Memoirs of Museum Victoria 77:1-14 (2018) Published 2018

1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.Org/10.24199/j.mmv.2018.77.01

Burrowing lobsters mostly from shallow coastal environments in Papua New

Guinea (Crustacea: Axiidea: Axiidae, Micheleidae)

(http://zoobank.org/urn:lsid:zoobank.org:pub:876F855F-AF2E-41BC-8CF6-BB87875BA074)

GARY C. B. Poore (http://zoobank.org/urn:lsid:zoobank.org:author:893860FE-E71B-4F69-A46A-8B89834579FC)

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

Abstract Poore, G.C.B. 2018. Burrowing lobsters from shallow coastal environments in Papua New Guinea (Crustacea: Axiidea:

Axiidae, Micheleidae). Memoirs of Museum Victoria 77: 1-14.

Surveys of coral reefs and associated habitats have discovered nine species of Axiidae and one of Micheleidae in

Papua New Guinea. Only the micheleid is new to science. The collection provides an opportunity to provide colour

photographs of some and to revisit their taxonomy. Two species are synonymised with others: Alienaxiopsis lizardensis

Sakai, 2011 with A. clypeata (De Man, 1905) and Allaxiopsis bougainvillensis Sakai, 2011 with Axiopsis Picteti var.

spinimana De Man, 1905, now Allaxiopsis spinimana (De Man, 1905). Axiopsis pica Kensley, 2003 is recognised as

distinct from A. serratifrons, with which it co-occurs. Michelea papua sp. nov. is described as new.

Keywords Crustacea, Axiidae, Micheleidae, Alienaxiopsis, Allaxiopsis, Axiopsis, Parascytoleptus, Paraxiopsis, Ralumcaris,

Michelea, Papua New Guinea, taxonomy

Introduction

The Our Planet Reviewed (La Planete revisitee) expeditions in

Papua New Guinea in 2012 and 2014, coordinated by Phillipe

Bouchet for the Museum nationale d’Histoire naturelle, Paris,

discovered a diverse fauna of axiidean ghost shrimps and

burrowing lobsters in shallow water. Ghost shrimps of the family

Callianassidae are being studied separately, but here burrowing

lobsters belonging to Axiidae and Micheleidae are dealt with.

While only one of the ten species recorded is new, four are

first records for Papua New Guinea and provide new

morphological information that illuminates the complicated

and sometimes confusing taxonomy of some species. Seven

species were photographed in colour. The discoveries have

provided information for a reassessment of genera of Axiidea

(work in progress) that were most recently reviewed (as

Axioidea) by Sakai (2011).

The material comes from two sampling series in shallow

water environments in Papua New Guinea using a variety of

methods, including divers in shallow water brushing coral

rubble under water (PB and KB prefixes), hand-dredging (PD

and KD prefixes) or sampling individually by hand (PR, KR

and KZ prefixes). The expedition in November-December

2012 near Madang, Madang Province, provided samples in the

PAPUA NIUGINI series. The expedition in April 2014 based

in Kavieng, New Ireland, provided samples in the KAVIENG

2014 series. These collections were augmented for comparative

purposes with others from shallow waters of Papua New

Guinea in other museums, and by material from the Kimberley

region of Western Australia.

Material and methods

Material is deposited in the Museum national d’Histoire

naturelle, Paris (MNHN, IU-prefixes), Museum fur

Naturkunde, Berlin (ZMB), Zoological Museum, Hamburg

(ZMH), Museums Victoria, Melbourne (NMV), the Australian

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

(WAM) and Northern Territory Museum and Art Gallery,

Darwin (NTMAG). All measurements are of carapace length

(cl.) including rostrum. Photographs of fresh specimens that

were taken in the lab shortly after collection were made by

Tin-Yam Chan (TYC) and Arthur Anker (AA).

Results

Four of the ten species recorded, Alienaxiopsis clypeata (De

Man, 1888), Allaxiopsis spinimana (De Man, 1905), Axiopsis

pica Kensley, 2003, and Paraxiopsis austrinus Sakai, 1994,

have been recorded from Papua New Guinea for the first time.

Axiopsis ( Axiopsis) pitatucensis De Man, 1925 (= Calaxius

pitatucensis), described from Papua New Guinea, was not

collected but is probably a higher water species. It is surprising

that, despite hundreds of dredge and diving samples targeting

axiidean habitats in shallow water, more species were not

captured. Most of the species found are widespread in the Indo

West-Pacific; in particular, Axiopsis serratifrons and

Paraxiopsis brocki have been recorded numerous times from

many places. The possibility that these last two are species

complexes cannot be discounted.

G.C. B. Poore

Infraorder Axiidea de Saint Laurent, 1979

Axiidae Huxley, 1879

Alienaxiopsis Sakai, 2011

Alienaxiopsis Sakai, 2011: 32-33.

Type species. Alienaxiopsis lizardensis Sakai, 2011, by

original designation.

Remarks. Alienaxiopsis was erected to include two species,

Alienaxiopsis lizardensis Sakai, 2011 (type species) and A.

clypeata (De Man, 1888). Sakai’s (2011) key differentiated the

two species but, as explained below, the two are synonymous.

Alienaxiopsis clypeata (De Man, 1888)

Figs la, b, 3

Axius clypeatus De Man, 1888: 470, pi. 20 fig. 2.

Axiopsis (Axiopsis) clypeata.—De Man, 1925: 70.

Allaxius clypeatus.— Sakai and de Saint Laurent, 1989: 73-74.—

Poore and Collins, 2009: 237.

Alienaxiopsis lizardensis Sakai, 2011: 34-36, fig. 2. Syn. nov.

Material examined. Papua New Guinea. Madang Province. PAPUA

NIUGINI stations. Tab I., 05° 09.9' S, 145° 50.4' E, 20 m (stn PB06),

IU-2013-7096 (ovigerous female, 3.4 mm).

New Ireland Province, Kavieng lagoon, KAVIENG 2014 stations.

E side of Ral I., 02°36.7'S, 150° 42.6' E, 3-10 m (KZ22), IU-2014-1142

(ovigerous female, 2.5 mm; male, 4.2 mm). Mouth of Albatross

Passage, E side, 02° 35.2' S, 150° 43.1’ E, 13 m (KB72), NMV J71641

(female, 4.2 mm). New Ireland mainland, N coast, 02° 35.2’ S,

150° 50.3' E, 17 m (KB66), IU-2016-8134 (female, 3.5 mm).

Australia. Queensland, Great Barrier Reef, Yonge Reef, near

Lizard I., 14° 38’ S, 145° 38' E, AM P.25014 (holotype of Alienaxiopsis

lizardensis, male, 4.7 mm).

Photographed specimens not seen. Papua New Guinea. Madang

Province. PAPUA NIUGINI stations: location not specified, 15

m (stn PR89), 1 specimen. Kranket I., 05° 12' S, 145° 48.8' E

(stn PR86), 2 ovigerous females.

Type locality. Ambon, Indonesia.

Supplementary description. Rostrum acute, depressed, 0.3

length of rostral base-cervical groove, with pair of erect lateral

spines at midpoint and larger pair at base. Median gastric

carina obsolete, with 1 tooth, ending in broad triangular plate;

submedian gastric carina obsolete, with 2 erect teeth;

supraocular spine oblique, not marginal; lateral gastric carina

obsolete, with 1 erect tooth. Eyestalk reaching end of rostrum.

Antenna article 2 with broadly triangular distal spine;

scaphocerite 5 times as long as greatest height (lateral view),

reaching to midpoint of article 5. Major cheliped ischium,

merus and carpus each with minute distal tooth on lower

margin; propodus swollen, upper margin 1.35 times greatest

height, carinate, with distal tooth; fixed finger 0.5 times length

of upper margin of propodus, with blunt distal tooth near tip of

cutting edge; dactylus 2.3 times as long as wide, cutting edge

with 2 low rounded teeth in proximal half. Minor cheliped as

long as major cheliped, propodus about 0.75 times as high as on

major cheliped; ischium and merus each with minute distal

tooth on lower margin; propodus swollen, upper margin 1.2

times greatest height, carinate, with distal tooth; fixed finger as

long as upper margin of propodus, with distal tooth on cutting

edge; dactylus 3.2 times as long as wide, cutting edge smooth.

Telson 1.1 times as wide as long at level of most anterior lateral

teeth; distal margin 0.5 times telson greatest width; lateral

margin with 4 teeth; distal margin convex, with 1 or 2 lateral

articulating robust setae, with median spine; face with 2 pairs

of spines. Uropodal endopod twice as long as wide; anterior

margin strongly lobed proximally (as rounded shoulder),

otherwise concave with strong tooth at midpoint and elevated

distal spine; posterior margin convex, unarmed; distal margin

straight, oblique, with strong elevated spine at anterior end,

smaller spine at posterior end; facial rib with 3 spines. Uropodal

exopod oval, twice as long as wide; anterior margin of article 1

with 4 teeth; posterior margin convex; distal margin irregular,

3 marginal spines, stronger spine defining posterior corner,

strong articulating spine near anterior corner; article 2 oval,

with distal spine almost as long as body of article.

Colour. Translucent with bright red band anteriorly and

dorsolaterally on carapace, dorsolaterally on pleon, and on

upper margins of cheliped carpus and propodus; maxilliped 3

bright red; cheliped fingers white.

Distribution. Indo-West Pacific (Guam, Papua New Guinea,

Indonesia, Fiji); to 20 m depth.

Remarks. De Man’s (1888) description was extensive but his

drawings few. Here, colour photographs and figures of the

carapace, pereopods 1 and tail fan are included.

Sakai (2011) based a new species Alienaxiopsis lizardensis

on the specimen from Lizard I., Great Barrier Reef, Australia

(AM P.25014) that was examined and identified by Poore and

Collins (2009) as Allaxius clypeatus. Sakai listed Poore and

Collins’ record under the synonymy and distribution of both

species. This specimen has been re-examined and redrawn; no

differences in the gastric ornamentation (characters alleged to

differentiate the two) could be detected between it, material

from Papua New Guinea or De Man’s (1888) figures. Sakai’s

(2011: fig. 2B) figure of the dorsal carapace is quite misleading.

Alienaxiopsis lizardensis is here synonymised with A. clypeata.

Until now, Alienaxiopsis clypeata was known only from

Ambon, Indonesia (type locality), Guam and Fiji (Kensley,

2003), but is here recorded from Papua New Guinea. The

record from Fiji is based on unpublished data from the USNM

online database http://collections.nmnh.si.edu/search/iz/).

Allaxiopsis Sakai, 2011

Allaxiopsis Sakai, 2011: 34-35.

Remarks. Sakai (2011) included three species in Allaxiopsis but

confused their records. The type species, Paraxius picteti

Zehntner, 1894, was described from a single female (cl. 10 mm)

collected at Ambon, Indonesia. Two males (the larger cl. 8.5

mm) were recorded from Siboga station 209 at Kabaena I.,

Indonesia, by De Man (1905) and then re-illustrated (De Man,

1925). More specimens were recorded from Guam, Marshall

Islands, and Fiji by Kensley (2003). Allaxiopsis spinimana (De

Burrowing axiidean lobsters from Papua New Guinea

Figure 1. Alienaxiopsis clypeata (De Man, 1888): a, b, MNHN unregistered (stn PR86). Allaxiopsispicteti (Zehntner, 1894): c, MNHNIU-2013-

1209. Axiopsis pica Kensley, 2003, MNHN IU-2013-7048: d, preserved; e, living. Axiopsis serratifrons A. Milne-Edwards, 1873: f, MNHN

IU-2013-638; g, h, MNHN IU-2013-7052; i, MNHN IU-2013-7046; j, MNHN IU-2013-302. Photo credits: AA, a-c, e, g-j; TYC, f; GCBP, d.

G.C. B. Poore

Figure 2. Parascytoleptus papua Poore and Collins, 2010: a, MNHN IU-2014-2735; b, MNHN IU-2013-7128. Paraxiopsis brocki (De Man,

1888): c, MNHN IU-2013-7108; d, MNHN IU-2014-2736. Ralumcaris bisquamosa (De Man, 1905): e, MNHN IU-2013-7120. Photo credits: AA,

b, c, e; TYC, a, d.

Man, 1905) was originally described as a variety of A. picteti

from Siboga station 209 based on a 9.25 mm female and a

smaller male. The variety was treated at the species level by

Sakai and de Saint Laurent (1989) and has been rediscovered

(see below). Allaxiopsis bougainvillensis Sakai, 2011 (type

species of the genus) was described from much smaller

specimens (4.5 mm and 4.9 mm) from Bougainville, Papua New

Guinea and is treated here as a synonym of A. spinimana. Sakai

included Kabaena I., Sulawesi, Indonesia, in the distribution of

all three species and as type locality of the last two, contradicting

the data provided with his type specimens. Each of the two

species is diagnosed here with a minimal character suite.

Allaxiopsis picteti (Zehntner, 1894)

Figs lc, 4

Paraxius picteti Zehntner, 1894: 196-199, pi. 9 fig. 25.

lAxiopsis picteti.— Borradaile, 1903: 539.

Axiopsis ( Axiopsis) picteti.—De Man, 1925: 6,70, 92-96, pi. 7 fig.

16.

Allaxius picteti .—Sakai and de Saint Laurent, 1989: 75.—

Kensley, 2003: 361, pis 5, 6.

Allaxiopsis picteti.— Sakai, 2011: 39-40.

Material examined. Papua New Guinea. Madang Province. PAPUA

NIUGINI stations. S of Urembo I., outer slope, 05° 15.9' S, 145° 47. V E,

3 m (stn PB43), IU-2013-1209 (male, 8.3 mm). N of Bil Bil I., 05° 17.7 S,

145° 46.9’ E, 5 m (stn PB51), IU-2013-7014 (male, 2.5 mm).

New Ireland Province, Kavieng lagoon, KAVIENG 2014 stations.

S side of Patio I., 02° 36.2’ S, 150° 31.6' E, 6-8 m (stn KB38), IU-2014-

2526 (male, 6.8 mm). NW point of Nusa I., 02° 33.9' S, 150° 46.7' E,

8-10 m (stn KS3), IU-2014-2041 (female, 2.0 mm).

Australia. Western Australia, Kimberleys, Echuca Shoal, <23 m,

13° 53.781’ S, 123° 53.686' E (Woodside Kimberley Survey stn 107/

K12), WAM C50773 (female, 5.5 mm).

Type locality. Ambon, Indonesia.

Burrowing axiidean lobsters from Papua New Guinea

Figure 3. Alienaxiopsis clypeata (De Man, 1888), male, MNHN IU-2016-8134: a, b, anterior carapace, dorsal and lateral views; c, telson and

uropod; d, major left cheliped; e, minor right cheliped. Scale bar = 1 mm.

Diagnosis. Major cheliped, propodus with blunt tubercles on

lateral and mesial faces, more prominent nearer upper margin.

Minor cheliped, propodus tuberculate on lateral and mesial faces.

Supplementary description. Rostrum acute, depressed, 0.3 length

of rostral base-cervical groove, with pair of erect lateral spines

near apex and larger pair at base. Gastric carina difficult to

differentiate; median gastric carina obsolete except near base of

rostrum, with sequence of 1, 1, 2, 2, 2 teeth; submedian gastric

carina obsolete, with 1 tooth anteriorly and 2 or 3 teeth posteriorly;

supraocular spine oblique, not marginal; lateral gastric carina

with 2 or 3 blunt teeth. Eyestalk reaching beyond end of rostrum.

Antenna article 2 with small distal spine; scaphocerite 4 times as

long as greatest height (lateral view), reaching third length of

article 4. Major cheliped coxa-carpus unarmed; propodus upper

margin 1.4 times greatest height, carinate, with 5 spines, lateral

face tuberculate proximally near upper margin; fixed finger 0.5

times length of upper margin of propodus, cutting edge with 2

blunt distal teeth; dactylus cutting edge with 3 low rounded teeth

in proximal half. Minor cheliped coxa-carpus unarmed; propodus

upper margin 1.5 times greatest height, carinate, with 2 distal

spines, lateral face with few proximal tubercles; fixed finger

almost as long as upper margin of propodus, with distal teeth near

G.C. B. Poore

Figure 4. Allaxiopsis picteti (Zehntner, 1894), males, MNHN IU-2013-1209: a, b, anterior carapace, dorsal and lateral views; c, telson and

uropod; d, minor right cheliped. MNHN IU-2013-2526: e, major left cheliped. Scale bar = 1 mm.

tip of cutting edge; dactylus cutting edge smooth. Telson 1.4 times

as wide as long at level of most anterior lateral teeth; distal margin

0.8 times telson greatest width; lateral margin with 3 teeth; distal

margin straight, with 1 lateral articulating robust seta, lateral

fixed spine, with median spine; face with 2 pairs of spines.

Uropodal endopod 1.6 times as long as wide; anterior margin

strongly lobed proximally (as rounded shoulder), otherwise

concave with or without spine at midpoint, with subdistal and

distal spine; posterior margin convex, with 5 spines along distal

third; distal margin straight, transverse, with depressed spine at

anterior end, 2 marginal spines, 1 stronger spine and another

superior, at posterior end; facial rib unarmed. Uropodal exopod

semicircular, 1.5 times as long as wide; anterior margin of article

1 convex, with 5 or 6 marginal teeth, 1 submarginal; posterior

margin straight, with 3 distal spines set obliquely; distal margin

transverse, 3 marginal spines, strong articulating spine near

anterior corner; article 2 with 5 teeth along distal margin.

Colour. Carapace high red-brown; pleon with patches of green-

brown and scattered red chromatophores; antennal flagellum with

alternating white and brown stripes; cheliped high blue; pereopods

with transverse blue bands on major articles, otherwise white.

Distribution. Indonesia: Ambon (type locality), Kabaena I.,

Sulawesi (De Man, 1925); Guam; Marshall Islands; Fiji

(Kensley, 2003); Papua New Guinea: Madang, Bougainville,

New Ireland; Australia, N Western Australia; 3-20 m depth.

Remarks. Kensley (2003) reported on material from Guam and

included a photograph with colours similar to the one here.

Kensley also reported on unpublished records of the species

from Papua New Guinea, Fiji and the Marshall Islands

identified by him (see USNM online database http://collections.

nmnh.si.edu/search/iz/). Kensley’s record of the species from

Malaysia is not on the database. The carapace, tail fan and the

never-before-illustrated chelipeds are figured here.

Allaxiopsis spinimana (De Man, 1905)

Fig. 5

Axiopsis Picteti var. spinimana De Man, 1905: 597.

Axiopsis ( Axiopsis ) Picteti var. spinimana.—De Man, 1925: 6,70,

96, pi. 7 fig. 17.

Allaxius spinimanus.— Sakai and de Saint Laurent, 1989: 75.

Allaxiopsis spinimana.— Sakai, 2011: 40.

Allaxiopsis bougainvillensis Sakai, 2011: 37-39, fig. 3. Syn. nov.

Material examined. Papua New Guinea. Bougainville, Teop I.,

05° 34.3' S, 155° 4.7’ E, (as ‘Tiop Bougainville, German New Guinea’,

H. Schoede, ZMB 14440 (holotype female, 4.9 mm; paratype female,

4.5 mm of Allaxiopsis bougainvillensis Sakai, 2011) (both photographed

by C.O. Coleman).

Madang Province, Channel between Pik I. and Kranket I.,

05° 09.6’ S, 145° 49.7' E, 3-8 m, coll. R. Hanley, NMV J67992 (2

ovigerous females, 7.5, 7.8 mm; 3 males, 3.2-5.0 mm; part of larger

collection, NTMAG Cr.0100212).

Burrowing axiidean lobsters from Papua New Guinea

Figure 5. Allaxiopsis spinimana (De Man, 1905) male, NMV J67992: a, habitus; b, c, anterior carapace, dorsal and lateral views, with detail of

rostrum; d, telson and uropod; e, major right cheliped, lateral; f, major right cheliped, fingers, mesial; g, minor left cheliped; h, maxilliped 3; i—1,

pereopods 2-5; m, n, pleopod 2, with details of appendices interna and masculina. Scale bars = 1 mm. Bases of many setae indicated by small ovals.

G.C. B. Poore

Type locality. Indonesia, off south point of Kabaena I., 22 m

(,Siboga stn 209).

Diagnosis. Major cheliped, propodus with blunt tubercles on

proximal lateral face, more prominent nearer upper margin

becoming spine-like and more hooked towards distal upper

margin. Minor cheliped, propodus with few tubercles on

proximal lateral face, upper margin with 3 spines.

Supplementary description. Rostrum acute, depressed, 0.3

length of rostral base-cervical groove, with pair of erect lateral

spines near apex, larger pair at midpoint, and ventral tubercle.

Gastric carina difficult to differentiate; median gastric carina

obsolete except near base of rostrum, with sequence of 1, 1, 2

teeth; submedian gastric carina obsolete, with 1 tooth anteriorly

and 2 or 3 teeth posteriorly; supraocular spine oblique, not

marginal; lateral gastric carina, with 2 blunt teeth. Eyestalk

reaching beyond end of rostrum. Antenna article 2 with small

distal spine; scaphocerite 4 times as long as greatest height

(lateral view), reaching sixth length of article 4. Major cheliped

ischium, merus and carpus each smooth on lower margin;

propodus swollen, lateral face tuberculate over proximal half,

tubercles larger closer to upper margin, upper margin about

equal to greatest height, with 2 rows each of 6 spines, the larger

ones sharper and more hooked distally; fixed finger 0.45 times

length of upper margin of propodus, with 2 blunt distal teeth

along cutting edge, with short tuberculate mesial ridge; dactylus

2.1 times as long as wide, with lateral carina near upper margin,

cutting edge with 2 low rounded teeth in proximal half. Minor

cheliped as long as major cheliped, propodus about 0.5 times as

high as on major cheliped; ischium and merus smooth on lower

margin; propodus swollen, propodus upper margin 1.5 times

greatest height, lateral face with 3 tubercles on proximal lateral

face, upper margin with 3 teeth, second and third sharp; fixed

finger almost as long as upper margin of propodus, with 2 distal

teeth on cutting edge; dactylus 4 times as long as wide, cutting

edge smooth. Telson 1.4 times as wide as long at level of most

anterior lateral teeth; distal margin 0.8 times telson greatest

width; lateral margin with 3 teeth; distal margin excavate, with

1 lateral articulating robust seta, lateral fixed spine, with

median spine; face with 2 pairs of spines. Male pleopod 2

appendix masculina 1,3 times as long as appendix interna, stiff

setae on posterior face. Uropodal endopod 1.8 times as long as

wide; anterior margin strong lobed proximally (as rounded

shoulder), otherwise concave with spine at midpoint, with

subdistal and distal spine; posterior distal margin curved, with

10 spines, the most distal 2 submarginal, anterodistal angle

marked by 2 spines; facial rib with 3 spines. Uropodal exopod

semicircular, 1.8 times as long as wide; anterior margin of

article 1 convex, with 6 marginal teeth; posterior margin

straight, with 5 distal spines set obliquely; distal margin

transverse, 2 marginal spines, strong articulating spine near

anterior corner, facial rib with 3 spines; article 2 with 5 teeth

along distal margin.

Colour. Traces of purple on pereopod 1 propodus and dactylus on

preserved material. De Man (1925) described the colour as being

similar to A. picteti but more violet in parts. Juveniles and adults

of A. picteti appear to differ so this species may also differ.

Distribution. Indonesia, S Sulawesi; Papua New Guinea,

Central Province, Bougainville; shallow heights.

Remarks. De Man (1905) based Axiopsis picteti var. spinimana

on two syntypes from the ‘Anchorage off the south point of

Kabaena-island’, Indonesia {Siboga stn 209), the same locality

at which he also recorded A. picteti. De Man (1925) described

his two syntypes in moderate detail but illustrated only the

distinctive cheliped.

Allaxiopsis bougainvillensis Sakai, 2011, is based on two

female specimens (ZMB 14440) from Papua New Guinea,

which Sakai referred to as the holotype with both chelipeds

(‘lectotype’ in fig. 3 caption) and a paratype without chelipeds.

They are correctly identified on the ZMB label as Axiopsis

( Axiopsis ) picteti var. spinimana De Man, 1905’, possibly by

H. Schoede. In describing his new taxon, Sakai (2011) correctly

stated that these specimens are not types but used this

observation to justify a new species without stating how it

differed from A. (A.) picteti spinimana. He did not illustrate or

describe the distinctive spinose chelipeds of the holotype

(photographs of which were provided to me by C.O. Coleman),

which are clearly identical to those in De Man’s (1925) figure

of A. spinimana and to those figured here from other material.

The supposed differences in gastric sculpture between A.

bougainvillensis and A. picteti are small and not relevant.

The type locality of A. bougainvillensis was given by

Sakai (2011) as ‘Triop Bougainville, German New Guinea’, a

mistranscription of Tiop written on the label, which is now

spelled Teop.

The species shares the trifid rostrum, regular pattern of

blunt gastric spines, short scaphocerite, broad telson, uropodal

endopod with shouldered anterior margin, and spinose uropodal

rami with A. picteti. The most significant difference is the

presence of chelipeds with spinose palms, the distal spines on

the upper margin of the palm having a characteristic hooked

form, characters that formed the basis of the identification of the

Papua New Guinea specimens. Most of the characters of Sakai’s

(2011) diagnosis of this species are of generic value only.

Axiopsis Borradaile, 1903

Axiopsis pica Kensley, 2003

Figure Id, e

Axiopsis pica Kensley, 2003: 363, figs 1, 2, pi. 1.—Ngoc-Ho,

2005: 51-55, fig. 2.

Axiopsis serratifrons.— Sakai, 2011: 56-63 (part).

Material examined. Papua New Guinea. Madang Province, PAPUA

NIUGINI stations. Kranket I., outer slope, 05° 11.3' S, 145° 49.5' E,

1-24 m (stn PR129), IU-2013-7048 (female, 14.4 mm).

Mariana Islands. Guam Island, Apra Harbour, Middle Shoal,

among coral rubble and rocks, 1 m, IU-2016-8007 (UF 2782), (1

ovigerous female, 16 mm); near Harbour entrance, among rocks, 8-12

m, IU-2016-8008 (UF 3021) (female, 13.5 mm).

Distribution. Guam (type locality), Papua New Guinea, French

Polynesia; to 24 m depth.

Remarks. The single female from Papua New Guinea was first

identified by its striking colour pattern, similar to that published

Burrowing axiidean lobsters from Papua New Guinea

by Kensley (2003: pi. 1). Kensley (2003) noted that, as well as a

distinctive colour, Axiopsis pica has ‘a broader and more robust

larger cheliped of pereopod 1 bearing flattened scale-like

tubercles’ than A. serratifrons with which it co-occurred. The

upper margin of the propodus of the holotype and of the Papua

New Guinea female is 1.5 times its greatest height. Kensley

(2003) also compared his new species with material identified as

A. serratifrons from Hawaii which has more slender chelipeds.

Ngoc-Ho (2005) compared specimens that she identified

as A. pica from French Polynesia with a syntype of A.

serratifrons also from Hawaii. The major cheliped of this

syntype is twice as long as wide and smooth. Following

Sakai’s (2011) selection of the other syntype from Tonga as the

lectotype (see below), comparison with Hawaiian specimens

may be irrelevant.

Axiopsis pica co-occurs with A. serratifrons in both

French Polynesia and Papua New Guinea. The major cheliped

of the largest specimen is similarly proportioned, 1.5 times as

long as wide, as of similarly-sized A. serratifrons. The most

reliable morphological distinction between the two species

can be found in the carapace. The carapace and pleon of A.

serratifrons is smooth and flexible, with few scattered long

setae, while that of A. pica is sclerotised, almost calcified and

pitted with short stiff setae associated with the pits (Fig. Id).

Axiopsis serratifrons (A. Milne-Edwards, 1873)

Figs lf-i, 6a

Axia serratifrons A. Milne-Edwards, 1873: 263, pi. 13 figs 6, 6a.

Axiopsis serratifrons.— Sendler, 1923: 44, pi. 21 fig. 10.—Sakai

and de Saint Laurent, 1989: 76.—Sakai, 2011: 56-63, fig. 9 (extended

synonymy).

Material examined. Paralectotype. Hawaii, IU-2016-8115 (Thl47)

(male, 10 mm).

Papua New Guinea. Madang Province, PAPUA NIUGINI

stations. Kranket I., outer slope, 05° 12.1' S, 145° 49.3' E, 17 m (stn

PB02), IU-2013-302 (female, 9.3 mm); 05° 11.3’ S, 145° 49.5’ E, 1-11

m (stn PR225), IU-2013-7051 (male, 22 mm), NMV J71638 (ovigerous

female, 22 mm); 05° 12' S, 145° 49’ E, 10 m (stn PR99), IU-2013-7033

(ovigerous female, 9.5 mm). Rempi Area, S of Barag I., 05° 01.3’ S,

145° 47.9’ E, 2-13 m (stn PR61), IU-2013-638 (ovigerous female, 9.3

mm); S of lagoon inside bay, 05° 01.6’ S, 145° 48.P E, 2-15 m (stn

PR69), IU-2013-7116 (male, 4.3 mm); outer slope, 05° 01.6' S,

145° 48.1' E (stn PR65), IU-2013-637 (male, 11.5 mm). Alexishafen,

05° 05.3' S, 145° 48.1' E, 1-6 m (stn PD31, IU-2013-7019 (male, 6.3

mm). W of Panab I., 05° 10.3' S, 145° 48.5' E, 1-18 m (stn PR147), IU-

2013-7052 (female, 15.8 mm). Riwo waters, 3-15 m (stn PR109), IU-

2013-7061 (male, 11.9 mm). S of Yabob I., 05° 15.5’ S, 145° 47.3' E,

2-6 m (stn PD66), IU-2013-7098 (male, 5.8 mm). Ulimal I., 05° 05.6' S,

145° 48.7' E, 6 m (stn PS16), IU-2013-15308 (male, 10.0 mm).

New Ireland Province, Kavieng region, KAVIENG 2014 stations.

Edmago I., 02° 36.9’ S, 150° 44.4’ E, 9 m (KZ2), IU-2014-826 (male,

8.6 mm); IU-2014-2685 (male, 9.3 mm). New Ireland mainland near

N Cape, 02° 33.3’ S, 150° 47.7' E, 1-20 m (stn KZ18), IU-2016-1011

(female, 12.5 mm). W side of Edmago I., 02° 37.1' S, 150° 44.2' E,

5-6 m (stn KZ20), NMV J71639 (ovigerous female, 10.5 mm). E side

of Ral I., 02° 36.7' S, 150° 42.6' E, 3-10 m (stn KZ22), IU-2014-1090

(female, 10.6 mm). Byron Channel, SE Patio I., 02° 36.6’ S,

150° 32.9' E, 2-7 m (stn KB40), IU-2014-2577 (ovigerous female,

10.9 mm). NE of Big Nusa I., entrance to Kavieng Harbour,

02° 33.7' S, 150° 49.1' E, 10 m (stn KZ11), IU-2014-2625 (ovigerous

female, 10.6 mm). Mouth of Albatross Passage, E side, 02° 35.2' S,

150° 43. P E, 13 m (KB72), IU-2016-8136 (juv., 5.0 mm). Between Big

Nusa and Little Nusa Islands, 02° 34.6' S, 150° 46.3' E, 13-14 m

(KB16), IU-2014-17688 (female, 6.9 mm). Eickstedt Passage W of

Usien I., 02° 40.3' S, 150° 39.1' E, 9-11 m (KR70), IU-2014-17691

(male, 13.6 mm). Albatross Passage, 02° 44.6’ S, 150° 42.8’ E, 12-15

m (KD12), IU-2014-17692 (juv., 3.6 mm).

Colour. Variable. Generally reddish-orange, stronger colour on

gastric carina; pleonal pleura with white patch anteroventrally;

chelipeds similar or steel-blue, colour stronger at base of fingers

(see figs lf-i and Kensley [1981]).

Distribution. Widespread in the Indo West-Pacific, eastern

Pacific (Hendrickx, 2008), south-west Atlantic (Sakai, 2011,

2015) and south-east Atlantic (Wirtz, 2009); subtidal.

Remarks. Of the two syntypic specimens from Samoa and

Hawaii recorded by A. Milne-Edwards (1873), Sakai (2011)

selected that from Samoa as the lectotype, not the one from

Hawaii erroneously applied to the ‘type locality’ by Kensley

(2003) and called ‘holotype’ by Ngoc-Ho (2005). This

confusion was discussed by Komai and Tachikawa (2008).

Sakai’s (2011: figs 8A, B, 9) illustrations of the Samoan

lectotype (ZMB K8405: checked for me by A. Brandt) are

indistinguishable from Ngoc-Ho’s (2005: fig. 3) of the Hawaiian

paralectotype (MNHN IU-2016-8115 [Thl47]). Sakai’s (2011:

fig. 8C) illustration of the Hawaiian paralectotype differs from

both in appearing to have larger rostral teeth, the rostrum less

evenly tapering, more teeth on the median carina (shown by my

re-examination to have two on the rostrum, c. 15 on gastric

region; fig. 6a), almost no spines on the lateral gastric carina

(actually 13, 15), and fewer intermediate gastric tubercles

(actually c. 20). The cheliped of the paralectotype lacks

tuberculation on the propodal faces and the spine on the upper

border of the merus, but these absences are common in

juveniles of this size.

In an extensive synonymy, Sakai (2011) synonymised

four species with A. serratifrons. The synonymy of Axius

affinis De Man, 1888 (type locality, Ambon, Indonesia),

Axiopsis sculptimana Ward, 1942 (type locality, Diego

Garcia, Chagos Archipelago) and Axiopsis brasiliensis

Coelho and Ramos-Porto, 1991, has not been disputed

although a species with such a wide distribution suggests

further examination is warranted as Komai and Tachikawa

(2008) suspected. Kensley (1981) discussed the species in the

Americas but his synonymy was limited. Ngoc-Ho (2005)

recognised A. pica Kensley, 2003 (type locality, Guam), the

fourth species synonymised by Sakai (2011), following a

detailed justification and recorded it from French Polynesia.

This synonymy is not recognised here (see A. pica above for

discussion of differences).

Sakai’s (2011) key to species of Axiopsis relied on the

presence of a tooth on the upper margin of the merus and a

smooth propodus of the cheliped to distinguish A. consobrina

from A. serratifrons (without a tooth, with squamose

propodus). Many smaller individuals, including ovigerous

females, identifiable as A. serratifrons based on colour

resemble A. consobrina De Man, 1905 in these features. De

G.C. B. Poore

Figure 6. Axiopsis serratifrons A. Milne-Edwards, 1873: a, male, MNHN IU-2013-637, anterior carapace, dorsal view. Parascytoleptus papua

Poore and Collins, 2010, male, MNHN IU-2013-7128: b, major right cheliped. Paraxiopsis brocki (De Man, 1888), male, MNHN IU-2013-7108:

c, anterior carapace, lateral view; d, antenna with scaphocerite; e, cheliped, merus; male, MNHN IU-2014-2736; f, g, pleopods 1, ventral and lateral

views. Ralumcaris bisquamosa (De Man, 1905), male, MNHN IU-2013-7120: h, i, anterior carapace, dorsal and lateral views. Scale bars = 1 mm.

Man (1905) distinguished A. consobrina on the absence of

intermediate gastric teeth between the carinae and the palm of

the smaller cheliped as long as the fingers but this is true only

for the type. Axiopsis consobrina occurs usually from 75 m to

a maximum of 310 m depth (Sakai, 2011; Vaitheeswaran,

2014) but Ngoc-Ho (2005) recorded one individual from 2.5 m

depth. Axiopsis serratifrons is more immediately subtidal.

Parascytoleptus Sakai and de Saint Laurent, 1989

Parascytoleptus papua Poore and Collins, 2010

Figs 2a, b, 6b

Parascytoleptus papua Poore and Collins, 2010: 614-618, figs 1, 2.

Material examined. Papua New Guinea. Madang Province, PAPUA

NIUGINI stations. N of Riwo mangrove and seagrass, 05°08.7'S,

Burrowing axiidean lobsters from Papua New Guinea

145°48.2'E, 2 m (stn PB48), IU-2013-7100 (male, 2.6 mm). N of Sek I.,

inner slope, 05°04.7'S, 145°48.9'E, 3 m (stn PB50), NMV J71642 (2

males, 2.8,3.1 mm).

New Ireland Province, Kavieng region, KAVIENG 2014 station.

NW side of Ral I., coral wall, 19 m, 02° 36.4' S, 150° 42.4' E (stn

KB62), IU-2014-2735 (female, 3.0 mm); IU-2014-17694 (male, 3.6

mm).

Distribution. Papua New Guinea, Madang and New Ireland

provinces; 2-19 m depth.

Remarks. The types were collected not far from the new

material. The major cheliped of the adult male figured here is

longer and more elongate than that of the female figured by

Poore and Collins (2010). Sakai (2011) diagnosed the genus

with the male pleopod 1 ‘a small unsegmented protrusion’

based on its presence on two males of 4.2 mm and 5.3 mm

length. This has not been observed on the holotype of P. papua

(4.2 mm) or the smaller males reported here. The pleopod 1

may appear only in larger specimens.

Paraxiopsis De Man, 1905

Remarks. Two of the 16 known species were found in Papua

New Guinea and are diagnosed here with a minimal character

suite.

Paraxiopsis austrinus (Sakai, 1994)

Eutrichocheles austrinus Sakai, 1994: 185, figs. 6, 7.—Sakai

2011 : 111 .

Paraxiopsis austrinus.— Kensley 2003, 373.—Poore and Collins

2009: 266, fig. 29.

Material examined. Papua New Guinea. New Ireland Province,

Kavieng region, KAVIENG 2014 stations. Mouth of Albatross

Passage, E side, 02° 35.2' S, 150° 43.1' E, 13 m (stn KB72), IU-2014-

1046 (male, 6.1 mm). S coast of Baudison I., 02° 45.2’ S, 150° 41.7’ E,

22-27 m (stn KB68), IU-2014-1153 (male, 7.6 mm; ovigerous female,

6.3 mm). E of Albatross Passage, 02° 45.2’ S, 150° 43.4' E, 13-17 m

(stn KB24), IU-2014-2364 (male, 9.0 mm).

Diagnosis. Carapace smooth, with tomentum of short and

longer setae. Rostrum with 0-3 small lateral teeth; lateral

gastric carina with supraorbital spine plus 2 teeth; submedian

gastric carina with 6-8 teeth; median gastric carina without

spines. Telson with 3 or 4 pairs of dorsal spines. Cheliped

merus with 2 spines on upper margin, palm unornamented.

Male pleopod 1 absent in small specimens, single article in

adults.

Distribution. Northern Australia; New Ireland Province, Papua

New Guinea; to 27 m depth.

Remarks. The new material extends the range of this species

from Darwin, northern Australia, to Papua New Guinea. The

species differs from the original description only in having no

rostral spines (three small spines in Australian specimens) and

in having one (rather than two) post-supraocular spine on the

lateral gastric carina. All of the males lack pleopod 1 but

possess a minute tubercle in its place. Kensley (2003) and

Poore and Collins (2009) justified the generic placement of

this species.

Paraxiopsis brocki (De Man, 1888)

Figs 2c, d, 6c-g

Axius brocki De Man, 1888: 475, pi. 20 fig. 3.

Axiopsis ( Paraxiopsis ) brocki.— De Man, 1905: 597.—Tirmizi,

1983: 88-90, fig. 3.

Eutrichocheles brocki.— Sakai and de Saint Laurent ,1989: 52, fig.

4B.—Ngoc-Ho, 1998: 365-368, fig. 1.

Paraxiopsis brocki.— Kensley, 1996.—Poore and Collins, 2009:

266.—Sakai, 2011: 158-161, fig. 27C (full synonymy).

Material examined. Papua New Guinea. Madang Province, PAPUA

NIUGINI stations. W of Panab I., 05° 10.3’ S, 145° 48.5' E, 1-18 m (stn

PR147), IU-2013-7108 (male, 6.8 mm). Riwo, mangrove, 05° 09’ S,

145° 48.2' E, 1-2 m (stn PR235), IU-2013-7118 (ovigerous female, 5.5

mm). Kranket I., Cape Jantzen, 05° 12.5’ S, 145° 49.P E, 13 m (stn

PB11), IU-2013-7126 (male, 3.1 mm).

New Ireland Province, Kavieng region, KAVIENG 2014 stations.

S coast of Baudison I., 02° 45.2' S, 150° 41.7’ E, 22-27 m (stn KB68),

IU-2014-990 (female, 5.5 mm); IU-2014-1032 (female, 5.9 mm). NW

point of Nusa I., 02° 33.9’ S, 150° 46.7' E, 15-17 m (stn KB04), IU-

2014-2054 (juvenile, 3.1 mm). Marthas Shoal, sand and coarse rubble

in gutter, 20 m, 02° 32.5' S, 150° 35.3’ E (stn KB60), NMV J71640

(male, 7.5 mm; 2 ovigerous females, 6.8 mm); IU-2014-2736 (male,

6.8 mm). NW point of Nubis I., 02° 37.2' S, 150° 31.8' E, 20 m (stn

KB39), IU-2014-17693 (juvenile, damaged).

