ridge, 37°32.90' S, 177°6.12' E, 325—327 m, 14

January 1995; NMNZ P.031923 (114) Bay of Plenty, southern

Colville Ridge, 36°57.12' S, 176°15.90' E, 336-339 m, 9

January 1995; NMNZ P.033156 (340) Wellington, north end

of Mana Island, 41°4.63' S, 174°47.40' E, 10-15 m, 8 March

1996; NMNZ P.033318 (160) Wellington, north end of Mana

Island, 41°4.63' S, 174°47.40' E, 10-15 m, 8 March 1996;

NMNZ P.033569 (245) Bay of Plenty, Long Point, Mahia

Peninsula, 39°10' S, 177*49' E, 8 m, 4 January 1995; NMNZ

P.034786 (520) Hawke's Bay, White Cliffs, northern end of

Ocean Beach, 39°41' S, 177°3' E, 20 m, 27 April 1997; NMNZ

P.036897 (440) South Auckland, Anchor Reef, southeast

corner Whale Island, 37°51' S, 176°59' E, 1 July 1998; NMNZ

P.036907 (375) South Auckland, east side of White Island,

37°31' S, 177°12' E, 160 m, 4 July 1998; NMNZ P.041342

(380) Northland, 232 m, 25 January 2005. South Island:

NMNZ P.005345 (2, 127) Nelson, Central Eastern Challenger

Plateau, west of Farewell Spit, 40°37.5' S, 171°37.5' E, 247 m,

3 March 1971; NMNZ P.007707 (353) Canterbury, Mernoo

Bank, 43°23.0' S, 175°6.8' E, 124—129 m, 12 January 1979;

NMNZ P.008031 (489) Marlborough, Kaikoura region, 42°25'

S, 173°43' E, 109-128 m; NMNZ P.009008 (2, 290—380)

Westland, Hokitika Canyon, 42°13' S, 170*34' E, 305—380 m,

12 August 1979; NMNZ P.009465 (420) Marlborough,

Kaikoura region, 42°25' S, 173°43' E; NMNZ P.016345 (300)

Snares Islands, off North Promontory, 48*0' S, 166°36' E, 110

m, 5 December 1984; NMNZ P.016889 (422) Southland, west

of Richards Point, Bradshaw Sound, Fiordland, 45°16.7' S,

167°0.7' E, 15-21 m, 27 February 1985; NMNZ P.016933 (2,

150-177) Southland, off Seymour Island, Doubtful Sound,

Fiordland, 45°18' S, 167°0' E, 12 m, 28 February 19855; NMNZ

P.016970 (460) Southland, off Seymour Island, Doubtful

sound, Fiordland, 45°18' S, 167°O' E, 12 m, 28 February 1985;

NMNZ P.017874 (65) Westland, NW off Hokitika, 40°38.48'

S, 170°11.42' E, 169 m, 11 December 1978; NMNZ P.021159

(620) Southland, mouth of Caswell Sound, Fiordland, 44°58.9'

S, 167°7.6' E, 48 m, 15 February 1987; NMNZ P.023949 (430)

Stewart Island, off Hebe Island, Port Pegasus, 47°11.8' S,

167°38.5' E, 10-16 m, 28 January 1989; NMNZ P.027612 (2,

195—256) Stewart Island, SE Point Rosa Island, Port Pegasus,

47°9.9' S, 167742.0 E, 0-9 m, 10 March 1992; NMNZ

P.027622 (390) Stewart Island, 13 Fathom Point, Whale

Passage, Port Pegasus, 47°11.00' S, 167°43.05' E, 20 m, 13

March 1992; NMNZ P.0278770 (225) Stewart Island, SE Point

Rosa Island, Port Pegasus, 47°9.9' S, 167°42.0' E, 0—9 m,

March 1992; NMNZ P.030270 (133) Southland, “Little Cove’,

Acheron Passage, Fiordland, 45°39.88' S, 166°44.78' E, 0—10

m, 21 March 1993; NMNZ P.030520 (320) Southland, “46

fathom point", Broughton Arm, Breaksea Sound, Fiordland,

45°33.35' S, 166°57.95' E, 2-18 m, 21 March 1993; NMNZ

P.030554 (330) Southland, Peninsula opposite Oak Island,

Wet Jacket Arm, Fiordland, 45?38.65' S, 166°51.90' E, 0—33

m, 28 March 1993; NMNZ _ P.030559 (310) Southland,

Stephens Cove, Fiordland, 45*36' S, 166°40' E, 7 m, 18 March

1993; NMNZ P.030940 (3, 84—183) Tasman, Taupo Point,

Abel Tasman National Park, 40*47.5' S, 172*57.0' E, 4—8 m, 19

November 1993; NMNZ P.032260 (157) Southland, | nautical

mile south of Deas Cove, south side of Thompson Sound,

Fiordland, 45°12.79' S, 166°57.31' E, 8-24 m, 28 March 1995;

NMNZ P.033439 (475) Stewart Island, south of Stewart

Island, Campbell Plateau, 48°47.78' S, 166°47.26' E, 158 m, 5

March 1996; NMNZ P.035083 (140) Westland, Bridget Point,

Milford Sound; Fiordland, 44°38.61' S, 167?54.86' E, 6—14 m,

5 April 1998; NMNZ P.035981 (462) Westland, point south of

Teer Creek, 44^0' S, 168°29' E, 15-18 m, 13 February 1999.

Chatham Rise: NMNZ P.020957 (2, 265) east of Chatham

Islands, 43°30.95' S, 176°9.25'W, 176—205 m, 22 May 1987;

NMNZ P.021008 (378) east of Chatham Islands, 43?31' S,

176°7'W, 206-226 m, 24 May 1987; NMNZ P.026557 (5,

143—255) channel between Point Munning and Te Whakuru

Island, Chatham Island, 43°44.3' S, 176°12.0'W, 0—3 m, 15

February 199]; NMNZ P.026670 (245) Cape Fournier,

Owenga, Chatham Island, 44°2.0' S, 176°19.5'W, 15—17 m, 10

February 1991; NSMT 32967 (537) Wanganella Bank, 32°41.6'

S. 167°42' E, 123—405m, 12 December 1989.

Pseudophycis breviuscula

(636 specimens, 15-196 mm SL). Australia, New South

Wales: AMS 1.3982 (152) off Newcastle, 32?56' S, 151°56' E,

4 March 1898; AMS 1.3983 (170) off Newcastle, 32°58' S,

151°55' E, 4 March 1898; AMS A.10105 (138) Port Jackson,

33°50' S, 151°10' E, 1881; AMS I.16879-001 (105) Jervis Bay,

off Darling Road, 35°3' S, 150°44' E, 22 September 1971;

AMS 1.17178-016 (65) Port Jackson, near Manly, 33°50' S,

151°16' E, 13 August 1972; AMS 1.19893-024 (4, 67-128)

south of Nadgee River mouth, north end of Blackpoint, 37°30!

S, 149°58' E, 0—5 m, 26 August 1976; AMS 1I.20065-002 (27,

35—46) off Ulladulla, 35°21' S, 150°49' E, 0—250 m, 27

October 1977; AMS I.20472-002 (2, 40-43) east of Green

Cape, 36°24' S, 150°18' E, 0-128 m, 1 November 1977; AMS

1.20568-002 (2, 130-170) east of Tweed Heads, 28°14' S,

153°50' E, 132-137 m, 2 June 1978; AMS [.20653-008 (119)

and AMS 1[.20653-013 (59) east of Hat Head, 31°5' S, 31°2' S,

M.F. Gomon, C.D. Struthers & J. Kemp

153°13' E, 155 m, 24 August 1977; AMS I.21366-019 (5, 35—

45) off Newcastle, 33°17' S, 153?5' E, 28 November 19791;

AMS 1.21797-003 (127) east of Tuggerah Lake, 33°8' S,

151°54' E, 115 m, 17 July 1978; AMS 1.21798-002 (4, 100-120)

east of Tweed Heads, 28°14' S, 153?50' E, 132-137 m, 2 June

1978; AMS L22873-002 (115) southeast of Cape Byron,

28?43' S, 153?49' E, 131 m, 1 November 1978; AMS 1.23379-

006 (42) Coffs Harbour, boat harbour, 30?20' S, 153°20' E,

0—6 m, 3 May 1977; AMS 1I.23685-001 (132) Smoky Cape,

Coffs Harbour, Scott Point, Nambucca Heads, 30°43' S,

153°16' E, 134-151 m, 10 October 1978; AMS 1.23687-001

(82) Ballina - Tweed Heads, 28°4' S, 153°50' E, 137 m, 16

August 1978; AMS LI.23688-001 (3, 117—165) Ballina - Tweed

Heads, east of Brunswick Heads, 28?25' S, 153?48' E, 119 m, 3

June 1978; AMS 1.23692-001 (2, 122-145) Ballina - Tweed

Heads, 28?34' S, 153°50' E, 146—150 m, 18 August 1978; AMS

1.23870-006 (106) Sydney-Newcastle, 33°40' S, 152°56' E, 720

m, 20 December 1976; AMS L.24367-003 (2, 37—48) off

sydney, 33°43' S, 151°54' E, 430—541] m, 27 October 1983;

AMS 1.24440-002 (4, 35-45) east of Twofold Bay, 37?03' S,

150 20' E, 149 m, 1 November 1977; AMS I.25865-003 (145)

off Port Stephens, 32?49' S, 152?2' E, 40—46 m, 10 April 1985;

AMS 1.25893-001 (196) off Twofold Bay, 37°5' S, 149°55' E,

18—46, 25 November 1984; AMS 1.26239-005 (118) east of

Wooli, 29?49' S, 153?24' E, 36—54 m, 25 March 1985; AMS

1.26444-002 (112) northeast of Broken Bay, 33°29' S, 151°49'

E, 142-137 m, 13 March 1986; AMS 1I.26451-005 (4, 51—99)

Sydney, off Broken Bay, 33°35' S, 151?4]' E, 134—135 m, 10

February 1986; AMS 1.26906-009 (93) Iluka, 29°24' S,

153°21' E, 1987; AMS 1.26919-002 (42) northeast of Jervis

Bay, off Point Perpendicular, 35?5' S, 150°55' E, 128 m, 13

December 1984; AMS I.27064-006 (91) 35?0' S, 150?45' E,

2—7 m, January 1987; AMS 1.27179-010 (43) 30°40' S, 159°0

E, 0-229 m, 20 September 1987; AMS 1.27322-005 (141) off

Tuncurry, 32°08' S, 152°31' E, 1 October 1985; AMS 1.27323-

004 (104) east of Camden Head, 31?42' S, 152?49' E, 2 October

1985; AMS L.27670-001 (93) off Coffs Harbour, 30?26' S,

153°22' E, 25 July 1981; AMS 1.31483-003 (135) off Evans

Head, 29°00' S, 153°49' E, 6 May 1990; AMS 1.32120-001

(109) off Clarence River, 29?24' S, 153?35' E, 1-2 May 1990;

AMS 1.34467-001 (2, 143—162) off Wollongong, 34?28' S,

151?2' E, 100 m, 6—7 May 1993; AMS I.34472-002 (3, 90—150)

off Wollongong, 34?26' S, 150°58' E, 7-8 May 1993; AMS

1.34474-001 (94) off Wollongong, 34°28' S, 151°2' E, 100 m,

7-8 May 1993; AMS 1.34475-001 (128) off Wollongong,

34°28' S, 151?2' E, 100 m, 7-8 May 1993; AMS 1I.34557-003

(96) off Wollongong, 34?26' S, 150°57' E, 50 m, 6-7 May

1993; AMS I.34715-001 (2, 85-115) Botany Bay channel off

revetment, 33?59' S, 151°12' E, 16 December 1978; AMS

1.34894-001 (125) off Wollongong, 34°0' S, 151°0' E, 28-29

March 1994; AMS I.35422-001 (2, 95-120) northeast of Coffs

Harbour, 30°15.75' S, 153°21.98' E, 98 m, 12-13 August 1993;

AMS I.35424-001 (3, 120—135) east of Coffs Harbour, 30°17'

S, 153°13' E, 50 m, 8-9 September 1994; AMS I.35427-001

(2, 86—93) east of Coffs Harbour, 30°17' S, 153°13' E, 50 m, 9

september 1994; AMS I.40321-006 (130) southeast of Yamba,

29°39' S, 153°41' E, 5 July 1999; AMS I.40491-001 (6, 854—125)

off Wollongong, 32°26' S, 150°57' E, 50 m, 6-7 May 1993;

Australasian red cods

AMS 1.44627-019 (79) Tathra, Baronda Headland, south side,

36°41'11" S, 149°59'54" E, 8 April 2008; AMS 1.44627-020

(81) Tathra, Baronda Headland, south side, 36°41'11" S,

149*59'54" E, 8 April 2008; AMS 1.44632-006 (2, 100)

Tathra, Kianniny boat ramp, 36°44'15" S, 149°58'60" E, 9

April 2008; AMS 1.44823-022 (97) north side of Moon Island,

33°05'12" S, 151°40'16" E, 5 May 2009; AMS 1.45025-004 (4,

96-115) Colliers Beach, inlet near Mollymook Golf Course,

35°20'46" S, 150?28'36" E, 10 March 2010; AMS I.45025-005

(2, 100-115) Colliers Beach, inlet near Mollymook Golf

Course, 35?20'46" S, 150?28'36" E, 10 March 2010; AMS

1.45027-018 (2, 65-75) Jones Beach, Mollymook, 35°19'19" S,

150?29' E, 11 March 2010; AMS L.45633-057 (6, 71—95)

Washerwomans Beach, 35?14'39" S, 150?32'09" E, 16 March

2011; AMS IA.1955 (72) off Green Cape, 13—35 km northeast

of cape, 37°4' S, 150°9' E, 71-84 m, June 1924; AMS IA.1956

(80) and AMS IA.1957 (60) 13-35 km northeast of Green

Cape, 37°4' S, 150°9' E, 71-84 m, June 1924; AMS IA.2966

(125) 16-19 km northeast of Sydney Harbour, 33°' S, I51°E,

110-146 m, 1926; AMS IA.6898 (75) 5 km off Broughton

Island, 32?37' S, 152?22' E, 82-91 m, 29 May 1936; AMS

[B.4359 (110) east of Tuggerah Lakes, 33?20' S, 151°35' E,

229—260 m, 19 June 1959; AMS IB.7005 (147) off Port

Stephens, 32° S, 152° E, 1964; CSIRO A 1622 (84 mm TL)

Pambula Bay, 11 m, 13 May 1950; CSIRO A 1639 (97 mm TL)

Twofold Bay, off Eden, 10.8 m, 5 May 1950; CSIRO A 1640

(83 mm TL) Twofold Bay, off Eden, 10.8 m, 5 May 1950;

CSIRO H 3575-01 (2, 60—70) east of Merimbula, 36°55.4' S,

149°56.8' E—36°54.6' S, 149°58.0' E, 33—40 m, 4 August 1993;

CSIRO H 4258-04 (107) south of Green Cape, 37°21.23' S,

150°06.03' E-37?20.1' S, 150°06.52' E, 90—94 m, 7 May 1996;

CSIRO H 4773-01 (148) east of Yamba, 29°24' S, 153?35

E-29*23' S, 153?35' E, 68—71 m, 17 April 1996; CSIRO H

4773-02 (4, 132-158) east of Yamba, 29?24' S, 153?35'

E-29*23' S, 153?35' E, 68—71 m, 17 April 1996; CSIRO H

5962-01 (2, 105-142) northeast of Coffs Harbour, 29°50' S,

153°27' E-29*48' S, 153°27' E, 66-68 m, 15 April 1996;

CSIRO H 6838-08 (4, 103—160) east of Broken Bay, 33°32.3'

S. 151°30.9' E-33?28.3' S, 151?32.5' E, 60—62 m, 27 June

2006; CSIRO H 0845-01 (3, 117-181) east of Broken Bay,

33°29.4' S, 151?32.2' E-33?33.7' S, 151?29.8' E, 60—62 m, 29

June 2006; CSIRO T 1434 (1) off Port Stephens, 32°44' S,

152°16' E-32?43' S, 152°23' E, 83-104 m, 20 August 1979;

NMV A6437 (31.0) 65 km east of Nowra, 34?52.9' S, 151°22.7'

E, 23 October 1988; NMV A6438 (31.3) 52 km east-southeast

of Nowra, 34°56.1' S, 151°14.7' E, 21 October 1988; NMV

A13051 (2, 52.4—68.5) 12 km east-northeast of Bermagul,

36°23.3' S, 150°10.7' E, 72-69 m, 15 August 1993. Victoria:

NMV A7531 (55.9) Batison Point, tide pool, 37°34.2' S,

149°46.2' E, I m, 6 April 1989; NMV A7500 (55.9) Gabo

Island, harbour on southeast side, 37°34.2' S, 149°55.2' E, 4 m,

4 April 1989; NMV A13050 (68.8) Disaster Bay, 37°16.3' S,

149*57.9' E, 24—29 m, 11 August 1993; NMV A2682 (28.1)

central Bass Strait, 26 km southeast of Aireys Inlet, 38°39.8'

S, 144°18.2' E, 79 m, 19 November 1981; NMV A3776 (2,

72.2—T1.2) eastern Bass Strait, 24 km southwest of Lakes

Entrance, 38?03' S, 147?49.8' E, 45 m, 1 October 1983; NMV

A384] (4, 58.9-79.3) eastern Bass Strait, 28 km south-

southwest of Marlo, 37°58.8' S, 148°27' E, 51 m, 30 July 1983;

NMV A5812 (3, 46.8—61.8) eastern Bass Strait, 40 km south-

southwest of Lakes Entrance, 38°18' S, 147°37' E, 55 m, 31

July 1983; NMV A835 (108) Bass Strait, 24 km (16 miles)

south-southwest of Lakes Entrance, 38°7' S, 147°51.8' E, 29—

37 m, 27 April 1948; NMV A8775 (38.3) Bass Strait, 30 km

east of Gabo Island, 37°37.2' S, 150°16.7' E, 432 m, 14 October

1984; NMV A20817 (74.5) southeast of Lakes Entrance,

38°10.2' S, 148°4.8' E, 1971; NMV A20818 (4, 48.3—80.0) no

collection data; NMV A29091-001 (80.5) Portland, inner

harbour, South Henty, 38°21' S, 141°36' E, 13 m, 1 May 1996.

Tasmania: AMS 1.43935-001 (78) East of Fortescue Bay,

north of Hippolyte Rocks, 43°0642" S, 148°03'27" E, 9-10

April 1994; CSIRO H 6980-04 (42) southern Tasman Sea,

40°48.97' S, 150?58.30' E-40?49.16' S, 150°56.54' E, 392-185

m, 21 June 2009; CSIRO H 6991-01 (36) southern Tasman

Sea, 40°48.35' S, 152?16.42' E—40?48.53' S, 152°15.25' E,

174-91 m, 21 June 2009; CSIRO T 1952 (150) Frederick

Henry Bay, 25 m, 19 September 1983; NMV A2626 (47.2)

eastern Bass Strait, 100 km northeast of North Point, Flinders

Island, 38?52.6' S, 148?25.2' E, 130 m, 15 November 1981;

NMV A2676 (35.4) central Bass Strait, 35 km north of Cape

Wickham, King Island, 39°13.6' S, 143°55.6' E, 85 m, 23

November 198]; NMV A3789 (40.7) eastern Bass Strait, 18

km north-northeast of Deal Island, 39°19.2' S, 147?277 E, 63

m, | October 1983; NMV A6435 (25.8) 68 km east of Cape

Tourville, 42°3.9' S, 149°11.9' E, 27 October 1988; NMV

A6439 (2, 24.3—34.0) 52 km east-northeast of Cape Tourville,

42°2.4' S, 148°58.3' E, 28 October 1988; NMV A6440 (3,

17.7—22.2) 54 km east-northeast of Cape Tourville, 42?02.1' S,

148°58.4' E, 28 October 1988; NMV A6909 (30.5) 78 km

south of Point Hicks, 38°30.1' S, 149°15.5' E, 25 October 1988;

NMVA26688-003 (2, 65.0—76.4) Port Phillip Bay, off Seaford,

artificial reef, 38705.25' S, 145°05.95' E; NMV A31158-001

(101) Port Phillip Bay, off Carrum, 7 April 2011; NMV

A31159-001 (95.5) and NMV A31159-002 (87.9) Port Phillip

Bay, 28 March 2011; WAM P.27554.021 (127) Spring Bay,

42?32' S. 147°55' E, 1-8 m. South Australia: AMS 1.20167-

021 (69) Kangaroo Island, Knob Point, 2 km east of Stokes

Bay, 35°37' S, 137°15' E, 15 m, 5 March 1978; AMS 1.20168-

013 (77) Kangaroo Island, Hanson Bay, 36°OI' S, 137°52' E, 6

March 1978; AMS I.20171-022 (41) Kangaroo Island, Vivonne

Bay, 36?0' S, 137°11' E, 0-2 m, 6 March 1978; CSIRO H 5337-

Ol (2, 73—86) Great Australian Bight, 33?16.00' S, 130°43.15'

E—33?16.60' S, 130°48.63' E, 137 m, 11 May 2000; CSIRO H

5338-01] (4, 69-125) Great Australian Bight, 31°50.05' S,

130°45.90' E-31*50.32' S, 130°45.10' E, 55 m, 14 May 2000;

SAMA AMSTACS89 (4) Great Australian Bight, 8 nautical

miles south of Point Weyland, 33°22'49" S, 134°37'53" E, 18

September 1981; SAMA AMSTAC1660 (1) Great Australian

Bight, 6 miles off Venus Bay, 33°18'S, 134°35'E, 1 July 1982;

SAMA F3948 (1) northern Great Australian Bight, 32°24' S,

133°30' E, 42 m, 5 May 1973; SAMA F9096 (1) Kangaroo

Island, Nepean Bay, 35°38' S, 137°46' E, 54 m, 28 April 1981;

SAMA F9098 (1) Great Australian Bight, Anxious Bay,

33°20'03" S, 134°38'47" E, 25 February 1981; SAMA F9099

(1) Great Australian Bight, Nuyts Archipelago, off Goat

Island, 32°17'00" S, 133°30'00" E, 40 m, 28 February 1981;

SAMA F10955 (1) Investigator Strait, 35°26'43" S, 137°56 42"

E, 17 May 2007; SAMA F10990 (1) Investigator Strait,

35°24'28" S, 137°54'40" E, 17 May 2007; SAMA F14583 (1)

22 May 1905. Western Australia: AMS 1.20229-021 (90)

Cockburn Sound, | km south of Carnac Island, 32°10' S,

115?40' E, 6-8 m, 26 March 1978; AMS I.20247-010 (2, 58—

63) Rottnest Is, Kingston Reefs, 31°59' S, 115?33' E, 12 April

1978; CSIRO H 4384-01 (110) Albany, Princess Royal

Harbour, 35°02.02' S, 117°53.05' E, 7 m, 28 February 1996;

WAM P.2999.001 (120) Mandurah, 32?32' S, 115°43' E; WAM

P.7409.001 (174) Augusta, 34°19' S, 115°09' E; WAM

P.21913.001 (135) Wilson Inlet, 34?59' S, 117?26' E; WAM

P.25195.019 (3, 47—56) Cape Naturaliste, 33°32' S, 115?02' E,

l-5 m; WAM P.26006.003 (46) 34?07' S, 122°16' E, 13 m;

WAM P.26009.006 (2, 65—70) Lucky Bay, 34?00' S, 122? |4' E,

13 m; WAM P.26600.008 (4, 30—76) Albany, 35?08' S, 117°38'

E; WAM P.26608.016 (2, 50-75) Cheyne Beach, 34°53' S,

118°25' E, 12-15 m; WAM P.26621.003 (117) Porpoise Bay,

32°00' S, 115°30' E, 1 m; WAM P.28292.008 (9, 45-121)

Lucky Bay, 34°00' S, 122°14' E, 7-10 m; WAM P.28293.011

(67) Lucky Bay, 34°00' S, 122°14' E, 5-7 m; WAM

P.28296.013 (5, 45—96) 34?08' S, 122°15' E, 5-6 m; WAM

P.28298.004 (24, 42-118) Lucky Bay, 34°08' S, 122°15' E, 24

m; WAM P.28513.006 (92) Duke of Orleans Bay, 33°54' S,

122°37' E, 5 m; WAM P.28523.006 (3, 38—70) Augusta, 34°19'

S. 115°10' E, 12-13 m. New Zealand, North Island: AMS

I.18281-005 (6, 75—90) Auckland, Goat Island, 18 m, 1975;

AMS 1.18282-003 (8, 65-105) Auckland, Goat Island, 57°S,

168°E, 20 m, 31 March 1975; NMNZ P.002281 (4, 80—106)

southeast of Mayor Island, 37°22.50' S, 176°22.00' E, 207—219

m, 27 February 1957; NMNZ P.002289 (69) south of Mayor

Island, 37°19.5' S, 176°16.5' E, 102 m, 27 February 1957;

NMNZ P.002299 (102) Manakau Harbour, above Weymouth,

37°2.45' S, 174°50.25' E, 0-5 m, 5 January 1957; NMNZ

P.003151 (10, 78—117) Urupukapuka Island, Bay of Islands,

35°12.32' S, 174°14.40' E, 2-7 m, February 1961; NMNZ

P.003616 (2, 73—79) 22.5 km east of White Island, 37?30' S,

177°26' E, 192 m, 3 April 1963; NMNZ P.003756 (130) Napier

breakwater, 39°28.50' S, 176°55.25' E, 9 m, March 1964;

NMNZ P.005275 (132) off Kaipara Harbour, 36°26.0' S,

173*50.5' E, 115 m, 9 January 1971; NMNZ P.005866 (2, 93—

109) northeast of Motiti Island, 37°39' S, 176°33' E, 64 m, 14

April 1972; NMNZ P.005969 (2, 93—99) main channel off

Kopumiti Point, Whangaroa Harbour, 35°2' S, 173*45' E, 9-15

m, 22 February 1974; NMNZ P.006801 (75) northern

Taranaki Bight, Aotea Seamount, 38°13.5' S, 173°54.0' E,

135-137 m, 13 November 1971; NMNZ P.007794 (8, 51-72) 9

km north of Motuhara Island, 37°48.15' S, 177°1.10' E, 72-84

m, 20 January 1979; NMNZ P.007865 (10, 60—83) north of

Mayor Island, 37°10.90' S, 176°10.75' E, 198-273 m, 22

January 1979; NMNZ _ P.007899 (3, 55-110) between

Motuhara Island and Rurima Inlets, 37°51.75' S, 176°55.90' E,

34—39 m, 21 January 1979; NMNZ P.008349 (4, 68-121)

north of Mayor Island, 37°10.90' S, 176°10.75' E, 198—273 m,

22 January 1979; NMNZ P.008362 (9, 54—79) east-northeast

of Tolaga Bay, 38°15.2' S, 178°38.6' E, 139 m, 16 January

1979; NMNZ P.008364 (12, 15-86) off Mayor Island, 37°9.1'

S. 176°24.4' E, 753-826 m; NMNZ P.008461 (108) off

M.F. Gomon, C.D. Struthers & J. Kemp

Kohinga point, Bay of Islands, 35710' S, 174°10' E, 5 December

1973, NMNZ P.008463 (74) Goat Island, Leigh, 36°16' S,

174°48' E, 1970; NMNZ P.008472 (120) Whatawhiwhi,

Doubtless Bay, 34°53.00' S, 173°24.25' E, 17 November 1963;

NMNZ P.009003 (2, 73—76) Spirits Bay, Northland, 34°27' S,

172°50' E, 4 m, 14 November 1963; NMNZ P.009796 (56)

southeast of Three Kings Islands, 34°20.2' S, 172*21.8' E, 121

m, 2 February 1981; NMNZ P.009802 (75) Ranfurly Bank,

East Cape, 37°38.4' S, 178°51.7' E, 79—83 m, 22 January 1981;

NMNZ P.009809 (3, 49—71) off Ninety Mile Beach, 34°41.9'

S, 172°33.5' E, 103 m, 10 January 1981; NMNZ P.009813 (61)

off Raglan Harbour, 37°48.0' S, 174°14.7' E, 103 m, 13 January

1981; NMNZ P.009817 (4, 68—90) off Parengarenga Harbour,

34°32' S, 173°6' E, 93-102 m, 27 January 1981; NMNZ

P.009828 (6, 59—122) northwest Ahipara, 34°50.0' S, 172°46.1'

E, 90 m, 10 January 1981; NMNZ P.010431 (63) off Mahia

Peninsula, 39°10' S, 178*0' E, 15 m, 14 April 1981; NMNZ

P.011874 (95) 2.4 km west of Cape Colville, Hauraki Gulf,

36°37' S, 175°28' E, 40-70 m, 17 February 1976; NMNZ

P.012089 (58) northeast of Ninepin Rock, Bay of Islands,

35*8.8' S, 174°10.9' E, 66—79 m, 1 December 1971; NMNZ

P.012091 (103) North Cape, 34°25' S, 173°3' E, 109-146 m, 8

October 1964; NMNZ P.012093 (108) north of the Noises,

Hauraki Gulf, 36°42' S, 174*58' E, 46 m, 17 May 1965; NMNZ

P.014301 (12, 72-120) Matai Bay, south side of middle

headland, Othngahunga Bay, 34°50! S, 173°25' E, 7-13 m, 18

August 1983; NMNZ P.014349 (2, 90—97) Waikato Bay, south

end of Matai Bay, Northland, 34^50' S, 173°25' E, 3-5 m, 19

August 1983; NMNZ P.014380 (2, 100—109) north end of

Taupo Bay, 34°59.17' S, 173°43.00' E, 7-8 m, 20 August 1983;

NMNZ P.014397 (3, 75—92) Oakura Bay, Northland, 35°23' S,

174°21' E, 4-6 m, 21 August 1983; NMNZ P.015199 (108)

channel between Henry Island and Cape Home, Oakura Bay,

35°23.0' S, 174?22.2' E, 14-17 m, 23 February 1984; NMNZ

P.015305 (110) south of Tutukaka Harbour entrance,

Northland, 35°37.36' S, 174°32.55' E, 13 m, 19 February 1984;

NMNZ P.016983 (34) seawater intake, New Plymouth

Powerhouse, 39°3' S, 174°5' E, 1983; NMNZ P.018115 (110)

off marine laboratory, Island Bay, 41721' S, 174°45.89' E, 15

m, 22 January 1986; NMNZ P.018200 (3, 850—111) reef in

northern Crater Bay; Mayor Island, 37°17.2' S, 176°16.4' E, 12

m, 19 February 1986; NMNZ P.018223 (96) Mount Manganui,

Bay of Plenty, 37°37.5' S, 176°10.4' E, 0—3 m, 13 February

1986; NMNZ P.018258 (88) Tauranga Bay, Mayor Island,

37°18.4' S, 176715.8' E, I2 m, 18 February 1986; NMNZ

P.018290 (3, 62-100) off Rabbit [Motuotou] Island, Mount

Maunganui, 37°38.0' S, 176°11.6' E, 6-15 m, 14 February

1986; NMNZ P.021081 (7, 83-134) off Bell Block, New

Plymouth, 39°1' S, 174°9' E, 9 m, 13 December 1986; NMNZ

P.O21541 (4, 52-83.2) rocks off Whangamata Beach,

Coromandel, 37°12.88' S, 175°53.65' E, 17 m, 29 November

1987; NMNZ P.021645 (72.8) east side of Great Mercury

Island, Coromandel, 36°37.05' S, 175°50.23' E, 15 m, 2

December 1987; NMNZ P.021767 (3, 59—89.3) pools between

Jackson and Fantail Bays, Coromandel, 36°32' S, 175°20' E,

0-3 m, 8 December 1987; NMNZ P.021797 (10, 72.1—99.8)

rocks off Whangamata Beach, Coromandel, 37°12.88' S,

175°53.65' E, 17 m, 29 November 1987; NMNZ P.021883 (95)

Australasian red cods

Firth of Thames, 37°0' S, 175°20' E, 30 January 1965; NMNZ

P.023179 (93) Matakaoa, East Cape, 37°34' S, 178°20' E, 16 m,

25 June 1988; NMNZ P.023211 (6, 81—113) Waiaka Bay near

Lottin Point, East Cape, 37°33' S, 178°9' E, 6 m, 24 June 1988;

NMNZ P.024349 (72) Higgins Wharf, Napier, 39°30' S,

176*50' E, 22 March 1988; NMNZ P.026295 (110) Charity

Reef, Pourerere, 40°6.2' S, 176°53.5' E, 12 m, 19 January 1991;

NMNZ P.026356 (66) Pauanui Point Reef, East Coast, 40°4.4'

S, 176°53.8' E, 7-10 m, 16 January 1991; NMNZ P.028100 (4,

509—120) Okakari Point, Northland, 36°15' S, 174°46' E, 2—3 m,

9 April 1992; NMNZ P.028204 (5, 86-151) Midway Point,

south of Lottin Point, East Cape, 37°32.6' S, 178°12.9' E, 15—

17 m, 2 May 1992; NMNZ P.028237 (5, 59-110) west side of

Orete Point, Te Kaha, 37°35.25' S, 177°53.15' E, 11-12 m, 3

May 1992; NMNZ P.028271 (2, 95-131) bay west of Lottin

Point, Waiaka Bay, East Cape, 37°32.9' S, 178°8.7' E, 15-20

m, | May 1992; NMNZ P.028389 (2, 62—68) south of Tohora

Pirau, Lottin Point, 3732.7 S, 178°10.0' E, 7-10 m, 2 May

1992; NMNZ P.028428 (4, 74—87) Kaipiro Reef, Maraetai

Bay, Te Kaha, Bay of Plenty, 37*42.6' S, 177°41.7' E, 19-21 m,

30 April 1992; NMNZ P.028460 (98) Kaipiro Reef, Maraetai

Bay, Te Kaha, Bay of Plenty, 37°42.6' S, 177°41.7' E, 19-21 m,

30 April 1992; NMNZ P.029790 (2, 105—130) off Horoera

Point, East Cape, 37°37.8' S, 178°28.9' E, 8—10 m, 24 January

1993; NMNZ P.029846 (2, 57—64) between Moutahiauru

Island and Koutunui Head, East Cape, 38°3.35' S, 178°22.20'

E, 9 m, 25 January 1993; NMNZ P.029855 (3, 60-75)

Whanarua Bay, Bay of Plenty, 37°40.0' S, 177*46.7 E, 15-21

m, 28 January 1993; NMNZ P.029875 (2, 62-75) between Te

Araroa and Horoera, East Cape, 37°37.6' S, 178°25.0' E, 03

m, 24 January 1993; NMNZ P.029969 (8, 45—65) northeast

Waipiro Bay, East Cape, 38°0.0' S, 178°23.1' E, 21 m, 25

January 1993; NMNZ P.030070 (3, 63—120) Tatapouri Beach,

Gisbourne, 38°39.2' S, 178*9.9' E, 6-8 m, 22 January 1993;

NMNZ P.030121 (5, 60—75) eastern Whanarua Bay, Bay of

Plenty, 37740.5' S, 177°47.4' E, 8 m, 28 January 1993; NMNZ

P.030607 (2, 70—117) Port of Napier, 39°28' S, 176°55' E, 23

March 1993; NMNZ P.030609 (7, 70—92) Gisborne Wharf,

Poverty Bay, 38°40.5' S, 178°1.5' E, 8 m, 4 December 1992;

NMNZ P.030615 (3, 78—117) Gisborne Wharf, Poverty Bay,

385'4]' S, 178*2' E, 18 May 1993, NMNZ P.030617 (2, 62—73)

Gisborne Wharf, Poverty Bay, 38*40' S, 178°1' E, March 1993;

NMNZ P.030631 (4, 71—104) Port of Napier, 39°29' S, 176°55'

E, 25 June 1993; NMNZ P.030685 (2, 87—106) Port of Napier,

39*21.9' S, 176°54.1' E, 5 m, 22 September 1992; NMNZ

P.030692 (19, 49—103) Gisborne Wharf, Poverty Bay, 38°40.8'

S, 178°1.1' E, 23 June 1993; NMNZ P.033570 (119) Long

Point, Mahia Peninsula, 39°10' S, 177°49' E, 8 m, 4 January

1995; NMNZ P.033644 (6, 50-110) Whangawehi, Mahia

Peninsula, 39°7' S, 177°54' E, 14 m, 25 January 1995; NMNZ

P.034527 (90) off Great Exhibition Bay, Northland, 34°40.4' S,

173°31.0' E, 182 m, November 1977; NMNZ P.035537 (10,

77-105) Archway, east end of Whale (Moutohora) Island,

37*5].44' S, 176°59.39' E, 12-17 m, 1 June 1998; NMNZ

P.035560 (3, 90-140) Rurima Islets, 37°49.78' S, 176°52.63' E,

7-10 m, 2 June 1998; NMNZ P.035584 (8, 82-139) North Bay,

Whale (Motuhora) Island, 37°51.05' S, 176°58.57' E, 13-15 m,

3 June 1998; NMNZ P.036587 (23, 66—112) south Whale

(Motuhora) Island, 37*51.70' S, 176°58.43' E, 8—12 m, 22 April

1999; NMNZ P.036641 (8, 82-118) Nursery Cove, White

Island, 37°31.42' S, 177°10.37' E, 5-16 m, 23 April 1999;

NMNZ P.036667 (7, 77-94) off Homestead Point, White

Island, 37°31.77' S, 177°10.68' E, 8-14 m, 23 April 1999;

NMNZ P.036698 (5, 77-116) west end of White Island,

37°51.05' S, 176°57.63' E, 12-16 m, 24 April 1999; NMNZ

P.036723 (22, 65-95) southeast MacEvans Bay; Whale

(Motuhora) Island, 37°51.57' S, 176°59.12' E, 12-17 m, 26

April 1999; NMNZ P.048348 (2, 62.1—67.9) off Kaweura,

midway between South Head, Omapere and Maunganui Bluff,

35°38.14' S, 173°26.09' E, 0—10 m, 14 February 2011; NMNZ

P.048431 (65.8) first coast beach south of southern headland

of Hokianga Harbour, 35°32.93' S, 173°22.01' E, 16 February

2011. South Island: NMNZ P.033548 (2, 110-133) Foul

Point, Abel Tasman, 40°54' S, 173°4' E, 8 m, 3 December

1993.

Memoirs of Museum Victoria 80: 101-112 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.05

Review of the Australian endemic odontocerid genus Barynema and status of

Australian Marilia (Trichoptera)

(http://zoobank.org/urn:|sid:zoobank.org:pub:827 E8E]E-E6C4-424C-A4A 2-2124128332F0)

ALICE WELLS" (http://zoobank.org/urn:|sid:zoobank.org:author:0D7A8359-1249-4DED-9D5F-DBF5FCD17876)

AND ROSALIND M. ST CLAIR? (http://zoobank.org/urn:|sid:zoobank.org:author:cO04d784-e842-42b3 -bfd3-317d359f8975)

' Australian National Insect Collection, CSIRO, PO Box 1700, Canberra, ACT 2601 Australia.

* Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia

* ‘To whom correspondence should be addressed. Email: alice.wells@csiro.au

Abstract

Wells, A. and St Clair, R.M. 2021. Review of the Australian endemic odontocerid genus Barynema and status of

Australian Marilia (Trichoptera). Memoirs of Museum Victoria 80: 101—112.

In this review of Australian Odontoceridae, we revise details of the two established species 1n the endemic genus

barynema Banks — B. costatum Banks and B. australicum Mosely — and describe six new congeners: B. paradoxum sp.

nov., B. lorien sp. nov., B. lobatum sp. nov., B. dilatum sp. nov., B. dolabratum sp. nov. and B. goomburra sp. nov. For the

only other odontocerid genus recorded for Australia, Marilia Müller, we discuss the present status of the three described

species — M. bola Mosely, M. aenigmata Neboiss and M. fusca Kimmins — and outline our efforts and those of others to

resolve problems of species delimitation based on morphology. We select a new replacement name for Marilia fusca and

provide brief notes on the larvae of both genera.

Keywords

venation, CO1 data, distributions, homonym

Introduction

The family Odontoceridae is one of several groups among

Australian Trichoptera, upon which Arturs Neboiss (late of

Museums Victoria) made some preliminary studies but then

set them aside. In 2003, he published a new Tasmanian

species of Marilia Müller, 1878 (M. aenigmata), thus bringing

to three the Australian species reported for this genus.

Barynema Banks, 1939, the only other odontocerid genus

recorded for Australia, 1s endemic, comprising two established

species. Discrimination between adult males of these two

Barynema species and a further six species, newly described

here, is reasonably easy based on male genitalic features,

augmented for some by thoracic features, but this 1s not so for

discrimination between adult specimens referrable to Marilia.

The genus Marilia has almost a worldwide distribution.

Features of the genus are discussed in detail in works by Oláh

and Johanson (2010) and Yang et al. (2017). Oláh and

Johanson (2010) found that "[m]any species have similar male

genitalia, and it is difficult to differentiate all species by

examining the phallic apparatus alone" (p. 73). Working with

east Asian species, they advocated discrimination of species

based on combined genitalic features and cephalic setal wart

patterns.

The three species of Marilia described from Australia —

Marilia bola Mosely, 1953, and M. fusca Kimmins, 1953

(both described in the work by Mosely and Kimmins, 1953),

and M. aenigmata — all have very uniform male and female

genitalia. Differentiation of the first two appears, on paper, to

be relatively simple. Males of Marilia bola have the eyes

holoptic (almost touching the vertex) and a tuft of long setae

on the anal lobe of the hind wing, whereas males of M. fusca

have the eyes dichoptic (not approximated) and lack the tuft

of setae on the wing. The female of M. bola was described

and illustrated, but not the female of M. fusca. Neboiss, 1n

describing the Tasmanian M. aenigmata, gave a small table of

comparison of features of all three species, noting that M.

aenigmata exceeds M. bola in size and, in comparison with

M. bola, lacks Fork 3 in the female forewing and has hindwing

Fork 1 sessile, not stalked (petiolate).

Confusing any simple separation of species, however, in

preliminary studies on Australian Marilia specimens in the

considerable Museums Victoria collection, Brian Armitage

(in 1990) recognised two major sets among Marilia specimens

from sites along the length of the Great Dividing Range, from

central Victoria to far north-eastern Queensland, and labelled

Specimens accordingly. He designated these Marilia Sp. A

and Marilia Sp. B, based mainly on wing features. In several

samples from a single site, he recognised both forms. Males

of both sets have the compound eyes holoptic and, 1n their

genitalic features, exhibit no clearly consistent differences.

Females are inseparable on genitalic features. Both sexes of

Marilia Sp. B have Fork | in the hind wing petiolate, and the

associated females have forewing Fork 3 present. Thus,

Marilia Sp. B could be Marilia bola.

102

Armitage’s Sp. A has Fork 1 in the hindwing sessile and the

associated females lack forewing Fork 3. It shares these wing

features and genitalic characteristics with M. fusca and M.

aenigmata. However, the eyes of the type specimen of M. fusca

as described by Kimmins (in the work by Mosely and Kimmins,

1953) are dichoptic; thus, Marilia sp. A 1s not conspecific with

M. fusca. Or, conceivably, it could be if the situation is as Olah

and Johanson (2010) determined for their newly described

South-East Asian species Marilia malickyi — that there 1s “[1]

ntraspecific variation in the interocular distance” (p. 74).

Armitage’s Sp. A does conform morphologically with M.

aenigmata. The body size of mainland specimens varies

considerably, with many being much smaller than the

Tasmanian specimens.

The only specimens in the available collections that

concur completely with the description of M. fusca are in

several small samples collected from streams in the

Mount Spec State Forest (to the north of Townsville in north-

east Queensland) and a stream near Tully. Surprisingly,

no dichoptic males have been identified among samples

from further south to the Sydney region or the Oxley Rivers

region of New South Wales from whence the type and

paratypes, respectively, were taken, nor further to the north

in Queensland.

None of this is particularly satisfactory and does not

corroborate the equally unsatisfactory preliminary results

from the limited patchy and geographically scattered COl

data from adults and larvae that are available on the

Biodiversity of Life Database (BOLD; http://www.

boldsystems.org). The BOLD data suggest there are at least

eight species of Marilia in Australia, most with large genetic

distances of at least 10% between them. This is apparent from

the very limited number of specimens available for genetic

analysis. More collecting and analysis will probably reveal

further species. Three specimens from north-east Queensland,

from localities from which specimens identified by Armitage

as sp. B were collected, have venation that differs from both

M. aenigmata and M. bola.

similarly puzzling 1s the available information on larvae

of Australian species of Marilia. Larvae and pupae from the

Yarra River at Reefton, Victoria, were described by

Drecktrah (1990) as those of M. fusca. He gave no indication

of how the identification was made, and it 1s impossible to be

certain from the anterior view if the pupal head illustrated by

Drecktrah would have emerged as female or male, holoptic or

dichoptic. It now seems very unlikely that the larvae

described by Drecktrah are M. fusca. In 2004, Dean et al.

published a key for discrimination of late instar larvae of

Barynema and Marilia, and separation of three species of

Marilia, identified as M. bola, M. fusca and M. aenigmata;

photographic images are given to illustrate diagnostic

features used in the key. Marilia fusca and M. aenigmata

were separated in the key simply by geography, and the

authors comment that their larvae are similar and may

eventually be found to be the same species. In recent

collecting of Marilia larvae from New South Wales and

Queensland, St Clair (unpublished) has noticed further small

differences. Identification of larvae of Marilia species 1s

A. Wells & R.M. St Clair

therefore not possible until larvae can be associated with

known adults. All available specimens can be assigned to

two larval types that align with the names given by Dean et

al. (2004). These are probably better labelled “Marilia larval

sp. 1" (was bola) and “Marilia larval sp. 2” (was fusca) since

such confusion is now recognised. Further studies involving

association of adults and close morphological examination of

larvae are required.

Marilia fusca 1s a junior secondary homonym of the name

Anisocentropus fuscus Banks, 1905 (synonymised with

Marilia flexuosa Ulmer, 1905, by Betten, 1934), so here we

select a replacement name.

Overall this 1s a confusing mix, probably only solved by a

thorough and comprehensive combined morphological and

molecular phylogenetic study. For the present, it 1s not possible

to assign Australian specimens of Marilia to species. Thus,

we can offer no further insight into Australian Marilia. The

late Arturs Neboiss would understand.

Material and Methods

Most of the material studied is in the collection of Museums

Victoria, Melbourne (NMV) and bears the standard NMV

registration number. Genitalic specimens that were

macerated and cleared by Neboiss are tagged with PT-

numbers and several others have WTH (Wet ‘Tropics

Heritage) numbers or BOLD identifications. A small number

of specimens are in the Australian National Insect Collection

(ANIC), Canberra, or will be lodged in the Queensland

Museum (QM). Images of types in the Natural History

Museum, London (BMNH) were provided by Dr Ben Price.

Other images were taken at ANIC by AW and were prepared

using a stereomicroscope linked to a Leica Application Suite

(version 4.2) to integrate multiple images; some of the images

are of cleared, others of intact, specimens. Plates were

prepared in Adobe Photoshop (version 12).

The terms applied to wings follow Mosely and Kimmins

(1953), and the terms applied to genitalic structures follow

Yang et al. (2017), save for structures that 1n ventral view

appear beside the phallus, for which we follow Mosely and

Kimmins in using the term “upper penis cover’. This latter

structure could represent tergite X or the phallobase, but it

appears to arise further ventrad than one would expect for the

former and to be independent of the phallus, so probably is

not homologous with the latter.

