Memoirs of Museum Victoria 81: 1-23 (2022) Published 2022
1447-2594 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2022.81 .01
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae)
from Australia
(http://zoobank.org/urn:|sid:zoobank.org:pub:53 DA FE4A-6C49-42A0-9D44-F37BO3FBAT7FO)
CHUNG CHENG LU” AND TAKASHI OKUTANT
! Museum Victoria, Melbourne, Victoria 3000, Australia
* Department of Life Sciences, National Chung Hsing University, Taichung, Taiwan 402
* #714, Takaishi 4-17-1, Asao-ku, Kawasaki, Japan 215-0003
* ‘To whom correspondence should be addressed. Email: cclu@nchu.edu.tw
Two new genera and species of sepioline squid (Cephalopoda: Sepiolidae) are described from Australian waters.
Dextrasepiola taenia ıs characterised by having copulatory organs (1.e. the hectocotylised arm ın the males and the bursa
copulatrix in the females) 1n the right side of the body. All other known sepiolinids have copulatory organs in the left side
of the body. Amutatiola macroventosa 1s characterised by the absence of a hectcotylised arm in mature males; instead, it
possesses many enormously enlarged suckers on some of the arms of the males. The bursa copulatrix is in the left side of
the female body, as in other known sepioline squids. The discovery of these two new taxa indicates that the present
Abstract Lu, C.C., and Okutanı, T. 2022. Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from
Australia. Memoirs of Museum Victoria 81: 1—23.
definition of Sepiolinae needs to be broadened to accommodate these two new genera.
Keywords Sepiolinae; Dextrasepiola taenia, Amutatiola macroventosa; Australia.
Introduction
Members of the family Sepiolidae are small to medium size
benthic or pelagic cephalopods and are common in tropical,
subtropical and temperate waters. Currently, 18 genera are
recognised in the following three subfamilies: (1)
Heteroteuthinae: Heteroteuthis Gray, 1849, Stoloteuthis Verrill,
1881, Nectoteuthis Verrill, 1883, /ridoteuthis Naef, 1912, and
Sepiolina Naef, 1912; (11) Sepiolinae: Sepiola Leach, 1817,
Inioteuthis Verrill, 1881, Euprymna Steenstrup, 1887,
Rondeletiola Naef, 1921, Sepietta Naef, 1912, Adinaefiola
Bello, 2020, Boletzkyola Bello, 2020, Lusepiola Bello, 2020,
and Eumandya Bello, 2020; and (111) Rossiinae: Rossia Owen,
1834, Semirossia Steenstrup, 1881, Austrorossia Berry, 1918,
and Neorossia Boletzky, 1971. For Chonneteuthis Lu and
Boucher-Rodoni, 2006, the subfamilial placement is uncertain
(Bello, 2020; Lu and Boucher-Rodoni, 2006; Nesis 1987). The
three subfamilies are well defined. Table 1 shows the diagnostic
characters of these subfamilies.
A member of Sepiolinae was first recorded from Australia
by Lu and Phillips (1985) and assigned to the old genus Sepiola
without specific identification. A dorsal view of a specimen
was also presented in Lu and Dunning (1998). The record was
accidentally omitted in Lu (2001). In this paper, we describe
two new sepioline species and placed them in two new genera
in light of Bello’s (2020) revision. These new genera and
species appear to belong to Sepiolinae based on the main
characters of this subfamily, apart from their peculiar male
arm modifications that do not correspond to the normal
sepioline hectocotylisation.
Material and methods
All materials examined are listed under each relevant taxon. All
except one lot are in the collection of the Museum of Victoria.
The materials were all formalin fixed and preserved in 70%
ethyl alcohol. All measurements and indices are standard
teuthological measurements and indices, following Roper and
Voss (1983) and Lu and Boucher-Rodoni (2006), except for
nuchal commissure index, which is the width of nuchal
commissure expressed as a percentage of the width of mantle at
the position of nuchal commissure. Nuchal commissure width is
called occipital band width by some authors (e.g. Nesis, 1982).
Roper and Voss (1983) used the mantle length as the standard
reference size for free funnel index and nuchal commissure
index. For arm lengths, arm sucker counts and sucker diameters,
left arms were measured or counted. In males, the right arm I
was also measured and suckers and sucker stalks counted to
ascertain 1f any modification occurs ın right arm I. The maturity
stages used in the Tables are those used in Lu and Roper (1979).