Indonesia. Maluku Province, Pulau Wuliaru, 7° 27’ S, 131° 3.7’ E,

IU-2014-12081 (female, 9.5 mm).

Diagnosis. Carapace smooth, without tomentum of setae.

Rostrum with 4-6 lateral teeth; lateral gastric carina with

supraorbital spine plus 2 teeth; submedian gastric carina with 1

or 2 teeth; median gastric carina without spines. Telson with 3

or 4 pairs of dorsal spines. Cheliped merus with 1-3 spines on

upper margin, palm unornamented. Male pleopod 1 absent in

small specimens, single article in adults.

Distribution. Widespread in the Indo West-Pacific, from eastern

Africa, Western Australia, northern Australia, to southern

Japan and French Polynesia; to 91 m depth (Sakai, 2011).

Remarks. The new material contributes little to knowledge of

this widespread and frequently taken species. The species was

well illustrated by Ngoc-Ho (1998). Sakai (2011: fig. 27C)

figured a simple male pleopod 1 on a male from Darwin,

Australia; pleopod 1 is absent in the smallest male and minute

in the larger ones from this collection. Most of the specimens

at hand have a minute anterior tooth on pleonal pleura 2-5.

Sakai (2011) himself discussed variation in this character and

in the presence or absence of the male pleopod 1 after

presenting an extensive diagnosis.

Ralumcaris Sakai, 2011

Ralumcaris Sakai, 2011: 182-183.

Ralumcaris bisquamosa (De Man, 1905)

Figs 2e, 6h, i

Axiopsis ( Paraxiopsis ) bisquamosa De Man, 1905: 597.—De

Man, 1925: 7, 72, 109, pi. 8 fig. 20-20c, pi. 9 fig. 20d-m.-Holthuis,

1953: 51.

Eutrichocheles bisquamosa.—Sakai and de Saint Laurent, 1989:

53, fig. 15.—Kensley, 1994: 822.

G.C. B. Poore

Paraxiopsis bisquamosa.— Kensley, 1996: 711, 712.—Kensley,

2003: 372, table 2.

Ralumcaris bisquamosa.— Sakai, 2011: 183-185, figs 33, 34.

Material examined. Papua New Guinea. Madang Province, PAPUA

NIUGINI stations. Rempi Area, W of Barag I., 05° 01.2' S, 145° 47.9' E,

5-10 m (stn PD45), IU-2013-7130 (male, 2.6 mm). N of Kranket I.,

05° 11.3' S, 145° 49.6' E, 5 m (stn PB47), IU-2013-7037 (ovigerous

female, 3.6 mm). Cape Barschtch, 05° 03.9’ S, 145° 48.8’ E, 12 m (stn

PB27), IU-2013-7119 (male, 3.2 mm). Tab I., inner slope, 05° 10.1’ S,

145° 50.2’ E, 1-4 m (stn PR162), IU-2013-7120 (male, 4.1 mm).

New Ireland Province, Kavieng region, KAVIENG 2014 stations.

Steffen Strait, W side of Wadei I., 02° 39.5’ S, 150° 37.7’ E, 15 m (stn

KS31), IU-2014-2451 (juvenile damaged), IU-2016-8135 (juvenile, 2.9

mm). W side of Tsoilaunung I., 02° 32.8’ S, 150° 30.8’ E, 6 m stn

KB48), IU-2014-2619 (ovigerous female, 5.9 mm). Tab I., N.L. Bruce,

05° 10' S, 145° 51' E, NMV J34090 (ovigerous female, 4.2 mm).

Distribution. Indonesia, Papua New Guinea, Mariana Is; 1-36

m depth.

Remarks. Kensley (1996) pointed out the differences between

Paraxiopsis bisquamosa and the remaining species of this

genus and excluded the species from Paraxiopsis as redefined

by him. Sakai (2011) described and figured De Man’s holotype

(ZMA Crust. De. 102.674) but labelled it as Tectotype’.

Micheleidae Sakai, 1992

Michelea Kensley & Heard, 1991

Michelea papua sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:37FAF4Cl-2B8E-

44AA-B357-783EDE48B9FD

Figure 7

Material examined. Holotype. Papua New Guinea. New Ireland

Province, N of KobotteronL, 02° 36.4’ S, 150° 42.4’ E, 2-3 m, reef wall

and rubble, (KAVIENG 2014 stn KB62), IU-2013-2781 (male, 3.2 mm).

Diagnosis. Gills fully developed. Pleopods 2, 3, 5 with 18/6,

18/8 and 25/13 marginal lamellae on endopods/exopods,

respectively (pleopod 4 unknown). Telson tapering to rounded

apex, length 1.15 width. Maxilliped 3 ischium with obsolete

crista dentata; merus with mesial tooth.

Description. Cephalothorax 0.4 total length, about 1.65 times

as long as greatest height; rostrum triangular, about half as long

as basal width, slightly depressed distally, about 0.4 as long as

eyestalks; cervical groove weakly defined, reaching 0.6 length

of cephalothorax; longitudinal setal-row level with lateral

margin of eyestalk, of 5 setae; vertical setal-row of 5 setae

below horizontal row and 2 setae near cervical groove.

Pleomere 1 with dorsolateral longitudinal setal-row of 10

setae. Pleomeres 2-6 each with transverse setal-rows of 6-7

setae near midpoint; all somites also with groups of long

simple setae, none with marginal setal-rows.

Antennule with elongate waisted article 1, 0.6 length of

cephalothorax; articles 2 and 3 subequal, each about 0.25

length of article 1; flagella with 11 and 9 articles, longer than

peduncle. Antenna with distinct articulating scaphocerite,

about half length of article 2; article 4 reaching to middle of

article 3 of antennule; article 5 short; flagellum missing.

Mandible, maxillules, maxillae, maxillipeds 1 and 2

typical of genus. Maxilliped 3 ischium with obsolete crista

dentata; merus with strong mesial tooth on right of pair only

(absent on left); exopod 1.6 times ischium length.

Chelipeds equal; ischium with weak lower tooth; merus

with weak tooth on slightly convex lower margin, upper margin

more convex proximally than distally, 1.8 times as long as high;

carpus unarmed; propodus almost cylindrical, 3.6 times as long

as high; fixed finger 0.35 total length of propodus, its cutting

edge with long obsoletely bicuspid tooth at midpoint; dactylus

cutting edge straight, curved distally, equal to fixed finger.

Pereopod 2 unknown. Pereopod 3 propodus 2.5 times as

long as wide, with 2 spiniform setae on distal-upper mesial

face, 6 on distal-lower face; and 2 transverse setal-rows of 1

and 2 setae; dactylus with 2 spiniform setae on upper margin.

Pereopod 4 propodus 3.8 times as long as wide, with 7 spiniform

setae on upper margin, 5 on lower margin; with 2 transverse

setal-rows each of 2 setae; dactylus with 5 spiniform setae on

upper-mesial margin. Pereopod 5 subchelate; fixed finger with

4 distal spiniform setae; dactylus without spiniform setae.

Pleopod 1 of male lobed mesially, expanded distally, with

c. 8 minute hooks, setose around midpoint and laterally, and

with 5 simple seta laterally. Pleopod 2 with appendix interna

sac-like, 2.5 times as long as wide; appendix masculina

narrow, about third long as endopod; with 18 lamellae on

lateral margin of endopod, 6 on distolateral margin of exopod.

Pleopod 3 with 18 lamellae on lateral and distomesial margin

of endopod, 8 on lateral margin of exopod; pleopod 4 unknown;

pleopod 5 with 25 lamellae on endopod, 13 on exopod.

Telson tapering to rounded apex from one-third length; 1.15

times as long as wide. Uropodal endopod ovate, 1.5 times as long

as wide, anterior margin straight, distal margin semicircular,

without distal tooth, posterior margin convex; exopod ovate, 1.7

times as long as wide, anterodistal margin with 16 short

spiniform setae, posterior margin with 6 blade-like setae.

Branchial formula as in M. kalbarri Poore and Collins, 2015.

Distribution. Papua New Guinea. New Ireland Province (03° S,

151° E), 2-3 m. depth (known only from type locality).

Etymology. From Papua New Guinea; noun in apposition.

Remarks. Michelea papua resembles M. imperieusae Poore

and Collins, 2015, from north-western Australia in having

similar numbers of pleopodal lamellae but differs in the short

broad rostrum (not spine like), longer antennae, and more

elongate maxilliped 3 and pereopodal articles.

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 Planete Revisitee

expeditions organised jointly by Museum National d’Histoire

Naturelle, Pro-Natura International and Institut de Recherche

pour le Developpement, with support from Papua New Guinea’s

National Fisheries Authority. The organisers acknowledge

supporting funding from the Total Foundation, the Laboratoire

Burrowing axiidean lobsters from Papua New Guinea

Figure 7. Michelea papua sp. nov., holotype: a, lateral carapace, antenna, antennule, maxilliped 3; b, carapace, antenna, antennule; c, pleomere

6, telson, uropod; d, maxilliped 3; e, f, g, pereopods 1, 3, 4; h, pereopod 5 dactylus; i, j, pleopods 1, 2. Scale bar = 1 mm.

d’Excellence Diversites Biologiques et Culturelles (LabEx

BCDiv), the Laboratoire d’Excellence Diversites Biologiques et

Culturelles (LabEx BCDiv, ANR-10-LABX-0003-BCDiv), the

Programme Investissement dAvenir (ANR-ll-IDEX-0004-02),

the Fonds Pacifique, and CNRS Institut Ecologie et

Environnement (INEE). The expedition was endorsed by the

New Ireland Provincial Administration and operated under a

Memorandum of Understanding with the University of Papua

New Guinea (UPNG). I am grateful to Laure Corbari, Paula

Lefevre-Martin and Anouchka Krygelmans-Sato at MNHN and

Zdenek Duris, University of Ostrava, Czech Republic for help in

making the collections available. Tin-Yam Chan, National

Taiwan Ocean University, Taiwan, and Arthur Anker kindly

provided the colour photographs. I thank C. Oliver Coleman

(ZMB) for information on the type locality and photographs of

the holotype of Allaxius bougainvillensis. Thanks also to S.

Keable (AM) and Andrew Hosie (WAM) for arranging loans.

Finally, I acknowledge the support of Philippe Bouchet,

expedition funding, MNHN generally, all the divers who

collected samples, the expeditioners who sorted them in the

field, and the Crosnier Fund for financial support in Paris.

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Memoirs of Museum Victoria 77:15-28 (2018) Published 2018

ISSN 1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI: https://doi.Org/10.24199/j.mmv.2018.77.02

The Indo-West Pacific species of Neaxiopsis and Neaxius (Crustacea: Axiidea:

Strahlaxiidae)

(http://zoobank.org/urn:lsid:zoobank.org:pub:9CAD42D5-099D-4BA9-9CFF-E377A8D422CA)

GARY C. B. Poore 1 (http://zoobank.org/urn:lsid:zoobank.org:author:c004d784-e842-42b3-bfd3-317d359f8975) and

PETER C. DwORSCHAK 2 (http://zoobank.org/urn:lsid:zoobank.org:author:4BCD9429-46AF-4BDA-BE4B-439EE6ADC657)

1 Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia gpoore@museum.vic.gov.au

2 Dritte Zoologische Abteilung, Naturhistorisches Museum, Burgring 7, Wien, Austria Peter.Dworschak@nhm-wien.ac.at

Abstract Poore, G.C.B., and Dworschak, P.C. (2018). The Indo-West Pacific species of Neaxiopsis and Neaxius (Crustacea:

Axiidea: Strahlaxiidae). Memoirs of Museum Victoria 77: 15-28.

The synonymy of Axius ( Neaxius ) gundlachi var. orientalis De Man, 1925, with Axius (Neaxius?) euryrhynchus De

Man, 1905, now Neaxiopsis euryrhynchus (De Man, 1905), is confirmed. The synonymy of Axia acantha (A. Milne

Edwards, 1879), Eiconaxius taliliensis Borradaile, 1900, and Axius acanthus mauritianus Bouvier, 1914, is confirmed; they

are a single species, Neaxius acanthus. They and a second species from the Indo-West Pacific, Neaxius trondlei Ngoc-Ho,

2005, are not synonyms of Neaxius glyptocercus (von Martens, 1868), as was proposed in Sakai’s (2011) family synthesis.

Instead, a second species (from southern Queensland, Australia, Fiji and French Polynesia) close to Neaxius glyptocercus

from north-eastern Australia is diagnosed as Neaxius capricornicus sp. nov.

Keywords Crustacea, Strahlaxiidae, Neaxiopsis, Neaxius, taxonomy

Introduction

Attempts to identify specimens of Strahlaxiidae, one from

the western Indian Ocean and others from throughout the

Indo-West Pacific, led us into a web of confused names and

errors in the most recent catalogue and review of the family

(Sakai, 2011). Here, this confusion is resolved after

examination of a wide range of material from across the

Indo-West Pacific.

We test the assumption that N. acanthus (A. Milne

Edwards, 1879) is widespread by reviewing collections from

a wide geographic range, and re-diagnose the three known

Indo-West Pacific species: N. acanthus, N. glyptocercus

(von Martens, 1868) and N. trondlei Ngoc-Ho, 2005. In the

process, a fourth undescribed species is uncovered.

Species are differentiated, in part, by the number of

spines along certain margins. These can vary between

individuals, and between left and right sides. Here, for each

character we provide median numbers of spines followed by

a range or outlying value in parentheses.

The bulk of the material is lodged in the Museum

nationale d’Histoire naturelle, Paris (MNHN) (IU-

prefixes; former registration numbers with Th

prefix). Others are from the collections of the Australian

Museum, Sydney (AM); Museums Victoria, Melbourne

(NMV); Naturhistorisches Museum, Vienna (NHMW);

National Museum of the Philippines, Manila (NMCR);

University Museum of Zoology, Cambridge (UMZC);

Zoologisches Museum, Berlin (ZMB); Zoological Museum,

Hamburg (ZMH); and the Zoological Reference Collection,

Lee Kong Chian Natural History Museum (previously

known as Raffles Museum of Biodiversity Research),

National University of Singapore (ZRC). Size of specimens

is given as carapace length (cl) unless otherwise stated (total

length, tl).

Strahlaxiidae Poore, 1994

Strahlaxiidae Poore, 1994: 100.—Sakai, 2011: 319-320.

Remarks. The diagnosis of the family stands. It was elaborated

by Sakai (2011) without providing any more diagnostic

characters. Poore (1994) and Sakai (2011) both provided keys

to the three genera.

Neaxiopsis Sakai and de Saint Laurent, 1989

Neaxiopsis Sakai and de Saint Laurent, 1989: 32.—Poore, 1994:

100.—Sakai, 2011: 320.

Remarks. The genus is recognisable from the broad plate-like

rostrum with an apical notch. Sakai (2011) provided a key to

distinguish the two species but confused their synonymies.

G.C.B. Poore & PC. Dworschak

Neaxiopsis euryrhynchus (De Man, 1905)

Axius ( Neaxius ?) euryrhynchus De Man, 1905: 590.—De Man,

1925c: 3, 12, 31, pi. 1 fig. 2.

Axius ( Neaxius ) gundlachi var. orientalis De Man, 1925b: 122—

125, fig. 2, 2b.—De Man, 1925c: 4, 12, 31 (type locality: Matupi [now

Matupit I.] near Rabaul, New Britain, Papua New Guinea).

Axius ( Neaxius ) euryrhynchus.— Miyake, 1982: 90, 192 (list), pi.

30 fig. 5.

Neaxiopsis euryrhynchus.—Sakai and de Saint Laurent, 1989: 33.

Neaxiopsis euryrhyncha.— Sakai, 2011: 321-323, fig. 60.

Material examined. Reunion. Off Sainte Anne, 21° 00.6' S, 55° 43.8' E, 45

m (Expediton MD32 stn DR154), MNHNIU-2016-8079 (male, 3.5 mm).

Type locality. Anchorage off Dongala, Palos-bay, Sulawesi,

Indonesia, 36 m ( Siboga stn 86).

Distribution. Japan; Indonesia, Sulawesi; Papua New Guinea,

New Britain; Reunion; to 36 m depth.

Remarks. De Man (1925c) believed that the syntypes of Axius

(Neaxius?) euryrhynchus are “a very young stage” of Axius

(Neaxius) gundlachi orientalis, both described by him from

the south-western Pacific. He synonymised the two names.

Sakai (2011) argued first (p. 321) that Neaxiopsis euryrhynchus

and N. gundlachi von Martens, 1872, a species from the

Caribbean, are distinct species but then argued (pp. 323-324)

for the synonymy of ‘TV. orientalis’’'’ and N. gundlachi. He said

both share a row of tubercles along the carina of the pereopod

1 palm but did not explain the state of this character in N.

euryrhynchus. On purely biogeographic criteria, De Man’s

synonymy is the more probable.

Miyake (1982) recorded a female with total length of 69

mm from Japan. Sakai and de Saint Laurent (1989) doubted

this was the same species on the basis of its size. The syntypes

of A. (A.?) euryrhynchus are 11 mm long juveniles, while the

syntypes of A. (N.) gundlachi orientalis range in length from

48 to 74 mm. Miyake’s (1982) specimen is within this range.

The specimen from Reunion is very small but shares the

characteristic rostrum, cheliped dentition and tail fan.

Assuming cryptic species are not involved, the record extends

the species’ range throughout the Indo West-Pacific.

The specific name is a noun and does not follow the gender

of the genus name.

Neaxius Borradaile, 1903

Axius (Neaxius) Borradaile, 1903: 537.—De Man, 1925c: 12.

Neaxius.— Sakai and de Saint Laurent, 1989: 29.—Poore, 1994:

100.—Sakai, 1994: 176,-Sakai, 2011: 324-325.

Type species. Axia acantha A. Milne-Edwards, 1879, by original

designation.

Remarks. Specimens of the type species, now N. acanthus (A.

Milne-Edwards, 1879) from the type locality, New Caledonia,

have never been illustrated, and Milne-Edwards’ (1879)

description is too general to be certain of the species’ identity:

he described the antenna as having four or five lateral spines, the

anterior carapace margin with four or five spines, the cervical

groove with three or four spines, and the scaphocerite with one

mesial and four lower spines. This description applies to many

specimens throughout the Indo West-Pacific and the species has

been assumed to be widespread. The standard reference for

details of this species is De Man’s (1898) description and

illustrations of specimens from Sulawesi, Indonesia, not those

from the type locality. Sakai and de Saint Laurent (1989) were

the first to include type material in their appraisal of N. acanthus.

Borradaile (1903) synonymised without comment

Eiconaxius taliliensis Borradaile, 1900, with N. acanthus.

Axius acanthus var. mauritiana Bouvier, 1914, has also long

been thought to be a junior synonym. This synonymy was

accepted by Ngoc-Ho (2006) who tabulated differences

between the six accepted species of Neaxius, three from the

Indo-West Pacific and three from the Atlantic Ocean.

Sakai (2011: 329-330, figs 61, 62) took a different view and

treated all nominal Indo-West Pacific species and subspecies

- Axia acantha (type locality: New Caledonia), Eiconaxius

taliliensis (New Britain), Neaxius trondlei Ngoc-Ho, 2005

(Marquesas Islands) and Axius acanthus var. mauritianus

(Mauritius) - as synonyms of Axius glyptocercus von Martens,

1868 (Cape York, Qld, Australia), which he believed to be a

variable species. He described and figured the antenna,

carapace spination and scaphocerite of specimens from Fiji,

Tahiti, Palau, Sulawesi and Ryuku, Japan, to justify that only

one species, Neaxius glyptocercus, was distributed widely in

the Indo-West Pacific. He argued that the variability in a

population of N. acanthus from Motupore, Papua New Guinea,

studied by Mukai and Sakai (1992) (5-7 spines on the cervical

groove, 1 or 2 mesial and 3-6 lateral spines on the second

antenna article, spinose merus on pereopod 2) supports his

view, but in reality, they confirm the opposite. This population

differs consistently from N. glyptocercus in having the cervical

groove, antenna article and pereopod 2 unarmed as in all

Australian specimens examined by Poore and Griffin (1979)

and in more recently examined examples (AM, NMV, NHMW,

ZMH). To these characters can be added differences in the

shape and ornamentation of the telson. The telson of N.

glyptocercus is c. 1.3 times as wide as long, moderately

tapering, with 1 or 2 small spines along the lateral margin, with

the anterior transverse ridge reaching the lateral margins, and

the posterior concave face without ornamentation. The telson

of N. acanthus is 1.5 times as wide as long, strongly tapering,

with 1-6 tubercles above each posterolateral margin, with the

second transverse ridge one-third of the way between the first

and the posterior margin, and with a third short obsolete

transverse ridge and longitudinal lateral buttresses emerging

from the ends of the second transverse ridge.

This separation is, however, confused by the discovery that

southern and Pacific representatives of “A. glyptocercus ” are

morphologically distinct and warrant description of another

species, Neaxius capricornicus sp. nov. This confusion has led

to errors in the identification of species for which sequences are

registered in Genbank at the National Center for Biotechnology

Information (https://www.ncbi.nlm.nih.gov/nuccore/neaxius)

(Table 1).

Tsang et al. (2008) showed on the basis of three rRNA

sequences that N. acanthus from Taiwan differs from N.

capricornicus from Australia (wrongly identified as N.

glyptocercus) with 100% probability.

Species of Neaxiopsis and Neaxius

Table 1. Present identifications of species of Neaxius recorded in Genbank (National Center for Biotechnology Information)

Accession no.

Sequences

Citation

Locality/voucher

number

Identification

EF585463.1

EF585474.1

EF585452.1

18S, 28S, 16S subunits

ribosomal RNA

Tsang et al., 2008

(as N. acanthus )

Taiwan/NTOU A00421

N. acanthus

NC_019609

JN897379.1

mitochondrion, complete

genome

Lin et al., 2012

(as N. glyptocercus )

Kensley et al., 2000

(as N. acanthus)

Renting, Taiwan/NTOU

N. acanthus

KC107821.1

mitochondrion, partial

genome

Shen et al., 2013

(as N. acanthus)

Indonesia, Sulawesi/

no voucher

N. acanthus

EU874994.1

EU874944.1

18S, 16S subunits

ribosomal RNA

Tudge and Cunningham, 2002;

Tsang et al., 2008; Robles et al., 2009

(as N. glyptocercus)

Australia, S Qld/

NMV J39643

N. capricornicus

Anker et al. (2015) illustrated in colour specimens of what

are clearly N. acanthus from Lombok, Indonesia, as N.

glyptocercus, but expressed confusion over Sakai’s synonymy.

Sakai (2017) repeated his incorrect diagnosis of “ N.

glyptocercus ” and figured a cheliped from Japan clearly of the

N. acanthus form.

The four Indo-West Pacific species of Neaxius are here

diagnosed with the same character suite. Major diagnostic

characters of N. acanthus, N. capricornicus sp. nov. and N.

glyptocerus are compared in fig. 8. The distributions of all

species in the Indo-West Pacific are shown in fig. 9.

Neaxius acanthus (A. Milne Edwards, 1879)

Figures 1-5, 8a-f

Axia acantha A. Milne-Edwards, 1879: 110.

Eiconaxius acanthus.—Tit Man, 1896: 491-497.—De Man, 1898:

700, pi. 34 fig. 57 (West-Celebes = Indonesia, Sulawesi).

Eiconaxius taliliensis Borradaile, 1900: 420-421, fig. 15a-c.

Axius acanthus.— Borradaile, 1903: 537 (listed as type species of

Neaxius).

Axius taliliensis.— Borradaile, 1903: 537 (as synonym of Axius

( Neaxius ) acanthus A. Milne-Edwards, 1879).

Axius acanthus var. mauritiana Bouvier, 1914: 704.

Axius ( Neaxius) acanthus var. mauritianus.— Bouvier, 1915: 196-

198, fig. 7.—Fourmanoir, 1955 31, fig. 4.—De Man, 1925c: 3, 10, 14.

Axius ( Neaxius ) acanthus.—Tit Man, 1925a: 50-55.—De Man,

1925c: 3, 14 (part).-Poore and Griffin, 1979: 235-236, fig. 7 (Qld,

Australia).—Tirmizi, 1983: 85-88, figs 1, 2 (Maluku, Indonesia).—

Holthuis, 1953: 51 (Marianas Is, Saipan).—Miyake, 1982: 93 (Japan).—

Sakai, 1987: 303,304 (Japan).

Neaxius acanthus.—Sakai and de Saint Laurent, 1989: 30-31.—

Mukai and Sakai, 1992: 47-52, fig. 1 (Papua New Guinea).—Sakai,

1994: 200.—Kensley et al., 2000: 212, figs 5, 7F (Taiwan).—Kensley,

2003: 383 (Guam).—Kneer, 2006; Kneer, Asmus, Ahnelt and Vonk,

2008; Kneer, Asmus and Vonk, 2008 (Bone Batang I., S Sulawesi,

Indonesia).—Sakai and Sawada, 2006: 1357 (Japan).—Tsang et al.,

2008: 218-219 (Taiwan).—Tan et al., 2017: 5 (comment on name).

Neaxius glyptocercus.— Sakai, 2011: 326-331 (part) figs 61B,

62C, D (not figs 61 A, C-E, 62A, B, E, F [interchanged with 61B]), = N.

glyptocercus (von Martens, 1868.—Lin et al., 2012: 2-9 (Taiwan,

misidentification).—Anker et al., 2015: 335, figs 25, 26.—Sakai, 2017:

188, fig. 3A.

Type material examined. Lectotype of Axia acantha. New Caledonia.

MNHNIU-2014-11315 (Th812) (female, tl 72 mm, dry). Paralectotype,

MNHN IU-2014-11316 (Thl90) (male, cl 27 mm) (Fig. 1).

Syntypes of Axius acanthus mauritiana. Mauritius. Le Chaland,

MNHN IU-2014-11317 (Thl91), ovigerous female, tl 69 mm (Bouvier,

1915, listed 2 specimens from this locality). Port Louis, MNHN IU-

2014-11318 (Thl92), 1 male, tl 58 mm; 2 ovigerous females, tl 67, 62

mm (as listed by Bouvier, 1915) (Fig. 2).

Syntypes of Eiconaxius taliliensis. Papua New Guinea, New

Britain, Talili Bay, UMZC 1.57590 (male, 22.3 mm; ovigerous female,

19.4 mm) (Fig. 3).

Other material examined. Specimens marked * were listed by Sakai

and de Saint Laurent (1989).

Tanzania. Mombasa, Levin Reef, MNHN IU-2016-8073 (Th780*)

(1 individual).

Madagascar. Nosy Iranja, IU-2014-22792 (Th454*) (1 individual).

Nosy Be, IU-2014-22793 (Th456*) (2 individuals); IU-2016-8071

(Th455*) (3 individuals); NHMW 19385 (male, 26.3 mm, broken);

NHMW 19386 (1 male, 24.5 mm, telson damaged); NHMW 19387

(fragments of 2 specimens); NHMW 24999 (female, 23.7 mm +

exuvia). Nosy Be, Palm Beach Hotel Bay, sand between coral rubble,

3 m. Sainte Luce, W of Ilot Babet, 24° 46.2' S, 47° 12.4' E, 1-10 m

(ATIMO VATAE stn TA63) IU-2010-4331 (1 damaged individual).

Glorieuse Is, Grande Glorieuse I.. IU-2016-8072 (Th451*) (1

individual).

Mayotte. IU-2014-22789 (Thl565) (1 individual); IU-2014-22790

(Thl564) (1 individual).

Indonesia. Sulawesi, Bone Batang I., NHMW 25859 (male, 10.3

mm). Bali, Nusa Dua, intertidal seagrass, NHMW 25854-25858

(male 15.3 mm; 4 juveniles, 7.1-11.1 mm).

Papua New Guinea. Central Province, Motupore I., 09° 32' S,

147° 17' E, NMV J17235 (3 males, 21.9-27.5 mm; 2 ovigerous females,

20.3, 27.5 mm). Madang Province, Jais Aben Resort, Riwo, seagrass.

G.C.B. Poore & PC. Dworschak

05° 09' S, 145° 48.2' E, 1-3 m (PAPUA NIUGINI stn PR195-A),

MNHN not registered (photo only seen).

Malaysia. Sipadan, IU-2016-8070 (1 individual).

Palau. ZMH K8411 (female, 17.1 mm).

Philippines. Bohol, Panglao I., Balicasag I., NMCR 39107 (25.8

mm); NHMW 25860 (male, 25.7 mm); ZRC 2017.0415 (male, 15.5

mm); ZRC 2017.0416 (male, 17.4 mm); MNHN-2016-3495 (male, 20

mm); NMCR 39108 (female, 20.6 mm). Momo Beach, 9° 36 .V N,

123° 45.2' E, NHMW 25861 (male, 20 mm); NHMW 25862 (male,

25.8 mm). Looc, sand and seagrass with coral patches, 9° 35.7’ N,

123° 44.4’ E, 4 m, NHMW 25863 (male, 19.5 mm).

Taiwan, Pingtung County, Banna Bay, 10 m, NHMW 25919

(male, 10.4 mm), NHMW 25923 (juvenile, 7.1 mm).

Japan. Ryuku Is., Ishigakaki I., IU-2016-8078 (Th865*) (9

individuals); IU-2016-8075 - 8077 (3 dry individuals).

New Caledonia. IU-2016-8080 (Th511*) (13 individuals). Bourail,

IU-2016-8074 (Thl488) (1 individual). Ouano Plage, IU-2014-22791

(Thl486) (1 individual).

Diagnosis. Carapace supra-antennal margin without anteriorly

directed spine; anterolateral margin with 6 (4, 5) prominent

spines, dorsalmost anteriorly directed; branchiostegite anterior

margin unarmed; cervical groove with 4 (0-7) sharp spines

along posterior margin. Telson 1.3-1.5 times as wide as long;

tapering strongly from widest point to posterior margin,

posterior margin about half greatest width; anterior transverse

ridge straight, curving laterally but not reaching lateral margin

at its widest point; posterior transverse ridge situated at 0.35-

0.4 distance between anterior ridge and posterior margin;

posterolateral margin with 2-6 obsolete submarginal

tubercles; posterior face concave, with obsolete third transverse

ridge, with pair of sublateral longitudinal ridges subtended

from ends of second transverse ridge, each with 2 (1-3)

rounded tubercles, sometimes obsolete. Antenna article 2 with

1 (2) upper-mesial spine, 3 (0-6) lateral spines; scaphocerite

with 1 (2) mesial sharp spine, 5 (2-6) sharp ventral spines;

Figure 1. Axia acantha A. Milne-Edwards, 1879, lectotype, MNHN IU-2014-11315: a, dorsal view; b, lateral left view; c, lateral right view; d,

telson and uropods. Paralectotype, MNHN IU-2014-11316: e, dorsal view; f, lateral left view; g, telson and uropods.

Species of Neaxiopsis and Neaxius

article 4 lower margin with 3 (0-5) sharp spines. Cheliped

merus, lower margin with 4 (2-5) spines, lateral face with

curved row of 8 (5-10) spines. Pereopod 2 merus, lower

margin with row of 10 (8-17) spines. Pereopod 3 merus, lower

margin without row of spines.

Remarks. A. Milne-Edwards (1879) did not specify how many

specimens he had. Two remain in MNHN, one of which Sakai

and de Saint Laurent (1989) called the type. This was an

effective lectotype designation. This and many more from New

Caledonia were examined.

Figure 2. Axius acanthus mauritiana Bouvier, 1914, syntype, MNHN IU-2014-11317: a, dorsal view; b, lateral left view; c, telson and uropods.

Syntypes, MNHN IU-2014-11318: d, dorsal views; e, lateral views, f, telsons and uropods.

G.C.B. Poore & PC. Dworschak

Bouvier (1914, 1915) distinguished his variety from

Mauritius, Axius acanthus mauritianus, on differences in the

denticulation of the median rostral ridge, the number and

nature of the spines on the anterolateral margin of the

carapace and cervical groove, spination of the merus of the

cheliped, and ornamentation of the pleonal epimera.

Fourmanoir (1955) recorded the variety from the Comores.

Consistent differences from material from New Caledonia

could not be detected during our examination of numerous

specimens from the western Indian Ocean. Spines on the

carapace and antenna varied in number and strength, even

within a small geographic range. Spination along the ventral

margins of pleonal epimera 2-4 ranged from non-existent to

prominent, as it does in other populations. The longitudinal

ridges extending from the second transverse ridge usually

carried one or two small tubercles. In some specimens from

Japan a single large tubercle dominated but not in others from

the same location.

Figure 3. Eiconaxius taliliensis Borradaile, 1900, syntype, male, UMZC 1.57590: a, lateral view; b, telson.

Figure 4. Neaxius acanthus (A. Milne-Edwards, 1879): a, b, Papua New Guinea (PAPUA NIUGINI stn PR195-A), MNHN unregistered, photos,

A. Anker; c, d, Philippines, Looc, NHMW 25863, photos, T.-Y. Chan.

Species of Neaxiopsis and Neaxius

De Man (1896) was the first to apply the name acanthus to

specimens of Neaxius from Sulawesi, Indonesia (as Celebes)

and later from New Britain, Papua New Guinea (as Nouvelle

Pomeranie) (De Man, 1898, 1925a). Borradaile (1900)

introduced Eiconaxius taliliensis for representatives from this

region but soon synonymised it with A. acanthus (Borradaile,

1903). We examined the syntypes of E. taliliensis and several

specimens from nearby but could not detect consistent

differences between them, nor could we detect differences

from individuals from Malaysia, Palau, Philippines, Taiwan or

Japan. Similarly, the syntypes of Axius acanthus mauritiana

resembled others from the western Indian Ocean no more than

they did those from the western Pacific. The numbers of spines

on the carapace and cheliped varied over a small range

between individuals and from one side of an individual to the

other, as did the expression of spines, some individuals having

more prominent spines than others (see figs 1-4 and especially

5). This variability was not correlated with locality.

Sakai (2011) listed Axius acanthus mauritianus,

Eiconaxius taliliensis and Neaxius trondlei Ngoc-Ho, 2005 in

the synonymy of N. glyptocercus. While the synonymy of

Axius acanthus mauritianus and Eiconaxius taliliensis with

N. acanthus is supported on morphological grounds, N.

acanthus differs from N. glyptocercus in the many ways

tabulated by Ngoc-Ho (2005). Neaxius trondlei from French

Polynesia differs in spination from N. glyptocercus and N.

acanthus for the reasons given by Ngoc-Ho (2005) (see

diagnosis below).

Colour photos of live specimens indicate that the species is

generally orange with stronger pigmentation on the chelipeds,

anterior carapace and tailfan (Fig. 4; Anker et al., 2015: fig. 25).

Neaxius capricornicus sp. nov.

(http://zoobank.org/urn:lsid:zoobank.org:act:0FEDB7A0-7265-

4E5B-A26D-861FFFB7F7B6)

Figures 6, 8g-l

Axius ( Neaxius ) glyptocercus .—Poore and Griffin, 1979: 236-238

(partim), figs 8g-i.

Neaxius glyptocercus.— Tudge and Cunningham, 2002: 841.—

Tsang et al., 2008: 218-219.-Robles et al., 2009: 316,-Sakai, 2011:

326-331 (partim) figs 62A, B, E.

Figure 5. Neaxius acanthus (A. Milne-Edwards, 1879), anterior carapace: a. New Caledonia, IU 2014-22791; b, Madagascar, IU 2014-22792; c, Japan,

IU 2016-8076; d, Tanzania, IU 2016-8073. Papua New Guinea, NMV J17235: e, habitus; f, dorsal carapace; g, telson and uropods. Various scales.

G.C.B. Poore & PC. Dworschak

Material examined. Holotype. Australia, Qld, North Stradbroke I.,

Deanbilla Bay, Dunwich, IT 30' S, 153° 24’ E, NMV J39643 (female,

26 mm; see Tudge and Cunningham [2002]),

Paratypes. Collected with holotype. NMV J71641 (female, 23

mm); NMV J40714 (2 females, 26 mm; male, 21 mm).

Australia, Qld, North Stradbroke I., Dunwich, 27° 30' S,

153° 24' E, AM P.13723 (male, 27 mm). Capricorn Group, North West

I., 23° 18’ S, 151° 42’ E, AM P.10060 (female, 38 mm), AM P.11829

(female, 30 mm).

Other material. Fiji, Viti Levu, ZMH K8392 (Godeffroy No. 7430)

(Sakai, 2011: fig. 62A, E) (male, 14.3 mm).

French Polynesia, Tahiti. ZMH 41226 (Sakai, 2011: fig. 62B)

(male, 27 mm).

Figure 6. Neaxius capricornicus sp. nov., holotype, NMV J39643: a, lateral carapace, merus of cheliped; b, anterior carapace; c, telson; d, telson,

right uropod; e, f, pereopods 2, 3. Paratype, NMV J71643: g, pereopod 4; h, habitus lateral; i, dorsal carapace j, telson and uropods. All

pereopods, lateral faces. Scale bars = 5 mm.

Species of Neaxiopsis and Neaxius

Diagnosis. Carapace supra-antennal margin with anteriorly

directed spine; anterolateral margin with 6 (5-7) spines,

dorsalmost anterolaterally directed; branchiostegite anterior

margin with 1 spine; cervical groove without spines along

posterior margin. Telson 1.3-1.5 times as wide as long; tapering

from widest point to posterior margin; anterior transverse ridge

straight, curving laterally to reach lateral margin at its widest

point; posterior transverse ridge situated at 0.5 distance

between first transverse ridge and posterior margin, ends

sharply rounded, almost overhanging; lateral margin with 0-2

marginal teeth; posterior face concave, with shallow median

groove, smooth sublaterally. Antenna article 2 without upper-

mesial spine, without lateral spine; scaphocerite with 1 mesial

sharp spine, 2 (1-4) sharp ventral spines; article 4 lower margin

without spines. Cheliped merus, lower margin with 5 (4-6)

spines, distolateral face with row of 4 (3-5) spines. Pereopod 2

merus, lower margin with 6 (3-11) spines. Pereopod 3 merus,

lower margin with 8 (4-12) spines.

Supplementary description of holotype. Rostrum with 5

pairs of erect blunt spines; sharp hiatus before smooth lateral

carina; median carina with 5 tubercles; anterior gastric

region rugose; cervical groove defined posteriorly by sharp

carina; branchiostegal groove separating smooth cardiac

region from punctate branchiostegal region. Anterolateral

margin with 6 spines on right, 5 on left, first flaring laterally,

longer gap between third and fourth, between fifth and sixth;

anterior branchiostegal margin with 1 short spine. Pleomere

1 pleuron with 3 tubercles; pleomere 2 with 7 tubercles.

Telson 1.35 times as wide as long; widest at prominent lateral

lobes, at c. 0.4 of length; tapering sharply then gradually

from widest point to posterior margin; posterior margin c.

0.6 times greatest width; anterior transverse ridge at c. 0.25

length, straight, curving laterally to reach lateral margin at

its widest point; posterior transverse ridge situated at 0.5

distance between first transverse ridge and posterior margin,

ends sharply rounded, almost overhanging; lateral margin

without marginal teeth; posterior face concave, with shallow

median groove, smooth sublaterally. Antenna article 2

without mesial spine, without lateral spine; scaphocerite with

1 mesial sharp spine, 3 sharp ventral spines; article 4 lower

margin without spines. Maxilliped 3 merus with 1 short, 2

longer distal spines. Cheliped coxa with 2 spines; basis with

1 spine; ischium with 3 spines; merus, lower margin with 4

spines on right, 5 on left, distolateral face with row of 3

spines, upper margin with 4 spines; carpus lower margin

with 1 distal spine. Pereopod 2 coxa with 2 spines; basis with

2 spines; ischium with 3 spines; merus, lower margin with 7

spines on right, 6 on left, more proximal one minute.

Pereopod 3 coxa with 2 spines; basis with 2 spines; ischium

without spines; merus, lower margin spines in 2 rows: 6

spines mesially, last 3 minute, 6 laterally, last minute.

Pereopods 4 and 5 without spines.

Etymology. For the Tropic of Capricorn, which marks the

species’ northern limit in Queensland, Australia. The name is

a noun in apposition.

Distribution. Australia, Queensland, 23°S-27° 30' S; Fiji;

French Polynesia.

Figure 7. Neaxius glyptocercus (von Martens, 1868), ovigerous female, AM P.18842: a, lateral carapace, merus of cheliped; b, anterior carapace;

c, pereopod 2. Female, AM P.16177: d, e, pereopods 3, 4; f, telson. All pereopods, lateral faces. Scale bar = 10 mm.