Taxonomy

Marilia Miller

Marilia fusca Kimmins (in the work by Mosely and Kimmins),

1953: 167, fig. 112.

Marilia disjuncta nom. nov., new replacement name for Marilia

fusca Kimmins, preoccupied by Anisocentropus fuscus Banks, 1905,

a junior synonym of Marilia flexuosa Ulmer, 1905.

Australian Odontoceridae reviewed

Barynema Banks

Figures 1—35

Barynema Banks 1939: 483, type species: Barynema costatum

Banks, 1939 by monotypy.

In their treatment of the Australian endemic genus Barynema,

Mosely and Kimmins (1953) discussed its early placement in

the family Calamoceratidae and their rationale for transferring

the genus to the family Odontoceridae on the grounds that

Barynema lacks the median cell in the forewing. Adults of

Barynema species are distinguished from Australian

representatives of Marilia by having a pair of setose warts on

the mesoscutum, whereas Marilia has two rows of setae;

males of many specimens of Marilia have holoptic compound

eyes, not seen in any Barynema. Two species of Barynema —

B. australicum Mosely, 1953, and B. goomburra sp. nov. —

also have a pair of setate warts posteriorly on the

mesoscutellum, a characteristic that appears to be absent in

all other Odontoceridae. This could be considered a

distinguishing feature at the genus level but given the very

close similarity of other male and female features of species

assigned to Barynema, we believe all eight species are

appropriately referred to a single genus. Similar variation in

presence and absence of mesoscutal and mesoscutellar setal

warts and their arrangement is seen in the closely related

Philorheithridae (Neboiss, 1977).

In the inferior appendages of the males of some species of

Barynema (e.g. B. costatum and B. australicum), the harpago

appears to be fused with the coxopodite (or lost?). In other

species, a rather similar structure (when viewed laterally)

appears to be simply an extension of the coxopodite that 1s

probably homologous with the structure termed

"subapicodorsal lobe of an inferior appendage” by Yang et al.

(2017, p. 88). The socketed harpago forms a smaller, but

morphologically similar, structure (e.g. B. lobatum sp. nov.)

or is reduced to a small triangular (B. dilatum sp. nov.) or

subquadrate (B. dolabratum sp. nov.) lobe laterad of the

extended coxopodite; it usually bears an area of short peg-like

black setae. A similar situation appears to occur in the east

Asian species Psilotreta malickyi Oláh and Johanson, 2010,

although in that species the reduced harpago is a mesal

structure. In females of Barynema, the apical lobes are stout

structures on abdominal segment X, while in Marilia they are

absent or fused with tergite X.

Here, new records and images are given for each of the

two established species of Barynema: B. costatum from

Victoria and B. australicum from north-eastern New South

Wales. Thoracic and female genitalic features are described

and illustrated for these species. In addition, six new species

are described, based on male and female thoracic and

cenitalic features, and a key is provided for the genus. The

genus has been collected along the Great Dividing Range of

eastern Australia but has not been recorded further west; in

the south, the genus 1s not recorded from west of the central

ranges of Victoria or from Tasmania. Genetic data from

BOLD suggest that additional species are present in Australia.

Larvae of a few species can be associated with the adult

103

using the COI barcode. The larvae show subtle differences

that may enable species separation, but this 1s best left until

more species are associated. The larval and pupal descriptions

of Barynema costatum by Cartwright and Dean (1987) may

not be that species and may not be adequate to separate larvae

of related species.

Key to males of Barynema Banks.

l. Mesoscutellum subcircular to rectangular, bearing small

paired setate warts separate from each other and close to

posterior margin (figs 29, 33) 2

- Mesoscutellum ovoid, with large paired setate warts

fused and covering most of mesoscutellum (fig. 5) 3

2. Inferior appendages, each with harpago in ventral view

bearing abbreviated area of short stout black peg-like

setae subapicomesally; coxopodite with short mesal lobe

bearing similar, but tapered, setae (fig. 32) l..

B. australicum Mosely

- Inferior appendages, each with harpago in ventral view

bearing elongate area of short stout black peg-like setae

subapicomesally; coxopodite without mesal lobe (fig. 35)

B. goomburra sp. nov.

3. Inferior appendages, each with brush of short, stout black

setae on harpago subapicomesal, elongate, extending

along distal half of mesal margin (figs 3, 6) oon

B. costatum Banks

- Inferior appendages, each with brush of short stout black

setae on harpago apical and rounded, not elongate (figs 9,

13, 18, 21, 25) 4

4. Lobes of upper penis cover in ventral view expanded

apicolaterally (figs 21, 25) 5

- Lobes of upper penis cover in ventral view tapered,

acuminate to rounded apically, but not expanded

apicolaterally (figs 9, 13, 15, 18) 6

5. Apicolateral lobes of upper penis cover in ventral view

rounded (fig. 21) B. dilatum sp. nov.

- Apicolateral lobes of upper penis cover in ventral view

sharply triangular, pick-shaped (fig. 25) oe

B. dolabratum sp. nov.

6. Inferior appendages, each with coxopodite in ventral

view about 2 times as long as wide (figs 13, 15)

B. lorien sp. nov.

- [nferior appendages each with coxopodite in ventral view

subquadrate (figs 9, 18) 7

7. Coxopodites in ventral view, each with mesal margin

rounded (fig. 18) B. lobatum sp. nov.

- Coxopodites in ventral view, each with mesal margin

angled obliquely from base (fig. 9) . B. paradoxum sp. nov.

104

Barynema costatum Banks

Figures 1, 3—8

Barynema costatum Banks, 1939: 484, figs 43, 51, 54.

Material examined. Holotype. male, Victoria, Mount Donna Buang,

6—7 December, Darlington (ANIC, ex CZM, dry on a pin). Victoria: 1

male (dissected), Cement Creek near Warburton, 4.x1.1972, P. Zwick

(NMV Tri-26514, PT-1389). Victoria: 1 adult female, same data

(NMV Tri-26517, PT 1890); 1 male (pharate adult, dissected), small

trib. Snobs Creek, Snobs Road crossing bridge, 11.6 km u/s Eildon

Road, 31.x.1981, J. Dean (NMV Tri-26506); 1 female (reared), Back

Creek, 1 km NE of Noorinbee, 13.x.1982, A. Bolton, reared (NMV

Tr1-26518).

Diagnosis. Males, when freshly caught or dried, have on the

black forewing a band of white and golden hairs angled across

the wing at about 3/5 length and a streak of golden scales and

hairs between Al and Cu2; on the inferior appendages, a rather

elongate hairbrush-like cluster of short and blunt black setae

line the inner subapical or apical region of the harpago, features

that distinguish them from B. paradoxum sp. nov. with similar

black wings, but with the distal band of hair on the forewing

wing only slightly curved and the harpago with a rounded

cluster of short black setae apically; in ventral view, the basal

section of the coxopodite is broadly subrectangular in B.

costatum, but more rounded in B. paradoxum, and the upper

penis cover with each lobe expanded laterally toward the apex,

rather than tapered or rounded.

Description (revised after Mosely and Kimmins, 1953).

Mesothorax with scutellum subquadrate, without

mesoscutellar setate warts.

Male. Body and wings (figs 1, 3, 4) black, forewings each

with gold band along proximal section of Al and white band

across vein anastomoses at about 3/5 wing length; length of

each forewing 9.5 mm (n = 1). Mesoscutellum ovoid, without

setate warts. Abdominal sternite VII bearing median

subquadrate lobe. Genitalia: Pre-anal appendages in dorsal

view stoutly conical, about length of inferior appendages; upper

penis cover elongate, wrapped lateroventrad of phallus, flared

and widest towards apex; inferior appendages in ventral view

with coxopodite stout, subquadrate basally; harpago in ventral

view slender, with elongate apicomesal brush of sharp, stout

black setae lining distal half, giving hairbrush-like appearance.

Female. Terminalia: Distal abdomen bearing pair of short,

stout, apically truncate apical lobes; striated plates ventrally

on segment IX.

Distribution. Found in central and eastern Victoria.

Remarks. Very few specimens of B. costatum have been

collected — many of the specimens previously identified as B.

costatum are assigned here to the new species Barynema

paradoxum sp. nov. BOLD data groups the two, with only a

short distance between them. One of the few confirmed B.

costatum specimens 1s a pharate pupa from a tributary of Snobs

Creek on the north-western edge of the Great Dividing Range

in central Victoria, images of which are included here (figs

5—8), particularly to show the form of the mesoscutellum that is

obscured by the pin on the dried type.

A. Wells & R.M. St Clair

Barynema paradoxum sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:62A5E978-BCFE-

49A3-829D-4FO8FB542881

Figures 2, 9-12

Material examined. Holotype. male, New South Wales, 8 km N of

Nowra, 21.x.1966, N.D. (NMV Tri-27118, P'T-1646).

Paratypes. New South Wales: 2 males, data as for holotype,

21.x.1966, N.D. (NMV Tri-27113); 4 males, data as for holotype,

21.x.1966, N.D. (NMV Tri-27115); 1 male, Minnamurra Falls, N of

Kiama, 25.111.1973, A. Neboiss (NMV Tri-27114); 2 males, Mumbulla

Creek, near Bega, 20.x1.1978, I. Campbell (NMV Tri-27110); 3 males,

| female, Minnamurra Falls, 22.x.1982, G. Theischinger (NMV Tri-

27139); 2 males, Bundanoon, 34° 39' S 150° 18' E, 1x.1983, G.

Theischinger (NMV Tri-27137).

Other material. New South Wales: 1 male [pinned], Brown

Mountain, 2.x11.1956, E.F. Riek (ANIC); 2 males [pinned], Macquarie

Falls, 14.1x.1960, D.H. Colless (ANIC); 3 males [pinned], upper

Kangaroo Valley, 24.1x.1960, E.F. Riek (ANIC); 1 male, eastern Fall,

Clyde Mountain, 26.x.1960, E.F. Riek (ANIC); 2 males, 1 female,

Minnamurra Falls, 23.x.1962, D.K. McAlpine (NMV Tri-27062; 2

males, 1 female, Minnamurra Falls (NMV TRI-27062); 1 male, 1

female, New South Wales, Macquarie Pass, 13x.1986, "G. Thei. & L.

Mu." [G. Theischinger and L. Muller] (NMV Tri-27133); 1 male, New

South Wales, Macquarie Rivulet, W of Shell Harbour, 34° 34' S

150° 41' E. 4.1.1990, G. Theischinger (NMV Tri-27134); 1 male, same

data (NMV Tri-27070); 1 male, Bundanoon, Fairy Bower, 35° 39' S

150° 18' E, 1x.1991, G. Theischinger (NMV Tri-27135); 2 male, same

data (NMV TRI-27137); 1 male, same data (NMV TRI-27138); 1

female, New South Wales, Yadboro Creek, Budawang Range, 40 km

SW of Nowra, 35° 22' S 150° 03' E, x1.1991, G. Theischinger (NMV

1r1-27136); 1 female, Yadboro Creek, Budawang Range, 40 km SW of

Nowra, x1.1991, G Theischinger, 35° 22' S 150° 03' E (NMV TRI-

27136); 1 male, South East Forests National Park, trail near Monaro

Highway, malaise in Hopping Joe Creek, 37° 12' 58" S 149° 18' 37" E,

5.x11.2004—12.1.2005, C. Lambkin, N. Starick (ANIC); 16 males 1

female, East Boyd State Forest, Anteaters Road, 55 km SE Bombala

Forest, 37? 12' 18" S 149? 42' 24" E, 6.x11.2004—-12.1.2005, C. Lambkin,

N. Starick (ANIC); 1 male, New South Wales, Polblue Creek,

Barrington Tops Forest Road, 31° 57' S 151° 26' E, 2.x11.2007, A.

Glaister, J. Dean and R. St Clair (NMV Tri-54560); 1 male, tributary

of Macquarie Rivulet off Clover Hill Road, 29.1x.2017, Z. Billingham

(NMV Ento 2018-246, JOS-424); 1 male, Minnamurra Creek, off

Minnamurra Rainforest Walk, 3.x.2017, Z. Billingham (NMV Ento

2018-246, JOS-423); 13 males, 1 female, Budawang National Park,

Mount Budawang Road, 35.4563 S 149.4455 E, 16.1.2019, K.M.

Bayliss, J. Lumbers and D.K. Yeates (ANIC). Victoria: Coopracambra

National Park, malaise trap over Beeline Creek, 27 km NNE of Cann

River, 5.x11.2004—12.1.2005, C. Lambkin, N. Starick (ANIC).

Diagnosis. Males of Barynema paradoxum resemble those of

B. costatum in having distinctive black wings with a median

white band across the area of vein anastomoses and a band of

bright yellow hairs between veins Cu2 and Al. But in B.

paradoxum, the white band 1s slightly curved, not angled, and

the area of golden setae is smaller. Like B. costatum, B.

paradoxum shares with B. dilatum sp. nov., B. lorien sp. nov.

and B. dolabratum sp. nov. the absence of setate warts on the

mesoscutellum and, as with those three species, the male of

this new species has a rounded club of short, stout black setae

apically on the distal extremity of each inferior appendage, not

an elongate brush as in B. costatum. Together with B. lorien

Australian Odontoceridae reviewed 105

LI »

"s i

EI is ^

Figures 1—8, Barynema species. 1, 3, 4, Barynema costatum Banks holotype male: 1, body and wings, dorsal; 3, 4, genitalia, ventral and lateral,

respectively. 2, B. paradoxum sp. nov., New South Wales, Upper Kangaroo Valley, body and wings, dorsal; 5-8, B. costatum, male, Victoria,

tributary of Snobs Creek (NMV TRI-27406): 5, mesothorax, dorsal; 6—8, genitalia, ventral, lateral, and dorsal, respectively.

Abbreviations: cox. = coxopodite; har. = harpago; ph. = phallus; pre. app. = pre-anal appendages; up. p. = upper penis cover.

106

and B. lobatum sp. nov., B. paradoxum has the ventrolateral

lobes of the upper penis cover tapered apically, rather than

expanded. Barynema paradoxum is characterised by the

coxopodite of each male inferior appendage subquadrate, not

longer than wide in ventral view and not produced to form a

rounded lobe at its apicomesal angle.

ore. app.

har.

COX.

A. Wells & R.M. St Clair

Description. Mesothorax without mesoscutellar setate warts.

Body and wings black, each forewing with slightly curved band

of white setae across vein anastomoses.

Male. Length of each forewing 6.2-9.7 mm (n = 10).

Abdominal segment VII bearing small, sharp, sclerotised

median spur. Genitalia: Pre-anal appendages stout, apices

pre.app. 1l

Figures 9-17, Barynema species. 9-12, B. paradoxum sp. nov., holotype, New South Wales, Nowra (NMV TRI 27118, PT-1646): 9-11, male

genitalia, ventral, lateral, and dorsal, respectively; 12, female, New South Wales, Yadboro Creek, Budawang Range, 40 km SW of Nowra (NMV

T RI-27136) terminalia, ventral. 13—17, B. lorien sp. nov.: 13, 14, holotype male, New South Wales, Lansdowne via Taree (NMV TRI-27061, PT-

1564), genitalia ventral and lateral, respectively; 15, 16, paratype male, Queensland, Lamington National Park (NMV TRI-27061), genitalia,

ventral and lateral, respectively; 17, paratype female (NMV TRI-27061), terminalia, ventral.

Abbreviations: api. lob. = apical lobe; cox. = coxopodite; har. = harpago; ph., phallus; pre. app. = pre-anal appendages; up. p. = upper penis cover.

Australian Odontoceridae reviewed

narrowly rounded in dorsal view; lobes of upper penis cover

in ventral view tapered to acuminate apices; inferior

appendages, each with coxopodite in ventral view stout,

subquadrate basally, harpago about length of coxopodite,

Slender with a cluster of short, bristle-like dark setae on

rounded club-shaped apex.

Female. Length of each forewing 9.0 mm (n = 1).

Terminalia: Distal abdomen bearing pair of short apical lobes,

stout basally, rounded laterally and tapered to rounded apices;

striated plates ventrally on segment IX.

Distribution. Widespread in eastern New South Wales,

extending into far eastern Victoria, possibly overlapping with

B. costatum.

Etymology. Named for the past confusion over this species, and

the paradox presented by the variability in male genitalic

structures. Most specimens were identified previously as

b. costatum.

Barynema lorien sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: E8CB61 BO-BA99-

4CC9-8A92-E94F3037EISD

Figures 13-17

Material examined. Holotype. male, New South Wales, Lansdowne

via Taree, 29.1x.1985, G.W. Williams (NMV TRI-27071, PT-1564)

Paratypes. Queensland: 2 males, 1 female, [Lamington| National

Park, 1.x1.1954, T.E Woodward (NMV TRI-27060); 1 male,

Lamington National Park, 15.x1.1955, Yeo (NMV TRI-27069); 1 male

(reared from pupa), Tamborine Mountain, 111.1962, K. Korboot (NMV

TRI-27068); 1 female (reared from pupa), same data (NMV TRI-

27067); 3 males, | female, Lamington National Park, x1.1982, T.

Hinger (NMV TRI-27061); 1 male, Lamington National Park, x1.1982,

G. Theischinger (NMV TRI-27065).

Other material. New South Wales: 1 male 1 female, Wentworth

Falls, 22.x1.1960, C.N. Smithers (NMV TRI-27059); 1 male, same

data (NMV TRI-27070); 1 male, [Barrington Tops State Forest],

Manning River, Pheasant Creek Road, 3.x11.2007, A. Glaister, J. Dean

and R. St Clair (NMV TRI-54568, JOS-237).

Diagnosis. Resembling B. paradoxum, B. costatum, and

B. lobatum sp. nov. in having males with the pre-anal

appendages stout, apically rounded. The male differs from

that of B. costatum in having each inferior appendage

terminating in a small rounded area of short black setae, not

an elongate brush. It is distinguished from B. costatum by

having the lobes of the upper penis cover in ventral view

acute apically, not flared distally as in B. costatum, and 1s

distinguished from B. paradoxum by having the basal region

of each inferior appendage subrectangular, not subquadrate,

and from B. lobatum sp. nov. which has the mesal angle more

strongly produced, rounded. Females resemble those of

B. lobatum, having the apical lobes short, stout, and stepped

(obliquely truncate).

Description. Mesothorax without mesoscutellar setate warts.

Male. Length of each forewing 6.8-7.5 mm (n = 4).

Abdomen with small sharply pointed spur medially on

sternite VII. Genitalia: Pre-anal appendages stout, rounded

apically. Upper penis cover forming pair of elongate lobes of

107

more or less uniform width for most of length, tapered

slightly to apex. Inferior appendages each with coxopodite

basally long, subrectangular to ovoid in ventral view, without

any tufts of setae or mesal lobes, harpago slender at base,

slightly dilated distally, with round pad of short, stout black

setae apically.

Female. Length of each forewing 8.3—9.6 mm (n = 3).

lerminalia: Apical lobes, slightly stepped toward rounded

apices.

Distribution. Found from eastern New South Wales, from the

Blue Mountains to the west of Sydney to south-eastern

Queensland.

Etymology. Named after the Lorien Wildlife Refuge and

Conservation Area near Lansdowne, New South Wales.

Barynema lobatum sp. nov.

http://zoobank.org/urn:Isid:zoobank.org:act:5985 E223-FCó65$-

48BA4-A1C4-19D51 BDS5E0C2

Figures 18—20

Material examined. Holotype. male, Queensland, Montville, Bon

Accord Falls, 27.1x.1955, A.N. Burns (NMV Tri-27064).

Paratypes. Queensland: 3 male, 1 female, Montville, Bon Accord

Falls, 29.1x.1955 A. N. Burns (NMV TRI-27063); 1 male, same data

(NMV '11-27066); 1 male, south-east Queensland, Stony Creek,

Conondale Range, 10.vii1.1988, S. Bunn (NMV Tri-27120).

Diagnosis. In most features, 5. lobatum sp. nov. resembles B.

lorien sp. nov. but alone among congeners, this species has the

apicomesal angle of the otherwise subquadrate coxopodite of

the inferior appendages produced and rounded distally; the

pre-anal appendages are tapered distally, and a small rounded

mesal process occurs on sternite VII. The female is

indistinguishable from those of B. lorien, both having stepped

(obliquely truncate) apical lobes.

Description. Thorax with mesoscutellum almost subquadrate,

pale, margin dark, lacking setate warts.

Male. Length of each forewing 5.2-7.2 mm (n = 5).

Sternite VII with small, apically rounded median tab.

Genitalia: Pre-anal appendages stout in dorsal view, tapered

to rounded apices, in lateral view almost same width

throughout, apically rounded. Lobes of upper penis cover in

ventral view constricted distally to form sharp tips, in lateral

view stout, rounded apically. Inferior appendages each with

coxopodite in ventral view stout, apicomesal angle slightly

produced, rounded; harpago narrow near base, cluster of

short, dark, bristle-like setae apically.

Female. Length of each forewing 8.6 mm (n = l).

Terminalia: Distal abdomen bearing pair of apical lobes, stout

basally, narrowed laterally, and produced to rounded apex.

Distribution. Known only from north of Brisbane in the

Sunshine Coast region of south-eastern Queensland.

Etymology. Named for the expanded apicomesal angles of the

inferior appendages.

108 A. Wells & R.M. St Clair

pre. app.

d. lob.

pre. app.

har.

| d. lob.

m v- 7 Pa

7 "

-s A p. -.

Figures 18-26, Barynema species. 18-20, B. lobatum sp. nov.: 18, holotype male, Queensland, Montville, 27.1x.1955, (NMV Tri-27064), genitalia

ventral; 19, paratype male (NMV 27120), lateral; 20, paratype female (NMV TRI-27063) terminalia, ventral. 21—24, B. dilatum sp. nov.: 21—23,

holotype, male (NMV WTH-2966, PT-2033) genitalia ventral, lateral, and dorsal, respectively; 24, paratype female (Python Creek NMV),

terminalia, ventral. 25, 26, B. dolabratum sp. nov. holotype male, (Oliver Creek NMV), genitalia, ventral and lateral, respectively.

Abbreviations: api. lob. = apical lobe; d. lob. = dorsal lobe of inferior appendages; har. = harpago; ph. = phallus; pre. app. = pre-anal appendages;

up. p. = upper penis cover.

Australian Odontoceridae reviewed

Barynema dilatum sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:26539E9C-AFA3-

4F1B-BD16-B4C2F76A38E8

Figures 21—24

Material examined. Holotype. male, north Queensland, Carron Creek,

Kirrama State Forest, April 1993, G. Theischinger, 18° 06'S 145° 41' E

(NMV WTH-2966 PT-2033).

Paratypes. North Queensland: 5 males, Carron Creek, Kirrama

State For., 17° 50' S 145° 35' E, April 1993 (NMV WTH-1171); 5

males, same data (NMV WTH-1174); 5 males, same data (NMV

WTH-1172); 5 males, same data (NMV WTH-1173); 5 males, same

data (NMV WTH-1174); 5 males, same data (NMV WTH-1175); 5

males, same data (NMV WTH-1176); 3 males, same data (NMV

WTH-1345); 7 males, north Queensland, Goodard Creek, Kirrama

state Forest, April 1993, G. Theischinger, 18° 06' 5 145° 41' E(NMV

WTH-1040); 7 males, same data (NMV WTH-1041); 17 males, 6

females, stream on Tully Gorge Road, 6.7 km d's of power station, M.

Shackleton and J. Mynott, 8.v.2011 (NMV Tri-54556).

Other material. North Queensland: 3 females, upper Mulgrave

River via Gordonvale, 29-30 April 1970, S.R. Curtis (NMV WTH-

0786); 5 male, same data (WTH-2654); 1 female, Lock-Davies Creek

Road, Lamb Range, Mareeba District, 10 November 1974, M.S.

Moulds (WTH-0437 PT-2028); 1 female, upper Freshwater Creek,

Whitfield Range, Cairns, 24 August 1974, MV-light, MS Moulds

(WTH-0615); 1 male, same data (NMV WTH-0603); 1 male, same

data (NMV WTH-0604); 1 male, upper Freshwater Creek, Whitfield

Range, near Cairns, 15 December 1974, MS Moulds (NMV WTH-

0606); 1 male, 1 male, same data (NMV WTH-0605); 1 male, same

data (NMV WTH-0607); 2 females, same data (NMV WTH-06014); 1

male, the Crater, near Herberton, 18 December 1974, M.S. Moulds

(NMV WTH-0602); 1 female, Mareeba, Davies Creek Road, 21

January 1976, Waltord-Huggins (NMV WTH-0616); 1 male, State

Forest, 24 km along Goldsborough Road near Gordonvale, 27.x11.1980,

M.S. and B.J. Moulds (NMV WTH-1350); 1 male, Behana Gorge,

Cairns, 16 November 1982, T. Hinger, 17° 11' S 145° 50' E (NMV

WTH-0608); 2 females, same data (NMV WTH-0609); 1 male,

Mossman Gorge, Daintree National Park, 17 November 1988, MV It,

K. Walker (NMV WTH-0595); 1 male, river on Lake Morris Road,

Cairns, 3.v.2011, 16.9412 145.71762 (NMV EPAVTI22); 2 males, 5

females, river on Lake Morris Road, Cairns, 16.9412 145.71762,

3.v.2011, J. Mynott and M. Shackleton (NMV Tri-54557); 1 male,

small creek beside Josephine Falls, M. Shackleton and J. Mynott,

6.v.2011 (NMV Tri-54559); 1 male, 1 female, Josephine Falls, 17.4338

145.8630, Shackleton and Mynott 110506-7, 6.v.2011 (NMV Tri-

54558); 3 males, Dunn? Creek at Bridge 11, Kirrama Range Rd, ~14

km from national park sign, —18.2138 S 145.7982 E, 24.x.2017, D.

Cartwright and R. St Clair; 1 male, Mulgrave Road, Goldsborough

Valley Campsite, Wooroornoonan National Park, 17.2374 S

145.7733 E, 26.x.2017 (NMV JOS 451); 1 male, Python Creek in Tully

Gorge Road, ~52 km NW of Tully, 17.76628 145.5895E, 2 November

2017, D. Cartwright and R. St Clair (QM JOS-448).

Diagnosis. The male of Barynema dilatum closely resembles

that of B. dolabratum sp. nov. 1n having the inferior appendages

each with the harpago reduced to a small apically setose ventral

lobe and the coxopodite in lateral view produced and curved

ventrad distally. In B. dolabratum, the upper penis cover in

ventral view appears to form two lobes apically, with the

ventral lobe expanded and rounded laterally, in contrast with B.

dolabratum, which has the upper penis cover in ventral view

pick-shaped apically.

109

Description. Mesothorax with mesoscutellum shield-shaped,

without setate warts; fresh adult males with body and wings

dark except white area at anastomosis on wings.

Male. Length of each forewing 9.3—10.3 mm (n = 10).

sternite VII without median tab or spur. Genitalia: Pre-anal

appendages in dorsal view broadest at about midpoint,

constricted towards base. Upper penis cover distally bilobed,

dorsal lobe tapered apically in ventral view, rounded in

lateral view, ventral lobe expanded and rounded laterally

towards apex. Inferior appendages each with coxopodite base

short, stout, produced to form a dorsal lobe with slender neck

before slightly dilated apex bearing short stubby black setae,

harpago short, rod-shaped, bearing dense brush of short

black setae apically.

Female. Length of each forewing 11.8—17.7 mm (n = 10).

Terminalia: Apical lobes conical, tapered to darkly sclerotised

apices.

Distribution. From north-eastern Queensland, west of Tully to

the Cairns district.

Etymology. The name is derived from the Latin dilato, to

extend, for the lateral swellings on the upper penis cover lobes.

Barynema dolabratum sp. nov.

http://zoobank.org/urn:|sid:zoobank.org:act:/OCBASA9-B547-

4657-9FF7-F5D89A A BE20E

Figures 25, 26

Material examined. Holotype. male, north Queensland, Gap Creek,

10.11.1982, M.S. Moulds (NMV W'TH-0599, P'T-1176).

Paratypes. North Queensland: 1 male, Gap Creek, Mount

Finlayson Range, S of Cooktown, 25 November 1974, M.S. Moulds

(NMV WTH-00601); 1 male, Gap Creek, 10 February 1982, M.S.

Moulds (NMV WTH-0600); 1 male, Oliver Creek on Bloomfield

Road, 8 km S of Cape Tribulation, 16.1377 S 145.4408 E, 29.x.2017, D.

Cartwright and R. St Clair (QM); 1 male, data as above, D. Cartwright

and R. St Clair, NMV (JOS-452).

Diagnosis. This species, known only from males, closely

resembles that of B. dilatum, but 1s distinguished by the slightly

smaller body, darker body, mesoscutal setate warts elongate-

ovoid, and the sharp, pick-shaped form of the upper penis cover

lobes in ventral view.

Description. Mesothorax with mesoscutellum shield shaped,

without setate warts.

Male. Length of each forewing 9.0—10.6 mm (n = 5). Small

white area around lower part of wing at anastomosis. Sternite

VII without median spur or tab. Genitalia: Pre-anal

appendages stout, apically rounded in ventral view. Upper

penis cover lobes produced distally, pick-shaped apicolaterally

in ventral view, in lateral view triangular apico-dorsally.

Inferior appendages each with coxopodite broad at base, 1n

lateral view produced in slender, curved dorsal lobe bearing

cluster of stubby black setae apically; harpago forming short

apically truncate lobe bearing stout peg-like black setae at tip.

Female. Unknown.

Distribution. Collected from Cape York, Queensland, from the

two sites to the north of Cairns and south of Cooktown.

110

Etymology. From the Latin dolabratus, meaning pick-shaped

for the appearance of the upper penis cover lobes.

Barynema australicum Mosely

Barynema australicum Mosely, in Mosely and Kimmins, 1953:

162, figs 108, 109.

Figures 27—32

Material examined. Holotype. male, New South Wales, Ebor, 5.1.1916,

R.J. Tillyard, (BMNH) [head and abdomen mounted in balsam,

images courtesy B. Price, BMNH]

New South Wales: 1 female, Wilson River Reserve near Bellangry,

5.x11.1986, G. Theischinger (NMV TRI-26493); 1 male, Coomboodja

Creek, Washford National Park, 29° 16' S 152° 22' E, 5.1.1986,

Theischinger (NMV Tri-27119, P1-1655); 2 females, same data (NMV

Tri- 27116); 1 male ,1 female, “Cockerawo... Creek", 23 km WNN

Bellangry, 8.x11.1986, 730 m, G. Theischinger (NMV TRI-26505); 19

males 1 female, Orara West State Forest, Tuckers Knob, 29° 41' S

152° 48' E, 27.x1.1990, G. Theischinger (NMV TRI-27117); 1 male,

Manning River, Pheasant Creek Road, A. Glaister, J. Dean and R. St Clair,

3.x11.2007, 31° 53' S 151° 29' E (NMV TRI-54569, JOS-238); 1 male,

Never Never River, Whitneys Road, —30.33001S 152.86222E, 10.

x1.2010, 10110-5, MS 747 [M. Shackleton] (NMV TRI-54570, JOS- 108).

Diagnosis. Distinguished from most other species of Barynema

by having a pair of setate warts on the mesoscutellum, a feature

shared with B. goomburra sp. nov., from which it differs by

having male genitalia in ventral view with the coxopodite of

each inferior appendage subquadrate, and in both ventral and

lateral views the harpago with a short, slender basal portion

before the swollen terminal section that bears a broader

hairbrush-shaped area of short blunt black setae, rather than an

elongate toothbrush-shaped region as in B. goomburra sp. nov.

Description. Male. Length of each forewing 6.4—7.8 mm (n =

10). Sternite VII bearing apically rounded median tab.

Genitalia: see Mosely and Kimmins, 1953: 162).

Female. Length of each forewing 8.4—-10 mm (n = 3).

Terminalia: Apical lobes on abdominal segment X short,

broad, almost quadrate.

Distribution. Found in north-east New South Wales, in the

Barrington Tops-Wauchope and mid north coast regions.

Remarks. Two specimens among those that have been

sequenced for the BOLD project can be referred to B.

australicum. Each appears in a different cluster on the current

BOLD ‘Taxon Identification Tree. This is due to either

contamination of the COI gene or the presence of sibling

species. Several more specimens are required before this can

be resolved. This species, together with specimens here referred

to B. dilatum sp. nov., show no presently resolvable relationships.

Barynema goomburra sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:3BS54EDD-SAEB-

4E71-903C-DEEOAEEF26EE

Figures 33—35

Material examined. Holotype. male, Queensland, Goomburra State

Forest, NE of Warwick, 28° 03'S 152° 07' E, 20.1.1986, G. Theischinger

(NMV TRI- 27112).

A. Wells & R.M. St Clair

Paratypes. New South Wales: 1 male, 4 females, Styx River, Hyatt

Flat, 8.x11.1998, G. Theischinger (ANIC).

Diagnosis. Adults of this species closely resemble those of B.

australicum in having a pair of setose warts on the

mesoscutellum; the males of both species have a brush of short,

blunt black setae terminally on each harpago and, in lateral

view, the pre-anal appendages are broad-based and gradually

tapered distally. The male of B. goomburra 1s characterised by

having the inferior appendages narrower than those of B.

australicum and the coxopodites lacking apicomesal clusters of

setae that are present in B. australicum. The male of B.

goomburra resembles that of B. costatum with the harpago

bearing an elongate and narrow brush of stout black setae

occupying almost its entire inner distal margin, but differs

from B. costatum in that B. goomburra has a subcircular to

rectangular mesoscutellum bearing small paired setate warts

separate from each other and close to the posterior margin,

whereas the mesoscutellum of B. costatum is ovoid and with

large paired setate warts fused and covering most of the

mesoscutellum.

Description. Mesothorax with mesoscutellum bearing pair of

setate warts close to distal margin.

Male. Length of each forewing 6.5-7.2 mm (n = 2)

Abdominal segment VII bearing stoutly rounded midventral

tab. Genitalia: Pre-anal appendages in lateral view elongate

triangular; upper penis cover in ventral view forming pair of

short, stout, apically expanded structures; inferior appendages in

ventral view each with coxopodite about equal width throughout

length, without apicomesal brush of setae, harpago not clearly

delineated, with somewhat stout, short, blunt black setae lining

entire mesal side, giving toothbrush-like appearance.

Female. Length of each forewing 8.0—9.6 mm (n = 4).

Terminalia: Apical lobes broad-based, arising close to each

other, then tapered to narrowly rounded apices.

Distribution. Collected from just north of the New South

Wales-Queensland border, and from the Styx River in the

Northern Tablelands area of New South Wales.

Remarks. No specimens recognisable as B. goomburra are among

material for which molecular (COI) data are available at present.

Etymology. Named for the holotype locality.

Acknowledgements

We thank, particularly, staff of Museums Victoria Entomology

section for making both of us welcome, for making available

their extensive odontocerid collection and for providing

laboratory space to RSC. The Australian National Insect

Collection, CSIRO, Canberra, is thanked for providing

laboratory facilities for AW. Dr Ben Price, Natural History

Museum, London, kindly supplied images of the type of

Barynema australicum.

Material from Queensland collected by Michael

Shackleton was taken under collecting permits WITKIO27111

and that collected by David Cartwright and Ros St Clair under

the Entomological Society of Queensland collecting permits

WITK15549915 and TWB/02/2015. Material from New South

Australian Odontoceridae reviewed 111

Figures 27-35, Barynema species. 27, 28, B. australicum Mosely, holotype male (BMNH): 27, genitalia, lateral (on microscope slide); 28, part

of body and wings of holotype (images, courtesy B. Price, BMNH). 29-32, B. australicum, New South Wales, Orara West State Forest (NMV-

27117): 29, male mesothorax dorsal; 30, female, terminalia ventral; 31, 32, male, genitalia, lateral and ventral, respectively. 33—35, B. goomburra

sp. nov., holotype male, Queensland, Goomburra State Forest, NE of Warwick (NMV TRI-27112): 33, mesothorax dorsal; 34, 35, genitalia lateral

and ventral, respectively.

Abbreviations: har. = harpago; mes. lob. = mesal lobe of coxopodite; pre. app. = pre-anal appendages; set. wt. = setate warts on mesoscutellum;

up. p. = upper penis cover.

112

Wales was collected by Michael Shackleton under Scientific

Research Permit PO7/0095 and National Parks Service

scientific License 512404; by Alena Glaister, Ros St Clair,

and John Dean under Scientific Research Permit PO7/0095-

1.0, and National Parks Service Scientific License S12404;

and by Zac Billingham as bycatch under National Parks

Service Scientific License SL101930.

Material from Queensland was collected under Scientific

Research Permit numbers WITK06190909 (May 2010)

and WITK10277111 (8 Nov 2011—7 Nov 2013). Material

from Victoria was collected under Scientific Research

Permits 10005961 (19 Aug 2012-31 Aug 2013).

Material from Queensland was collected under Scientific

Research Permit numbers WITK006190909 (May 2010)

and WITKIO277111 (8 Nov 2011—7 Nov 2013). Material

from Victoria was collected under Scientific Research

Permits 10005961 (19 Aug 2012-31 Aug 2013).

Material from Queensland was collected under Scientific

Research Permit numbers WITK06190909 (May 2010)

and WITKIO277111 (8 Nov 2011—7 Nov 2013). Material

from Victoria was collected under Scientific Research

Permits 10005961 (19 Aug 2012-31 Aug 2013).

References

Banks, N. 1939. Trichoptera. Bulletin of the Museum of Comparative

Zoology, Harvard 85: 439—504.

Betten, C. 1934. The caddisflies or Trichoptera of New York State. New

York State Museum Bulletin No. 292. The University of the State

of New York: Albany. 576 pp. doi.org/10.5962/bhl.title.132984

A. Wells & R.M. St Clair

Cartwright, D.C., and Dean, J.C. 1987. Descriptions of the immature

stages of Barynema costatum Banks from Australia (Trichoptera:

Odontoceridae). Aquatic Insects 9: 27-32. dol.

org/10.1080/01650428709361267

Dean, J.C., St Clair, R.M., and Cartwright, D.I. 2004. /dentification

keys to Australian families and genera of caddis-fly larvae

(Trichoptera). Identification and ecology guide. No 50.

Cooperative Research Centre for Freshwater Ecology: Albury,

NSW. 131 pp.

Drecktrah, H.G. 1990. Larval and pupal descriptions of Marilia fusca

(Trichoptera: Odontoceridae). Entomological News 101: 1-108.

doi.org/10.11646/zootaxa.1362.1.3

Mosely, M.E., and Kimmins, D.E. 1953. The Trichoptera (Caddis-flies)

of Australia and New Zealand. British Museum (Natural History):

London. 550 pp. doi.org/10.5962/bhl.title.118696

Müller, F. 1880[“1878”]. Sobre as casas construidas pelas larvas de

insectos Trichopteros da Provincia de Santa Catharina. Archivos

do Museu Nacional, Rio de Janeiro 3(1878): 99-134, 209—214.

Neboiss, A. 1977. A taxonomic and zoogeographic study of Tasmanian

caddis—flies (Insecta: Trichoptera). Memoirs of Museum Victoria

38: 1-208. doi.org/10.24199/1.mmv.1977.38.01

Neboiss, A. 2003 [72002"]|. New genera and species, and new records,

of Tasmanian Trichoptera (Insecta). Papers and Proceedings of

the Royal Society of Tasmania 136: 43—82. doi.org/10.26749/

rstpp.136.43

Olah, J., and Johanson, K.A. 2010. Description of 33 new species of

Calamoceratidae, Molannidae, Odontoceridae and

Philorheithridae (Trichoptera), with detailed presentation of their

cephalic setal warts and grooves. Zootaxa 2457: 1-128. doi.

org/10.11646/zootaxa.2457.1.1

Yang, L.-F., Yuan, H.-Y., and Morse, J.C. 2017. Lannapsyche and

Marilia species of China (Trichoptera: Odontoceridae). Zootaxa

4320(1): 81-99. doi.org/10.11646/zootaxa.4320.1.5

Memoirs of Museum Victoria 80: 113-152 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.06

New species of squat lobsters of the genus Munida from Australia

(http://zoobank.org/urn:|sid:zoobank.org:pub:EA21667A-77A5-411D-9C1A-23ECFFF3D505)

ANNA W. McCCALLUM!, SHANE T. AHYONG? AND NIKOS ANDREAKIS>

! Museums Victoria, GPO Box 666, Melbourne VIC 3001, Australia. Email: amccallum(?museum vic.gov.au

* Australian Museum Research Institute, 1 William St, Sydney NSW 2010, Australia, and School of Biological, Earth

and Environmental Sciences, University of New South Wales, Kensington NSW 2052, Australia.

Email: shane.ahyong@austmus.gov.au

* College of Science and Engineering, James Cook University, Townsville QLD 4814, Australia.

Abstract

McCallum, A.W., Ahyong, S.T. and Andreakis, N. 2021. New species of squat lobsters of the genus Munida from

Australia. Memoirs of Museum Victoria 80: 113—152.

This study reports on new squat lobsters of the genus Munida collected during recent surveys of

Australia's continental margins. We report on 33 species of Munida including seven new species and 14

new range extensions for Australia. More than 500 specimens were collected, mostly from the western

continental margin of Australia, but also including a new species from deep water (22000 m) off Tasmania.

We provide new data on the colour patterns of some species and include molecular data from two

mitochondrial markers (16S rRNA and COI) to support the taxonomic status of the new species.

Introduction

Australia has a vast marine realm, the third largest jurisdiction

in the world, encompassing tropical and temperate biomes and

connected to the highly biodiverse Indo-West Pacific region.

Over the last decade, efforts to document the biodiversity of

Australia's deep seafloor habitats have grown, as a system of

Marine National Parks were established and monitored.

sampling of Australia’s deep seafloor has revealed a high

proportion of new species and significant range extensions, but

most of these records await detailed taxonomic description.

Squat lobsters (Chirostyloidea and Galatheoidea) are

among the most diverse groups of decapod crustaceans in

deep waters, and there have been intensive efforts to describe

and revise species and higher taxa from the group

(Macpherson and Baba, 2011). This has been assisted by the

availability of taxonomic resources such as synoptic works

(e.g. Baba, 2005; Baba et al., 2008) and keys (Taylor and

Poore, 2010). In addition, molecular-based approaches are

helping to identify overlooked species that show only subtle

morphological differences (Poore and Andreakis, 2012).

Munida is one of the largest genera of squat lobsters,

currently comprising 297 species (WoRMS, 2021). The most

recent major taxonomic study of Munida in Australia

recorded 19 species (Ahyong and Poore, 2004). Additional

records have since been added (e.g. Ahyong, 2007; Baba,

2005; Poore et al., 2008); so, to date, 32 species of Munida

have been documented from Australia.

Here, we report the specimens of Munida collected during

several deep-water surveys from 2005 to 2015, including

surveys of the continental margin of Western Australia, the

Great Australian Bight, Tasmanian seamounts, and the eastern

Australian continental margins and abyss. We describe and

illustrate new species and record new occurrences of

previously described taxa. The synonymies for each species

are restricted to papers published after the compilation of

Baba et al. (2008). Where available, we include molecular

data from two mitochondrial markers (168 and COI) to

support the taxonomic status of the species.

Materials and methods

Sampling

Specimens were collected using beam trawls and epibenthic

sleds ın surveys aboard the FRV Southern Surveyor of the

south-western continental margin of Australia (SS10/2005) and

north-western continental margin (8505/2007); and surveys

aboard the RV Investigator of the Great Australian Bight

(IN2015 COI. 042) and the continental margin and abyss of

eastern Australia (IN2017 V03). A small number of specimens

were collected from south-west Tasmania (TT 01/2008) using

the remotely operated vehicle Jason aboard the RV Thomas G

Thompson. Specimens collected from well-sampled sites on

Tasmanian seamounts (SSO1/2008, SSO2/2007) were reviewed

and represent well-known species previously reported for the

area; they are not redescribed herein.

114

Morphological methods

Specimen measurements in Material examined refer to

postorbital carapace length (pcl), measured dorsally from the

base of the rostrum (situatued at the level of the orbit) to the mid-

posterior margin of the carapace, and where the rostrum 1s intact,

carapace length (cl) is measured from the rostral apex to the mid-

posterior margin of the carapace. The angle of the frontal-margin-

from-the-midline is measured as the angle of the intersection

between an imaginary line along the frontal margin and the

median axis. Thus, a transverse margin corresponds to an angle

of 90°. The lengths of articles of the cheliped are measured along

the dorsal margin and those of ambulatory legs and maxilliped 3

are measured along the extensor margin and exclude spines.

Abbreviations used are Mxp3, maxilliped 3; P1, pereopod 1; P2—

4, pereopods 2-4; M, male; F, female; ovig., ovigerous.

Specimens are deposited in the collections of Museums

Victoria, Melbourne (NMV), the Western Australian Museum

(WAM), the Australian Museum, Sydney (AM), Muséum

national d'Histoire naturelle, Paris (MNHN) and the South

Australian Museum, Adelaide (SAM). Other abbreviations: WA,

Western Australia; acq. refers to the Commonwealth Scientific

and Industrial Research Organisation (CSIRO) acquisition

number allocated to the specimen on board the vessel at the

time of collection.

Molecular analysis

Material collected from south-western Australia (SS10/2005),

being mostly preserved 1n formalin, was not sequenced.

Results

A.W. McCallum, S.T. Ahyong & N. Andreakis

Total DNA was extracted from 50-100 mg of abdominal

tissue or pereopod of the target specimen preserved in ethanol

following the salt-based extraction procedure described by

Aljanabi and Martinez (1997) with minor modifications. The

partial mitochondrial cytochrome oxidase subunit I (COI) and

165 rDNA gene (165 fragment 2) were polymerase chain reaction

(PCR) amplified following primer combinations, reaction

conditions and thermal cycling profiles established previously

(Cabezas et al., 2012). Quantity and length of the PCR products

were examined by 1% gel electrophoresis against known DNA

standards. PCR reactions were sent to Macrogen Inc. (Korea,

www.macrogen.com) for purification and direct sequencing in

both directions. | Electropherograms were assembled in

sequencher v4.9 and sequences were initially aligned in Bioedit

v7.0.9 (Hall, 1999) together with 16S and COI sequences from

previous works (Cabezas et al., 2009, 2011; Machordom and

Macpherson, 2004; Macpherson and Machordom, 2005;

Rodriguez-Flores et al., 2019). Final alignments of each of the

loci were performed using MAFFT (Katoh et al., 2002).

Uncorrected divergences (p) were calculated using MEGA 10.04

(Kumar et al., 2018). GenBank accession numbers for the new

sequences are given in Table |. We have not provided the

divergence values among all pairs of closely related species

because in the genus Munida the amount of molecular data 1s still

scarce (excluding the present paper, the sequences of only 72

species are available). Nevertheless, when the molecular data

from a new species and its morphological closest relative are

available, we have included the divergence values in the Remarks.