2 C.C. Lu & T. Okutani
Table 1. Comparison of distinguishing characters of subfamilies Heteroteuthinae, Rossiinae and Sepiolinae of the family Sepiolidae (after Nesis, 1982)
L0 |RHeteroteuthinae
not extended forward, not covering
funnel
not extended forward, not covering
funnel
Anterior ventral mantle edge | extended into projecting ventral
shield covering funnel from below
and sometimes reaching level of eyes
or farther forward; ventral shield
with an incision in centre for funnel
opening
Anterior dorsal mantle edge | fused with head, except in not fused with head fused with head
Heteroteuthis
Nuchal cartilage developed in Heteroteuthis; absent in | developed no nuchal cartilage
other genera
Arms connected by web 3 dorsal pairs of arms joined by deep | 2 dorsal pairs of arms not connected | 2 dorsal pairs of arms not connected
web or connected only by shallow web or connected only by shallow web
Arm suckers in 2 series; some suckers on lateral
arms in males in some taxa greatly
enlarged
Club narrow or slightly widened, with
very small suckers
Fins large, fin length 60-100% of mantle
length
Photophores on ventral side of ink sac
Hectocotylisation usually both dorsal arms
hectocotylised
present
Habitat benthic or pelagic benthic, some species ascend to
surface during reproduction
Abbreviations:
CSIRO: Commonwealth Scientific and Industry Research
Organisation of Australia
IYGPT: International Young Gadoid Pelagic Trawl
MOV: Museum of Victoria, Melbourne, Victoria, Australia
Q: FRV Courageous of CSIRO
Q47/51: FRV Courageous of CSIRO cruise 47, station 51
SO: FRV Soela of CSIRO
FRV Soela SO1/85/124 (and similar): FRV Soela of CSIRO,
1985 cruise 1, station 124
Systematic description
Class Cephalopoda Cuvier, 1797
Subclass Coleoidea Bather, 1888
Order Sepiolida Fioroni, 1981
Family Sepiolidae Leach, 1817
Subfamily Sepiolinae Leach, 1817
Dextrasepiola n. gen.
Type Species: Dextrasepiola taenia n. gen.,n. sp. by monotypy
Diagnosis: Sepiolinae with fins rounded with large anterior
lobes, which do not reach the anterior mantle margin; fin length
in 2 or 4 series; no greatly enlarged
suckers
narrow or widened, with very small,
medium or large suckers
moderate size, much shorter than
mantle
only one genus, Semirossia, 1s
known to have a bilobed photophore
on ink sac
left or both dorsal arms left dorsal arm hectocotylised
rn
in 2 or 4 series; some suckers in
males maybe enlarged
narrow or slightly widened, with
small suckers
moderate size, much shorter than
mantle
saddle-shaped, two ear-shaped,
lens-shaped or absent (/nioteuthis)
‘rudimentary orabsent — or absent
benthic, some species ascend to
surface during reproduction
about 50-80% mantle length. Suckers biseriate on all arms.
Tentacular club suckers in 4—8 longitudinal series. Nuchal
commissure narrow, not reaching over the ocular globes. A pair
of dumbbell-shaped photophores on ventral surface of ink sac.
Gladius absent. Ventral mantle margin slightly sinuate, without
any deep funnel indentation. Right arm I of male hectocotylised.
Hectocotylus tripartite: basal part with five suckers in two
series, two suckers 1n dorsal series and three in ventral series;
copulatory apparatus long, fleshy, tape like, formed by fusion of
two adjoining very elongate sucker stalks, no additional
modified structure on the arm (1.e. hook-like stalks); distal to
tape-like copulatory apparatus biserial suckers to arm tip (19
suckers in holotype, 23 suckers in paratype 2). Female bursa
copulatrix on right side of mantle cavity, open type (cf. Bello,
2020), roughly ear shaped.
Etymology: Generic name Dextrasepiola 1s derived from Latin
dextra meaning right or on the right side plus sepiola meaning
a small cuttlefish. The name denotes the unique feature among
the Sepiola and its kin having the right dorsal arm hectocotylised
instead of the left dorsal one in mature males and the bursa
copulatrix in females on the right side of the mantle cavity.