G.C.B. Poore & PC. Dworschak

Remarks. Neaxius capricornicus and N. glyptocercus are

immediately differentiated from N. acanthus in having a

prominent supra-antennal spine on the anterior margin of the

carapace, and both the cervical groove and second antenna

article unarmed. Neaxius capricornicus differs from N.

glyptocercus in having: 2-4 spines on the lower margin of the

scaphocerite (vs. usually none, rarely 1 or 2 in N. glyptocercus ),

3-5 spines along the distolateral ridge of the merus of the

cheliped (vs. usually none, rarely one), 3-11 spines on the

lower margin of the merus of pereopod 2 (vs. none), 8-12

(rarely fewer) spines on the lower margin of the merus of

pereopod 3 (vs. none), one spine on the anterior margin of the

branchiostegite (vs. 2 or 3) and the dorsalmost spine of the

anterolateral carapace margin directed anterolaterally

(vs. anteriorly).

The two species were confused by Poore and Griffin

(1979) who illustrated both in their fig. 8.

Specimens from Fiji (ZMH K8392; Sakai, 2011: fig. 62A,

E) and Tahiti (ZMH K41226; Sakai, fig. 62B) have 2 spines

on the lower margin of the scaphocerite, 4 and 3 spines

respectively along the distolateral ridge of peropod 1 merus,

4 and 10 spines respectively on the lower margin of pereopod

2, 8 and 10 spines respectively on the lower border of

pereopod 3 merus and 1 spine on the anterior margin of the

branchiostegite, within the range of the Australian material.

Neaxius glyptocercus (von Martens, 1868)

Figures 7, 8m-r

Axius glyptocercus von Martens, 1868: 613-614.—Haswell, 1882:

165-166.

? Axius ( Neaxius ) glyptocercus.— Borradaile, 1903: 537.

Axius ( Neaxius ) glyptocercus.— De Man, 1925a: 50-56, fig. 1.—

De Man, 1925c: 4, 13.—Poore and Griffin, 1979: 236-238 (partim),

figs 8a-f, k.

Neaxius glyptocercus.— Sakai, 1994: 200.—Sakai, 2011: 326-331

(partim), figs 61A, C-E, fig. 62F (pereopod 2 of holotype, mislabelled)

(not figs 62A, B, E = N. capricornicus-, not fig. 61B [interchanged with

62F] = N. acanthus-, not figs 62C, D = N. acanthus).

Material examined. Holotype. Australia, Qld, Cape York, ZMB 2973

(described and anterior carapace figured by De Man (1925c: 50-56,

fig. 1); rostrum, antenna, pereopods 2,4, 5, pleopod 2 figured by Sakai

(2011: 326-331, figs 61A, C-E, 62A, B, F).

Other material. Australia, Qld, Cape York, Fly Point, 10° 45' S,

142° 37’ E, AM P.24813 (2 females, 13, 18 mm; male, 17 mm).

Townsville area, 19° 16' S, 146° 49' E, AM P.16176 (male, 20 mm).

Mossman, Cooya Beach, 16° 26' S, 145° 24’ E, NHMW 19591 (3

females, 13.5-24.7 mm). Cannonvale Beach, near Bowen, 20° OP S,

148° 15’ E, AM P.16177 (female, 22 mm).

NT, Darwin, Lee Point, 12° 20’ S, 130° 54’ E, AM P.20358

(female, 21 mm). Port Darwin, 12° 27' S, 130° 48' E, AM P.15030

(male, 19 mm). Nightcliff, Darwin, 12° 23’ S, 130° 50’ E, AM P.18842

(ovigerous female, 28 mm).

Diagnosis. Carapace supra-antennal margin with anteriorly

directed spine; anterolateral margin with 4 (5-7) spines,

dorsalmost anteriorly directed; branchiostegite anterior

margin with 2 or 3 spines cervical groove without spines

along posterior margin. Telson about 1.3 times as wide as

long; tapering slightly from widest point to posterior margin;

anterior transverse ridge straight, curving laterally to reach

lateral margin at its widest point; posterior transverse ridge

situated at 0.5 distance between first transverse ridge and

posterior margin, ends sharply rounded, almost overhanging;

lateral margin with 0-2 marginal teeth; posterior face

concave, with shallow median groove, smooth sublaterally.

Antenna article 2 without upper-mesial spine, without lateral

spine; scaphocerite with 1 mesial sharp spine, 1 (2) sharp

ventral spine; article 4 lower margin without spines. Cheliped

merus, lower margin with 3 or 4 spines, distolateral face

without or rarely with 1 spine. Pereopod 2 merus, lower

margin without spines. Pereopod 3 merus, lower margin

without spines.

Distribution. Australia: Northern Territory, E of Darwin;

Queensland, Cape York to Bowen; 10° 45' S-20° S.

Remarks. Differences between N. glyptocercus and N.

capricornicus were outlined above. The species is confined to

north and north-eastern Australia. The holotype was

photographed for us by C. Oliver Coleman and we were able to

confirm the spination of the pereopods.

Neaxius trondlei Ngoc-Ho, 2005

Neaxius trondlei Ngoc-Ho, 2005: 59-63, figs 6, 7.

Material examined. French Polynesia, Marquesas Is, Ua Huka, Hane

Bay (MUSORSTOM 9 stn 19), MNHN Thl419 (holotype male, 29

mm), MNHN Thl427 (paratypes: male, 30 mm; female, 21 mm). W of

Haamamao Bay, MNHN Thl428 (paratype female, 13.5 mm).

Diagnosis. Carapace supra-antennal margin without

anteriorly directed spine; anterolateral margin with 3 or 4

spines, dorsalmost anterolaterally directed; branchiostegite

anterior margin unknown; cervical groove with 2 or 3 spines

along posterior margin. Telson 1.5 times as wide as long;

tapering strongly from widest point to posterior margin,

posterior margin about 0.6 greatest width; anterior transverse

ridge straight, curving laterally but not reaching lateral

margin at its widest point; posterior transverse ridge situated

at half distance between anterior ridge and posterior margin;

posterolateral margin without tubercles; posterior face

concave, with well-defined third transverse ridge, without

pair of sublateral longitudinal ridges subtended from ends of

second transverse ridge. Antenna article 2 with 1 or 2 upper-

mesial spines, without lateral spines; scaphocerite with 1

mesial sharp spine, 5 sharp ventral spines; article 4 lower

margin without spines. Cheliped merus, lower margin with 3

or 4 spines, lateral face without row of spines. Pereopod 2

merus, lower margin without spines. Pereopod 3 merus, lower

margin without spines.

Remarks. Neaxius trondlei is distinguished by the unique

combination of no spines on the lateral margin of the second

article of the antenna and no spines on the lateral face of the

cheliped or on the lower margins of pereopods 2 and 3. The

species and N. capricornicus both occur in French Polynesia

but c. 1400 km apart. They can be differentiated by the presence

of spines along the cervical groove and spines on article 2 of

the antenna.

Species of Neaxiopsis and Neaxius

Figure 8. Neaxius acanthus (A. Milne-Edwards, 1879), NMV J17235, male, 26 mm: a-f. Neaxius capricornicus sp. nov., holotype, NMV J39643:

g-k. Paratype, NMV J71641: 1. Neaxius glyptocercus (von Martens, 1868), AM P.16177: m-r. Note: a, b, g, h, m, n, dorsal and lateral views of

anterior carapace; c-f, i—1, o-r, pereopods 1-4. Pereopods 1, 2 in lateral view, pereopods 3, 4 in mesial view. Scale bar = 5 mm.

C Cccc

G.C.B. Poore & PC. Dworschak

Figure 9. Distribution of four species of Neaxius in the Indo-West Pacific (based on material examined).

Acknowledgements

We are grateful to Laure Corbari, Paula Lefevre-Martin and

Anouchka Krygelmans-Sato for help in making the collections

available at MNHN. Matthew Lowe (UMZC) and Sammy De

Grave (Oxford) arranged for access to the syntypes of

Eiconaxius taliliensis. Angelika Brandt and Kathrin Philips-

Bussau (ZMH) provided loans. Noemy Malloret (MNHN) and

C. Oliver Coleman (ZMB) photographed type specimens.

Arthur Anker provided a colour photograph of an individual

from Papua New Guinea and Tin-Yam Chan provided a colour

photograph of an individual from Panglao.

Part of the material studied was collected during the

Panglao Biodiversity Project 2004, which was made feasible

through grants from the Total Foundation for Biodiversity and

the Sea, the French Ministry of Foreign Affairs, and the

ASEAN Regional Centre for Biodiversity Conservation. The

Philippines Bureau of Fisheries and Aquatic Resources is

acknowledged for issuing a research permit. PCD thanks the

principal investigators, Philippe Bouchet (MNHN) and Danilo

Largo (University of San Carlos, Cebu, Philippines) for the

invitation to participate in the field work.

Finally, GCBP acknowledges the support of Philippe

Bouchet (MNHN) for supporting participation during and

after the Madang Lagoon Biodiversity Survey, part of the of

La Planete Revisitee expedition to Papua New Guinea, and the

Crosnier Fund for financial support in Paris.

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Memoirs of Museum Victoria 77:29-40 (2018) Published 2018

1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.Org/10.24199/j.mmv.2018.77.03

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria:

historical records and contemporary uncertainty

Nick Clemann 1 ’ 2 , Timothy Stranks 2 , Rebecca Carland 2 , Jane Melville 2 , Bianca op den Brouw 3 and

Peter Robertson 4

1 Arthur Rylah Institute for E nvironmental Research, Department of Environment, Land, Water and Planning, PO Box 137,

Heidelberg VIC 3084 (corresponding author. Email address: nick.clemann@delwp.vic.gov.au)

2 Museums Victoria, 11 Nicholson St, Carlton VIC 3053 (Email addresses: TStranks@museum.vic.gov.au; rcarland@

museum.vic.gov.au; jmelv@museum.vic.gov.au)

3 Venom Evolution Lab, School of Biological Sciences, University of Queensland, St Lucia QLD 4072 (Email address:

b.opdenbrouw@uq.net.au)

4 Wildlife Profiles Pty Ltd, PO Box 572, Hurstbridge VIC 3099 (Email address: wildlife.profiles@bigpond.com)

Abstract Clemann, N., Stranks, T., Carland, R., Melville, J., op den Brouw, B. and Robertson, P. 2018. The death adder

Acantophis antarcticus (Shaw & Nodder, 1802) in Victoria: historical records and contemporary uncertainty. Memoirs of

Museum Victoria 77: 29-40.

The south-eastern distributional limit of many Australian species coincides with northern, and sometimes far-

eastern, Victoria. In the mid-19th century, Blandowski's Lower Murray Expedition sought to study the natural history of

this area, specifically north and north-western Victoria. The expedition collected many specimens that are now registered

with Museums Victoria, including species that are now extinct, extinct in the state or greatly reduced in distribution.

During the expedition, a specimen of the death adder Acanthophis antarcticus was collected at Lake Boga in north¬

western Victoria. During the 20th and 21st centuries, there has been debate about whether this species persists in Victoria.

We review early records of this species, including voucher specimens held by Museums Victoria, one of which we confirm

as the specimen collected during Blandowski’s Lower Murray Expedition. We also explore recent claims of sightings of

this species in Victoria. We collate names for the death adder used by Aboriginal people in northern and north-western

Victoria. Death adders undoubtedly occurred in north-western Victoria in the 19th century and were known to the

Aboriginal people, but it is probable that they no longer occur in that part of the state. It is possible that death adders persist

in far East Gippsland, east of the Wallagaraugh River, although no substantiating material, such as photographs or

specimens, has been collected in that area.

Keywords Gerard Krefft, Blandowski Lower Murray Expedition, voucher specimen, venomous snake

Introduction

The death adder Acanthophis antarcticus is a distinctive and

iconic venomous snake found over much of eastern and

southern continental Australia, except for the cooler parts of

the south-east (Cogger, 2014, fig. 1). The south-eastern limit of

the species’ distribution is generally accepted to be in or

abutting northern Victoria and far East Gippsland. The pre¬

eminent national guide to the herpetofauna of Australia since

the 1970s is Reptiles & Amphibians of Australia by Harold

Cogger (1979). Early editions of this book described the

species’ distribution as “throughout continental Australia,

except central desert regions and wetter parts of Vic and

south-eastern NSW” (Cogger, 1979, p. 373), and the

accompanying map included the species in far north-western

Victoria. However, the latest edition (Cogger, 2014) describes

this part of the species’ range only as “through parts of

southern and south-eastern Australia”, and the accompanying

map no longer includes north-western (or any) parts of Victoria

(p. 856). There are few reliable accounts of the species’

occurrence in the state (the species is listed as Data Deficient

by the Department of Sustainability and Environment, 2013),

and there is uncertainty about the provenance of the three

specimens held by Museums Victoria that are labelled as

being from Victoria. We sought to document references to

death adders in Victoria from the literature, review the

provenance of putative Victorian death adder voucher

specimens in the collection of Museums Victoria and, lastly,

review selected verbal accounts of death adders from Victoria.

Acceptable records of the death adder in Victoria

Voucher specimen from Blandowski’s Lower Murray Expedition

(1856-57)

N. Clemann, T. Stranks, R. Carland, J. Melville, B.Op Den Brouw & P. Robertson

Figure 1. South-eastern Australia, showing records of the death adder Acanthophis antarcticus (black dots; Atlas of Living Australia, year) and

key localities discussed in the text.

The occurrence of the death adder in Victoria was

confirmed during Blandowski’s Lower Murray Expedition in

the mid-19th century. Confirmation came “from the diary of

Gerard Krefft who, in 1856, drew the head and the tail of a

specimen from Lake Boga” (Coventry and Robertson, 1991, p.

22; however, see below for a correction to the year of this

record). Our recent re-examination of the relevant text in

Krefft’s narrative (which is not really a diary per se ) hinted at

the possibility of there being more information regarding this

record and prompted a re-evaluation of this species in Victoria.

Krefft (c. 1858) provides a rather detailed description of the

finding, capture and documentation of this snake in 1857, the day

after the expedition arrived at the Lake Boga Mission Station:

Sunday Morning the 8th of March found us busy as ever for

Mr Blandowski begged of us to remember the holy day we

had the night before which as usual silenced all opposition. I

had been up early and was busy transferring a toad to paper

which I had caught when my attention was roused in repeated

“queeing” (a loud noise used in the bush to call at one another).

First I took no notice of it but the Captain told me to see what

was wrong and so I left the hut.

I found our Cook in the deserted garden of the mission

wrangling with a snake, the head of which he had secured to

the ground like a sensible fellow with a forked stick. He

informed me that he had been looking for some tomatoes and

lifting one of the bushes he nearly touched the snake. A stick

with a prongue to it was soon found and there he was.

I took charge of the reptile and told him to report to the

Captain who immediately came up to the field of action and

gave directions to take the snake alive. Now I have had a great

deal of experience in snakes and secured many a one alive,

but as the specimen in question was to all appearances a

Death Adder, supplied with a poisonous sting on the end of

the tail; I did not like to take it up. So after a few words with

the Captain, I asked him to show me how to catch it, which he

instantly did by taking hold of the snake behind the jaws. I

had tied a bit of cord around the tail to prevent the snake from

making use of it and so we carried the ugly customer to the

hut. Mr B1 made several vain attempts to pass the snake off on

myself but as I thought that he had a good grip of it, I begged

to be excused. We pinned it down on a board as I set

immediately to work to secure its colours while alive. I might

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria

as well state here a fact often observed, that every snake or

part of a snake, however mutilated will live until the sun goes

down and though the neck had been cut through this snake

lived until evening.

Our commander considered the reptile to be a new species

which I doubt as it is by all appearance Brown’s Death Adder.

Having made a minute drawing of it I am able to describe all

its peculiarities. It measured about 30’ and was rather thick in

proportion to its length. The head flat and the scales distributed

in the same manner as in most all other Australian snakes. Of

a Brown colour, all the scales on the back were divided or

riveted vide sketch in the margin. The borders of the scales on

the head turned upwards. The belly was of a pale pink colour

with the side scales dotted with darker pink spots not unlike

those which the edges of books are ornamented vide sketch.

From the abdomen to the end of the tail about 5’, the tail all at

once becoming very thin. The specimen is now in the

Melbourne Museum although much destroyed by the dirty

fluid in which Mr B1 attempted to preserve his specimens.

Blandowski (1858) mentions an incident that probably relates

to this encounter: “At Lake Boga I was exposed to some danger

in presence of my men, by a very poisonous snake, on which I

had inadvertently placed my feet” (p. 135). The statement in

the final sentence of the excerpt from Krefft’s narrative - that

a specimen had been lodged with the Museum in Melbourne

- aroused our interest. Cursory inspection of the death adders

in the Museum’s collection immediately revealed the most

promising candidate, specimen D4349 (fig. 2). The label on the

jar that holds this specimen mentions a date and locality that

approximates the era and localities of the Lower Murray

Expedition: the “Banks of the Murray” in 1859. The Lake

Boga Moravian Mission Station was established by German

missionaries Brother Andrew Frederick Charles Taeger and

Brother Frederich W. Spieseke on the south-eastern shore of

Lake Boga in 1851 and abandoned on 27 May 1856 (Kenny,

2003, which contains a sketch of the mission location on page

104). This location is approximately seven to eight kilometres

south of the Murray River, and a little over five kilometres

south of the Little Murray River channel (fig. 1).

Two other features of specimen D4349 accord well with

Krefft’s narrative. First, the length of the snake was estimated

by Krefft to be “about 30” inches (this measurement is repeated

on Krefft’s illustration of the Lake Boga adder, pencilled

adjacent to the full-body image). Using a piece of string to run

along the body of specimen D4349, we measured its total

length at 728 mm (28.7 inches), consisting of 635 mm (25

inches) snout to vent and 93 mm (3.7 inches) tail length.

However, the tip of the tail has been lost, either deliberately cut

off or broken, since collection (fig. 3). Given Krefft’s concerns

about the “poisonous sting on the end of the tail”, it is plausible

that the tail tip was removed while the snake was being

processed by Blandowski and Krefft. With a complete tail,

this specimen would be very close to 30 inches in total length

and the tail would be very close to the five inches mentioned

by Krefft. Second, Krefft notes that “the neck had been cut

through”. Specimen D4349 has an obvious broad wound on

the dorsal surface of the neck (fig. 4). Certainty that this is the

ELAPfDAH

^ c &nthophis ant& rcti

D 4349

Banks of it * 0

1859

Ex NMV S930

Figure 2. Specimen D4349, a death adder Acanthophis antarcticus in

the collection of Museums Victoria.

death adder collected at Lake Boga in 1857 required careful

comparison with the detailed and accurate illustration

prepared by Krefft on the day the snake was captured.

Krefft’s illustration of the death adder is held in the

collection of the Historische Arbeitsstelle (Historical

Collections department) at the Museum fur Naturkunde,

Berlin (reg. no. MfN, BVIII / 56). The illustration (fig. 5) is in

pencil, ink and watercolour, and includes an image of the

whole animal, with details of the head and tail along with

some short explanatory notes. It is clearly by Krefft’s hand and

in his style (Stranks, in prep.).

Krefft’s illustration was retained by Blandowski among

the bound portfolios of illustrations assembled during his time

in Australia, and it would have stayed with Blandowski during

1857 to 1859 while he was back in Melbourne after the Lower

Murray Expedition. Although all of the natural history

specimens collected during the 1856-57 Lower Murray

N. Clemann, T. Stranks, R. Carland, J. Melville, B.Op Den Brouw & P. Robertson

Figure 3. The truncated tail tip of death adder specimen D4349 (left) compared with the full tail tip of specimen D3579 (right).

Figure 4. Dorsal perspective of the head and neck (including neck

wound) of specimen D4349.

Expedition were lodged at the National Museum in Melbourne,

Blandowski retained possession of most of the illustrations

and papers pertaining to the expedition (including the death

adder illustration); he was eventually granted permission from

the Victorian Government to retain this material for further

study, and it went with him to Europe in March 1859 (T.

Stranks, unpublished data).

Blandowski’s portfolios returned with him to his hometown

in Gleiwitz, Upper Silesia (now Gliwice, Poland) in 1860

(Darragh, 2009). They were part of the collection that stayed

with Blandowski’s family in Gleiwitz after he was committed

to the Bunzlau mental asylum in September 1873. Not long

after Blandowski’s death in December 1878, his sister

Clementine donated the collection to the Konigliche Bibliothek

zu Berlin (Royal Library, Berlin) in August 1881. The natural

history-related material from the collection was eventually

transferred to the Zoologisches Museum in Berlin in a series of

moves in 1882, 1884 and 1885 (and has remained with that

institution, which is now known as the Museum fur Naturkunde;

see Darragh, 2009; Landsberg and Landsberg, 2009).

We have a high-resolution version of the illustration (200

MB TIF / 17 MB JPEG; fig. 5), which allows the fine detail of

the illustration and annotations to be seen. Consequently, we

can read Blandowski’s handwriting in pencil with “Lake

Boga” (bottom left), “sehr giftig, nur 2 bis 2 1/2 ‘ lang” (very

poisonous, only 2-2.5 feet long; bottom right), and Krefft’s

handwriting, with “Snake A” and “30 inches” (centre right)

and “Death Adder” (bottom centre).

On 17 July 2017, we compared Krefft’s illustration with

the following six specimens of death adder from the collection

at the Melbourne Museum: D3579, D51857, D15392, D15394,

D76869 and D4349. These specimens were chosen because

their labels suggested that they might have come from Victoria,

or their collection locality is not mentioned on the label of the

receptacle in which they are kept at the Museum (i.e. all the

other death adder specimens in the Museums Victoria

collection are labelled as being collected from other states).

Using a stereomicroscope, we examined details of scalation

and colour, primarily on the ventral surface of the heads of

each snake, comparing them to the illustration. Scale and

colour were highly variable between specimens. D4349 was

the only exact match for the illustration of the death adder

from Lake Boga, confirming beyond doubt that this was the

specimen collected in 1857 by Blandowski, Krefft and the

expedition’s cook (fig. 6, which includes images of the two

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria

Figure 5. Illustration by Gerard Krefft of the death adder collected at Lake Boga in north-western Victoria on the 8 March 1857. (Photograph by

Rebecca Carland; Museum of Natural History Berlin. Historical collection of pictures and writings. [Sigel: MfN, HBSB.] Bestand: Zool. Mus.

Signatur: B VIII/56.)

N. Clemann, T. Stranks, R. Carland, J. Melville, B.Op Den Brouw & P. Robertson

Figure 6. Ventral view of the head scales of Death Adder specimens from Melbourne Museum, and close up of Gerard Krefft’s illustration of the

ventral head scales of the Death Adder collected at Lake Boga in 1857 (bottom right). Top left is specimen D3579. Top right is D51857. Bottom

left is D4349.

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria

other death adder specimens with “Victoria” on their labels,

allows comparison of the three specimens with Krefft’s

illustration and shows that only D4349 is a precise match for

the snake collected at Lake Boga in 1857).

There are minor discrepancies around the dates of this

record. Coventry and Robertson (1991) state the year of the

record as 1856; however, this was the year that Blandowski’s

Lower Murray Expedition commenced (departing Melbourne

on 6 December 1856; Allen, 2009a; Darragh, 2009), and

Krefft’s narrative is clear that the death adder was found and

captured on 8 March 1857. Specimen D4349 is labelled in the

old National or Public Museum register as dating from 1859.

Although two years later than the collection date of the Lake

Boga adder, it is plausible that either the final digit of the date

was transposed incorrectly (the handwriting of the digit 7 in

the museum register resembles a 9 to some degree) or - more

likely - that the more recent date represents the date the

specimen was registered or catalogued at the museum. Many

of the collections from Blandowski’s Lower Murray Expedition

in the Museums Victoria collection were simplistically

labelled and registered as coming from the “Junction of the

Murray and Darling Rivers” (the expedition’s official

destination), even for specimens collected considerable

distances from the location (Wakefield, 1966); the date given

in the register is often the date of registration rather than the

date of collection (discussed by Wakefield, 1966). The

collection from Lake Boga and Mondellimin, the expedition’s

camp (now known as Chaffey Landing, near Mildura; fig. 1),

would have arrived in Melbourne by mid-1857. By this stage,

the Public Museum was under Frederick McCoy’s control, and

was located in the north wing of the Quadrangle Building at

the University of Melbourne. The material may have been

stored there, largely unworked, until further sorting took place

in readiness for the Museum’s first collection registration

system in c.1858-1859.

Alternatively, perhaps due in part to his feud with McCoy

(Allen, 2009a), Blandowski was loath to hand over specimens

to the Museum, instead transferring them to his private

lodgings (Pescott, 1954, cited in Wakefield, 1966). Allen

(2006) states:

In 1858, Blandowski received a letter from the Surveyor-

General requesting him to surrender all drawings and

memoranda relating to the Natural History of the country,

made during the period he held an appointment in the

Government Service. Blandowski’s reply states his position,

“I deny ... the justice of this demand, and as I regard these

papers and drawings my private property, I must decline to

allow them to pass out of my hands” (Paszkowski 1967:160,

quoting Blandowski letter of 23 November, 1858) (p. 33).

Thus, it is plausible that Blandowski begrudgingly released

specimens over the period spanning his return from

Mondellimin and his departure for Germany in 1859, perhaps

resulting in the specimen now labelled as D4349 not entering

the collection until 1859.

Interestingly, Krefft perpetuates two myths about snakes

in his narrative. Krefft was a knowledgeable zoologist, so his

belief that it is “a fact often observed, that every snake or part

of a snake, however mutilated will live until the sun goes down”

is odd for an experienced collector who had presumably killed

and prepared many reptiles. Less surprising (for the era) is

Krefft’s belief that death adders have “a poisonous sting on the

end of the tail”. By the mid-19th century, the European

colonisers still had much to learn about the biology of many

Australian species, and the specialised tail of the death adders

was clearly believed to be a venomous adjunct to the snake’s

fangs (the tail tip of death adders usually has a terminal spike

that resembles a venomous sting; fig. 3). Death adders use their

specialised tails to lure small vertebrate prey, and early

observers who witnessed the outcome for animals lured to the

snake’s tail may have believed that the tail indeed contained a

venomous sting. However, Krefft later corrected this fallacy in

his book on Australian snakes, where he states that the tip of

the death adder’s tail, “which is so much dreaded by many

persons, is neither a weapon of attack or defence” (Krefft,

1869, p. 80).

Intriguingly, the text on the receptacle holding specimen

D4349 is a precise match for the words used by another natural

history illustrator from that period in relation to some snake

specimens. Ludwig Becker was the artist and naturalist who

accompanied the Burke and Wills exploring expedition in

1860-61, and he wrote the following letter (held in the

collection of the State Library of Victoria) to Dr John

Macadam (Honorary Secretary of the Royal Society of

Victoria’s Exploration Committee) while the expedition was

staying at Camp 15 at Swan Hill on 8 September 1860,

regarding a bottle of three snakes to be donated to the Museum

by Dr Benjamin Gummow (doctor at Swan Hill from 1857-72):

Camp at Swan Hill

Sept. 8. 1860

To Dr J Macadam MLA

Hon. Secretary

Royal Society in Victoria.

PS. By this mail I have the honor to forward to you a bottle

containing 3 snakes presented to the Museum by Dr

Gommow [sic] of Swan Hill. The 2 larger snakes were found

near Swan Hill on the Banks of the Murray, the small one in

the Mallee Scrub, 40 miles from Swan Hill, but still in

Victoria.

I have etc.

L Becker.

The words “Banks of the Murray” is a match with the details

in the specimen register for D4349, although it is plausible that

“Banks of the Murray” may have been like “the Junction of

the Murray and Darling Rivers”, a term used in that era to

describe various locales in northern Victoria close to the

Murray River (see Wakefield, 1966).

Becker does not appear to have illustrated these preserved

specimens from Swan Hill (or illustrated any death adder from

the Burke and Wills expedition), but he does provide another

illustration and short accompanying note of a live specimen of

death adder (unknown locality and date, but presumably from

around the same region and period):

N. Clemann, T. Stranks, R. Carland, J. Melville, B.Op Den Brouw & P. Robertson

Deaf [sic] adder, 1/5 nat. size, full grown, whole length exactly

3 feet, greatest diameter 2 l h inches, colour of upper part of

head and body: brown, with transverse bands (about 30) of a

lighter tint; under part pale yellow. Scales spined. Spine on the

end of tail sometimes half an inch long. The one I killed crossed

the road in front of my horse and moved slowly towards the

grass, and when attacked by me with a stick, rose in self

defence, resting upon the broadest part of his body. L. Becker.

Becker’s pencil, ink and watercolour illustration and hand¬

written note can be seen online at the website Caught and

Coloured: Zoological Illustrations from Colonial Victoria

(Kean et al., 2006)._Becker later transformed his illustration

into a beautiful lithograph for McCoy’s Memoirs of the

Museum ; this was never published, but the illustration

eventually appeared as Plate 12 in the Prodromus of the

Zoology of Victoria (McCoy, 1878). Becker’s information

provided much of the source information for McCoy’s account

of the species, which McCoy said was restricted to “hot tracts

near the Murray River” (p. 12).

William Lockhart Morton s death adder record (1861)

William Lockhart Morton provided an account of finding and

killing a death adder on the eastern edge of the Big Desert,

west of Pine Plains and Patchewollock, in July 1861 (the

locality description by Morton notes entering Wirringren

Plain, then turning west, crossing the plain over four miles

before he began to “mount the acclivity of the sandy desert

country” where he found the death adder “on the sloping bank

on a spot exposed to the warm morning sun”; Morton, 1966, p.

43). After killing the snake, Morton cut off the “tail-like

prolongation”, which he later examined under a microscope,

noting that “the reputed sting appears blunt, and though a dark

line runs along it, giving it seemingly the character of a tube,

no opening at that point can be detected. It is by its bite that the

death-adder proves so instantaneously destructive to animal

life” (Morton, 1966, p. 44). Morton notes that his “companion,

who had resided for eight years in South Africa, remarked that

this specimen bore a strong resemblance to the puff-adder of

that country” (p. 43).

Other claimed Victorian records of the death adder

Voucher specimens

As well as the Lake Boga adder, there are two death adder

voucher specimens in the collection of Museums Victoria that

are labelled as being from Victoria (fig. 6). One of these

specimens (D51857) was given to the Museum by Steve

Wilson but was not collected in Victoria (email from Steve

Wilson to JM, 16 May 2017).

According to the hand-written specimen register, death

adder specimen D3579, with the locality listed only as

“Victoria”, was lodged with the Museum by “Prof. Halford

23/7/1878”. George Britton Halford was a British anatomist

and physiologist who founded the first medical school in

Australia at the University of Melbourne. From 1866, Halford

conducted research in Melbourne on the effects of snake

venom, inducing snakes to bite dogs, cats and pigeons

(Hobbins, 2013). After experimenting with animals, by 1868

Halford (who believed that venom comprised living germinal

matter) began trials using injected ammonia as an antivenom

(Hobbins, 2013). This treatment remained in use for some time

(The Victorian Naturalist, Vol. IX, No. 1, 1892, p. 2, under

‘Exhibition of Specimens’ reports a death adder collected at

Monduval, NSW, “after having bitten Mr. J. M. Simson, of

Toorak, who was successfully treated with strychnine and

ammonia”). Given his research using Australian elapid snakes,

it is probable that specimen D3579 was in Halford’s possession

for this purpose. In his account of the death adder in the

Prodromus of the Zoology of Victoria, McCoy (1878) states

that: “a large dog bitten by a captive Death Adder in one of Dr.

Halford’s experiments was dead in 18 minutes”, confirming

that at least one death adder was used by Halford. It is likely

that Halford was supplied with snakes by others. Even in the

19th century, death adders had a restricted distribution in

Victoria, and it is unclear whether or not specimen D3579 was

collected in Victoria.

Further references to Victorian death adders in the literature

References to death adders in Victoria in the literature show

that death adders were an accepted component of the

Victorian fauna by the late 19th century; however, these

references do not add any more specific, substantiated

records than those discussed above. Krefft (1866) again

mentions the death adder collected during Blandowski’s

Lower Murray Expedition by stating: “of this highly

venomous snake, I obtained but a single specimen at Lake

Boga; it brings forth about 10 or 12 young” (p. 31). McCoy

(1878) states that death adders were “not found in the

southern parts of Victoria, but common in the hot tracts near

the Murray” (p. 12). A poster presenting the Dangerous

Snakes of Victoria, produced in 1877 by the Museum and the

Education Department for distribution to all Victorian

schools and railway stations, features the death adder along

with another four species of well-known venomous snakes

that remain abundant in Victoria (fig. 7). A second edition of

this poster, containing the same five species with updated

illustrations that were more life-like, was produced in the

1890s (https://museumvictoria.com.au/caughtandcoloured/

deadoralive.aspx). McCoy (1867) notes that, in Victoria the

death adder was “confined to the northern boundary” (p.

182). Le Souef (1884) lists the “Death or Deaf Adder” (p. 87)

in his Catalogue of Victorian Fauna. Soon after this, Le

Souef (1887; based on a trip undertaken in December 1886)

writes that near Lake Hindmarsh, “a farmer lately ploughed

up six death adders when ploughing up new land, but they

were not numerous, and we saw none, although a good look¬

out was kept ...” (pp. 44-45). It is possible that other snake

species that occur in this area - in particular the Bardick

Echiopsis curta - could be mistaken for death adders because

they look superficially similar when posing defensively.

However, given the relative proximity of this area to where

Morton (1966) recorded a death adder in 1861, it is not out of

the question that farmers ploughing this new land did indeed

encounter death adders.

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria

BUKBHS SUKES IF VICmiA.

INDICATED BY PROFESSOR M=COY.

TIGER SNAKE (Hoploeephalus Cnrtus)

Length, about 4 to 6 lest; Brown above banded with darker; Yellow below.

COPPER-HEADED SNAKE (Hoplocephalus Superbus).

BROWN SNAKE (Dienaenla SnpenciliOBa),

BLACK SNAKE

(Peaudechys Porphyraicns).

Length, about 6 feet; Slate color above; Pmk below.

DEATH ADDER

(Acantbophis Antarcticus).

Length, about E to 3 feet; Brown above banded with darker; paler below.

Figure 7. First edition of the Dangerous Snakes of Victoria poster, produced in 1877.

N. Clemann, T. Stranks, R. Carland, J. Melville, B.Op Den Brouw & P. Robertson

Early in the 20th century, French (1901), commenting on

the fauna of the Victorian Mallee, notes that the death adder

is one of the snakes of the region (although he did not

encounter this species during the trip that forms the basis of

his article) but “appears to be rather rare in Victoria” (p. 14).

At around the same time, Best (1901) notes that “with respect

to snakes, at our last meeting Mr. Le Souef mentioned that in

the Mallee the Death Adder was more numerous than is

generally supposed, as owing to its sluggishness it is often

passed over” (p. 93). Kershaw (1927) states that the death

adder “in Victoria, is restricted to the dry areas of the north¬

west” (p. 337). Worrell (1963) states that death adders are

“rare in Victoria” (p. 108); identical words are used by Gow

(1976). Wilson and Knowles (1992) suggest that the species

“penetrates Vic. only in far east and north-west” (p. 379), and

their accompanying map reflects that statement. Wilson and

Swan (2013) provide a distribution map but no description of

the species’ range; however, they do report the death adder’s

current status in Victoria (Data Deficient), with the

Department of Sustainability and Environment (2013) being

the source for this listing.

Aboriginal knowledge of the death adder in Victoria

Blandowski and Krefft worked with Aboriginal people during

the 1856-57 Lower Murray Expedition (engaging them as

guides, collectors and sharers of knowledge; Allen, 2009b),

and at least one of the groups they encountered along the

Murray River knew of the death adder. On his illustration of

the death adder from Lake Boga, Krefft has added a note at the

bottom about its Koorie name: Pelletoak - Yarree Yarree.

Pelletoak was the animal name provided by the Yarree Yarree

[Nyeri Nyeri] people who worked with the Blandowski Lower

Murray Expedition at Mondellimin from April to December

1857. Blandowski and Krefft were in the practice of showing

various animal illustrations to local Aboriginal people along

the way, trying to gather native names for particular species

(Stranks, in prep.). Pelletoak appears to be a specific name for

death adder (as opposed to a more generic word for snake such

as Cournvil or Cumvill ) recorded from the Yarree Yarree. This

indicates that the species was known in the Mondellimin area

at the time the expedition visited, although no specimens were

collected there.

Krefft gave a talk on the “vertebrated animals” (he

discussed mammals only) from the “Lower Murray Expedition”

to the monthly meeting of the Philosophical Society of NSW

on 10 September 1862 (reported in the Sydney Morning Herald,

24 October 1862, page 2). This was followed by a continuation

of the former paper - a talk on the reptiles, amphibians and

fishes from the Lower Murray Expedition to the monthly

meeting of the Philosophical Society of NSW on 16 September

1863 (meeting report in the Sydney Morning Herald, 21

September 1863, page 13). In this second talk, the newspaper

reports Krefft stating: “The Death Adder (Acanthophis

antarctica) [sic], not very common, as I have never seen but one

single individual at Lake Boga; there is no difference in the

coloration of this snake from those inhabiting the east coast;

the natives [at Mondellimin] never brought one though high

rewards had been offered” (p. 13). It is notable that the

Aboriginal people did not collect a specimen of the death adder

during the eight months that the expedition spent at

Mondellimin. Perhaps the Aboriginal people could not find the

species, or would not collect it because it was venomous and

dangerous, or there was a cultural consideration such as a taboo

that prevented them from harming or killing it.

It does not appear that Blandowski and Krefft worked with

any Aboriginal people during their relatively brief stay Lake

Boga in March 1857, so there is no record of a Koorie name for

the species there. Curiously, however, there is another reference

to the death adder at Lake Boga. Mr A. Chas. Stone worked as

the baker at Lake Boga for more than 18 years, and was in close

contact with the “Lake Boga tribe”, known as Gourrmjanyuk

(meaning “along the edge of trees”; Stone, 1911, p. 433), from

about 1890 to 1910. He published a paper on The Aborigines of

Lake Boga (Stone, 1911) that has a list of Koorie names for

animals, including ten varieties of snake, with the name “Llerk”

for “Deaf Adder” (p. 446, noting the frequently used - and

technically erroneous - variation on the common name). This

paper was a primary source for the Wemba Wemba Dictionary

compiled by linguist Luise Hercus in the 1960s (Hercus, 1992);

she has listed an alternative spelling of “Lirrk” for death adder

(p. 106).

Claimed sightings of death adders in Victoria in recent

decades

Most claims of sightings of death adders in Victoria in the late

20th and early 21st centuries have been from either the far

north-west or from the eastern tip of the state. After two

credible reports from beside the Murray River in the far north¬

west, in 2005 two of us (PR and NC) led a brief survey in the

area. Instead of death adders, that survey resulted in the capture

of a snake previously unknown from Victoria: De Vis’ Banded

Snake Denisonia devisi (Clemann et al., 2007). It appears that

those most recent claims of death adders in the riverine area

surveyed by Clemann et al. (2007) were actually mis-identified

De Vis’ Banded Snakes.

In far south-eastern New South Wales there are death adder

records east of the Princes Highway in Nadgee State Forest,

less than one kilometre from the border with Victoria (Atlas of

NSW Wildlife, accessed online, 28 May 2017; Swan et ah,

2004; fig. 1). Most (but not all) claims of death adders in far

East Gippsland are from east of Mallacoota Inlet. One of us

(BodB) spent much of her life in Mallacoota, observing snakes

in that area. BodB’s father - an experienced naturalist - has

seen what he believes to be death adders in that area on two

occasions; both observations, and the majority of others she has

heard of, including some recent (2016) reports, were east of the

Wallagaraugh River. The few potential sightings she is aware

of that occurred west of the Wallagaraugh all occurred in the

1980s. No substantiating material such as photographs or a

voucher specimen have been produced from eastern Victoria.

Death adders possess traits that predispose them to

population losses (Reed and Shine, 2002). Because Victoria is

the southern extreme of this species’ range, it is here that

declines are particularly likely. Land use changes and the

The death adder Acanthophis antarcticus (Shaw & Nodder, 1802) in Victoria

introduction of stock animals and exotic predators associated

with the push into northern Victoria by European people had a

profound impact on native fauna, resulting in the extinction of

some mammals (Menkhorst, 2009). These impacts were

already evident to Krefft in the 1850s, and it is sobering to

acknowledge that the ecocidal impacts of European incursion

proclaimed by Krefft 160 years ago (Kean, 2009; Menkhorst,

2009) continue to this day. As well as the losses of mammals,

it is probable that around this time there were deleterious

impacts on various reptile species. Frequent burning of native

vegetation on public land in the Victorian Mallee has resulted

in changes to the fauna (Robertson et al., 2012); both the Mallee

and far East Gippsland are currently subject to frequent fuel

reduction burning, and death adders may be particularly

susceptible to fire (McDonald et al., 2012; Smith et al., 2012).

The Mallee region is popular with natural historians,

particularly ornithologists and herpetologists, and the fact that,

despite the frequent activities of these people, no reliable death

adder records have been produced in north-western Victoria

since the mid-19th century suggests that this species no longer

occurs in that part of the state. However, due to their lower

energetic needs, some reptile species may persist in small

isolated areas. The part of Victoria east of the Wallagaraugh

River is comparatively under-surveyed, and consequently, it is

not entirely out of the question that death adders persist in the

far east of the state.