Table 1. Specimens studied for mitochondrial DNA sequences (COI and 165), including registration codes, survey, station, and GenBank accession numbers

Species Reg. no. 16S COI Surve

Munida agave NMV J56099 MK847949 MK848000 S5505/2007 161

Munida agave NMV J56008 MK847961 MK84s8011 5305/2007 116

Munida asprosoma NMV J56393 MK847955 MK848006 S5505/2007 155

Munida asprosoma NMV J56016 MK847971 MK848022 5505/2007 68

Munida asprosoma NMV J57258 MK847950 MK848001 S5505/2007 155

Munida babai NMV J55042 na MK848021 SS 10/2005 170

Munida distiza NMV J56487 MK847947 MK847998 5505/2007 11

Munida leeuwin sp. nov. NMV J55046 MK847968 MK848018 SS 10/2005 170

Munida leeuwin sp. nov. NMV J56100 MK847954 MK848005 5305/2007 82

Munida leeuwin sp. nov. NMV J55046 MK847967 MK848017 SS 10/2005 170

Munida gracilis NMV J55107 MK847963 MK$848013 SS 10/2005 32

Munida gracilis NMV J55107 MK847964 MK848014 SS 10/2005 32

Munida gracilis SAM C7658 MK847932 na BPZ 2010 500

Munida compressa NMV J55980 MK847946 na 5505/2007 176

Munida compressa NMV J5598 1 MK847944 MK847996 S5505/2007 189

Munida endeavourae NMV J59312 MK847934 MK847986 1101/2008 J2-390-015

Munida endeavourae NMV J59310 MK847933 na 1101/2008 J2-389-006

Munida gordoae NMV J56420 MK847957 MK848008 3305/2007 62

Munida gordoae NMV J56420 MK847956 MK848007 5505/2007 62

Munida abrolhos sp. nov. NMV J55279 MK847962 MK848012 3305/2007 70

Munida abrolhos sp. nov. NMV J55262 MK847943 MK847995 5305/2007 1

Munida leagora NMV J56018 MK847951 MK848002 SS05/2007 77

Munida leagora NMV J56004 MK847970 MK848020 S5505/2007 143

Munida leagora NMV J56004 MK847969 MK848019 S5505/2007 143

Munida leagora NMV J55999 MK847942 MK847994 5505/2007 143

Munida pectinata NMV J55268 MK847939 MK847991 S5505/2007 62

Munida philippinensis NMV J56003 MK847953 MK848004 5505/2007 143

Munida philippinensis NMV J56003 MK847952 MK848003 SS05/2007 143

Munida compacta NMV J56411 MK84794]1 MK847993 SS05/2007 57

Munida compacta NMV J56402 MK847937 MK847989 SS05/2007 15

Munida compacta NMV J56412 MK847938 MK847990 SS05/2007 56

Munida roshanei NMV J5599] MK847958 MK848009 S5505/2007 97

Munida roshanei NMV J56009 MK847960 na 5505/2007 125

Munida roshanei NMV J56400 MK847940 MK847992 5305/2007 116

Munida julumunyju sp. nov. NMV J55064 MK847966 MK848016 SS 10/2005 152

Munida julumunyju sp. nov. NMV J55979 MK847945 MK847997 S5505/2007 34

Munida heteracantha NMV J55038 MK847965 MK848015 SS 10/2005 146

Munida galalala sp. nov. WAM C78558 MK847959 MK848010 SS05/2007 156

Munida jurunjurun sp. nov. WAM C78561 MK847948 MK847999 SS05/2007 63

Munida lutruwita sp. nov. NMV J67480 MK847935 MK847987 1101/2008 J2-387-022

Munida lutruwita sp. nov. NMV J67474 MK847936 MK847988 1101/2008 J2-387-023

Australian Munida

Systematics

Munididae Ahyong, Baba, Macpherson and Poore, 2010

Munida abrolhos sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:14DA5265-EOES5-

47B5-B4B0-93296A355921

Figures 1, 2, 3A

Munida sp. MoV 5214. — Poore et al., 2008: 21 (south-western

Australia, 201—206 m).

Munida aft. rubiesi. — Poore et al., 2008: 20 (part), unnumbered

fig. — McEnnulty et al., 2011: app. 1.

Munida sp. MoV 5526. — Poore et al., 2008: 21.

Munida keiensis. — McEnnulty et al., 2011: app. 1.

Type material. Holotype: WAM C78556, ovigerous female (cl 23 mm, pcl

14 mm), Abrolhos, Western Australia, 29° 00.594' S, 113° 42.78' E to

29° 01.512' S, 113°43.32' E, 700—704 m, SS10/2005/85, 2 December 2005.

Paratypes (all Western Australia): NMV J55036, 6 ovigerous

females (cl 18.9 mm, pcl 11.2 mm to cl 25.9 mm, pcl 15.7 mm), 6

females (cl 14.0 mm, pcl 8.1 mm to cl 19.0 mm, pcl 12.5 mm), 10 males

(cl 15.0 mm, pcl 8.9 mm to cl 22.8 mm, pcl 14.6 mmy), collected with

holotype; NMV J55262, 1 ovigerous female (cl 27.5 mm, pcl 18.3 mm),

| female (cl 14.5 mm, pcl 8.4 mm), Barrow L1 transect, 20° 59.412' S,

114° 07.896' E to 20° 59.13' S, 114? 08.394' E, 700 m, S805/2007/1, 9

June 2007; NMV J55279, 1 specimen, Mermaid L24 transect,

16° 44.286' S, 119° 15.042' E to 16? 43.794' S, 119° 15.48' E, 693—698

m, $5505/2007/70, 17 June 2007; WAM C78557, 1 male (cl 22.6 mm, pcl

13.0 mm), Ningaloo South, 22° 03.57' S, 113° 43.74' E to 22° 04.026' S,

113° 43.26' E, 658—754 m, SS10/2005/149, 10 December 2005.

Description. Carapace. Length and width subequal, widest at

midlength. Dorsal surface with main transverse ridges mostly

uninterrupted, without secondary transverse striae between

main ridges; ridges and striae lined with short, non-iridescent

setae. Gastric region with large pair of epigastric spines behind

supraocular spines and 2 or 3 pairs of small epigastric spines;

without median row of spines behind rostrum. Hepatic region

eranular; parahepatic spine present. Anterior part of branchial

region between cervical groove and postcervical groove with |

or 2 ridges and sometimes lor 2 short striae; postcervical spine

present; posterior part of branchial region with 5 transverse

ridges (excluding posterodorsal ridge). Cardiac region with 3

main transverse ridges. Intestinal region with 2 lateral striae;

posterodorsal ridge distinct, without secondary stria. Frontal

margin inclined posteriorly at 110° from midline. Lateral

margin slightly convex; anterolateral spines parallel, directed

slightly upwards, reaching sinus between rostrum and

supraocular spine; hepatic marginal spine anterior to cervical

eroove, much shorter than anterolateral spine; branchial margin

with 4, occasionally 5, spines (additional third spine small).

Rostrum spiniform; curving dorsally, 0.6—0.7 x pcl; supraocular

spine length 0.3 length x rostrum length. Epistomial ridge

straight, ending at antennal gland; mesial protuberance distinct.

Thoracic sternum. Median length of sternal plastron

(sternites 4—7) 0.6 x width of sternite 7. Thoracic sternites

suface smooth, sternites 4 with few striae. Sternite 3 0.5 x

width of sternite 4. Sternite 4 anterior margin subtriangular,

narrowly contiguous with sternite 3.

115

Abdomen. Somite 2 with 8—10 large spines, evenly spaced

along anterior ridge; with stria behind anterior ridge. Somites

3-4 with 2 striae behind anterior ridge. Somite 6

posteromedian margin slightly concave. Telson with numerous

transverse squamae; greatest width 1.6 x midlength;

anterolateral margin concave.

Eye. Maximum corneal diameter 0.35 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded) not

overreaching corneae; distolateral and distomesial spines

subequal; 2 lateral spines, proximal smaller, longer lateral

spine reaching end of distal spines.

Antenna. Article | with distomesial spine reaching distal

margin of article 2. Article 2 distomesial spine nearly

reaching distal margin of article 4; distolateral spine reaching

midlength of article 3. Articles 3 and 4 unarmed.

Maxilliped 3. Ischium 1.8 x length of merus, with small

flexor distal spine. Merus flexor margin with large spine and

small distal spine; extensor margin without distal spine.

P1. 2.0-2.5 pcl, with plumose setae on inner margins, and

some iridescent setae, merus 0.8—0.9 pcl, with row of 6 spines

on dorsal margin; distomesial spine reaching midlength of

carpus. Carpus 0.4 x length of merus, length 0.9—1.1 x width,

with 5 spines along the mesial margin. Propodus 1.0—1.4 x

merus length, fingers longer than palm, 0.5-0.6 x total

propodus length; pollex with subdistal spine, outer margin

unarmed; dactylus outer margin unarmed.

P2-4. Long, slender, with few small scales on lateral

surfaces of meri, carpi and propodi; extensor margin with

plumose setae and few iridescent setae. P2 and P3 similar in

length, P4 shorter (P3 merus 0.9—1.0 x length of P2 merus, P4

merus 0.8 x length of P3 merus). P2 1.92.1 x pcl; merus

0.6—0.8 x pcl, length 6 x width, 3—4 x carpus length and 1.6 x

propodus length, extensor margin with 9 spines including

distal spine, flexor margin with 4 spines and well-developed

distal spine; carpus with 2 extensor spines a distal flexor

spine; propodus length 5.5—6.5 x height, with 5 movable

flexor spines; dactylus 0.8—0.9 x propodus length, gently

curved, length 6—7 x width, extensor margin densely lined

with stiff short setae on distal half, flexor margin with 8 small

moveable spines, space between spines increasing distally,

unarmed along distal 1/5. End of P2 carpus overreaching end

of PI merus. P3 with similar spination and article proportions

as P2; merus, propodus and dactylus as long as those of P2.

PA Jength 0.85 x P2 length; merus 0.5—0.6 x pcl; merocarpal

articulation reaching hepatic marginal spine of carapace.

Egg diameter. 0.5 mm.

Colour. Carapace and abdominal somites 2—3 pink. Rostrum

and supraocular spines white. Pl pale pink with white fingers,

P2—4 pale pink on meri and carpi, white on propodi and dactyli.

Genetic data. COI and 16S; see Table 1.

Etymology. After the type locality near the Abrolhos Islands;

used as a noun in apposition.

Remarks. Munida abrolhos sp. nov. 1s similar to M. andamanica

Alcock, 1894, M. rosula Macpherson, 1994, and M. curvirostris

Henderson, 1885, with a dorsally curving spiniform rostrum, P1

116 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figue 1. Munida abrolhos sp. nov., holotype, ovigerous female cl 23 mm, pcl 14 mm (WAM C78556): A, carapace and abdomen, dorsal view; B,

rostrum, lateral view; C, right chela, dorsal view; D, left antenna and antennule, ventral view; E, sternum; F, abdominal somite 6, telson and right

uropod. G, ovigerous female cl 22.9 mm, pcl 14.4 mm (NMV J55036), anterolateral margin of carapace. Scale: A-C, F = 5 mm, D, E, G = 2 mm.

Australian Munida 117

Figure 2. Munida abrolhos sp. nov., holotype, ovigerous female cl 23 mm, pcl 14 mm (WAM C78556): A, right P2 dactylus, lateral view; B, right

P2, lateral view; C, right P3, lateral view; D, right P4 lateral view; E, left maxilliped 3, lateral view. Scale A, E = 2 mm, B-D = 5 mm

118 A.W. McCallum, S.T. Ahyong & N. Andreakis

EA cA

t :

— o aes tars

Figure 3. A, Munida abrolhos sp. nov., female paratype, cl 27.5 mm, pcl 18.3 mm (NMV 755262); B-D, Munida julumunyju sp. nov., undetermined

paratype from NMV J55979, juvenile paratype, cl 9.2 mm, pcl 10.8 mm (NMV J55061), juvenile paratype cl 10.9 mm, pcl 6.6 mm (NMV

J55062); E, Munida lutruwita sp. nov. ovigerous female, cl 14.3 mm, pcl 10.6 mm, (AM P103229). Photos: K. Gowlett-Holmes, CSIRO.

Australian Munida

fixed finger with only subterminal spine (without marginal spines

on lateral margin), and abdominal somite 2 with a row of 6—8

spines and at most 4 transverse ridges. Most clearly the new

species differs from these species as it usually has only 4 branchial

spines, although sometimes a smaller third branchial spine is

present. In addition, postcervical spines, present in M. abrolhos

sp. nov., are always absent in the aforementioned species.

Of those species with 4 branchial spines, M. abrolhos sp.

nov. is most similar to M. keiensis Baba, 2005, in having

subequal distal spines of the basal antennular article, and the

distomesial spine of antennal article 2 overreaching article 3

but not distinctly overreaching article 4. The two species can

be distinguished by the shape of the anterior margin of

thoracic sternite 4, which is broadly rounded in M. keiensis

but subtriangular in M. sp. nov.

Sequence divergence between M. abrolhos sp. nov. and M.

rosula is 3% for COI and 0.8% for 168.

Distribution. Presently known only from off Western Australia,

658—754 m.

Munida galalala sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:83168BC4-60EA-

4710-A32B-42A AFFF985CA

Figures 4, 5

Type material. Holotype. WAM C78558, male (cl 17.8 mm, pcl

11.5 mm), Western Australia, Leveque L27 transect, 14^ 33.432' S,

121° 20.388' E to 14° 2.772 S, 121° 19.644' E, 924-1101 m,

$S05/2007/156, 3 July 2007 to 4 July 2007.

Description. Carapace. Length 1.1 x greatest width, widest at

midlength. Dorsal surface with main transverse ridges mostly

uninterrupted and some short secondary transverse striae

between main ridges on anterior half; ridges and striae lined

with short, non-iridescent setae. Gastric region slighly elevated,

with 2 pairs of epigastric spines, longest pair behind supraocular

spines. Hepatic region granular; parahepatic spine absent.

Anterior part of branchial region between cervical groove and

postcervical groove with few short scale-like ridges and

tubercles; postcervical spine absent; posterior part of branchial

region with 5 main transverse ridges (excluding posterodorsal

ridge) and few short secondary striae laterally between main

ridges. Cardiac region with 4 main transverse ridges. Intestinal

region with short median stria; posterodorsal ridge distinct,

slightly convex medially, without secondary stria. Frontal

margin inclined posteriorly at 110° from midline. Lateral

margins slightly convex; anterolateral spines parallel,

horizontal, reaching sinus between rostrum and supraocular

spine; marginal hepatic spine anterior to cervical groove also

strong, although distinctly shorter than anterolateral spine;

branchial margin with 5 spines. Rostrum spiniform, gentle

curving upwards, length 0.7 pcl; supraocular spine 0.45 x

length of rostrum. Epistomial ridge curved ending anterior to

antennal gland; mesial protuberance distinct.

Thoracic sternum. Surface smooth, without striae. Sternite

3 0.4 width of sternite 4; median length of sternal plastron

(sternites 4—7) 0.6 width of sternite 7. Sternite 4 anterior

margin subtrapezoid, broadly contiguous with sternite 3.

119

Abdomen. Somite 2 with 7 prominent spines along

anterior transverse ridge. Somites 2-4 each with 1

uninterrupted stria behind anterior ridge, without striae at

lateral margins. Somite 6 posteromedian margin almost

straight. Telson with numerous transverse squamae; greatest

width 1.5 x median length; anterolateral margin concave.

Eye. Maximum corneal diameter 0.15 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded)

overreaching corneae; well-developed distolateral spine much

longer than short distomesial spine; 2 lateral spines, proximal

smaller, not exceeding distal spines.

Antenna. Article | with strong distomesial spine not

reaching distal margin of article 2. Article 2 distomesial spine

not overreaching distal margin of article 3; distolateral spine

reaching distal margin of article 3. Article 3 distomesial spine

almost reaching distal margin of article 4. Article 4 unarmed.

Maxilliped 3. Ischium 1.3 x merus length, with small

flexor distal spine. Merus with 2 large spines on flexor margin

and | small spine; extensor margin unarmed.

P1. 3.0 x pcl, covered in plumose setae, without iridescent

setae, merus 1.2 x pcl, with row of 6 dorsal spines and 2 rows of

spines on mesial margin; distal spines strong, distomesial spine

not reaching midlength of carpus. Carpus 0.6 x merus length,

length 2.6 x width, with spines along mesial and dorsal margins.

Propodus 1.2 x merus length, fingers slightly shorter than palm,

0.5 x total propodus length; pollex with small subdistal spine,

margin unarmed; dactylusouter margin unarmed.

P2—4. Moderately long, slender, with numerous scales on

lateral surfaces of meri; extensor margin with row of plumose

setae. P2 length 2.1 x pcl; merus as long as pcl, length 7x

height, 3.5 x carpus length and 1.4 x propodus length, with

well-developed spines on extensor margin, increasing in size

distally, flexor margin with row of small spines and well-

developed distal spine; carpus with large spine and distal spine

on extensor margin, distal spine on flexor margin; propodus

length 6.5 x height, with 5 small movable flexor spines;

dactylus compressed, slightly curved, 0.6 x propodus length,

length 5.5 x width, with 9 movable spines evenly spaced along

flexor margin including spine at base of unguis. End of P2

carpus reaching end of P1 merus. P3 missing. P4 length 0.8 x

P2 length; merus 0.7 x pcl; merocarpal articulation almost

reaching hepatic marginal spine of carapace.

Genetic data. COI and 16S; see Table 1.

Etymology. Named galalala, meaning lobster in Dambimangari

language, used as a noun in apposition.

Remarks. Munida galalala sp. nov. belongs to the group of

species having five spines on the lateral branchial margins of

the carapace, smooth thoracic sternites, small eyes, and spines

on the anterior ridge of abdominal somite 2. The antennal

peduncle is most similar to Munida parvioculata Baba, 1982,

both species having a distinct distomesial spine on antennal

peduncle article 3. The new species differs in:

- the generally smooth surface of the carapace with widely

spaced transverse striae, compared to rugose with interrupted

transverse ridges in M. parvioculata

120 A.W. McCallum, S.T. Ahyong & N. Andreakis

B j C Re

eomm

2.232?) )

Figure 4. Munida galalala sp. nov., holotype, male 17.8 mm (WAM C78558): A, carapace and abdomen, dorsal view; B, right chela, dorsal view;

C, left antenna and antennule, ventral view; D, abdominal somite 6, telson and right uropod. E, sternum. Scale: A-E = 2 mm.

Australian Munida

121

Figure 5. Munida galalala sp. nov., holotype, male 17.8 mm (WAM C78558): A, left P2 dactylus, lateral view; B, left P2, lateral view; C, left P4,

lateral view; D left maxilliped 3, lateral view. Scale A = 1 mm, B-D = 2 mm

- the unarmed lateral margin of the cheliped pollex (with 2

spines in M. parvioculata)

- arow of 7 spines on abdominal somite 2 versus a single

pair in M. parvioculata.

Munida galalala can be distinguished from all other

species in this group (1.e. M. clevai Macpherson, 2009, M.

ampliantennulata Komai, 2011, M. profunda Macpherson and

de Saint Laurent, 1991, M. endeavourae Ahyong and Poore,

2004, M. pollioculus Komai and Higashiji, 2016, and M. typhle

Macpherson, 1994) by the presence of a spine on antennal

peduncle article 3.

Distribution. Presently known only from off north-western

Australia, 924—1101 m.

Munida julumunyju sp. nov.

http://zoobank.org/urn:lIsid:zoobank.org:act: A4C FA E49-1832-

4C56-BDED-A68F843BB9F4

Figures 3B—D, 6, 7

Munida rubridigitalis. — Poore et al., 2008: 20, unnumbered fig.

(upper right) — McEnnulty et al., 2011: app. 1,2.

Type material. Holotype: WAM C78559, female cl 22.4 mm, pcl 15.8

mm), Western Australia, Onslow L19 transect, 20° 07.962' S, 114? 587]' E

to 20? 07.584' S, 114? 58.416' E, 415—470 m, SS05/2007/15, 11 June 2007.

Paratypes (all Western Australia): NMV J56082, 1 female (cl 24.4

mm, pcl 16.3 mmy), collected with holotype; NMV J55979, ] male (cl

22.0 mm, pcl 15.0 mm), | ovigerous female (cl 18.8 mm, pcl 13.3 mm),

Dampier L20 transect, 19° 43.776' S, 115° 21.216' E to 19° 43.56' S,

115? 20.598' E, 389—423 m, SS05/2007/34, 12 June 2007; NMV

J55061, 1 juvenile (cl 9.2 mm, pcl 10.8 mm), Carnarvon, 24^ 33.12' S,

112? 15.12? E to 24? 33.804' S, 112? 15.18' E, 396—404 m,

5510/2005/122, 7 December 2005; NMV J55062, 1 juvenile (cl 10.9

mm, pcl 6.6 mm), Red Bluff, 23° 59.196' S, 112° 32.04' E to

23? 597724' S, 112? 31.74' E, 411 m, SS10/2005/130, 8 December 2005;

WAM C78560, 2 males (cl 22.6 mm, pcl 14.8 mm; cl 27.0 mm, pel 19.2

mm), 1 female (cl 22.6 mm, pcl 16.0 mm), Point Cloates, 22° 50.808' S,

113? 20.28' Eto 22° 51.228' S, 113? 19.98' E,420—430 m, SS10/2005/137,

9 December 2005; NMV J55060, 1 male (cl 24.2 mm, pcl 16.7 mm),

Ningaloo South, 22° 04.374' S, 113° 45.36' E to 22° 04.968' S,

113? 45.36' E, 391—396 m, SS10/2005/148, 10 December 2005; NMV

J55064, 1 male (broken rostrum; pcl 10.8 mm), Ningaloo South,

22? 04.314' S, 113? 45.36' E to 22? 04.854' S, 113? 45.36' E, 387—399 m,

$S10/2005/151, 10 December 2005; NMV J55059, 1 female (cl 23.2

mm, pcl 15.9 mm), Ningaloo North, 21° 58.212' S, 113° 47.46' E to

21° 58.806' S, 113° 47.1' E, 373—382 m, SS10/2005/165, 12 December

2005; NMV J55058, 2 males (cl 17.0 mm, pcl 11.1 mm; cl 25.8 mm, pcl

18.4 mm), Barrow Island, 21° 00.402' S, 114° 22.86' E to 21° 00.042' S,

114? 22.5' E, 399—408 m, SS10/2005/172, 13 December 2005.

Other material examined. New Caledonia: MNHN-IU-2014-15475, 2

females (cl 14.6 mm, pcl 9.1 mm; cl 18.5, pcl 11.5), 3 males (cl 14.3—

16.0 mm, pcl 8.5—10.1 mm), east coast, 21° 14.84' S 165° 55.49' E, 450—

122 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 6. Munida julumunyju sp. nov., holotype, ovigerous female cl 22.4 mm, pcl 15.8 mm (WAM C78559): A, carapace and abdomen, dorsal

view; B, right chela, dorsal view; C, right antenna and antennule, ventral view; D, sternum; E, telson; F, rostrum, lateral view. Munida julumunyju,

New Caledonia, male cl. 18.2, pcl 12.2 (MNHN-IU-2014-15475): G, rostrum, lateral view. A-D, G = 5 mm, E-F = 2 mm.

Australian Munida 123

Figure 7. Munida julumunyju sp. nov., holotype, ovigerous female cl 22.4 mm, pcl 15.8 mm (WAM C78559). A, right P2 dactylus, lateral view; B,

right P2, lateral view; C, right P3, lateral view; D, right P4, lateral view; E, left maxilliped 3, lateral view. Scale A-E = 2 mm.

124

490 m, HALIPRO 1 CP869, 23, March 1994; MNHN-IU-2014-15474,

| female (cl 15.5 mm, pcl 10.3 mm), Norfolk Ridge, 22° 10' S

167° 15.2' E, 495-515 m, BIOCAL CP109, 9 September 1985.

Description. Carapace. Length 0.9 x greatest width, widest at

midlength. Dorsal surface with main transverse ridges mostly

uninterrupted, with secondary transverse striae between main

ridges; ridges and striae lined with short, non-iridescent setae.

Gastric region with 3—6 pairs of epigastric spines, longest pair

behind supraocular spines. Hepatic region with short striae; small

parahepatic spine sometimes present. Anterior part of branchial

region between cervical groove and postcervical groove with

about 6 ridges and some shorter striae; posterior part of branchial

region with about 12 transverse ridges (excluding posterodorsal

ridge). Cardiac region with 3 or 4 main transverse ridges. Intestinal

region with 2 or 3 striae; posterodorsal ridge distinct, with

secondary stria. Frontal margin inclined posteriorly at 105° from

midline. Lateral margin slightly convex; anterolateral spines

parallel, horizontal, just reaching sinus between rostrum and

supraocular spine; marginal hepatic spine anterior to cervical

eroove, much shorter than anterolateral spine; branchial margins

with 5 spines. Rostrum laterally compressed; dorsal margin

straight, inclined dorsally, ventral margin strongly convex, deepest

slightly anterior to midlength, length 0.4—0.5 x pcl; supraocular

spine 0.4 x length of rostrum. Epistomial ridge straight, ending at

antennal gland; mesial protuberance distinct.

Thoracic sternum. Median length of sternal plastron

(sternites 4—7) 0.5 x width of sternite 7. Sternites 4—5 with

Striae; sternite 6 with few striae. Sternite 3 0.4 width of

sternite 4. Sternite 4 anterior margin subtrapezoid, narrowly

contiguous with sternite 3.

Abdomen. Somite 2 with 8 small spines evenly spaced along

anterior ridge. Somites 2—4 with about 5, 7 and 12 uninterrupted

striae behind anterior ridge, respectively, and squamae at lateral

margins. Somite 6 posteromedian margin almost straight.

Telson with numerous transverse squamae; greatest width 1.7 x

median length; anterolateral margin concave.

Eye. Maximum corneal diameter 0.3 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded) not

overreaching corneae; distolateral and distomesial spines

subequal; 2 lateral spines, proximal smaller, longer lateral

Spine exceeding distal spines.

Antenna. Article | with distomesial spine reaching

midlength of article 2. Article 2 with distomesial spine

reaching distal margin of article 4; distolateral spine almost

reaching distal margin of article 3. Article 3 and 4 unarmed.

Maxilliped 3. Ischium 1.8 x merus length, with flexor

distal spine. Merus with large spine and small distal spine on

flexor margin; extensor margin unarmed.

Pl. 1.6-1.9 pcl (females), 1.7—2.1 pcl (males), covered in

plumose setae and iridescent setae in inner margin; merus

0.6—0.7 pcl (females), 0.7—0.8 pcl (males), with row of 5 small

spines on dorsal margin; distomesial spine reaching midlength

of carpus. Carpus 0.4 (females), 0.4—0.5 (males) x merus

length, length 0.9—1.] x width, with 5 spines along the mesial

margin. Propodus 1.2-1.3 (females), 1.1—1.6 (males) x merus

length, dactylus longer than palm, 0.5 x total propodus length;

dactylus and pollex outer margin unarmed.

A.W. McCallum, S.T. Ahyong & N. Andreakis

P2-4. Relatively stout, with scales on lateral surface of

meri, carpi and propodi; extensor margin with plumose setae

and iridescent setae. Meri shorter posteriorly (P3 merus 0.9 x

P2 merus length, P4 merus 0.8 x P3 merus length). P2 1.5—2.2

x pcl; merus 0.6—0.8 x pcl, length 4 x width, 2.6—3.0 x carpus

length and 1.3-1.6 x propodus length, extensor margin with

11-17 spines, flexor margin with acute ridges and well-

developed distal spine; carpus with 2 large spines and 2 or 3

smaller spines on extensor margin, distal spine on flexor

margin; propodus length 5 x height, with 6—8 movable flexor

spines; dactylus 0.7—0.9 x propodus length, curved, length 5—6

x width, extensor margin densely lined with stiff long setae on

distal half, flexor margin with 8—12 spines, space between

spines increasing distally, unarmed along distal 1/5. End of P2

carpus almost reaching end of P1 merus. P3 with similar

spination and article proportions as P2; merus slightly shorter

than P2 merus (0.9); propodus and dactylus as long as those of

P2. P4 length 0.8 x P2 length; merus 0.5 x pcl; merocarpal

articulation reaching hepatic marginal spine of carapace.

Egg diameter. 0.5 mm.

Colour. Carapace and abdominal somites 2-3 orange,

remaining abdomen and telson white. Rostrum white with red

subdistal spot, supraocular spines orange. Chelipeds and

walking legs with transverse white/pale orange and dark orange

bands; distal half of palm and proximal half of cheliped fingers

orange, distal half of fingers of cheliped white; dactyli of

walking legs white.

Genetic data. COI and 16S; see Table 1.

Etymology. Named julumunyju with approval of the Kariyarra

people. The word julumunyju means prawn in Kariyarra

language; used as a noun in apposition.

Remarks. Munida julumunyju sp. nov. is very similar to M.

rubridigitalis Baba, 1994, described from Queensland, Australia.

Both species have a laterally compressed rostrum with a red

mark and red cheliped fingers, the second abdominal somite with

8 or 9 spines along the anterior ridge and subequal distal spines

on the basal antennular article. Comparison of M. julumunyju

with paratypes of M. rubridigitalis and additional specimens

from eastern Australia found that the two species differ chiefly in

the shape of the rostrum. The rostrum of both species is laterally

compressed, but shallower and more evenly tapered in M.

rubridigitalis. In M. rubridigitalis, the ventral rostral margin 1s

gently convex with the dorsal and ventral margins parallel for the

proximal half or slightly more, after which they gently converge

to the apex. In M. julumunyju sp. nov., however, the rostrum is

deeper, with the dorsal and ventral margins distally diverging,

reaching the greatest depth slightly anterior to the midlength and

then converging to the apex, making the ventral margin more

strongly convex than in M. rubridigitalis. In addition, M.

rubridigitalis usually has a spine on the mesial margin of the

antennal article 2 (occasionally present only on one side), which

is always absent in M. julumunyju. Colour-in-life may also be

helpful 1n distinguishing these species. The new species has a red

patch on the pterygostomian flap below the linea anomurica,

which is noted as absent in M. rubridigitalis by Baba (1994). In

M. julumunyju sp. nov. the fingers are completely red with a

Australian Munida

white tip, whereas M. rubridigitalis is described with only the

distal half of the fingers red.

We examined six specimens identified as M. rubridigitalis

from New Caledonia in the collections of the MNHN, and

herein we identify these specimens as M. julumunyju sp. nov.

These specimens agree with M. julumunyju sp. nov. 1n rostral

form and lack of a spine on the mesial margin of antennal article

2. Specimens reported and figured from New Caledonia and the

Loyalty Islands as Munida sp. by Macpherson (1994: fig. 13b,

90) also agree with M. julumunyju 1n rostral form and the colour

of the fingers, which are described as red with a white tip.

sequence divergence between M. julumunyju sp. nov. and a New

Caledonian specimen of “M. rubridigitalis" (MNHN-

[U-2014-15474) is 4% (COI) and 0.6% (16S). Thus, further study

is required to determine if other reported specimens of "M.

rubridigitalis" from the New Caledonian region are conspecific

with, or distinct from, M. julumunyju sp. nov. The apparently

disjunct distribution of M. julumunyju sp. nov. suggests that it

can be expected to occur at intermediate localities, possibly off

southern Indonesia and Papua New Guinea.

Distribution. Presently known from off Western Australia, 373—

450 m. New Caledonia, 450—515 m. Loyalty Islands, 540 m.

Munida jurunjurun sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act: CD77DA72-B760-

4983-9252-6BC62CC7CE50

Figures 8,9

Type material. Holotype: WAM C78561, ovigerous female (cl 14.7

mm, pcl 9 mm), Western Australia, off Imperieuse Reef, 17° 35.982' S,

118? 59.076' Eto 17? 38.574' S, 119° 01.26' E,222-319m,S8805/2007/63,

16 June 2007.

Description. Carapace. Length 1.1 x greatest width, widest at

midlength. Dorsal surface with numerous uninterrupted

transverse ridges and secondary transverse striae between main

ridges; ridges and striae lined with short, non-iridescent setae.

Gastric region with 5 pairs of epigastric spines, longest pair

behind supraocular spines. Parahepatic, postcervical and dorsal

branchial spine present. Anterior part of branchial region

between cervical groove and postcervical groove with 3 main

ridges; posterior part of branchial region with 6 main transverse

ridges (excluding posterodorsal ridge) and 5 secondary striae

laterally between main ridges. Cardiac region with 5 main

transverse ridges. Intestinal region with transverse ridge,

posterodorsal ridge and secondary stria. Frontal margin inclined

posteriorly at 112° from midline. Lateral margins slightly

convex; anterolateral spines parallel, horizontal, overreaching

sinus between rostrum and supraocular spine; small hepatic

marginal spine in front of cervical groove (0.15 x length of

anterolateral spine); branchial margin with 4 spines. Rostrum

spiniform, horizontal, 0.7 x pcl; supraocular spine 0.5 x length of

rostrum. Epistomial ridge straight ending slightly anterior to

antennal gland; mesial protuberance distinct.

Thoracic sternum. Sternal surface smooth, sternite 4 with

few short striae. Sternite 3 0.4 width of sternite 4; midlength

of sternal plastron (sternites 4—7) 0.5 width of sternite 7.

Sternite 4 anterior margin broadly rounded.

125

Abdomen. Somite 2 with 2 pairs of small spines at lateral

margins of anterior transverse ridge. Somites 2—4 each with 2

or 3 uninterrupted striae behind anterior ridge and some

shorter striae. Somite 6 posteromedian margin almost

straight. Telson with numerous transverse squamae; greatest

width 2.0 x median length; anterolateral margin almost

straight or shallowly concave.

Eye. Maximum corneal diameter 0.36 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded)

overreaching or reaching end of corneae; 2 well-developed

distal spines, distomesial slightly longer than distolateral; 2

lateral spines, proximal small, distolateral spine exceeding

both distal spines.

Antenna. Article 1 with strong distomesial spine

overreaching distal margin of article 4. Article 2 with strong

distomesial spine overreaching distal margin of article 4;

distolateral spine reaching distal margin of article 4; 2 small

lateral spines on mesial margin. Articles 3 and 4 unarmed.

Maxilliped 3. Ischium 1.45 x merus length, with strong

flexor distal spine; merus with strong spine on flexor margin and

small spine distally, with small spine on distal extensor margin.

P1. length 3.0 x pcl, with iridescent setae on inner margin.

Merus 1.25 x pcl, with a row of 5 dorsal spines and row of 5

long, close set spines on mesial margin; distal spines strong,

distomesial spine not reaching midlength of carpus. Carpus 0.4

x merus length, length 2.6 x width, with spines along mesial

and dorsal margins. Propodus 1.1 x merus length, fingers 0.5 x

total propodus length, with 2 small distolateral spines; dactylus

with small proximal spine small subdistal spine.

P2-4. Long and slender, with numerous scales on lateral

surfaces of meri; margins with row of plumose setae and

iridescent setae on ischium and merus. P2 length 2.6 x pcl; merus

as long as pcl, length 6 x height, 4.0 x carpus length and 1.2 x

propodus length, extensor margin spinose, flexor margin with

small spine near distal one-fifth and well-developed distal spine;

carpus extensor margin with 5 small spines (broken on right P2

of the holotype) and prominent distal spine, flexor margin with

distal spine; propodus length 9.3 x height, with 11 small movable

flexor spines; dactylus compressed, slightly curved, length 0.5 x

propodus length and 5 x height, with 10 evenly spaced spines

along distal 0.8 of flexor margin. End of P2 carpus reaching end

of Pl merus. P3 with similar spination and article proportions as

P2; merus slightly shorter than P2 merus (0.85); propodus and

dactylus as long as those of P2. P4 length 0.9 x P2 length; merus

0.7 x pcl, 0.8 x P3 merus length; propodus 0.85 and dactylus

0.95 x length of those of P3; merocarpal articulation slightly

overreaching anterolateral corner of carapace.

Genetic data. COI and 16S; see Table 1.

Etymology. Named jurunjurun with the approval of Bardi

elders, after Jurun jurun, meaning crayfish 1n Bardi; used as a

noun in apposition.

Remarks. Munida jurunjurun sp. nov. is closest to M. acantha

Macpherson, 1994, from New Caledonia, both species having

antennal articles | and 2 with long mesial spines overreaching

article 4, antennule with distomesial spine longer than

126 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 8. Munida jurunjurun sp. nov., holotype, ovigerous female cl 14.7 mm, pcl 9 mm (WAM C78561). A, carapace and abdomen, dorsal view; B,

right chela, dorsal view; C, right antenna and antennule, ventral view; D, sternum; E, abdominal somite 6, telson and right uropod. Scale: A-E = 2 mm.

Australian Munida

distolateral spine, a smooth thoracic sternum, and abdominal

somite 2 with small spines limited to the lateral margins of the

anterior ridge. The new species differs from M. acantha in

having 4 rather than 5 branchial carapace spines. In addition,

the new species has a distinct row of spines on the mesial

margin of the Pl merus, which are close-set on the distal half

of the article, rather than more evenly spaced along the distal

0.6 as in M. acantha. Sequence divergence between M. acantha

and M. jurunjurun sp. nov. is 8% for COI and 3.5% for 16S.

Of those species with 4 branchial spines, M. jurunjurun

127

sp. nov. is similar to M. albiapicula Baba and Yu, 1987, M.

zebra Macpherson, 1994, M. volantis Macpherson, 2004, and

M. nesiotes Macpherson, 1999, all of which have a long mesial

spine on antennal articles 1 and 2. In these species, however,

the spines on the anterior ridge of abdominal somite 2 are

evenly spread along the entire margin, rather than placed at

the lateral extremities.

Distribution. North-western Australia, off Imperieuse Reef,

222—319 m

Figure 9. Munida jurunjurun sp. nov., holotype, ovigerous female cl 14.7 mm, pcl 9 mm (WAM C78561): A, right P2 dactylus, lateral view; B,

right P2, lateral view; C, right P3, lateral view; D, right P4, lateral view; E, left maxilliped 3, lateral view. Scale A= 1 mm, B-E = 2 mm.

128

Munida leeuwin sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:6D615718-C84B-

4]6F-891 A-4D9FA BFB7592

Figures 10, 1]

Munida sp. MoV 5176. — Poore et al., 2008: 20: unnumbered

colour fig. (lower right) (south-western Australia, 101—100 m).

Type material. Holotype. WAM C78562, male (cl 8.1 mm, pcl 5.3

mm), Western Australia, Barrow Island, 20° 59.082' S, 114° 54.42' E

to 20° 59.67 S, 114° 54.54' E, 100-101 m, SS10/2005/170,

13 December 2005.

Paratypes (all Western Australia) WAM C78563, 1 ovigerous

female (cl 7.2 mm, pcl 5 mm), 7 females (cl 6.0 mm, pcl 3.8 mm to cl

5.5 mm, pcl 5.5 mm), 7 males (cl 6.3 mm, pcl 3.8 mm to cl 9.1 mm, pcl

6.2 mm), collected with holotype; NMV J55046, 1 male (cl 8.7 mm,

pel 5.9 mm), 1 ovigerous female (cl 7.9 mm, pcl 5.2 mm), collected

with holotype; NMV J55100, 2 ovigerous females (cl 7.7 mm, pcl 5.0

mm; cl 7.8 mm, pcl 5.1 mm), | female (cl 7.8 mm, 5.0 mm), collected

with holotype; NMV J56100, 1 juvenile (cl 4.4 mm, pcl 3.0 mm), 1

ovigerous female (broken rostrum; pcl 4.8 mm), Imperieuse L23

transect, 18° 27.612' S, 120° 08.682' E to 18° 27.72' S, 120° 08.682' E,

80—81 m, SS05/2007/82, 16 June 2007.

Description. Carapace. Length 1.1 x greatest width, widest at

midlength. Dorsal surface with main transverse ridges mostly

uninterrupted, without secondary transverse striae between

main ridges; ridges and striae lined with short, non-iridescent

setae and few long iridescent setae. Gastric region with 5 or 6

pairs of epigastric spines, longest pair behind supraocular spines,

with median row of 2 or 3 small spines behind rostrum. Hepatic

region with 3 or 4 spines on dorsal surface; parahepatic spines

present. Anterior part of branchial region between cervical

eroove and postcervical groove with 3 ridges and | or 2 small

spines; | postcervical spine; posterior part of branchial region

with 4 or 5 main transverse ridges (excluding posterodorsal

ridge) and few shorter secondary striae between main ridges.

Cardiac region with 3 main transverse ridges. Intestinal region

without striae; posterodorsal ridge distinct, without secondary

stria. Frontal margin inclined posteriorly at 115° from midline.

Lateral margin slightly convex; anterolateral spines parallel,

horizontal, not reaching sinus between rostrum and supraocular

spine; hepatic marginal spine shorter than anterolateral spine;

branchial margin with 5 spines. Rostrum spiniform, slightly

sinuous in profile, length 0.5 x pcl; supraocular spine 0.3 x length

of rostrum. Epistomial ridge curved, ending anterior to antennal

gland; mesial protuberance distinct.

Thoracic sternum. Sternital surface smooth, sternite 4 with

few long striae. Sternite 3 0.4 width of sternite 4; midlength of

sternal plastron (sternites 4—7) 0.7 width of sternite 7. Sternite

4 anterior margin truncate, entirely contiguous with sternite 3.

Abdomen. Somite 2 without spines. Somites 2—4 each with

2 or 3 uninterrupted striae behind anterior ridge and striae at

lateral margins. Somite 6 posteromedian margin almost

straight. Telson with numerous transverse squamae; greatest

width 2.0 x median length; anterolateral margin concave.

Eye. Maximum corneal diameter 0.35 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded) not

overreaching corneae; distolateral and distomesial spines

A.W. McCallum, S.T. Ahyong & N. Andreakis

subequal; 2 lateral spines, proximal smaller, longer lateral

spine exceeding distal spines.

Antenna. Article 1 with distomesial spine reaching

midlength of article 2. Article 2 distomesial spine slightly

overreaching distal margin of article 5; distolateral spine almost

reaching distal margin of article 3. Article 5 unarmed or with

small distolateral spine. Article 4 unarmed.

Maxilliped 3. Ischium 1.6 x merus length, with distal

flexor spine. Merus with 3 well-developed spines on flexor

margin; extensor margin with small distal spine.

Pl. length 3.1—4.4 pcl (males), 2.8—3.0 pcl (females), with

dense covering of iridescent setae on inner margins of merus

(without plumose setae), merus 1.3—1.4 (males), 1.0—1.1 (females)

x pcl, with row of 5 subtriangular spines on dorsal and mesial

margin; distomesial spine not reaching midlength of carpus.

Carpus 0.3-0.4 x merus length, length 1.9 (males), 2.02.4

(females) x width, with spines along mesial margin. Propodus

1.6-1.7 (males), 1.4 (females) x merus length; fingers 0.4—0.6

(males), 0.5—0.6 (females) x total propodus length; pollex with

row of small spines along outer margin; dactylus with 3 small

spines on outer margin.

P2—4. Moderately long and slender, with scales on lateral

surfaces of meri; extensor margin with plumose setae and

iridescent setae. P2 length 2.1-2.6 x pcl; merus as long as

carapace, length 6.4 x height, 3.4 x carpus length and 1.3 x

propodus length; extensor margin spinose; flexor margin with

acute ridges, | or 2 spines and well-developed distal spine;

carpus with 4 small extensor spines, distal spine on extensor and

flexor margin; propodus length 7 x height, with 10—14 movable

flexor spines; dactylus compressed, almost straight, 0.7—0.9 x

propodus length, length 6—7 x height, flexor margin with 7—12

spines, unarmed along distal 1/3. End of P2 carpus not reaching

end of Pl merus. P3 with similar spination and article

proportions as P2; merus slightly shorter than P2 merus (0.8);

propodus and dactylus as long as those of P2. P4 length 0.8 x P2

length; merus 0.6—0.7 x pcl, 0.9 x P3 merus length; merocarpal

articulation almost reaching marginal hepatic spine of carapace.

Genetic data. COI and 16S; see Table 1.

Etymology. Named after the Leeuwin current which flows off

the west coast of Australia; used as a noun in apposition.

Remarks. Munida leeuwin sp. nov. is most similar to M. roshanei

Tirmizi, 1966, M. janetae Tirmizi and Javed, 1992, and M.

arabica Yirmizi and Javed, 1992, described from the western

Indian Ocean. These three species are very similar to each other

and have been distinguished by the length of the pereopod 1

chela, the ratio of the length of the fingers to the propodus palm,

and the shape of sternite 3. In his key, Baba (2005) characterises

M. janetae as having cheliped fingers that are distinctly longer

than the palm, while the fingers are shorter than the palm in M.

roshanei and M. arabica. In M. leeuwin sp. nov., the fingers are

usually as long as the palm but are occasionally longer or shorter

than the palm (0.4—0.6 x propodus length). We tentatively

describe this species as new based on a combination of subtle

differences from the aforementioned species and significant

molecular divergence from material we identify as M. roshanei.

In M. leeuwin sp. nov., the dactlyli of P2—4 are slender and

Australian Munida 129

Figure 10. Munida leeuwin sp. nov., holotype, male cl 8.1 mm, pcl 5.3 mm (WAM C78562): A, carapace and abdomen, dorsal view; B, right chela,

dorsal view; C, left antenna and antennule, ventral view; D, sternum; E, abdominal somite 6, telson and right uropod. Scale: A-E = 2 mm.

130

unarmed on the distal one-third of the flexor margin, with the

ultimate spine closer to the penultimate spine than the unguis.

This differs from M. janetae in which the dactyli have spines

regularly arranged along the entire flexor margin (Tirmizi and

Javed, 1992). Illustrations of M. arabica show the dactyli to be

similar to those of M. leeuwin sp. nov., but M. arabica can be

distinguished by the presence of distal spines on antennal article

4, which are always absent in M. leeuwin sp. nov. The type

description of M. roshanei lacks a description or illustration of

the dactyli, as does that of Lewinsohm (1969) for nearby Red

sea specimens. Specimens identified as M. roshanei from the

Phillipines (Baba, 1988) and Australia (present study), however,

A.W. McCallum, S.T. Ahyong & N. Andreakis

have the dactyli with spines regularly arranged along the entire

flexor margin. Although the spination of the P2—4 dactyli of the

type material of M. roshanei remains to be confirmed, given the

brevity of the original description and without access to the

type, we fully describe M. leeuwin as new to improve the

taxonomy of this difficult group.

The genetic sequences of M. leeuwin are highly divergent

from all other species analysed. Despite their morphological

similarities, M. leeuwin and our specimens of M. roshanei are

divergent by 12% in COI.

Distribution. Presently known only from off Western Australia,

658—754 m.

Figure 11. Munida leeuwin sp. nov., holotype, male cl 8.1 mm, pcl 5.3 mm (WAM C78562): A, right P2 dactylus, lateral view; B, right P2, lateral

view; C, right P3, lateral view; D, right P4, lateral view; E, left maxilliped 3, lateral view. Scale A-E = 1 mm.

Australian Munida

Munida lutruwita sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:14904374-57EC-

4548-A4D7-ACA713F712C9

Figures 3E, 12, 13

Munida cf. manqingae. — Farrelly and Ahyong, 2019: 13,55, fig. 99.

Type material. Holotype: NMV J67481, ovigerous female (cl 11.7 mm,

pcl 7.8 mm), Tasmania, south of Hobart, Z39 Seamount, 44° 23.346' S,

147° 16.376' E, 2040 m, TT 01/2008/J2-387-003, 25 December 2008.