Acknowledgements

Laura Cook (Museums Victoria) facilitated access to the

specimens at the Melbourne Museum. Steve Wilson

(Queensland Museum) provided helpful information on the

death adder specimen he donated to the Museum. Peter

Menkhorst (Arthur Rylah Institute for Environmental

Research [ARI]) provided helpful additional information and

perspectives on Blandowski’s Lower Murray Expedition, and

provided a detailed and insightful review of a draft of this

paper. Lindy Lumsden (ARI) also provided helpful comments

on a draft of this paper. Harry Allen (University of Auckland)

kindly provided access to his transcription of Krefft’s

expedition narrative held at the Mitchell Library, State Library

of New South Wales. Anita Hermannstadter, Sabine Hackethal

and Sandra Miehlbradt (Museum fur Naturkunde, Berlin)

kindly hosted visits by Rebecca Carland in July 2016 and

Timothy Stranks in May 2017, providing access to collections

and permission to reproduce Krefft’s illustration of the death

adder. Michael Scroggie (ARI) prepared the map, and Matt

White and Steve Sinclair (both ARI) provided helpful

guidance on the geography around Morton’s death adder

record. We thank Joanna Sumner for an excellent review of an

earlier version of this paper and for photographing some of the

specimens, and we thank Richard Marchant for guiding

improvements to this paper.

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Memoirs of Museum Victoria 77:41-61 (2018) Published 2018

1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.Org/10.24199/j.mmv.2018.77.04

Diversity in Australia’s tropical savannas: An integrative taxonomic revision of

agamid lizards from the genera Amphibolurus and Lophognathus (Lacertilia:

Agamidae)

(http://zoobank.org/urn:lsid:zoobank.org:pub:22334107-0784-466E-8288-D6E29F87F6E2)

Jane Melville 1 *, Euan G. Ritchie 12 , Stephanie N. J. Chapple 1 , Richard E. Glor 3 and James A. Schulte II 4

1 Department of Sciences, Museum Victoria, GPO Box 666, Melbourne VIC 3001, Australia

2 School of Life and Environmental Sciences, Deakin University, Burwood VIC 3125, Australia

3 Herpetology Division, Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas,

Lawrence, KS 66045, USA

4 Beloit College, 700 College Street, Science Center 338, Beloit, WI 53511, USA

* Corresponding author. Email: jmelv@museum.vic.gov.au

Abstract Melville, J., Ritchie, E.G., Chappie, S.N.J., Glor, R.E.and Schulte II, J.A. 2018. Diversity in Australia’s tropical savannas:

An integrative taxonomic revision of agamid lizards from the genera Amphibolurus and Lophognathus (Lacertilia:

Agamidae). Memoirs of Museum Victoria 77: 41-61.

The taxonomy of many of Australia’s agamid lizard genera remains unresolved because morphological characters

have proved to be unreliable across numerous lineages. We undertook a morphological study and integrated this with a

recent genetic study to resolve long-standing taxonomic problems in three genera of large-bodied Australian agamid

lizards: Amphibolurus, Gowidon and Lophognathus. We had broad geographic sampling across genera, including all

currently recognised species and subspecies. Using an integrative taxonomic approach, incorporating mitochondrial

(. ND2 ) and nuclear ( RAG1) genetic data, and our morphological review, we found that both generic and species-level

taxonomic revisions were required. We revise generic designations, creating one new genus ( Tropicagama gen. nov.) and

confirming the validity of Gowidon , giving a total of four genera. In addition, we describe a new species ( Lophognathus

horneri sp. nov.) and reclassify two other species. Our results provide a significant step forward in the taxonomy of some

of Australia’s most iconic and well-known lizards and provide a clearer understanding of biogeographic patterns across

Australia’s monsoonal and arid landscapes.

Keywords Agamid lizards, Amphibolurus horneri sp. nov., Lophognathus, Gowidon, Tropicagama gen. nov., integrative taxonomy,

Australia, monsoon tropics

Introduction

Tropical savannas constitute one of Earth’s major biomes,

covering 20-30% of the land surface (Myers et al., 2000).

Australian tropical savannas are particularly important

because they are the largest and least modified tropical

savanna woodlands in the world, comprising approximately

25% of the Earth’s remaining savannas that are in good

ecological condition (Woinarski et al., 2007). The Australian

monsoonal tropics, which span the northern third of the

continent, are home to a major component of biodiversity, with

some areas, particularly the sandstone escarpments, having

similar biodiversity levels to Australian rainforests (Bowman

et al., 2010). Yet, only recently has research started to uncover

unexpected levels of diversity and phylogeographic structure

across the monsoonal tropics (Melville et al., 2011; Moritz et

al., 2016; Oliver et al., 2014; Potter et al., 2016; Smith et al.,

2011). As a result of these research findings, the current

taxonomy in many groups does not reflect actual species

diversity.

One lizard group that is in immediate need of a taxonomic

revision is the large-bodied agamids of the tropical savannah

woodlands. Despite their ubiquity and ecological significance

in this biome, major taxonomic problems characterise the

group at both the generic and the species levels. Molecular

work suggests major taxonomic problems within Amphibourus,

Gowidon and Lophognathus (Hugall et al., 2008; Melville et

al., 2011; Schulte et al., 2003). Melville et al. (2011) identified a

clade containing five species in three genera: Amphibolurus

muricatus (White, 1790), A. norrisi Witten and Coventry,

1984, A. burnsi (Wells and Wellington, 1985), L. gilberti Gray,

1842, and Chlamydosaurus kingi Gray, 1825. According to

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

Table 1. Current taxonomic designations of Agamidae genera under revision, including details of synonyms and primary types, from Department

of the Environment and Energy (2014). All types have been examined for morphological analysis, except those where they are presumed lost or

status unknown.

Species plus junior synonyms

Described

Details of types

1 .

Amphibolurus muricatus

[White (1790)]

BMNH 1946.9.4.44

Lacerta muricata

White (1790)

BMNH 1946.9.4.44

Agama jaksoniensis

Cloquet (1816)

RMNH 3117

Amphibolurus norrisi

Witten and Coventry (1984)

NMV D51499

Chlamydosaurus kingii

Gray (1825)

Type presumed lost.

Amphibolurus burnsi

[Wells and Wellington (1985)]

AMR 116981

Lophognathus gilberti

Gray (1842)

BMNH 1946.8.28.69

Redtenbacheria fas data

Steindachner (1867)

Type not found.

Physignathus incognitus

Ahl (1926)

ZMB 30086

Physignathus gilberti centralis

Loveridge (1933)

MCZ 35207

Gowidon longirostris

[Boulenger (1883)]

BMNH 1946.8.12.64-65, 1946.8.28.73

Physignathus eraduensis

Werner (1909)

Status unknown

Physignathus longirostris quattuorfasciatus

Sternfeld (1924)

SMF 10366

Gowidon temporalis

[Gunther (1867)]

BMNH 1946.8.12.73/63, 1946.8.28.72

Lophognathus lateralis

Macleay (1877)

AM R31882

Lophognathus labialis

Boulenger (1883)

BMNH 1946.8.12.72, 1946.8.12.63

Note: BMNH, British Museum of Natural History, London; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden, Holland; AM,

Australian Museum, Sydney; NMV, Museum Victoria, Melbourne; ZMB, Zoologisches Museum, Universitat Humboldt, Berlin,

Germany; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, United States; SMF, Senckenberg Naturmuseum,

Frankfurt-am-Main, Germany.

these results, the frill-necked lizard ( Chlamydosaurus kingii ),

which has remained monotypic since its description, belongs to

this clade, despite its stunning morphological distinctness.

Further complicating these results is the possibility that L.

gilberti is actually a species complex. Molecular work (Melville

et al., 2011; Schulte et ah, 2003) and field observations

(Melville, unpublished data) led to the hypothesis that a number

of populations that have been relegated to L. gilberti’s

synonomy (Table 1, e.g. Physignathus gilberti centralis

Loveridge 1933) may represent a valid species.

There has been a complex taxonomic history for the genus

Lophognathus, which was originally erected for a single

species ( Lophognathus gilberti ), with the inclusion of multiple

species at different times, including Amphibolurus burnsi

(Wells and Wellington 1985), Gowidon longirostris

(Boulenger, 1883) and G. temporalis (Gunther, 1867). The

current catalogue (Department of the Environment and

Energy, 2014) lists one species in Lophognathus (L. gilberti )

and two in Gowidon (G. temporalis and G. longirostris ),

although a formal revision of the generic placement of G.

temporalis into Gowidon has not occurred. Molecular work

suggests that G. temporalis and G. longirostris are not closely

related to L. gilberti, and that A. burnsi is more closely related

to Amphibolurus, with more recent publications reflecting this

placement (e.g. Wilson and Swan, 2017). Based on this

confusing taxonomic history and that the morphological

characters previously used to define Gowidon, Lophognathus

and Amphibolurus do not seem to be diagnostic, a complete

review of these species is required.

We undertook a morphological study and an integrative

taxonomic review of the genera Gowidon, Amphibolurus and

Lophognathus species from northern Australia, incorporating

results from the most recent molecular study (Melville et al.,

2011). We examined all primary types for the study species,

including junior synonyms, and conducted detailed

morphological analyses using additional museum specimens

that included many of the specimens sequenced in the

molecular study (Melville et al., 2011). This comprehensive

review of these genera provides a complete taxonomic revision

and contributes significantly to our understanding of generic

and species-level diversity in the Australian tropical savannas.

Taxonomy in Australia’s tropical savanna lizards

Figure 1. MtDNA phylogenetic tree for the genera Lophognathus, Amphibolurus and Chlamydosaurus reproduced from Melville et al. (2011).

Tree presented is a Bayesian 50% majority-rule consensus tree based on ~1200 bp mitochondrial DNA ( ND2 ). Bayesian posterior probabilities

and ML boostraps are provided on branches. Sample identification numbers are either Genbank accession numbers for previously published

sequences or museum IDs (shown in brackets) for samples sequenced previously. Vertical bars indicate species following the taxonomic revision.

Materials and methods

Morphology

Primary types, including all junior synonyms (Table 1), were

examined for morphological analysis and taxonomic revision.

We also examined additional museum specimens for a

morphological analysis of all species currently belonging to

Lophognathus (see Supplementary Appendix SI for details).

Ten morphometric characters were measured using calipers for

all specimens examined: snout-vent length (mm), tail length

(mm), upper hindlimb length (proximal hindlimb; mm), lower

hindlimb length (distal hindlimb; mm), hindfoot length to end

of fourth toe (autopod hindlimb; mm), head length (mm), head

width at widest point behind ear (mm), head depth at deepest

point between eyes and ears (mm), number of femoral pores

and number of preanal pores. In addition to these morphometric

measurements, specimens were examined for scalation

patterns, colour patterns and other synapomorphies.

Univariate and multivariate analyses were used to examine

differences in the morphometric characters between the

species. We used SYSTAT Version 10.2 (SYSTAT Software

Inc., Richmond, California, USA) for analyses. Before analysis,

all morphological variables were regressed against snout-vent

length and the residuals were used for subsequent analyses to

remove the effect of body size. First, all ten morphometric

characters were analysed using analysis of variance with

multiple comparisons (Tukey’s procedure). Then, principal

components analysis was used to reduce the dimensionality of

the morphological data (FACTOR procedure of SYSTAT).

Principal components were extracted from a correlation matrix

of the raw data. Principal components were named by the

correlations of the original variables to the principal component;

correlations with absolute values greater than 0.5 were

considered important. Resultant principal components were

explored using analysis of variance with multiple comparisons

(Tukey’s procedure) to determine whether there were

interspecific differences in morphometric characters.

Species delimitation assessment

We used an integrative taxonomic approach for species

delimitation assessment by following the principle that as

many lines of evidence as available should be combined to

delimit species (Miralles and Vences, 2013), which has been

successfully used in Australian dragon lizards (Melville et al.,

2014). We first used the mtDNA phylogenetic tree in association

with the nuclear tree to determine that no mtDNA introgression

exists. Thus, the mtDNA was used to define a starting

hypothesis for the clustering of specimens (Miralles and

Vences, 2013). Species delimitation was then based on

additional support from at least two of the following: (a)

sympatric occurrence without admixture, as revealed by

consistent differences in morphological or molecular

characters at the same geographic location; (b) congruent

diagnostic differences between sister lineages in morphological

characters; and (c) the absence of haplotype sharing in nuclear

loci. Integrative taxonomy therefore minimises the alpha error

by only taking into account the most unambiguous species

evidence provided by a variety of approaches and attempts to

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

keep the beta error low by seeking evidence from as many

different approaches as possible (Miralles and Vences, 2013).

Results

Taxonomic implications of phylogenetic relationships

Results from the comprehensive molecular study (Melville et

al., 2011) provide strong evidence that a taxonomic revision of

Lophognathus, Gowdon and Amphibolurus is warranted

(fig. 1). Within the clade containing Amphibolurus,

Chlamydosaurus and Lophognathus (fig. 1), Chlamydosaurus

forms a monophyletic group that is sister to the other genera.

Based on these results, Chlamydosaurus is a well-supported

genus. However, revision of Amphibolurus and Lophognathus

is required. Molecular results show that Amphibolurus, for

which A. muricatus is the type specimen, consists of L. gilberti

centralis, A. burnsi, A. muricatus and A. norrisi. The genus

Lophognathus contains only L. gilberti, but there is strong

molecular evidence that L. gilberti is two species, a northern

taxon and a more southern taxon (fig. 1). These two L. gilberti

lineages are analysed further in the morphological section

below, with the southern lineage referred to as Lophognathus

sp. nov. Additionally, Gowidon temporalis and G. longirostris,

which have previously been placed in Lophognathus, form

two independent lineages that fall outside the Amphibolurus,

Chlamydosaurus and Lophognathus clade. Based on these

phylogenetic results, G. temporalis and G. longirostris should

be placed in two separate genera. The name Gowidon (Wells

and Wellington, 1984) has been used in recent publications

(e.g. Wilson and Swan, 2017) based on molecular data

(Melville et al., 2011). However, there is no available name for

G. temporalis and a new genus is required (see taxonomic

revision below). These generic designations are analysed

further in the morphological section below.

Morphological analysis

Museum specimens for species in Lophognathus, Gowidon and

Amphibolurus were analysed using ten morphometric characters

and examined to identify morphological synapomorphies.

Initially, the specimens that were sequenced in the molecular

study (Melville et al., 2011) were examined to determine

diagnostic characters for the six species identified in the

phylogenetic analyses: L. gilberti, L. sp. nov., L. gilberti

centralis, G. longirostris, G. temporalis and A. burnsi. Once the

diagnostic characters had been established, we then went through

museum specimens (Supplementary Appendix SI) and primary

types (Table 1) and assigned each of them to one of these species.

All specimens were then measured and scored for morphometric

and synapomorphic characters. A summary of morphometric

characters for each species is presented in Table 2. While there is

extensive variation within species in body colour, patterns and

scalation, we were able to determine some consistent and

diagnostic characters that differed between the taxa. These

synapomorphies were particularly related to scalation and colour

patterning on the head and upper body. These synapomorphies

are covered in detail in the taxonomic revision section below.

Morphometric analyses - multivariate methods were used

to examine the morphological differences between the six

Taxonomy in Australia’s tropical savanna lizards

Table 2. Mean morphological measurements (mm ± standard error)

for study species

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Lophognathus species. We conducted a principal components

analysis on the measured morphological characters (Table 3).

Three morphological measurements were found not to vary

significantly between species: proximal hindlimb; distal

hindlimb and head length. Consequently, these characters

were not included in the multivariate analyses.

Principal components analysis of the morphometric

variables revealed that body proportions accounted for almost

half of the variance in the data. This first principal component

(PCI) explained 46.43% of the variance in morphology.

Lizards scoring high on PCI are large with relatively long

hind feet, long tails, and deep or wide heads, while lizards

scoring low on PCI are small with short body proportions

(Table 3). The second principal component (PC2) explained

19.69% of the variance in morphology. Lizards with high

scores on PC2 had lower numbers of femoral and preanal

pores, while lizards scoring low on PC2 had higher numbers

of pores (Table 3). The third principal component (PC3)

explained 14.40% of the variance in morphology. Lizards with

high scores on PC3 had proportionally short hind feet, while

lizards scoring low on PC3 had longer hind feet. Analysis of

variance indicated a statistically significant difference between

species on PCI (F s 136 = 12.61, P < 0.001), and Tukey’s honest

significant difference post hoc test showed that L. gilberti and

L. sp. nov. scored significantly lower on this principal

component than L. gilberti centralis ( P = 0.029 and P - 0.003,

respectively), A. burnsi (both P < 0.001) and G. longirostris (P

= 0.039 and P - 0.010, respectively) but not G. temporalis. In

addition, G. temporalis had a significantly lower score than G.

longirostris (P < 0.001) on PCI. Thus, relative to body size, L.

gilberti, L. sp. nov. and G. temporalis have shorter or smaller

body proportions than the other species.

Analysis of variance of PC2 also indicated a statistically

significant difference between species (F 5 = 56.49, P <

0.001). Tukey’s honest significant difference post hoc test

indicated that G. temporalis scored significantly higher on PC2

than all of the other taxa (all P < 0.001), indicating lower pore

numbers, and G. longirostris scored significantly lower than all

other taxa (all P < 0.001), indicating higher pore numbers.

Analysis of variance of PC3 indicated a statistically significant

difference between species (F 5 136 = 21.08, P < 0.001) and

Tukey’s honest significant difference post hoc test indicated

that on PC3, G. temporalis and G. longirostris scored

significantly higher than all of the other taxa (all P < 0.001),

indicating shorter hind feet length, relative to body size.

Although there is a statistically significant separation of

these species in morphospace, it is visually difficult to

ascertain (fig. 2). There is extensive overlap in morphospace,

with G. temporalis and G. longirostris being the only species

with visible separation from the others and that do not overlap

each other in morphospace.

Species delimitation assessment

All available specimens were separated into groups based on

the independent lineages supported in the mtDNA tree and

then assessed for fixed and unambiguous morphological

character states. We identified five morphologically

diagnosable lineages that were also resolved as independent

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

Table 3. Principal components analysis of lizard morphology. Correlations with absolute values greater than 0.5 are in bold and are considered

important.

PCI

PC 2

PC 3

% Variance explained

46.43

19.69

14.40

Eigenvalues

3.25

1.38

1.01

Component loadings

SVL

0.81

0.37

-0.05

Tail

0.78

0.13

0.32

Hind foot

0.55

0.18

-0.77

Head width

0.80

0.10

0.27

Head depth

0.83

0.08

0.02

Femoral pores

0.40

-0.75

-0.42

Preanal pores

0.45

-0.79

0.30

Species

Amphibolurus burnsi

0.51 ±0.34

0.65 + 0.22

-0.71 + 0.15

Lophognathus gilberti centralis

0.34 + 0.24

0.44 + 0.14

-1.09 + 0.20

Lophognathus gilberti gilberti

-0.51+0.16

0.10 + 0.13

-0.29 + 0.20

Lophognathus sp. nov.

-0.54 + 0.17

-0.12 + 0.10

-0.24 + 0.14

Gowidon longirostris

0.85 + 0.10

-1.32 + 0.10

0.65 + 0.12

Gowidon temporalis

-0.28 + 0.13

1.13+0.13

0.83 + 0.14

Note: PC, principal component; SVL, snout-vent length.

lineages in the mtDNA and nuclear trees, which were equated

to generic level divisions (fig. 3). Within lineages, further

morphologically diagnosable lineages, which were also well-

supported in the mtDNA and nuclear trees, could be identified.

A number of these lineages have wholly or partially overlapping

geographic distributions within the Australian tropical

savannah (fig. 4): Lophognathus gilberti gilberti, L. gilberti

centralis, L. gilberti sp. nov. and Chlamydosaurus kingi.

Using this species delimitation method, five genera and nine

species could be delineated across Amphibolurus, Gowidon

and Lophognathus sensu lato.

Taxonomic revision

Diagnoses below are given only in terms of synapomorphies.

Species diagnoses are only provided for those species for

which taxonomic revision is required.

Genus Amphibolurus Wagler, 1830

Gemmatophora Kaup, J.J., 1827. Zoologische Monographien. Isis

Von Oken, Jena 20: 610-625 [621] [nom. oblite, described as subgenus

of Calotes Cuvier, 1817]. Type-species Lacerta muricata White,

1790 by original designation.

Amphibolurus Wagler, J.G., 1830. Natiirliches System der

Amphibien, mit vorangehender Classification der Sdugethiere und

Vogel. Miinchen, Cotta’schen vi 354 pp. [145] [replacement for

Gemmatophora Kaup, 1827, which Wagler rejected as an invalid

hybrid name (“vox hybrida”)].

Grammatophora Dumeril, A.M.C. and Bibron, G., 1837.

Erpetologie Generate ou Histoire Naturelle Complete des Reptiles.

Paris, Roret 4: ii 571 pp. [468] [non Grammatophora Stephens, 1829

{nom. nud)\ emendation of Gemmatophora Kaup, 1827].

Petroplanis Fitzinger, L.J., 1843. Systema Reptilium. Vienna,

Braiimuller u. Seidel vi 106 pp. [83, 84] [nom. nude, introduced in

synonymy of Amphibolurus Wagler, 1830]. Types species Petroplanis

jacksoniensis Fitzinger, 1843 (=? Agama jacksoniensis Cloquet, 1816)

by monotypy.

Polylophus Fitzinger, L.J., 1843. Systema Reptilium. Vienna,

Braiimuller u. Seidel vi 106 pp. [83, 84] [nom. nude, introduced in

synonymy of Amphibolurus Wagler, 1830]. Types species Polylophus

jacksoniensis Fitzinger, 1843 (=? Agama jacksoniensis Cloquet, 1816)

by monotypy.

Synonymy that of: Cogger, H.G. 1983, in Cogger, H.G., Cameron,

E.E., and Cogger, H.M. Amphibia and reptiles. Pp. 108-116 in:

Walton, D.W. (ed.) Zoological catalogue of Australia. Vol. 1. Netley,

South Australia: Griffin Press Ltd. vi 313 pp. [108]

Diagnosis. A genus consisting of large agamid lizards in the

subfamily Amphibolurinae with exposed tympanum, long

robust limbs and a long tail. Gular scales smooth to weakly

keeled, ventral scales smooth to strongly keeled. Head wide

and deep in comparison with length. Heterogenous body scales,

dorsal surface scattered with many spinose scales. Well-

developed spinose nuchal and vertebral crest. Two broad pale

dorsolateral stripes running from ear or neck to the hip,

Taxonomy in Australia’s tropical savanna lizards

pci

PC2

" L. temporalis

* L longirostris

L. sp. nov.

- L. gilbert!

- L bumsi

- L. g. centralis

Figure 2. The distribution of Lophognathus sensu lato samples included

in this study along the first three morphological principal components

axes. Distribution of each taxon is delineated by a 95% confidence ellipse.

Lophognathus gilberti has been separated into L. sp. nov., L. gilberti

centralis , and L. gilberti.

discontinuous with pale lip scales. Dorsolateral stripes

intersected by multiple wedges of brown or grey along their

length.

Included species. Amphibolurus norrisi Witten, G.J. and

Coventry, A.J., 1984; Lacerta muricata White, J., 1790;

Amphibolurus bumsi Wells, R.W. and Wellington, C.R.,

1985; Physignathus gilberti centralis Loveridge, A., 1933.

Distribution. Continental Australia, including south-eastern

Australia extending west across Nullabor Plain, eastern

Australia extending north into Queensland, central Australia

incorporating Northern Territory, Western Australia and

western Queensland. A broad range of habitats occupied,

including arid and semi-arid woodlands, dry sclerophyll forests

and woodlands, and coastal heathlands.

Diagnosis. Large robust member of the Amphibolurus genus.

Large wide head with extensive covering of spinose scales.

Posterior ventral portion of head heavily covered with spinose

scales. Well-developed spinose nuchal and vertebral crest,

which continues down back to hips. At least two more spinose

dorsal crests on each side of vertebral crest. Scalation on back

strongly heterogeneous, with two dorsolateral rows of spinose

scales running from shoulders to hips. Scales on thighs strongly

heterogeneous with scattered spinose scales. Prominent row of

spinose scales running along the posterior edge of thighs.

Shades of brown, grey to almost black. Two broad pale

dorsolateral stripes running from ear or neck to the hip,

discontinuous with lip scales. Dorsolateral stripes intersected

by multiple wedges of brown or grey along their length.

Femoral pores 3-5; preanal pores 4-6.

Description ofholotype. Adult. Large robust lizard with distinct

neck, limbs long and robust; canthus well defined; nasal scale

below canthal ridge, nare slightly to the posterior-dorsal

section of the nasal scale; visible tympanum. Infralabials 12;

supralabials 13. Labials elongate somewhat keeled. Scales on

dorsal surface of head heterogenous and strongly keeled. Well-

developed spinose nuchal crest. Posterior portion of head

heavily covered with spinose scales. Well-developed vertebral

crest, which continues down back to hips. Two paravertebral

rows of enlarged and prominent spinose scales on each side of

vertebral crest, running from shoulders to hips. Scales on

thighs strongly heterogeneous with scattered spinose scales.

Row of enlarged spinose scales running along posterior edge of

thighs. Scales on the dorsal surface of body and tail are strongly

keeled and scales on the ventral surface are weakly keeled.

Colour dorsally is light to dark brown and grey, with scattered

black markings.

Variation. Considerable variation in the number and size of

spinose scales between males, females and juveniles. In adult

males there are numerous long spines (> 2 mm) and the spinose

scales are dense across the back of the head, nuchal and ventral

crests, and rear of the thighs. In females and juveniles, spinose

scales are still present and diagnostic but they are smaller and

less dense, providing an overall appearance of the lizards being

less spiny. Some individuals, particularly adult males, have a

broad pale stripe running along the full extent of the lower lip.

However, a white stripe along the upper lip is not present and a

well-defined pale stripe between the eyes and ears is not present.

Distribution and ecology. Occurs in dry woodlands and

associated with eucalypts along inland watercourses.

Distributed across southern and central-western Queensland

and northern inland New South Wales.

Amphibolurus bumsi

(fig- 8)

Amphibolurus bumsi Wells, R.W. and Wellington, C.R., 1985. A

classification of the Amphibia and Reptilia of Australia. Australian

Journal of Herpetology Supplementary Series 1: 1-61 [18].

Designation that of Melville, J., this work.

Holotype. AM R116981 (previously AMF 28917), Collarenebri,

New South Wales.

Remarks. The distribution of Amphibolurus bumsi potentially

overlaps with A. centralis and A. muricatus. A. bumsi has been

included in the genus Lophognathus but DNA sequencing has

confirmed that it is unrelated to Lophognathus species and

demonstrates a clear sister-species relationship with A.

centralis. Morphologically, A. centralis and A. bumsi can be

distinguished by the latter having heterogeneous scales on the

thighs, spinous scales on the thigh and enlarged spinous scales

along the rear of the thigh.

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

Sister mtDNA lineages - not necessarily sympatric - with absence of gene flow for nuclear gene

i | Sister mtDNA lineages - not necessarily sympatric - with correlated evidence from at least two

independent lines of evidence (e.g-, morphological, nuclear mtDNA)

Sister mtDNA lineages with geographically overlapping distributions - with correlated evidence from at

least two independent lines of evidence (e.g., morphological, nuclear. mtDNA)

Figure 3. Results from the integrative taxonomic approach to species delimitation. Genera are designated by a multicoloured horizontal bars at

the top of the figure, and species within genera are designated by a number in black above a coloured vertical segment. Segment colours and

species numbers correspond to those in fig.l.

Amphibolurus centralis

(Figs. 5 & 8)

Physignathus gilberti centralis Loveridge, A. 1933. New

agamid lizards of the genera Amphibolurus and Physignathus from

Australia. Proceedings of the New England Zoological Club, Boston

13: 69-72 [71]. Designation that of Wells, R.W. and Wellington, C.R.,

1983. A synopsis of the Class Reptilia in Australia. Australian Journal

of Herpetology 1: 73-129 [80],

Holotype. MCZ 35207, Anningie 30 mi[les] W of Teatree Well,

Northern Territory.

Paratype. AM R10993 (formerly MCZ 35208), Australia,

Northern Territory, Tea Tree Well, (22° 8' S, 133° 24' E). Juvenile.

Note: Original MCZ catalogue lists locality information identical to

MCZ 35207.

Diagnosis. Large robust member of the Amphibolurus genus.

Large wide head in proportion to body size. Well-developed

spinose nuchal and vertebral crest. Scalation on back

heterogeneous. Scales on thighs relatively homogeneous,

lacking row of large spinose scales. Shades of light to dark

brown and grey. Two broad pale dorsolateral stripes running

from ear or neck to hip, discontinuous with pale lip scales.

Dorsolateral stripes intersected by multiple wedges of brown or

grey along their length. Most individuals have a broad pale or

white stripe running along extent of the lower lip. Femoral

pores 2-6; preanal pores 3-6.

Description of holotype. Adult male. Large robust lizard with

large wide head in proportion to body size and well-developed

spinose nuchal crest comprising a row of eight enlarged scales.

Taxonomy in Australia’s tropical savanna lizards

a. L gilberti

b. L horneri sp. nov.

d. A. burnsi

Figure 4. Distribution of study species included in this study, compiled from museum records.

Figure 5. Primary type specimens: a ,Amphibolurus centralis (MCZ 35207);

b, Gowidon longirostris (BMNH 1946.8.28.73); c, Lophognathus gilbert

(BMNH1946.8.28.69); d, Tropicagama temporalis (BMNH 1946.8.28.72).

Additional (4-6) enlarged spinose scale protruding from rear

of head, posterior to the jaw. Scales on thighs relatively

homogeneous, lacking row of large spinose scales. Distinct

neck, limbs long and robust; canthus well defined; nasal scale

below canthal ridge, nare slightly to the posterior-dorsal

section of the nasal scale; visible tympanum. Infralabials 13,

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

I**;' jiijF

ik. 7' ~ risg

(e) L. sp.nov. NMV D73663

fb) L giibertimv D74260

<c) L . gilberticentmUs N MV D72661 (d) L gift&rii NMV D 74 281

(e) L. gilberticentralis NMV D74288

Figure 6. Colour pattern variation in the lateral head views of

Amphobolurus centralis, Lophognathus gilbert and L. horneri sp. nov.

Museum registration numbers for the individual lizards photographed

are provided.

supralabials 14. Labials elongate without obvious keels. Scales

on dorsal surface of head heterogenous and strongly keeled.

Scattered enlarged, keeled mucronate scales present on side of

head posterior to the eye. Scales on the dorsal surface of body

and tail are strongly keeled and scales on the ventral surface

are weakly keeled. Tail long, robust at base, tapering distinctly

from approximately one-third along its length to a fine tip.

Shades of cream, light to dark brown and grey. Two broad pale

dorsolateral stripes running from rear of head to back of rear

legs; pale stripes bordered by narrow discontinuous dark brown

stripes along entire length and flecks of dark brown within the

posterior two-thirds of the pale stripes. Irregular dark brown

colouration between the pale dorsolateral stripes on the anterior

one-third of the torso. Lacking pale stripe running along the

extent of the lower lip.

Variation. Marked variation in colour pattern between

individuals. Broad white lip stripe occurs along the upper lip in

some individuals, mostly adult males. Alternatively, some

individuals do not have a white stripe on either the upper or the

lower lips. Diffuse pale stripe between eye and ear in some

individuals, but it is not a well-defined stripe bordered dorsally

and ventrally by a row of darker scales extending the full span

of eye-ear.

Distribution and ecology. Arid northern-central and central

Australia, particularly associated with mulga woodlands but

also occurring in eucalypt woodlands. Western Queensland,

Northern Territory and Western Australia.

Taxonomy in Australia’s tropical savanna lizards

Table 4. Summary of morphometric measurements (mm) for the primary types of Amphibolurus burnsi, A. centralis, Gowidon longirostris,

Lophognathus gilbert, L. horneri sp. nov. and Tropicagama temporalis

Amphibolurus

burnsi

Amphibolurus

centralis

Gowidon

longirostris

Lophognathus

gilberti

Lophognathus

horneri sp. nov.

Tropicagama

temporalis

AM R116981

Holotype

MCZ 35207

Holotype

BMNH

1946.8.28.73

Lectotype

BMNH

1946.8.28.69

Holotype

NTM R16472

Holotype

BMNH

1946.8.28.72

Lectotype

SVL

105.2

99.6

88.8

112.9

102

102.7

39.4

38.3

29.1

40.2

36.1

31.3

31.0

29.6

18.9

27.5

23.2

19.5

13.3

12.2

12.8

14.5

14.4

13.2

LegL

88.6

85.2

83.9

105.6

102.0

87.1

Note: SVL, snout-vent length; HL, head length; HW, head width at widest point; HD, head depth; LegL, hindlimb length.

nose tail nose tail

nov (NMVD73663) (b) Lophognathus sp nov. {NMVD7381i8)

nose tail nose tail

Figure 7. Variation in white pigmentation on tympanums of

Lophognathus sp. nov., L. gilberti andL. gilberti centralis. Orientation

of the tympanums is provided under images. Museum registration

numbers for the individual lizards photographed are also provided.

Remarks. As detailed above, the distribution of Amphibolurus

centralis potentially overlaps with A. burnsi Wells and

Wellington, 1985, Gowidon longirostris Boulenger, 1883,

Lophognathus gilbert Gray, 1842 and L. horneri sp. nov. DNA

sequencing has confirmed that Amphibolurus centralis is

unrelated to the Gowidon and Lophognathus species and

demonstrates a clear sister-species relationship with A. burnsi.

Morphologically, A. centralis can be distinguished from A.

burnsi by having mostly homogeneous scales on the thighs,

lacking spinous scales on the thigh and having no enlarged

spinous scales along the rear of the thigh.

Genus Gowidon Wells and Wellington, 1984.

Gowidon Wells, R.W. and Wellington, C.R., 1983. A synopsis of

the Class Reptilia in Australia. Australian Journal of Herpetology 1:

73-129. Type-species Gowidon longirostris Boulenger, G.A., 1883 by

monotypy.

Diagnosis. A monotypic genus consisting of a large agamid lizard

in the subfamily Amphibolurinae with exposed tympanum, gular

scales smooth to weakly keeled, ventral scales smooth to weakly

keeled. Long-limbed, very long tail, long snout and distinct

nuchal crest. Head narrow and shallow in depth compared with

length of snout. Dorsal scales uniform, with keels converging

posteriorly toward midline. Prominent pale dorsolateral stripes

and pale stripe along lower jaw. One to three small white spots on

a black background positioned directly posterior to the ear.

Preanal pores 4-7; femoral pores range 11-22.

Included species. Gowidon longirostris Boulenger, G.A., 1883.

Distribution. Arid western interior of Australia. Semi-arboreal,

occurring in a broad range of habitats across arid and semi-arid

habitats, particularly associated with inland arid watercourses, gorges

and river beds.

Gowidon longirostris

(Figs. 5 & 8)

Gowidon longirostris Boulenger, G.A., 1883. Remarks of the

lizards of the genus Lophognathus. Annals and Magazine of Natural

Historyl2(5): 225-226 [225].

Physignathus eraduensis Werner, F., 1909. Reptilia exkl.

Geckonidae and Scincidae. In Michaelsen, W. and Hartmeyer, R.

(eds.) Die fauna siidwest-Australiens. Jena: Gustav Fischer 2: 251-278

[275]. Type data, holotype, status unknown, from Eradu, Western

Australia.

Physignathus longirostris quattuorfasciatus Sternfeld, R., 1924.

Beitrage zur herpetology inner-Australiens. Abhandlungen der

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

Figure 8. Photos in life of species under revision: a, Lophognathus horneri sp. nov., adult male with breeding colouration, 80 mile beach. Western

Australia (photo: R. Glor); b, Lophognathus gilberti, Katherine, Northern Territory (photo: R. Glor); c, Amphibolurus centralis, adult male with

breeding colouration. West MacDonnell Ranges, Northern Territory (photo: J. Melville); d, Amphibolurus burnsi, adult male with breeding

colouration, Westmar, Queensland (photo: S. Wilson); e, Tropicagama temporalis, Jabiru, Northern Territory (photo: S. Wilson); f, Gowidon

longirostris, adult male with breeding colouration, Ormiston Gorge, Northern Territory (photo: R. Glor).

Taxonomy in Australia’s tropical savanna lizards

Senckenbergischen Naturforschenden Gesellschaft 38: 221-251

[236]. Type data, lectotype SMF 10366, from Hermannsburg Mission,

Northern Territory. Designation by Mertens, R., 1967. Die

Herpetologische Section des Natur-Museums und Forschungs-

Institutes Senckenberg in Frankfurt-a-M. nebst einem Verzeichnis

ihrerTypen. 1. Senckenberg Biology 48: 1-106.

Synonymy that of: Melville, J., this work; Cogger, H.G., 1983, in

Cogger, H.G., Cameron, E.E., and Cogger, H.M. Amphibia and

Reptiles. Pp. 121-122 in: Walton, D.W. (ed.) Zoological catalogue of

Australia. Vol. 1. Netley, South Australia: Griffin Press Ltd. vi 313 pp.

[ 122 ],

Lectotype. BMNff 1946.8.28.73, Champion Bay, Western

Australia. Designation by Wells, R.W. and Wellington, C.R., 1985. A

classification of the Amphibia and Reptilia of Australia. Australian

Journal of Herpetology Supplementary Series 1: 1-61. Paralectotypes

BMNH 1946.8.12.64-65, Nickol Bay, Western Australia.

Diagnosis. As for genus.

Description of Lectotype. Adult. Moderately sized slender

agamid lizard with relatively long snout and dorsoventrally

compressed head. A distinct neck, very long limbs and very

long whip-like tail; canthus well defined; nasal scale below

canthal ridge, nare slightly to the posterior-dorsal section of

the nasal scale; visible tympanum. Infralabials 13; supralabials

15. Labials elongate unkeeled. Scales on dorsal surface of head

moderately heterogeneous and weakly to moderately keeled.

Low nuchal crest of slightly enlarged scales, extending as a row

of enlarged vertebral scales down the back to base of tail.

Lacking enlarged spinose scales on head or torso. Dorsal scales

on body and tail mostly homogeneous and moderately keeled.

Scales on thighs homogeneous and strongly keeled. Scales on

ventral surface of head strongly keeled and weakly keeled on

the body. Colour dorsally is light to dark brown and grey. Broad

white lip stripe, widest on lower jaw and narrow on upper lip,

extends below ear and as two broad white dorsoventral stripes

extending to mid-way down the back. A dark diffuse area of

pigmentation behind ear with diffuse pale spot immediately

behind the tympanum but the characteristic well-defined white

spot or spots on a black background behind the ear is not

apparent. Ventral surface of head, throat and upper chest darkly

pigmented with the dark pigmentation extending to the lateral

surfaces of the throat and up over the shoulders, bordering the

white dorsolateral stripes.

Variation. A few specimens examined lack the broad

dorsolateral stripes and are a pale grey colour, with a few rust-

brown coloured markings between the shoulders. An example

of this colour-morph is NMVD74317 collected on the Great

Northern Highway, 1 km E of Roebuck Roadhouse, Western

Australia (17° 48' 57"S, 122° 40' 44" E). However, these pale

colour morphs still retain the white spot or spots on a black

background behind the ear.

Distribution and ecology. Arid western interior of Australia.

Semi-arboreal, occurring in a broad range of habitats across

arid and semi-arid habitats, particularly associated with inland

arid watercourses, gorges and river beds.

Remarks. The distribution of Gowidon longirostris overlaps

with Amphibolurus centralis and Lophognathus horneri sp.

nov. but can be distinguished morphologically by having more

than 10 femoral pores, > 1 white spot on a black background

behind the ear, a relatively long snout and dorsoventrally

compressed head, and a very long whip-like tail.

Genus Lophognathus Gray, 1842

Lophognathus Gray, J.E., 1842. Description of some hitherto

unrecorded species of Australian reptiles and batrachians. Pp. 51-57 in:

Gray, J.E. (ed.). The zoological miscellany. London: Treuttel, Wiirz &

Co. [53]. Type-species Lophognathus gilberti Gray, 1842 by monotypy.

Redtenbacheria Steindachner, F., 1867. Redtenbacheria fasciata

Steindachner, F„ 1867. Reptilien. Pp. 1-98 in: Reise der

Osterreichischen Fregatte Novara um die Erde in den Jahren 1857,

1858, 1859 unter den Befehlen des Commodore B. von Wiillerstorff-

Urbair. Zoologie 1(3). Vienna: State Printer. [1869 on title page] [31].

[junior homonym of Redtenbacheria Schiner, 1861]. Type species.

Redtenbacheria fasciata Steindachner, 1867 by monotypy.

Synonymy that of: Cogger, H.G. 1983, in Cogger, H.G., Cameron,

E.E., and Cogger, H.M. Amphibia and Reptiles. Pp. 121-122 in:

Walton, D.W. (ed.) Zoological catalogue of Australia. Vol. 1. Netley,

South Australia: Griffin Press Ltd. vi 313 pp. [121]

Diagnosis. A genus consisting of large agamid lizards in the

subfamily Amphibolurinae, with exposed tympanum, gular

scales smooth to weakly keeled, ventral scales weakly to

strongly keeled. Stoutly built with moderately long legs and

tail. Broad white stripe on the upper and lower lips, extending

along the full extent of the jaw, pale stripe from behind the eye

to the top of the ear, which is cream, white, grey or yellow in

life. This pale stripe is well defined ventrally and dorsally by a

row of darkly pigmented scales (fig. 6). Heterogenous scales on

the back both at the midline and doroslaterally, associated with

a weak to prominent row of enlarged, strongly keeled scales.

Colour patterns of grey, brown, rust-brown and black. Well-

developed nuchal crest continuous with the enlarged row of

vertebral scales. Broad pale dorsolateral stripes, which may

extend from top of ear or back of head to hips. Dorsolateral

stripes are not continuous with the pale lip stripes. On the back,

dorsolateral stripes may be intersected by wedges of brown or

grey. Preanal pores 3-6; femoral pores 2-8.

Included species. Lophognathus gilberti Gray, J.E., 1842;

Lophognathus horneri sp. nov.

Distribution. Northern Australia, extending from northern-

central and western Queensland, through the northern regions

of the Northern Territory and across northern Western

Australia. Occurs in woodlands and river courses.

Lophognathus gilberti

(fig- 5)

Lophognathus gilberti Gray, J.E., 1842. Description of some

hitherto unrecorded species of Australian reptiles and batrachians. Pp.

51-57: in Gray, J.E. (ed.). The zoological miscellany. London: Treuttel,

Wiirz & Co. [53],

Redtenbacheria fasciata Steindachner, F. 1867. Reptilien. Pp.

1-98 in: Reise der Osterreichischen Fregatte Novara um die Erde in

den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B.

von Wiillerstorff-Urbair. Zoologie 1(3). Vienna: State Printer. [1869

on title page] [31]. Type data. Holotype whereabouts unknown (not

found), Australia.

Physignathus incognitus Ahl, E. 1926. Neue Eidechsen und

Amphibien. Zoologischer Anzeiger 67: 186-192 [190]. Type data.

Holotype ZMB 30086, Australia.

Synonymy that of: Melville, J., this work; Cogger, H.G. 1983, in

Cogger, H.G., Cameron, E.E., and Cogger, H.M. Amphibia and Reptiles.

Pp. 121-122 in: Walton, D.W. (ed.) Zoological catalogue of Australia.

Vol. 1. Netley, South Australia: Griffin Press Ltd. vi 313 pp. [121].

Holotype. BMNH 1946.8.28.69 from Port Essington, NT.

Diagnosis. As for genus. Lophognathus gilberti is distinguished

from Lophognathus horneri sp. nov. by lacking a distinct white

spot on the tympanum (fig. 7) that is surrounded by or adjacent

to black pigmentation.

Description of Holotype. A large robust male dragon lizard

with large robust limbs and tail. Large head in comparison with

body size, prominent nuchal crest of 18 enlarged spinose scales,

extending from anterior of ear to shoulders. Small nasal scale

and nares, below canthal ridge. Supralabials 13; anterior point

of lower jaw damaged. Head scales heterogeneous and strongly

keeled; 3-5 enlarged white spinose scales protruding from rear

of head, posterior to the jaw. Dorsal scales on body and tail

strongly keeled and heterogeneous. Gulars smooth and ventrals

weakly to strongly keeled. Scales on thighs heterogeneous and

strongly keeled. Very broad white lip stripes, extending under

jaw and up to anterior border of ear. Broad pale dorsolateral

stripes, continuous from neck to hips, bordered and well

defined by row of darker scales. Dorsolateral stripes

discontinuous with lip stripes. Poorly defined and discontinuous

pale stripe between eye and top of ear, bordered dorsally and

ventrally by darker scales. No clearly defined white spot on a

dark background on the tympanum, although there is a patch of

pale pigment in the upper-back quadrant of the tympanum.

Colour dorsally is light to dark brown and grey. Ventral surface

of head, throat and upper chest darkly pigmented with the dark

pigmentation extending to the lateral surfaces of the throat and

up over the shoulders, bordering the white dorsolateral stripes.

Femoral pores 6; preanal pores 4.

Variation. Some specimens of L. gilberti do have white areas

on the tympanum but they are not a well-defined spot

surrounded by the black pigmented area (fig. 7); instead, they

are a diffuse white or off-white smear or patch of pale pigment

without the associated black pigmentation. An example of this

is specimen NMVD74026 collected from Mt Wells Road, near

Grove Hill in the Northern Territory (13° 28' 47" S,

13° 132' 41" E), which has a smear of white pigmentation across

the posterior half of the tympanum.

Distribution and ecology. Far northern Australia in woodlands

and tropical savannahs. In the Northern Territory north of

Katherine, in Arnhem Land, and across coastal areas into

Western Australia and western Queensland. In Western

Australia, occurs north of Kununurra and extending up into the

eastern coastal Kimberley.

Remarks. Lophognathus gilberti shares similar body

proportions and meristic characters with L. horneri sp. nov.,

with extensive distributional overlap (fig. 4) but is readily

separated by the lack of a well-defined white spot on the

tympanum. Lophognathus gilberti is also superficially similar

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

to Amphibolous centralis and potentially has distributional

overlap, but it differs in mostly having a well-defined white or

pale stripe extending the full length between the ear and the

eye, and a broad white stripe running the extent of the upper lip

being mostly present.

Lophognathus horneri sp. nov.

ZooBank LSID: urn:lsid:zoobank.org:act: 4E027CDD-F9B2-

451B -B 08E-26D6A0B8A8ED.

(Figs. 8 & 9)

Grammatophora temporalis (part.) Gunther, A., 1867. Additions to

the knowledge of Australian reptiles and fishes. Annals and Magazine

of Natural History 20(3): 45-68 [52].

Synonymy that of: Melville, J., this work.

Holotype. NTM R16472 Sambo Bore, Wave Hill Station, Northern

Territory (18° 52' 48" S, 130° 40’ 12" E).

Paratypes. NMV D72658 Wave Hill Homestead, Northern Territory

(17° 23' 08" S, 131° 06' 44" E); NMV D73846 King Edward River Camp,

Mitchell Plateau, Kimberley, Western Australia (14° 52’ 57" S,

126° 12’ 10" E); NMV D74687 road to Davenport Ranges National Park,

Northern Territory (20° 37' 34" S, 134° 47' 14" E); WAM R131990

Kununurra, Kimberley, NE Western Australia (15° 48’ 0.00" S,

128° 43’ 0.12" E); WAM R108806 Mabel Downs Station, Calico

Springs, NE Western Australia (17° 16’ 59.88" S, 128° 10' 59.88" E);

WAM R132850 Kununurra, NE Western Australia (5° 47' 37.68" S,

128° 43' 10.92" E); BMNH 1946.8.12.73 Nickol Bay, Western Australia

[paralectotype Grammatophora temporalis (part.) Gunther, 1867].

Diagnosis. A member of the Australian genus Lophognathus

Gray, 1842, characterised by broad white stripe on the upper

and lower lips, extending along the full extent of the jaw, a pale

stripe from behind the eye to the top of the ear, which is cream,

white, grey or yellow in life. This pale stripe is well defined

ventrally and dorsally by a row of darkly pigmented scales (fig.

6). It is a large robust dragon with long head and well-built

moderately long limbs. It has heterogenous scales on the back,

both at the midline and dorsolaterally, associated with a weak

to prominent row of enlarged strongly keeled scales.

Lophognathus horneri is distinguished from Lophognathus

gilberti by the presence of a distinct white spot on the

tympanum (fig. 7). This well-defined white spot is wholly

surrounded or bordered dorsally and to the anterior by an area

of black pigmentation that is positioned on the upper posterior

quarter of the tympanum. This area of black pigmentation also

runs along a raised ridge that extends from the outer

dorsoposterior edge of the tympanum towards its centre (fig. 9).

Description of holotype. A large robust male dragon lizard

(snout-vent length: 102 mm; head length: 36.1 mm; head width

at widest point: 23.2 mm; head depth: 14.4 mm; hindlimb length:

102 mm). Head moderately long and wide, slightly rounded

profile of snout and slightly dorsolaterally compressed. Nuchal

crest low, extending from anterior of ear to shoulders and

composed of enlarged strongly keeled scales. Gulars smooth and

ventrals weakly to strongly keeled. Dorsal scales strongly keeled

and heterogeneous in size. Broad pale dorsolateral stripes,

continuous from neck to the hips, bordered and well-defined by

row of darker scales. Dorsolateral stripes discontinuous with lip

stripes. Well-defined pale stripe between eye and top of ear,

Taxonomy in Australia’s tropical savanna lizards

Figure 9. Holotype of Lophognathus horneri sp. nov. (NTM R16472).

bordered dorsally and ventrally by row of darker scales. Well-

defined white spot on the tympanum, which is adjacent to an area

of black pigmentation that runs along a raised ridge that extends

from the outer dorsoposterior edge of the tympanum towards its

centre. Femoral pores 2L/3R; preanal pores 4.

Variation. The white stripe on the lower lip and the white spot

on the tympanum are always present. The white stripe between

the eye and the ear is almost always present. The broad white

stripe on the upper lip can be pale and not prominent in a few

individuals. The pale dorsolateral stripes are not continuous

with the white lip stripes. Dorsolateral stripes are often absent

between the ear and neck or are intersected by wedges of

darker scales along the back. Three specimens collected from

80 Mile Beach (fig. 8), Western Australia (NMV D74362-

D74364), lacked a distinct stripe between the eye and the ear -

superficially similar to Amphibolurus centralis - but they still

had the white spot on the tympanum. A range of specimens

were examined from large adult males, females, juveniles and

hatchlings. All these specimens had a white stripe on the lower

lip and the white spot on the tympanum.

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

Coloration in alcohol. Specimens retain key diagnostic characters

in preservative. For example, the paralectotype Grammatophora

temporalis (part.) BMNH1946.8.12.73 from Nickol Bay, Western

Australia, which has also been designated as a paratype for L.

horneri sp. nov., was collected before 1867. This specimen,

which has been in preservative for over 150 years, still retains

diagnostic characters: broad white stripe on upper and lower lips,

white spot on the tympanum and a pale stripe between eye and

ear, which is bordered ventrally and dorsally by a row of darkly

pigmented scales.

Distribution and ecology. Arid and semi-arid eucalypt woodlands

and tropical savannahs of the central and western portions of

northern Australia. Specimens were collected in 2009 as far

south as the Davenport Ranges in the Northern Territory, which

are south-east of Tennant Creek. This species extends north of

Threeways but south of Katherine, west through Timber Creek

and Wave Hill and into Western Australia. They have been

collected in the Kununurra area, along the Gibb River Rd and in

the Mitchell Plateau area. In Western Australia they extend south

of Halls Creek and down to the northern Pilbara coast, south¬

west to Coral Bay and offshore islands.

Comparison with other species. Lophognathus horneri shares

similar body proportions and meristic characters with L. gilberti

with extensive overlap (Tables 2 and 3; fig. 4). It is readily

separated from this species by the presence of a well-defined

white spot on the tympanum (fig. 7), which is wholly surrounded

or bordered by an area of black pigmentation. Lophognathus

horneri is also superficially similar to Amphibolurus centralis,

but it differs in having proportionally shorter tail, hindlimbs

and head. Additionally, L. horneri has a well-defined white spot

on the tympanum and a well-defined stripe between the eye and

the ear (figs 6 and 7), which are lacking in A. centralis. Some

specimens of L. gilberti and A. centralis do have white areas on

the tympanum but they are not a well-defined spot surrounded

or adjoining the black pigmented area; instead, they are a

diffuse white or off-white smear or a patch of pale pigment

without the associated black pigmentation. Lophognathus

horneri can be distinguished from Tropicagama temporalis

gen. nov. by having a well-defined stripe between the eye and

the ear and heterogeneous dorsolateral scales along the back. In

addition, Lophognathus has > 2 preanal pores, whereas

Tropicagama temporalis gen. nov. has only two. Gowidon

longirostris differs from L. horneri by having very long limbs

and tail, being dorsoventrally compressed, having 1-3 white

spots on a black background behind the ear and having >10

femoral pores.

Etymology. This species is named in honour of Paul Horner, the

Curator of Terrestrial Vertebrates at the Museum and Art

Gallery of the Northern Territory, in recognition of his

contributions to the knowledge of the tropical lizard fauna of

Australia and his instrumental role in the taxonomic review of

agamid lizards from this region.

Genus Tropicagama gen. nov.

ZooBank LSID: urn:lsid:zoobank.org:act: F534B4D5-CBD7-

41E0-950A-B95B14F1D858.

Type-species. Grammatophora temporalis (part.) Gunther, A.,

1867. Additions to the knowledge of Australian reptiles and fishes.

Annals and Magazine of Natural History 20(3): 45-68 [52] [one of the

original syntype series, BMNFI 1946.8.12.73 (Nickol Bay), represents

Lophognathus horneri sp. nov.].

Diagnosis. A monotypic genus consisting of a large agamid

lizard in the subfamily Amphibolurinae, with exposed

tympanum, gular scales smooth to weakly keeled, ventral

scales smooth to weakly keeled. Very long-limbed, prominent

erectable nuchal crest. Long tail and head relatively narrow for

length. Dorsal scales uniform, with keels converging posteriorly

toward midline. Prominent pale dorsolateral stripes that are

broadly continuous with wide pale stripe along upper and lower

jaw. Lacks well-defined pale stripe between eye and ear. Upper

portion of head usually dark grey or black and uniformly

coloured. Under the head, on the chin, gular and neck areas,

there is dark grey or black uniform pigmentation in adult males,

with two narrow white stripes extending from the back of the

jaw anteriorly under the chin, parallel to the jaw, ending

approximately half way along the jaw. Femoral pores 1-6;

preanal pores 2 (range 1-3).

Included species. Grammatophora temporalis (part.)

Giinther, A., 1867.

Distribution. Far northern Australian coastal regions in the

Northern Territory and western Cape York. Also occurs on

northern offshore islands, including Indonesian islands close to

Australian waters and southern Papua New Guinea.

Tropicagama temporalis

(Figs. 5 & 8)

Grammatophora temporalis (part.) Gunther, A., 1867. Additions

to the knowledge of Australian reptiles and fishes. Annals and

Magazine of Natural History 20(3): 45-68 [52] [one of the original

syntype series, BMNH 1946.8.12.73 (Nickol Bay), represents

Lophognathus horneri sp. nov.].

Lophognathus lateralis Macleay, W., 1877. The lizards of the

Chevert Expedition. Second paper. Proceedings of the Linnean Society

of New South Wales 2: 97-104 [1878 on title page] [103]. Type data.

Holotype AM R31882, Mawatta, Binaturi River (as Katow), Papua

New Guinea.

Lophognathus labialis Boulenger, G. A., 1883. Remarks on the

lizards of the genus Lophognathus. Annals and Magazine of Natural

History 5 12: 225-226 [225], Type data. Syntype(s)' BMNH

1946.8.12.72, Port Essington, NT; BMNH 1946.8.12.63.

Lophognathus maculilabris Boulenger, G. A., 1883. Remarks on

the lizards of the genus Lophognathus. Annals and Magazine of

Natural History 12(5): 225-226 [226]. Type data. Syntypes: BMNH

1946.8.28.70-7i (BMNH 1919.8.26.13-14), Timor Laut Islands,

Indonesia.

Synonymy that of: Melville, J., this work; Cogger, H.G. 1983, in

Cogger, H.G., Cameron, E.E., and Cogger, H.M. Amphibia and

Reptiles. Pp. 121-122 in: Walton, D.W. (ed.) Zoological catalogue of

Australia. Vol. 1. Netley, South Australia: Griffin Press Ltd. vi 313 pp.

[ 122 ],

Lectotype. BMNH 1946.8.28.72, Port Essington, Northern

Territory. Designation by Cogger, H.G., 1983, in Cogger, H.G.,

Cameron, E.E., and Cogger, H.M. Amphibia and Reptiles. Pp. 121—

122 in: Walton, D.W. (ed.) Zoological catalogue of Australia. Vol. 1.

Taxonomy in Australia’s tropical savanna lizards

Netley, South Australia: Griffin Press Ltd. vi 313 pp. [122].

Paralectotype. BMNH 1946.8.12.63, Nickol Bay, WA.

Diagnosis. As for genus.

Description of Lectotype. Adult. Moderately sized slender

agamid lizard with relatively short rounded snout; head narrow

and moderately elongated. Prominent canthal ridge consisting

of row of enlarged heavily keeled scales. Small nasal scale and

nares below ridge. Visible tympanum. A distinct neck, long

limbs, long and slender tail, which is damaged and missing its

end. Infralabials 11; supralabials 12. Labials elongate unkeeled.

Scales on dorsal surface of head moderately heterogeneous,

strongly keeled. Prominent nuchal crest of 10 enlarged scales,

from ear to shoulder and extending along the back as a row of

enlarge scales to base of tail. Head scales heterogeneous and

strongly keeled; 4-5 enlarged white spinose scales protruding

from rear of head, posterior to the jaw. Dorsal scales on body

and tail mostly homogeneous and weakly to moderately keeled.

Scales on thighs homogeneous and strongly keeled. Scales on

the ventral surface of head strongly keeled and are weakly

keeled on the body. Upper portion of head dark grey-brown and

uniformly coloured. Under the head, on the chin, gular and

neck dark grey pigmentation, with two pale stripes extending

from back of the jaw anteriorly under chin, parallel to jaw,

ending approximately half way along jaw. Broad white lip

stripe, equally wide on lower and narrow upper lip, extends

below ear and continues as two broad white dorsoventral

stripes, extending to shoulder. Broad dorsoventral stripes

intersected by three dark bands at neck, shoulders and upper

back.

Variation. Adult males have dark grey to black uniform colour

on top of head and below chin, onto neck and ventral surface of

shoulders. However, females and juveniles often lack this

uniform colour and instead have brown and black patterning on

top of the head and only have flecks of grey, brown or black on

their ventral surface. However, they still have two continuous

dorsolateral stripes from the jaw onto the back and lack a well-

defined pale stripe between the eye and the ear. Also, it is

common for Tropicagama temporalis to have one or more

broad dark lateral bands across the back at the shoulders.

Commonly, the white lower lip stripe does not extend the full

length of the jaw; instead, it is only present on the posterior

section of the lower jaw.

Distribution and ecology. Far northern Australian coastal

regions in the Northern Territory and western Cape York. Also

occurs on northern offshore islands, including Indonesian

islands close to Australian waters and southern Papua New

Guinea. Semi-arboreal, occurring in dry tropical woodland

habitats, particularly associated with coastal pandanus and

paperbark watercourses. Genetic data has not yet found

evidence of this species occurring on the Western Australian

mainland. However, future work may find this species

occurring in coastal Kimberley regions.

Remarks. Tropicagama temporalis is superficially similar to,

and has extensive distributional overlap with, Lophognathus

gilbert (fig. 4). T. temporalis is readily separated from this

species on body proportions (fig. 2) and by having two or fewer

preanal pores, uniform dorsal scales with keels converging

posteriorly toward midline, a prominent pale dorsolateral stripe

that is broadly continuous with stripe along jaw, and lacking a

well-defined pale stripe between the eye and the ear.

Identification key for Amphibolurus, Chlamydosaurus and

Lophognathus

1. No large “frill” of skin around neck.2

A large, loose “frill” of skin around neck.

Chlamydosaurus kingii

2. Fewer than 10 femoral pores, no white spots on a black

background behind the ear.3

More than 10 femoral pores, > 1 white spot on a black

background behind the ear, relatively long snout and

dorsoventrally compressed head, very long whip-like tail

. Gowidon longirostris

3. More than 2 preanal pores, dorsal scales heterogeneous,

dorsolateral stripes discontinuous with wide pale stripe

along upper or lower jaw (if present).4

Two or fewer preanal pores, dorsal scales uniform, with

keels converging posteriorly toward midline. Prominent

pale dorsolateral stripes that are broadly continuous with

wide pale stripe along upper and/or lower jaw. Lacks well-

defined pale stripe between eye and ear.. Tropicagama

temporalis

4. Lacks well-defined white or pale stripe extending full

length between ear and eye (fig. 6), which is defined

dorsally and ventrally by darker scales. Broad white stripe

running extent of upper lip mostly absent.5

Well-defined white or pale stripe extending full length

between ear and eye present (fig. 6). Broad white stripe

running extent of upper lip mostly present.6

5. Scales on thighs strongly heterogeneous, with scattered

spinous scales and a row of small to large spinous scales

running along rear of thigh.7

Scales on thighs mostly homogeneous, lacking spinous

scales on the thigh and row of enlarged spinous scales

running along rear of thigh absent. Amphibolurus

centralis

6. Well-defined white spot on tympanum is present (fig. 7). It

is wholly surrounded or bordered by an area of black

pigmentation that runs along a raised ridge, which extends

from the outer dorsoposterior edge of the tympanum

towards its centre. Lophognathus horneri

Well-defined white spot on tympanum absent (fig. 7). If

white or pale pigmentation is present on the tympanum, it

is not a well-define spot or not wholly surrounded or

bordered by an area of black pigmentation that runs along

a raised ridge, which extends from the outer dorsoposterior

edge of the tympanum towards its centre. Lophognathus

gilberti

7. Prominent spinose nuchal and vertebral crest, and two or

more additional dorsal crests. A row of large spinose

scales running along rear of thigh. Amphibolurus burnsi

Lacking multiple additional dorsal crests and row of only

small spinose scales running along rear of thigh

8. Dark strip between nostril and eye ..Amphibolurus norrisi

Dark stripe between nostril and eye absent.

Amphibolurus muricatus

Discussion

Molecular work, incorporating both mitochondrial and

nuclear gene regions, provided compelling evidence that

taxonomic revision of the genera Amphibolurus, Gowidon

and Lophognathus was required (Melville et al., 2011). This

molecular work also provided a starting point for

morphological analyses, allowing us to determine diagnostic

characters for each species. Until now, without the aid of

molecular data, it has been difficult for taxonomists to

determine which morphological characteristics were

diagnostic. These past difficulties and our current study

have shown that there is significant morphological

homoplasy across these agamid species, with significant

overlap in morphometries, colour, patterning and scalation.

We have determined that there are four independent

evolutionary lineages in the former genus Lophognathus

(now Gowidon, Lophognathus, Amphibolurus and

Tropicagama ). Despite the deep divergences between the

four lineages, there are extensive morphological similarities

in the constituent species. In addition, the fact that we have

found that the junior synonym of one Lophognathus species

was an unrelated species, now Amphibolurus centralis,

demonstrates the extent of homoplasy in these dragons.

It seems that most of the difficulties in determining

species in these genera have occurred in northern Australia,

with Amphibolurus centralis, Gowidon longirostris,

Tropicagama temporalis, Lophognathus gilberti and

Lophognathus horneri all occurring in northern and north¬

western Australia. This geographic concentration of

taxonomic problems may result from several reasons. A lack

of dedicated taxonomic research into the genera

Lophognathus and Amphibolurus in northern Australia may

be an important factor. There has been a paucity of

taxonomic research undertaken on the Australian agamid

lizards, particularly those of the Northern Territory and

western Queensland. Glenn Storr from the Western

Australian Museum provided much of the foundational work

on the taxonomy of dragons in the arid and semi-arid regions

of Australia but his taxonomic work did not include

Amphibolurus and Lophognathus, except for his taxonomic

review of bearded dragons and the dismemberment of the

Amphibolurus genus (Storr, 1982). Since Storr, there has

been little taxonomic work conducted on northern Australian

agamids. Similar taxonomic problems have arisen in

dragons occurring in the eastern states of Australia, where

more research is focused. For example, Witten (1972)

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

described Amphibolurus nobbi from Queensland, New

South Wales and Victoria, but more recent work has

demonstrated that this species is in the genus Diporiphora

(Edwards and Melville, 2011). Following Witten’s taxonomic

placement of the nobbi dragon into Amphibolurus, Greer

(1989) noted that the colour patterns of the nobbi dragon

were similar to other species of Diporiphora, rather than

Amphibolurus, with a pink or rose flush to the base of the

tail and yellowish sides. Thus, it appears that Witten (1972)

selected characters that were not correlated with evolutionary

relatedness when placing the nobbi dragon into

Amphibolurus. Consequently, the problems with the

taxonomic resolution in Amphibolurus and Lophognathus

have probably resulted from a combination of little research

and difficulties in morphological taxonomic research into

these genera.

There is a reoccurring theme of non-diagnostic

characters and morphological homoplasy in the taxonomy of

Australian agamids. The use of molecular tools in the

taxonomy of Amphibolurinae has greatly improved

resolution and nomenclatural stability that has not been

possible with morphology alone. In addition, these molecular

data allow insight into evolutionary patterns in this diverse

group of lizards, where there is both a high degree of

morphological diversity and homoplasy across the subfamily.

Our study has demonstrated the high level of morphological

homoplasy across independent evolutionary lineages. Our

results provide significant scope for future research into the

evolutionary processes underlying the morphological

convergence or parallelism in Australian agamid lizards.

Supporting information

Supplementary Appendix SI List of museum specimens

examined morphologically.

(PDF)

Acknowledgments

JM examined specimens for morphological review at the

Australian Museum, Western Australian Museum, British

Museum of Natural History, Museum Victoria, Museum of

Comparative Zoology (Harvard University), Senckenberg

Naturmuseum (Frankfurt-am-Main) and Zoological

Research Museum Alexander Koenig (Bonn, Germany). JM

would like to thank staff and researchers from these

institutions for their help and assistance, including W.

Longmore, R. O’Brien, D. Bray, P. Doughty, R. Sadlier, J.

Rosado, C. McCarthy and, in particular, P. Wagner for

organising type loans from other museums within Germany

to be examined at the Zoological Research Museum

Alexander Koenig (Bonn, Germany). We thank S. Wilson

for use of images. We thank G. Shea for advice on

nomenclature and discussions regarding synonymies.

Research funding provided to JM by Australian Research

Council and to JM, RG and JS by the Australian Biological

Resources Study.

Taxonomy in Australia’s tropical savanna lizards

References

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Supplementary Appendix SI.

Voucher specimens examined

Museum abbreviations are: AM for Australian Museum

Sydney, NMV for Museum Victoria Melbourne, WAM for

Western Australian Museum.

Amphibolurus burnsi

NMV D52082,8 km E of Surat, Queensland, 27° 09' S, 149° 05' E;

NMV D56415, St George, Queensland, 28° 03' S, 148° 35' E;

NMV D56416, St George, Queensland, 28° 03' S, 148° 35' E;

NMV D74134,10 km N of Chinchilla, Queensland, 26° 40' 23" S,

150° 35' 52" E; NMV D74135, 15 km W of Cecil Plains,

Queensland, 27° 33' 29" S, 151° 03' 04" E; NMV D74137,15 km

W of Cecil Plains, Queensland, 27° 32' 44" S, 151° 04' 55" E; AM

R151557, Gingham Floodplain, Te Mona, 3 km N of Allombie

Bridge over Gwydir River, New South Wales, 29° 19' S,

149° 27' E; AM R151838, Kwiambai National Park, New South

Wales, 29° 11' 13" S, 150° 57' 51" E; AM R155887, Brewarrina,

Barwon River at West Brewarrina, New South Wales, 29° 58' S,

146° 52' E; AM R166772, Nocoleche Nature Reserve, New South

Wales, 29° 51' 15" S, 144° 8' 8" E; AM R166873, AM R166874,

Wanaaring township, New South Wales; AM R166910, Nocoleche

Nature Reserve, New South Wales, 29° 51' 26" S, 144° 8' 9" E;

AM R137612, Byerawering Property, Site 1, Banks of Culgoa

River, 30 km from Goodooga Road, New South Wales, 29° 5' S,

147° 8' E; AM R148373, Macquarie Marches, Sandy Camp

Property, 8 km N of Homestead on Carinda River, New South

Wales, 30° 49' S, 147° 43' E; AM R153324 Yuleba State Forest

(South of Condamine Hwy.), Coup 105, Queensland,

26° 54' 13" S, 149° 44' 18" E.

Amphibolurus centralis

NMV D11166, Attack Creek, Northern Territory, 18° 18' S,

134° 35' E; NMV D175, NMV D176, NMV D182, NMV D184,

NMV D189, NMV D196, Alice Springs, Northern Territory,

23° 42' S, 133° 52' E; NMV D492, Illamurta, Northern Territory,

24° 18' S, 132° 41' E; NMV D72659, Buntine Highway, 20 km E

of Kalkaringa, Northern Territory, 17° 24' 08" S, 130° 56' 56" E;

NMV D72661, NMV D72680, Buchanan Highway, 101 km E of

Top Springs, Northern Territory, 16° 45' 25" S, 132° 40' 20" E;

NMV D72709, 108 km S of Cape Crawford on Tablelands

Highway, Northern Territory, 17° 32' 24" S, 135° 41' 17" E; NMV

D72710, 3 km S of Heartbreak Inn on Tablelands Highway,

Northern Territory, 16° 42' 14" S, 135° 43' 46" E; NMV D73981,

Kimberley, Duncan Road, N of Spring Creek Station, Western

Australia, 16° 25' 02" S, 129° 02' 54" E; NMVD73988, Buchanan

Highway S of Jasper Creek, Northern Territory, 16° 06' 55" S,

130° 53' 23" E; NMV D74005, Nutwood Road, 3 km SE, N of

Daly Waters, near Scarlet Hill, Northern Territory, 16° 11' 03" S,

133° 26' 28" E; NMV D74021, Carpentaria Highway, W of

Northern Territory-Queensland border, SE of Robinson River

Aerodrome, Northern Territory, 16° 51' 43" S, 137° 14' 39" E;

NMV D74023, Carpentaria Highway, 125 km E of Hi Way Inn,

Northern Territory, 16° 28' 59" S, 134° 25' 26" E; NMV D74052,

Lawn Hill Road via Gregory Downs Station, Queensland,

18° 38' 28" S, 139° 05' 55" E; NMV D74283, road to Hamilton

Station Youth Camp, off Tanami Track, Northern Territory,

J. Melville, E.G. Ritchie, S.N.J. Chappie, R.E. Glor & J.A. Schulte II

23° 36' 07" S, 133° 34' 36" E; NMV D74288, Red Bluff Gorge,

Northern Territory, 23° 36' 07" S, 132° 30' 38" E; NMV D74673,

Hodgson River Road, near Tennant Creek, 2 km E of Stuart

Highway, Northern Territory, 16° 12' 18" S, 133° 25' 50" E; NMV

D74695, West MacDonnell Ranges, Haast Bluff Road, Northern

Territory, 23° 33' 14" S, 132° 18' 21" E.

Lophognathus gilberti

NMV D290, Amhem Land, Oenpelli, East Alligator River,

Northern Territory, 12° 19' S, 133° 03' E; NMV D34178, NMV

D34179, NMV D34180, Ord River Dam area, Western Australia,

17° 24' S, 128° 52' E; NMV D5096, NMV D5123, Borroloola,

Northern Territory, 16° 04' S, 136 18' E; NMV D5150, Amhem

Land, Upper Roper River, Northern Territory, 14° 44' S,

134° 31' E; NMV D5222, Amhem Land, Oenpelli, Northern

Territory, 12° 19' S, 133° 03' E; NMV D72579, West Amhem

Land, Yirrkakak, Northern Territory, 12° 12' 14" S, 133° 48' 04" E;

NMV D72590, West Amhem Land, Gubjekbinj, Northern

Territory, 12° 15' 09" S, 133° 48' 07" E; NMV D74026, Mt Wells

Road, near Grove Hill, Northern Territory, 13° 28' 47" S,

131° 32' 41" E; NMV D74027, Wells Road, near Grove Hill

Station, Northern Territory, 13° 28' 42" S, 131° 32' 13" E; NMV

D74258, 20 km N of Katherine, Stuart Highway, Northern

Territory, 14° 18' 57" S, 132° 06' 18" E; NMV D74260, Mount

Wells, Northern Territory, 13° 40' 25" S, 132° 48' 26" E; NMV

D74263, road to Umbarrumba Gorge, S of Pine Creek, Northern

Territory, 13° 51' 19" S, 131° 49' 14" E; NMV D74280, Mount

Wells Road, Northern Territory, 13° 29' 31" S, 131° 34' 25" E;

NMV D74281, NMV D74282, Mount Wells Road, Northern

Territory, 13° 29' 33" S, 131° 36' 44" E; NMV D74286, off Stuart

Highway, 42 kmN of Katherine, Northern Territory, 14° IT 12" S,

132° 01' 52" E; NMV D74289, NMV D74293, Umbrawarra

Gorge, Northern Territory, 13° 57' 14" S, 131° 41' 39" E; NMV

D74298, Marrakai Road, about 8 km SW of Amhem Highway,

Northern Territory, 12° 47' 59" S, 131° 26'38" E. WAMR126027,

WAM R126029, WAM R126031,4 km SW of Point Spring Yard,

WestemAustralia, 15° 25' 35" S, 128° 51' 09" E; WAM R163559,

Anjo Peninsula, Western Australia, 14° 04' 19" S, 126° 24' 27" E.

Gowidon longirostris

NMV D222, NMV D224, NMV D231, NMV D235, Central

Australia, Dalhousie (original label in Spencer's writing

“Dalhousie"), Northern Territory, 26° 30' S, 135° 28' E; NMV

D2336, NMV D3481, Finke River, Northern Territory, 25° 02' S,

134° 24' E; NMV D239, Derwent Creek, Northern Territory,

17° 34' S, 145° 13' E; NMV D4945, NMV D4946, Central

Australia, Barrow Creek, Northern Territory, 21° 32' S, 133° 53' E;

NMV D50509, NMV D50510, NMV D50511, Todd River, Alice

Springs, Northern Territory, 23° 48' S, 134° 25' E; NMV D50537,

Emily Gap, MacDonnell Ranges, Northern Territory, 23° 45' S,

133° 57' E; NMV D509, NMV D510, NMV D511, Charlotte

Waters, Northern Territory, 25° 56' S, 134° 53' E; NMV D5383,

NMV D5384, no data; NMV D56308, Alice Springs, Northern

Territory, 23° 42' S, 133° 52' E; NMV D67487, Finke River,

Northern Territory, 25° 02' S, 134° 24' E; NMVD67644, Simpsons

Gap, Northern Territory, 23° 42' 07" S, 133° 43' 05" E; NMV

D67663, NMV D67665, NMV D67668, Ellery Creek, Northern

Territory, 23° 47' 10" S, 133° 04' 07" E; NMV D67669, NMV

D67670, NMV D67671, NMV D67672, NMV D67703,

Taxonomy in Australia’s tropical savanna lizards

Serpentine Gorge, Northern Territory, 23° 45' 25" S,

132° 58' 24" E; NMV D67713, NMV D67714, Ormiston Pound,

near creek, Northern Territory, 23° 37' S, 132° 48' E; NMV

D67758, Finke River, Palm Valley, old rangers' station. Northern

Territory, 24° 03' 21" S, 132° 45' 22" E; NMV D67764, NMV

D67765, NMV D67766, Palm Valley, Finke River, Northern

Territory, 24° 02' 33" S, 132° 42' 22" E; NMV D67777, NMV

D67778, Finke River, Palm Valley, Red Gum site near old rangers'

station, Northern Territory, 24° 03' 06" S, 132° 45' 18" E; NMV

D67786, Central Australia, waterhole, 3.4 km from Boggy Hole,

Northern Territory, 24° 08' 07" S, 132° 50' 07" E; NMV D69,

NMV D70, Tennant Creek, Northern Territory, 19° 39' S,

134° ll l E; NMV D72733, Todd River, East MacDonnell Ranges,

Northern Territory, 23° 47' 45" S, 134° 18' 42" E; NMV D74269,

Ormiston Gorge, Northern Territory, 23° 37' 56" S, 132° 43' 39" E;

NMV D74317, Great Northern Highway, 1 km E of Roebuck

Roadhouse at entrance to Kilto Station, Western Australia,

17° 48' 57" S, 122° 40' 44" E; NMV D74368, NMV D74370,

NMV D74371, Great Northern Highway, N of Sandfire

Roadhouse, Stanley picnic stop, Western Australia, 19° 02' 35" S,

121° 39' 57" E; NMV D74391, Pilbara, Ingee Station, off Great

Northern Highway, Western Australia, 20° 46' 42" S,

118° 31' 33" E; NMV D74407, Great Northern Highway, South

Gascoyne River, 64 km S of Kumarina Roadhouse, Western

Australia, 25° 12' 09" S, 119° 20' 05" E; NMV D74408, Great

Northern Highway, South Gascoyne River, 64 km S of Kumarina

Roadhouse, Western Australia, 25° 12' 09" S, 119° 20' 05" E;

NMV D74426, Tjukayiria Roadhouse, about 3 km W, Warburton

Hwy, Western Australia, 27° 09' 57" S, 124° 32' 54" E; NMV

D95, NMV D96, NMV D97, Oodnadatta, South Australia,

27° 33' S, 135° 27' E.

Lophognathus horneri sp. nov.

NMV D10440, NMV D10441, NMV D10442, NMV D10443,

NMV D10444, NMV D10838, NMV D10887, Timber Creek,

Northern Territory, 15° 39' S, 130° 29' E; NMV D2360, NMV

D2365, Port George IV, Western Australia, 15° 22' S, 124° 39' E;

NMV D2934, NMV D5630, Tennant Creek, Northern Territory,

19° 39' S, 134° IP E; NMV D72638, NMV D72652, Montejinni

Creek, Buntine Highway, Northern Territory, 16° 38' 06" S,

131° 45' 20" E; NMV D72643, Willaroo Station, Top Springs,

track off Victoria Highway, Northern Territory, 15° 18' 55" S,

131° 34' 10" E; NMV D72658, Kelly Creek, Wave Hill

Homestead, Northern Territory, 17° 23' 08" S, 131° 06 ‘44" E;

NMV D73807, NMV D73809, NMV D73810, NMV D73811,

Kimberley, Gibb River Road crossing of the Durack River,

Western Australia, 15° 58' 26" S, 127° 09' 13" E; NMV D73818,

Kimberley, Gibb River Road, 10 km W of Ellenbrae Station,

Western Australia, 15° 59' 27" S, 126° 57' 10" E; NMV D73820,

Kimberley, Gibb River Road, 8 km W of Ellenbrae Station,

Western Australia, 15° 59' 24" S, 127° 00' 16" E; NMV D73845,

NMV D73846, Kimberley, Mitchell Plateau, King Edward River

Camp, Western Australia, 14° 52' 57" S, 126° 12' 10" E; NMV

D73851, Kimberley, Gibb River Road, 1 km W of the Kalumburu

tum-off, Western Australia, 16° 09' 19" S, 126° 30' 10" E; NMV

D73863, Kimberley, Mt Elizabeth, Western Australia,

16° 13' 59" S, 125° 59' 21" E; NMV D73867, Kimberley, Gibb

River Road, W of Snake Creek, Western Australia, 16° 32' 15" S,

126° 15' 29" E; NMV D73886, Kimberley, Derby Caravan Park,

Western Australia, 17° 18' 30" S, 123° 37' 44" E; NMV D73967,

Kimberley, Duncan Road, N of Spring Creek Station, Western

Australia, 16° 19' 43" S, 129° 03' 33" E; NMV D73984, Buchanan

Highway S of Jasper Creek, Northern Territory, 16° 02' 45" S,

130° 51' 59" E; NMV D74259, Stuart Memorial, Stuart Highway,

Northern Territory, 19° OP 24" S, 132° 08' 30" E; NMVD74271,

Stuart Memorial, 48 km N of Three Ways, Stuart Highway,

Northern Territory, 19° OP 24" S, 134° 08' 30" E; NMV D74309,

NMV D74312, Wolfe Creek, Tanami Road, Western Australia,

18° 59' 44" S, 127° 41' 52" E; NMV D74334, Derby, road to

Prison BoabTree, WestemAustralia, 17° 21'04" S, 123° 40' 09" E;

NMV D74362, NMV D74363, NMV D74364, 80 Mile Beach

Caravan Park, 50 km S of Sandfire, Western Australia,

19° 45' 16" S, 120° 40' 20" E; NMV D74683, road to Karundi, 35

km E of Stuart Highway, Northern Territory, 20° 28' 22" S,

134° 28' 51" E; NMV D74687, road to Davenport Ranges

National Park, near Tennant Creek, Northern Territory,

20° 37' 34" S, 134° 47' 14" E; NMV D74690, road to Davenport

Ranges National Park, near Tennant Creek, Northern Territory,

20° 38' 28" S, 134° 46' 42" E; WAM R102296, WAM R102307,

Hermite Island (South), Montebello Islands, Western Australia,

20° 28' 00" S, 115° 31' 00" E; WAM R102321, Ah Chong Island,

Montebello Islands, Western Australia, 20° 31' 00" S,

115° 33' 00" E; WAM R108806, Calico Springs, Western

Australia, 17° 17' 00" S, 128° IP 00" E; WAM R113222, South

Muiron Island, Western Australia, 21° 42' 00" S, 114° 48' 00" E;

WAM R113987, King Edward River, Western Australia,

14° 55' 00" S, 126° 12' 00" E; WAM R114385, Coulomb Point

Nature Reserve, WestemAustralia, 17° 22' 00" S, 122° 09' 00" E;

WAM R131275, Fitzroy Crossing, Western Australia,

18° 10' 58" S, 125° 36' 00" E; WAM R131990, Kununurra,

WestemAustralia, 15° 48' 00" S, 128° 43' 00" E; WAM R132850,

Kununurra, Western Australia, 15° 47' 38" S, 128° 43' 11" E;

WAM R132851, Kununurra, Western Australia, 15° 47' 38" S,

128° 43' 11" E; WAM R139477, WAM R139481, Potter Island,

WestemAustralia, 20° 56' 15" S, 116° 09' 10" E; WAMR141302,

Cape Preston Area, Western Australia, 20° 52' 59" S,

116° IP 41" E.