Paratypes (all Tasmania): NMV J67474, 2 ovigerous females (cl 9.5

mm, pcl 7.1 mm; cl 9.6 mm, pcl 7.4 mm), Z39 Seamount, 44° 23.32' S,

147? 15.349' E, 1599 m, TT 01/2008/J2-387-023, 26 December 2008;

NMV J67475, 2 damaged ovigerous females (cl 10.6 mm, pcl 7.3 mm;

cl 11.2 mm , pcl 7.9 mm), Z39 Seamount, 44° 22.994' S, 147° 15.09' E,

1893 m, TT 01/2008/J2-387-008, 25 December 2008; NMV J67480, 1

ovigerous female (cl 11.5 mm, pcl 79 mm), Z39 Seamount,

44? 23.295' S, 147? 15.348' E, 1616 m, TT 01/2008/J2-387-022, 26

December 2008; NMV J67472, | male (abdomen missing; cl 10.9 mm,

pel 6.3 mm), Z39 Seamount, 44° 23.345'S 147° 16.277'E, 1990—2004

m, TT 01/2008/J2-387-004, 25 December 2008.

Other material examined. AM P103229, 1 ovigerous female (cl 14.3

mm, pcl 10.0 mm), Great Australian Bight, 35° 22.627' S,

132° 19.166' E, 1689-1784 m, from crevice in dead coral,

IN2015-C01-042, 6 November 2015.

Description. Carapace. As long as greatest width, widest at

midlength. Dorsal surface with main transverse ridges mostly

uninterrupted, without secondary transverse striae between

main ridges; ridges and striae lined with short, non-iridescent

setae. Gastric region with 3 pairs of epigastric spines and 1 or

2 small additional spines, without median row of spines behind

rostrum. Hepatic region without spines on dorsal surface.

Anterior part of branchial region between cervical groove and

postcervical groove with 2 or 3 short tuberculate ridges and

often | small spine anteriorly; posterior part of branchial region

with 5 transverse ridges (excluding posterodorsal ridge).

Cardiac region with 2 main transverse ridges. Intestinal region

without striae; posterodorsal ridge distinct, without secondary

stria. Frontal margin strongly oblique, inclined posteriorly at

115° from midline. Lateral margin slightly convex; anterolateral

spine very small, far from reaching sinus between rostrum and

supraocular spine; hepatic marginal spine slightly smaller than

anterolateral spine; branchial margin with 5 spines. Rostrum

spiniform, 0.5 x pcl; supraocular spine 0.25 x length of rostrum,

exceeding eyes. Epistomial ridge straight ending at antennal

gland; mesial protuberance distinct.

Thoracic sternum. Sternal suface smooth, sternite 4 with

only few striae. Sternite 3 0.4 x width of sternite 4. Sterinte 4

anterior margin triangular, narrowly contiguous with sternite

3. Midlength of sternal plastron (sternites 4—7) 0.5 x width of

sternite 7.

Abdomen. Somites smooth without spines, distinct ridges

or striae. Somite 6 posteromedian margin slightly concave.

lelson with few striae; greatest width 1.2 x median length;

anterolateral margin weakly concave.

Eye. Maximum corneal diameter 0.18 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded)

overreaching corneae; distolateral spine much longer than

131

distomesial spine; 2 lateral spines, proximal smaller, longer

lateral spine not reaching end of distolateral spines.

Antenna. Article | distomesial spine almost reaching

distal margin of article 2. Article 2 distomesial spine reaching

distal margin of article 3; distolateral spine not reaching

midlength of article 3. Articles 3 and 4 unarmed.

Maxilliped 3. Ischium 1.9 x merus length, without flexor

distal spine. Merus with large median spine and distal spine

on flexor margin; extensor margin without distal spine.

P1. Length 2.4—3.2 x pcl, covered in rows of short plumose

setae. Merus length 0.9—1.1 x pcl, with row of 2 large spines

and 2 small spines on dorsal margin, 1 strong spine on

dorsolateral margin, and 4 spines on mesial margin,

distomesial spine not reaching midlength of carpus. Carpus

0.5 x merus length, length 3.0 x width, with 6 spines along

mesial margin. Propodus 1.3 x merus length, palm with row

of 3 or 4 spines on dorsal surface of palm, fingers 0.4—0.5 x

total propodus length, without spines on outer margins.

P2-4. Long and slender, with few small scales on lateral

sides of meri and carpi; extensor margin with short plumose setae

and few longer setae. P2 1.8—2.3 x pcl; merus 0.7-0.8 x pcl,

length 8.0 x width, 3.0 x carpus length and 1.5 x propodus

length, extensor margin with 5—7 spines, flexor margin with 3

spines and well-developed distal spine; carpus extensor margin

with spine at midlength and distal end, flexor margin with distal

spine; propodus length about 8 x height, with 5 movable flexor

spines on flexor margin; dactylus gently curved distally, 0.6—0.7

x propodus length, length about 7 x height, extensor margin

densely lined with stiff short setae on distal half, flexor margin

armed along entire length with 12—14 movable spines including

spine at base of unguis. End of P2 carpus not reaching end of PI

merus. P3 with similar spination and article proportions as P2;

merus 0.9 x P2 merus length, propodus and dactylus as long as

those of P2. P4 length 0.7—0.8 x P2 length; merus length 0.3—0.5

x pcl; propodus and dactylus similar in length to those of P3;

merocarpal articulation reaching hepatic marginal spine carapace.

Egg diameter. 0.65 mm.

Colour in life. Carapace pink anteriorly fading to white at

posterior, abdominal somite 2 white, somites 3—6 pink. P1 and

P2-4 white.

Genetic data. COI and 16S; see Table 1.

Etymology. Named lutruwita, the original name of Tasmania in

palawa kani,the language of the Tasmanian Aboriginal people,

with the approval of the Tasmanian Aboriginal Centre; used as

a noun in apposition.

Remarks. Munida lutruwita sp. nov. is very close to M.

manqingae Liu, Lin and Huang, 2013, described from a single

female specimen collected from hydrothermal vent fields at

2218 m in the south-western Indian Ocean. The spination of the

antenna and antennule is identical between the two species, but

the new species differs from M. manqingae 1n the following:

- the anterior branchial surface of M. lutruwita is rugose,

with 2 or 3 rows of small tubercles and a small spine

anteriorly, but smooth in M. manqingae.

- the P2 dactylus has 12-14 movable flexor spines in M.

lutruwita, but only 10 in M. manqingae.

132 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 12. Munida lutruwita sp. nov., holotype, ovigerous female, cl 11.7 mm, pcl 7.8 mm (NMV J67481): A, carapace and abdomen, dorsal view;

B, right chela, dorsal view; C, right antenna and antennule, ventral view; D, sternum; E, telson. Scale: A-B = 1 mm, C-E = 2 mm.

Australian Munida

Munida lutruwita is also very similar to Munida

magniantennulata Baba and Türkay, 1992, described from

hydrothermal vents in the Lau Basin near Fiji, and the recently

described, M. alba Liu, Li and Lin, 2020, from the Eastern

Pacific Rise. Differences between the three species are as follows:

- the antennal article | mesial spine reaches the end of

article 2 1n M. lutruwita and M. alba, but does not reach as far

as the midpoint of article 2 1n M. magniantennulata.

- the supraocular spines are parallel or slightly convergent,

overreaching the eyes in M. lutruwita, while in M.

magniantennulata and M. alba, the suproacular spines are

133

slightly divergent and do not overreach the eyes.

- the frontal margins of the carapace are oblique in M.

lutruwita and M. alba rather than near-transverse in M.

magniantennulata.

- M. lutruwita and M. magniantennulata further differ

from M. alba in the presence of the distal flexor spine on the

maxilliped 3 merus (absent in M. alba) and unarmed outer

margin of the cheliped pollex (spinose in M. alba).

Distribution. Off southern Tasmania and the Great Australian

Bight, 1599-2040 m.

Figure 13. Munida lutruwita sp. nov., holotype ovigerous female, cl 11.7 mm, pcl 7.8 mm (NMV J67481): A, right P2 dactylus, lateral view; B,

left P2, lateral view; C, left P3, lateral view; D, left P4, lateral view; E, left maxilliped 3, lateral view. Scale = 1 mm.

134

Munida maatijadakurnaaku sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:354E2119-60D1I-

498B-9595-32D05B1968BF

Figures 14, 15

Munida aff. rubiesi Macpherson, 1991. — Poore et al., 2008.

Type material. Holotype: WAM C78564, male (rostrum broken; pcl

9.0 mm), Western Australia, south-west of Kalbarri, 27° 56.106' S

113? 4.86' E to 27? 56.646' S 113? 5.28' E, 417—428 m, beam trawl,

$510/2005/98, 4 December 2005.

Description. Carapace. Length 1.2 x greatest width, widest at

midlength. Dorsal surface with numerous uninterrupted

transverse ridges and secondary transverse striae between

main ridges; ridges and striae lined with short, non-iridescent

setae and scattered long setae. Gastric region with 4 pairs of

epigastric spines, longest pair behind supraocular spines.

Hepatic region with short scales; parahepatic spine present.

Anterior part of branchial region between cervical groove and

postcervical groove with dorsal spine and 5 or 6 main ridges; |

postcervical spine; posterior part of branchial region with 8

main transverse ridges (excluding posterodorsal ridge) and 5

secondary striae laterally between main ridges. Cardiac region

with 5 main transverse ridges. Intestinal region with 2

transverse ridges, posterodorsal ridge distinct, with secondary

stria. Frontal margin inclined posteriorly at 106° from midline.

Lateral margins slightly convex; anterolateral spine slightly

divergent, horizontal, overreaching sinus between rostrum and

supraocular spine; hepatic marginal spine distinctly smaller

than anterolateral spine; branchial margin with 5 spines.

Rostrum broken; supraocular spines 0.4 x pcl, exceeding eyes.

Epistomial ridge straight ending slightly anterior to antennal

gland; mesial protuberance distinct.

Thoracic sternum. Sternal surface smooth, sternites 4 and

5 with few short striae. Sternite 3 0.4 x width of sternite 4;

median length of sternal plastron (sternites 4—7) 0.7 x width

of sternite 7. Sternite 4 anterior margin subtriangular,

narrowly contiguous with sternite 3.

Abdomen. Somite 2 with 5 pairs of spines on anterior

transverse ridge. Somites 2—4 each with 5 or 6 uninterrupted

striae behind anterior ridge. Somite 6 posteromedian

marginslightly convex, posterolateral margins slightly produced.

Telson with numerous transverse squamae; greatest width 1.6 x

median length; anterolateral margin slightly concave.

Eye. Maximum corneal diameter 0.4 x distance between

anterolateral spines.

Antennule. Basal article (distal spines excluded) overreaching

corneae; 2 long, subequal distal spines; 2 lateral spines, proximal

smaller, distolateral spine not exceeding distal spines.

Antenna. Article 1 with distomesial spine overreaching

distal margin of article 2. Article 2 with strong distomesial

spine significantly overreaching distal margin of article 4;

distolateral spine reaching distal margin of article 3. Articles

3 and 4 unarmed.

Maxilliped 3. Ischium 1.5 x merus length, with strong flexor

distal spine; merus with strong spine on flexor margin and small

spine distally, without spine on distal extensor margin.

Pl. Length 2.8 x pcl, with long plumose setae and

A.W. McCallum, S.T. Ahyong & N. Andreakis

iridescent setae, setae longest on dorsal surfaces. Merus length

1.1 x pcl, with a row of 8 dorsal spines and row of 4 spines on

mesial margin; distal spines strong, distomesial spine not

reaching midlength of carpus. Carpus 0.4 x merus length,

length 1.9 x width. Propodus 1.1 x merus length, fingers 0.6 x

total propodus length; pollex with row of 5 spines on lateral

margin; dactylus with row of 3 spines on dorsal margin, 5

spines on lateral margin and | small subdistal spine.

P2-4. Long, slender, with numerous scales on lateral sides

of meri; extensor margins with row of plumose and iridescent

setae. P2 length 2.3 x pcl, merus as long as carapace, length

about 7 x height, 4.0 x carpus length and 1.9 x propodus

length, row of spines on extensor margin, flexor margin row

of spines and well-developed distal spine; carpus with 2 small

spines and 2 large spines on extensor margin, distal spine on

flexor margin; propodus length 5.3 x height, with 9 small

movable flexor spines; dactylus compressed, almost straight,

as long as propodus length, length 8.6 x height, with 7

movable spines along the flexor margin, distal one-quarter

unarmed. P3 with similar spination and article proportions as

P2; merus slightly shorter than P2 merus (0.75); propodus and

dactylus slightly shorter than those of P2. P4 length 0.6 x P2

length; merus 0.4 x pcl, length 0.6 x P3 merus length;

propodus 0.8 and dactylus 0.9 x as long as those of P3.

Etymology. Named maatijada for crawling and kurnaaku for

crayfish/yabbie in Nhanda language at the Nhanda Language

Day (2 July 2019) at Bundiyarra-Irra Wanga Language Centre

in Geraldton; used as a noun in apposition.

Remarks. Despite the broken rostrum in the holotype, Munida

maatijadakurnaaku sp. nov. 1s clearly distinguished by a

number of diagnostic characters. It is most similar to M.

aequalis Ahyong and Poore, 2004, from eastern Australia,

which also has long and subequal distal spines on the basal

antennular article, a subtriangular sternite 4, slender P2—4

dactyli and similar antennal spination. The two species can be

distinguished by:

- the supraocular spines overreach the eyes in M.

maatijadakurnaaku but do not reach the end of the eyes in M.

aequalis

- antennal article 2 has the distomesial spine significantly

overreaching article 4 in M. maatijadakurnaaku, but only

slightly overreaching article 4 1n M. aequalis

- the P2—4 dactyli are as long as their respective propodi

in the new species, with only the distal one-quarter unarmed,

rather than slightly shorter than the propodus and unarmed on

the distal one-third in M. aequalis.

Distribution. Known only from Western Australia, 417—428 m.

Munida agave Macpherson and Baba, 1993

Figures 16A

Munida agave Macpherson and Baba, 1993: 387, figs 1, 2 (type

locality: Philippines). — Baba et al., 2008: 84. — Baba et al., 2009:

139, fig. 115.

Material examined. Western Australia: NMV J56008, 1 female (cl

11.6 mm, pcl 10.4 mm), Broome L25 transect, 16° 45.156' S,

Australian Munida 135

Figure 14. Munida maatijadakurnaaku sp. nov., holotype, 1 male with missing rostrum, pcl 9.0 mm (WAM C78564): A, carapace and abdomen,

dorsal view; B, right chela, dorsal view; C, right antenna and antennule, ventral view; D, sternum; E, telson. Scale A-E = 2 mm.

136

A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 15. Munida maatijadakurnaaku sp. nov., ] male with missing rostrum, pcl 9.0 mm (WAM C78564): A, right P2 dactylus, lateral view; B,

left P2, lateral view; C, left P3, lateral view; D, left P4, lateral view; E, left maxilliped 3, lateral view. Scale: A-E = 1 mm.

121° 027796 E to 16° 44.592' S, 121° 02.208' E, 100-108 m,

$505/2007/116, 30 June 2007; NMV J56099, Adele L28 transect,

14° 33.732' S, 122? 55.092' E to 14? 33.684' S, 122? 54.906' E, 95-105

m, $505/2007/161, 4 July 2007.

Colour. Carapace and abdominal somites 2—5 pale orange with

orange striae and spines, telson whiteish. Rostrum and

supraocular spines orange. Pl and P2-4 pale orange with

orange striae.

Genetic data. COI and 16S; see Table 1.

Remarks. This specimen agrees well with the description of the

holotype from the Phillipines, having arcuate striae on sternites

3—6 and two pairs of spines on the anterior ridge of abdominal

somite 2. This 1s the first record of the species from Australia.

Distribution. Japan, Philippines, Indonesia, Taiwan, 89-549 m.

North-western Australia, 55—108 m.

Australian Munida 13/

Figure 16. A, Munida agave Macpherson and Baba, 1993, female, cl 11.6 mm, pcl 10.4 mm (NMV J56008); B, Munida armilla Macpherson, 1994,

male, cl 16.1 mm, pcl 10.2 mm (NMV J55081); C-D, Munida asprosoma Ahyong and Poore, 2004, male, cl 13.2 mm, pcl 8.0 mm (J55076),

ovigerous female, cl 17.4 mm, pcl 10.7 mm (NMV J56016). Photos: K. Gowlett-Holmes.

138

Munida armilla Macpherson, 1994

Figure 16B

Munida armilla Macpherson, 1994: 446, figs 6, 65 [type locality:

New Caledonia]. — Baba et al., 2008: 87. — Macpherson, 2013: 296.

Munida aff. amathea — Poore et al., 2008: 20.

Material examined. Western Australia: NMV J55081, 1 male (cl 16.1

mm, pcl 10.2 mm),

Abrolhos, 28° 29.37' S, 113° 25.14' E to 28° 30.06' S, 113° 25.5' E,

416—431 m, SS10/2005/95, 2005.

Colour. Carapace and abdominal somites pink/white with

orange/reddish striae and spines. Rostrum white with red tip,

supraocular spines red at base and white distally, anterolateral

spines red. Walking legs with orange/red bands. Chela with

orange bands.

Genetic data. Not available.

Remarks. This single specimen differs slightly from the type

material of M. armilla Macpherson, 1994. In our specimen the

basal article of the antennule has subequal distal spines while

in the holotype the distomesial spine is slightly shorter than the

distolateral. In our specimen the carapace has interupted striae

while the holotype has numerous scales. This is the first record

of the species from Australia.

Distribution. New Caledonia, Matthew and Hunter Islands,

Tuscaroa Bank, Tonga, French Polynesia (Tarava seamounts,

Tuamoto and Society Archipelagos), 233—710 m. Western

Australia, 416—431 m.

Munida asprosoma Ahyong and Poore, 2004

Figure 16C—D

Munida asprosoma Ahyong and Poore, 2004: 20, fig. 3 (type

locality: east of Broken Bay, New South Wales, Australia). — Baba et

al., 2008: 87. — Baba et al., 2009: 146, figs 123—126. — Poore et al.,

2011: pl. 16D. — McEnnulty et al., 2011: app. 1.

Material examined. Western Australia: NMV J56016, 2 ovigerous

females (cl 16.3 mm, pcl 10.2 mm; cl 17.4 mm, pcl 10.7 mm), Mermaid

L24 transect, 16° 38.064' S, 119° 09.216' E to 16° 38.766' S,

119? 08.034' E, 983—993 m, 8805/2007/68, 17 June 2007;

NMV J56096, 1 female (cl 13.9 mm, pcl 8.3 mm), 1 juvenile (cl

10.3 mm, pcl 6.0 mm), Mermaid L24 transect, 16° 44.286' S,

119° 15.042' E to 16° 43.794' S, 119° 15.48' E, 693-698 m,

SS05/2007/70, 17 June 2007; NMV J56393, 2 females (pcl 6.7-8.1

mm; #B, ZA), 1 male (pcl 5.0 mm; #C), Leveque L27 transect,

14° 33.06' S, 121° 15.366' E to 14? 33.588' S, 121° 16.56' E, 1021—1023

m, SS05/2007/155, 3 June 2007; NMV J57258, 1 ovigerous female (cl

20.6 mm, pcl 12.7 mm), $805/2007/155; NMV J55077, 1 male (cl 20.4

mm, pcl 12.2 mm), Perth Canyon, 31° 57.9' S, 115° 06.3' E to

31° 56.982' S, 115? 07.08' E, 928-1170 m, SS10/2005/73, 30 November

2005; NMV J55078, 4 ovigerous females (cl 18.7 mm, pcl 10.7 mm to

cl 22.1 mm, pcl 13.6 mm), 19 females (cl 14.4 mm, pcl 8.6 mm to cl

23.5 mm, pcl 14.5 mm), 11 males (cl 12.4 mm, pcl 7.4 mm to cl 26.5

mm, pcl 15.9 mm), SS10/2005/73; NMV J55076, 3 males (cl 13.2 mm,

pcl 8.0 mm to cl 21.5 mm, pcl 12.8 mm), Abrolhos, 29° 00.594' S,

113? 42.78' E to 29? 01.512' S, 113? 43.32' E, 700—704 m, SS10/2005/85,

2 December 2005; NMV J55079, 1 female (cl 16.9 mm, pcl 10.4 mm),

5 males (cl 15.9 mm, pcl 9.6 mm to cl 26.1 mm, pcl 16.1 mm),

SS10/2005/85.

A.W. McCallum, S.T. Ahyong & N. Andreakis

Colour. Carapace, abdomen, chela and P2—4 pale pink. Reddish

on distal portion of fingers, antenna and posterior margin of

carapace. One specimen photographed is different in colour

from the other two photographed specimens. This specimen

was collected from the Abrolhos station at ~/00 m (male, cl

13.2 mm, pcl 8.0 mm, NMV J55076). The carapace is white

with orange on the striae and abdominal somites 2—3, P2—4 are

white, the chelae are orange with white fingers.

Genetic data. COI; see Table 1.

Remarks. The present specimens of M. asprosoma from Western

Australia accord well in most respects with those from eastern

Australia (Ahyong and Poore, 2004). Three juveniles (NMV

J56393; pcl 5.0—8.1 mm), however, differ from adults in having

lesser developed abdominal spination: the median pair of spines

on the anterior ridge of abdominal somite 2 is well developed but

spines laterad are either incipent or minute, and the anterior ridge

of somite 3 is unarmed. Thus, the diagnostic spination of the

abdominal somites may not be fully developed in juveniles and

care should be taken when identifying small specimens.

The COI sequences of M. asprosoma were only 1.3%

divergent from Munida hoda Macpherson, Rodriguez-Flores

and Machordom, 2017, from Mozambique. Intraspecific

divergence between the three specimens sequenced here was

0.3—0.6%. These species share a number of morphological

characters including: the presence of 5 spines on the branchial

lateral margins of the carapace, strongly oblique frontal

carapace margins, absence of secondary striae between the

widely spaced primary striae, spines along the anterior ridge

of the abdominal somite 2, large eyes, and the distomesial

spine of the antennular article | shorter than the distolateral

spine. They can easily be distinguished by the presence of

spines on abdominal somite 3 in adults of M. asprosoma, and

the flexor spination of the P2 dactyli, with 7—9 spines in M.

asprosoma and only 4 spines in M. hoda. One of the

specimens photographed had a different colour pattern from

the others (NMV J55076, male pcl 8.0 mm; see above) but we

could find no other morphological characters to distinguish

this specimen from the others. Similarly, Baba et al. (2009)

showed a number of distinct colour morphs of M. asprosoma

from Taiwan, not dissimilar to the variation reported here,

and in some cases different from that of the holotype (Poore

et al. 2011: pl. 16D).

Distribution. Eastern Australia (New South Wales and

Queensland), Vanuatu, Taiwan, 495—1802 m. Western Australia,

700—1170 m.

Munida babai Tirmizi and Javed, 1976

Munida babai Tirmizi and Javed, 1976: 81—85, fig. 1, 2 (type

locality: off Natal, South Africa). — Baba et al., 2008: 88. —

McEnnulty et al., 2011: app. 1.

Material examined. Western Australia: NMV J55044, 2 males (cl 7.7

mm, pcl 5.1 mm to cl 7.8 mm, pcl 5.3 mm), Ningaloo North,

21? 58.704' S, 113? 49.2' E to 21? 59.052' S, 113? 492' E, 170—177 m,

$510/2005/152, 10 December 2005; NMV J55041, 2 males (cl 7.8 mm,

pel 5.3 mm; cl 9.2 mm, pcl 5.9 mm), Ningaloo North, 21° 59.172' S,

113? 49.2' E to 21° 59.79' S, 113? 49.14' E, 165—166 m, SS10/2005/153,

Australian Munida

11 December 2005; NMV J55042, 3 females (cl 6.0 mm, pcl 3.8 mm;

cl 6.1 mm, pcl 4.0 mm), 4 males (cl 7.7 mm, pcl 5.0 mm; cl 8.1 mm, pcl

5.7 mm), Barrow Island, 20° 59.082' S, 114° 54.42' E to 20° 59.67' S,

114° 54.54' E, 100-101 m, SS10/2005/170, 13 December 2005.

Genetic data. CO1; see Table 1.

Remarks. Originally described from off South Africa (off Natal),

M. babai has subsequently been reported from shelf depths

across the Indian Ocean as far east as Hong Kong (Baba, 1988).

It can be distinguished by the presence of spines on the anterior

margin of abdominal somite 4 and very short supraocular spines.

As noted by Baba (1988), the number of spines on the abdominal

somites is variable. We found abdominal somite 2 with 5—9

spines, somite 3 with 2—5, and somite 4 with 2—5 spines. The P2

dactylus is 0.75 the length of the propodus in Australian

specimens compared to 0.64 for the type (based on Tirmizi and

Javed, 1976: fig. 2D), and 0.84 for the Albatross material from

Hong Kong and the Phillipines (Baba, 1988). This is the first

record of the species from Australia.

Distribution. South Africa, Madagascar, Hong Kong, Malaysia,

70—456 m. Western Australia, 100—177 m.

Munida benguela de Saint Laurent and Macpherson, 1988

Munida benguela de Saint Laurent and Macpherson, 1988: 106,

figs 1, 2a, 2c, 3a, 3d, 3f-1 (type locality: south of Namibia). — Baba et

al., 2008: 89.

Material examined. Western Australia: NMV 3756405, 1 male

(damaged; cl 11.6 mm, pcl 8.2 mm), SSO5/2007/176, Kulumburu L29

transect, 13? 13.482' S 123? 23.742' E to 13? 13.338' S 123? 23.286' E,

392 m, 5 July 2007.

Genetic data. Not available.

Remarks. Although this specimen is damaged, most diagnostic

characters are evident. In M. benguela, the P2 dactylus has 6—8

spines with the ultimate being distant from the unguis. Similarly,

our specimen has 8 spines and is unarmed on the distal one-

third. In M. benguela, the fixed finger of the chela has at least 4

spines on the lateral margin which agrees with our specimen.

However, our specimen lacks any spines on the mesial margin

of the moveable finger while M. benguela has a subdistal spine

and an additional 2—4 spines on the mesial margin.

Distribution. South African coast between S Namibia and

Natal, and Madagascar; 352-1000 m. North-west Australia,

302 m.

Munida compacta Macpherson, 1997

Figure 17A-C

Munida compacta Macpherson, 1997: 605—606, fig. 2 (type

locality: Kei Islands, Indonesia). — Baba et al., 2008: 90.

Munida andamanica. — Poore et al., 2008: 19. — McEnnulty et

al., 2011: app. 1,2.

Material examined. Queensland. AM P89035, 2 ovigerous females (pcl

11.7-13.5 mm), east of Cape York, 10° 29.810' S, 144° 01.380' E, 596 m,

RV Franklin, FRO688-02, 20 August 1988. Western Australia. AM

P72138, 1 ovigerous female (cl 26.5 mm, pcl 19.7 mm), 240 km north-

west of Port Hedland, 18° 06' S, 117? 45' E, 500 m, RV Soela, SO2/82/31,

139

coll. J. Paxton, 7 April 1982; NMV J56402, 1 male (cl 14 mm, pcl 7.0

mm), Onslow L19 transect, 20° 07.962' S, 114° 58.71' E to 20° 07.584' S,

114° 58.416' E, 415—470 m, S805/2007/15, 11 June 2007; NMV J56409,

4 females (cl 18.8 mm, pcl 11.6 mm; cl 26.5 mm, pcl 18.0 mm), Dampier

L20 transect, 19° 43.548' S, 115° 20.604' E to 19? 43.764' S,

115? 21.144' E, 415—428 m, S805/2007/28, 12 June 2007; NMV J56414,

] male (cl 14.5 mm, pcl 7.5 mm), SSO5/2007/28; NMV J56408, 1 male

(cl 17.9 mm, pcl 11.3 mm), Hedland L22 transect, 18° 34.194' S,

117? 27.864 E to 18? 34.062' S, 117° 28.626' E, 401—405 m,

SS05/2007/52, 14 June 2007; NMV J55266, 1 female (cl 13.5 mm, pcl

5.5 mm), Imperieuse L23 transect, 17° 21.492' S, 118° 57.312' E to

17? 20.88' S, 118° 56.802' E, 437—446 m, SS05/2007/56, 15 June 2007;

NMV J56412, 2 males (cl 24.7 mm, pcl 16.5 mm; cl 25.0 mm, pcl 16.6

mm), SSO5/2007/56; NMV J56411,3 males (cl 15.7 mm, pcl 9.5 mm to

cl 18.0 mm, pcl 11.2 mm), | ovigerous female (cl 27.3 mm, pcl 18.1

mm), Imperieuse L23 transect, 17° 31.734 S, 118° 50.6l' E to

17? 32.508' S, 118? 50.352' E, 403-407 m, SS05/2007/57, 15 June 2007;

NMV J56426, | female (cl 14.4 mm, pcl 7.6 mm), 5805/2007/57; NMV

J55260, 3 ovigerous females (cl 23.9 mm, pcl 16.0 mm to cl 25.9 mm,

pel 24.0 mm), Imperieuse L23 west transect, 17° 31.044' S,

118° 51.162' Eto 17? 31.716' S, 118° 50742 E,405—406 m, SS05/2007/64,

16 June 2007; NMV J55265, 1 male (cl 12.4 mm, pcl 7.5 mm),

$505/2007/64; NMV J55990, 1 female (18.9 mm, pcl 13.4 mm), 1

Juvenile (cl 9.5 mm, pcl 8.4 mm), SSO05/2007/64; NMV J56406, 1 male

(cl 11.5, pcl 10.5 mm), SSO5/2007/64; NMV J56410, 1 ovigerous female

(cl 24.5 mm, pcl 16.7 mm), Leveque L27 transect, 14° 50.814' S,

121° 26.436' E to 14° 5094' S, 121° 28.164' E, 382-401 m,

3305/2007/102, 26 June 2007; NMV J56080, 8 males (cl 19.9 mm, pcl

14.7 mm to cl 26.7 mm, pcl 16.9 mm), 3 females (cl 21.9 mm, pcl 14.8

mm to cl 25.7 mm, pcl 18.3 mm), Leveque L27 transect, 14° 51.198' S,

121° 25.878 E to 14° 50724' S, 121° 27.012" E, 396-403 m,

SS05/2007/144, 2 July 2007; NMV J56081, 2 females (cl 20.6 mm, pcl

12.6 mm; cl 22.4 mm, pcl 15.1 mm), | juvenile (cl 14.9 mm, pcl 9.2

mm), Ashmore L30 transect, 12° 28.884' S, 123° 25.062' E to

12? 29.958' S, 123? 25.002' E, 397—405 m, SS05/2007/189, 6 July 2007;

NMV J55994, 4 ovigerous females (cl 22.4 mm, pcl 15.1 mm to cl 23.5

mm, pcl 15.6 mm), | female (cl 24.1 mm, pcl 16.3 mm), 3 males (cl 19.6

mm, pcl 12.9 mm to cl 24.4 mm, pcl 16.3 mm), Ashmore L30 transect,

12? 31.77' S, 123? 25.638' E to 12? 30.828' S, 123? 25.362' E, 401—404

m, SS05/2007/192, 6 July 2007; NMV J55051, 1 female (cl 14.4, pcl 9.0

mm), Barrow Island, 21° 00.402' S, 114° 22.86' E to 21° 00.042' S,

114? 22.5' E, 399-408 m, SS10/2005/172, 13 December 2005.

Colour. Carapace and abdominal somites 2—4 pale orange/

pink. Rostrum red, supraocular spines white or red and white

on distal half. Pl pale orange/pink with white fingers, P2—4

pale orange on meri and carpi, white on propodi, dactyli white

with red tips.

Genetic data. COI and 16S; see Table 1.

Remarks. Munida compacta 1s very close to M. rubridigitalis

Baba, 1994, and M. julumunyju sp. nov. from eastern and

western Australia, respectively, and M. rhodonia, from New

Caledonia. These species can be distinguished by:

- the dactylus of P2 is similar in length (0.8—1.0) to the

propodus in M. compacta and distinctly shorter (0.8) in

M. rhodonia

- chelipeds with well-developed spines on the dorsal face

of the palm in M. compacta, which are instead very small in

M. rhodonia.

While the length of the distomesial spine of the Pl merus

was used to differentiate these species (Baba, 1997), we found

140 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 17. A-C, Munida compacta Macpherson, 1997, female, dorsal view and lateral (NMV J56409), female 18.9 mm, pcl 13.4 mm (NMV

J55990); D, Munida endeavourae Poore and Ahyong, 2004, ovigerous female, cl 21.7 mm, pcl 13.9 mm (NMV J55052); E, Munida gordoae

Macpherson, 1994 (NMV J56420). Photos: K. Gowlett-Holmes, CSIRO.

Australian Munida

that that the length of this spine varied in our specimens of M.

compacta and did not always overreach the midlength of the

carpus. Also, the P2 dactylus 1n M. compacta 1s described as

clearly shorter than the propodus, but in our specimens, the

P2 dactylus was 0.8—1.0 times the propodus length.

The sequences of M. compacta from north-western

Australia and M. rhodonia from New Caledonia were 1.2%

divergent for COI and 0.2% divergent for 165.

The present specimens represent the first confirmed

records of M. compacta from Australia.

Distribution. Indonesia, Kei Islands, 246—694 m. Queensland

and Western Australia, 397—470 m.

Munida compressa Baba, 1988

Munida compressa Baba, 1988: 91, figs 33, 34 (type locality:

Moluccas, Indonesia). — Baba et al., 2008: 91. — Baba et al., 2009:

152, figs 130—134. — Poore et al., 2011: pl. 16H (colour).

Material examined. Western Australia: NMV J55980, 1 male (cl 21.2

mm, pcl 13.0 mm), Kulumburu L29 transect, 13° 13.482' S,

123? 23.742' E to 13° 13.338' S, 123° 23.286' E, 392 m, SS05/2007/176,

5 July 2007; NMV J55981, 4 males (cl 12.0 mm, pcl 8.2 mm; cl

18.6 mm, pcl 12.1 mm), Ashmore L30 transect, 12° 28.884' S,

123? 25.062" E to 12? 29958' S, 123? 25.002' E, 397-405 m,

SS05/2007/189, 6 July 2007.

Genetic data. 16S and COI; see Table 1.

Remarks. This 1s the first record of the Munida compressa

from Australia. Sequences from our material and a specimen

from the Solomon Islands (Machordom and Macpherson,

2004) were 0.2% divergent for COI and invariant for 165.

Distribution. Indonesia, Japan, Taiwan, Arafura Sea, 180—668

m. North-west Australia, 392—405 m.

Munida distiza Macpherson, 1994

Munida distiza Macpherson, 1994: 459, figs 14, 68, 69 (type

locality: New Caledonia). — Baba et al., 2008: 93. — Macpherson,

2013: 300.

Material examined. NMV J56487, 1 ovigerous female (cl 20.3 mm,

pel 12.6 mm), 1 male (cl 18.7 mm, pcl 12.4 mm), Barrow L1 transect,

21° 00.816' S, 114° 39.15' E to 21° 00.78' S, 114° 38.898' E, 258—271 m,

SS05/2007/11, 10 June 2007.

Genetic data. COI and 16S; see Table 1.

Remarks. COI sequences from this material were 4-5%

divergent from sequences of Munida distiza Macpherson,

1994, from New Caledonia. However, we could not find any

distinguishing morphological characters.

Distribution. Philippines, New Caledonia, Loyalty Islands,

Matthew and Hunter Islands, French Polynesia, 150—540 m.

North-west Australia, 258—271] m.

Munida endeavourae Ahyong and Poore, 2004

Figure 17D

Munida endeavourae Ahyong and Poore, 2004: 26, fig. 5 (type

locality: south-east of Green Cape, New South Wales, Australia). —

141

Baba et al., 2008: 94. — Yaldwyn and Webber, 2011: 211. — Farrelly

and Ahyong, 2019: 13,54. — Yan et al. 2020: 2, tab. 1.

Munida grievae Vereshchaka, 2005: 140, fig. 3A—F (type locality:

Bay of Plenty, New Zealand).

Munida sp. MoV 5199. — Poore et al., 2008: 21, unnumbered

figure (lower right). —McEnnulty et al., 2011, app. 1, 2.

Material examined. Western Australia: NMV J55052, 1 ovigerous

female (cl 21.7 mm, pcl 13.9 mm), Albany, 35° 26.046' S, 118° 21.06' E

to 35? 26.19' S, 118? 20.64' E, 912—922 m, SS10/2005/28, 23 November

2005. Tasmania: MV J59312, 1 specimen, Cascade Plateau,

43° 49/706' S, 150° 29.999' E, 1061 m, TT 01/2008/J2-390-015, 4

January 2009; MV J59310, 1 specimen, Hill off St. Helens,

41° 14.349' S, 148° 49.293' E, 1309 m, TT 01/2008/J2-389-006, 1

January 2009.

Colour. Carapace, abdomen, chela and P2—4 pink. Eggs red.

Genetic data. COI and 16S; see Table 1.

Remarks. Poore et al. (2008) and McEnnulty et al. (2011)

recorded M. endeavourae from south-western Australia as

"Munida sp. MoV 5199”; the present record is the first

confirmation of the species from the area.

Distribution. South-eastern Australia from south-east of Green

Cape to Tasmania, 620—1700 m. South-western Australia, 912—

022? m

Munida gordoae Macpherson, 1994

Figure. I7E

Munida gordoae Macpherson, 1994: 469, fig. 18 (type locality:

Chesterfield Islands). — Baba et al., 2008: 97.

Munida aff. volantis. — Poore et al., 2008, 20, unnumbered fig.

— McEnnulty et al., 2011: 26, app. 1,2.

Material examined. Western Australia: NMV J56420, 3 ovigerous

females (cl 5.7 mm, pcl 3.5 mm to cl 8.2 mm, pcl 5.3 mm), 8 males (cl 5.1

mm, pcl 2.9 mm to cl 8.1 mm, pcl 6.1 mm), 2 juveniles (cl 4.8 mm, pel 3.1

mm; cl 5.1 mm, pcl 3.2 mm), Imperieuse L23 east transect, 17° 35.706' S,

118° 58.902' E to 17° 35.346' S, 118? 58.794' E, 140—108 m, SS05/2007/62,

16 June 2007; NMV J55087, 3 males (cl 10.0 mm, pcl 6.3 mm to cl 11.2

mm, pcl 7.2 mm), Two Rocks, 31° 37.452' S, 115° 00.24' E to 31° 37.71' S,

115° 00.18' E, 205-210 m, SS10/2005/11, 19 November 2005; NMV

J55085, 1 male (cl 12.0 mm, pcl 6.3 mm), Perth Canyon, 31^ 55.26' S,

115° 12.12' E to 31° 55.446' S, 115? 11.76' E, 194—232 m, SS10/2005/69,

29 November 2005; NMV J55088, | female (cl 11.0 mm, pcl 6.3 mm),

Abrolhos, 28° 59.328' S, 113? 47.04' E to 28° 59:784' S, 113° 47.28' E,

180—183 m, SS10/2005/91, 3 December 2005; NMV J55086, 5 ovigerous

females (cl 7.3 mm, pcl 4.8 mm to cl 11.2 mm, pcl 7.4 mm), 1 damaged

male, Kalbarri, 27? 55.716' S, 113? 08.28' E to 27° 56.022' S, 113° 08.64" E,

252—253 m, SS10/2005/99, 4 December 2005.

Colour. Carapace and abdominal somite 2—3 pink and orange

patterning. Abdominal somite 4—6 with white markings.

Rostrum and supraocular spines orange/pink. Chela and P2-4

orange/red with white bands.

Genetic data. COI and 16S.

Remarks. These are the first records of M. gordoae from

Australia. Sequence divergence between our north-western

Australian specimens of M. gordoae and those from New

Caledonia was only 0.2% for COI and 0.4% for 16S.

142

Distribution. New Caledonia, Loyalty Islands, Matthew and

Hunter Islands, and Chesterfield Islands, Vanuatu, Fiji, 850—500

m. Western Australia, 180—232 m.

Munida gracilis Henderson, 1885

Figure 18

Munida gracilis Henderson, 1885: 412 (type locality: New

Zealand, 38° 50' S, 169? 20' E). — Baba et al., 2008: 97. — Yaldwyn

and Webber, 2011: 212. — Yan et al. 2020: 2, tab. 1.

Munida chydaea Ahyong and Poore, 2004: 24, fig. 4 (type locality:

east of Brush Island, New South Wales, Australia). — Poore, 2004:

234, fig. 64b. — Baba, 2005: 260. Syn. nov.

Munida disgrega Baba, 2005: 103, figs 40, 41 (east of Victoria,

Australia). — Poore et al., 2008: 19. — McEnnulty et al., 2011: app. 1.

Syn. nov.

Material examined. South Australia: SAM C7658, 1 ovigerous female

(cl 32.3 mm, pcl 18.5 mm), | damaged specimen (cl 18.5 mm, pcl 10.6

mm), Great Australian Bight, 33° 27.09' S, 130° 41.19' E, 500 m, BPZ

2010/500, 2010.

Western Australia: NMV J55107, 2 females (cl 24.8 mm, pcl 13.7

mm; cl 26.1 mm, pcl 15.5 mm), 3 males (cl 20.6 mm, pcl 11.2 mm to cl

21.9 mm, pcl 12.1 mm), Bald Island, 35° 14.022' S, 118° 39.84' E to

35° 13.674' S, 118° 40.5' E, 710—728 m, SS10/2005/32, 2005; NMV

J55106, 6 females (cl 11.4 mm, pcl 6.3 mm to cl 25.4 mm, pcl 14.3

mm), 10 males (cl 12.4 mm, pcl 7.4 mm to cl 24.0 mm, pcl 13.6 mm),

Bald Island, 35? 14.022' S, 118? 39.84' E to 35? 13.674' S, 118? 40.5' E,

710—728 m, SS10/2005/32, 23 November 2005; NMV J55083, 3 males

(cl 27.9 mm, pcl 19.5 mm to cl 47.8 mm, pcl 34.9 mmy), Pt Hillier,

35° 22.902' S, 117° 12.42' E to 35° 22.902' S, 117° 12.42' E, 539 m,

SS10/2005/19, 2005; NMV 755080, 1 female (pcl 7.9 mm), Albany,

35? 22.29' S, 118? 19.98' E to 35? 22.644' S, 118? 19.26' E, 685—695 m,

A.W. McCallum, S.T. Ahyong & N. Andreakis

SS10/2005/27, 2005; NMV J55084, 16 females (cl 20.9 mm, pcl 11.9

mm to cl 36.6 mm, pcl 19.6 mm), 19 males (cl 14.3 mm, pcl 9.0 mm to

cl 30.2 mm, pcl 17.9 mm), Albany, 35° 22.14' S, 118° 20.1' E to

35? 22.518' S, 118? 19.32' E, 676—680 m, SS10/2005/29, 2005; NMV

J55082, 1 female (cl 9.1 mm, pcl 7.0 mmy), 2 males (cl 17.4 mm, pcl 7.8

mm; cl 15.1 mm, pcl 9.9 mm), Bald Island, 35° 12.81' S, 118° 39.06' E

to 35? 12.24' S, 118° 40.14' E, 408—431 m, SS10/2005/34, 2005.

New Zealand: NIWA 53772, | female (cl 23.3 mm, pcl 15.8 mm),

44? 09.47' S, 174? 33.32' W to 44? 09.48' S, 174? 33.56' W, 520—650 m,

TANO905/103, 26 June 2009; NIWA 54087, 1 ovigerous female (cl

21.0 mm, pcl 12.2 mm), Diamondhead Peak B, Andes Seamounts,

44° 08.97' S, 174° 45.41' W to 44? 09.02' S, 174? 45.63' W, 519—609 m,

TANO905/113, 27 June 2009; NIWA 54108, 1 female (cl 24.4 mm, pcl

13.9 mm), Diamondhead Peak B, Andes Seamounts, 44° 08.99' S,

174? 46.09' W to 44? 09.01' S, 174° 46.3' W, 830—900 m, TAN0905/114,

27 June 2009; NIWA 63655, 4 males (pcl 5.4-7.7 mm), Site 3a

seamount, Hikurangi Margin, 41° 19.16'S, 176° 11.84' Eto 41° 19.28'S,

176° 11.53' E, 495 m, TAN1004/66, 21 April 2010.

Genetic data. CO1 and 16S; see Table 1.

Remarks. Munida gracilis 1s a common species in New

Zealand, usually from depths of 300—600 m with a recorded

range of about 100-1200 m (Yan et al. 2020), that is

morphologically closest to the Australian Munida chydaea

Ahyong and Poore, 2004. COI sequences of specimens

corresponding to Munida chydaea Ahyong and Poore, 2004,

from south-western Australia, however, were only 0.9-1.0%

divergent from M. gracilis from New Zealand and 0.2%

divergent for 165. This low level of sequence divergence led us

to further examine the morphological distinctions between M.

gracilis and M. chydaea. Ahyong and Poore (2004)

distinguished the two species based on the number of anterior

(FA 0 Xu

| s RN

South-west Australia

Great Australian Bight

Tasmania

New Zealand

Figure 18. Munida gracilis Henderson, 1885. Variation of P2 dactyli. A. female cl 32 mm, 18.3 mm; B. male cl 26.2 mm, pcl 15.5 mm (NMV

J55048); C, ovigerous female, cl 32.3 mm, pcl 18.5 mm (SAM C7658); D, male cl 21.2 mm, pcl 11.9 mm (NMV J53574); E, female, cl 30.0 mm

(J53574); F, female cl 23.5 mm, pcl 15.8 mm (NIWA 53772); G, ovigerous female, cl 21.0 mm, pcl 12.2 mm (NIWA 54087).

Australian Munida

spines on abdominal somite 3 (1 or 2 spines in M. chydaea; 4

spines in M. gracilis), the length of the unarmed flexor margin

of the P2 dactyli (distal one-quarter in M. chydaea versus

distal one-third in M. gracilis), spinules on the outer margin of

antennal articles 3 and 4 in M. chydaea (absent in M. gracilis)

and more numerous transverse abdominal striae in M.

chydaea. Our comparisons between new Australian and New

Zealand specimens, however, failed to identify consistent

distinguishing features. Although the many specimens

examined by Ahyong and Poore (2004), primarily from from

eastern Australia, were consistent in the aforementioned

features, Specimens examined herein from southern Australia

(Tasmania, South Australia and Western Australia) and New

Zealand exhibit morphological overlap. Thus, the shape and

armature of the P2 dactyli in southern Australian specimens is

variable, with the flexor margin unarmed on the distal one-

fifth to one-third (fig. 21 A—E). Similar variation was seen in

the eight New Zealand specimens we examined; in most

specimens, the distal one-third of the P2 dactylus was unarmed

(fig. 21F) but in one ovigerous female (pcl 12.2 mm; NIWA

54087; Fig. 21G), the distal one-quarter was unarmed.

similarly, abdominal somite 3 1s variously armed with 2—4

spines, albeit usually 4 in adult New Zealand specimens, and

almost always 2 in adult southern Australian specimens.

Another character used to distinguish M. chydaea was the

presence of outer spinules on the third and fourth antennal

article, but such spinules were present on all eight of the

specimens of M. gracilis we examined from New Zealand.

Thus, considered as a whole, Australian material is not reliably

morphologically separable from New Zealand specimens, and

given the minimal divergence between New Zealand and

Western Australian sequences, we consider both populations

as probably conspecific.