Tropicagama temporalis

NMV D14455, NMV D14456, NMV D14457, NMV D14458,

NMV D14459, NMV D14460, NMV D14461, Gulf Province,

Balimo, Papua New Guinea, 08° 00' S, 142° 55' E; NMV D2289,

NMV D2293, NMV D3732, NMV D3733, NMV D5924, NMV

D5925, Borebada (as “Bora Bada, New Guinea"), 09° 28' S,

147° 12' E; NMV D4551, Northern Territory; NMV D49838,

NMV D49839, NMV D49840, NMV D49841, NMV D49842,

NMV D49889, NMV D50083, NMV D50084, NMV D50085,

NMV D50086, NMV D50087, NMV D50088, NMV D50089,

Gulf Province, Aramia River, Awaba, Papua New Guinea,

08° OP S, 142° 45' E; NMV D5234, Melville Island, Northern

Territory, 11° 27' S, 130° 47' E; NMV D5517, Darwin, Northern

Territory, 12° 27' S, 130° 50' E; NMV D6655; NMV D74299,

Marrakai Road near Heather Lagoon, Northern Territory,

12° 54' 40" S, 131° 12' 24" E; NMV D74306, NMV D74307,

Darwin, Malack, Wescombe Court, Northern Territory,

12° 23' 26" S, 130° 54' 24" E; NMV D74311, NMV D74313,

NMV D74314, NMV D74318, NMV D74319, Darwin, gardens

at Museum and Art Gallery of the Northern Territory, Northern

Territory, 12° 26' 16" S, 130° 50' 03" E.

Memoirs of Museum Victoria 77:63-104 (2018) Published 2018

1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.Org/10.24199/j.mmv.2018.77.05

Recognising variability in the shells of argonauts (Cephalopoda: Argonautidae): the

key to resolving the taxonomy of the family

Julian K. Finn

Sciences, Museums Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia. Email: jfinn@museum.vic.gov.au

Abstract Finn, J.K. 2018. Recognising variability in the shells of argonauts (Cephalopoda: Argonautidae): the key to resolving the

taxonomy of the family. Memoirs of Museum Victoria 77: 63-104.

Argonauts (Cephalopoda: Argonautidae) are a family of pelagic octopuses that are most commonly recognised by the

beautiful white shells of females (known as paper nautiluses), prized by beachcombers the world over. Taxonomic delineation

of the group has historically relied exclusively on features of the shells of females and has resulted in more than 50 species

names being coined worldwide. This approach has created considerable confusion in the taxonomy of the family because

argonaut shells are not true molluscan shells and display considerable variation in form. This study closely examined a large

number of argonaut shells from museum collections throughout the world. Two types of shell formation that had been

previously attributed to separate argonaut species were recognised within individual shells. It is proposed here that the

different shell forms reflect the effects of ecological or biological factors or events, often manifesting as dramatic changes

in shell growth and shape within the development of an individual shell. The resulting combinations of shell formation types

clearly explain both the extreme variation observed across large numbers of argonaut shells and the high number of nominal

species names coined in the past. Researchers coining new fossil argonaut species based solely on shell characters are

advised to proceed with caution. This study supports parallel morphological and molecular research recognising the

existence of only four extant argonaut species worldwide: Argonauta argo, A. hians , A. nodosus and A. nouryi.

Keywords paper nautilus, Argonauta argo, Argonauta hians, Argonauta nodosus, Argonauta nouryi, Coleoidea, Octopoda, shell

morphometries

Introduction

Argonauts (Cephalopoda: Argonautidae) are a family of

pelagic octopuses that inhabit tropical and temperate oceans

of the world (fig. 1). Derived from benthic octopus ancestors,

argonauts have departed the sea floor to carry out their entire

life cycle in the open ocean (Young et al., 1998). Argonauts are

most widely recognised by the beautiful white shells of

females that are commonly known as paper nautiluses and are

prized by beachcombers the world over. These shells function

as both brood chambers for the females’ eggs (Naef, 1923) and

hydrostatic structures by which female argonauts are able to

attain neutral buoyancy (Finn and Norman, 2010).

Images of argonaut shells have a long history, adorning

artefacts dating back to Minoan civilisations (3000-1050 BC;

Walters, 1897; Mackeprang, 1938; Hughes-Brock, 1999) and

featuring in the earliest conchological works (e.g. Rumphius,

1705; Argenville, 1742; Gualtieri, 1742; Seba, 1758; Martini,

1769). By contrast, the identity of the occupant of the shell

(i.e. the argonaut) has remained largely unknown or

misinterpreted. For example, in the early 1800s it was widely

believed that the octopus commonly found in the argonaut

shell was not the rightful owner, but was a parasite having

devoured the original occupant (Sowerby and Sowerby,

1820-1825; Broderip, 1828).

In the absence of knowledge about the animals that created

the shells, a taxonomic system that relied completely on shell

features arose for the family. Variations in shell shape and

appearance formed the basis of new species descriptions,

giving rise to 53 species names and 11 subspecies names

worldwide (Sweeney and Young, 2004).

At the core of this taxonomic system is the issue that

argonaut shells display a considerable degree of variability.

This variability has been observed across shells produced by

individuals of the same species (Voss and Williamson, 1971)

and even between opposing faces of the same shell (Cotton and

Godfrey, 1940; Trego, 1992). This variability is likely to be

exacerbated by the female argonaut’s ability to repair (Power,

1856; Hoyle, 1886; Boletzky, 1983; Trego, 1993) and completely

rebuild the shell (Holder, 1909a, 1909b; Alliston, 1983).

Argonaut shells are not true molluscan shells. Unlike the

shells of other molluscs (e.g. gastropods), argonaut shells are

not produced by the derivatives of the shell field (the mantle

epithelium responsible for shell secretion in other molluscs;

see Kniprath, 1981). In argonauts, the shell field disappears

during embryonic development (Kniprath, 1981). The argonaut

J.K. Finn

Figure 1. Live female argonaut (Argonauta argo) observed swimming close to the ocean surface and holding her white paper nautilus shell that

functions as a brood chamber for the female’s eggs and as a hydrostatic structure for maintaining neutral buoyancy.

shell is a secondary calcium carbonate structure secreted from

webs on the distal ends of the female argonaut’s first (dorsal)

arm pair.

Female argonauts commence forming shells approximately

12 days after hatching (A. argo: Power in Roberts, 1851; Power

in Catlow, 1854) at a size of approximately 5-7 mm mantle

length (A. argo : Jatta, 1896; Naef, 1923; A. hians : Nesis, 1977;

A. nouryi : Finn, 2009). The initial shell is formed without

sculpturing (Jatta, 1896). By the time the female argonaut

reaches 10 mm mantle length, the shell (which is now 14 mm

in length) is fully formed (A. hians : Nesis, 1977). The webs on

the female’s dorsal arms overlap the edge of the shell and add

to it as the female grows. Irregularities in the lay of the web

along the shell edge is presumed to cause the undulations in

the surface of the shell, visible as radiating ridges (or ribs) in

fully formed shells (Mitchell et al., 1994).

Once the female argonuat reaches maturity, she lays long

strands of eggs that are attached to the internal central axis of

the shell. Female argonauts are continuous spawners (Boletzky,

1998; Rocha et al., 2001; Laptikhovsky and Salman, 2003) with

asynchronous ovulation and monocyclic spawning (i.e. egg-

laying occurring over an extended and continuous spawning

period in relation to the animal’s life; Rocha et al., 2001).

Spawning is thought to extend over several months (Boletzky,

1998) and based on published counts, proposed spawning

frequencies and proposed spawning durations, it has been

surmised that the potential fecundity of a female A. argo could

exceed one million eggs (Laptikhovsky and Salman, 2003).

To stabilise argonaut taxonomy, the aim of this study was

to examine the inter- and intra-specific variation in argonaut

shell shape. Four key species (identified from morphological

studies; see Finn, 2013, 2016) and shells at the centre of

taxonomic confusion for these species were targeted. These

target groups were the Argonauta nouryi!cornutus complex,

the A. hians/boettgeri complex, the A. nodosus/tuberculatus

complex and A. argo. This study supports parallel

morphological and molecular research recognising the

existence of only four argonaut species worldwide: A. argo

Linnaeus, 1758; A. hians [Lightfoot], 1786; A. nodosus

[Lightfoot], 1786; and A. nouryi Lorois, 1852.

Materials and methods

More than 1500 argonaut shells were examined over the

course of this project. Most of the shells examined reside in

museum collections within Australia, South Africa, Europe

and the United States. Institutions visited include: Australian

Museum, Sydney, Australia (AMS); Academy of Natural

Argonaut shell variability

Sciences, Philadelphia, USA (ANSP); The Natural History

Museum, London, UK (BMNH); Museum National d’Histoire

Naturelle, Paris, France (MNHN); Museums Victoria,

Melbourne, Australia (NMV); Queensland Museum, Brisbane,

Australia (QMB); South African Museum, Cape Town, South

Africa (SAM); South Australian Museum, Adelaide, Australia

(SAMA); Santa Barbara Museum of Natural History, Santa

Barbara, USA (SBMNH); Tasmanian Museum and Art

Gallery, Hobart, Australia (TMAG); National Museum of

Natural History, Smithsonian Institution, Washington, USA

(USNM); Western Australian Museum, Perth, Australia

(WAM). Material loaned from the Museum and Art Gallery of

the Northern Territory, Darwin, Australia (NTM) was

examined at NMV.

While all shells examined ultimately helped in the

formation of ideas and an understanding of shell shape

variation, two large collections were pivotal in enabling

argonaut shell variability to be interpreted.

The first lot, is a large collection of beach-cast argonaut

shells collected by Andres Gonzalez-Peralta ( Departamento

de Biologia Marina, Universidad AutUnoma de Baja

California Sur, MEXICO) on the beach at El Mogote, La Paz,

Baja California Sur, Mexico, North America, 24° 10' 00" N,

110° 24' 00" W, during the winters of 2000 and 2005. These

shells are lodged in the collection of SBMNH with the

following registration numbers: 172 Argonauta shells collected

on 15 January 2000 - SBMNH 345766 (93 shells), SBMNH

345767 (15 shells) and SBMNH 345768 (64 shells); 92

Argonauta shells collected on 31 January 2005 - SBMNH

357476 (77 shells) and SBMNH 357475 (15 shells).

The second lot was obtained by chance when researchers

on a research expedition off Rowley Shoals, Western Australia

left a pelagic trawl net in the water while steaming between

two stations: north-east of Mermaid Reef (Stn. 10, 17° 23' S,

118° 52' E) and south-west of Imperieuse Reef (Stn. 11,

16° 53' S, 119° 53' E). This occurred on board the FV

Courageous on 18-19 August 1983. On recovering the net,

researchers P. Berry and N. Sinclair were surprised to find 73

female argonauts with intact shells. Two specimens were

lodged in NMV while the remainder were retained by WAM.

The collection records of these lots are as follows: 71 female

argonauts - WAM S31520; 2 female argonauts - NMV F87104.

Shell terminology and measurements follow Finn (2013).

The opening of the shell is termed the aperture while the left

and right sides of the shell are termed lateral faces. An

extension of the axial thickening beyond the surface of the

lateral face of the shell is termed an ear. The lateral faces are

adorned with ribs radiating from the central axis of the shell

towards the keel. Ribs may be smooth (i.e. continuous) or

tuberculated (i.e. consisting of raised separate tubercles). The

keel is bordered by two opposing rows of keel tubercles. The

keel surface may be concave , straight or convex. The presence

of tubercles on the keel surface is known as inter-keel

tuberculation. To allow quantitative comparison of a large

number of shells, a set of standard measurements was taken.

These measurements included: shell length (ShL), maximum

length of shell (note that P indicates that the ShL measurement

was taken from a scaled digital photograph of the shell, not

directly from the shell); shell weight (ShW), weight (grams) of

dry shell; shell breadth (ShB), maximum breadth of the shell;

rib count (RC), number of ribs adorning a single lateral face,

counted around the keel and aperture edge; ear width (EW),

external measurement between lateral tips of opposing ears;

aperture length (ApL), internal distance from the axial

thickening to the ventral keel surface; aperture width (ApW),

internal measurement between two opposing lateral walls at

widest point; keel width (KW), external measurement of keel

at ventral most position; keel tubercle count (KTC), number of

keel tubercles counted around a single face (see fig. 2).

Scatter plots of measurements against ShL were used to

assess differences across large numbers of shells. Regression

lines were plotted using Microsoft Excel for Mac 2011.

Analysis of covariance (ANCOVA) was performed using

Systat 13.2 to assess the significance of the difference between

the slopes of the regression lines.

Scanning electron microscopy was used to examine shell

microstructure and allow accurate measurement of shell

thickness. Shell sections were placed in a sonicator bath for

short periods (5-10 seconds) to dislodge any debris, allowed to

air-dry, then placed onto double-sided carbon tabs (Ted Pella,

Redding) and sputter coated with gold. Scanning electron

micrographs were taken using a Zeiss EVO 40 XVP (Zeiss,

Cambridge) housed at SBMNH.

Where female argonauts could be definitively linked with

shells, a set of soft body measurements was taken following Finn

(2013). These measurements included: dorsal mantle length

(DML), length from posterior tip of mantle to furrow between

mantle edge and base of first arms; mantle width (MW), lateral

width of mantle at widest point; head width (HW), lateral width

of head measured between the opposing eye surfaces; arm length

(AL), length of arm from the edge of the mouth to arm tip,

measured along the face of the arm using a piece of string (for

arms 2-4); funnel length (FL), distance from the anterior tip of

the funnel to the posterior medial margin. The relationship

between features of female argonauts and their shells were

examined using scatter plots and linear regression.

Results

Argonauta nouryi Lorois, 1852; the A. nouryilcornutus

complex

In spring each year, small argonauts wash up in large numbers

on beaches in the southern Gulf of California (Gonzales-

Peralta in Saul and Stadum 2005). These small argonauts are

regularly attributed to two species: A. nouryi Lorois, 1852 and

A. cornutus Conrad, 1854 1 (Garcia-Dominguez and Castro-

Aguirre 1991; Gonzales-Peralta 2006).

A. nouryi was described by Lorois in 1852. The

identification of this species resides solely in features of the

shell, which is described as elliptical with numerous fine

lateral ribs and weak keel tubercles. Fig. 3 incorporates a

1 A third large form also washes up on southern Gulf of California

beaches in spring and is regularly attributed to the species A. pacificus,

a synonym of A. argo\ see Finn (2013) for details.

J.K. Finn

tuberculation

Figure 2. Argonaut shell measurements and terminology, following Finn (2013): a, Argonauta nodosus aperture view (NMV F164695);

b, A. nodosus lateral view (NMV F164695); c, A. argo aperture view (WAM S31503); d, A. nouryi aperture view (SBMNH 345766, specimen

#074); e, A. nouryi aperture view (SBMNH 345768, specimen #109). Abbreviations: ApL = aperture length; ApW = aperture width; EW = ear

width; KW = keel width; ShB = shell breadth; ShL = shell length. Illustrations: R. Plant.

Argonaut shell variability

Figure 3. Comparison of a shell from the examined SBMNH lot with an illustration taken from the original description of Argonauta nouryi

Lorois, 1852: a, reproduction of the illustration from the original description of A. nouryi Lorois, 1852, plate 1, fig. 5; b, illustrations of a shell

matching the description of A. nouryi taken from the examined lot (shell #109, 66.5 mm shell length, SBMNH 345768). Illustration: R. Plant.

Scale bar = 1 cm.

J.K. Finn

Figure 4. Comparison of a shell from the examined SBMNH lot with the type specimen and illustrations taken from the original description of

Argonauta cornutus Conrad, 1854: a, reproduced illustration taken from the original description of A. cornutus Conrad, 1854, plate 34, fig. 2; b,

photographs of the type specimen illustrated in the original description (58.6 mm shell length, ANSP 63496; please note, the original description

illustrations mirror the characters of the shell, most likely due to the engraving and printing process of the era); c, illustrations of a shell matching the

description of A. cornutus taken from the examined lot (shell #74, 65.0 mm shell length, SBMNH 345766). Illustration: R. Plant. Scale bar = 1 cm.

Argonaut shell variability

reproduction of the illustration presented by Lorois, 1852

(plate 1, figure 5), and illustrations of a shell from the Gulf of

California that is consistent with the original description (shell

#109, SBMNH 345768). According to Keen (1971) “the ‘shell’

is more elliptical than that of A. cornutus, with only the early

part of the coil moderately well tinged with brown along the

wide and weak tuberculate keel. The surface is delicately

ribbed and has a finely granular texture” (p. 895). Voss (1971)

believed that “ Argonauta nouryi is a distinctive species [...].

The shells are longer than in any other species of Argonauta,

the ribs are more numerous, there are no distinct tubercles

marking the edges of the carinal area; the carina is wide, very

convex, and covered by numerous, small, blunt tubercles

formed by the crisscrossing of the ribs” (p. 32).

Argonauta cornutus was described by Conrad in 1854. The

identification of this species also resides solely in features of the

shell, which is described as having a broad keel, large keel

tubercles and large ears. Fig. 4 incorporates a reproduction of

the illustration presented by Conrad, 1854 (plate 34, figure 2),

photographs of the type specimen (ANSP 63496) and

illustrations of a shell from the Gulf of California that is

consistent with the original description (shell #74, SBMNH

345766). According to Keen (1971), “the surface of the

yellowish-white ‘shell’ is finely granular, the spines and part of

the spire dark brown, the keel relatively broad, and the two long

axial expansions suffused with purplish brown” (p. 894). Voss

(1971) summarised that “ Argonauta cornutus seems best

characterised by the few radial ribs, the presence of fine sharp

tubercles or papillae over the sides of the shell, the few rather

sharp, large carinal tubercles on each side, the convex carinal

surface, and the few, large, blunt tubercles on the carinal surface

between the two rows of carinal boundary tubercles” (p. 32).

The distributions of these two species are reported to

overlap, with A. cornutus known from the Gulf of California

to Panama and A. nouryi being widespread in the equatorial

Pacific, ranging from the west coast of Southern California to

Peru (Keen 1971).

A mixed lot

As described in the Materials and Methods section above, the

157 shells in the collection at SBMNH were collected on the

same beach in Baja California on the same day. These shells

had previously been identified as representing both A. cornutus

and A. nouryi and were registered accordingly: SBMNH

345766, Argonauta cornutus 93 shells; SBMNH 345768,

Argonauta nouryi, 64 shells.

Initial examination of the lots indicated that the shells had

been attributed to either A. cornutus or A. nouryi based on the

presence or absence of ears - a character historically attributed

to only A. cornutus. Further examination of the lot revealed that

separation of the shells into two distinct groups (i.e. either A.

cornutus or A. nouryi ) was not as straightforward as first thought.

While some shells within the lot displayed all the characters

associated with either A. cornutus or A. nouryi, the lot also

appeared to contain shells with combinations of the attributes of

the two shell types. To illustrate this variation, three shells of

similar size but varied appearance were selected. Fig. 5 presents

photographs of these three shells from multiple perspectives:

• Shell #74 (SBMNH 345766), cornutus-type voucher (fig.

5a, i-iv and fig. 4c). Shell morphometries: ShL 65.0; ShW

4.0; ShB 40.7; RC 45; EW 58.1; ApL 45.9; ApW 28.4; KW

15.6; KTC 27.

• Shell #42 (SBMNH 345766), intermediate voucher (fig.

5b, i-iv). Shell morphometries: ShL 61.2; ShW 3.1; ShB

36.4; RC 47; EW 36.1; ApL 43.1; ApW 30.9; KW 14.0;

KTC 32.

• Shell #109 (SBMNH 345768), nouryi-type voucher (fig.

5c, i-iv and fig. 3b). Shell morphometries: ShL 66.5; ShW

2.4; ShB 39.9; RC 54; EW (28.3); ApL 48.8; ApW 32.5;

KW 15.8; KTC 54.

While it would be straightforward to attribute shell #74 (fig.

5a) to A. cornutus Conrad, 1854, and shell #109 (fig. 5c) to A.

nouryi Lorois, 1852, placement of shell #42 (fig. 5b) presents

problems. While shell #42 possesses the aperture shape of A.

cornutus, it lacks its protruding ears. While shell #42 possesses

the keel tuberculation and reduced ventral keel tubercles of A.

nouryi, its dorsal keel tubercles are large and pronounced.

To determine whether there was a significant difference

between eared and earless shells within the lot, a quantitative

approach was undertaken. All intact shells within the lot were

individually measured and weighed. All shells were designated

as being either eared or earless based on the relative EW and

ApW measurements. Because EW is an external measurement

(i.e. measured across the extremities of the opposing ears) and

ApW is an internal measurement (i.e. measured between the

lateral walls of the shell), 1.0 mm was added to the ApW to

accommodate for the thickness of the lateral walls of the shell.

Shells were classified as follows:

• eared = EW > ApW + 1.0 mm (103 shells)

• earless = EW < Apw + 1.0 mm (35 shells).

Scatter plots were generated to compare eared and earless

shells for all measured characters. Characters of primary

interest were those previously reported to distinguish A.

cornutus and A. nouryi.

Shell shape. The most universally recognised character of A.

nouryi is reportedly the elliptical shape of the shell: “The

whorls increase in size very rapidly and the last is very

elongate. Viewed laterally it is much shallower than is usual in

the genus” (Robson, 1932, p. 198). The shells are said to be

“more elliptical than that of A. cornutus ” (Keen, 1971, p. 895)

and “longer than in any other species of Argonauta” (Voss,

1971a, p. 33).

To investigate variation in shell shape across the lot, ShB

was plotted against ShL (fig. 6). Probability plots indicate that

both ShB and ShL follow normal distributions. An ANCOVA

was used to determine if the slopes of the linear regression

lines, generated for eared and earless shells, were the same or

different. Shell type (eared or earless) was the independent

variable, ShB the dependent variable and ShL the covariate.

The ANCOVA revealed that the slopes of the regression lines

are not equal and hence a significant difference in shell shape

exists between eared and earless shells (F (1 136) = 5.58, p = 0.02).

J.K. Finn

Figure 5. Three similarly sized shells of varied appearance selected from the examined SBMNH lot: a-c, three similarly sized shells of varied

appearance selected from the single lot collected at El Mogote, La Paz, Baja California Sur, Mexico (24° 10' 00" N, 110° 24' 00" W) on 15 January

2000; a, shell #74 (65.0 mm shell length, SBMNH 345766) assigned the name cornutus-type voucher, b, shell #42 (61.2 mm shell length, SBMNH

345766) assigned the name intermediate voucher, c, shell #109 (66.5 mm shell length, SBMNH 345768) assigned the name nouryi-type voucher,

i-iv, multiple perspectives of each shell; i, left lateral view; ii, anterior aperture view; iii, posterior keel view; iv, ventral view. Scale bar = 1 cm.

Argonaut shell variability

Rib count. Argonauta nouryi shells are reported to have more

ribs than A. cornutus shells: the ribs in A. nouryi are “more

numerous” than in other species of Argonauta, while A. cornutus

is reported to have “few radial ribs” (Voss, 1971, p. 32-33).

To investigate variation in the number of ribs per shell

across the lot, RC was plotted against ShL (fig. 7). Probability

plots indicate that both RC and ShL follow normal distributions.

An ANCOVA was used to determine if the slopes of the linear

Figure 6. Variation in shell shape across the examined SBMNH lot.

Scatter plot of shell breadth (ShB) against shell length (ShL) for the

single shell lot collected at El Mogote, La Paz, Baja California Sur,

Mexico (24° 10' 00" N, 110° 24’ 00" W) on 15 January 2000 (SBMNH

345766 & 345768). Eared shells (solid circles) and earless shells (open

circles) distinguished. Linear regression lines for eared shells (dashed)

and earless shells (dot dashed) with corresponding equations and

coefficients of determination (i.e. R 2 values) presented.

regression lines, generated for eared and earless shells, were

the same or different. Shell type (eared or earless) was the

independent variable, RC the dependent variable and ShL the

covariate. The ANCOVA revealed that the slopes of the

regression lines are not equal and hence a significant difference

in the number of ribs per shell does exist between eared and

earless shells (F (1 m) = 21.2, p < 0.001).

Other features. To investigate the full range of quantifiable

shell characters across the lot, scatter plots were similarly

generated to investigate KTC, ApL, ApW and KW.

Keel tubercle count. To investigate variation in the number of

keel tubercles per shell across the lot, KTC was plotted against

ShL (fig. 8). Probability plots indicate that both KTC and ShL

follow normal distributions. An ANCOVA was used to determine

if the slopes of the linear regression lines, generated for eared

and earless shells, were the same or different. Shell type (eared

or earless) was the independent variable, KTC the dependent

variable and ShL the covariate. The ANCOVA revealed that the

slopes of the regression lines are not equal and hence a significant

difference in the number of keel tubercles per shell does exist

between eared and earless shells (F (1 136) = 51.66, p < 0.001).

Aperture length. To investigate variation in the length of the

shell apertures across the lot, ApL was plotted against ShL (fig.

9). Probability plots indicate that both ApL and ShL follow

normal distributions. An ANCOVA was used to determine if

the slopes of the linear regression lines, generated for eared and

earless shells, were the same or different. Shell type (eared or

earless) was the independent variable, ApL the dependent

variable and ShL the covariate. The ANCOVA revealed that

the slopes of the regression lines are not equal and hence a

significant difference in the length of the aperture does exist

between eared and earless shells (F (1 136) = 18.63, p < 0.001).

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Shell Length [ShL] (mm)

Figure 7. Variation in rib number across the examined SBMNH lot.

Scatter plot of rib count (RC) against shell length (ShL) for the single

shell lot collected at El Mogote, La Paz, Baja California Sur, Mexico

(24° 10' 00" N, 110° 24’ 00" W) on 15 January 2000 (SBMNH

3045766 & 345768). Eared shells (solid circles) and earless shells

(open circles) distinguished. Linear regression lines for eared shells

(dashed) and earless shells (dot dashed) with corresponding equations

and coefficients of determination (i.e. R 2 values) presented.

Figure 8. Variation in keel tubercle number across the examined

SBMNH lot. Scatter plot of keel tubercle count (KTC) against shell

length (ShL) for the single shell lot collected at El Mogote, La Paz, Baja

California Sur, Mexico (24° 10' 00" N, 110° 24’ 00" W) on 15 January

2000 (SBMNH 345766 & 345768). Eared shells (solid circles) and

earless shells (open circles) distinguished. Linear regression lines for

eared shells (dashed) and earless shells (dot dashed) with corresponding

equations and coefficients of determination (i.e. R 2 values) presented.

J.K. Finn

Aperture width. To investigate variation in the width of the

shell apertures across the lot, ApW was plotted against ShL

(fig. 10). Probability plots indicate that both ApW and ShL

follow normal distributions. An ANCOVA was used to

determine if the slopes of the linear regression lines, generated

for eared and earless shells, were the same or different. Shell

type (eared or earless) was the independent variable, ApW the

Shell Length [ShL] (mm)

Figure 9. Variation in aperture length across the examined SBMNH

lot. Scatter plot of aperture length (ApL) against shell length (ShL) for

the single shell lot collected at El Mogote, La Paz, Baja California Sur,

Mexico (24° 10' 00" N, 110° 24' 00" W) on 15 January 2000 (SBMNH

345766 & 345768). Eared shells (solid circles) and earless shells (open

circles) distinguished. Linear regression lines for eared shells (dashed)

and earless shells (dot dashed) with corresponding equations and

coefficients of determination (i.e. R 2 values) presented.

Figure 10. Variation in aperture width across the examined SBMNH

lot. Scatter plots of aperture width (ApW) against shell length (ShL)

for the single shell lot collected at El Mogote, La Paz, Baja California

Sur, Mexico (24° 10' 00" N, 110° 24' 00" W) on 15 January 2000

(SBMNH 345766 & 345768). Eared shells (solid circles) and earless

shells (open circles) distinguished. Linear regression lines for eared

shells (dashed) and earless shells (dot dashed) with corresponding

equations and coefficients of determination (i.e. R 2 values) presented.

dependent variable and ShL the covariate. The ANCOVA

revealed that the slopes of the regression lines are not equal and

hence a significant difference in the width of the aperture does

exist between eared and earless shells (F (1 136) = 4.07, p = 0.046).

Keel width. To investigate variation in the width of the shell

keels across the lot, KW was plotted against ShL (fig. 11).

Probability plots indicate that both KW and ShL follow normal

distributions. An ANCOVA was used to determine if the slopes

of the linear regression lines, generated for eared and earless

shells, were the same or different. Shell type (eared or earless)

was the independent variable, KW the dependent variable and

ShL the covariate. The ANCOVA revealed that the slopes of

the regression lines are equal and hence a significant difference

in the width of the keel does not exist between eared and earless

shells (F (1 m = 0.87, p = 0.353).

Statistical analysis indicates that significance differences

in shell dimensions was associated with the presence or

absence of ears. Eared shells have significantly lower RC (p <

0.001), lower KTC (p < 0.001), shorter ApL (p < 0.001),

increased ShB (i.e. shortened; p = 0.02) and increased ApW (p

= 0.046). Earless shells have significantly higher RC (p <

0.001), higher KTC (p < 0.001), longer ApL (p < 0.001),

reduced ShB (i.e. elongate; p = 0.02) and reduced ApW (p =

0.046). KW was found to not be significantly different between

shell types (p = 0.353).

Historically, the features of eared and earless shell types

have been considered to represent separate species such that

features of eared shells are considered characteristic of A.

cornutus, while features of earless shells are considered

characteristic of A. nouryi.

Figure 11. Variation in keel width across the examined SBMNH lot.

Scatter plot of keel width (KW) against shell length (ShL) for the

single shell lot collected at El Mogote, La Paz, Baja California Sur,

Mexico (24° 10' 00" N, 110° 24’ 00" W) on 15 January 2000 (SBMNH

345766 & 345768). Eared shells (solid circles) and earless shells (open

circles) distinguished. Linear regression lines for eared shells (dashed)

and earless shells (dot dashed) with corresponding equations and

coefficients of determination (i.e. R 2 values) presented.

Argonaut shell variability

Two types of shell formation

Close examination of individual shells revealed that features

considered characteristic of each shell type could occur on a

single shell. While individual shells could display features of

both eared and earless shell types, the characters did not

appear in isolation. Sequential growth sections of the shells

appeared to display all the characteristics of one shell type or

another. For example, the initial component of the shell (the

smallest whorl) may display all the characters historically

associated with an A. cornutus shell while the latter component

(the larger final whorl) may display all the features associated

with an A. nouryi shell.

The most dramatic examples were shells that appeared to

have been repaired over the course of the argonaut’s life. Fig.

12 presents photographs of one such shell from lot SBMNH

357476 (52.3 mm ShL). The initial component of the shell

clearly displays the features historically attributed to A. nouryi

(numerous fine ribs, reduced keel tubercles and no apparent

ears), while the later component, following the clear repair

line, displays a transition to features historically attributed to

A. cornutus (ribs reduced in number and more pronounced,

keel tubercles reduced in number and of larger size, and

initiation of ears).

The presence of both shell types on a single shell clearly

demonstrates that they represent different types of shell

formation, not different argonaut species. This observation is

supported by morphological evidence; despite full examination

of nine female argonauts with shells (six historically identified

as A. cornutus and three A. nouryi ), no morphological

characters could be found to separate specimens with different

shell types (see Finn, 2013).

The realisation that the two shell morphs represented two

shell formation types, not two argonaut species, required that

they be defined independent of previous species association:

• Type 1 shell formation (historically attributed to A.

cornutus shells) - formation of ears, few pronounced ribs,

few large keel tubercles, appearance of more pronounced

arch in the keel resulting in a tighter final whorl (i.e.

increased ShB, reduced ApL).

• Type 2 shell formation (historically attributed to A. nouryi

shells) - absence of ears, numerous less pronounced ribs,

numerous small keel tubercles, appearance of less

pronounced arch in the keel resulting in the appearance of

a shallower final whorl and elliptical shell (i.e. reduced

ShB, increased ApL).

An important character associated with Type 2 shell

formation is inter-keel tuberculation (tubercles on the keel

surface; see fig. 2e). The appearance of inter-keel

tuberculation on the keel of a shell flags a shift to Type 2

shell formation, while a loss of inter-keel tuberculation

signifies a shift to Type 1 shell formation.

Based on this realisation, it became clear that this large lot,

and all other material examined of these shell morphs, belonged

to a single species. Because A. nouryi Lorois, 1852, has date

priority over A. cornutus Conrad, 1854; this study treats A.

nouryi as the available name. See Finn (2013) for full synonymy.

The key to understanding shell variation

The realisation that individual shells may be composed of

combinations of two types of shell formation provided the key

to understanding the huge variation in shell shape across the

single large collection of argonaut shells from Baja California.

Combinations of sequential shell formation could be

recognised in all shells and hence their varied appearance

could be understood. Shells were recognised within this single

lot that display a single type of shell formation plus those with

one, two or three transformations between the two shell

formation types.

The initial whorl of most of the shells displayed Type 1

formation. Shell #37 displays a single change from Type 1 to

Type 2 shell formation (fig. 13). Shell #72 displays a change

from Type 1 to Type 2 shell formation and then a change back

to Type 1 (fig. 14). Shell #41 displays a change from Type 1 to

Type 2 shell formation and then a change back to Type 1 and

then to Type 2 (fig. 15). Damage to shells normally results in a

conversion to Type 2 shell formation.

In a transition between shell formation types, ears may be

formed or subsumed. This is displayed across many shells

within the lot. For examples, shell #139 displays subsumed

ears as a result of a transition from Type 1 to Type 2 shell

formation (fig. 16), while shell #136 displays ear formation,

separate from the axis of the shell, as a result of a transition

from Type 2 to Type 1 shell formation (fig. 17).

Type material. Available type material for additional species

synonymised with A. nouryi Lorois, 1852, was also examined

for shifts in shell formation type. The holotype of A. dispar

Conrad, 1854 (54.9 mm ShL, ANSP 129978) displays a single

change from Type 2 to Type 1 shell formation (fig. 18). The

holotype of A. expansus Dali, 1872 (80.2 mm ShL [P], USNM

61368), displays two changes - from Type 1 to Type 2 and then

back to Type 1 (fig. 19).

Shell thickness. Preliminary observations suggested that the

shell walls of Type 1 formation are thicker than the walls of

Type 2 formation. To investigate this phenomenon, a scanning

electron microscope was used to examine variation in shell

thickness across recognisable shell breaks that corresponded

with a switch between shell types (a single damaged shell from

lot SBMNH 357476 was sacrificed). Preliminary results

indicate a reduction in shell wall thickness between Type 1 and

Type 2 formation. Fig. 20 presents two scanning electron

micrographs displaying a reduction in thickness across a break

signifying transition from Type 1 to Type 2. Shell thickness on

the lateral face drops from approximately 220 to 140 pm (fig.

20a), while thickness at the keel drops from approximately 275

to 210 pm in this shell (fig. 20b).

A lack of material that could be fragmented for examination

with a scanning electron microscope limited the extent to

which this phenomenon could be investigated. The lots housed

in the SBMNH collection are too valuable to be considered for

this style of destructive investigation.

A reduction in shell wall thickness may be related to

producing a larger shell area with less shell material. The

resulting thinner walled shell (Type 2) would therefore consist

J.K. Finn

Figure 12. Repaired shell displaying components consistent with Argonauta nouryi and A. cornutus: a-d, four perspectives of a single shell (52.3

mm shell length, SBMNH 357476) displaying an initial component consistent with A. nouryi Lorois, 1854 (“nouryi”) followed by a subsequent

component consistent with A. cornutus Conrad, 1854 (“cornutus”); a, right lateral view; b, oblique right lateral view; c, anterior aperture view;

d, oblique ventral keel view. Dashed line represents repair line separating two visually different components. Scale bar = 1 cm.

Argonaut shell variability

Figure 13. Argonauta nouryi shell displaying a single change in shell formation type: a-d, four perspectives of shell #37 (65.5 mm shell length,

SBMNH 345766) displaying a single change from Type 1 (Tl) to Type 2 (T2) shell formation; a, left lateral view; b, oblique left lateral view; c,

close-up oblique left lateral view; d, posterior keel view. Scale bar = 1 cm.

J.K. Finn

Figure 14. Argonauta nouryi shell displaying two changes in shell formation type: a-d, four perspectives of shell #72 (55.4 mm shell length,

SBMNH 345766) displaying two changes from Type 1 (Tl) to Type 2 (T2) shell formation and back to Type 1; a, right lateral view; b, left lateral

view; c, oblique left lateral view; d, close-up oblique left lateral view. Scale bar = 1 cm.

Argonaut shell variability

of less calcium carbonate and weigh less than an equivalently

sized thicker walled shell (Type 1). The relative weights of the

three shells presented in fig. 5 appear to support this theory.

The Type 1 shell ( cornutus-type voucher; 4.0 g) is 1.3 times

the weight of the Type 1/Type 2 shell ( intermediate voucher ;

3.1 g) and 1.7 times the weight of the Type 2 shell ( nouryi-type

voucher, 2.4 g), despite the shells having similar ShL. Weight

(g) to length (mm) ratios of the three shells were: 1:16 for the

Type 1 shell ( cornutus-type voucher); 1:20 for the Type 1/Type

2 shell ( intermediate voucher)-, 1:28 for the Type 2 shell

{nouryi-type voucher ). These ratios suggest that per gram of

calcium carbonate, Type 2 shell production results in a shell

1.8 times the length of a Type 1 shell.

To investigate this relationship across the lot, ShW was

plotted against ShL with eared and earless shells distinguished

(fig. 21). The scatter plot indicates a separation between eared

and earless shells based on weight. This difference was

analysed statistically to determine significance. Probability

plots indicate that both ShW and ShL follow normal

distributions. An ANCOVA was used to determine if the

slopes of the regression lines, generated for eared and earless

shells, were the same or different. Shell type (eared or earless)

was the independent variable, ShW the dependent variable and

ShL the covariate. The ANCOVA revealed that the slopes of

the regression lines are not equal and hence a significant

difference in weight exists between eared and earless shells

(F ( , 136) = 86.7, p< 0.001).

Argonauta hians [Lightfoot], 1786; the A. hianslboettgeri

complex

Recognition of shell form transformations in A. nouryi

provided a new perspective on shell variation in another highly

variable group of small argonauts, the A. hianslboettgeri

complex.

The original description of A. hians [Lightfoot], 1786,

refers to a single image in Rumphius (1705): plate 18, figure B

(fig. 22a), designated as a lectotype by Moolenbeek (2008) in

the absence of type material. Shells of A. hians can be

recognised by smooth lateral ribs and a keel that increases in

width with shell growth. Inter-keel tuberculation is absent.

Figure 15. Argonauta nouryi shell displaying three changes in shell formation type: a-b, two perspectives of shell #41 (64.7 mm shell length,

SBMNH 345766) displaying three changes in shell formation type from Type 1 (Tl) to Type 2 (T2) shell formation, back to Type 1 and then to

Type 2; a, left lateral view; b, oblique left lateral view. Note that the key to recognising the different shell formation types (challenging in this shell)

is to look for reductions in the size of sequential keel tubercles (that would normally increase in size), a change in the relative distance between keel

tubercles, a change in the ratio of lateral ribs to keel tubercles, and the appearance or disappearance of inter-keel tuberculation. Scale bar = 1 cm.

J.K. Finn

Figure 16. Argonauta nouryi shell displaying subsumed ears: a-d, four perspectives of shell #139 (72.2 mm shell length, SBMNH 345768)

displaying subsumed ear (E) associated with a shift from Type 1 (Tl) to Type 2 (T2) shell formation; a, left lateral view; b, close-up of subsumed

ear, left lateral view; c, close-up of subsumed ear, oblique left lateral view; d, anterior aperture view. The shell added to the aperture edge in Type

2 shell formation does not expand the ear, instead subsuming it. The resulting aperture edge is not eared. Scale bars = 1 cm.