This brings Munida disgrega Baba, 2005, to our attention,

which was described from 4 small specimens (cl 10.1—13.0

mm) collected from south-eastern Australia and noted as very

close to M. chydaea and M. gracilis. In M. disgrega, the flexor

margin of the dactylus 1s unarmed on the distal one-quarter to

one-fifth, which is encompassed by the newly observed

variation in M. gracilis and M. chydaea. Other distinguishing

features of M. disgrega, such as the length of the chela, the

lesser striation and broader anterior margin of the thoracic

sternum, and proportionately wider eyes apppear to be

allometric differences given the small size of the type

specimens. Consequently, we cannot reliably distinguish M.

disgrega from M. gracilis or M. chydaea

Although we do not have molecular data from eastern

Australia (type region of M. chydaea), on the basis of the low

molecular divergence (between New Zealand and south-

western Australian specimens) and morphological variability

within southern Australian specimens that encompasses those

from New Zealand and eastern Australia, we regard M.

chydaea and M. disgrega as junior synonyms of M. gracilis.

Munida gracilis 1s closely related to, but clearly distinct from,

M. haswelli, with 7.6% divergence in COI and 3.3% in 165.

Distribution. Southern Australia (Sydney to south-western

Australia), 146—700 m. New Zealand, 365—610 m.

143

Munida haswelli Henderson, 1885

Figure 19A

Munida haswelli Henderson, 1885: 411 (type locality: off southern

New South Wales, Australia). — Baba et al., 2008: 98. — McEnnulty

et al., 2011: app. 1, 2. — Farrelly and Ahyong, 2019: 13, 54, fig. 96.

Material examined. Western Australia: NMV J60017, 2 males (cl 19.5

mm, pcl 12.3 mm; cl 19.6 mm, pcl 12.5 mm), Ashmore L30 transect,

12? 26.058' S, 123? 36.078' E, 125 m, 8805/2007/186, 6 June 2007;

NMV 3755102, 1 male (cl 12.1 mm, pcl 7.4 mm), Two Rocks,

31° 37.452' S, 115? 00.24' E to 31? 37.71' S, 115? 00.18' E, 205—210 m,

$S10/2005/11, 19 November 2005; NMV J55103, 1 male (cl 19.6 mm,

pel 12.8 mm), SS10/2005/11; NMV J55104, 3 ovigerous females (cl

14.0 mm, pcl 9.0 mm to cl 15.4 mm, pcl 10.2 mm), 2 males (cl 11.4 mm,

pel 7.2 mm; cl 16.4 mm, pcl 10.4 mm), Bald Island, 35° 11.442' S,

118° 38.7' E to 35? 11.25' S, 118° 39' E, 147—157 m, SS10/2005/35, 24

November 2005; NMV J55105, 5 ovigerous females (cl 11.5 mm, pcl

7.5 mm to cl 15.2 mm, pcl 8.9 mm), 5 males (cl 12.3 mm, pcl 7.9 mm

to cl 16.4 mm, pcl 11.3 mm), Bald Island, 35° 11.406' S, 118° 38.94' E.

to 35? 11.514' S, 118? 38.76' E, 161-169 m, SS10/2005/38, 24 November

2005; NMV J55108, 4 ovigerous females (cl 14.4 mm, pcl 9.3 mm to cl

16.0 mm, pcl 10.8 mm), 9 females (cl 11.3 mm, pcl 7.7 mm to cl 16.1

mm, pcl 11.2 mm), 18 males (cl 13.6 mm, pcl 9.1 mm; cl 21.4 mm, pcl

14.5 mm), Albany, 35° 21.276' S, 118° 17.94' E to 35° 21.348' S,

118° 17776' E, 179 m, SS10/2005/47, 25 November 2005; NMV J55109,

4 females (largest with broken rostrum; pcl 6.2-9.5 mm), 3 males (cl

11.9 mm, pcl 7.6 mm; cl 13.9 mm, pcl 9.0 mm), Pointt Hillier,

35? 22 A1' S, 117? 11.82' E to 35? 22.404' S, 117? 12.24' E, 195-196 m,

SS10/2005/57, 27 November 2005; NMV J55099, 3 ovigerous females

(cl 11.2 mm, pcl 7.5 mm to cl 11.5 mm, pcl 7.7 mm), 11 males (one with

rhizocephalan externa; cl 8.6 mm, pcl 6.1 mm to cl 11.8 mm, pcl 8.7

mm), Ningaloo North, 21° 59.172' S, 113° 49.2' E to 21° 59.79" S,

113° 49.14' E, 165-166 m, SS10/2005/153, 11 December 2005; NMV

J55101, 1 ovigerous female (cl 10.4 mm, pcl 7.2 mm), Ningaloo North,

21° 56.628' S, 113° 50.46' E to 21? 57.09' S, 113° 50.28' E, 132-134 m,

SS10/2005/163, 12 December 2005.

Colour. Carapace and somites 2—4 pale orange, somites 5—6

and telson white. Rostrum and supraocular spines orange.

Chela orange/pink with red bands on distal half of palm. P2—4

whitish pink with red bands on middle of propodus.

Genetic data. COI and 16S; see Table 1.

Remarks. Munida haswelli and Munida gracilis are closely

related, with 7.6% divergence in COI and 3.3% divergence in

165.

Distribution. Central New South Wales, south to Victoria,

lasmania, South Australia, the Great Australian Bight and

Western Australia, 121—329 m.

Munida heteracantha Ortmann, 1892

Munida heteracantha Ortmann, 1892: 255, pl. 11 fig. 12, 121, 12k

(type locality: Sagami Bay, Japan). — Baba et al., 2008: 99. — Poore

et al., 2008: 19. — McEnnulty et al., 2011: app. 1. — Castro, 2011: 16

Munida sp. MoV 5214. — Poore et al., 2008: 21.

Material examined. Western Australia: NMV J55037, ovigerous

female (cl 9.4 mm, pcl 6.35 mm), | ovigerous female (cl 9.5 mm, pcl

7.2 mm), 4 females (cl 8.3 mm, pcl 5.3 mm to cl 9.3 mm, pcl 6.6 mmy),

4 males (cl 6.9 mm, pcl 4.4 mm to cl 10.5 mm, pcl 6.7 mm), 3 juveniles

(cl 6.1 mm, pcl 4.1 mm to cl 6.6 mm, pcl 4.5 mm), Ningaloo South,

144 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 19. A, Munida haswelli Henderson, 1885, male cl 19.55 mm (NMV J55103), pcl 12.81 mm; B-D, Munida leagora Macpherson, 1994, |

Juvenile cl 12.2 mm, pcl 7.5 mm (NMV J56018), undetermined specimen (NMV J55999; NMV J56004). Photos: K. Gowlett-Holmes.

Australian Munida

22° 04.77' S, 113° 47.76' E to 22° 05.238' S, 113° 47.64' E, 201-206 m,

SS10/2005/146, 10 December 2005; NMV J55038, 1 male (cl 10.7, pcl

7.4 mm), 1 female (cl 7.5 mm, pcl 5.0 mm), 5510/2005/146; NMV

J55040, 1 male (cl 10.5 mm, pcl 7.0 mm), | ovigerous female (cl 9.9

mm, pcl 7.1 mm), 1 female (cl 7.6 mm, pcl 5.2 mm), Ningaloo North,

21° 58.704'S 113° 49.2 'E to 21° 59.052'S 113° 49.2'E, 170—177 m,

S$$10/2005/152, 10 December 2005.

Genetic data. Not available.

Remarks. Munida heteracantha Ortmann, 1892 has been

recorded widely across the western Pacific, including from

Queensland, Australia. Our specimens show some variation

from the lectotype from Japan described by Macpherson and

Baba (1995). The lectotype has a pair of small parahepatic

spines, which are absent in all our material. In most of our

specimens the distolateral spine of the basal antennular article

is slightly longer than the distomesial spine, while in the

lectotype the distal spines are subequal. Macpherson and Baba

(1993) noted that some specimens of M. heteracantha have 2

small median spines on abdominal somite 3, and half of the

specimens here have 1—6 spines. Only three specimens had the

P2 attached, and these show the dactyli with 7—8 spines on the

flexor margin, compared to only 5 spines in the lectotype.

Munida heteracantha 1s also similar to Munida macphersoni

Cabezas, Lin and Chan, 2011, but differs in the length of the

supraocular spines, which are very short in M. heteracantha

(0.34 x rostral length compared to 0.5 in M. macphersoni).

Distribution. Japan, Philippines, and Indonesia, off Central

Queensland, New Caledonia, Fiji, 100—322 m. South-western

Australia, 170—177 m.

Munida leagora Macpherson, 1994.

Figure I9B-D

Munida leagora Macpherson, 1994: 485, figs 26,76 (type locality:

New Caledonia). — Baba et al., 2008: 104.

Material examined. Western Australia: NMV J56004, 1 ovigerous

female (cl 13.8 mm, pcl 8.1 mm), | male (cl 13.0 mm, pcl 8.0 mm),

Leveque L27, 14° 58.362' S, 121° 38.556' E to 14° 57.762' S,

121° 39.264' E, 228—232 m, 5805/2007/143, 2 July 2007; NMV J55999,

5 males (cl 15.7 mm, pcl 9.5 mm to cl 20.2 mm, pcl 12.3 mm), 1 male

with bopyrid (cl 14.9 mm, pcl 8.6 mm), | ovigerous female (cl 16.5

mm, pcl 10.2 mm), 8805/2007/143; NMV J56018, 1 juvenile (cl 12.2

mm, pcl 7.5 mm), Mermaid L24 east transect, 17^ 02.838' S,

119° 39.684' E to 17° 03.708' S, 119° 41.358' E, 424—456 m,

SS05/2007/77, 18 June 2007.

Colour. Carapace and abdominal somites 2—4 pale orange,

somites 5—6 and telson white. Rostrum and supraocular spines

pale orange. Pl and P2—4 meri with orange spots at base of

spines and on striae.

Genetic data. COI and 16S; see Table 1.

Remarks. Munida leagora was described from New Caledonia,

and the present specimens represent the first records of the

species from Australia. Our specimens show some minor

variation compared to the type description, including the

presence of | or 2 small spines on the lateral margins of the

anterior ridge of abdominal somite 2, and the presence of

145

parahepatic spines in some specimens. Munida leagora 1s

very similar to Munida alia Baba, 1994, described from a

single specimen in Queensland. We compared our specimens

of M. leagora to the holotype of M. alia and confirmed that

these species are distinct. In Baba's (2005) key to Munida,

these two species are separated by the length of the distomesial

spine of antennal article 1, which reaches the end of article 2

in M. alia and reaches or overreaches the distal end of article

3 in M. leagora. In addition, M. alia lacks a pair of postcervical

spines, which are always present in our specimens of

M. leagora.

Divergence in COI sequences between specimens of M.

leagora from north-western Australia and New Caledonia is

<0.5%. The wide transverse yellow and purple bands on the

carapace described for New Caledonian specimens are absent

in all the specimens examined here.

Distribution. New Caledonia, Loyalty Islands, Chesterfield

Islands, Vanuatu, Bayonnaise Bank, Fiji and Tonga, 240—610

m. North-west Australia, 232—450 m.

Munida magniantennulata Baba and Türkay, 1992

Figure 20A

Munida magniantennulata Baba and Türkay, 1992: 205, figs 2,3

(type locality: Lau Basin). — Baba et al., 2008: 105. — Farrelly and

Ahyong, 2019: 13,54. fig. 97.

Not Munida magniantennulata. — Baba, 1994a: 12 (2M. typhle

Macpherson, 1994)

Material examined. New South Wales: NMV J73001, 3 ovigerous

females (cl 10.6 mm, pcl 7.2 mm to cl 13.4 mm, pcl 12.5 mm), Central

Eastern CMR, 30° 05.862' S, 153° 53.922' E to 30° 07.158' S,

153° 52.47' E, 2429-2518 m, IN2017. v03/086, 5 June 2017.

Colour. Carapace, abdomen, cheliped and P2—4 pink. Eggs red.

Genetic data. Not available.

Remarks. The type specimens of M. magniantennulata are

described as having the P2 dactylus 0.5—0.6 times propodus

length, 6.6 times as long as high, and lined with 10 movable

spines. Our specimens have a very similar P2 dactylus, which

is 0.6—0.7 times the propodus length, 5—6 times as long as high,

with 9—10 movable spines on the flexor margin. The chelipeds

of all our specimens lack spines on the outer margin of the

fingers or on the dorsal face of the palm, which agrees with the

type material. All the specimens have two small tubercle-like

spines on abdominal somite 2.

These specimens differ from the holotype in the length of

the mesial spine on the first antennal article, which reaches

the midlength of article 2 in these specimens but is very short

in the holotype.

The specimens recorded by Baba (1994) as M.

magniantennulata from off central Queensland are referred

to Munida typhle Macpherson 1994 (see Remarks for that

species).

Distribution. Lau Basin, 1750—2003 m. Eastern Australia,

2429-2518 m.

146 A.W. McCallum, S.T. Ahyong & N. Andreakis

Figure 20. A, Munida militaris Henderson, 1885, ovigerous female cl 15.0 mm, pcl 9.4 mm, (NMV J55033); B, Munida magniantennulata Baba

and Türkay, 1992, ovigerous female (NMV J73001); C, Munida pectinata Machordom and Macpherson, 2005, male (NMV J55268); D. Munida

roshanei Tirmizi, 1966, male 11.1 mm, pcl 7.2 mm (NMV J55261). Photos: K. Gowlett-Holmes, CSIRO.

Australian Munida

Munida militaris Henderson, 1885

Figure 20B

Munida militaris Henderson, 1885: 410 (type locality: off Mataku,

Fiji Islands). — Baba, 2008: 108 (synonymies, type data). — Baba et

al., 2009: 173, figs 153—155 (in part) (Taiwan, 190—1183 m). — Poore

et al., 2011: pl. 17E (colour).

Not Munida militaris Baba et al., 2009: 173 (1n part) (Taiwan) (=

M. lanciaria Cabezas et al., 2011).

Munida aff. Rubiesi — Poore et al., 2008, 20 (part), unnumbered fig.

Munida MoV 5183. — McEnnulty et al., 2011: app. 1,2.

Material examined. Western Australia: NMV J55033, 1 ovigerous

female (cl 15.0 mm, pcl 9.4 mm), 31° 36.528' S 114° 58.86' E to

31° 37.038' S 114° 58.26' E, 329—370 m, SS10/2005/06, 19 November

2005; NMV J55035, 1 ovigerous female (cl 15.0 mm, pcl 10.2 mm),

near Abrolhos islands, 28° 29.37' S 113° 25.14' E to 28? 30.06' S

113? 25.5' E, 416—431 m, SS10/2005/95, 4 December 2005.

Genetic data. Not available.

Colour. Carapace and somites 2—4 orange and white, somites

5—6 and telson white. Rostrum and supraocular spines orange

on base and tips and white in the middle. Chela white and

orange, and orange/red bands on distal half of fingers. P2—4

transparent white/orange with orange/red across the middle of

propodus and tips of dactyli.

Remarks. The specimens here agree with the lectotype

description of M. militaris by Baba and Macpherson (1991).

Munida militaris closely resembles M. benguela de Saint

Laurent and Macpherson, 1985, but can be distinguished by the

following features:

- moveable finger of the cheliped bears a single basomesial

spine

- P2-4 dactyli with about 10 ventral spines, the untimate

at the base of the unguis.

Our specimens differ slightly from the illustrated

lectotype material from Fiji and Ambon. The lectotype has

the distomesial spine of antenna article 2 described as

occasionally overreaching the end of the peduncle, while in

both our specimens, the distomesial spine distinctly

overreaches the distal end of the peduncle.

Distribution. Taiwan, Indonesia, Queensland, New Caledonia,

Vanuatu, Wallis and Futuna, Fiji, T-onga, 190-1183 m. South-

western Australia, 329—431 m.

Munida pectinata Machordom and Macpherson, 2005

Figure 20C

Munida pectinata Machordom and Macpherson, 2005: 828, fig. 3

(type locality: New Caledonia). — Baba, 2008: 112. — Komai, 2011:

359, fig. 10, 11D (colour) (Japan 96-221 m). — Komai, 2012: 38 (Izu

islands, 174—192 m).

Material examined. Western Australia: NMV 755268, 4 males (2

damaged; cl 10.8 mm, pcl 6.5 mm; cl 18.4 mm, pcl 10.7 mm),

17° 35.706' S, 118? 58.902' E to 17? 35.346' S, 118? 58.794' E, 108—140

m, SS05/2007 stn 62, 16 June 2007.

Colour. Carapace and somites 2—4 pale orange with dark

orange markings, somites 5—6 and telson pale orange. Rostrum

147

and supraocular spines orange. Chela merus pink with red

bands, remaining chela mottled orange. P2—4 white with

orange/red bands.

Genetic data. COI and 16S; see Table 1.

Remarks. These Australian specimens of M. pectinata agree

well with illustrations of type material from New Caledonia,

extending the range to eastern Indian Ocean. The north-western

Australian sequences of COI (639 bp) and 165 were identical to

two sequences from New Caledonia. This is the first record of

M. pectinata from Australia.

Distribution. New Caledonia, Japan and north-western

Australia, 96—240 m.

Munida pherusa Macpherson and Baba, 1993

Munida pherusa Macpherson and Baba, 1993: 408, fig. 15 (type,

locality: Philippines). — Baba et al., 2008: 112. — Baba et al., 2009:

178, figs 158—159. — Kim et al., 2016: 9, fig. 9 (Korea, 112—117 m).

Material examined. Western Australia: NMV J55097, ] specimen,

Ningaloo South, 22° 04.002' S, 113° 48.66' E to 22° 04.248' S,

113? 48.9' E, 101-106 m, SS10/2005/144, 10 December 2005; NMV

J13307, 9 ovigerous females (cl 7.7 mm, pcl 4.5 mm to cl 11.4 mm, pcl

7.3 mm), 2 females (cl 8.3 mm, pcl 4.9 mm; cl 9.1 mm, pcl 7.3 mm), 7

males (cl 7.3 mm, pcl 4.9 mm to cl 12.7 mm, pcl 7.7 mm), North-west

Shelf, between Port Hedland and Dampier, 18° 50' S, 117° 39' E to

18? 50' S, 117? 39' E, 175 m, NWA stn 29, 6 June 1983.

Remarks. In addition to one specimen of M. pherusa collected

in 2005 in north-western Australia, another 17 specimens are

reported here from a survey of the North-West Shelf in 1983.

The specimens show that the length of the disto-mesial spine

on antennal article 2 1s slightly variable in length, sometimes

just reaching the end of the peduncle and in some specimens,

including the holotype, it slightly overreaches the peduncle. In

the illustrated holotype, the cheliped finger is distinctly shorter

that the palm, while in all of our specimens, the cheliped finger

and palm are subequal in length.

These are the first records of M. pherusa from Australia.

Distribution. Japan, Philippines, Indonesia, Korea, 73—167 m.

North-western Australia, 101—178 m.

Munida philippinensis Macpherson and Baba, 1993

Munida philippinensis Macpherson and Baba, 1993: 410, fig. 16

(type locality: Philippines). — Baba et al., 2008: 112.

Material examined. Western Australia: NMV J56003, 4 females (cl

7.9 mm, pcl 4.8 mm to cl 10.1 mm, pcl 9.0 mm), 12 males (cl 7.5 mm,

pel 4.8 mm to cl 11.2 mm, pcl 7.4 mm), Leveque L27, 14? 58.362' S,

121° 38.556' E to 14° 57.762' S, 121° 39.264' E, 228—232 m,

SS05/2007/143, 2 July 2007.

Genetic data. COI and 16S; see Table 1.

Remarks. This 1s the first record of the species from Australia.

Distribution. Philippines, Indonesia, 146—297 m. North-west

Australia, 228—222 m.

148

Munida pilorhyncha Miyake and Baba, 1966

Figure 21

Munida pilorhyncha Miyake and Baba, 1966: 81, figs 1, 2 (type

locality: Tosa Bay, Japan). — Baba et al., 2008: 112 (synonomies, type

data). — Baba et al., 2009: 180, figs 160—162 (Taiwan).

Munida sp. MoV 5200. — Poore et al., 2008: 20, unnumbered fig.

(lower right). —McEnnulty et al., 2011: app. 1,2.

Material examined. Western Australia: NMV J55145, 1 female (cl 21.5

mm, pcl 16.8 mm), Bunbury, 33° 00.504' S, 114° 34.26' E to 33° 00.12' S,

114° 34.5' E, 414—421 m, SS10/2005/13, 20 November 2005.

Colour. Carapace and somites 2—4 pale orange. Rostrum

orange. Supraocular spines white. Cheliped with orange/red

spots at base of spines and orange/red at distal ends of chela

fingers.

Genetic data. Not available.

Remarks. Poore et al. (2008) and McEnnulty et al. (2011) reported

the present specimens as “Munida sp. MoV 5200”; this is the first

confirmed record of Munida pilorhyncha from Australia.

Distribution. Japan, South China Sea, Philippines, Taiwan, Kei

Islands, Indonesia, 200—366 m. South-western Australia, 414—

42] m.

Munida roshanei Tirmizi, 1966

Figure 20D

Munida roshanei Tirmizi, 1966: 192, fig. 13 (type locality: Gulf of

Oman). — Baba, 2008: 116.

Material examined. Western Australia: NMV J55261, 1 male (cl 11.1

mm, pcl 7.2 mm), Barrow L1 transect, 21° 02.148' S, 114° 53.28' E to

21° 01.992' S, 114? 53.142' E, 90-100 m, S805/2007/8, 10 June 2007;

NMV J55991, 16 ovigerous females (cl 8.4 mm, pcl 5.0 mm to cl 10.9

mm, pcl 7.0 mm), 7 females (cl 8.5 mm, pcl 5.3 mm to cl 10.9 mm, pcl

6.7 mm), 22 males (cl 8.4 mm, pcl 4.9 mm to cl 11.2 mm, pcl 6.4 mm),

Mermaid L24 transect, 17? 46.104' S, 120? 43.152' E to 17? 45.948' S,

120? 42.942' E, 97-109 m, SS05/2007/97, 20 June 2007; NMV J56400,

2 males (cl 8.9 mm, pcl 5.9 mm; cl 13.5 mm, pcl 8.0 mm), 2 ovigerous

females (cl 9.9 mm, pcl 5.6 mm; cl 11.8 mm, pcl 6.8 mm), 3 females (1

damaged; cl 9.5 mm, pcl 5.6 mm to cl 10.0 mm, pcl 5.7 mm), | male (cl

$5.0 mm, 5.9 mm), Broome L25 transect, 16? 45.156' S, 121° 02.796' E

to 16? 44.592' S, 121? 02.208' E, 100-108 m, SS05/2007/116, 30 June

2007; NMV J56009, 2 ovigerous females (cl 10.9 mm, pcl 7.2 mm; cl

11 mm, pcl 7.8 mm), 1 male (cl 7.8 mm, pcl 4.7 mm), Lacepede L26

transect, 15? 48.672' S, 121? 03.738' E to 15? 48.426' S, 121? 03.654' E,

90—107 m, SSO5/2007/125, 1 July 2007; NMV J56098, 4 males (cl 8.4

mm, pcl 5.1 mm to cl 11.7 mm, pcl 7.0 mm), 1 female (cl 10.6 mm, pcl

6.7 mm), Adele L28 transect, 14° 33.732' S, 122° 55.092' E to

14° 33.684' S, 122° 54.906' E, 95-105 m, 8805/2007/161, 4 July 2007;

NMV J56395, 1 ovigerous female (cl 14.1 mm, pcl 8.6 mm), | female

(cl 9.1 mm, pcl 5.2 mm), 3 males (cl 11.3 mm, pcl 6.9 mm to cl 11.8

mm, pcl 7.8 mm). Ashmore L30 transect, 12° 26.7' S, 123° 36.048' E

to 12° 26.964' S, 123° 36.588' E, 95 m, SS05/2007/188, 6 July 2007;

NMV J56007, 1 ovigerous female (cl 11.3 mm, pcl 7.2 mm), | female

(cl 10.0 mm, pcl 6.3 mm), Ashmore L30 transect, 12° 25.944' S,

123° 35.784' E to 12° 26.064' S, 123° 35.868' E, 111 m, 5805/2007/196,

6 July 2007; NMV J55096, 1 male (missing chelipeds and rostrum; pcl

7.3 mm). Kalbarri, 27° 48.48' S, 113° 17.82' E to 27? 48.756' S,

113? 17.94' E, 112-123 m, SS10/2005/96, 4 December 2005; NMV

A.W. McCallum, S.T. Ahyong & N. Andreakis

J55047, 1 ovigerous female (missing chelipeds; cl 11.6, pcl 7.1 mm), 1

male (cl 12.8 mm, pcl 8.0 mm), 1 juvenile (cl 9.0 mm, pcl 5.4 mm),

Barrow Island, 21° 01.896' S, 114° 53.52' E to 21° 02.064' S,

114° 53.28' E,93m, SS10/2005/169, 13 December 2005; NMV J55045,

| female (cl 7.0 mm, pcl 5.0 mm), 6 males (cl 8.0 mm, pcl 4.3 mm to cl

12.7 mm, pcl 5.5 mm), Barrow Island, 20° 59.082' S, 114° 54.42' E to

20° 59.67' S, 114° 54.54’ E, 100-101 m, SS10/2005/170, 13 December

2005; NMV J55048, 1 male (cl 10.2 mm, pcl 5.8 mm), SS10/2005/170.

Genetic data. COI and 16S; see Table 1.

Remarks. Munida roshanei 1s reported for the first time from

Australia. Our specimens have much longer supraocular spines

(0.36 times rostrum length) than figured for the type (0.2 times

rostrum length) (Tirmizi, 1966: fig. 13). Longer supraocular

spines are also present on the specimen from the Red Sea

illustrated by Lewinsohn (1969). Our specimens all lack a row of

median gastric spines behind the rostrum, whereas the type has

a row of 3 median gastric spines behind the rostrum, and in other

specimens 1 or 2 spines are reported (Baba, 1988; Tirmizi and

Javed, 1993). There is clear sexual dimorphism in this species

with males having large, gaping cheliped fingers. All of our

specimens have cheliped fingers subequal to the palm in length.

Distribution. Indian Ocean (Gulf of Oman and Gulf of Aden,

Red Sea, Andaman Sea and Mozambique Channel), and

Phillipines, 16—528 m. Western Australia, 93—123 m.

Munida rubridigitalis Baba, 1994

Figure 22

Munida rubridigitalis Baba, 1994a: 13, fig. 6 (type locality: off

Central Queensland). — Ahyong and Poore, 2004: 41. — Poore, 2004:

234 (compilation). — Baba, 2005: 273. — Baba et al., 2008: 117.

Not Munida rubridigitalis — Poore et al., 2008: 20 (south-western

Australia, 396—411 m) (= Munida julumunyju sp. nov.).

? Munida rubridigitalis. — Macpherson, 1997: 610.

Munida sp. — Macpherson, 1994: 558, figs 13b, 90 (2 Munida

julumunyju sp. nov.).

? Munida rubrodigitalis [sic]. — Macpherson, 1999: 423, fig. 4e.

Type material. Paratypes: QM W19726, 2 males (pcl 10.0-12.7 mm), 3

ovigerous females (pcl 10.9—12.1 mm), 1 female (9.5 mm), Queensland,

Coral Sea, 17? 51.13' S 147° 8' E, 497—503 m, Cidaris Station 47-2, 16

May 1986.

Other material examined. Queensland: AM P42278, 1 male (pcl 14.7

mm, cl 20.5 mm), 1 ovigerous female (pcl 16.6 mm, cl 23.0 mm),

north-east of Tweed Heads, 27? 55—57' S, 154° 03' E, 549 m, trawl,

K78-23-09, 6 November 1978.

New South Wales: AM P88982, 1 male (pcl 12.6 mm, cl 17.8 mm),

4 ovigerous females (pcl 12.4 mm, 17.3 mm to pcl 14.7 mm, cl 20.6

mm), north-east of Tweed Heads, 28° 01.8' S, 153° 58.8' E, 549 m,

trawl, K78-09-05, 2 June 1978; AM P31504, 2 ovigerous females (pcl

13.1 mm, 18.9 mm; pcl 15.7 mm, cl 22.6 mm), south-east of Cape

Byron, 28° 41—44' S, 153° 51' E, 156 m, K78-17-21, 5 April 1978; AM

P31423, 1 female (pcl 16.0 mm, cl 22.5 mm), north-east of Sugarloaf

Point, 32? 15' S, 153* 02' E, 457 m, trawl, K78-03-04, 5 April 1978;

AM P31502, 1 male (pcl 13.4 mm, cl 19.5 mm), 1 female (pcl 13.1 mm,

cl 18.4 mm), north-east of North Solitary Island, 29° 53—50' S,

153° 42—43' E, 457 m, trawl, K78-06-07, 24 April 1978; AM P66658,

2 females (pcl 10.4 mm, cl 14.9 mm; pcl 12.2 mm, cl 18.0 mm), north-

east of Long Reef, 33° 42'S, 151° 42' E, K85-21-06, 439—466 m, K85-

21-06, 19 December 1985.

Australian Munida

149

Figure 21. Munida pilorhyncha Miyake and Baba, 1966, female, cl 21.5 mm pcl 16.8 mm, (NMV J551451).

l )

va yx

2mm

Figure 22. Munida rubridigitalis Baba, 1994. Baba, 1994, paratype, male cl 16.2 mm, pcl 10.2 mm (QM W19727): A, rostrum; B, right P2

dactylus and propodus. Scale: A-B = 2 mm

Genetic data. Not available

Remarks. Munida rubridigitalis, described from Queensland,

has been also reported from New South Wales, north-western

Australia, New Caledonia, Vanuatu, Indonesia (Kei islands)

(Macpherson, 1994, 1997, 1999; Ahyong and Poore, 2004; Baba

et al., 2008; Poore et al., 2008). The north-western Australian

records of M. rubridigitalis are herein shown to be referable to

a separate species, M. julumunyju sp. nov., which differs chiefly

in having a deeper, more strongly curved rostrum (see account

of that species). We compared paratypes and other Australian

material of M. rubridigitalis to six MNHN specimens identified

as M. rubridigitalis from New Caledonia (see material listed

under M. julumunyju). These New Caledonian specimens are

herein referred to M. julumunyju sp. nov., as are specimens

reported from New Caledonia and the Loyalty Islands as

Munida sp. by Macpherson (1994). Thus, other records of M.

rubridigitalis from New Caledonia, Kei Islands and Vanuatu

could also be referable to M. julumunyju sp. nov. (see Remarks

for that species). At present, M. rubridigitalis sensu stricto 1s

known only from eastern Australia.

150

Distribution. Eastern Australia, from Queensland and New

South Wales, 156—549 m.

Munida sacksi Macpherson, 1993

Munida sacksi Macpherson, 1993: 438 (part), fig. 6 (type locality:

Philippines). — Baba et al., 2008: 119.

Munida sp. MoV 5217. — Poore et al., 2008: 21. —McEnnulty et

al., 2011: app. 1.

Material examined. Western Australia: NMV J55039, 1 male

(cl 11.6 mm, pcl 8.2 mm), Ningaloo South, 22° 04.314' S, 113° 45.36' E

to 22? 04.854' S, 113° 45.36' E, 387—399 m, SS10/2005/151, 10

December 2005.

Genetic data. Not available.

Remarks. Munida sacksi was described from the Philippines,

based on a single female, missing the chelipeds. The present

specimen, reported by Poore et al. (2008) as "Munida sp. MoV

5217”, represents the first record of the species from Australia.

Distribution. Philippines, 300—330 m. Western Australia, 387—

390 m.

Munida shaula Macpherson and de Saint Laurent, 2002

Munida shaula Macpherson and de Saint Laurent, 2002: 474, figs

3A-C, E-H [type locality: Réunion Island]. — Baba et al., 2008: 117.

Munida sp. MoV 5215. — Poore et al., 2008: 21. — McEnnulty et

al., 2011: app. 1.

Material examined. Western Australia: NMV J55111, 1 ovigerous

female (cl 15.3 mm, pcl 9.9 mm), Ningaloo North, 21° 58.212' S,

113? 47.58' E to 21? 58.752 S, 113? 47.46' E, 324—356 m, SS10/2005/157,

11 December 2005.

Genetic data. Not available.

Remarks. This specimen, reported as "Munida sp. MoV 5215”

by Poore et al. (2008), agrees well with Munida shaula,

previously recorded from the western Indian Ocean (Reunion

Island, Madagascar and Zanzibar). This is the first confirmed

record of M. shaula from Australia.

Distribution. Zanzibar, Madagascar, La Réunion and

Mozambique, 263—510 m. Western Australia, 324—356 m.

Munida typhle Macpherson, 1994

Munida magniantennulata. — Baba, 1994: 12.

Munida typhle Macpherson, 1994: 549, fig. 60 (type locality: New

Caledonia). — Baba et al., 2008: 126. — Baba et al., 2009: 200, figs

181—182. — Poore et al., 2011: pl. 18H (colour). — Macpherson, 2013:

306. — Macpherson et al., 2017: 54.

Munida cf. magniantennulata. — Farrelly and Ahyong, 2019: 13, 55,

fig. 98.

Material examined. Queensland: QM W 19722, 1 female (cl 12 mm, pcl

7.5 mm), Coral Sea, 17° 45.04' S, 147? 48.014' E, 1223-1228 m, CIDARIS

I stn 20-4, 10 May 1986; OM W 19721, 1 female (cl 7.5 mm, pcl 5.0 mm),

Coral Sea, 16? 50.133' S, 147? 10.1' E, 1607-1609 m, CIDARIS I stn 35-3,

14 May 1986; NMV J72915, 1 ovigerous female cl 21.9 mm, pcl 13.8

mm, Coral Sea CMR, 23? 37.872' S, 154? 39.582' E to 23° 39.54' S,

154° 38.628' E, 1761-1770 m, IN2017_v03 stn 128, 13 June 2017.

A.W. McCallum, S.T. Ahyong & N. Andreakis

Genetic data. Not available

Remarks. The present specimens referred to M. typhle were

recorded by Baba (1994) from central Queensland as M.

magniantennulata. These two species are very similar overall,

but the cheliped of M. typhle has spines present on the dorsal

face of the palm and 2 lateral spines on the proximal half of the

fixed finger, which are absent in M. magniantennulata. In

addition, the P2—4 dactyli are longer in M. typhle (0.8 times the

propodus compared to 0.5—0.6 in M. magniantennulata); and

abdominal somite 2 has 4 spines on the anterior ridge while M.

magniantennulata has 2 spines. In the Queensland specimens

examined here, the P2 dactylus has 7—8 spines along the flexor

margin compared to the holotype of M. typhle, with 9 spines.

The QM registration numbers published by Baba (1994) were

listed incorrectly and have been corrected here. The ovigerous

female (NMV J72915; Ahyong and Farrelly, 2019: fig. 98)

collected during the recent [nvestigator voyage 1s much larger

than most specimens collected previously. This large female

differs from other specimens in lacking spines on the cheliped

dactylus, and the spines on abdominal somite 2 are proportionally

longer (similar in length to the branchial margin spines) and

limited to the median part of the anterior ridge.

Distribution. Somalia coast, Seychelles, southern Madagascar,

Taiwan, New Caledonia, Vanuatu, French Polynesia, 500—1940

m. Queensland, 1607—1761 m.

Acknowledgments

This work was supported by a Postdoctoral Fellowship from

the Australian Biological Resources Study. Funding for the

surveys was provided by the Commonwealth Department of

the Environment and the CSIRO Wealth from Oceans

Flagship. Thanks to Karen Gowlett-Holmes (CSIRO) for

permission to reproduce her colour images of squat lobsters.

We are grateful to Chris Rowley and Melanie Mackenzie

(both Museum Victoria) for collection management support.

We thank the following staff and their institutions for the loan

of specimens: Andrea Crowther (South Australian Museum),

Kareen Schnabel (NIWA), and Peter Davie (Queensland

Museum). Thanks to Laure Corbari (MNHN) for access to

collections and the loan of specimens. We are grateful to the

following indigenous organisations and community members

for their assistance and for sharing their language to name the

new species: the Tasmanian Aboriginal Centre, Kevin George,

the Kariyarra Aboriginal Corporation, the Dambimangari

Aboriginal Corporation, Dr Steve Kelly and all those from the

Nhanda community who attended the Nhanda Language Day,

Bundiyarra - Irra Wangga Language Centre and the Wangka

Maya Pilbara Aboriginal Language Centre. Finally, we

acknowledge the extraordinary work of Indigenous Language

Centres across Australia in revitialising and preserving

Aboriginal languages.

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Memoirs of Museum Victoria 80: 153-157 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.07

Phylogenetic placement of a recently discovered population of the threatened alpine

She-oak skink Cyclodomorphus praealtus (Squamata: Scincidae) in Victoria

JOANNA SUMNER!*, MARGARET L. HAINEsS!, PETER LAWRENCE’, JENNY LAWRENCE? AND NICK CLEMANN*

! Museums Victoria, GPO Box 666, Melbourne VIC 3001

? 2 Rose Street, Heyfield VIC 3858

* Parks Victoria, Licola Road, Heyfield VIC 3858

* Arthur Rylah Institute for Environmental Research, Department of Environment, Land, Water and Planning, PO Box 137,

Heidelberg VIC 3084 (Nick.Clemann@delwp.vic.gov.au)

* to whom correspondence should be addressed (jsumner@museum.vic.gov.au)

Abstract

sumner, J., Haines, M.L., Lawrence, P., Lawrence, J. and Clemann, N. 2021. Phylogenetic placement of a recently

discovered population of the threatened alpine she-oak skink Cyclodomorphus praealtus (Squamata: Scincidae) in

Victoria. Memoirs of Museum Victoria 80: 153—157.

The alpine she-oak skink Cyclodomorphus praealtus is a threatened alpine endemic lizard from the mainland of

Australia. The species 1s previously known from disjunct populations in Kosciuszko National Park in New South Wales

and three isolated localities in the Victorian Alps. The New South Wales and Victorian populations represent separate

evolutionarily significant units. In 2011, a fourth Victorian population was discovered. We conducted a phylogenetic

analysis and determined that the newly discovered population is discrete and may have been separated from other

populations since the end of the last glacial maxima. This population requires separate management.

Keywords

Introduction

The alpine she-oak skink Cyclodomorphus praealtus Shea

(1995) is restricted to sky island plains (sensu Koumoundouros

et al., 2009) above 1500 m elevation in the mainland

Australian alpine bioregion (Cogger, 2014). The species has

been recorded in New South Wales (NSW) at Kosciuszko

National Park and in Victoria at the Bogong High Plains,

around Mt Hotham, Mt Higginbotham and Mt Loch, and on

the Dargo High Plains at Lankey and Omeo Plains (Clemann,

2011; fig. 1). The Victorian sites are all within approximately

30 km of each other (Table 1). The species has specific habitat

requirements, occurring in alpine heathland and alpine grassy

heathland. These habitats generally have a disjunct occurrence

and are within a matrix of other alpine and sub-alpine habitats

in which the alpine she-oak skink is not known to occur.

Within some locations, the species distribution is further

fragmented by ski resort buildings and groomed ski slopes,

roads and tracks. The species is listed as nationally

endangered under the federal Environment Protection &

Biodiversity Conservation Act 1999, threatened in Victoria

under that state's Flora and Fauna Guarantee Act 1988,

critically endangered in Victoria by the Department of

Sustainability and Environment (2013) and endangered in

NSW under the Threatened Species Conservation Act 1995.

Cyclodomorphus praealtus, phylogenetic relationship, conservation, threatened species

The species is considered at risk from climate change

(Clemann, 2003; Koumoundouros et al., 2009) and is a high

priority for conservation management.

Populations of C. praealtus in NSW and Victoria are

reciprocally monophyletic at the mtDNA ND4 loci, with a

4.28% net sequence divergence between regions, and it was

recommended that they be treated as separate evolutionarily

significant units (Koumoundouros et al., 2009). The three

known Victorian populations exhibit strong genetic structure,

with no current inter-population gene flow, despite evidence

for historical connectivity (Koumoundouros et al., 2009).

This concurs with their present disjunct distribution on high

elevation plateaux and peaks in alpine meadows that are

separated by valleys of eucalypt forests.

In 2011, C. praealtus was discovered at a fourth Victorian

locality — the Wellington Plains (Clemann et al., 2016). This is

the southern-most area where this species has been recorded

and is 58 km from the closest Victorian population at Lankey

Plain. Here we present a phylogenetic analysis to determine

the relationship between this new population and those

studied by Koumoundouros et al. (2009). Our results provide

information that will assist the conservation management of

this species.

154 J. Sumner, M.L. Haines, P. Lawrence, J. Lawrence & N. Clemann

Table 1. Divergence among five populations of Cyclodomorphus praealtus in Victoria (VIC) and New South Wales (NSW). Uncorrected

p-distance (number of base substitutions per site averaged over all sequence pairs between groups; bottom) and straight-line distance (km)

between locations (top).

Bogong High Plains, | Mt Hotham, Lankey Plain, Wellington Plains, Mt Kosciuszko,

VIC VIC VIC VIC NSW

Bogong High Plains, VIC |- — — o |31 — — a — 96 a

Mt Hotham, VIC pool — — — [|- — Oy — — J7l — [T

Elevation

| Om

[zx] 400 m

|» | 800m

—3 1200 m

EE 1600+ m

Mt Kosciuszko}

^ » + ,

| " Op)

ns E

r è la

. a y

y exwWellinaton]Plains

Tasman Sea

Figure 1: Elevation map of the mainland Australian alpine region indicating the five major populations of Cyclodomorphus

praealtus, including the new location of Wellington Plains. Elevation is indicated by light, mid and dark grey areas at 500 m

intervals, with highest elevations at or above 1500 m above sea level. Adapted from fig. 1; Koumoundouros et al. (2009).

Phylogenetic placement of a recently discovered population of the alpine she-oak skink

Methods

Discovery of this population of C. praealtus and collection of

tissue samples is described by Clemann et al. (2016). Tissue

samples (tail tips) were collected from two sub-adult C.

praealtus from the Wellington Plains (1530 m elevation; fig.

1) in November 2011.

We extracted genomic DNA using a Qiagen DNeasy®

Blood and Tissue Kit according to the manufacturer’s

instructions (www.qgiagen.com). An 850 base pair region of

the mtDNA NADH dehydrogenase subunit 4 (ND4) gene was

chosen for sequencing because it has useful levels of

variability in this species and in related skink groups (Chapple

and Keogh, 2004; Koumoundourus et al., 2009). ND4 was

amplified according to the protocols and conditions in

Koumoundouros et al. (2009) using the primer pair ND4I

159

Chapple and Keogh, 2004). The resulting polymerase chain

reaction product was purified using ExoSAP-IT®

(Affymetrix), and the purified products were sent to Macrogen

(South Korea) for sequencing.

Forward and reverse sequences were assembled in Geneious

R6.1.8 (Biomatters Ltd.) and were aligned against 103 published

sequences and outgroups for this species (Koumoundouros et

al., 2009) using default settings in ClustalW. We translated the

protein-coding regions into amino acids and did not observe

stop codons or indels. IUPAC ambiguity codes were used for

heterozygous sites. We trimmed the alignments to 758 base

pairs because the end of the sequences could not be aligned due

to the length-variable loops and stems of the tRNA genes. The

Wellington Plains sequences were deposited in GenBank.

Tiliqua adelaidensis

Cyclodomorphus melanops

C. gerrardii

0.99

C. praealtus

-— IE

0.05

C. michaeli

Kosciuszko NP Cp105

L Kosciuszko NP Cp104

Ir Kosciuszko NP Cp107

l Kosciuszko NP Cp103

Bogong HP Cp077

NEW SOUTH

WALES

jog |! Bogong HP Cp028

Bogong HP Cp022

Bogong HP Cp023

Lankey Plain Cp042

r Lankey Plain Cp0 12

Mt Hotham Cp004

| Bogong HP Cp016

| Bogong HP Cp001

Wellington Plains CpTile5

Bogong HP Cp109

Bogong HP Cp015

Bogong HP Cp026

Mt Hotham Cp005

Bogong HP Cp054

Mt Hotham Cp002

Bogong HP Cp035

Bogong HP Cp009

VICTORIA

Figure 2: A Bayesian consensus tree of the mtDNA ND4 sequences, depicting the relationship between unique haplotypes at

each locality (Bogong High Plains, Lankey Plain, Mt Hotham, Wellington Plains and Kosciuszko National Park) for

Cyclodomorphus praealtus. Bayesian posterior probabilities are shown at major nodes.

156

To determine the relationship of the Wellington Plains

population to all other known populations, we created a

phylogenetic tree. Only unique haplotypes were used. We

used Modeltest 3.7 (Felsenstein, 1985) with the PAUP* plug-

in run within Geneious R6.1.8 to determine which of the 56

potential models of DNA sequence evolution was most

suitable for our data. The GTR+G model of evolution was

selected as the most appropriate model using the Akaike

information criterion (Akaike, 1974). The MrBayes version

3.2.6 (Huelsenbeck et al., 2001; Ronquist and Huelsenbeck,

2003) plug-in within Geneious R6.1.8 was used to conduct a

Bayesian Markov chain Monte Carlo phylogenetic analysis

using two million generations sampled every 500 generations,

with four heated chains sampling independently and a 25%

burn-in. Chain convergence was confirmed by examining the

log trace in Geneious and estimated sample size values > 200.

The mean between group distance was calculated among all

five populations in MEGAO (Tamura et al., 2006).

To infer the relationship between the mitochondrial ND4

sequences from Wellington Plains and the published

haplotypes in Victoria, we created a statistical parsimony

network using the program TCS (Clement et al., 2000) using a

95% statistical confidence limit for the maximum number of

nucleotide substitutions between two haplotypes.

Results

The ND4 sequences comprised 758 base pairs, including 63

variable sites and 50 parsimony informative sites. The

B Lankey Plains

L] Mt Hotham

O Bogong High Plains

Wellington Plain

J. Sumner, M.L. Haines, P. Lawrence, J. Lawrence & N. Clemann

sequences from the two Wellington Plains individuals were

identical and differed from all previously known haplotypes.

The Bayesian consensus tree (fig. 2) revealed that the ND4

haplotype from Wellington Plains falls ^ within the

monophyletic Victorian lineage and did not group with any

other populations. The Wellington Plains haplotype differed

from the populations in Victoria by 1.6-1.9% and from the

NSW population by 5.9% (Table 1).

A haplotype parsimony network of all ND4 sequences

generated in TCS (fig. 3) indicates that the Wellington Plains

haplotype forms a third distinct haplogroup cluster, separate

from all other Victorian haplotypes, and that they cluster

more closely with the north-eastern haplogroup that includes

individuals from Bogong High Plains and Mt Hotham. Eight

nucleotide changes separate the Wellington Plains sample

from the closest haplotype from the Bogong High Plains.

Discussion

Analysis of mtDNA ND4 sequences of two C. praealtus

individuals from Wellington Plains indicates the presence of a

haplotype that is not found in any other population across the

restricted range of this alpine endemic species. Further work

using multiple nuclear genes is needed before we can confirm

that the Wellington Plains population is isolated from the

other Victorian populations as the current data suggests. It 1s

not possible to determine the level of genetic diversity within

the Wellington Plains population due to the small sample size.

Figure 3: Population structure of Victorian Cyclodomorphus praealtus according to ND4 mitochondrial haplotype network.

Black indicates individuals from Lankey Plain, dark grey indicates those from Mt Hotham, white indicates those from Bogong

High Plains and diagonal stripes indicate those from Wellington Plains. The network structure indicates 3 haplogroups (1, 11 and

111) within Victoria. Each circle represents a unique haplotype, with the circle size indicative of frequency and sample sizes

within each circle. Empty circles represent missing haplotypes and differ by one base pair from the closest haplotype.

Phylogenetic placement of a recently discovered population of the alpine she-oak skink 157

The Wellington Plains population is the most divergent of

the populations in Victoria, with 1.6—1.9% sequence divergence

from the other populations, compared with the 0.9-1.2%

divergence among the other Victorian populations. Using the

conventional vertebrate mtDNA clock calibration of 2%

sequence divergence per million years (Avise, 2004), suggests

a divergence time between Wellington Plains and the other

populations in Victoria of between 500 OOO and one million

years before present. This places the divergence during the late

Pliocene to early Pleistocene, during which climatic

oscillations intensified resulting in rapid fluctuation between

cool-dry and warm-wet conditions across south-eastern

Australia (Markgraf et al., 1995). However, this predates the

most recent glacial period from approximately 25 000—18 000

years before present, during which the alpine habitat extended

into the valleys connecting the mountain peaks, allowing gene

flow between populations of other alpine endemics.

In 2019 severe bushfires burned across the entire Wellington

Plains plateau and, as a result, the status of this population is

now uncertain. Surveys to determine whether the population

still remains on the Wellington Plains are an urgent priority,

both to confirm the status of this population and to identify any

other specific threats. Threats already identified for C. praealtus

in the Victorian Alps include climate change, loss and

degradation of habitat (due to a range of processes, including

Several severe fires in the last decade, construction and

maintenance of infrastructure and roads, grazing and trampling

by exotic herbivores, and recreational activities), and predation

by exotic carnivores (Clemann, 2003, 2013). Historically, loss

and degradation of the habitat of C. praealtus due to cattle

erazing also occurred in the Alpine National Park, and damage

from this impact remains evident in parts of this lizard’s range

in Victoria (N. Clemann pers. obs.). Other reptiles endemic to

the mainland alpine region also occur on the Wellington Plains,

including threatened species such as the alpine bog skink

Pseudemoia cryodroma and the tussock skink P. pagenstecheri.

The fact that this population of C. praealtus remained unknown

until so recently demonstrates that further surveys of Victorian

alpine environments are needed if we are to document and

effectively manage the biodiversity of this region, especially in

the face of increasingly frequent and devastating bushfires.

Acknowledgements

Tissue samples were collected under a permit (number

100006167) granted by the Victorian Department of

Environmental Research, Department of Environment, Land,

Water and Planning (DELWP), and with approval from

Animal Ethics Committees at DELWP (approval number

15/08). We thank Wayne McCallum (Ranger, Parks Victoria,

Heyfield) who assisted in transporting survey tiles and

establishing tile transects and Tarmo Raadik for helpful

comments on a draft of this paper.

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Shea, G.M. 1995. A taxonomic revision of the Cyclodomorphus

casuarinae complex (Squamata: Scincidae). Records of the

Australian Museum 4T. 83-115.

Tamura, K., Stecher, G., Peterson, D., Filipski, A., and Kumar 5. 2013.

MEGAO: Molecular Evolutionary Genetics Analysis version 6.0.

Molecular Biology and Evolution 30: 2725-2729.

Memoirs of Museum Victoria 80: 159-168 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.08

The unusual genus Ramtheithrus (Trichoptera: Philorheithridae)

RosALIND M. ST CLAIR

Museums Victoria, GPO Box 666, Melbourne VIC 3001. rstclair@museum.vic.gov.au

St Clair, R.M.. 2021. The unusual genus Ramiheithrus (Trichoptera: Philorheithridae). Memoirs of Museum Victoria 80:

The descriptions of males of the two described species of Ramiheithrus (R. virgatus Neboiss and R. kocinus

Neboiss) are expanded using additional material. The female, pupa and larva of Ramiheithrus virgatus are described for

the first time for the genus. Preliminary genetic barcoding analysis suggests the presence of additional undescribed species

that are only known as larvae. The unusual larval habitat 1s described and issues relating to this are discussed.

Abstract

159-168.

Keywords Australia, groundwater habitat, COI gene, pectinate antennae

Introduction

The genus Ramiheithrus was created for two species, one each

in Victoria and Tasmania, by Neboiss (1974) with only adult

males known at the time. Information here brings the number of

species in the genus to four but the two additional species are

known only as larvae. The descriptions of males of the two

described species are revised here, partly based on specimens

from additional sites. Figures indicating differences from the

original descriptions, or those showing additional information,

are included for males. The adult female, pupa and larva of

Ramiheithrus virgatus Neboiss are described. The larva of R.

virgatus is compared with the larvae of two additional species.

The "rami" in Ramiheithrus refers to the distinctive

pectinate antennae. Pectinate antennae are otherwise unknown

in Trichoptera, except for three species of the Odontoceridae

genus Phraepsyche from South East Asia (Olah and Johanson,

2010) although the form of the pectination is apparently

different. A fossil trichopteran with yet another form of

pectination is discussed in Gao et al. (2016).

Few females of Philorheithridae are described and there is

variation between species in genera where more than one species

is known. Philorheithrus now has females of four species

described, and these show considerable variation in shape of

genitalia (Henderson and Ward, 2006; Mosely and Kimmins,

1954). This suggests that females of other Ramiheithrus species

may differ from the description below. The vaginal apparatus of

R. virgatus is illustrated below but there is no information on

other Philorheithridae females to compare with.

Larvae of Ramiheithrus have very rarely been collected.

The few available specimens provide information on variation

in the larvae and the likely reason for this rarity.

DNA barcoding data is preliminary but has assisted

taxonomic decisions.

Material and methods

The terminology here follows Henderson and Ward (2006)

and Holzenthal et al. (2007) but not Neboiss (1974).

All specimens are lodged in the Museum of Victoria (NMV)

and have been assigned registration (TRI) numbers. Reared

specimens usually include the case with pupal closure

membranes and larval sclerites, pupal skin and adult. The adult

has not emerged from the pupal skin in some cases and wing

venation cannot be discerned. Tissue samples sent to the

Barcode of Life Data System (BOLD; http://www.boldsystems.

org) for sequencing and analysis have been allocated L'TUT

numbers. Information on the methods used for sequencing and

cene segment sequenced are available on the BOLD website.

Data for specimens included in fig. 24 and Table | are available

from the BOLD website. The neighbour joining tree (fig. 24)

was produced on the BOLD website using a Kimura 2 parameter

and BOLD aligner (amino acid—based HMM) and produced in

Newick format using MEGA 7 on 26 October 2019.

Taxonomy

Ramiheithrus Neboiss

Type species: Ramtheithrus virgatus Neboiss

The information here uses and extends that given by Neboiss

(1974).

Diagnosis. This genus 1s strikingly unique with the antennae of

the adult pectinate on the basal half (fig. 1). In males, the superior

appendages are fused to segment IX and in basal half to each

other to form a hood over the genitalia (figs 2, 3, 22). This 1s like

the hood in Kosreithrus and, to a lesser extent, some species of

Aphilorheithrus. In Ramiheithrus, the superior appendages have

160

the ventral surfaces covered with moderately long spines in the

distal third. The inferior appendages are short and very simple,

most similar to those of Kosrheithrus. As with most

Philorheithridae, the forewing has vein R1 joining R2 just before

the wing margin in both sexes. However, this join is not present

in some specimens in some populations of R. virgatus (fig. 5) and

R. kocinus (fig. 23) and seems to be absent in at least some

species in other genera. The forewing has O to 5 crossveins

present between Sc and the costa proximal to the level of the r

vein in both species, fewer than most Aphilorheithrus species but

otherwise unusual in the family (fig. 5). The hind wing of both

sexes with Sc and RI merged for a short distance before

separating and diverging again (fig. 5). Spiracles on pleura

ventrally on segments 2 to 4 enlarged, elongate and variably

fleshy, smaller and more rounded on segments 5 to 8, only lightly

sclerotised around them (fig. 4).

In the larva, the anterior part of the frontoclypeal apotome

is almost twice as wide as posterior to the constriction (figs 10,

24, 26). On the mid leg, the strong dark distal spine on the tibia

remains despite the segment being fused with the tarsus (fig.

14). This spine is not present in other Australian Philorheithridae

genera, although long and short pale setae are often present in

this region. The larval case has the posterior opening almost

completely closed by larger sand grains and large flat mineral

particles but with gaps between these particles (fig. 17).

Generic diagnoses are not given for females or pupae

because they are known from one species only.

Description. Head: pilifers absent; maxillary palps 5-segmented,

long and slender in both sexes; labial palps short, about as long

as maxillary palps segments one and two combined; antennae

about as long as forewing, first segment stout, slightly longer

than vertical diameter of eye, second segment very short, next

14 to 16 segments with long lateral branches, becoming shorter

on the more distal segments, first 2 or 3 segments may have

branched rami (fig. 1); a pair of small anterior setal warts near

midline, pair of large, elongate, kidney-shaped posterior setal

warts and pair of posterolateral setal warts encircling posterior

half of eye. Thorax: 2 pairs of warts on pronotum, most medial

pair elevated; | pair of rounded scutal warts, usually larger than

scutellar warts; male forewing length 9 to 12 mm and lobe on

anal margin strongly sclerotised, apex broad apically truncate,

forks 1, 2, 3 and 5 present, male hind wing with forks 1, 2, 3 and

5, fork 3 footstalk of variable length. Abdomen: a small

projection on sternite VII, variable in size.

Male genitalia: superior appendages moderately long, deeply

excised, very broad and fused basally to segment IX and each

other forming a hood, ventral surface of each with short pointed

spines covering apical half; segment X (trilobed hood-like plate

of Neboiss, 1974) trilobed, lateral lobes similar to each other but

different to median lobe; inferior appendages a simple short

broad plate, sometimes scooped out apically, almost bilobed in

some specimens (figs 2, 3, 22). Phallus narrow, tapering apically.

Larva. Final instar about 9 to 12 mm long. Head: rounded

and somewhat flattened dorsally; strong carinae present just

above the eye from the anterior margin to about three quarters

head length, forming the lateral margin of the head on each

side; region between the carinae sloping ventrally near each

R.M. St Clair

carina which are raised above the head surface, roughened by

minute protuberances; antennae very short, close to front of

head and below carinae; labrum with 3 pairs of setae at about

3/4 length and another 3 pairs of short stout setae on front

margin, lateral two pairs curved along margin; ventral apotome

about 1/15 length of ventral ecdysial line; head setation as in

fig. 10, most setae on the dorsal surface (but not on the carina)

are short and very pale usually bent over or lying flat. Thorax:

metanotum with 3 pairs of sclerites, anteriomedial pair about 4

times as long as posterior pair (figs 11, 24); prosternum with a

large posterior central sclerite (figs 12, 25); mesosternum with 2

pairs of closely aligned sclerites medially, smaller and paler

than pronotal sclerite, posterior pair rectangular, anterior pair

tapering laterally (fig. 12); forelegs each with cluster of more

than 12 stout black setae centrally on anterior face of femur, all

arising from same point (fig. 13), tibial spur about 1/3 tarsal

length, tarsal claw about half as long as tarsus, apical spine of

tarsal claw almost as long as tarsal claw and lying closely along

its length; mid leg tarsal claw about 1/5 tibiatarsus, basal spine

of claw almost as long as tarsal claw and lying closely along its

length; hind legs much longer than forelegs, tarsal claw about

half tarsal length, 2 tibial spines on each tibiae about 1/3 tarsal

length, 1 on anterior face, 1 ventral, a group of short spines

apicoventrally (fig. 14). Abdomen: tergite IX short, not

completely covering the segment, lightly sclerotised, with

fringe of about 14 long setae and about 30 short or moderate

length setae (fig. 15). The lightly sclerotised channel running

ventrally from the base of anal claw towards the ventral

junction of the prolegs 1s deeper and more obvious than in some

other genera (fig. 16). This channel is darkly sclerotised at the

deepest point near the base of the anal claw. Anal claws with

tooth almost 1/2 length of claw.

Ramtheithrus virgatus Neboiss, 1974:

Figures 1-21

Type Material. Holotype male, Victoria. McKay Creek, Sassafras Gap,

36° 36'S 147? 4T E, 2 February 1974, A. Neboiss (NMV TRI-4558).

Paratypes 8 males, collected with holotype (NMV TRI-4559 to 4566).

Other Material examined. New South Wales. Alpine Creek

Kiandra, 35? 52' S 148° 29' E, 20 January 1966, 3 males 2 females

(NMV TRI-29289), 16 January 1968, 1 male (NMV TRI-29288), 13

December 1974, 2 males, 3 females (NMV TRI-29290), E.F. Riek.

Victoria. Roadside trickles 1.2 km N of Sassafras Gap, —36.6° E

14779? S, 10 February 2010, 1 larva (LTUT720-12) (NMV TRI-

54703), D. Cartwright and R. St Clair, 19 December 2013, 1 larva

(NMV TRI-55169), D. Cartwright; Roadside seep 1.6 km N of Sassafras

Gap, 10 February 2010, | larva (NMV TRI-55167), 11 February 2010,

preserved assorted dates to 21 February 2010, 3 reared males, 1 reared

female, 2 male pupae, 1 female pupa all with larval case and sclerites,

6 larvae (NMV TRI-54806 to 54815), D. Cartwright and R. St Clair;

Whites Bridge Mt. Buller Road 5 km SE Mirimbah, 37° 08' S

146° 26' E, 22 Mar 1985, 1 male A. Neboiss (NMV TRI-29291), 1 male,

2 Jan 2011, M. Shackleton (LTUT723-12) (NMV TRI-54702).

Diagnosis. As for the generic diagnosis with the following

additions. Male genitalia with segment X median lobe elongate,

longer than lateral lobes (figs 2, 3).

Larva distinguishable from larvae of the two other species

in the genus by the absence of small anterior sclerites on the

The unusual genus Ramiheithrus (Trichoptera: Philorheithridae)

spiracle

161

Figures 1—4. Ramiheithrus virgatus male: 1, Sassafras Gap body lateral; 2, Mt Buller genitalia ventral; 3, Kiandra genitalia lateral; 4, Mt Buller

abdomen lateral showing elongate spiracles on segments. ia = inferior appendage, sa = superior appendage, IX = segment IX, X = segment X.

prosternum, the more strongly developed brush of small

spines apically on the hind legs (fig. 14) and the reddish-brown

head and thorax (fig. 11). Collection of additional larval

specimens of the two undescribed species may show these

larval characters to not be as diagnostic.

Description. Revised after Neboiss (1974). As for the generic

description with the following. Antennae rami dark brown,

antennal segments yellowish. Forewings irregularly mottled

ereyish brown, a distinct pale oval spot close to wing margin

within fork 3, smaller and less conspicuous ones between the

veins above and below it (fig. 1); R1 joins R2 in males and females

although R1 joins the wing margin instead in most specimens

collected near Kiandra (fig. 5) and a few from Sassafras Gap.

Male genitalia: segment X median lobe laterally

compressed, lobe longer than lateral lobes; lateral lobes flat,

somewhat truncate in lateral view; phallus comes to a point

apically, curved downwards ventrally; inferior appendages

somewhat truncate distally in ventral view, irregular or with a

concavity (rarely 2) apically in some specimens (figs 2, 3).

Female. Antennal rami on segments 3 to 11, 12, or 13, rami

less than half the length of those in males. Forewings with

forks | to 5 present, forks 3 and 4 on long footstalks, hind

wings similar to male (fig. 5). Sternite VIII with a band of

short pale setae along posterior margin (figs 6, 7). Genitalia:

tergite IX very broad and sclerotised dorsally, narrowly cleft

mesally, medial notch on posterior margin. With a pair of

depressions laterally on segment, with a weak carina along

dorsal margin of each. Sternite [X with triangular sclerotised

area on anterior half of segment, fleshy laterally and posteriorly

(figs 6—9). Simple, apical lobes fused with segment X or lost.

Pupa. Head with a pair of prominent rounded, tuberculate

warts on the frons taking up most of the width of the frons

which are similar to, but more prominent than, those of

Aphilorheithrus and Kosrheithrus; a pair of long setae

dorsally medially on the head between the antennae, a second

pair just dorsal to the tuberculate warts (fig. 15), labrum with

3 pairs of setae in each dorsolateral corner; mandibles blade-

like in apical half, narrowing to a point, straight and serrate

on inner margin, rounded and smooth on outer margin, with a

pair of medium length setae basally (fig. 19); each scape with

medium length setae, 1 pair mesal dorsal and | pair distal

ventral (fig. 18). Legs without swimming fringes, rarely a

weak fringe on at least one mid leg. Abdomen with pairs of

162 R.M. St Clair

Figures 5—9. Ramiheithrus virgatus female: 5, Kiandra right fore and hind wings; 6, Kiandra genitalia lateral; 7, Sassafras Gap genitalia lateral;

8, Sassafras Gap genitalia dorsal; 9 Sassafras Gap genitalia ventral. d = depression.

The unusual genus Ramiheithrus (Trichoptera: Philorheithridae) 163

Figures 10-17. Ramiheithrus virgatus Sassafras Gap larva: 10, head dorsal; 11, head, thorax and first abdominal segment dorsal; 12, head, thorax

and first abdominal segment ventral; 13, foreleg anterior face lateral; 14, legs posterior face lateral; 15, abdominal segment IX dorsal; 16, last

abdominal segment ventrolateral; 17, case and posterior closure. cs = cluster of setae, pc = particles closing posterior opening.

164

R.M. St Clair

Figures 18-21. Ramiheithrus virgatus Sassafras Gap pupa: 18, shed cuticle head and thorax dorsal; 19, anterior of head ventral; 20, shed cuticle

abdomen dorsal; 21, last abdominal segment ventral.

anterior dorsal hook plates on segments 3 and 4 each with 2 to

4 teeth, segments 5, 6 and 7 each with 3 to 6 teeth; pair of

posterior hook plates on segment 5 each composed of a row of

16 to 25 spines, many bifid (usually different number on each

side) covering most of the width of the segment and with a

short gap between each plate, spines as long as plates (fig. 20).

Last abdominal segment elongate subtriangular, with 2 pairs

of long setae mesally, ventrally (fig. 21). Anal processes as for

the family: lightly sclerotised, long, very narrow and slightly

curved apically, each with a short dark pigment band close to

the apex and without setae (figs. 20, 21). Larval case with

anterior pupal closure a disc of silk without any openings

holding on a capping stone.

Larva. As for diagnosis. Final instar larva head capsule

width across eyes 1.3 to 1.4 mm.

Remarks. Note that the variation found in wing and genitalic

structures of the male were found in all populations.

The unusual genus Ramiheithrus (Trichoptera: Philorheithridae)

Ramiheithrus kocinus Neboiss, 1974.

Figures 2223

Type Material. Holotype male: Tasmania. Small creek in forest,

Corinna. 5 November 1972. A. Neboiss and G. Kocins (NMV TRI-

4567). 1 Paratype collected with holotype (NMV TRI-4568).

Other Material Examined. lasmania. Creek crossing Gordon

River Road. 42.887° S 146.379" E, 11 January 2012, 1 male

(LIUTI103-12) (NMV TRI-54701) M. Shackleton and J. Mynott.

Diagnosis. As for the generic diagnosis with the following

additions. Male genitalia with segment X median lobe broad,

shorter than lateral lobes (fig. 22).

Description. Revised after Neboiss (1974). As for the generic

description with the following. Type specimens blackish with

faint paler irregular mottling on forewings of which the most

conspicuous is a pale area covering cross veins s, r-m and m,

and extending along back along M1 about half the length of the

wing. Gordon River Road specimen greyish brown with

indistinct white mottling and spots, forewing 9 to 9.5 mm long.

R2 joining R3 in type specimens only, fork | with or without a

short footstalk. Forewings of male with vein A2 long in type

specimens, short in Gordon River Road specimen (fig. 23).

Male genitalia. Phallus broad apically; segment X lateral

lobes with a few moderately long spines medially; inferior

appendages come to a slight point laterally (fig. 22).

Female, pupa and larva unknown.

Remarks. This species is known from only two specimens from

the type locality and one specimen from a site (Gordon River

Road) about 170 km southeast of the type locality. DNA

information is only available for the southern specimen. There

is some variation in wing colour and venation and male

genitalia but with so few specimens available they are

considered conspecific for now.

Additional presumed species of Ramiheithrus

Larvae found at two localities in the Yarra Ranges National

Table 1. Specimen details for sequenced Ramiheithrus

169

Park are included here and are the only other species known

in the larval stage. This enables preparation of the generic

diagnosis and description. The distinctness of these two

Species 1s based on the genetic distance, as discussed in the

preliminary genetic analysis below.

Ramtheithrus sp. 1

Figures 24—25

As for the larval generic diagnosis and description with the

following additions.

Description. Larval head capsule yellowish brown, width

across eyes 1.4 mm, body length 12 mm. The only mid leg on

the one specimen has the remnant tibial spine about 1/3 as long

as that of R. virgatus; prosternum with pair of small sclerites

immediately anterior to the large central sclerite (fig. 24). These

characters may not enable separation from larvae of other

species when variation within each species is known.

Philorheithrus larvae also show almost no variation between

the three species recognised (Henderson and Ward, 2006).

Material examined. Victoria: Alderman Creek at Track 32, Yarra

Ranges National Park, 37.72]? S 145.94]? E, 5 November 2009, |

larva, J. Dean and R. St Clair (LTUT721-12) (NMV TRI54435).

Ramtheithrus sp. 2

Figure 26

Description. Adult unknown. Larva. As for generic diagnosis

and description. Head capsule yellowish brown, width across

eyes 1.5 mm, body length 12 mm. Case posterior closure

comparatively flat with attached particles smaller and sparser

than in the other two species.

Material examined. Victoria: Contentment Creek site ETT4DS RS,

CAPIM Black Spur Weir Project, 10 January 2013, 1 larva (LTUT1385-

14) (NMV TRI-55176); Contentment Creek, CAPIM Black Spur Weir

Project, 10 January 2015, | larva (NMV TRI-54816).

MV Registration Accession No. Life stage

No. (TRI)

54701

BOLD Process ID | COI GenBank BOLD BIN No.

Accession No.

ABV8840

LTUT721-12 JOS152 Larva

54702 LTUTI385-14 EPAVT54 Larva

54437 LIUT720-12 JOSI51 Larva

54703 LIUT723-12

KX292939

JOS154 Adult male

54435 LIUTIIOS-12 JOS316 Adult male

KX293205 ACC8741

166 R.M. St Clair

Figures 22 — 23. Ramiheithrus kocinus Gordon River Road male: 22, genitalia ventral: 23, left forewing (rotated 180° for comparison with other

wings). Figures 24 — 26. Ramiheithrus sp. 1 larva: 24, head and thorax dorsal; 25, head and thorax ventral. Ramiheithrus sp. 2 larva: 26, head

dorsal. ia = inferior appendage, sa = superior appendage, X = segment X.

The unusual genus Ramiheithrus (Trichoptera: Philorheithridae)

Discussion of COI data

An abbreviated COI taxon identification tree is shown in fig.

27 and information on specimens is provided in Table |. Only

one specimen was barcoded from each of five localities.

Usually, at least five specimens over the range of the species

are required to confirm genetic distinctness. Thus, more

sequencing is required to confirm the current interpretation.

The COI taxon identification tree indicates that the two

larvae from the Yarra Catchment (LTUT721-12 and

LTUT1385-14) are genetically very different from each other

and from other specimens (fig. 27). Because there is no adult

specimen available from either locality, these species are not

being formally described.

The genetic distance between the specimen of

Ramiheithrus virgatus from near the type locality (LTUT720-

12) and the specimen from near Mt Buller (LTUT723-12) is

close to 5%. This is greater than often considered necessary to

indicate distinct species. However, no consistent

morphological features were found to support recognition of

167

the Mt Buller population as a new species. The generally large

distances between all specimens indicate that the commonly

applied 2% or 3% threshold for species separation may not

apply to this genus. As a result, all the Victorian (apart from

the larvae from the Yarra catchment) and New South Wales

specimens are here assigned to the one species.

Discussion of larval habitat and its implications

Larvae of Ramiheithrus virgatus have been collected for the

first time. The first larva collected was in a roadside trickle

that is only present after rain and which was dry the day

following collection. The trickle appeared as a point where a

narrow conduit of groundwater reached the surface, rather

than just a point where a broad area of groundwater happened

to reach the surface. Investigation found pupae and more

larvae in saturated coarse gravel behind a large mound of

moss. The seep was less than two square metres on a vertical

area of road cutting. Water flowed into the roadside ditch from

this area in rainy conditions. This locality was very close to

Ramtheithrus|L TUT 721-12] RCUAIdermanCreekatT rack3 2|BOLD\0/2ABV8839

Ramiheithrus|LT UT 1103-12|CreekcrossingGordonRRd|BOLD WO /2ACC8/41

Ramtiheithrus|L T UT /723-12]White sBnidgeonMtBullerRd|BOLD\0/2ABV8340

— Ramiheithrusvirgatus|LT UT 7 20-12|Roadsideseep 1.2 kmNEustaceG apRd|BOLD'07 2ABV9442

Ramiheithrus|LT UT 1385-14|ContentmentCKk054BlackSpur.Sweep|BOLD V0 72ACM3556

I—1

0.0100

Figure 27. BOLD Taxon identification tree.

168

the type locality and an adult was collected at light at this new

locality soon after finding the larvae and pupae. Some pupae

were reared to the adult stage. No similar localities were

found nearby, despite extensive searching.

After the habitat was recognised, attempts were made to

find similar habitats on the Mt Buller Road, the only other

locality in Victoria where adults have been collected. No such

habitat or Ramiheithrus larvae were found. Areas where

eround water was flowing into roadside ditches following rain

were found but not associated with a particular moss and not

appearing to lead to a gravelly conduit. Presumably, such

areas are present but not on the roadside, at least at the time of

searching. Being so small, such habitats would be extremely

difficult to locate hidden within the vegetation.

The only specimen of Ramiheithrus sp. 1 was collected at a

standard riverine habitat. However, as only one larva was

collected, despite the site being sampled on numerous occasions

by the Environment Protection Authority Victoria over a 20-year

period, this clearly is not the preferred habitat of the species.

This larva was presumably washed in from its usual habitat. A

similar but smaller area of moss was noted on a road cutting

about 100 metres uphill from the site but not investigated.

The specimens of Ramiheithrus sp. 2 were collected from

very small trickles, a slightly different habitat to that of

Ramiheithrus virgatus. Several specimens were collected;

therefore, this appears to be their habitat.

The assumption is made that the habitat is similar for all

species of Ramiheithrus. The fact that only a single larva has

been found in a true riverine habitat supports the fact that the

main habitat is unusual, and soaks like that where R. virgatus

was collected near Sassafras Gap are considered the most likely.

Although found in alpine areas, this genus has not been

found above 1500 metres. This also could be related to the

habitat as it would occur lower down the mountain where

eround water seeps out of conduits and associated with a

particular type of moss.

The fact that collection of the larvae is so difficult means

that species in this genus will be difficult to manage.

Ramiheithrus virgatus 1s listed as vulnerable 1n Victoria and

R. kocinus is listed as threatened in Tasmania. The adults are

rarely found and in low numbers when they are collected.

However, E. Riek collected adults at the one locality four

times over 1] years. Adult specimens have been collected

from the White Bridge locality on more than one occasion,

although collecting on other occasions was unsuccessful. It 1s

possible that adults usually remain near the larval habitat.

largeted collecting would be extremely difficult due to the

very patchy nature of the habitat.

With the larvae and adults probably living in small areas

of habitat spread over a wide area, could this be the reason

R.M. St Clair

why adults have rami-bearing antennae. Presumably they

have a sensory function. Gao et al. (2016) suggested rami

enable greater sensory function for detection of pheromones

or volatile chemicals from food. It is possible the rami in

Ramiheithrus are used to locate other adults in highly

localised habitat patches.

The absence of swimming fringes on the mid legs of the

pupae may be an adaptation to the larval habitat as the pupae

were not collected in free water. At this stage, the pupa of

only one species 1s known so this may not be typical of all

Ramiheithrus pupae. Examination of pupae from most

Australian Philorheithridae genera found they have swimming

fringes. One of the two species of Philorheithridae pupae

described from New Zealand, Philorheithrus agilis 1s found

in swift streams has swimming fringes while Philorheithrus

litoralis has a very weak fringe of few pale setae and is found

in small streams or seepages (Henderson and Ward, 2006).

Two females of Ramiheithrus virgatus reared from pupae

had many well-developed eges 1n the abdomen. This may be

another adaptation to the unusual habitat, but because it 1s not

known how many philorheithrid species are similarly

advanced when emerging, this cannot be confirmed.

Acknowledgements

I am very grateful to David Cartwright for organising our

collecting trip and finding the larval habitat of Ramiheithrus

virgatus. John Dean is thanked for improving the manuscript

and David Cartwright for very helpful reviewer comments.

References

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D. 2016. Convergent evolution of ramified antennae in insect

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Proceedings of the Royal Society B 283: 20161448. http://dx.doi.

org/10.1098/rspb.2016.1448

Henderson, I.M., and Ward, J.B. 2006. Four new species of the caddis

genus Philorheithrus (Trichoptera: Philorheithridae) from New

Zealand. Records of the Canterbury Museum 20, 21-33.

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of Australia and New Zealand. British Museum (Natural History):

London. 550 pp. https://doi.0rg/10.5962/bhl.title.118696

Neboiss, A. 1974. A new caddis-fly genus from Victoria and Tasmania

(Philorheithridae: Trichoptera). Victorian Naturalist 49: 322—325.

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Memoirs of Museum Victoria 80: 169-181 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.09

George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-

century Australian entomology

DEIRDRE COLEMAN

https://hindanexpert.unimelb.edu.au/profile/50297-deirdre-coleman

Robert Wallace Chair of English, English and Theatre Studies, School of Culture and Communication, University of

Melbourne, Parkville VIC 3010, Australia. (colemand(?unimelb.edu.au)

Abstract

Coleman, D. 2021. George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-century

Australian entomology. Memoirs of Museum Victoria 80: 169-181.

This article focuses on the correspondence and careers of two lepidopterists, George Lyell and F. P. Dodd. Drawing

on Dodd's unpublished letters to Lyell during the late nineteenth-century rage for butterflying, it examines how private

acquisition gave way to the professional activity of collecting and, in Lyell's case, the eventual gifting of a large and

significant collection of moths and butterflies to the National Museum of Victoria from 1932 through to 1946. The article

also examines how issues of authority and expertise were measured and contested among collectors in this period.

Keywords

Introduction

Professionalisation in nineteenth-century life sciences 1s a lively

topic within British and North American histories of science,

but it is a topic less examined in the Australian context. This

article extends current debates by examining the correspondence

between two self-taught Australian lepidopterists, Frederick

Parkhurst Dodd (1861—1937) and George Lyell (1866-1951).

Both were private collectors, born in the shadow of the

Grampians in south-west Victoria, but that is where the

similarities end. Lyell built a large collection of Lepidoptera

while working full-time in business, whereas Dodd supported a

large family through selling his specimens. The article begins

with Lyell’s decision in 1932 to donate his large collection to the

National Museum of Victoria. It then moves back in time to an

examination of Dodd’s letters to Lyell from 1897—1904. These

letters, while reflecting informal one-to-one transactions of

exchange, donation and purchase, operate within a much larger

push and pull of external factors. These external factors include

a wide network of people who were themselves subject to a

thicket of protocols, depending on their perceived status within

the group. Furthermore, the complexity and interdependence of

the various people involved in the science of entomology led to

many disputes about who exactly qualified as professionals.

Arguments as to who held the most requisite authority and

expertise were particularly acute during what has been dubbed

the Period of the Amateurs, 1890—1930 (Mackerras, 1949).

F. P. Dodd, George Lyell, lepidoptery, collecting, correspondence, professionalisation, National Museum of Victoria

The George Lyell collection

In March 1946, Richard Pescott (1905—1986), Director of the

National Museum of Victoria, informed the Australian

Broadcasting Corporation’s Melbourne office that the first

instalment of a magnificent collection of Australian butterflies

and moths was about to be exhibited in the main hall of the

museum on Russell Street. The collector was George Lyell

(1866-1951; fig. 1), and his gift to the museum was so large

that it had to be exhibited in relays, from | April 1946 until

mid-year. The announcement contained an overall description

of the collection as well as information gleaned from an

interview with Lyell about his gift to the state of Victoria.

Although the whole appeared under Pescott’s name, the real

author was journalist and naturalist Charles Barrett (1879-

1959). After the Australian Broadcasting Corporation

published the piece, Barrett then circulated it to The Herald,

The Sun, The Age and The Argus.

We learn from Barrett’s interview that Lyell had decided to

donate his collection after suffering a serious illness in 1932.

In the 14 years since then, he had devoted himself to the

colossal task of preparing more than 50 000 individual

specimens for the handover, remounting and resetting many of

them in the process. His aim was to build for the museum “the

most comprehensive collection of Australian moths and

butterflies ever known”. At the time of Barrett’s writing, the

collection numbered 51 216 specimens, consisting of 11 721

butterflies and 39 495 moths, representing 6177 species all

told. World-renowned authorities had already named 534 type

170

specimens, but new species were still being discovered and

named within the collection. In addition to praising Lyell’s

ereat scientific knowledge and technical skill, Barrett

emphasised that the gift had been unconditional, Lyell having

taken the “broadminded scientific view" to amalgamate the

Museum’s collection with his own. In this way, gaps would be

filled, ensuring that the collection would be “truly

representative of Australian entomology’. Barrett concluded

with a brief overview of the collection’s highlights, such as the

“particularly beautiful” swallowtail butterflies and the “lovely

Blue butterfly, Papilio ulysses". It was in the moths, however,

Figure 1. George Lyell as a young man.

D. Coleman

that the collection approached perfection (Melbourne

Museum, Notice to Manager, ABC, AB 576, 29 March 1946).

Lyell was born at Ararat, Victoria, in 1866, the fourth of

eight children (Hewish, 2014). His father was a printer born in

scotland and his mother was English. Apparently, he showed

little interest in. natural history until he caught his first

butterfly, a caper white, at Albert Park in 1888. Wanting to

know more about this creature, the 22-year-old contacted

Frank Spry (1858—1922) who immediately introduced him to

the Field Naturalists’ Club of Victoria, which was founded in

1880. While attending meetings regularly over the course of

George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-century Australian entomology 171

the following year, Lyell continued to collect at Albert Park,

where he distinguished himself by collecting 13 lesser

wanderers, a butterfly that had not been recorded in southern

Victoria for well over a decade. He also made occasional

visits to other well-known hunting grounds: Murrumbeena,

Springvale, Hampton and Cheltenham.

In 1890, Lyell moved to Gisborne in central Victoria to

work as bookkeeper for the town’s largest business, Cherry &

sons Pty. Ltd., timber merchants and manufacturers of dairy

equipment. Later, as Lyell rose to Manager and then Director,

the firm developed an export business 1n entomological supplies

such as nets, mounting boards, pins, forceps, cotton wool, boxes

and cabinets (Hewish, 2014). Here, in this small country town

outside Melbourne, Lyell lived for the rest of his life. His

collecting was confined to Victoria and New South Wales,

except for one trip each to South Australia, Tasmania and

southern Queensland. Nevertheless, he corresponded and

exchanged Australia-wide with scores of notable entomologists

such as A. J. Turner and R. Illidge of Brisbane, G. M. Goldfinch

of Sydney, G. B. Lower of Adelaide and F. P. Dodd of Kuranda,

north Queensland. He also co-authored, with G. A. Waterhouse,

The Butterflies of Australia (1914). This extensive exchange

network across Australia resulted in long series of individual

species, a notable and highly valuable scientific feature of

Lyell's collection.

In outlining Lyell's career, Barrett paid particular attention

to Lyell's triumph over the largest obstacles facing collectors,

namely housing his collection and protecting it from the

scourges of pest infestation and mould. Lyell achieved this by

devising a small display cabinet of six or eight drawers,

constructing it in such a way that each subsequent unit could

be fitted together to make more cabinets, similar to the way in

which a bookcase might be enlarged. In the 1890s, at the start

of his collecting career, Lyell could only afford to build one of

these cabinets annually, but by 1932 he owned more than fifty,

built by Cherry & Sons. What Barrett did not know, or chose

not to disclose, was that Lyell had approached Sir Macpherson

Robertson in 1934 for £500 to build the many extra cabinets he

needed to house his gift. Robertson, founder of the

MacRobertson Confectionery Company in the Melbourne

inner-city suburb of Fitzroy, was one of Australia's richest and

most successful businessmen. Whereas Cherry & Sons

struggled after the Depression, business remained strong for

Robertson who contributed generously to the upcoming

centenary of Melbourne's founding in 1835, including a

substantial donation towards the building of the city's

herbarium. In essence, Lyell asked Robertson to do for

Australian Lepidoptera what he had recently done for botany.

signing off anonymously as a collector and fellow-Scot, the

normally modest Lyell took the opportunity to boast that his

collection would be "an object lesson to all the museums of the

world and a lasting scientific attraction to Melbourne". Lyell

finished by inviting Robertson to make enquiries of the

Museum Director, D. J. Mahony, an invitation promptly taken

up by the confectionary magnate (Melbourne Museum,

Archive Box 579, 27 December 1934). [n his response, Mahony

revealed Lyell's identity and confirmed that a shortage of

money was indeed hampering the progress of his gift. In

praising the collection as “the best of its kind’, Mahony

described the meticulous way in which Lyell cared for his

insects. Not only did he perform yearly stocktakes to check on

their condition, he also compiled annual balance sheets

showing acquisitions, as well as insects discarded or

exchanged. Mahony confirmed that "Every specimen is

therefore fully documented" (Melbourne Museum, Archive

Box 579, 9 January 1935). In the end, Robertson declined to

support Lyell, arguing that he had already over-committed

himself in support of the city's centenary.

George Lyell and Frederick Parkhurst Dodd

When interviewing the elderly Lyell in 1945, Barrett asked him

which was the favourite of his 6177 insect species. Lyell at first

parried the query, declaring “you have set me a poser. They are

all beautiful’. In the end he confessed that it was the Queensland

wood moth Dudgeonea actinias Turner, bred "from a rotting log

in a creek bed at Townsville" by Frederick Parkhurst Dodd (fig.

2). In 1905, Cyclone Leonta buried the log under 30 feet of flood

debris. Of the 13 specimens bred by Dodd, seven are in the Lyell

collection (fig. 3). While Dodd was just one of Lyell's many

Australian correspondents, we know a lot about their relationship

because there is an extensive run of letters from Dodd to Lyell in

the Melbourne Museum archive. Although, unfortunately,

Lyell's letters to Dodd do not appear to have survived, the

correspondence reveals the many interdependent players and

complex processes at work in shaping the Australian

entomological community during the late nineteenth century.

The collecting, exchanging and selling of insects was

serious business in the years leading up to Australian Federation

in 1901, evident in the brisk and lively circulation, both locally

and internationally, of collectors, publications, specimen boxes

and letters. Given the enthusiasm for Lepidoptera, there was

serious money to be made, especially on large and brightly

coloured tropical specimens. The story of Conrad Kelsall, an

English immigrant farmer who settled 1n the rainforest of north

Queensland, is instructive. Within four short months in 1903,

we see the rise and fall of Kelsall’s hopes for a tidy profit from

butterflying on the Little Mulgrave River. In letters to his sisters

back in Devon, he began with great confidence, declaring his

tropical home as “so new & unexplored that one is almost

certain of making new finds”. With the help of an indigenous

man nicknamed Paddy, who captured about a dozen males of

the large birdwing butterfly Ornithoptera euphorion Gray,

Kelsall asked 6d for each of them from Alfred Bell, an insurance

agent and butterfly enthusiast based in Cairns. Later, for a box

of 100 butterflies, Bell paid Kelsall 25/-, proposing that, instead

of cash transactions, his profit would double if he agreed in

future to “run on halves" and "take some risks". Once the

business was “in full swing", Bell envisaged that similar boxes

might average out at “£3 or £4 per hundred". Although new to

the collecting business, Bell boasted international contacts with

the famous lepidopterist Walter Rothschild, as well as with

Watkins and Doncaster, the English natural history dealers. Bell

was also supplying insects to Lyell and Waterhouse in Sydney,

at that time preparing their catalogue of Australian butterflies.

But within a few months, Kelsall confessed to his sisters that he

172

and his business partner needed “to modify our castles

considerably". After failing to catch anything in an expedition

to Lake Barrine, they returned to discover that all but 14 of the

100 insects they had sent to the Australian Museum had been

rejected. Worse was to come in the shape of an insulting and

caddish letter from the English dealers who purchased only two

of the butterflies sent over. Claiming that the rest were too

damaged, they promised to return them all to Queensland but

no box arrived, leaving Kelsall to conclude, ruefully, that “It 1s

easy swindling a person 12 000 miles away" (Lambkin, 2013).

While Kelsall was one of many minor part-time enthusiasts,

Frederick. Parkhurst Dodd is now regarded as a leading

Queensland collector. Born in Wickliffe, Victoria, one

stagecoach stop from Lyell's birthplace in Ararat, he was the

son of a pound-keeper and the eldest of eight children

(Monteith, 1991; Neboiss, 1986). After the family moved to

Stawell, on the edge of the Mallee, Dodd was educated at the

same local state school as Lyell. Here the similarities stop.

While Lyell went on to lead a settled life in a rural town within

easy reach of Melbourne, Dodd joined the bank in Victoria and

was then shunted around to various bank jobs 1n Queensland,

starting in remote Townsville in 1884. He never returned to live

in Victoria. Apart from six years in Brisbane, where he met and

learned much from a number of prominent entomologists

connected to the Queensland Museum and the Natural History

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Society, Dodd’s early life as a bank clerk took place a long way

from the metropolitan scientific societies of Brisbane,

Melbourne and Sydney. Eventually, hating the confinement of

office work, and determined to become a full-time collector, he

left the bank and eventually settled in 1904 in Kuranda, a tiny

town on the Atherton Tableland. The area was rich in insects,

but all his entomological books and journals had been destroyed

by Townsville’s Cyclone Leonta the year before. In Kuranda

there were no local libraries and no entomological societies or

meetings in which he could participate.

In Dodd’s letters to Lyell, we see the value, and indeed the

necessity, of correspondence that connected him to

Melbourne’s scientific and collecting community, including

the Victorian Field Naturalists’ Club and the National

Museum of Victoria. Lyell kept Dodd connected to

professional societies and their activities, including access to

the all-important journals. Membership of these societies and

subscriptions to their publications were expensive, often

beyond Dodd’s means, but when business was good, he turned

to Lyell for suggestions as to who might propose and second

his admission. In addition to Waterhouse and other prominent

entomologists at the Australian Museum in Sydney, Lyell

counted Walter Froggatt (1858—1937) as a friend. Founder of

the Naturalists’ Society of New South Wales in 1891, Froggatt

published regularly on Australian entomology in

Figure 2. Dudgeonea actinias ‘Turner.

174

the Proceedings of the Linnean Society of New South Wales.

After Froggatt was appointed government entomologist to the

New South Wales Agricultural Department in 1896, Dodd

asked Lyell to mention him in case the Department wanted to

purchase some of his specimens (Museums Victoria Archives,

OLDERS YSTEM~03023, AB 00368, 24 March 1901).

Dodd’s correspondence with Lyell is rich in self-description,

shedding light on the so-called divide between mere collectors

and entomologists. The label of mere collector had been hurled

by Gerard Krefft, controversial Curator of the Australian

Museum, at William John Macleay during a parliamentary

investigation into the museum in 1874 (Ville, Wright, and Philp,

2020). Although this divide between the true scientist and the

mere collector was more rhetorical than real, it was deeply

embedded in nineteenth-century thinking about who exactly

possessed the authority and expertise to speak for entomology.

In 1838, the British entomologist John Obadiah Westwood

(1805-1893) described as the very “lowest class of

entomologists” those "whose sole object 1s the procuring, either

by capture or by purchase, of a collection of handsome insects,

to be placed in drawers without any arrangement other than

that of beauty and colour or size" (Wale, 2019, pp 405—406). As

an insect breeder and naturalist, Dodd was far from this lowest

class. His deep knowledge of the bush around him and pride in

his technical skills are clear in his letters, which are full of

observations about the habits and life histories of 1nsects— their

location and distribution, their food plants, their relationship to

other species and genera, their enemies and their mechanisms

for self-defence. Many of Dodd's fine-grained observations

stemmed from tireless watching of the insects with which he

lived intimately, both at home in breeding boxes, in the bush

beyond, and in his Kuranda garden, planted with especially

chosen flower and tree species.

Despite his expertise, it is notable that Dodd never describes

himself as an entomologist. Instead, his insects are

"entomological material” and he himself is a “worker in

Entomology”. He even apologised to Lyell in his first letter for

addressing him as entomologist on the envelope, explaining

that he wanted to ensure the letter reached him (December

1897). There are several explanations for why Dodd refused to

call himself, or anyone else he admired, an entomologist. The

first reason was his dislike of the entomological fraternity: “I

have a very poor opinion of Entomologists generally", he tells

Lyell. In particular, he had little time for the growing number of

sedentary and salaried museum men whom he regarded as far

less knowledgeable and skilful than himself. Writing to Lyell 1n

1901 about J. A. Kershaw (1866-1946), later curator and then

Director of the National Museum of Victoria, Dodd places him

contemptuously amongst the “amateur Entomologists", with the

word "amateur" doubly underlined for emphasis (National

Museum of Victoria, 15 July 1901). The reason for his dismissal

of Kershaw and others stemmed from his pride as an insect

breeder with first-hand eye-witness experience. Even Froggatt,

author of Australian Insects (1907), the first comprehensive

textbook on Australian entomology, failed to come up to scratch

in Dodd’s opinion. While he conceded to Lyell that Froggatt

“may be a good entomologist” he added that “he had better

drop writing the life histories of moths”. According to Dodd,

D. Coleman

Froggatt had been duped by a Newcastle collector called

Thornton into believing that the larvae of Endoxyla encalypti

had bored as deep as five feet into the wattle tree roots, “high

class rubbish” that had been published in the Proceedings of the

Linnean Society of New South Wales (National Museum of

Victoria, 1 January 1898). Elsewhere, Dodd refers to Froggatt

(anonymously) as “an Australian Munchhausen” for producing

the same exaggerated observations (Dodd, 1916).

Keen to read the latest essays, notes, and pamphlets about

insects, Dodd published some important discoveries of his

own in entomological and natural history journals, both

Australian and international. Geoff Monteith gives two

examples of the ways in which Dodd was ahead of his time in

understanding the life histories of insects. Dodd 1s now known

for his breakthrough insights into the symbiosis between

ereen tree ants and the highly prized butterfly Liphyra

brassolis Westwood. Also notable was his detection of the

mimicry between the rare swallowtail butterfly, Papilio

laglazei Depuiset, and the poisonous day-flying moth, Alcides

agathyrsus Kirsch (Monteith, 1991). However, with no access

to large reference collections and short on time, Dodd was, as

he confessed to Lyell, “but poorly acquainted with the names

of our [Australian] insects’ (National Museum of Victoria, 11

October 1897). A few months later, having picked up on the

fact that Lyell was “well posted in the names”, he asked him

for help, confessing that

unfortunately hundreds of my species are yet

unknown to me by name. I shall do all I can to

get them identified + hope to get a list compiled

such as yours. When furnishing particulars of

specimens sent, or even acknowledging

exchanges, can you kindly add names of sub

family + family of individual specimens; in

many cases the generic name + view of the moth

does not tell me what the insect 1s (National

Museum of Victoria, 11 December 1897).

Before long, he was thanking Lyell for the trouble he had

taken in supplying him with names (National Museum of

Victoria, 12 February 1898).