Argonaut shell variability

Figure 17. Argonauta nouryi shell displaying ear formation: a-c, three perspectives of shell #136 (63.1 mm shell length, SBMNH 345768)

displaying ear (E) formation associated with a shift from Type 2 (T2) to Type 1 (Tl) shell formation; a, left lateral view; b, close-up of ear, left

lateral view; c, anterior aperture view. The shell added to the aperture edge in Type 1 shell formation produces a new ear separate from the axis

of the shell. The new ear becomes the widest point on the aperture edge. Scale bars = 1 cm.

J.K. Finn

Figure 18. Holotype of Argonauta dispar Conrad, 1854 (synonym of A. nouryi Lorois, 1852) from the Academy of Natural Sciences, Philadelphia:

a-d, four perspectives of A. dispar Conrad, 1854 Holotype (54.9 mm shell length, ANSP 129978) displaying a single change from Type 2 (T2)

to Type 1 (Tl) shell formation; a, left lateral view; b, right lateral view; c, anterior aperture view; d, posterior keel view. Scale bar = 1 cm.

Argonaut shell variability

Figure 19. Holotype of Argonauta expansus Dali, 1872 (synonym of A. nouryi Lorois, 1852) from the National Museum of Natural History

(Smithsonian Institution) Washington: a-c, three perspectives of A. expansus Dali, 1872 Holotype (80.2 mm shell length [P], USNM 61368)

displaying two changes from Type 1 (Tl) to Type 2 (T2) shell formation and back to Type 1; a, left lateral view; b, anterior aperture view; c,

posterior keel view. Scale bar = 1 cm.

J.K. Finn

Figure 20. Scanning electron microscope images of Argonauta nouryi shell displaying variation in shell thickness: a-b, scanning electron

microscope images of shell cross-sections (SBMNH 357476) across shell repairs (R) representing a shift from Type 1 (Tl) to Type 2 (T2) shell

formation; a, lateral face of shell, inner surface facing up; b, keel, outer surface facing up. Scale bars = 1 mm.

Argonaut shell variability

Figure 21. Variation in shell weight across the examined SBMNH lot.

Scatter plot of shell weight (ShW) against shell length (ShL) for the

single shell lot collected at El Mogote, La Paz, Baja California Sur,

Mexico (24° 10' 00" N, 110° 24' 00" W) on 15 January 2000 (SBMNH

345766 & 345768). Eared shells (solid circles) and earless shells (open

circles) distinguished. Linear regression lines for eared shells (dashed)

and earless shells (dot dashed) with corresponding equations and

coefficients of determination (i.e. R 2 values) presented.

Argonauta hians has long been recognised as displaying

considerable variation in shell form. Voss and Williamson

(1971) noted that “In the series from Hong Kong the sides of

the aperture at the umbilicus range from strongly eared or

auriculate with very large few knobs on the keel to specimens

with no trace of auriculation and with rather more numerous,

smaller knobs” (p. 105). They found that “if the 30 shells are

laid out graded from large few knobs and strong auricles to

smaller, more numerous knobs and flat sides there is an even

gradation with no breaks or sudden changes” (p. 105). They

concluded that all shells “belong to the same species” (p. 105).

As part of this study, 274 A. hians shells were directly

examined in museum and private collections in Australia,

United States, Europe, South Africa and Japan. With

knowledge gained from examining shells of A. nouryi, all

shells from all sites were examined for an abrupt change in

keel tubercle height or ears that had been formed or subsumed

in single shells. Because inter-keel tuberculation is not

expressed in argonaut shells other than A. nouryi, this

character could not be used.

Two shell formation types. Shells of A. hians were found to

display two clear shell formation types:

• Type 1 shell formation - few pronounced ribs, large

prominent keel tubercles, formation of ears.

• Type 2 shell formation - numerous less-pronounced ribs,

small and greatly reduced, keel tubercles, absence of ears.

These shell formation types are similar to those expressed in

A. nouryi except that variation in the arch of the shell was not

observed and inter-keel tuberculation was not present.

This variation had been noted by Voss and Williamson

(1971) who stated: “The knobs on the keel are very large and

prominent in the first half of the shell and may remain large on

the last half or may become considerably smaller” (p. 105).

Two shells are presented as examples:

• A shell from the Philippines (79.6 mm ShL [P], BMNH

unreg., “Cuming, i.”) (fig. 23). This shell displays a clear

shift from Type 1 to Type 2 shell formation indicated by a

reduction in the size and spacing of the keel tubercles, a

reduction in the ratio of ribs to keel tubercles from

approximately 1.5:1 to 1:1 and ears subsumed.

• A shell from the North West Shelf, Western Australia (53.0

mm ShL, WAM S31510) (fig. 24). This shell displays a

shift from Type 1 to Type 2 shell formation. This transition

occurred when the shell was at a smaller size and hence

the ears are less developed. The resultant aperture shape

(fig. 24c) is extremely similar to that observed in Type 2 A.

nouryi shells; see fig. 5c, ii for comparison.

Variation also occurs between the opposing faces of individual

shells, further highlighting the plasticity of shell characters in

this species. A single shell is presented here as an example:

• A shell from Madagascar (60.8 mm ShL, NMV F164734,

“Madagascar”) displays a large ear on the right side only;

the left side is earless (fig. 25).

The A. hians/boettgeri complex. Small, earless A. hians shells

have regularly been attributed to the species A. boettgeri

Maltzan, 1881 (fig. 22b, c). Smith (1887) outlined the diagnostic

characters of A. boettgeri: “The distinguishing features of this

species are the numerous ribs and tubercles, the total absence of

auricular expansions at the sides, its constantly small size, and

the fine granulation (a feature not remarked upon by Maltzan),

which more or less covers the whole surface, producing a dull

non-glossy appearance” (p. 409). Berry (1914) similarly noted

that the shell of A. boettgeri “seems unique in its small size,

compact coil, and the circumstance that the auricular expansion

at the sides of the aperture, so frequently developed in other

species of the genus, are here notable only for their entire

absence” (p. 280). Robson (1932) added “the almost invariable

absence of colouring on the carinal knobs” to the distinguishing

characters of A. boettgeri (p. 197). While Smith (1887) concludes

that “the shell of this species must not be confounded with

young stages of A. hians\ the more numerous ribs and tubercles

and the rougher granular surface will separate it” (p. 410).

Unfortunately, this dichotomy is not so straightforward.

Of the 274 A. hians shells examined, 41 can be attributed

to A. boettgeri based on the above description. While it is

possible to select a subset of shells possessing these

characteristics, which in isolation appear distinct, examination

of the entire range of material quickly dissolves the parameters

on which this subset is based. All features mentioned above

are variable in A. hians: ribs and keel tubercles can be

numerous or scarce, pronounced or reduced, consistent across

the shell or variable; ears can be present or absent, produced or

subsumed, expressed on one side of the shell or both; the shell

surface can be granular or smooth, pigmented or white. Two

shells, displaying variation across the growth of the shell, are

presented as examples:

J.K. Finn

Figure 22. Reproduced illustrations referenced in the descriptions of Argonauta hians [Lightfoot], 1786 and A. boettgeri Maltzan, 1882: a,

illustration of A. hians [Lightfoot], 1786, designated as alectotype by Moolenbeek (2008), Rumphius, 1705: pi. 18, fig. B; b-c, illustrations of A.

boettgeri Maltzan, 1881, featured in the original publication, Maltzan, 1881: 163, pi. 6 fig. 7; b, right lateral view; c. anterior aperture view.

Argonaut shell variability

Figure 23. Argonauta hians shell from the Philippines: a-d, four perspectives of an A. hians shell from the Philippines (79.6 mm shell length [P],

BMNH unreg., “Cuming, i.”) displaying a clear shift from Type 1 shell formation (Tl) to Type 2 shell formation (T2) indicated by a reduction in

the size and spacing of the keel tubercles, a reduction in the ratio of ribs to keel tubercles (from approximately 1.5:1 to 1:1) and subsumed ears;

a, right lateral view; b, anterior aperture view; c, posterior keel view; d, ventro-posterior keel view. Scale bar = 1 cm.

J.K. Finn

Figure 24. Argonauta hians shell from the North West Shelf, Western Australia: a-d, four perspectives of an A. hians shell from the North West

Shelf, Western Australia (53.0 mm shell length, WAM S31510) displaying a clear shift from Type 1 shell formation (Tl) to Type 2 formation (T2)

indicated by a reduction in the size and spacing of the keel tubercles and a reduction in the ratio of ribs to keel tubercles (from approximately

1.5:1 to 1:1); a, right lateral view; b, oblique right lateral view; c, anterior aperture view; d, posterior keel view. Scale bar = 1 cm.

Argonaut shell variability

Figure 25. Single eared Argonauta hians shell from Madagascar: a-c, three perspectives of a single eared A. hians shell from Madagascar (60.8

mm shell length, NMV F164734); a, left lateral view; b, right lateral view; c, anterior aperture view. Scale bar = 1 cm.

J.K. Finn

• A shell from the British Museum (76.1 mm ShL [P],

BMNH unreg., locality unknown, “B698, t”; fig. 26).

This shell displays an aperture shape and axial region

consistent with the original description of A. boettgeri

(fig. 22b, c) yet defies the description of A. boettgeri by

showing signs of possessing ears at an earlier growth

stage. While the ears have been subsumed with a shift

from Type 1 to Type 2 shell formation, only the keel

tubercles on the right side show a reduction in size (fig.

26b); the left keel tubercles have remained large (fig. 26a).

• A shell from Museums Victoria (25.0 mm ShL [P],

NMV F164767, locality unknown; fig. 27). This shell

would historically have been attributed to A. boettgeri

due to its small size and distinctive earless aperture.

This shell displays a dramatic change in keel tubercle

size and spacing associated with a shift from Type 2 to

Type 1 shell formation, thus highlighting the plasticity

of these characters.

In the absence of any consistent and definable diagnostic

shell characters (in combination with a lack of diagnostic

morphological characters or distinct distributions; see Finn,

2013), no evidence exists to justify maintaining A. boettgeri as

a separate species. Consequently A. boettgeri Maltzan, 1881,

is treated here as a synonym of A. hians [Lightfoot], 1786.

Insight from whole animals. As described in the Materials and

Methods section above, a single specimen lot of 73 female A.

hians, most with intact shells, exist in the collections of the

Western Australian Museum and Museums Victoria. On initial

examination, it was found that the lot included submature,

mature and spawned (i.e. females with eggs attached to the

central axis of the shell) individuals. The shells of the spawned

females tended to show a shift to Type 2 shell production in the

last components of the shells (all other shells were composed

entirely of Type 1 shell production). This led to the consideration

that shell shape and transformation may be triggered by

changes in reproductive stage or condition.

To understand the underlying cause of a change in shell

formation type at the point of egg laying, a subset of 33 intact

and measurable individuals were selected and fully measured.

The subset included submature, mature and spawned

individuals, with a size range of 13-27 mm DML and 21-36

mm ShL. Two larger females, also collected over the North

West Shelf, were incorporated into the analysis to expand the

size range (QM Mo77789: 39.9 mm DML and 51.8 mm ShL;

28.7 mm DML and 38.9 mm ShL).

Changes in shell morphometries relative to animal size. Shell

dimensions were plotted against DML to determine if the size

of the shell relative to the size of the female changes between

submature, mature and spawned individuals. Scatterplots

against DML were generated for ShL, ShB, ApL, ApW, KW

and EW. The scatter plots indicate a linear relationship between

shell dimensions and animal size, with linear regressions

returning coefficient of determination values (i.e. R 2 values)

between 0.72 and 0.90 (see Table 1). No discontinuities were

observed between the three maturity stages.

Table 1. Linear regression equations for scatter plots of shell dimensions (y) against dorsal mantle length (v) for 35 female Argonauta hians from

Australian waters (WAM S31520/NMV 87104/QM Mo77789) including submature, mature and spawned individuals. Corresponding coefficients

of determination (i.e. R 2 values) presented.

equation

R 2

Shell length (ShL)

Dorsal mantle length (DML)

y = 1.0936x + 7.9242

0.8980

Shell breadth (ShB)

Dorsal mantle length (DML)

y = 0.8705x + 0.6315

0.8705

Aperture length (ApL)

Dorsal mantle length (DML)

y = 0.8889x + 3.4049

0.8882

Aperture width (ApW)

Dorsal mantle length (DML)

y = 0.3875x + 9.3432

0.7697

Keel width (KW)

Dorsal mantle length (DML)

y = 0.1839x + 4.1154

0.7244

Ear width (EW)

Dorsal mantle length (DML)

y = 0.3360x + 11.1327

0.7268

Table 2. Linear regression equations for scatter plots of female argonaut dimensions (y) against dorsal mantle length (v) for 35 female Argonauta

hians from Australian waters (WAM S31520/NMV 87104/QM Mo77789) including submature, mature and spawned individuals. Corresponding

coefficients of determination (i.e. R 2 values) presented.

equation

R 2

Mantle width (MW)

Dorsal mantle length (DML)

y = 0.4166x + 6.6124

0.8156

Head width (HW)

Dorsal mantle length (DML)

y = 0.4594x + 4.5374

0.8609

Funnel length (FL)

Dorsal mantle length (DML)

y = 0.4905x + 3.2751

0.7559

Arm length 2 (AL2)

Dorsal mantle length (DML)

y = 1.8016x - 1.5714

0.8773

Arm length 3 (AL3)

Dorsal mantle length (DML)

y = 1.1550x + 5.5401

0.8169

Arm length 4 (AL4)

Dorsal mantle length (DML)

y = 0.8482x + 5.8121

0.8292

Argonaut shell variability

Figure 26. Argonauta hians shell from the British Museum: a-d, four perspectives of an A. hians shell from the British Museum (76.1 mm shell

length [P], BMNH unreg., locality unknown, “B698, t.”) which, while displaying an aperture shape and axial region consistent with the original

description of A. boettgeri (fig. 22B, C), shows signs of possessing ears (E) at an earlier stage of growth; a, left lateral view; b, right lateral view;

c, anterior aperture view; d, posterior keel view. A shift from Type 1 shell formation (Tl) to Type 2 shell formation (T2) is expressed by ears

subsumed and a reduction in keel tubercle size on the right side only. Scale bar = 1 cm.

J.K. Finn

Figure 27. Shell consistent with description of Argonauta boettgeri from Museums Victoria: a-d, four perspectives of a shell consistent with A.

boettgeri (treated here as a synonym of A. hians [Lightfoot], 1786) from Museums Victoria (25.0 mm shell length, NMV F164767) displaying an

increase in keel tubercle size consistent with a shift from Type 2 shell formation (T2) to Type 1 shell formation (Tl); a, left lateral view; b, right

lateral view; c, anterior aperture view; d, posterior keel view. Scale bar = 1 cm.

Argonaut shell variability

Table 3. Linear regression equations for scatter plots of shell dimensions (y) against shell length (x) for 35 female Argonauta hians from Australian

waters (WAM S31520/NMV 87104/QM Mo77789) including submature, mature and spawned individuals. Corresponding coefficients of

determination (i.e. R 2 values) presented.

equation

R 2

Shell breadth (ShB)

Shell length (ShL)

y = 0.7676x - 4.8377

0.9015

Aperture length (ApL)

Shell length (ShL)

y = 0.8013x - 2.6968

0.9613

Aperture width (ApW)

Shell length (ShL)

y = 0.3369x + 7.0488

0.7752

Keel width (KW)

Shell length (ShL)

y = 0.1701x + 2.7255

0.8254

Ear width (EW)

Shell length (ShL)

y = 0.2936x + 9.1011

0.7391

Ontogenetic changes in animal morphology. Dimensions and

characters of the female argonauts were plotted against DML to

determine if the relative proportions of the female changes

between submature, mature and spawned individuals.

Scatterplots against DML were generated for MW, HW, FL and

AL. The scatter plots indicate a linear relationship between

animal dimensions, with linear regressions returning coefficient

of determination values (i.e. R 2 values) between 0.76 and 0.88

(see Table 2). No discontinuities were observed between the

three maturity stages.

Ontogenetic changes in shell morphometries. Shell dimensions

and characters were plotted against ShL to determine if relative

shell proportions change between submature, mature and

spawned females. Scatterplots against ShL were generated for

ShB, ApL, ApW, KW and EW. The scatter plots indicate a linear

relationship between shell dimensions and characters, with

linear regressions returning coefficient of determination values

(i.e. R 2 values) between 0.74 and 0.96 (see Table 3). No

discontinuities were observed between the three maturity stages.

The scatter plots provided no evidence of a change in

relative shell and animal proportions between submature,

mature and spawned individuals. If the examined characters

underwent dramatic transformation with changes in state of

maturity, it was expected that discontinuities would be

observed in the plotted data. It is apparent that the visual

change in shell form observed across this lot was not reflected

in the relative measurements of the individuals measured.

Argonauta nodosus [Lightfoot], 1786; the A. nodosus/

tuberculatus complex

The original description of A. nodosus [Lightfoot], 1786, refers

to a single image in Rumphius (1705): plate 18, figure 1 (fig.

28a), designated as a lectotype by Moolenbeek (2008) in the

absence of type material. Shells of A. nodosus can be recognised

by the presence of lateral ribs composed of separate tubercles.

Two types of A. nodosus shells exist in collections: a finer shell

with more ribs and small rib tuberculations (fig. 29a), and a coarser

shell with fewer ribs and larger rib tuberculations (fig. 29b).

This variation has previously been used as justification for

splitting A. nodosus into two species. Kirk (1885), in recognising

the two forms, generated a new species name for the fine

tuberculated and earless form (A. gracilis) to separate it from the

coarse tuberculated and eared form (known to Kirk, 1885, as A.

tuberculata Shaw). Robson (1932) recognised the two shell types

as varieties, not separate species, stating: “Though the shell of

this species is clearly distinguished from its fellows by the rough

tuberculations, there are evidently two well marked varieties -

one with very large carinal knobs and coarse sculpture, the other

with low knobs and fine sculpture” (p. 200). Dell (1952) called

this the “nodosa-tuberculata complex” 2 and described it as

follows: “Group 1. The shell is eared laterally and the

tuberculations on the ribs are comparatively large - this is what

has been called nodosa. Group 2. The edge of the lip comes off

the previous whorl in an even curve without trace of an ‘ear’. The

tuberculations are much finer and more numerous than in Group

1 - tuberculata ” (p. 54). Dell (1952) considered both forms to

belong to a single species.

While both shell varieties are common, individual shells

displaying an obvious shift between fine and coarse shell

formation are extremely rare. A single shell from Moreton

Bay, Queensland (109.1 mm ShL, QM Mol4232) displays a

transition from fine shell formation to coarse shell formation

at a point of previous damage (fig. 30). While the later

component of the shell possesses ears, it is not clear whether

the earlier component was eared or earless. No obvious

changes were noted in shell thickness, curvature of the keel or

relative heights of sequential keel tubercles.

Examination of a large number of A. nodosus shells found

no examples displaying a marked change in keel tubercle

height or ears that had been formed or subsumed. While eared

and earless forms exist, transition between the two types

appeared more gradual than the sudden transformation

documented in smaller species. A shell in the British Museum

(109.0 mm ShL [P], BMNH unreg., locality unknown, “B395,

e.”) displays an ear on only one side, clearly demonstrating the

plasticity of this character in this species (fig. 31).

2 Following Finn (2013) it is necessary to correct the original spelling

of A. nodosa to A. nodosus. In accordance with the International Code of

Zoological Nomenclature, Article 34.2 “the ending of a Latin or latinized

adjectival or participial species-group name must agree in gender with the

generic name with which it is at any time combined [Art. 31.2]; if the

gender ending is incorrect it must be changed accordingly (the author and

date of the name remain unchanged)” (I.C.Z.N., 1999). As Argonauta is

masculine "from the final noun nauta (a sailor)” (I.C.Z.N., 1999, p. 34)

the species-group name must be changed from the feminine nodosa (-a

feminine) to the masculine nodosus {-us masculine).

J.K. Finn

Figure 28. Reproduced illustrations referenced in the descriptions of Argonauta nodosus [Lightfoot], 1786 and A. argo Linnaeus, 1758: a,

illustration of A. nodosus [Lightfoot], 1786, designated as a lectotype by Moolenbeek (2008), Rumphius 1705, pi. 18, fig. 1; b, illustration of A.

argo Linnaeus, 1758, considered a paralectotype following the designation of a lectotype by Moolenbeek (2008), Rumphius 1705, pi. 18, fig. A.

Argonaut shell variability

Figure 29. Coarse and fine Argonauta nodosus shells: a, fine A. nodosus shell from Mayor Is., Bay of Plenty, New Zealand (127.2 mm shell

length, NMV F164784); b. Coarse A. nodosus shell from the Indo Pacific (127.3 mm shell length, NMV F164774). Scale bar = 1 cm.

J.K. Finn

Figure 30. Repaired Argonauta nodosus shell from Moreton Bay, Queensland. Left lateral view of repaired A. nodosus shell from Moreton Bay,

Queensland (109.1 mm shell length, QM Mol4232) showing a transition from fine shell formation (Fine) to coarse shell formation (Coarse) at

point of previous damage. Scale bar = 1 cm.

Argonaut shell variability

Argonauta argo Linnaeus, 1758

An image referenced in the original description of A. argo

Linnaeus, 1758, is considered a paralectotype, with designation

of a lectotype by Moolenbeek (2008); Rumphius, (1705) plate

18 figure A (fig. 28b). Shells of A. argo can be recognised by

an extremely narrow keel of consistent width. The keel

tubercles are paired and the lateral ribs are continuous (i.e.

they are not broken into separate tubercles).

Shells of A. argo are extremely consistent in dimensions and

sculpturing. The area that has caused the most confusion for

naturalists defining the species has been the aperture edge. A.

argo can display huge variation in the shape of the aperture edge

near the axis. Note the variation in the aperture edge of the two

shells presented in fig. 32. Unlike ear formation, this variation

occurs on the edge of the lateral wall parallel with the longitudinal

axis of the shell; it is not a lateral extension. The expression of the

lateral ribs can vary slightly from fine to coarse, suggesting the

presence of two varieties (fig. 32). Transition between fine and

coarse shell formation on a single shell is extremely rare. An

illustrated shell from Monterey, California (81.9 mm ShL,

USNM 61374) displays a shift from finer to coarser shell

formation at the point of earlier damage (fig. 33). Small A. argo

shells can also display laterally protruding ears. A shell from

Venezuela (51.4 mm ShL, USNM 122208) highlights the

plasticity of this character, displaying an ear on only the right

side (fig. 34). The varied size and shape of the keel tubercles on

the opposing sides of this shell demonstrate the range of

variability of these structures in this species.

Discussion

Among molluscs, the shells of small argonaut species (in

particular, A. nouryi) display an unprecedented level of variability.

The extreme forms are so different that it initially seems

incomprehensible that they could be produced by the same

argonaut species. Given this apparent disparity, it is necessary to

emphasise that argonaut shells are fundamentally different in

nature from the true molluscan shells of non-cephalopod

molluscs; they are produced by different structures, for different

reasons and have a different construction.

Shell material laid down by females of small argonaut

species (A. nouryi and A. hians ) can take one of two distinct

morphologies. Firstly, the shell can be heavy and thick walled

with prominent sculpture (Type 1 shell formation). The large

corrugations of the lateral walls are displayed as distinct robust

lateral ribs. The thickened keel is defined by two rows of large

and distinct keel tubercles. The axis of the shell projects laterally

to form large ears, providing support to the lateral walls. In the

second form, the shell can be lightweight and thin walled, with

greatly reduced sculpture (Type 2 shell formation). The

corrugations of the lateral walls are downgraded to fine lateral

ribs. The convex keel is undefined, with the keel tubercles

diminished to slight projections of the lateral rib extremities.

The axis of the shell is rolled ventrally to join the aperture edge

without lateral projection (i.e. earless).

The shell morphology expressed by a growing female

argonaut does not follow a predetermined order. Shells of

female A. nouryi demonstrate that females can switch between

the two shell formation types at least three times during

production of a single shell. The initial shell formation type is

variable (it can be Type 1 or Type 2), as is the portion of shell

laid down before switching to another shell formation type.

It is largely impossible to determine the conditions an

argonaut was exposed to at the time that it switched shell

formation types. The exception is the response to shell breakage.

Individual shells retain evidence of earlier trauma in the form of

repairs and irregularities in shell form. A. nouryi shells almost

invariably display a shift to Type 2 shell formation following

major damage. At the time of shell breakage, an argonaut would

be exposed and vulnerable. As has been observed for A. argo,

shell integrity is critical in allowing the argonaut to attain

neutral buoyancy, free itself from the sea surface and undertake

rapid horizontal locomotion (Finn and Norman, 2010). In the

absence of shell-aided buoyancy, the female must remain in the

water column by siphon-jetting alone. As such, it would be

imperative for a female argonaut to rebuild her shell as quickly

as possible following any damage. It is assumed that the shift to

thinner walled Type 2 shell formation allows the female to

rebuild her shell more rapidly, spreading the available building

material (calcium carbonate) over a greater area.

The different morphologies expressed in an individual A.

nouryi shell are therefore considered to represent periods of

varied rate of shell formation. Components of a shell laid

down over longer periods are believed to exhibit thicker walls

and more prominent sculpture (Type 1), while rapidly produced

sections display thinner walls and reduced sculpture (Type 2).

The rate at which the female lays down the shell is believed to

determine the gross morphology of the shell. Based on this

presumption, two hypotheses are raised to explain the variable

shell production rate (expressed as the variable shell formation

type) evident in undamaged, unrepaired shells:

• Hypothesis 1: Rate of shell production correlates directly

with animal growth. Three factors are believed to

influence octopus growth rate: temperature, nutrition, and

maturation or reproduction (see Semmens et al., 2004 for

a review). While very little is known about the lives of

argonauts, they are known to occur in the open ocean

spanning huge geographical distributions. This wide-

ranging pelagic existence has the potential to expose them

to a mosaic of food availability and water temperatures.

Encountering a large school of prey or pocket of warmer

water may result in a period of increased growth.

Additionally, reproductive investment (i.e. egg production)

may slow body growth. This hypothesis suggests that

these periods of varied morphological growth of the

animal are reflected in the gross morphology of the shell.

• Hypothesis 2: Rate of shell production influenced by

external factors. The shells of female argonauts, in

addition to providing protection and buoyancy, primarily

function as a case for external brooding of the female’s

eggs. Strings of eggs are suspended from the inner core of

the shell. This strategy requires that the internal volume

of the shell accommodate both the female argonaut and

her eggs. This hypothesis proposes that the space

constraints associated with commencement of egg

J.K. Finn

Figure 31. Single eared Argonauta nodosus shell from the British Museum: a-c, three perspectives of a single eared A. nodosus shell from the

British Museum (109.0 mm shell length [P], BMNH unreg., locality unknown, “B395, e.”); a, left lateral view; b, right lateral view; c, anterior

aperture view. Scale bar = 1 cm.

Argonaut shell variability

Figure 32. Coarse and fine Argonauta argo shells: a-b, shells of A. argo displaying different degree of sculpturing and variation in the aperture

edge; a, fine A. argo shell from off San Clement Island, California (113.3 mm shell length [P], USNM 316580); b, coarse A. argo shell from Baja

California (128.1 mm shell length [P], ANSP 404279). Scale bar = 1 cm.

J.K. Finn

Figure 33. Repaired Argonauta argo shell from Monterey, California. Repaired A. argo shell from Monterey, California (81.9 mm shell length,

USNM 61374): a, left lateral view; b, oblique left lateral view; c, oblique anterior aperture view. Note change in direction of lateral ribs along

repair line. Scale bar = 1 cm.

Argonaut shell variability

Figure 34. Single eared Argonauta argo shell from Venezuela, South America: a-c, three perspectives of a single eared A. argo shell from

Venezuela (51.4 mm shell length, USNM 122208); a, left lateral view; b, right lateral view; c, anterior aperture view. Scale bar = 1 cm.

100

J.K. Finn

spawning triggers an increase in shell production rate.

The gross morphology of the shell therefore displays

intermittent periods of volume constraints as a result of

intermittent spawning or brooding events.

To gain insights into the relationships between features of the

argonaut shell and its female occupant, focus was directed at

A. hians. The large single lot of female A. hians (with

accompanying shells) from Western Australia includes three

classes: submature, mature (unspawned) and spawned

individuals. Examination of the lot revealed that all spawned

individuals (i.e. all individuals with eggs deposited within

their shells) displayed a shift from Type 1 to Type 2 shell

formation. No converse arrangements were observed. Based

on this qualitative observation, this large sample appeared to

support Hypothesis 2; the presence of spawned eggs within the

shell causing a space constraint and thus triggering a shift to

more expansive thin-walled shell production. Hypothesis 1

would predict slower body growth of the argonaut associated

with increased reproductive investment in egg production.

This would predict slower shell production and hence a shift to

Type 1 formation. This was not observed. Quantitative analysis

of argonauts in this lot did not provide further insight. A full

range of characters of both the animals and shells were

measured. Features of the shell, the female and the shell

relative to the female were compared, plotted and analysed

with linear regression. In all instances, scatter plots indicated

linear relationships between animal and shell dimensions,

with linear regressions returning high coefficients of

determination values (i.e. R 2 values) with no discontinuities

observed between submature, mature and spawned individuals.

Examination of shells of larger species (A. nodosus and A.

argo ) revealed that they are not subj ect to the extreme variability

in shell form identified in the smaller argonaut species. While

both large species display two distinct morphologies (coarse

and fine forms), the two shell types are never expressed as

alternations on individual shells. Rare examples of extremely

damaged shells can display a shift from fine to coarse

morphology, but a reversion (i.e. from coarse to fine) was never

observed. Because the two shell types do not vary in shell wall

thickness or amount of material used, the variation between the

two shell types appears fundamentally different from that

observed in smaller species. One possible explanation is that

the coarser shells of larger species represent reformed shells

produced by large individuals (i.e. new shells constructed to

replace lost or damaged shells), while the finer shells represent

the original shells that are produced as the animals grow.

In the absence of captive rearing studies and sequential

collections of the same argonaut species from the same

location, it is not possible to conclusively support either

hypothesis. Based on limited observational evidence, it is the

author’s opinion that the variation observed in the shells of

small argonaut species is the result of space constraints (i.e.

Hypothesis 2) and independent of argonaut growth. The prime

circumstantial evidence comes firstly from gross differences

in shell occupation between large and small species, and

secondly from the dramatic transformation or reversion

boundaries on the shells of small species.

Gross differences in shell occupation. At commencement of

egg laying, the shells of females of small argonaut species

possess an extremely small amount of available space for egg

storage. Fig. 35a shows a preserved female A. nouryi that had

already commenced egg laying with a DML of 15.2 mm

(SBMNH 64369). As can be seen from the image, the space at

the top of the shell where the eggs are to be stored is extremely

small. The shell has barely formed through 90 degrees. Storage

of egg-strings within this shell will have a significant impact

on the space available for this small female within the shell.

Fig. 35b shows a female A. hians with a DML of 28.7 mm (QM

Mo77789). Yellow eggs are clearly visible and occupy almost

half of the shell volume. While the shell has developed through

almost a complete rotation, the volume occupied by the eggs

significantly displaces the female. With the posterior tip of the

mantle firmly against the egg mass, the female is still only

partially within her shell. Note the distance of the eye from the

edge of the shell aperture. In the absence of eggs, female

argonauts typically retract well into their shells with their eyes

at the boundary of the lateral walls. Fig. 36a presents a

photograph of a live female A. hians photographed in an

aquarium (after Sukhsangchan and Nabhitabhata, 2007). With

a large volume of eggs in the initial whorl of the shell, the

female can only partially retract within. The aperture edge of

the shell sits posteriorly to the mantle edge and a considerable

distance from the female’s eye.

Displacement of the female from the shell would provide a

strong stimulus for rapid shell deposition, resulting in the

extended flange-like form of Type 2 shells. Subsequent

interruptions to egg production (or hatching) could explain a

return to full occupancy of the shell and Type 1 shell formation,

as demonstrated in A. nouryi.

The apparent space constraint observed in smaller species

is not evident in larger species. Females of A. nodosus, observed

live, appear uninfluenced by large volumes of eggs held within

their shells. Fig. 36b presents a photograph of a live female A.

nodosus. This female is positioned well within her shell; note

the proximity of the female’s eye to the edge of the shell

aperture. Although not apparent from this photograph, the

female is carrying a huge volume of eggs. Fig. 36c presents the

egg strings revealed on removal of the female from her shell.

It is possible that the increased size of the shell of larger

species at the commencement of egg laying enables egg and

female accommodation. The shells of female A. nodosus are

considerably more developed than those of smaller species

when spawning commences; five females with ShL ranging

from 54.6 to 62.1 mm (and DML ranging from 31.1 to 38.5 mm)

were found to still be immature (see Finn, 2013, for details).

Immediacy of transformations and reversions. Additional

qualitative support comes from the abrupt nature of shell

transformations and reversions. Shells of female A. nouryi

display obvious precise boundaries between shell formation

types. It is the author’s opinion that the distinct boundaries

between shell formation types indicates that the causal

stimulus acts instantaneously on the female. It is felt that

spawning of eggs would have an immediate effect, requiring

the female to abruptly change the way the shell material is laid

Argonaut shell variability

101

Figure 35. Preserved female Argonauta nouryi and A. hians with spawned eggs: a, preserved female A. nouryi from the Pacific Ocean (15.2 mm

dorsal mantel length, 18.4 mm shell length, SBMNH 64369) with spawned eggs attached to the axis of the shell; b, preserved female A. hians

from the North West Shelf, Western Australia (28.7 mm dorsal mantel length, 38.9 mm shell length, QM Mo77789) with yellow eggs visible in

dorsal component of shell. Scale bar = 1 cm.

102

J.K. Finn

Figure 36. Images of live female argonauts, Argonauta hians and A. nodosus, demonstrating the effect of spawned eggs on the position of the

females relative to their shells: a, live female A. hians from Andaman Sea, Thailand, photographed in an aquarium (photo: J. Nabhitabhata, after

Sukhsangchan and Nabhitabhata 2007); b-c, A. nodosus Phillip Bay, Victoria, Australia (photos: R. Kuiter); b, live female argonaut photographed

in the wild; c, eggs of same specimen, shown with argonaut removed from shell.

Argonaut shell variability

103

down to accommodate the increased volume. If a change in

the growth rate was responsible for the transformation between

shell formation types, it is believed that the transition would be

more gradual and the boundaries in the shell less pronounced.

Argonaut nomenclature and the fossil record.

Misinterpretation of intra-specific shell variation has hindered

the resolution of extant argonaut systematics. Historic

generation of species names based on individual malformed

shells, and shells of different formation types, has created

confusion and complication. Fortunately, this practice has

largely ceased. The last major erection of new species names

occurred in 1914 when Monterosato proposed three new

species names and one variety based on four shells of A. argo

(Monterosato 1914). Interpretation of the fossil record, however,

appears to be mirroring the historic approach applied to extant

argonauts. Variation in shell characters is continuing to be used

to designate new fossil argonaut species (Stadum and Saul,

2000), and many have been erected based on single fossilised

shells (e.g. Martill and Barker, 2006). Saul and Stadum (2005)

reviewed the current situation stating: “ten fossil argonaut

species have been placed into four genera based on the absence

or presence of keels and the degree of sculpture” (p. 520). If the

situation is at all similar to that of extant argonauts, great

caution should be undertaken when erecting fossil argonaut

species based solely on shell characters.

Acknowledgements

This study would not have been possible without the generous

financial support of Australian Biological Resources Study,

the Hermon Slade Foundation, the Malacological Society of

London, the Linnean Society of New South Wales, La Trobe

University and Museums Victoria. This study relied heavily

on previously collected argonaut material housed in museum

collections throughout the world. Special thanks go to the

museum staff that facilitated access to this material including

(but not limited to) Paul Callomon (ANSP), Bob Hamilton-

Bruce (SAMA), Eric Hochberg (SBMNH), Thierry Laperousaz

(SAMA), Ian Loch (AM), Melanie Mackenzie (NMV), Alison

Miller (AM), Darryl Potter (QM), Martina Roeleveld (SAM),

Clyde Roper (USNM), Chris Rowley (NMV), Shirley Slack-

Smith (WAM), David Staples (NMV), Joanne Taylor (NMV),

Liz Turner (TMAG), Corey Whisson (WAM), Genefor

Walker-Smith (TMAG/NMV), Michael Vecchione (USNM),

Kathie Way (BMNH) and Richard Willan (NTM). Special

thanks go to individuals who provided feedback on various

versions of this manuscript, including: Prema Finn, Eric

Hochberg, Chung-Cheng Lu, Bruce Marshall, Mark Norman,

Richard Young and an anonymous reviewer. Rhyll Plant

deserves special thanks for producing all new illustrations, as

do Rudie Kuiter (Aquatic Photographies) and Charuay

Sukhsangchan (Kasetsart University, Thailand) for providing

additional live photographs, Daniel Geiger (SBMNH) for

assistance with scanning electron microscopy, Richard

Marchant (NMV) for assistance with statistics and Steve

Reynolds (Marine Life Society of South Australia) for

providing essential references.

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Memoirs of Museum Victoria 77:105-120 (2018)

1447-2554 (On-line)

https://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

DOI https://doi.Org/10.24199/j.mmv.2018.77.06

Published 14 December 2018

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

(http://zoobank.org /urn:lsid:zoobank.org:pub:FFB0A3El-53D8-416B-8E22-49ED61081AE5)

GARY C. B. Poore 1 (http://zoobank.org/urn:lsid:zoobank.org:author:c004d784-e842-42b3-bfd3-3r7d359f8975) and

PETER C. DwORSCHAK 2 (http://zoobank.org/urn:lsid:zoobank.org:author:4BCD9429-46AF-4BDA-BE4B-439EE6ADC657)

1 Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia gpoore@museum.vic.gov.au

2 Dritte Zoologische Abteilung, Naturhistorisches Museum, Burgring 7, Wien, Austria Peter.Dworschak@nhm-wien.ac.at

Abstract Poore, G.C.B., and Dworschak, P.C. (2018). The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae).

Memoirs of Museum Victoria 77: 105-120.

Four species of Eiconaxius are known to possess a denticulate median rostral carina: E. antillensis Bouvier, 1905,

E. asper Rathbun, 1906, E. cristagalli Faxon, 1893, and E. indicus (De Man, 1907). They are reviewed and two similar new

species are described: E. dongshaensis sp. nov., and E. gololobovi sp. nov. A key to distinguish them is presented.

Keywords Crustacea, Decapoda, Axiidae, Eiconaxius, new species

Introduction

The axiid genus Eiconaxius Bate, 1888 comprises more than

30 species confined to deep water that are, as far as is

known, associates of sponges (Komai and Tsuchida, 2012).

A few species differ from all others in having a prominent

median denticulate crest on the rostrum reaching back to the

gastric region where it bifurcates. In all other species this

ridge is low and smooth, or at best only slightly serrate.

Faxon (1893), who used the species epithet cristagalli for the

first species in this group alluded to a cock’s comb and

described the ridge as bearing ‘prominent teeth’. Three

more similar species were described shortly thereafter

(Bouvier, 1905; Rathbun, 1906; De Man, 1907). These four

are rediagnosed and two new similar species are described

from the Indo-West Pacific.

Methods

The material comes from: the Museum nationale d'Histoire

naturelle, Paris (MNHN) expeditions to Guadeloupe,

(KARUBENTHOS 2016) and to the Mayotte-Glorieuses

region, 2017 (BIOMAGLO); four expeditions by MNHN-

ORSTOM (Office de la recherche scientifique et technique

outre-mer, now IRD Institut de recherche pour le

developpement) (see http://expeditions.mnhn.fr/ and Richer de

Forges et al., 2013); collections made by the National Taiwan

Ocean University, Keelung (NTOU) in the South China Sea;

and the IUCN Seamounts expedition to the southwestern

Indian Ocean, 2011 (Rogers and Taylor, 2012), material now

lodged in Naturhistorisches Museum, Vienna (NHMW).

Type material consulted and types of new species are

lodged in the Naturalis Biodiversity Center, Leiden (ZMA),

Museums Victoria, Melbourne (NMV) and the Museum of

Comparative Zoology, Harvard University, Cambridge (MCZ).

Other specimens were viewed in the National Museum of

Natural History, Washington (USNM).

Size is expressed as carapace length, including rostrum, in

mm. Relative lengths of fixed fingers of chelipeds are expressed

as (a-b)lb where a is the length of the lower margin of the

propodus, including the fixed finger, and b is the length of the

upper margin.

As part of this study and continuing discovery of new

species in the Indo-West Pacific diagnoses have been prepared

for all species of Eiconaxius 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 two respects from every other species.

The poorly known Eiconaxius asper Rathbun, 1906 is

diagnosed on the basis of its description but not included in

the key.

Family Axiidae Huxley, 1879

Eiconaxius Bate, 1888

Eiconaxius Bate, 1888: 40. - Poore, 2017: 365-366.