Dodd’s failure to master the precise names of insects did

not mean that he despised the systematists. In fact, he was

extremely proud of the great attention shown to his collection

by Brisbane-based Dr Alfred Jefferis Turner (1861-1947), an

expert in classification who would often travel to Townsville,

and later Kuranda, to study, describe and name Dodd’s insects

(National Museum of Victoria, 29 September 1900). You can

hear Dodd’s pride in a comment he made to Lyell of a small

syntomid he sent on to him. Turner, he wrote, regarded the

Specimen “as a great prize, and in fact he thinks very

favourably of all the kinds I am sending you. He often calls

upon me, to examine my collections, + learn if I have anything

fresh in the way of captures, or from my breeding boxes”.

Rowland Illidge, Dodd's mentor during his years in Brisbane,

was another expert identifier, helping him to name hundreds

of species. Despite his self-confessed deficiencies in this

arena, Dodd was never cowed. When Lyell returned a large

hawk moth, believing it was not the moth Dodd had promised

George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-century Australian entomology 175

him, Dodd exploded with irritation. In this instance, where

the differences between species were scarcely perceptible,

Dodd’s experience as a breeder gave him that extra degree of

self-certainty (National Museum of Victoria, 27 May 1901).

On balance, rarity was a strong possibility in the remote

Queensland tropics, but how could Dodd always be certain,

lacking expert associates and reference materials? Perhaps his

rare insects were common elsewhere? Furthermore, in several

letters, responding to Lyell’s doubts about identifying a

particular specimen, Dodd asked him to "refrain from getting

fresh species named" until he had corresponded with Turner on

the matter. If the insect was indeed rare, Dodd could only get

full credit for the discovery after Turner's identification. More

often, to his regret and frustration, he confessed to Lyell that he

carelessly sent away unnamed things, perhaps

some reared with great care + trouble, to find

some day that they have been described from

"Somebody" collection, having been taken or

bred in Queensland!! This is very annoying to

me. I have bred more Zeuzeridae than anyone

living or dead, and my name is never mentioned

in connection with the new things. Also I have

bred more Charagiae [Aenetus] than anyone else.

As to Xylos I have bred over 50 species in north

Q’land, including those I got at Charters Towers

years ago, perhaps 60 species. I don’t know how

many species I bred in south Q’land (National

Museum of Victoria, 26 November 1900).

Clearly, Dodd resented the anonymity that came with living

so far away from the centres of entomological research. Given

the toil of collecting rare and elusive insects, this was

understandable. Why should others, often wealthy purchasers,

be given the credit for his work? This sometimes made him

quite vain about being acknowledged for the discoveries he had

made. In a letter of June 1898, he followed up with Lyell on one

of his moths: "Oh, when getting that beautiful Oecophoridae

named did you credit me with breeding it? I have one left and

Dr Turner was charmed with it” (National Museum of Victoria,

9 June 1898). At other times, however, he begged Lyell to

conceal information about what and where he was collecting.

The wish to be acknowledged for the very details he needed to

conceal reveals the painful bind in which he found himself

(Monteith, personal communication, 6 July 2015).

Lyell understood his correspondent's dilemma and was

happy to observe the friendly protocol of sending all queries

about names to Turner before consulting anyone else. In

return, Dodd made a point of impressing on Lyell how much

he trusted him. For instance, Dr Turner was, he told Lyell,

anxious to name one of his insects as new but he only had two

specimens, one of which he had sent to Lyell. "No other

collector would have done that", he declared. Nevertheless,

Dodd's inability to name insects scientifically remained

something of a sore point, leading him many years later to

pronounce, defensively, to J. A. Kershaw:

It is too severe a mental strain for a professional

collector to attempt to learn the names of the

thousands of insects which pass through his

hands, so I seldom can supply names . . . Years

ago I had a long list of names of Lepidoptera,

but lost it in the Townsville cyclone. I could not

replace it, a busy life of collecting has prevented

my endeavouring to make another.

Dodd's boast was that he possessed other skills, such as

his detective work in locating and hatching out insects, then

setting them perfectly, reminding Kershaw at the end of his

letter: "I can send nicely set bugs ants &c &c 1f the Mus[eum|

cares for same" (National Museum of Victoria, | June 1911).

Building Trust

Dodd’s first letter to Lyell, dated 9 September 1897, enquired if

he might be included in his circle of “exchanging

correspondents” (National Museum of Victoria, 9 September

1897). Dodd had often seen Lyell’s “nicely set insects ... perfect

in every way" in Queensland, so he was confident that their

exchanges would not just be rewarding but (even more

importantly) equal. A perfectionist, Dodd often complained

about the low standards of others. In fact, he told Lyell he did

not want Melbourne entomologists to know that he had a large

number of insects for exchange, his reason being that "several

of them do not set well enough to please me - as a rule I get

inferior material to my own”. Kershaw, for instance, had proved

disappointing, palming off on him "faded, damaged, or

common specimens’ for the “rare or beautiful things" he had

been sent. Lyell was probably sympathetic; like Dodd, he was

proud of his meticulous standards of preservation and

mounting. Notably, although Barrett would later describe

Lyell’s gift to the National Museum as unconditional, there

was, in fact, one condition. When it came to amalgamating his

collection with the Museum’s, Lyell stipulated that his own was

to take precedence. While anything worth saving from “the

smaller and poorer old museum collection” would be remounted

for inclusion, he demanded that most of the museum’s

specimens be scrapped (Melbourne Museum, Archive Box 579,

27 December 1934; Lyell to Robertson).

Despite Lyell’s meticulously high standards, Dodd soon

found cause to chastise him for the arrival of 26 damaged

specimens, including some with “one or both antennae broken

and missing’. Suddenly, the mutual trust and reciprocity that

was to cement their relationship was threatened. The

geographical balance and complementarity that Dodd was

seeking between Victorian and Queensland specimens was

also jeopardised:

In looking through the exchange lately received

I regret to notice that many of the specimens are

imperfect, or rubbed, and others seem rather

old. You may remember my request for clean

and perfect insects, and I trust future lots will be

a great improvement on this ... I have a large

collection and the condition of same is first

class. I keep no damaged specimens that I can

replace with perfect ones and I am sorry to say

that my Victorian collection compares very

176

poorly with my own; in fact they spoil the

appearance of the others. Therefore I want no

more damaged things and no matter how rare a

species 1S, please do not send it to me unless

perfect in every particular. I cannot prize a thing

with a great gap or gaps in its wings, antennae

missing, or badly rubbed &c &c. Should you

care to continue these exchanges may I hope

that you will kindly place me upon your first.

class list ... (National Museum of Victoria,

12 February 1898).

Behind Dodd 's fastidiousness lay the fear that Lyell was not

taking him seriously enough. Perhaps Lyell regarded him as a

mere collector? This anxiety is evident in the way in which the

adjective first class shifts in this paragraph, from the insects in

Dodd’s collection to the list of Lyell’s correspondents. Keen to

reinforce his standing as a first-class collector with a first-class

collection, he reminded his Melbourne correspondent of the

abundance he enjoyed as a tropical collector. Unlike many who

were obliged to capture “almost everything that flies", he had

the geographical advantage of refusing hundreds when out in

the fields. The spectre of being branded as a mere collector,

isolated from professional networks, also prompted Dodd to

say that, should anyone enquire of a particular insect, then

perhaps you will have it mentioned they are in

collections Lyell et Dodd, not taken by me as if I

was a mere collector. I fear there is an

impression abroad that I want specimens for

others and not myself and that that is why I

“haggle” for only first class specimens, should

therefore you have any such impression pray

dismiss it.

In other words, Dodd wanted it to be known that, instead

of collecting insects solely for exchange, he was proud to

retain many of the most perfect and beautiful insects for his

own collection (National Museum of Victoria, 16 July 1898).

This early rupture was soon healed when Dodd received a

parcel whose contents were "perfect in every way”. Writing to

thank him, Dodd made no apology for being fastidious. In

fact, as if to underline the absence of any apology, he added

bluntly that Lyell’s new parcel contained “several common

things" he did not want. These he would return straight away.

Unable to resist reiterating the point about tropical abundance,

he had to concede, in fairness, that he was able to catch more

in 24 hours than Lyell could catch in five days, but that

discrepancy did not mean he had to accept "poor or broken

things from a Victorian collector". Nevertheless, the

relationship was important to him so that meant an on-going

commitment to the delicate task of building trust and

reliability. When Lyell received a rare and expensive butterfly,

Ogyris genoveva Hewitson, and one of its antennae was

missing, Dodd insisted that the specimen was absolutely

perfect when despatched from Cairns; nevertheless, he

immediately sent a damaged specimen with "one good

antenna to replace the lost one" (National Museum of

Victoria, 26 October 1900, 26 November 1901). As part of the

D. Coleman

rules governing their exchange, Dodd also assured Lyell that

he could return anything he did not want for credit.

This gentlemanly exchange of first-class specimens

continued for another couple of years until August 1900 when

Dodd informed Lyell that altered circumstances meant he was

unable to continue indulging in this pleasurable pastime. With a

erowing family and a costly relocation from Brisbane to

Townsville — a place of “higher prices for everything" — he must

now leave off his gentlemanly pursuits (National Museum of

Victoria, 27 August 1900). To Kershaw, whom he hoped would

purchase insects for the National Museum of Victoria, he wrote

that instead of gracing his friends’ cabinets, he must now look

upon his "beloved specimens ... from an £8 point of view”.

“Most of my best things are reared”, he boasted, adding "I keep

no rubbish, and no one need fear at any time that I will victimize

my correspondents". Despite the taint of trade — making dollars

— Dodd nevertheless insisted that his business was an honourable

calling. Furthermore, he would keep himself at arm's length

from commerce by employing an agent (National Museum of

Victoria, 17 September 1900, 31 May 1901). By 1904, he

declared that "business with the dealer fraternity is so

unsatisfactory" that he was badly 1n need a larger pool of reliable

correspondents. Dispensing with middlemen, he now preferred

to deal directly with collectors and museum personnel (National

Museum of Victoria, 4 May 1904).

Once Dodd had turned commercial dealer, he encountered

a host of rules and protocols governing selling, buying and

eifting. These activities were linked, not just to questions of

honour and trust between correspondents, but to perceptions

of social class and educational background (Lucas, 2013).

There was, for instance, the delicate matter of promising

certain correspondents first option on rare or large insects. So

great was the offence if these sought-after insects were

subsequently seen in others’ collections that, if Dodd was

planning to initiate a new contact, he would first ask

permission from his established correspondents. It was also

bad form to share the secret of a special location without

seeking permission. For instance, Dodd claimed to be the first

to let his mentor Rowland Illidge into the secret of where to

find two species of the extremely valuable Aenetus (A.

ramsayi Scott and A. lewinii Walker) outside Brisbane. To

Lyell he divulged: “I was the first 1n Queensland to find and

breed these and at once informed Illidge and we several times

went out together. | went away for 12 months + he took

[Reggie] Relton into ‘mateship’ without consulting me”.

Illidge had also found Aenetus exuviae when out in the bush

on his own and not let on to Dodd about his discovery. Finally,

Dodd liked a correspondent to give full details of unusual

insects. When it came to conveying such information, Oswald

Lower, a pharmacist in Broken Hill, was one of the very slim

ones, he complained. Lower also offended Dodd by lacking

an eye for beauty. The closest he got to praising Dodd's

settings, or a particular insect, was to ask him to "send another

pair”. Instead of such obliquity, Dodd preferred directness. He

liked correspondents “to express pleasure over a beauty or a

rarity’: “I get quite cross when I send away a lovely thing if

the receiver does not ‘enthuse’ a little”. Lyell must have

remonstrated over these complaints about Lower and others

George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-century Australian entomology 177

because Dodd conceded: "You are right, biz is biz and I

should get all I can. As a seller it matters nothing to me what

the buyer 1s like, I suppose (National Museum of Victoria, 12

August 1901, 6 July 1904).

As a commercial dealer Dodd touted his insects via

several selling points. Rarity, beauty and large size were chief.

So too, as we have seen, was endorsement by an expert

classifier like Turner. The other selling point was his talent as

an insect breeder. When circulating sale lists with prices,

some pages contained a banner heading stating that all

specimens had been bred. In particular, he was proud of his

high-priced things, such as his Xylos, stuffed wood-boring

moths, bred and reared by him. In his eyes these Xylos were

perfect in every way. With every skerrick of fat scrupulously

removed, they were (he boasted) very unlikely to turn greasy

(National Museum of Victoria, 17 September 1900). Dodd's

letters also contain detailed descriptions of his breeding

routines. Sometimes he would watch all day until late at night

for a large and rare wood-boring moth to hatch. Then, sleeping

only for a few hours, he would wake to find his vigilance had

been in vain: his valuable moth had emerged and rubbed its

wings. Moths in the "restless" but "handsome" family of

Notodontidae were particularly prone to offend in this regard,

often emerging after he had retired (National Museum of

Victoria, 24 March 1901). It was necessary to keep vigil 1n the

bush as well, tending the larvae of insects for years before

cutting and transporting the timber home for closer

monitoring on the eve of emergence. Sometimes, to Dodd’s

chagrin, local aboriginals ate the grubs he was watching over.

That these grubs were a traditional and highly nutritious food

source for Indigenous people cut no ice with Dodd. He was

always very testy in his letters when mentioning this so-called

theft of his livelihood. His son Walter D. Dodd (1891-1965),

also a naturalist, understood better than his father the

symbiosis between Indigenous people and country. From

Walter Dodd’s observations, it was clear that customary ways

of living on country entailed a balanced economy of nature.

Writing about some very large wood moths he had caught

south-east of Perth in 1912, he noted that “The blacks were

very fond of ‘the grub’”. Captive to the prevailing discourse

surrounding the “inevitable dying out” of the Aboriginal race,

Walter Dodd added that since indigenous people had become

extinct 1n certain localities, ^whole patches of wattle forests

were laid low, there being no check upon the breeding of these

insects" (The North Queensland Register, 13 April 1935).

The highest priced moth Dodd ever offered for sale was

Aenetus mirabilis Rothschild, a species found only in north

Queensland (fig. 4). His excitement at sourcing this large moth

can be heard in his warning to certain favoured correspondents

to "save up your pennies" for a pair (National Museum of

Victoria, 26 November 1900). He had read about this moth 1n

a Journal article published by Walter Rothschild 1n 1894, in

which the location was disclosed as Cedar Bay, North

Queensland. This was a very remote spot 40 kilometres south

of Cooktown and accessible only by boat or by foot

(Rothschild, 1894). Dodd spent weeks and “a great deal of

time and trouble" searching for this moth, at a cost of over

£60, so he was obliged to charge highly for it. He assured

Lyell, one of the first collectors he contacted regarding his

precious discovery, that A. mirabilis was unlikely to be found

in private collections, or even in the British Museum or any

Australian museum “for many a long day, unless through me”.

As for Lyells hint that the Victorian Government's

entomologist Charles French (1842-1933) owned an A.

mirabilis, Dodd was dismissive, imagining that it must be a

damaged specimen, not perfect like those now in his

possession. He asked that Lyell keep quiet about his discovery

as demand for specimens was going to be strong, and Dodd

wanted to prioritise overseas collectors because they would

pay considerably higher prices than those fetched in Australia.

At first, Dodd asked Lyell for 60/— to 100/— [£3—£5] per

pair for A. mirabilis. This approximates to a price range of

£350—£585, or AUD $660-$1100 in today’s purchasing power,

the wide range reflecting the size, condition and appearance

of the insects. Dodd believed this price range was fair and

reasonable. His justification lay in the relative pricing of

Aenetus ramsayi, a species of Aenetus that he had managed to

sell to English collectors for £4—£6 per pair, even though this

moth had been known for a long time and was well distributed,

unlike the rare and “very fine” A. mirabilis. A. ramsayi was

also smaller than A. mirabilis, the female of which averaged

wing spans of 6.5 inches. In the end, Dodd settled on a lower

quotation of 55/— to 75/— per pair for Lyell, reducing the cost a

little because he realised his friend would find it impossible to

obtain such rarities if he “stuck at their money value”. He

asked Lyell not to mention this discount as he also planned to

sell to his competitors (i.e. Australian collectors he disliked

such as Lower, a member of the questionable dealer fraternity;

National Museum of Victoria, 26 November 1900). Charles

French was another he distrusted, telling Lyell that French

had the nerve to ask for his precious Xylos but still hadn’t sent

on the beetles he owed him (National Museum of Victoria, 17

March 1901). Lyell, on the other hand, belonged to Dodd's

category of reliable correspondents. In fact, so reliable was

Lyell that when, a few months later, wealthy clients had failed

to send on remittances, Dodd asked him to pay up-front

before he had even despatched the specimen box (National

Museum of Victoria, 4 February 1901). He also asked that

Lyell pay him a little every month to help keep him afloat

between transactions.

When it came to selling his A. mirabilis, Dodd pitched his

highest price to Lower — £5 for a single pair. In the end, he

sold him a pair for £4, a price that he claimed pleased Lower

very much. Dodd himself was happy with the transaction.

Given Lower’s wide network and authority in the field, this

sale would prove a good advertisement for Dodd’s insects (13

June 1901). In general, Dodd figured that it made good

business sense to sell his A. mirabilis cheaper to Australian

correspondents because this would advertise the perfection of

his preservation techniques. Despite this, Dodd held himself

aloof and was never less than testy with most of the dealer

fraternity, believing that his southern brethren were all down

on him because, after 1900, he would only sell and not

exchange his grand tropical species for anything they could

supply (National Museum of Victoria, 12 August 1901).

178

Conclusion

In July 1910, G. A. Waterhouse travelled from Sydney to the

Atherton Tableland for a week of collecting. Staying in the

Kuranda Hotel, he spent a good deal of time with Dodd and his

family who were warmly hospitable. From here, Waterhouse

wrote to Lyell, describing his first impressions of Dodd. There

was much to like about the man. He was clearly an immensely

enthusiastic collector who generously lent his sons to visitors

like himself for collecting trips. He was pernickety, though, a

charge that Dodd would have been proud to acknowledge. In

running his business, however, Waterhouse declared him

unmethodical, with barely one per cent of his pinned insects

labelled with dates. Where there were dates, he suspected that

they were a “mere matter of recollection’. His classifications

were sloppy too, with “similar groups of insects ... mixed up

anyhow” (National Museum of Victoria, AB 369; 17 July 1910).

From Dodd’s correspondence with the English dipterist Colbran

Wainwright in the same year as Waterhouse’s visit, this

somewhat cavalier attitude about details can be seen in his

Figure 4. Aenetus mirabilis Rothschild (female).

D. Coleman

postscript to one specimen box: “The localities are roughly

marked off on the lids of the cigar boxes. The next lot will be

arranged better, and month of capture given” (Royal

Entomological Society, 24 August 1910).

As Waterhouse’s week in Kuranda neared the end, he

informed Lyell that Dodd’s “tourist business brings the most

money’ meaning that “attention to scientific detail suffers in

consequence’ (National Museum of Victoria, AB 369; 17 July

1910). By tourist business, Waterhouse was referring to Dodd's

recent commercial initiative: the opening of his house to

members of the public for an entrance fee. Before 1910, Dodd

had always welcomed visitors curious to view his collections

and see him at work, setting and preserving his specimens, but

with entomology a full-time business for supporting his large

family, he decided to charge a fee for this. Here we see him in a

three-piece suit, posing in his garden with a butterfly net for one

of his paying tourists (fig. 5). Dodd was quite the showman in

this new business venture, an empresario who used special

lighting effects and other tricks to impress his visitors with the

mystery and beauty of his insects, turning his home into a semi-

George Lyell and Frederick Parkhurst Dodd: authority and expertise in nineteenth-century Australian entomology 179

- -— b E

Figure 5. F. P. Dodd in his garden at Kuranda. Image courtesy of Queensland Museum.

180

public commercial site and museum. Into sober, scientific

scrutiny, he injected an older element of spectacle and magic,

performing his insects with dramatic exhibitions which mingled

natural science, wonder and magic. This was the case even when

his visitors were some of the top people in the field, such as

Walter Baldwin Spencer and A. J. Gilruth, who visited in 1911.

Margaret Fountaine (1862-1940), a globe-trotting British

lepidopterist who had recently settled with her lover not far

from Kuranda, left her own account of calling to see Dodd's

collection in 1916. Waiving the usual admittance fee of one

shilling each, Dodd greeted them heartily as fellow

entomologists, and they had a delightful time taking tea and

revelling in his exhibits. Fountaine was impressed by Dodd's

knowledge of Britain’s scientific scene. He had read her articles

in the Entomologist and in the Transactions of the

Entomological Society of London. After talking entomological

shop for a while, Dodd tried, with a “shrewd, penetrating”

look, to dissuade Fountaine from thoughts of farming in the

area. He argued, as others had done, that her prospects of

Success were dim. As she left, she invited him to come over

and see her Malay and Java butterflies, an idea that seemed to

delight him, but it would not be for a week or so (he claimed)

because he was so busy. Fountaine’s diary entry concludes:

We both liked Mr Dodd very much, and I

believe the old man wishes to be a good friend

to us, especially as he sees that we are not going

to be in any way rival dealers, which naturally,

as he makes his living out of this business, he

could not be expected to look upon with any

favour (L. Joanne Green, personal

communication, 30 June 2019).

Fountaine’s own eye was shrewd and penetrating in

concluding that their reception would not have been so

friendly had they planned to set up as business competitors.

We have seen Dodd move from exchange to commerce,

transacting a business in which there were no fixed prices to

guide him in determining the value at which he should trade a

specimen. Beauty, colour, size and rarity were key

determinants of cost, but perhaps most important was the

symmetry and neatness of the setting, allowing the specimens’

natural attributes to be seen. Since he was well known for his

personal skill in preserving specimens, he believed his

reputation in this regard entitled him to charge extra. At first

Dodd is uneasy about his new dealer persona, embarrassed to

be treating his insects as commodities instead of exchanging

them for pleasure. He also worried, at the outset, that seasoned

dealers and collectors like Lower would see him as green and

try to take advantage of him. But in general he regarded trade

as honourable. He needed to get his insects out through the

proper channels and did not see any of this as injurious to the

pursuit of science. In sum, he was well above J. O. Westwood's

definition of the very "lowest class of entomologists”, a

collector with inert drawers of “handsome insects”.

Despite Dodd's many promises to Lyell, Kershaw and

others that he would start to be more methodical in preserving

dates and locations, as well as keep lists of his insects, he kept

on failing to do so. As for keeping a collection of named

D. Coleman

specimens to guide him, this would not only be too cumbersome

but also beyond what he could afford (National Museum of

Victoria, | June 1911). It is difficult to determine if his

shortcomings regarding taxonomic identification. negatively

affected his pricing. Although many of the letters contain lists

with prices attached, we see him in his correspondence

attempting to establish equivalencies and differences in value

from one specimen to the next. Isolation, doubts about

identification and the difficulty of knowing for certain 1f one's

captures were rare were all inhibiting factors. The best he could

do was to offer the insects in as perfect a condition as possible,

along with close observation of their life histories. How did he

classify himself? In the end, Dodd saw himself, not as an

entomologist but as a professional collector, training up his four

sons to be useful, all-round collectors across the various orders

(National Museum of Victoria, 6 December 1912). One son,

Alan Parkhurst Dodd (1896-1981), would in fact become a

distinguished entomologist, collecting and importing live

specimens of the Cactoblastis moth from Argentina to destroy

the prickly pear that had spread over millions of acres of

Queensland farmland. In Alan, F. P. Dodd's legacy lived on.

Acknowledgements

My thanks to Dr Geoff Monteith, entomologist at the Queensland

Museum, for his generous and expert annotation of my

transcription of the Dodd—Lyell correspondence in Melbourne

Museum’s archives. Thanks also to L. Joanne Green for sharing

her transcriptions of Margaret Fountaine’s diaries, held in

Norwich Castle Museum, UK, and to Geoff Thompson at

Queensland Museum for speedily supplying the image of Dodd

in his Kuranda garden. My collaborators at the Melbourne and

Australian Museums, archivists Nik McGrath and Vanessa

Finney, and entomologist Simon Hinkley, must also be thanked.

This paper has been funded by the McCoy Seed Fund, a

collaboration between the University of Melbourne and

Museums Victoria, and the Australian Research Council Linkage

Grant, “Merchants and Museums’ (LP160101761).

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Memoirs of Museum Victoria 80: 183-191 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021.80.10

Parasitoids of the uniquely social colletid bee Amphylaeus morosus (Hymenoptera:

Colletidae) in Victoria

Lucas R. HEARN”, MARK I. SrEvENS^?, MICHAEL P. SCHWARz! and BEN A. PARSLOW?

! College of Science and Engineering, Flinders University, GPO Box 2100, Adelaide South Australia 5001, Australia

? Earth and Biological Sciences, South Australian Museum, GPO Box 234, Adelaide South Australia 5001, Australia

? School of Biological Sciences, University of Adelaide South Australia 5005, Australia

'" Corresponding author: Lucashearn7? gmail.com

Hearn, L.R., Stevens, M.I., Schwarz, M.P. and Parslow, B.A. 2021. Parasitoids of the uniquely social colletid bee

Understanding how nest parasites contribute to brood mortality rates 1n host species 1s an 1mportant step towards

uncovering the potential implications for host behaviour. This can be especially important for understanding the evolution

of social living, where defence against parasites is often posited as a major benefit of cooperative nesting. Only two

parasitoid species have previously been reported for the only known social colletid bee, Amphylaeus morosus: the

ecasteruptiid wasp, Gasteruption primotarsale, and the mutillid, Ephutomorpha tyla. Here we report six additional

parasitoid species of A. morosus: the gasteruptiid wasps G. atrinerve, G. globiceps, G. melanopoda and G. cinerescens;

the bombyliid fly Anthrax maculatus; and the mutillid wasp Ephutomorpha aff. varipes. The mechanisms of parasitism

for these eight parasitoid species are described in combination with how they operate throughout the host brood rearing

Abstract

Amphylaeus morosus (Hymenoptera: Colletidae) in Victoria. Memoirs of Museum Victoria 80: 183—191.

period and whether benefits of social nesting vary across the season.

Keywords Bombyliidae, Gasteruptiidae, host-association, Hylaeinae, Mutillidae, parasitism

Introduction

Detailed host and parasite association data are important

yet rarely explored for primitively social hosts, where

defence against parasites is posited to be a driver of

cooperative nesting. For insect parasitoids, the value of this

information is often emphasised but is heavily weighted

towards key taxon groups, such as Ichneumonidae,

Braconidae, Chalcididae, Encyrtidae, Tachinidae and

Bombyliidae, that are of economic importance (Heraty,

2017). Entomophagous insect associations with non-apid

bees largely get overlooked but can have substantial

consequences for brood mortality, behaviour and

demography of host species (Segers et al., 2016; Smith et

al., 2017). Additionally, parasite—host interactions in bees

are seldom observed over their full life history, meaning the

full series of parasites attacking a single host is often

understated. In the small carpenter bees Ceratina dupla

Say, 1837, and C. calcarata Robertson, 1900, Vickruck et

al. (2010) reared a total of eight parasite species from brood

cells after sampling nests across a six-month period. Wcislo

et al. (2004) similarly sampled nests of the halictine bee

Megalopta genalis (Meade-Waldo, 1916) across one year

and reported four brood parasites infiltrating nests.

However, Smith et al. (2017) noted that the full effect these

brood parasites may have on the social nesting of M. genalis

is hard to fully assess due to insufficient field data. For

analogous species, the need for descriptive biological

information combined with rigorous sampling across the

colony life cycle is often needed to determine the true

diversity and effect of brood parasitism.

The Australian hylaeine bee, Amphylaeus morosus

(Smith, 1879) (Colletidae: Hylaeinae), is the only species in

the hyper-diverse bee family Colletidae that is unambiguously

known to exhibit social nesting. Social nesting in this species

is very simple, with no known reproductive hierarchies or

morphological differentiation between nestmates. Amphylaeus

morosus has a semelparous and univoltine life cycle and may

form social or solitary nests within the same population

(Spessa et al., 2000). In the Dandenong Ranges, A. morosus

uses naturally abscissed dead fronds of the rough fern tree

Cyathea australis to construct linear nests out of the pithy

interior. Like many hylaeine bees, A. morosus mass provision

cells with a mixture of pollen and nectar before ovipositing

into the cells.

spessa et al. (2000) first reported a species of

Gasteruption wasp, later identified as Gasteruption

primotarsale Pasteels, 1957 (Gasteruptiidae) (Parslow et al.,

184

2020), attacking nests of A. morosus in the Dandenong

Ranges, Victoria, while a species of mutillid wasp,

Ephutomorpha tyla Hearn, Williams & Parslow, 2019

(Mutillidae), was recently reported entering nests late in the

brood rearing season (Hearn et al., 2019). Spessa et al. (2000)

found that social nesting in A. morosus seemed to provide

only minimal benefits in reducing parasitism by G.

primotarsale. However, that study did not consider possible

defence benefits against other parasites, and indeed no other

parasites were identified in that study. Gasteruption

primotarsale parasitises the earliest brood during nesting

period and may therefore only explain potential benefits of

social nesting during the early stages of brood provisioning.

Additionally, observations of E. tyla adults in the nest

towards the end of brood-rearing (Hearn et al., 2019) suggest

that E. tyla may exploit the mature larvae of A. morosus

towards the end of its nesting period. It is important to

understand the full suite of parasites that attack A. morosus

because if these parasites operate at other times in the nesting

period, that information will further our understanding of

why social nesting in A. morosus persists until brood

maturation. In south-eastern Queensland, Houston (1969)

reported a range of parasites and parasitoids attacking A.

morosus nests, including the wasps Agamerion pulchra

(Pteromalidae: | Cleonyminae), Coelopencyrtus | spp.

(Encyrtidae), Gasteruption sp. (Gasteruptiidae) and an acarid

mite. However, in the Dandenong Ranges, little is known

about the full range of parasites that might be involved.

Here, we identify eight different parasitoid species that

attack A. morosus, including novel host records for the

Gasteruption wasps G. atrinerve (Kieffer, 1911), G. globiceps

Pasteels, 1957, G. melanopoda Pasteels, 1957, and G.

cinerescens Schletterer, 1885, the bombyliid fly Anthrax

maculatus Macquart, 1846 (Diptera: Bombyliidae), and the

mutillid wasp Ephutomorpha aff. varipes (André, 1895). We

examine the specific mechanisms of parasitism and eclosion

timings of these parasitoids, which have been poorly

understood. It is important that the respective host associations

are well documented to fully recognise how these multiple

parasitoid species may influence the nesting behaviour of the

only known social bee in the colletid family.

Materials and methods

Nest collections

Nests of Amphylaeus morosus were collected from naturally

abscissed fronds that haphazardly fall around the base of

the rough tree fern Cyathea australis R.Br. Domin. These

nests were collected in the Gembrook, Cockatoo and

Marysville regions of the Dandenong Ranges, Victoria,

Australia. Nests were sampled over five collection periods:

4—6 December 2017, 6-8 December 2018, 21—22 November

2019, 11 January 2020 and 27 February 2020. The collection

region primarily consists of cool temperate montane habitat

dominated by tall canopies of Eucalyptus regnans F. Muell

and E. viminalis Labill, with a mixture of ferny, shrubby

and grassy understories. Nests were collected early in the

morning, late in the afternoon or during periods of light

L.R. Hearn, M.I. Stevens, M.P. Schwarz & B.A. Parslow

rain when bees were inactive to ensure all colony members

were present. Nest entrances were sealed upon collection

and all nests were stored in cool insulated boxes and

transported to the laboratory where they were stored at

10°C until processed.

Parasitism data

Nests were dissected lengthwise and the contents, including

adult female A. morosus, immatures and nest parasitoids,

were recorded. Where possible, parasitised cells were

carefully extracted from the nests and left to develop in Petri

dishes at ambient room temperature. Petri dishes were kept

moist by occasionally spraying filter paper rectangles inside

each dish with water. Once the specimens had reached

adulthood, defined as fully mature and mobile, they were

placed in 99% ethanol for preservation and later identification.

For parasitoids, where the pupal stages form cocoons (e.g.

Mutillidae), at least one pupal case from each nest was

dissected in coordination with the nest being opened to

determine the developmental stage of each immature.

Not all the Gasteruption specimens were successfully

reared to pupae or adulthood because developmental rates for

Gasteruption species lagged significantly behind host

maturation rates; due to this, the specific parasitisation rates

for each Gasteruption species were unable to be determined.

Further, the scarcity of larval descriptions based on

morphology for the genus Gasteruption made it impossible to

discriminate species at the larval stage (Bogusch et al., 2018).

Therefore, the parasitisation rates for each species are

combined and treated at the genus level Gasteruption.

Specimen identification

Specimens were examined using a Nikon SMZIOO0

stereomicroscope at the South Australian Museum, Adelaide,

Australia. Images were taken using a Visionary Digital LK

imaging system (Dun, Inc.) with a Canon EOS 5DsR camera

at Flinders University, Adelaide, Australia. Images were

produced using Zerene Stacker (Zerene Systems LLC)

software and cropped and resized in Photoshop CS5.

Material examined in this study were initially identified

using the following keys: Amphylaeus morosus specimens were

identified using Houston’s (1975) key to Australian hylaeine

bees; Gasteruption atrinerve, G. globiceps, G. melanopoda, G.

primotarsale and G. cinerescens were identified by B.A.

Parslow (South Australian Museum) using Pasteel’s (1957) key

to the Australian Gasteruption and comparison with type

material; Ephutomorpha tyla specimens were identified by

L.R. Hearn by comparing specimens with type material held at

the South Australian Museum; and Anthrax maculatus was

identified by X. Li (Florida Museum). Voucher specimens for

the examined species have been deposited at the Melbourne

Museum (Table S1).

DNA sequencing for the eight parasitoids of A. morosus

was performed by the Canadian Centre for DNA Barcoding at

the Biodiversity Institute of Ontario using standard protocols

(Ivanova et al., 2006; Table S1).

Parasitoids of the uniquely social colletid bee Amphylaeus morosus (Hymenoptera: Colletidae) in Victoria 185

Table 1. Prevalence of parasitoid species 1n host nests and brood cells at each collection period. Mean parasitised cells calculated from parasitised

nests only”.

Nest collection Parasitised nests

Year Total Total Gasteruption Anthrax Ephutomorpha Ephutomorpha

nests cells maculatus tyla

Earlysummer 85 400 16 0 9

2017

Early summer 6 29 0 2 0

2018

Late spring 27 | 124 7 0 V

2019

Mid-summer 26 109 l f 2

2020

Late summer 16 78 3 0 4

2020

Total 160 740 27 4 15

Results

Nest architecture and composition

In the Dandenong Ranges, females of A. morosus construct their

nests in dead abscissed fronds of the rough fern tree Cyathea

australis by excavating the pithy interiors of the stems to create

an unbranched linear tunnel. Nests were on average 212.63 +

59.39 mm long (mean + standard deviation; N = 156) with nest

entrance diameters of 5.65 + 0.61 mm (N = 52). In other regions,

nests also occur in the dead flower scapes of Xanthorrhoea spp.

(L. Hearn, personal observation). The tunnel is lined with a

cellophane-like material, a characteristic common to hylaeine

bees, that is thought to act as a waterproofing and antimicrobial

agent (Almeida, 2008). Provisioning of brood cells generally

begins in mid-spring, and cells are laid sequentially starting

from the distal end of the fern frond towards the proximal end.

Brood production can last until mid-summer across the

Dandenong Ranges populations, allowing different parasitoid

species to attack at different brood development stages (fig. 1).

Nest architecture in A. morosus can be broadly categorised

into nests with brood cells interspersed with vestibules and

nests with brood cells laid sequentially (fig. 2). Vestibules are

empty spaces containing no provisions or brood and are

common in many stem-nesting Hymenoptera (Danks, 1971).

In A. morosus, vestibular cells are frequently found 1n solitary

nests, but nest burrows are fully excavated before the first cell

is provisioned. This limitation to nest length once cell

provisioning has begun suggests that there may be a trade-off

between the number of potential brood cells and constructing

vestibular cells to deter nest parasitoids.

Host associations

Gasteruption Latreille, 1796 (fig. 3a). Gasteruption 1s the

most abundant parasitoid of A. morosus in terms of nests

parasitised, parasitising 16.9% (27/160) nests across all

collection periods (Table 1). Of the nests parasitised by

Gasteruption, only the first three brood cells were found to

contain parasitoid immatures. Gasteruption parasitised cells

were often followed by vestibular cells (11/35 parasitised cells),

Parasitised cells (total/mean)*

Gasteruption Anthrax Ephutomorpha Ephutomorpha

aff. varipes maculatus tyla aff. varipes

0 22/1.38 0/0 37/4.11 0/0

0 0/0 7/3.5 0/0 0/0

0 9/1.29 0/0 0/0 0/0

l 1/1 15/7.5 4/2 8/8

0 3/1 0/0 26/6.5 0/0

1 35/1.29 22/5.5 67/4.47 8/8

detritus (9/35) or provisioned cells that failed to develop (4/35).

The larvae of Gasteruption are predator-inquilines in the

nests of A. morosus where they kill the host egg or developing

host larva before consuming the entire host provisions

(Bogusch et al., 2018; Parslow et al., 2020). They then construct,

using anal secretions, dark semi-cocoons above and below the

larvae separating the parasitised cell from the rest of the nest

(Eardley and Daly, 2007; Malyshev, 1968; Westrich, 2018).

Although some species of Gasteruption have been reported to

consume the contents of multiple adjacent cells (Donovan,

2007; Malyshev, 1968; Westrich, 2018), Gasteruption

immatures were only observed to occupy a single host cell in

our nests of A. morosus.

Adult eclosion of Gasteruption occurred far later in the

season compared with A. morosus (fig. 4). In most cases,

Gasteruption brood were still at a late larval stage at the time

A. morosus cells were reaching adult eclosion. Five species of

Gasteruption were reared from host nests. One of these,

Gasteruption primotarsale Pasteels, 1957, has been previously

recorded from nests of A. morosus (Spessa et al., 2000). The

novel host association for G. atrinerve (Kieffer, 1911); G.

globiceps Pasteels, 1957; G. melanopoda Pasteels, 1957; and

G. cinerescens Schletterer, 1885, were determined using

adults reared from host nests.

Anthrax maculatus Macquart, 1846 (fig. 3d). Four nests were

found to be parasitised by the bombyliid fly Anthrax maculatus

across two separate sampling periods. Anthrax maculatus was

never directly observed ovipositing into or hovering around the

nests of A. morosus. Yet, developing A. maculatus immatures

were found occupying both single and consecutive A. morosus

brood cells, accounting for 24.1% (7/29) and 13.8% (15/109) of

the total brood cells across the early summer (December 2018)

and mid-summer (January 2020) collection periods, respectively.

The pupae of A. maculatus were primarily observed occupying

cells in the middle of the nests, but in one instance, every cell in

the nest contained parasitised immatures. All brood cells

parasitised by A. maculatus contained late-stage pupae when

the nest was opened, and in nests containing both A. maculatus

186 L.R. Hearn, M.I. Stevens, M.P. Schwarz & B.A. Parslow

Ephutomorpha aff. vanpes

= B.

+ =

T i J" d

n a

— ux. Ww

Ephutomorpha tyla

Anthrax maculatus "m ——

Gasteruption

Amphylaeus morosus

Brood

development

Brood

production

Nest founding

Overwintering

Mating

Adult

eclosion

AUTUMN WINTER SPRING. SUMMER

Figure 1. Annual cycle of Amphylaeus morosus across the full univoltine, semelparous life cycle in the Dandenong Ranges, Victoria (grey). The

timings of nest parasite attacks (coloured) were estimated from rates of occurrence in nests across the brood development period.

Figure 2. Nest dissections showing the two typical nesting structures for Amphylaeus morosus in the Dandenong Ranges: a, typical structure of a

solitary nest; b, close-up of brood cells directly followed by a pithy cell plug, interspersed with vestibular cells; c, typical structure of a social

nest; d, close-up of brood cells laid sequentially. Scale bar = 10 mm.

Parasitoids of the uniquely social colletid bee Amphylaeus morosus (Hymenoptera: Colletidae) in Victoria 187

Figure 3. Images of the parasitoids reared from Amphylaeus morosus nests: a, lateral habitus of Gasteruption globiceps; b, lateral habitus of

Ephutomorpha tyla; c, lateral habitus of Ephutomorpha aff. Varipes; d, lateral habitus of Anthrax maculatus e, Ephutomorpha aff. varipes larvae

feeding on A. morosus pupae (photo credit James Dorey, 2020). Scale bar = 1.0 mm.

188

immatures and unparasitised host immatures, the host brood

cells contained mid- to late-stage pupae. The pupae of A.

maculatus were packed into the host brood cells and eclosed to

adults within minutes of being extracted from the brood cell in

the nest. The emergence of adult A. maculatus therefore

appeared to be stimulated by disturbance but still synchronised

with host adult eclosion (fig. 4).

Ephutomorpha tyla Hearn, Williams, and Parslow, 2019

(fig. 3b). The host association for Ephutomorpha tyla has

already been described by Hearn et al. (2019). Here, we present

additional details based on new data. In two nests, a single

adult female E. tyla was observed trying to break through the

most proximal cell in the nest, while in a further 21 instances,

adult females were observed waiting in the nest entrance.

Ephutomorpha tyla does not appear to specifically target

certain nest types, attacking nests with up to 14 brood cells or

nests with only four brood cells interspersed with vestibules. A

total of 29 nests contained either parasitised immatures, an

adult female E. tyla in the entrance, or both. No adult female A.

morosus were present in 69% (20/29) of these nests.

Number of emergences

JANUARY

FEBRUARY

L.R. Hearn, M.I. Stevens, M.P. Schwarz & B.A. Parslow

Parasitised brood cells consisted of brown papery cocoons

that occupied the entirety of the brood cell. As the E. tyla

immatures developed, the brood cells became darker and

increasingly difficult to open. In nine affected nests, all brood

cells were parasitised by E. tyla, with damaged partitions

between cells. The body sizes of male and female E. tyla were

generally consistent. However, in some cases there were

marked differences in body size within each sex.

Ephutomorpha aff. varipes (André, 1895) (fig. 3c). Larvae of

Ephutomorpha aff. varipes were observed as external

parasitoids of the mature larvae, prepupae and pupae of A.

morosus and were observed feeding on young pupae (fig. 3e).

Larvae of E. aff. varipes remained on their host until mature

before producing a light-brown silk-like cocoon and eclosing

as an adult. The ectoparasitoid larva of E. aff. varipes were

observed on multiple consecutive A. morosus immatures in one

nest. The first three brood cells of the parasitised nest contained

fully healthy A. morosus immatures, whereas the next eight

brood cells were occupied by parasitised larvae, all of which

eclosed as males. Ephutomorpha aff. varipes was observed

ame /Amphylaeus morosus

ee œs» Anthrax maculatus

Ephutomorpha aff. varipes

****»*** Gasteruption

SS S o ae oe S

S aS S QoS S

MARCH APRIL MAY

Date reached adulthood

Figure 4. Eclosion phenologies of Amphylaeus morosus and its nest parasites. Emergence observations are only reported for the provisioning and

rearing phase during 2019. Host emergences are only reported for nests containing parasitised brood.

Parasitoids of the uniquely social colletid bee Amphylaeus morosus (Hymenoptera: Colletidae) in Victoria 189

parasitising several species that occupy excised Cyathea

australis fronds, including another mass provisioning bee,

Hylaeus sp., and a crabronid wasp, Pison sp. While, on

numerous occasions, females were observed in the nest

entrance of the allodapine bee, Exoneura robusta Cockrell,

1922, no parasitic association has been confirmed.

Other enemies

Of the 243 potentially viable A. morosus nests collected,

34.2% (83/243) contained no A. morosus adults or brood cells

and had been superseded or temporarily occupied by other

insect species. Twenty-four of these nests were occupied by

Pison sp. (Hymenoptera: Crabronidae), two nests were

occupied by the _ pteromalid wasp of the genus

Eupelmophotismus (Hymenoptera: Pteromalidae), while one

nest contained recently eclosed encyrtid wasps from the genus

Coelopencyrtus (Hymenoptera: Encyrtidae). Five nests

contained clerid beetle larvae (Coleoptera: Cleridae),

including two instances of nests with A. morosus immatures

and adults that contained beetle larvae in the nest entrance.

Discussion

Nest parasitoids are known to have detrimental effects on the

survival and productivity of bees (Vickruck et al., 2010; Wcislo

et al., 1994). Of the parasitoid species that attack A. morosus,

the predator-inquilines of the genus Gasteruption were the

most prevalent, but least detrimental, only parasitising a mean

of 1.29 cells per infiltrated nest, with 16.9% of nests being

infiltrated. We never observed Gasteruption entering nests of

A. morosus, possibly because our nest collections were

restricted to cool weather conditions when insect activity was

low; it is likely that Gasteruption wait for the adult bee to leave

before infiltrating the nest and ovipositing in an open brood cell

(Macedo et al., 2012; Parslow et al., 2020). Parasitisation by

Gasteruption was only observed in single female nests, which

suggests that multifemale nests are guarded by one female

while other individuals collect provisions, restricting the

Opportunity for wasps to oviposit (Parslow et al., 2020). The

extended development time of Gasteruption in A. morosus

nests has been similarly reported in Gasteruption

brachychaetum Schrottky, 1906a, larvae in nests of Hylaeus

aff. guaraniticus (Schrottky, 1906b) (Macedo et al., 2012). The

resulting asynchrony in adult emergence between Gasteruption

immatures and host could be attributed to delaying their

eclosion until hosts have started establishing new nests, which

may explain observations of Gasteruption pupae in

overwintering A. morosus nests (L. Hearn, personal

observation). For Gasteruption, parasitising the distal nest cells

could be a factor of this extended development period and

avoiding being disturbed by emerging bees and other emerging

parasitoids. There is limited information on Gasteruption host

specialisation, with a large proportion of records based on

single observations (Parslow et. al. 2020). It 1s unlikely that A.

morosus 1s the only host for G. atrinerve, G. globiceps, G.

melanopoda and G. cinerescens because their distributions

extend beyond the distribution of A. morosus (Atlas of Living

Australia 2021; Parslow, 2020; Pasteels, 1957).

In contrast, parasitisation by the dipteran Anthrax maculatus

had a greater effect when invading A. morosus nests, parasitising

5.5 brood cells on average across A. maculatus infiltrated nests.

Previous studies have suggested that adult bombyliid females

identify a host nest and oviposit directly into the nest entrance

(Bohart et al., 1960). However, there are also reports of adults

ovipositing haphazardly onto the ground, leaving the planidium

to search for a host nest (Yeates and Greathead, 2008). This latter

scenario could be likely in the Dandenong Ranges given the high

density of A. morosus nests that can occur. It was unknown if the

larvae of A. maculatus feed on both the immatures and provisions

in a cell. Gerling and Hermann (1976) reported early instar

larvae of the bombyliid fly, Xenox tigrinus (Evenhuis, 1984)

(previously Anthrax tigrinus), feeding on the pollen bread in

Xylocopa virginica (Linnaeus, 1771) nests. However, bombyliid

fly immatures are also known to feed on the mature larvae of

their hosts (Felicioli et al., 2017; Minckley, 1989). The delay in

adult eclosion by A. maculatus compared with its host is

consistent with observations by Minckley (1989) of Anthrax

xylocopae (Marston, 1970) delaying emergence to synchronise

with its host. This 1s because adults are not able to break through

the cell partitions and therefore must wait for hosts to emerge

(Gerling and Hermann, 1976). This could explain why A.

maculatus puparium in our current study appeared to be

stimulated by disturbance when removed from their cells.