Remarks. Poore (2017) provided a new diagnosis and discussed

the synonymy of the genus. The following key deals only with

species having a prominent median denticulate crest on the

rostrum reaching back to the gastric region.

106

G.C.B. Poore & PC. Dworschak

Key to species of Eiconaxius cristagalli species complex

1. East Pacific or Caribbean species.2

- Indo-West Pacific species.3

2. Rostrum triangular, tapering, median carina with 6 or 7

teeth; lateral carina without clear hiatus between posterior

extension of rostral margin and posterior section; East

Pacific. E. cristagalli Faxon, 1893

- Rostrum tapering more steeply anteriorly, median carina

with >10 teeth (Figs lb, 2b); lateral carina with a clear

hiatus between posterior extension of rostral margin and

posterior section (Figs lb, 2b); Caribbean.

. E. antillensis Bouvier, 1905

3. Major cheliped palms fixed finger cutting edge crenellate

(Fig. 5f, g); rostrum parallel-sided basally, tapering

steeply anteriorly; lateral carina continuous from lateral

margins of rostrum, with short overlap posteriorly (Fig.

5c); Western Indian Ocean. E. gololobovi sp. nov.

- Major cheliped palms fixed finger cutting edge with

simple blade (Figs 3e, f, 9g, h); rostrum tapering evenly

from base to acute apex; lateral carina usually with

considerable hiatus between lateral margins of rostrum

and posterior section, or absent (Figs 3c, 9c).4

4. Rostrum with ventral tooth (Fig. 3b); major cheliped,

distolateral margin of propodus with 2 teeth at base of

dactylus, lobe and narrow keyhole-shaped notch in gape

(Fig. 3e); East China Sea. E. dongshaensis sp. nov.

- Rostrum without ventral tooth (Fig. 9b); major cheliped,

distolateral margin of propodus straight at base of

dactylus, circular notch in gape (Fig. 9g); SW Pacific.

. E. indicus (De Man, 1907)

Eiconaxius antillensis Bouvier, 1905

Figures 1 ,2

Eiconaxius crista-galli var. antillensis Bouvier, 1905: 803.

Iconaxius cristagalli var. antillensis. - Balss, 1925: 210.

Axius (Eiconaxius) crista-galli antillensis. - Bouvier, 1925: 456-

458, pi. 8 fig. 3, pi. 9 fig. 1. - De Man, 1925: 4, 33.

Eiconaxius antillensis. - Sakai and de Saint Laurent, 1989: 21. -

Kensley, 1996: 475. - Sakai, 2011: 273. - Felder et al. 2009: 1063.

Figure 1 .Eiconaxius antillensis Bouvier, 1905, Syntype, MCZ 11964 (male, 6.1 mm) - a, carapace, pleon, lateral, b, anterior carapace, antenna,

antennule, dorsal view, c, rostrum, anterior median carina, left oblique view, d, telson, right uropod. e, f, major cheliped (left), mesial, lateral

views, g, h, minor cheliped (right), mesial, lateral views. Scale bar = 1 mm.

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

107

Figure 2. Eiconaxius antillensis Bouvier, 1905, IU-2016-8456 (ovigerous female, 6.3 mm) - a, b, anterior carapace, antenna, antennules, lateral,

dorsal views, c, d, major cheliped (left), lateral, mesial views, e, f, minor cheliped (right), lateral, mesial views, g, h, major cheliped (left), distal

propodus, dactylus, lateral, mesial views. NMV J71649 (ovigerous female, 7.6 mm) - i, major cheliped (left), lateral view. Scale bars = 1 mm.

Eiconaxius ?antillensis. - Poupin and Corbari, 2016: 32, fig. 8a

(photograph).

Material examined. Syntypes. Off Montana, Monserrat, 16.7°N,

62.2°W, 545 m, Blake stn 154, MCZ CRU-11946 (male, 6.1 mm).

Barbados, 12.9°N, 59.6°W, 527 m, Blake stn 281, MCZ CRU-11947

(ovigerous female, 6.7 mm).

Guadeloupe, N of Grande Terre, 16°37'N, 61°31'W, 432-482 m

(KARUBENTHOS stn DW4550), MNHN IU-2016-8456 (ovigerous

female, 6.3 mm), MNHN IU-2018-106 (3 females, 5.8-7.1 mm),

MNHN IU-2013-18924 (2 males, 4.3, 6.7 mm), NMV J71649

(ovigerous female, 7.6 mm). NMV J71655 (4 females, 53-7.3 mm).

Caribbean Sea, Mexico, NE of Yucatan, 22.72°N, 86.22°W, 1030

m, (Cruise: 65A9-21) USNM 1014172 (1 specimen, det. B. Kensley,

examined by C.C. Tudge). E of Guadeloupe, 16.55°N, 61.62°W, 466-

585 m ( Pillsbury stn P994), USNM 1081088 (3 specimens, examined

by C.C. Tudge).

Diagnosis. Rostrum tapering more over distal third than

proximal, with rounded apex in adult (acute in juvenile), 1.2-

1.3 times as long as wide. Median carina erect, with clear

dentition (10-20 teeth); lateral gastric carina with hiatus

between lateral rostral margin and short gastric section, visible

only posterior to confluence of submedian carinae. Major

108

G.C.B. Poore & PC. Dworschak

cheliped, palm wider distally than at midpoint, distolateral

margin with sharp tooth at base of finger; fixed finger about 3/4

length of upper margin of palm, cutting edge with basal notch,

blade-like proximal half, and distal concavity, dactylus cutting

edge with blunt tooth at midpoint, denticulate beyond. Minor

cheliped, palm upper margin as long as greatest width;

distolateral margin with prominent triangular toothed lobe at

base of dactylus; fixed finger cutting edge weakly crenellate,

with excavated distal quarter.

Distribution. Monserrat, Barbados, Guadeloupe, Caribbean

Sea, W Atlantic, S. of Jamaica; 432-1030 m.

Remarks. Bouvier (1905) listed no specimens when he erected

his new variety but mentioned a male and a female from two

stations later (Bouvier, 1925). We treat them as syntypes. The

male syntype differs significantly from all other specimens

identified by us and by others in the past, including the second

specimen identified by Bouvier (1925). Notably the distolateral

margin of the palm of the major cheliped bears a high and

sharp tooth (Fig. If) whereas in all other material there is a

blunt asymmetrical tooth (Figs 2h, i are typical; broken in Fig.

2c). The notch below this tooth is less pronounced in the male

syntype than in other specimens. These differences appear not

size-related - this specimen is within the size range of the other

material.

Eiconaxius antillensis is the only species of this group in

the Atlantic but is confined to the Caribbean or nearby, an area

with biogeographic affinities to the Indo-West Pacific where

the remainder live. The species is similar to E. cristagalli, the

Eastern Pacific species, in the dentition of the propodus and

fixed finger of the major cheliped, both with a deep notch in

the gape and a distal concavity on the finger. It differs in

having a narrow, parallel-sided rostrum with numerous teeth

on the median carina (cf. rostrum triangular, tapering, median

carina with six or seven teeth in Faxon’s and Kensley’s

accounts of E. cristagalli). Kensley (1996: fig. 7F) showed the

distolateral margin of the minor cheliped of E. cristagalli with

two teeth; only one is present on these specimens of E.

antillensis. The lateral carina has a clear hiatus between the

posterior extension of the rostral margin and the ridge level

with the submedian carina, as in E. gololobovi sp. nov.; Kensley

(1996) noted no such hiatus in E. cristagalli.

Eiconaxius asper Rathbun, 1906

Eiconaxius asper Rathbun, 1906: 895, fig. 52. - Sakai and de Saint

Laurent, 1989: 22. - Kensley, 1996: 475. - Sakai, 2011: 273.

Iconaxius asper. - Balss, 1925: 209.

Axius (Eiconaxius) asper. - De Man, 1925: 4, 14, 34.

Diagnosis. Median carina erect, with clear dentition. Major

cheliped, palm wider distally than at midpoint. Major cheliped,

palm lateral and mesial faces tuberculate near base of fingers-,

distolateral margin with 1 or 2 teeth in gape but without a deep

notch-, fixed finger about half as long as upper margin of palm,

cutting with basal notch, blade-like proximal half, and distal

concavity, dactylus cutting edge with basal molar-like tooth,

notch and straight beyond. Minor cheliped, palm lateral and

mesial faces tuberculate near base of fingers.

Distribution. Hawaii, Kauai I., 418-628 fm (765-1149 m)

(known only from type locality).

Remarks. Rathbun (1906) remarked that the species resembled

E. cristagalli in having the median carina denticulate but

differed in the ‘presence of a larger basal tooth on dactylus of

larger hand and a more prominent tooth not far from middle of

the pollex [fixed finger]’. She illustrated only the larger cheliped

where these differences are unconvincing compared to Faxon’s

(1895) figures of a type or Kensley’s (1996) figure of

E. cristagalli from the Galapagos. Without examining the types

we are unsure of the status of this species. It is omitted from

the key.

Eiconaxius cristagalli (Faxon, 1893)

Axius crista-galli Faxon, 1893: 193. - Faxon, 1895: 104, pi. 28 fig.

1-lh.

Axius (Eiconaxius) crista-galli. - Borradaile, 1903: 538 - De Man,

1925: 4, 14.

Eiconaxius crista-galli. - Rathbun, 1906: 895.

Iconaxius cristagalli. - Balss, 1925: 210.

Eiconaxius cristagalli. - Sakai and de Saint Laurent, 1989: 18. -

Hendrickx, 1995: 390. - Hendrickx, 2008: 1002, fig. 2. - Kensley,

1996: 475, 480-481, fig. 7. - Sakai, 2011: 276-278.

Diagnosis. Rostrum 1.5-2.0 times as long as wide. Median

carina erect, with clear dentition. Major cheliped, palm lateral

and mesial faces tuberculate near base of fingers; distolateral

margin with single blunt tooth in gape\ fixed finger 0.6 length of

upper margin of palm, cutting edge with broad blade over

proximal half, irregular beyond-, dactylus cutting edge with

basal molar-like tooth, notch and straight beyond. Minor

cheliped, palm upper margin as long as greatest width;

distolateral margin with prominent bifid triangular tooth at

base of dactylus. Uropod endopod anterolateral apex acute,

with 1 or few small teeth.

Distribution. Pacific coast, Panama (Albatross stn 3358), 465

fm (851 m) (type locality); Ecuador, Galapagos Is, 111 m

(Kensley, 1996), 1123-1378 m (Hendrickx, 2008).

Remarks. The species was redescribed by Kensley (1996) and

compared by us with E. antillensis above. Hendrickx (2008)

illustrated variation in the dentition of the rostrum of five

syntypes and recorded the species from much greater depths

than previously.

Eiconaxius dongshaensis sp. nov.

(http://zoobank.org/urmlsid: zoobank.org:act:7A965426-D789-

4C02-8B8C-4A3A84C17E3B)

Figures 3, 4a, b

Eiconaxius indicus. - Sakai and Ohta, 2005: 73-77, figs 3-5. -

Tsang et al., 2008: 363, fig. 2.

Material examined. Holotype. South China Sea, off Pratas Islands, S

of Hong Kong, 20°50.9'N, 117°27.17'E, 730-720 m (stn CD320),

NTOU A01439 (female, 10.0 mm).

Paratype. Collected with holotype, NTOU A01440 (female,

9.4 mm).

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

109

antenna, lateral, c, anterior carapace, antennule, antenna, dorsal, d, pleomere 6, telson, right uropod. e, f, right major cheliped, lateral and mesial,

g, h, left minor cheliped, lateral and mesial, i, maxilliped 3. j, pereopod 2. k, 1, left pereopod 3, detail of dactylus. m, n, left pereopod 4, detail of

dactylus. o, p, left pereopod 5, detail of dactylus. Scale = 2 mm (except b, c, 1, n, p).

Diagnosis. Rostrum 1.5-2.0 times as long as wide; with ventral

tooth. Median carina erect, with clear dentition; sublateral

gastric carinae present, diverging widely from base of median

carina; lateral gastric carina not running from rostrum,

commencing level with confluence of submedian carinae.

Major cheliped, palm wider distally than at midpoint, upper

margin denticulate; lateral and mesial faces of palm tuberculate

near base of fingers; distolateral margin with 2 teeth at base of

dactylus, lobe and keyhole-shaped notch in gape; fixed finger

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

basal notch, blade-like proximal half, and distal concavity;

dactylus cutting edge with basal molar-like tooth, notch and

110

G.C.B. Poore & PC. Dworschak

Figure 4. Eiconaxius dongshaensis sp. nov., holotype, NTOU A01439 (female, 10.0 mm) - a, habitus, in vivo, b, habitus in host sponge (photos,

Tin-Yam Chan). Eiconaxius gololobovi sp. nov., unspecified specimen - c, habitus, in vivo (photo, David Shale).

straight beyond. Minor cheliped, palm upper margin as long as

greatest width; distolateral margin with prominent triangular

tooth at base of dactylus (bifid); lateral and mesial faces

tuberculate near base of fingers.

Description. Carapace smooth. Rostrum 0.18 carapace length,

concave dorsally, tapering evenly to acute tip, twice as long as

wide at base, with 6 teeth on lateral margins, depressed below

level of median carina, not continuous with lateral carinae,

with ventral tooth. Lateral gastric carina unarmed, reaching

anteriorly to base of median carina. Submedian gastric carina

smooth, together curved and converging on median carina.

Median gastric carina prominent, erect, reaching two-thirds

along rostrum, with c. 13 uneven sharp teeth.

Pleuron 1 posteroventrally rounded; pleuron 2 truncate,

posteroventrally acute; pleura 3 and 4 truncate, posteroventrally

subacute; pleuron 5 rounded, all 5 pleura without anteroventral

tooth; pleuron 6 with acute posteroventral angle; pleonite 6

dorsal posterior margin denticulate.

Eyestalk, 0.4 length of rostrum; cornea white.

Antennular peduncle reaching to midpoint of antennal

article 4; article 1 unarmed. Antennal peduncle article 1

unarmed; article 2 with upper-distal elongate triangular

blade, reaching two-thirds along article 4; scaphocerite a

vertical blade, reaching just beyond end of article 4; article

3 lower margin with distomesial tooth; article 5 about half

length of article 4.

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

111

Maxilliped 3 basis with mesial spine; ischium unarmed;

crista dentata of about 16 small similar teeth; merus and carpus

unarmed; exopod with flagellum reaching to base of merus.

Major cheliped merus lower margin with c. 8 irregular

teeth, upper margin with 2 minute blunt teeth; carpus lower

margin with 1 distal tooth; propodus upper margin carinate,

obscurely dentate, angled distally, length 0.9 greatest height,

lower margin smooth, lateral face tuberculate near base of

fixed finger, mesial face with few tubercles near base of fixed

finger; fixed finger 0.6 times as long as upper palm, cutting

edge shallowly concave over proximal two-thirds, denticulate

beyond, with longitudinal mesial ridge; distolateral margin of

palm with 2 teeth at base of dactylus, lobe and keyhole-shaped

notch in gape; distomesial margin of palm with 2 teeth at base

of dactylus; dactylus distally curved, upper margin carinate,

cutting edge with basal tooth, straight beyond.

Minor cheliped shorter and more slender than major, palm

0.8 times height of major palm; merus lower margin with 8

sharp teeth, increasing in size distally; carpus lower margin

with 1 distal tooth; propodus dilating, upper margin carinate,

weakly denticulate, as long as greatest height, lower margin

smooth, lateral and mesial faces tuberculate near base of fixed

finger; fixed finger 1.25 times as long as upper palm, cutting

edge dentate, with longitudinal mesial and lateral ridges;

distolateral margin of palm oblique, with prominent bifid

triangular tooth in gape; distomesial margin of palm oblique,

with 2 triangular teeth in gape; dactylus distally curved, upper

margin carinate, cutting edge smooth.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus 0.8 length of propodus upper margin;

propodus upper margin 2.5 times as long as dactylus. Pereopod

3 merus unarmed; propodus 3.3 times as long as dactylus, with

7 rows of spiniform setae, of 2-4 setae; dactylus spatulate, with

10 spiniform setae along oblique margin, plus unguis, 4 facial

spiniform setae. Pereopod 4 similar to pereopod 3; propodus

3.5 times as long as dactylus, with 7 rows of spiniform setae, of

1-3 setae; dactylus spatulate, with 8 spiniform setae along

oblique margin, plus unguis, and 2 facial spiniform setae.

Pereopod 5 dactylus spatulate, with 8 spiniform setae along

oblique margin, plus unguis, with 1 facial spiniform seta.

Uropodal endopod 1.9 times as long as wide, oval, anterior-

distal margin with c. 20 evenly-spaced teeth, without

longitudinal ridge. Uropodal exopod 1.7 times as long as wide,

oval, anterior margin with many small irregular teeth over

most of length, without longitudinal rib.

Telson 1.35 times as long as wide, widest at midlength,

then tapering to rounded posterolateral angles, lateral margin

upturned, denticulate, distal margin evenly curved, with

posteromedian spine; dorsal face smooth, concave.

Etymology, dongshaensis, from Dongsha, the Chinese name of

the Pratas Islands near the type locality.

Distribution. Sulu Sea, Philippines, 688-2019 m (Sakai and

Ohta, 2005); Pratas Is., South China Sea, 720-730 m (Tsang et

al., 2008).

Remarks. Sakai and Ohta (2005) illustrated the habitus, tailfan

and chelipeds of this species as Eiconaxius indicus based on

nine specimens of both sexes from the Sulu Sea. Their figures

can by-and-large be reconciled with those published here. The

chelipeds are similar but the palmar distolateral and

mesiolateral armature is simpler in their figures than in the

types. Sakai and Ohta (2005) showed tubercles on the lateral

face of the palm of the major cheliped in fig. 3 but not in fig. 5,

and did not note the easily overlooked subrostral tooth.

Eiconaxius dongshaensis is the only species in this group

with a rostral ventral tooth. This tooth and the small keyhole¬

shaped notch in the gape of the cheliped are like no other species.

Eiconaxius gololobovi sp. nov.

(http://zoobank.org/urn:lsid:zoobank.org:act:6BA2620A-9AF9-

4536-AD2F-4C0E980F65D8)

Figures 4c, 5-8

Material examined. Holotype. SW Indian Ocean, Gololobov Bank,

‘Coral’ seamount, 41°21.0283'S, 42°55.145'E, 1117 m (RV James Cook

cruise JC606, code 996) NHMW 25659 (male, 7.8 mm).

Paratypes. SW Indian Ocean, Gololobov Bank, ‘Coral’ seamount,

41°21.767'S, 42°54.907'E, 686.5 m, NHMW 25658 (male, 6.8 mm),

NHMW 25660 (female, 6.1 mm), NHMW 25661 (male, 7.7 mm),

NHMW 25662 (male, 5.1 mm), NHMW 25663 (female, 5.4 mm),

NHMW 25664 (female, 6.8 mm), NHMW 25665 (male, 4.8 mm),

NHMW 25666 (male, 8.3 mm), NHMW 25667 (ovigerous female, 9.1

mm), NHMW 25668 (ovigerous female, 8.1 mm), NHMW 25669

(male, 7.0 mm), NHMW 25670 (ovigerous female, 8.0 mm), NHMW

25671 (male, 6.2 mm), NHMW 25672 (male, 5.6 mm), NHMW 25673

(female, 5.6 mm), NHMW 25674 (ovigerous female, 8.8 mm), MNHN

IU-2016-8156 (male, 7.4 mm), NMV J71648 (male, 7.8 mm), NHMW

26058 (male, 6.1 mm), NHMW 26059 (female, 6.8 mm).

Other material. Madagascar. S of Pt Barrow, 25°39’S, 44°16’E,

986-991 m (ATIMO VATAE stn CP3596), MNHN IU-2014-12083

(ovigerous female, 7.4 mm).

Mozambique Channel, Geyser Bank, between Malekula and

Ambrym islands, 12°18'S, 46°27'E, 920-935 m (BIOMAGLO stn

DW4791), MNHN IU-2017-530 (male, 6.4 mm).

Diagnosis. Rostrum parallel-sided proximally, with acute apex,

1.2-1.3 times as long as wide. Median carina erect, with clear

dentition; sublateral gastric carinae present, diverging widely

from base of median carina-, lateral gastric carina continuous

with short overlap of ridges posterior to confluence of

submedian carinae. Major cheliped, merus lower margin with 2

spines near midpoint, or with single denticle; palm wider

distally than at midpoint, upper margin smooth, carinate, or

denticulate (juveniles); fixed finger about half as long as upper

margin of palm, cutting edge crenellate, with row of diminishing

rounded teeth; dactylus cutting edge smooth. Minor cheliped,

palm upper margin significantly less than greatest width;

distolateral margin with prominent triangular tooth at base of

dactylus; fixed finger cutting edge smooth, straight.

Description of holotype. Carapace smooth, few obsolete

tubercles on gastric region between carinae. Rostrum 0.18

carapace length, concave dorsally, tapering distally to acute tip,

1.35 times as long as wide at base, with c. 8 obscure teeth on

lateral margins, depressed below level of median carina,

continuous with lateral carinae, without ventral tooth. Lateral

gastric carina unarmed, with posterior section weakly separated

112

G.C.B. Poore & PC. Dworschak

Figure 5. Eiconaxius gololobovi sp. nov., holotype, NHMW 25659 (male, 7.8 mm) - a, carapace, pleon, lateral, b, c, anterior carapace, antennule,

antenna, lateral and dorsal, d, median carina. e, pleomere 6, telson, right uropod. f, g, right major cheliped, lateral and mesial, h, i, left minor

cheliped, lateral and mesial. Scale bar = 1 mm (except b-d).

from, but almost overlapping anterior section. Submedian

gastric carina smooth, together curved and converging on

median carina, slightly longer than lateral gastric carina.

Median gastric carina prominent, erect, reaching midpoint of

rostrum, with 9 erect teeth becoming pentagonal anteriorly.

Pleuron 1 posteroventrally acute; pleura 2, 3 truncate,

posteroventrally acute; pleuron 4 truncate, posteroventrally

subacute, with anteroventral tooth; pleuron 5 rounded; pleuron

6 with acute posteroventral angle; pleonite 6 dorsal posterior

margin denticulate.

Eyestalk, reaching half length of rostrum; cornea white.

Antennular peduncle reaching two-thirds length of antennal

article 4; article 1 unarmed. Antennal article 1 unarmed;

article 2 with distal spine an elongate triangular blade,

reaching two-thirds length of article 4; scaphocerite a

vertical blade, reaching to end of article 5; article 3 lower

margin with distomesial tooth; article 5 about half length of

article 4.

Maxilliped 3 coxa unarmed; ischium with tubercle; crista

dentata of c. 15 similar obsolete teeth; merus and carpus

unarmed; exopod reaching beyond midpoint of merus.

Major cheliped merus lower margin convex, with 3

minute teeth, upper margin with minute blunt tooth; carpus

lower margin with 1 distal tooth; propodus upper margin

carinate, angled distally, length 1.15 greatest height, lower

margin smooth, lateral face with few minute tubercles;

mesial face smooth; fixed finger 0.7 times as long as upper

palm, cutting edge with proximal U-shaped notch, 10

diminishing triangular teeth, with longitudinal lateral

ridge; distolateral margin of palm weakly evenly convex;

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

113

Figure 6. Eiconaxius gololobovi sp. nov., holotype, NHMW 25659 (male, 7.8 mm) - a, b, maxilliped 3, anterior detail of merus. c, pereopod 2. d,

e, right pereopod 3, detail of dactylus. f, g, right pereopod 4, detail of dactylus, h, i, right pereopod 5, detail of dactylus. j, pleopod 2, proximal

endopod, appendices interna and masculina. Scale = 1 mm (except b, e, g, i)..

distomesial margin of palm convex; dactylus distally

curved, upper margin carinate, cutting edge smooth, weakly

convex proximally.

Minor cheliped shorter and more slender than major,

palm 0.9 times height of major palm; merus lower margin

with minute tooth at midpoint; carpus lower margin with 1

distal tooth; propodus dilating, upper margin carinate, with

obscure proximal notch, 0.85 times greatest height, lower

margin smooth, lateral and mesial faces smooth; fixed

finger 1.4 times as long as upper palm, cutting edge weakly

serrate, with longitudinal lateral ridge; distolateral margin

of palm oblique, with prominent triangular tooth in gape;

distomesial margin of palm oblique-convex, with 2 tubercles

in gape; dactylus distally curved, upper margin carinate,

cutting edge smooth.

Pereopod 2 ischium lower margin unarmed; merus

lower margin unarmed; carpus as long as propodus upper

margin; propodus upper margin 2.2 times as long as

dactylus. Pereopod 3 merus unarmed; propodus 2.8 times

as long as dactylus, with 5 rows of 1-4 spiniform setae;

dactylus spatulate, with 11 spiniform setae along oblique

margin, plus unguis, and 2 rows of 3 facial spiniform

setae. Pereopod 4 merus more slender than that of

pereopod 3; propodus 2.8 times as long as dactylus, with

6 rows of 2-5 spiniform setae; dactylus spatulate, with 12

spiniform setae along oblique margin, plus unguis, and 2

oblique rows of 3 facial spiniform setae. Pereopod 5

dactylus spatulate, with 11 spiniform setae along oblique

margin, plus unguis, without facial spiniform seta.

Pleopod 2 with appendices interna and masculina of similar

lengths, 0.8 length of proximal endopod. Uropodal endopod 1.9

114

G.C.B. Poore & PC. Dworschak

Figure 7. Eiconaxius gololobovi sp. nov., NHMW 25662 (male, 5.1 mm) - a, anterior carapace, antenna, lateral, b, anterior carapace, dorsal, c, d,

right major cheliped, lateral and mesial, e, f, left minor cheliped, lateral and mesial. NHMW 25658 (male, 6.8 mm) - g, anterior carapace, h,

pleomere 6, telson, right uropod. i, left major cheliped, j, right minor cheliped. Scales = 1 mm.

times as long as wide, oval, anterior-distal margin with 10

evenly-spaced teeth, last distal, without longitudinal ridge.

Uropodal exopod 1.4 times as long as wide, oval, anterior

margin with small irregular teeth over distal two-thirds, without

longitudinal rib.

Telson 1.5 times as long as wide, widest at third length,

then tapering to posterolateral angles, lateral margin upturned,

obscurely denticulate, distal margin obtusely angled, with

posteromedian spine; dorsal face smooth.

Etymology. For Ya. K. Gololobov (1909-1980), Russian

oceanographer, for whom the Gololobov Bank on the South¬

west Indian Ocean Ridge is named, of which the type locality,

‘Coral’ seamount, is part.

Distribution. Gololobov Bank, Mozambique Channel, south¬

west Indian Ocean; 686-1117 m depth.

Remarks. Eiconaxius gololobovi sp. nov. is known from 21

specimens ranging 4.8-9.1 mm in carapace length collected from

sponges by ROV on the Gololobov Bank, plus two from nearby

localities in the Mozambique Channel. The species is

distinguished from others in this group by the regularly dentate

cutting edge of the fixed finger of the major cheliped (simple in

other species) and from most species by the hiatus in the lateral

carina. The dentition of cheliped fingers is obsolete in the

ovigerous female from Madagascar (Fig. 81, m). This individual

has a more acute rostrum than others (Fig. 8i) but in this is similar

to at least one smaller male (Fig. 8b) from Mozambique Channel.

While several individuals display the distinctive pentagonal teeth

on the median carina (Fig. 5d) (like children’s drawings of little

houses), others have a similar number of more irregular teeth

(Figs 7g, 8a, h) but in the smallest specimen dentition is obsolete

(Fig. 7a). The upper margin of the propodus of the chelipeds often

has a clear proximal notch (Figs 5h, i, 7i, j) but it is not obvious on

the major cheliped of larger individuals (Figs 5f, g, 8d, e, 1). The

smallest individuals differ in having this margin serrate and a

prominent tooth on the upper margin of the dactylus (Figs 7c-f).

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae) 115

Figure 8. Eiconaxius gololobovi sp. nov., MNHN IU-2013-7046 (male, 6.4 mm) - a, carapace, pleon, lateral, b, carapace, dorsal, c, pleomere 6,

telson, right uropod. d, e, right major cheliped, lateral and mesial. NMV J71648 (male, 7.8 mm) - f, right minor cheliped, propodus distolateral

margin. MNHN IU-2016-8156 (male, 7.4 mm) - g, right minor cheliped, propodus distomesial margin. MNHN IU-2014-12083 (ovigerous

female, 7.4 mm) - h, carapace, antennule, antenna, lateral, i, carapace, antennule, antenna, dorsal, j, right epimera of pleomeres 1-6. k, pleomere

6, telson, left uropod. 1, m right major cheliped, lateral and mesial, n, o, left minor cheliped, lateral and mesial. Scale bars = 1 mm.

116

G.C.B. Poore & PC. Dworschak

Eiconaxius indicus (De Man, 1907)

Figures 9, 10

lconaxius crista-galli var. indica De Man, 1907: 128-129.

Iconaxius cristagalli var. indica. - Balss, 1925: 210.

Axius (Eiconaxius) crista-galli var. indica. - De Man, 1925: 4, 15,

31, pi. 2 fig 3.

Eiconaxius indica. - Sakai and de Saint Laurent, 1989: 22.

Eiconaxius indicus. - Kensley, 1996: 475. -Sakai, 2011: 279

(partim), fig. 52.

Not Eiconaxius indicus. - Sakai and Ohta, 2005: 73-77, figs 3-5.

- Tsang et al., 2008: 363, fig. 2 (= Eiconaxius dongshaensis sp. nov.)

Material examined. Holotype. Indonesia, E of Palau Kei Besar (Great

Kei L), 5°54'S, 132°56'7"E, 984 m ( Siboga stn 267), ZMA102465

(ovigerous female, 10.6 mm) - photographed by C.H.J.M. Fransen,

figured by Sakai (2011: fig. 52).

Vanuatu. Between Malekula and Ambrym islands, 16°30.7'S,

167°55.5'E, 550-565 m (BOA1 stn CP2468), MNHN IU-2014-10474

Figure 9. Eiconaxius indicus (De Man, 1907). MNHN IU-2014-10474 (male, 9.1 mm) - a, carapace, pleon, lateral, b, anterior carapace, antenna,

lateral, c, anterior carapace, antennule, antenna, dorsal, d, pleomere 6, telson, right uropod. e, f, right minor cheliped, lateral and mesial, g, h, left

major cheliped, lateral and mesial, i, pereopod 2. j, k, left pereopod 3, detail of dactylus. 1, left pereopod 5, detail of dactylus. Scale = 2 mm

(except b, c, k, 1).

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae) 117

Figure 10. Eiconaxius indicus (De Man, 1907). MNHNIU-2016-8020 (ovigerous female, 12.1 mm) - a, anterior carapace, antenna, b, telson, right

uropod. c, d, left major cheliped, lateral and mesial views, e, f, left minor cheliped, lateral and mesial views. MNHN IU-2014-7149 (ovigerous

female, 11.0 mm) - g, anterior carapace, antenna. MNHN IU-2014-7151 (male, 9.0 mm) - h, right major cheliped fingers, lateral view. NMV

J71631 - telsons of 4 individuals: i, male, 9.8 mm; j, male, 7.3 mm; k, ovigerous female, 9.0 mm; 1, female, 7.6 mm. Scale = 2 mm

(male, 9.1 mm). Between Malekula and Epi islands, 16°38.28'S,

167°58.38'E, 586-646 m (BOAO stn CP2307), MNHN IU-2014-7149-

7152 (2 ovigerous females, 10.8, 11.0 mm; 2 males, 9.0, 10.8 mm)

Solomon Islands, NW of San Cristobal, 09°56'S, 161°04'E, 418-

432 m (SALOMON 1 stn DW1826), MNHN IU-2016-8020 (ovigerous

female, 12.1 mm).

New Caledonia, BATHUS 3 stations. Loyalty Ridge, seamount K,

24°43'S, 170°07'E, 750-760 m (stn DW778), NMV J71631 (4 males,

6.1-9.8 mm; 3 females, 7.6-9.0 mm); seamount K, 24°44’S, 170°08’E,

770-830 m (stn DW776), MNHN IU-2016-8022 (2 females, 6.1, 9.3

mm); seamount D, 23°35’S, 169°37'E, 655 m (stn DW800), MNHN IU-

2016-8023 (female, 10.0 mm). S of lie des Pins, 23°09'S, 167°11'E, 650-

680 m (BIOCAL stn DW36), MNHN IU-2016-8021 (female, 7.8 mm).

Diagnosis. Rostrum tapering more over distal third than

proximal, with rounded apex in adult (acute in juvenile), 1.5-2.0

times as long as wide. Median carina erect, with clear dentition;

sublateral gastric carinae present, diverging widely from base of

median carina-, lateral gastric carina with hiatus between lateral

rostral margin and short gastric section, visible only posterior

to confluence of submedian carinae. Major cheliped, palm

upper margin smooth, carinate, or denticulate; lateral and

mesial faces tuberculate near base of fingers; distolateral

margin with 1 or 2 teeth in gape but without a deep notch; fixed

finger about half as long as upper margin of palm, cutting edge

118

G.C.B. Poore & PC. Dworschak

blade-like, with proximal notch and distal concavity, dactylus

cutting edge with basal molar, notch and straight beyond.

Minor cheliped, palm upper margin as long as greatest width;

distolateral margin with sharp spine at base of dactylus; lateral

and mesial faces tuberculate near base of fingers', fingers almost

as long to longer than upper margin of palm.

Description, (based on MNHN IU-2014-10474, male, 9.1 mm).

Carapace smooth. Rostrum 0.2 carapace length, concave

dorsally, tapering evenly to acute tip, twice as long as wide at

base, with c. 7 teeth on lateral margins, depressed below level

of median carina, separated from lateral carinae by long hiatus,

without ventral tooth. Lateral gastric carina visible only

posterior to confluence of submedian carinae, short, unarmed.

Submedian gastric carina smooth, together curved and

converging on median carina. Median carina prominent, erect,

reaching two-thirds along rostrum, with c. 12 uneven teeth.

Pleuron 1 posteroventrally rounded; pleura 2,3 posteroventrally

produced, acute; pleura 4, 5 less produced, posteroventrally

acute; pleura 3, 5 with anteroventral tooth; pleuron 6 with

rounded posteroventral angle; pleonite 6 dorsal posterior

margin denticulate.

Eyestalk third length of rostrum; cornea unpigmented.

Antennular peduncle reaching to midpoint of antennal article 4;

article 1 unarmed. Antennal article 1 unarmed; article 2 without

distomesial spine, stylocerite an elongate triangular blade,

reaching just beyond midpoint of article 4; scaphocerite a

vertical blade, reaching midpoint of article 5; article 3 lower

margin with mesial tooth; article 5 about half length of article 4.

Major cheliped ischium lower margin produced as spinose

ridge; merus lower margin finely denticulate, upper margin

with minute tooth; carpus lower margin with 1 distal tooth;

propodus upper margin carinate, denticulate, toothed distally,

length 1.25 greatest height, lower margin smooth, lateral face

tuberculate on lower palm, mesial face tuberculate at base of

fixed finger; fixed finger half as long as upper palm, cutting

edge concave with proximal and subdistal obtuse teeth, with

longitudinal mesial ridge; distolateral margin of palm

denticulate, with blunt tooth above circular gape; distomesial

margin of palm denticulate, angled tooth near gape; dactylus

distally curved, cutting edge with blunt proximal tooth,

otherwise straight, smooth.

Minor cheliped shorter and more slender than major, palm

0.9 times height of major palm; ischium lower margin with 2

subdistal spines; merus lower margin denticulate, most distal

spine-like; carpus lower margin with 1 distal tooth; propodus

dilating, upper margin carinate, denticulate, 0.95 times

greatest height, lower margin smooth, lateral and mesial faces

tuberculate at base of fixed finger; fixed finger as long as upper

palm, cutting edge denticulate, with longitudinal mesial and

lateral ridges; distolateral margin of palm oblique, with sharp

spine in gape; distomesial margin of palm oblique, with spine

in gape; dactylus almost straight, upper margin carinate,

cutting edge smooth, with subdistal notch.

Pereopod 2 ischium lower margin unarmed; merus lower

margin unarmed; carpus 0.85 length of propodus upper

margin; propodus upper margin twice as long as dactylus.

Pereopod 3 merus unarmed; propodus 3.0 times as long as

dactylus, with 9 rows of spiniform setae, of 2-4 setae; dactylus

spatulate, with 7 spiniform setae along oblique margin, plus

unguis, 3 facial spiniform setae. Pereopod 4 virtually identical

to pereopod 3. Pereopod 5 propodus with 8 rows of spiniform

setae; dactylus spatulate, with 6 spiniform setae along oblique

margin, plus unguis, 6 facial spiniform setae.

Uropodal endopod 1.6 times as long as wide, elongate-

oval, anterior-distal margin with 19 evenly-spaced teeth, last 3

teeth distal, without longitudinal ridge. Uropodal exopod 1.4

times as long as wide, oval, anterior margin with c. 30 small

teeth over most of length, without longitudinal rib.

Telson 1.1 times as long as wide, widest at mid-length,

tapering to square posterolateral angles, lateral margin

upturned, denticulate, distal margin concave each side of

posteromedian spine; dorsal face smooth.

Distribution. Indonesia, Solomon Is, New Caledonia, Vanuatu;

418-984 m.

Remarks. The holotype, photographed for us by Charles

Fransen, clearly shows the uneven row of c. 15 teeth on the

median carina, the short separate lateral carina on the right

side, two prominent teeth on the distomesial margin of the

propodus of the minor cheliped, the semi-enclosed notch at the

base of the fixed finger with two irregular teeth above, and the

obscurely denticulate blade on the distomesial margin of the

propodus of the major cheliped. The marginal denticulation of

the fingers and the tuberculation of the propodus are consistent

with the present material. Sakai’s (2011) figure 52 is consistent

with this except for the absence of the lateral carina.

The circular notch in the gape of the propodus of the major

cheliped is similar to that in E. dongshaensis sp. nov. but not as

enclosing; E. indicus lacks the ventral rostral tooth seen in the

new species.

The lateral gastric carina may be obvious (Fig. 10a) but

typically displays a strong hiatus between the lateral rostral

ridge (Fig. 9a-c) and is sometimes absent, especially in larger

specimens (Fig. lOg). In males, the circular notch in the gape

of the major cheliped sometimes appears almost keyhole¬

shaped. The specimens from New Caledonia exhibit some

variability that includes that of the Vanuatu material. The

telson has the same general appearance but ranges from 1.1 to

1.3 times as long as wide, males being broader than females

(cf. Figs 9d, lOi, j with 10b, k, 1). The armature of the

distolateral margins of the chelipeds also varies, some more

spinose than others.

Acknowledgements

The MNHN collections derived from six expeditions. The

Program Tropical Deep-Sea Benthos expeditions in the south¬

western Pacific were headed by Bertrand Richer de Forges

(BATHUS 3, BOAO, SALOMON) and Sarah Samadi (BOA1).

The BIOMAGLO cruise was conducted by MNHN (Pis Laure

Corbari, Sarah Samadi, Karine Olu). The KARUBENTHOS

cruise was conducted by MNHN (Pis Laure Corbari, Philippe

Bouchet), in conjunction with the National Park of Guadeloupe

and Universite des Antilles et de la Guyane, with support from

Institut Ecologie et Environnement (INEE) of the French

The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae)

119

Research Council (CNRS) and the AGOA Marine Sanctuary.

The Pis also acknowledge the UMS Flotte Oceanographique

Framjaise, Genavir and Institut de Recherche pour le

Developpement (IRD) for deploying the Research Vessel

Anted. We are grateful to Laure Corbari, Paula Lefevre-Martin

and Anouchka Krygelmans-Sato for help in making these

collections available at MNHN.

The NERC ‘James Cook’ cruise JC066 on the SW Indian

Ridge (funded through NERC Grant NE/F005504/1, PI A.D.

Rogers) was part of the Southwest Indian Ocean Seamounts

Project, supported by the EAF Nansen Project, the Food and

Agriculture Organization of the United Nations, the Global

Environment Facility and the International Union for the

Conservation of Nature. Our thanks are extended to the crew

and scientists involved in the cruise and to Sammy De Grave

(Oxford University Museum of Natural History) for making

the collection available for study.

We are grateful to Adam Baldinger, Museum of

Comparative Zoology, Harvard University, Cambridge, for

the loan of the syntypes of Eiconaxius antillensis, to C.H.J.M.

Fransen, Naturalis Biodiversity Center, Leiden, for

photographs of the holotype of Iconaxius crista-galli var.

indica and to Tin-Yam Chan, National Taiwan Ocean

University, Taiwan, for the loan of specimens from his

collections from the South China Sea.

We thank Tin-Yam Chan and David Shale who provided

photographs and Christopher C. Tudge, American University,

for providing illustrations of specimens in the USNM.

Finally, GCBP acknowledges the support of Philippe

Bouchet, MNHN, and the Crosnier Fund for financial support

in Paris.

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