In our study, Ephutomorpha tyla was the most abundant and

destructive nest parasitoid 1n terms of both the number of brood

cells parasitised and (potentially) adult hosts killed.

Ephutomorpha tyla was rarely observed in nests with an adult

host present, supporting similar reports of mutillids forcefully

removing or killing host guards before ovipositing (Brothers et

al., 2000). Adult E. tyla were generally observed in the nest

entrances and on some occasions attempting to break through

the cell closest to the entrance, suggesting that E. tyla may open

the brood cell closest to the entrance to check for host brood

before ovipositing. The size differences between reared adults of

E. tyla are likely due to differences in the consumed host

immatures. For instance, in A. morosus, the males are generally

smaller than the females (Spessa, 1999). Additionally, the

amount of provisions in each brood cell can vary dramatically,

which can influence the size of the host immatures and

subsequently the ectoparasitoid feeding on it (Brothers, 1989).

Comparatively, the size differences observed in adults of E. aff.

varipes support the notion that 1t 1s likely a parasitoid of multiple

insect hosts; this accords with reports by Mickel (1928) that

variation in body sizes within mutillid species is linked to them

exploiting a range of hosts of varying size. Ephutomorpha aff.

varipes was observed in nests of numerous different host species

including the allodapine bee, Exoneura robusta Cockerell,

1922. However, given Brothers’ Rule (Brothers et al., 2000),

which states mutillids will only attack fully enclosed immatures,

it is unlikely that such a condition is universal given that E.

robusta progressively rears its brood in open linear nests. These

observations suggest that E. aff. varipes 1s a generalist parasitoid

of hosts using Cyathea australis as a nesting substrate, rather

than any specific hymenopteran host, which accords with the

notion that some mutillid species are adapted to specific

situations rather than hosts (Brothers, 1989).

190

Attack by wingless parasitoids such as mutillid wasps is

uncommon in stem-nesting bees (Ronchetti and Polidori,

2020). Michener (1985) suggested that this may be because

predators need to search in a three-dimensional space looking

for sparsely distributed nesting sites, whereas ground-nesting

hymenopteran entrances can be found using a two-

dimensional search pattern. In A. morosus, the high rates of

parasitism might be attributable to the high density of their

nesting substrate around mature tree ferns (Groulx and

Forrest, 2018). This high density of nesting sites may act in a

similar system to the ground nesting aggregations of halictid

bees, where nesting sites are abundant, and parasitism can be

prevalent ( Wcislo, 1996).

The mechanisms of parasitism and bionomy of parasitoids

has generally been poorly understood. Our study finds that

parasitoids of A. morosus have adapted to host nesting

behaviour and suggests variation 1n the oviposition chronology

of each parasitoid species, with Gasteruption wasps ovipositing

at the beginning of brood provisioning, A. maculatus

ovipositing across the middle stages of brood development and

the two mutillid species ovipositing in nests at the end of the

season. This variation in parasitoid pressure could have

implications for understanding the factors driving social

nesting in the only known social bee in the family Colletidae.

Acknowledgements

We thank Terry Houston for his insightful and thorough

comments on this manuscript. This project was supported by

The Holsworth Wildlife Research Endowment & The

Ecological Society of Australia awarded to L.R.H. and in part

funded by an Australian Biological Resources Study National

Taxonomy Research Grant Programme grant RF217-14 to

B.A.P. Nests were obtained under the scientific collection

permit 10008943 for Victoria. Thank you to James Dorey

Photography for imaging of the parasitised bee pupa.

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Table S1. Table showing BOLD sequencing reference number and voucher number for deposited material. All material deposited at Melbourne

Museum.

Family Species Melbourne Museum Voucher number BOLD reference number

Gasteruptiidae Gasteruption primotarsale HY M-97074 OZBOL2593-21

Gasteruptiidae Gasteruption globiceps HY M-97075 OZBOL2607-2 1

Gasteruptiidae Gasteruption atrinerve HY M-97076 OZBOL2600-21

Gasteruptiidae Gasteruption cinerescens HY M-97077 OZBOL2605-21

Gasteruptiidae Gasteruption melanopoda HY M-97078 N/A

Bombyliidae Anthrax maculatus DIP-2590 OZBOL2594-21

Mutillidae Ephutomorpha tyla HY M-97079 OZBOL2596-21

Mutillidae Ephutomorpha aff. varipes HY M-97080 OZBOL2599-2]

Memoirs of Museum Victoria 80: 193-205 (2021) Published 2021

1447-2554 (On-line)

httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/

DOI https://doi.org/10.24199/j.mmv.2021 80.11

Early land plants from the Lower Devonian of central Victoria, Australia, including a

new species of Salopella

FEARGHUS R. McSwEENEY'*, JEFF SHIMETA* AND JOHN ST J.S. BUCKERIDGE '”

! Earth and Oceanic Systems Group, RMIT University, GPO Box 2476, Melbourne VIC 3001, Australia

* Centre for Environmental Sustainability and Remediation, School of Science, RMIT University, GPO Box 2476,

Melbourne VIC 3001

> Museums Victoria, GPO Box 666, Melbourne VIC 3001, Australia

*To whom correspondence should be addressed. Email: tidal75@gmail.com

Abstract McSweeney, F.R., Shimeta, J. and Buckeridge, J.S. 2021. Early land plants from the Lower Devonian of central Victoria,

Australia, including a new species of Salopella. Memoirs of Museum Victoria 80: 193—205.

Early land plants with elongate sporangia held in the palaeobotanical archives of Museums Victoria were examined.

The fossil plants are from Yea ("upper Silurian) and near Matlock (Lower Devonian) in central Victoria, and are of interest

because they contribute to our understanding of the evolution of early land plants in a region in which research has been

limited. Both Salopella australis and Salopella caespitosa were originally described over 30 years ago and this

reinvestigation has resulted in the emending of the diagnosis of Salopella australis and the erection of a new morphotaxon

Salopella laidae sp. nov. based primarily on differing branching architecture and sporangial morphology. Salopella

laidae comes from Yea Formation and possesses regular 1sotomous branching over at least two orders of branching,

terminating in elongate sporangia that are wider than their subtending axes, differing from S. australis, which possesses

only one dichotomy emanating from at least two erect parallel parent axes with sporangia that are the same width as their

subtending axes. A recently collected specimen of Salopella caespitosa was also examined and adds to our understanding

of this taxon, which was previously only known from one specimen. Consideration is given to the possible sources of these

early land plants based on other early land plants with a similar grade of organisation.

Keywords

embryophytes, rhyniophytoids, Baragwanathia flora, Devonian, Victoria

Introduction

Fossil plants have been known in Victoria since the 19th century

(Nicholas, 1875). Palaeobotanical studies on the Baragwanathia

flora (Jaeger, 1966) from the Silurio-Devonian of Australia have

primarily been limited to the first half of the twentieth century

(Cookson 1935, 1937, 1949; Lang and Cookson, 1927, 1930,

1935) and further expanded with Tims (1980), Tims and

Chambers (1984) and McSweeney et al. (2020). The majority of

plant fossils have been uncovered in the Melbourne Zone (fig.

1), and some further fragmentary remains have been found

further south in the Mathinnia Group of northeastern Tasmania

(Baillie et al., 1989; Banks, 1962; Cookson, 1937; Powell et al.,

1993). New data on early land plants from Australia, which was

once part of northeastern Gondwana during the Silurio-

Devonian periods, is of significance globally because it allows

comparisons between floral assemblages in Laurussia (Europe,

the United States and Canada), mid-palaeolatitude (Argentina)

and high palaeolatitude (Bolivia and Brazil) western Gondwana

sites and Chinese sites (North and South China Plates; Torsvik

and Cocks, 2019). Furthermore, the new data allow for more

data to be acrued to test the hypothesis of a Lower Devonian

northeastern Gondwana phytogeographic unit within which both

South China and Australia were parts or subunits of, according

to Hao and Gensel (1995, 1998, 2001) and Wang et al. (2002).

This paper records one new taxon after an investigation of

early land plant material with elongate sporangia collected by

Tims and Chambers (1984) from the ?upper Silurian-Lower

Devonian Yea Formation, the Humevale Formation near Yea

and the Lower Devonian Wilson Creek Shale Formation near

Matlock in central Victoria (fig. 1). Early land plants

possessing elongate sporangia with tapering apices of either

rounded or pointed tips and with fusiform or elongate parallel

sided sporangia characteristic of rhyniophytes/rhyniophytoids

(sensu Edwards & Edwards 1986) were described by Tims

and Chambers (1984) and assigned to Salopella australis and

Salopella caespitosa. The specimens described herein are

compressions/impressions with little anatomical detail

preserved and belong to the same morphogenus Salopella

Edwards and Richardson, 1974. One new specimen of S.

caespitosa, only the second so far recorded, was also

194

examined and is of siginifcance to our understanding of this

morphotaxon because it was preserved in semi-relief as a

coalified compression/impression.

Locality, stratigraphy and age

The Melbourne Zone is a triangular stuctural region occuping

central Victoria bounded by the Heathcote Fault in the west and

Governor Fault in the east, opening towards the south

(VandenBerg and Gray, 1988). The Wilson Creek Shale extends

across the Melbourne Zone (VandenBerg, 1975; VandenBerg et

al., 2006), and based on the conodont fauna, is considered to be

middle Pragian—Emsian (Mawson and Talent, 1994). It comprises

mudstone and shale units, characteristic of deep water and anoxic

conditions, with Uncinatograptus thomasi (=Monograptus

thomasi) Jaeger, 1966, found throughout and Neomonograptus

New South Wales

F.R. McSweeney, J. Shimeta & J.S. Buckeridge

notoaequabilis (=Monograptus aequabilis notoaequabilis)

Jaeger et al., 1969, in the upper half (Edwards et al., 1997, 1998;

Lenz 2013; VandenBerg, 1988; VandenBerg et al., 2006). The

outcrop examined occurs on Frenchmans Spur track,

approximately 10 km west of Matlock, and is the type location of

the rhyniophytoids Salopella australis and S. caespitosa, and the

early "trimerophyte-grade" (sensu lato Banks 1975) plant

Dawsonites subarcuatus (Tims and Chambers, 1984).

The Humevale Siltstone and Yea Formation occur in the

western part of the Melbourne Zone (Edwards et al., 1997;

VandenBerg, 1988) and both are exposed near Yea. The dating

of the exposures at Ghin Ghin Road and Limestone Road, Yea,

as Pridolian and Ludlovian respectively by Garratt and

Rickards (1984, 1987), Garratt et al. (1984), Rickards and

Garratt (1990), and Rickards (2000) are contentious for a

number of workers (Banks 1980; Cleal and Thomas, 1999;

Australia

500 km

"T : | » in.

Victoria 3. = o

2 N XO,

X i | A

> NE Mm

E ' | -NA

Melbeuyrne,

Y Vilsons Promontory

Darraweit Guim Province

Bass Strait

King

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. Flinders

) Island — Mount Easton Province

| d

0 30 100 iz : =<) e Fossil locations

m ^X. Tasmania Clarke Island 1, Yea 2. Matlock

Figure |. Map of Victoria, Australia, showing the fossil locations within the Melbourne Zone. Location |. Yea, Ghin Ghin Road, Limestone Road

(37° 12.38' S, 145° 25.39’ E). Location 2. Matlock, Frenchmans Spur (37° 25.82’ S, 146° 77.24’ E.), the type location of Salopella australis and

S. caespitosa (Tims and Chambers, 1984). Source: adapted from Moore et al. (1998: fig. 2).

Early land plants trom the Lower Devonian of central Victoria, Australia, including a new species of Salopella 195

Edwards et al., 1979; Edwards et al., 1997; Hao and Gensel,

2001; Hueber, 1983, 1992). Hueber (1983) considered the

morphological and anatomical complexity of Baragwanathia

longifolia more akin to a Lower to Middle Devonian age and

noted the complex nature of the conspecific taxa Yarravia

(=Hedeia) to further that point, McSweeney et al. (2021).

Hueber (1992) suggested differences between the

Baragwanathia from the two horizons at Yea were likely

intraspecific and considered the interpretation of graptolites by

Garratt and Rickards (1984) as subjective because preservation

was poor. Additionally, Hueber (1992) highlighted that the

purported 24 million year difference between the two horizons

had resulted in little evolutionary change, as did Hao and

Gensel (2001: 103), yet massive differences occurred over 3

million years between the Cooksonia spp. found in Ireland

(Edwards et al., 1983) and the younger Lower Plant Horizon of

Yea. Edwards et al. (1979: 321) questioned the interpretation of

the age of Maoristrophia banksi and the graptolites not

occurring on the same plane as the Baragwanathia in the

Lower Plant Horizon. Finally, Edwards et al. (1997: 22)

consider the Humevale and Yea Formations at Yea to be part

of the Norton Gully Sandstone Formation.

The Yea and Humevale Formations espoused by Garratt

(1978) are retained because they provide a stratigraphic

framework for the area (McSweeney et al., 2020: fig. 2 —

stratigraphical comparison chart and Ghin Ghin Road

exposure). However, a Lower Devonian age will be used when

assigning an age to specimens at Yea to account for these

differing interpretations.

Materials and methods

The collection site of early land plants from Limestone Road,

Yea, 1s heritage listed, with further collecting forbidden; in

addition, the Frenchmans Spur locality 1s heavily overgrown

and, for the most part, inaccessible. The specimens described

herein are from Museum Victoria's palaeontological archives

and were collected by J.D. Tims and J. Douglas in the 1970s

and early 1980s, and one specimen S. caespitosa (NMV

P235941) collected by C. Earp and M. Pepper, 19 m from the

Devil's Elbow road cutting on Ghin Ghin Road, Yea, central

Victoria, Australia.

The specimens collected by Tims and Douglas are

preserved as fine compressions/impressions, and even after

maceration with Schultze's solution and acetate film pulls (see

Tims and Chambers, 1984: 266), no evidence of internal

anatomy remains. The S. caespitosa collected by Earp and

Pepper is coalified and a fragment was taken from the lowest

part of the parent axis of NMV P23594].2 using a fine steel

needle. Additionally, a fragment was dislodged from the

centre of the basal dark presumed sporogenous region of

sporangia two and four on NMV P23594].1. These fragments

were placed onto aluminium stubs for scanning electron

microscope analysis under low vacuum on an FEI Quanta 200

ESEM at RMIT University’s RMIT Microscopy and

Microanalysis Facility (RMME).

Images were taken at Museums Victoria (Carlton) using a

Leica M205 C microscope and Leica Application Suite

software version 3.8.0. Images were edited using Adobe

Illustrator CC 2017 and morphological measurements were

taken using ImageJ software. Dégagement was undertaken on

the Museums Victoria material archived by Tims (1974,

1980), Tims and Chambers (1984) and J. Douglas.

Institutional abbreviations

NMV P, Museum Victoria Palaeontology Collection,

Melbourne, Australia. The use of .1 and .2 at the end of NMV

P refer to part and counterpart, respectively. All specimens

including the samples placed on stubs for scanning electron

microscope analysis are housed in the Palaeontological

Section, Museums Victoria, Melbourne.

Systematic palaeobotany

Plantae. /ncertae sedis

Genus. Salopella Edwards and Richardson, 1974

Type Species. Salopella allenii Edwards and Richardson, 1974

Salopella australis Tims and Chambers, 1984 (fig. 2A—G; line-

drawing fig. 6A)

Salopella australis Tims and Chambers (1984: pl. 32, figs. 1—6; pl.

34, figs. 4, 5; text-fig. 2A—C)

Emended diagnosis (after Tims and Chambers, 1984). New

information in bold. Axes 0.9—2.4 mm wide with central line,

at least two long aligned parallel parent axes with one

dichotomy each resulting in narrower shorter daughter

axes terminated by a single sporangium on each. Plant at

least 80 mm high. No obvious branching at the base of the

sporangia. Sporangia 6.5—14.0 mm high and 1.3—2.0 mm wide,

with parallel sides in the lower two-thirds of the presumed

fertile portion. Sterile sporangia apex tapering to a point in the

upper third. Spore characters unknown.

Locality. Frenchmans Spur Track, 10 km west of Matlock,

Victoria. 37° 25.82' S, 146° 77.24' E.

Stratigraphy and age. Wilson Creek Shale, mid-Pragian-

Emsian, Lower Devonian (Carey and Bolger, 1995; Mawson

and Talent, 1994).

Note: Two specimens assigned to S. australis by Tims and

Chambers (1984) from Limestone Road have been transferred

to Gen. et sp. indet. because the specimens are too poorly

preserved to unequivocally assign to any taxon. These are

NMV P229617 and NMV P157323; NMV P157323 (Jack

Douglas Private Collection) was figured by Tims and

Chambers (1984: pl. 32, figs 5, 6 and text-fig. 2b) but possesses

a completely different branching architecture to the holotype

for S. australis and has poorly defined sporangia. Specimen

NMV P157323 has been re-drawn here (fig. 6B) for

comparison to S. australis (ig. 2A-E). Specimen NMV

P50011 from Limestone Road has been moved out of S.

australis to a new Salopella species (see below). Most

specimens assigned to the Wilson Creek Shale by Tims and

Chambers (1984) on Frenchmans Spur track are considered S.

australis because they have subtending axes the same width

196

as their sporangia (e.g. NMV P50014, NMV P33219 and

NMV P50008 [holotype]). Only in cases where this character

is equivocal are specimens assigned to Gen. et sp. nov. (e.g.

NMV P50010).

Description. Salopella australis possessed an erect habit with

at least two long parent axes aligned parallel to each other and

with one distal dichotomy on each resulting 1n narrower shorter

daughter axes, each terminating in an elongate sporangium

with an acuminate apex. A slight indentation occurs on some

sporangia at the darker carbonaceous area and sterile interface,

and proximally the dark carbonaceous area tapers inwards on

the daughter axes.

Remarks. A reassessment of the morphospecies Salopella

australis as originally determined by Tims and Chambers

(1984) has necessitated the removal of some of the specimens

attributable to it, primarily due to differing branching

architectures and poor preservation. Tims and Chambers

(1984: 268) noted four species with two dichotomies, but only

three were found; presumably, the fourth is in a private

collection. Nonetheless, all the specimens figured in Tims and

Chambers (1984) were examined. Tims and Chambers (1984:

270) described S. australis as "open-branched" (Tims and

Chambers, 1984: pl. 32, fig. 5, 6) with at least two dichotomies

of widely dichotomising daughter axes emanating from one

parent axis, and up to at least 145 mm in length (fig. 6B).

However, this was found to occur on only three specimens

from Limestone Road and was not found on the holotype (fig.

2D, E — double arrows between aligned parent axes).

The holotype possessed two aligned parent axes, with one

of the parent axes dichotomising into shorter narrower

daughter axes. The other parent axis was not as well

preserved. However, another specimen with better

preservation (fig. 2A, C) shows both parent axes aligned and

dichotomising into shorter narrower daughter axes terminated

with elongate sporangia that are no wider than their

subtending axes. Tims and Chambers (1984: pl. 32, fig. 4)

partly illustrated this specimen with only one of the parent

axes shown. However, Tims and Chambers (1984) did not

mention or show the other parent axis aligned parallel to this

axis. Furthermore, there was an additional specimen in the

collection not figured by Tims and Chambers (1984), likely

due to its poor preservation, but which also had two aligned

almost parallel axes and is similar to the holotype with two

parent axes aligned and with one of the parent axes possessing

a single dichotomy with short daughter axes terminated in

elongate sporangia (fig. 2B). Even taking into account the

effects of degradation of the specimens before fossilisation

and the effects of ocean currents on orientation of the axes,

the Limestone Road specimens appears to have more in

common with S. caespitosa than S. australis, with two

dichotomies and relatively longer daughter axes from the

ultimate dichotomy (fig. 6B).

This emendation has resulted in the maximum known

length of S. australis being reduced from 145 mm to 80 mm

and the reduction of known dichotomies to one, with the

sporangial dimensions and morphology remaining the same.

Specimens attributed to S. australis herein (fig. 2A—E) were

F.R. McSweeney, J. Shimeta & J.S. Buckeridge

deposited in a deep marine quiescent environment of the

Wilson Creek Shale, and like the holotype, possessed parallel

erect axes. This branching architecture of erect parallel

aligned axes was found only in three specimens, including the

holotype. All the other specimens bar one, which is defined

below, are too poorly preserved to attribute to any taxon.

Parsimony suggests that parallel axes that dichotomise at the

same level, with axes of comparable widths, terminating in

elongate sporangia on short daughter axes also of equal

dimensions, that have been transported a considerable

distance belong to the same plant (sensu lato Wang and Hao,

2004; Edwards et al., 2015; Edwards and Li, 2018a) and are

representative of its true branching architecture. We

acknowledge that we have failed to show organic connection,

and current alignment could conceivably result in this axial

configuration, but the likelihood of this occurring to three

separate specimens with characters such as branching and

sporangia occurring at the same height and with the same

dimensions is unlikely. We postulate that the parallel aligned

axes may have emanated from a rhizomatous region because

there was no converging of the parent axes. Two of these

parallel aligned specimens were included in Tims and

Chambers (1984) but noticeably in both cases, only one of the

parent axes was visible (Tims and Chamber, 1984: text-fig.

2A, pl.32, figs. 1, 2, 4). Additionally, the line-drawing of S.

australis in Tims and Chambers (1984: text-fig. 2a) shows the

daughter axes of the parent axes with slightly differing

lengths, which was based on Tims and Chambers (1984: pl.

32, fig. 1). But a closer examination of this specimen

(holotype; fig. 2D) shows the daughter axes are preserved on

different levels of lamina, with the perceived shorter daughter

axis (upper arrow) partly obscured by another axis.

The specimens removed from Salopella australis (Yims

and Chambers, 1984: pl. 32, figs 3,5) and placed in Gen. et sp.

indet. include isolated poorly preserved sporangia from both

the Wilson Creek Shale on Frenchmans Spur (fig. 2F, G) and

specimens from Limestone Road because the preservations

were generally poor and the branching architectures greatly

differed to the holotype. One of the excluded specimens

(Tims and Chambers, 1984: pl. 32, fig. 3) with differing

branching architecture to S. australis is described below as a

new morphospecies. Thus, S. australis is no longer

represented in the flora of the Lower Plant Horizon.

Salopella caespitosa Tims and Chambers, 1984 (fig 3A—I

(holotype) and fig. 4A—E)

Material: NMV P50016 (holotype) and NMV P235941 both with

part and counterpart.

The circumscription of Salopella caespitosa was based on a

single specimen with its counterpart from the Wilson Creek

Shale Formation on Frenchmans Spur Track approximately 10

km west of Matlock in central Victoria. No new data have

been added on this taxon since its inception. The discovery of

a new specimen from the base of the Humevale Formation on

Ghin Ghin Road near Yea preserved in semi-relief contributes

information on its dimensions, gross morphology and

temporal range.

Early land plants from the Lower Devonian of central Victoria, Australia, including a new species of Salopella 197

Figure 2. Salopella australis displaying parallel parent axes dichotomising into much shorter daughter axes that are terminated with elongate

sporangia, delineated with a constriction just above dark sporogeneous region. All from Wilson Creek Shale on Frenchmans Spur track, 10 km

west of Matlock. A, erect parent axes parallel to each other, dichotomising into two elongate sporangia from Wilson Creek Shale. Re-photographed

here; originally published in Tims and Chambers (1984: pl. 32, fig. 4) and Tims (1980: fig. 4.1.9). Specimen NMV P50014. B, forked dichotomy

terminated by sporangia from Wilson Creek Shale. And to the right hand side of the forked axis is another long axis, which based on its

orientation may also be part of the same plant. Constriction at arrow, lower arrow at dichotomy and double arrow at two aligned axes. Specimen

NMV P33219. C, close-up of fructification in A, sporangia barely extend beyond the confines of their subtending axes, with slight constriction

present above sporogeneous region (at arrow). Specimen NMV P50014. D, E, holotype, part and counterpart. On part, constriction at arrow in

sporangium. On counterpart, both parent axes are parallel to each other (at dotted arrow). Re-photographed here, originally published in Tims

and Chambers (1984: pl. 32, figs. 1, 2). Specimens NMV P50008.1 and NMV P50008.2, respectively. F, Gen. et sp. indet. — short daughter axes

terminated 1n elongate sporangia. The cortex may be absent from subtending axes, with only the central line visible. The lack of cortex prevents

assigning to S. australis as width of subtending axis to sporangial width is required. Originally photographed by Tims (1980: fig. 4.1.13).

Specimen NMV P50010.2. G, S. australis, with two short daughter axes, with constriction at arrow of the sporangium, which is the same width

as its subtending axis. Specimen NMV P202886.

198

Description. The new specimen’s length is 135 mm and consists

of two synchronous dichotomies with longer than wide

sporangia terminating each axis. The naked slender axes are

0.8—1.5 mm wide, with the axial width decreasing equally after

each dichotomy. The first dichotomy occurs c. 80 mm from the

base of the fossil and c. 31 mm from the second dichotomy (fig.

4A at lower arrow). The subtending axes to the sporangia are

0.9-1.0 mm wide and 22-25 mm high with some slight

variation in the width along all four axes and a slight narrower

evident beneath sporangia three and four (fig. 4B, the subtending

axis to sporangium four pinches slightly about | mm below the

sporangium). The sporangia are fusiform 1.5—2.0 mm wide and

4.8—6.0 mm high with the subtending axis slightly widening

into the base of each sporangium. The widest part of the

sporangia occurs in the lowest quarter length, followed by a

gentle tapering to rounded sporangial tips. A sporangial wall is

evident primarily but 1s not confined to the basal parts of the

sporangia measuring 0.2—0.3 mm wide and is continuous with

the subtending axes. Sporangia two, three on the part and four

on the counterpart possess what appears to be an infilled oval

presumed sporogenous area, which are no greater than 1.2 mm

wide and 1.5 mm high. Sporangium four on the counterpart

shows a fine linear cavity 0.15 mm wide entering the base of

the sporogenous area and sporangium three on the part shows

similar but larger opening beneath the sporogenous area, and

beneath this is a fine central axis 0.1 mm wide; beneath

sporangium two on the subtending axis, this axis is 0.2 mm

wide. Sporangium two (part) upper half possesses longitudinally

oblique striations, 0.06—0.6 mm apart.

Remarks. Comparison to holotype. The new specimen

measures 50 mm longer than the holotype (fig. 3), and its axial

width 1s broadly in range with the holotype, which measures

1.1-1.9 mm wide according to Tims and Chambers (1984).

What appears to be the widening of the parent axis in the

specimen (fig. 4À) 1s where the first dichotomy occurs, with the

two daughter axes aligned for 26 mm before diverging for the

remaining 5 mm before the second dichotomy. From the

holotype, the best examples of branching to compare to the

specimen were found on the part specimen (right-hand side)

with isotomous dichotomies subtending sporangia one-seven

(fig. 3A, fig. 6D). Proximally on the axes ultimately terminate

in sporangia one-seven show the axes emanating from a

horizontal axis. This horizontal axis may represent part of a

rhizomatous system or a distorted axis. Tims and Chambers

(1984) appear to have considered this a dichotomy because no

other part of the fossil would give three dichotomies. However,

it is equivocal as to whether it is a distorted axis with a

dichotomy. Nonetheless, the axes of the new specimen

examined are noticeably less flexuous than the holotype; for

example, on the holotype axes leading to sporangia eight and

nine showing changes in orientation of about 45? in conjunction

with the daughter axes of dichotomies occurring at 10° and 45°.

These differences possibly represent differences in both current

strength during deposition and the level of degradation of each

of the plants. There is a noticeable variation in the distance

between the ultimate branching points and sporangia in the

holotype, from c. 6 mm as seen in sporangium 12, to c. 26 mm

F.R. McSweeney, J. Shimeta & J.S. Buckeridge

for sporangium seven, and up to at least 57 mm for sporangium

eight, where the presumed branching point is not visible. The

axes decreased after each dichotomy, as noted by Tims and

Chambers (1984), but with the specimen examined herein there

is a much greater decrease in axial width after the first

dichotomy. Examination of the holotype did reveal a slight

narrowing in the subtending axes (fig. 3D—F) of most of the

sporangia (Tims and Chambers 1984: pl. 34, fig. 3); this was

observed beneath two of the sporangia on the specimen, but in

both cases may be artefacts of preservation, and more

specimens are required to confirm this character.

The size range of the sporangia in the holotype, 3.2—6.0

mm high and 1.3—2.0 mm wide (Tims and Chambers, 1984),

is similar to the specimen examined herein, and both have a

comparable average height to width ratio of 2.6 (Morris and

Edwards, 2014: table 3) for the holotype and 3.2 herein. The

holotype shows slight variation in the location of the widest

point of the sporangia, occurring either in the lowest quarter

(fig. 4F), like with NMV P23594] (fig. 4C) or midway (fig.

3D). The minor difference 1s possibly accounted for by the

different types of preservation. The specimen NMV P2359]

shows these sporangial bodies 1n semi-relief and indicate they

were not flattened bodies. Furthermore, examination of

sporangia of the specimen revealed an oval presumed

sporogenous region in the basal part of the sporangia, which

is also present on sporangium 13 of the holotype (fig. 3H).

lims and Chambers (1984: pl. 33, figs 3—4) noted that on

sporangium 13 of the holotype it may either be a double

sporangium or dissociation of the sporangial wall giving the

appearance of a double sporangium. The evidence of a

sporogenous area basally and relatively thick sporangial walls

from the specimen examined herein (fig. 4B) suggests either a

crushed sporangium or stage of senescence pertaining to

dehiscence. Furthermore, some of the sporangia in the

holotype have a darkened presumed sporogenous area

completely filling insides of the sporangia (fig. D—F), which

may have resulted from the breakdown of the wall holding the

sporogenous area prior to fossilisation, as the sporogenous

area occurs as a basal oval body on all sporangia of the

specimen examined.

Tims and Chambers (1984: pl. 33, fig. 3) found central

lines of 0.2 mm diameter throughout the specimen, one of

which was beneath sporangium 13. On the specimen NMV

P235941, the central line was found to connect to base of the

sporogenous region (fig. 4B, C), which Tims and Chambers

(1984) had postulated. Tims and Chambers (1984) noted some

axes of the holotype to dichotomise (c. 6.0 mm) relatively

close to the base of the sporangia (fig. 3C); however, this was

not the case with NMV P23594].

The presence of longitudinal oblique striations on the upper

half of sporangium two may reflect cell orientation and possible

twisting as a means of dehiscence. Obliquely orientated cells in

the sporangia have been recorded for some S. allenii and absent

for S. marcensis (Fanning et al., 1992; Morris and Edwards,

2014), and throughout the entire sporangia and subtending axes

of Tortilicaulis (Edwards et al., 1994). There was no indication

of twisting found for the subtending axes of the specimens

sporangia, Suggesting it may only affect the upper part of the

Early land plants from the Lower Devonian of central Victoria, Australia, including a new species of Salopella 199

Qe wp "VIE. *

« m gt =

-—

aÃ- g

pmm

i -—

A — E t E “ae

9 A > z . B

Figure 3. Salopella caespitosa (holot

P202987.1) and counterpart (NMV P202987.2), respectively; numbering follows Tims and Chambers (1984, pl. 33, fig. 1). Counterpart images

are reversed to be in the same orientation as the part specimen. A, on right-hand side of part specimen, double isotomous dichotomies lead to

sporangia one—seven. Note, at the arrows, there 1s perpendicular branching that 1s suggestive of a rhizomatous system. C, sporangium 12; at the

arrow there is another axis that is terminated in a sporangium that is partially visible. Specimen NMV P202987.1. D—F, Evidence of a slight

constriction beneath sporangia at arrows. D, sporangia eight and nine (on the left). Widest part of each sporangium occurring approximately

midway along their length. Note, rephotographed; originally published in Tims and Chambers (1984: pl. 34, fig. 3). Specimen NMV P202987.1.

E, sporangium 22 is c. 4.06 mm long and 1.4 mm wide. The axis decreases from 1.2 mm proximally to 0.5 mm just beneath the sporangium.

Specimen = NMV P202987.2. F, sporangia four (right) and five, both sporangia are slightly wider in the lower quarter of each sporangia.

specimen = NMV P202987.2. G, sporangium 22, appears to be two immature sporangia juxtaposed. At arrow, the apex (rounded) of the smaller

fusiform sporangium is apparent. Specimen NMV P202987.1. H, sporangium two, lower arrow at walls surrounding presumed oval sporogeneous

area, which reaches approximately halfway the length of the sporangium to the upper arrow. Specimen NMV P202987.1. I, sporangium 13,

arrows at walls surrounding sporangeneous area and upper arrow showing extent of oval sporogenous body. The walls does not recombine

apically like in sporangium two, suggesting it may have been crushed, or hidden, beneath the matrix. Rephotographed; originally figured by Tims

and Chambers (1984: pl. 33, fig. 3). Specimen NMV P202987.2.

200

sporangia; however, this is conjectural. As noted by Morris and

Edwards (2014: 42), sporangia containing spores are required

for unequivocal identification and assessment of relationships.

F.R. McSweeney, J. Shimeta & J.S. Buckeridge

more data. The presence of this specimen at the base of the

Humevale Formation potentially suggests a late Silurian age

(Rickards, 2000). However, as outlined in the stratigraphy

section, we suggest a Lower Devonian age range for specimens

found at Ghin Ghin Road until unequivocal evidence of age

becomes available.

While the specimen described is three-dimensionally

preserved, we believe erecting a new taxon to segregate it from

the form genus would be premature and we should wait for

uz | Ty" MORTEM yee. ON, o,

Figure 4. Salopella caespitosa NMV P235941 from Devil's Elbow on Ghin Ghin Road, northwest of Yea. A, overview of whole specimen. NMV

P235941.1. B, arrow (Vt) at vascular trace entering base of oval presumed sporogenous body of sporangium four, and the subtending axis to

sporangium four pinches slightly about 1 mm below the sporangium. Lower arrow at region where subtending axis is continuous with sporangial

wall and upper arrow shows extent of sporangial body of sporangium three. Note, for both sporangia, the distal parts appear to be hidden in the

matrix. NMV P23594].2. C, lower arrow (Vt) at vascular trace entering base of oval sporogenous body, with upper arrow at distal extent of

sporogenous body on sporangium two on the counterpart. Note, no longitudinally oblique striations are evident in upper half of sporangium but

are evident on the part specimen. NMV P235941.2. D, sporangium two at arrow longitudinal oblique striations on the upper half of the

sporangium. NMV P23594].1. E, close-up of parent axis with longitudinal striations. NMV P235941.1.

Early land plants from the Lower Devonian of central Victoria, Australia, including a new species of Salopella 201

Plantae. /ncertae sedis

Genus. Salopella Edwards and Richardson, 1974

Salopella laidae McSweeney, Shimeta and Buckeridge, sp. nov.

(fig. 5A, B, fig. 6D)

Salopella australis Tims and Chambers (1984: pl. 32, fig. 3, text-

fig. 2C)

Diagnosis. Plant at least 55 mm long, tuft branching

architecture, with sporangia all reaching the same height, with

naked dichotomously synchronous branched axes up to 3.2 mm

wide basally and 1.3 mm below sporangia. Axial width

decreases after each dichotomy. Isotomous branching angles

small, 30°—40° for the ultimate dichotomies and 15?—50? for

basal dichotomies. At least two orders of branching. Branching

10.0—14.0 mm below sporangia. Eight sporangia, elongate sub-

fusiform 9.7—-11.0 mm high and up to 2.5 mm at widest section

in mid region, and wider than subtending axes, obtuse apices.

Etymology. Named for Mrs Peg Laid who kindly gave

permission to collect fossil specimens from her property on

Ghin Ghin Road, Yea.

— O

Holotype. NMV PS50011.1 and NMV P50011.2, part and

counterpart respectively, figured by Tims and Chambers, 1984:

pl. 32, fig. 3 and text-fig. 2C

Locality. Location 4, Brackley's cutting (Harris and Thomas,

1941) Limestone Road, 2 km southeast of Yea. Note: exposure

referred to as locality 62 in Couper (1965). 37° 12.38' S, 145°

25.39' E.

Stratigraphy and age. Location 4 (Harris and Thomas, 1941:

303) Yea Formation (Garratt 1978), Lower Devonian (Pragian;

Edwards et al., 1997; Rickards, 2000; Rickards and Garratt,

1990).

Salopella laidae sp. nov. (fig. 5A, B, fig. 6C)

Description. Specimen preserved as a faint white compression

on blue-grey claystone with six of the eight sporangia preserved.

There is no clear sporogenous area within the sporangia.

However, sporangia five, seven and eight (fig. 6C) on the part

(fig. 5A) possess darkened creamy-grey regions extending up

two-thirds of their lengths and possibly represent a

sporogeneous region. No internal anatomy or spores were

Figure 5. Salopella laidae sp. nov. (holotype) NMV P50011.1 and NMV P50011.2, part and counterpart, respectively, with counterpart reversed

to be in the same orientation as part specimen. From location 4 (Brackley’s cutting) on Limestone Road, Yea. A, double isotomous dichotomy

visible, terminated with eight elongate sporangia, five visible. At lower arrow central line and at F, folding of tissue. On sporangium seven

?sporogenous region highlighted. Rephotographed; originally figured by Tims and Chambers, 1984: pl. 32, fig. 3 and text-fig. 2C. B, arrow at

daughter axis missing on part present.

202

recovered when Tims and Chambers (1984) examined this

specimen. The sporangia are slightly wider than the widths of

their subtending axes and do not taper distally but come to a

rounded apical region. There are two synchronous dichotomies

clearly visible, with the axial width decreasing only after each

dichotomy. Folding of tissue appearing as a longitudinal

lineation along the edges of the axes and sporangia are visible;

no evidence of dehiscence mechanism is present (fig. 5A).

Comments and comparison with other species. The diagnosis

and description of this taxon is based on one specimen (part

and counterpart). While we are mindful of issues in naming a

new species on one specimen, we have done so because of the

shape of the sporangia and compact branching architecture that

set it apart from other rhyniophytoids (Table 1). The use of the

paraphyletic genus Salopella (Edwards et al., 1994) is to

emphasise its gross morphology and to follow previous workers

(Tims and Chambers, 1984) in identifying Victorian flora.

However, we note that work done by Edwards et al. (1994) on

in situ Salopella spores demonstrated significant diversity, that

can be difficult to ascertain because of convergence of some

sporangial shapes (Edwards et al. 2001: 11), making assertions

on palaeogeographic distribution and spread unwise without

palynological evidence. Tims and Chambers (1984) originally

assigned this specimen to S. australis, but Salopella laidae sp.

nov. differs to S. australis in a number of its characters (fig. 6

— comparative line drawings of Australian Salopella). Salopella

laidae sp. nov. possesses sporangia that are clearly delineated

from their subtending axes, differing noticeably from coeval S.

australis (fig. 2A—E) where the sporangia are as wide as their

= -e = = = «= &

A B

F.R. McSweeney, J. Shimeta & J.S. Buckeridge

subtending axes. Furthermore, the sporangia in S. laidae sp.

nov. do not taper from half-way to one-third from their apices

as in S. australis; instead, they hold their sporangial widths

before terminating with rounded apices. While the sporangial

apices appear obtuse, this cannot be assumed to have been the

case in life, as Morris and Edwards (2014: 40) found when

examining S. allenii that compressed specimens possessed

rounded apices of varying degrees, except that one specimen of

S. allenii that was preserved in 3D had a pointed apex. The

parent axes leading to the ultimate dichotomy in S. australis

are long, erect and aligned parallel to each other (fig. 2A), being

easily distinguished from S. /aidae, where the axes leading to

the ultimate dichotomy are at an angle to the vertical.

Furthermore, S. australis only possesses one dichotomy, while

S. laidae possesses two, or possibly three, as suggested in Tims

and Chambers (1984: text-fig. 2C). However, it should be noted

the proximal axes of S. laidae sp. nov. are orientated away from

each other, suggesting they may have come from a rhizomatous

axis instead of a dichotomising one.

One of the parent axes of S. laidae sp. nov. 1s incomplete,

but likely possessed four sporangia (fig. 6C). This 1s based on

the relative dimensions, orientation and positions of both

parent axes to each other, and that four sporangia terminate

the daughter axes of the other parent axis.

The sporangia of Salopella caespitosa are ovate to fusiform

and up to 6.0 mm high and 2.0 mm wide (Tims and Chambers,

1984), which is approximately half as high than S. laidae.

Furthermore, the axes of S. caespitosa are more gracile than

those of S. laidae, as the latter reached a maximum width of 1.9

Figure 6. Line-drawings of Victorian early land plants with longer than wider sporangia. A, Salopella australis from Wilson Creek Shale on

Frenchmans Spur track, near Matlock. Specimen NMV P50014. B, Gen. et sp. indet. from Limestone Road, Yea. Originally placed in S. australis

but branching architecture clearly differs. Specimen NMV P157323. C, Salopella laidae sp. nov. from Limestone Road, Yea. NMV P50011. D,

Salopella caespitosa (holotype) from Wilson Creek Shale on Frenchmans Spur track, near Matlock. Specimen NMV P202987. E, Salopella

caespitosa from Ghin Ghin Road, Yea. Specimen NMV P23594].

Early land plants trom the Lower Devonian of central Victoria, Australia, including a new species of Salopella 203

mm proximally and 1 mm beneath the sporangia, whereas for

S. laidae axes were at least 3.2 mm proximally and 1.3 mm

beneath the sporangia. In comparison to other Salopella

globally, the sporangia of S. marcensis and S. xinjiangensis are

much smaller than S. /aidae, reaching a maximum length of 3.4

mm and 3.5 mm long, respectively, and both much narrower

(Morris and Edwards, 2014: table 3). Moreover, the sporangia

of S. xinjiangensis are tongue-shaped (Cai et al., 1993; Dou and

sun, 1983), but S. marcensis sporangia are similar in gross

morphology to S. laidae, being elongate and tapered at their

apices according to Fanning et al. (1992). Salopella marcensis

are much smaller plant than S. laidae, reaching 6.38 mm high,

while S. laidae reached at least 55 mm high (Fanning et al.,

1992: 174). While disarticulation for S. marcensis during

transport seems likely, the isotomously branched axes bearing

the sporangia for S. marcensis are much smaller than for S.

laidae, ranging 0.05—0.5 mm wide (Fanning et al., 1992).

In Fanning et al. (1992: fig. 35), there is a silhouette of S.

marcensis that shows a sporangium occurring on a lateral

axis below a further bifurcation, indicating in this case at

least that sporangia did not necessarily occur at approximately

the same level, as is the case with S. /aidae sp. nov. Finally,

the type species of Salopella, S. allenii possessed fusiform or

spear-shaped sporangia (Edwards and Richardson, 1974;

Morris and Edwards, 2014) — significantly different to those

of S. laidae sp. nov.

Discussion

Edwards and Richardson (1974) established Salopella based on

one Lochkovian specimen of Salopella allenii from Brown

Clee Hill, Shropshire (England); they noted its affinities to

Rhyniaceae and that S. allenii possessed axes with dimensions

that were between the range seen for Rhynia gwynne-vaughani

and R. major (= Aglaophyton majus), with sporangia similar to

both but lacking internal anatomy as seen in the Rhynie chert

specimens (Edwards et al. 2017). Various workers have

hypothesised that rhyniophytes/rhyniophytoids occupied

ephemeral ruderal terranes, such as wetlands and deltas,

whereby the rhyniophytes/rhyniophytoids completed their life

cycles over short periods (Edwards, 2003; Edwards and Davies,

1990; Morris et al., 2011; Raymond et al., 2006; Wellman et al.,

2000), possibly with some forming dense monotypic stands,

such as the well-preserved aligned axes found near a fossilised

channel from the Lower Old Red Sandstone deposits of Brecon

Beacons (Edwards and Fanning, 1985: 156). According to

Lavender and Wellman (2002), Edwards and Richardson

(2004), and Raymond et al. (2006), seasonal conditions would

have suited the short life cycles of rhyniophytes/rhyniophytoids.

The location of the Melbourne Zone c. 30? south of the equator

during the Lower Devonian (Torsvik and Cocks, 2019: fig. 9)

suggests that the terrestrial sources for these deposits were

likely also influenced by a seasonal climate, suiting the short

life cycle of rhyniophyte/rhyniophytoids.

Furthermore, Edwards and Richardson (2004), when

investigating the Lower Devonian calcrete palaeosols on the

Avalonian Terrane, proposed rhyniophytes/rhyniophytoids as

likely pioneer taxa because they were found on calcrete

palaeosols that had formed in unstable interfluvial areas, with

more advanced coeval plants, such as the zosterophylls or

lycopsids, inhabiting more stable terranes to better

accommodate their longer life cycles (Wellman et al., 2000).

According to Powell et al. (2003: 29), there was a gradual

shallowing of marine facies from the western margins of the

Melbourne Zone in the Darraweit Guim Province, and

eastward movement of the shoreline during the upper Silurian—

Lower Devonian, which may have provided similar unstable

conditions for rhyniophytes/rhyniophytoids, because

palaeocurrents at areas in Yea and Matlock indicate a west or

southwesterly source from the western parts of Victoria, which

was terrestrial at the time (Powell et al., 2003: 23). Edwards

and Davies (1990: fig. 16.3) noted the decline of rhyniophytes/

rhyniophytoids from Laurussian sites during the late Pragian

and Emsian stages of the Lower Devonian, possibly due to the

proliferation of zosterophylls early in the Devonian. It is

possible the uplift of the Tabberaberan highlands towards the

end of the Lower Devonian in Victoria (Powell et al., 2003: 32)

may have provided ideal unstable conditions for rhyniophytes/

rhyniophytoids into the Emsian but work on the palaeoflora

from the Norton Gully Sandstone Formation has been limited.

Finally, there is a noticeable difference in the number of

rhyniophyte/rhyniophytoid taxa from Yunnan, China, in what

was once part of the South China Block located at and north

of the equator during the Lower Devonian (Torsvik and

Cocks, 2019: fig. 9a), and those found in the Baragwanathia

floral assemblage. According to Hao and Xue (2013: fig. 5.3),

rhyniophytes/rhyniophytoids make up a small percentage (c.

2.1%) of the taxonomic groups found in the Posongchong

Flora. Recently, Edwards and Li (2018b) erected

Polycladophyton gracilis and Jiangyounia gengi possessing a

erade of organisation belonging to Rhyniophytina. However,

Salopella specimens have not yet been recovered (Hao and

Xue, 2013: 193), suggestive of some level of 1solation between

the two regions.

Conclusions

Defining taxa based on gross morphological characters alone

has limitations because it excludes cryptic characteristics and

likely results in one morphospecies encompassing a number

of distinct species. However, this study and the work of Tims

and Chambers (1984) has provided a morphological

framework from which new Victorian taxa may be defined,

providing an impetus for further research.

Acknowledgements

We thank Museums Victoria and especially the Vertebrate

Palaeontology manager Tim Ziegler for facilitating access to

their collection of Salopella spp., and for organising the use of

the M205 C Leica microscope to photograph specimens. Dr

Michael Garratt for his help in Yea. Mrs Peggy Laid for access

to her property on Ghin Ghin Road, Yea, and Dr J. Tims who

met with FMcS and JB on 14 March 2018. Clem Earp in

corresponding with F. McSweeney on specimens he collected in

the Yea and Alexandra. This work was supported by an

Australian Government Research Training Program Scholarship.

204

Disclosure statement

No potential conflict of interest was reported by the authors.

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