Remarks: The hectocotylisation in Sepiolinae has been
thoroughly discussed and illustrated by Bello (2020), Naef
(1912a, b; 1923) and Nesis (1982). All known species have the
left dorsal arm hectocotylised and the sucker stalks of some
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) trom Australia 3
suckers are modified into a horn-like or hook-like copulatory
organ. The present genus is unique in having the right dorsal
arm hectocotylised and the copulatory organ as one thick tape-
like structure. In addition, the hectocotylus has the regular
tripartite structure typical of most sepioline genera except for
Euprymna and Eumandya (cf. Bello, 2020). The females of all
known species of Sepiolinae possess a bursa copulatrix on the
left side of the mantle cavity. The present genus 1s unique in
having the bursa copulatrix on the right side of the mantle
cavity. This right-handedness of the copulatory organs of both
sexes of this genus certainly facilitate copulation in a normal
fashion as they are on the same side of the animal.
Dextrasepiola taenia n.gen., n.sp.
Figures 1—5, 13; Table 2
Material examined: Holotype: MOV F80458: Redland Bay,
Queensland, 27° 36' S, 153° 19' E, 1.2 m, CSIRO Moreton Bay Survey,
J43, Location 31, 10 Aug 1951, 1 male, mature, 8.2 mm mantle length
(specimen #1).
Paratype 1: MOV F91359: Redland Bay, Queensland, 27° 36' S,
153° 19' E, 1.2 m, CSIRO Moreton Bay Survey, J43, Location 31, 10
Aug 1951, 1 female, mature, 7.8 mm mantle length (specimen #2).
Paratype 2: MOV F74469: Peel Island, Queensland, 27° 30' S,
153° 21' E, 1.2 m, CSIRO Moreton Bay Survey, J26, Location 41, 10
Aug 1951, 1 male, mature, 6.5 mm mantle length (specimen #3).
Paratype 3: MOV F91361: Peel Island, Queensland, 27° 30' S,
153° 21' E, 1.2 m, CSIRO Moreton Bay Survey, J26, Location 41, 10
Aug 1951, 1 female, mature, 8.7 mm mantle length (specimen #4).
Other material: MOV F91360: Peel Island, Queensland, 27° 30' S,
153° 21' E, 1.2 m, CSIRO Moreton Bay Survey, J26, Location 41, 10
Aug 1951, 2 specimens, poor condition, 1 female, juvenile, 5.4 mm
mantle length, 1 specimen, sex indeterminate, 3.9 mm mantle length.
Diagnosis: Small sepioline with right arm I of mature male
hectocotylised, copulatory apparatus in the form of a long tape-
like process, no hook-like structure on the arm. Females with
bursa copulatrix on right side of mantle cavity.
Description: Mantle (figs la—c) short dome-shaped, slightly
longer than wide, fused with head dorsally for about 25% of
width. Anterior ventral mantle margin shallowly concave with
lateral projections at position of mantle-funnel connectives.
Head wide, slightly narrower than mantle. Nuchal commissure
narrow, not reaching beyond level of medial border of eyeball,
approximately 25-40% of mantle width at level of nuchal
commissure. Eyes large, elliptical, located dorso-laterally on
head. Cornea membrane protecting eye attached to skin of
head along dorsal margin. Olfactory papilla located behind
posterior corner of eye orbit, ventral photosensitive vesicle not
found. Funnel long and slender, reaching the level beyond
anterior eye margin, free from head for 60-80% of funnel
length. Funnel connects to head by an oblique muscle band
extending from beneath anterior end of funnel locking cartilage
to ventral side of head.
Funnel locking cartilage (fig. If) elongated oval with
simple, slightly curved depression in the middle, mantle locking
cartilage long, low ridge. Dorsal element of funnel organ (fig.
lg) Y-shaped pad with a small papilla at the apex. Behind each
ramus is swelling that connects dorsal funnel organ with base
of funnel retractor. Ventral elements of funnel organ (fig. 1g) a
pronounced semi-spherical pad, becoming slightly narrower
anteriorly, with a mamillar projection slightly posteriorly to
centre. Funnel valve well developed on dorsal roof, tongue
shaped, located well behind funnel aperture.
Fins (figs la—c) circular in outline, anterior border of fins
projects forward prominently forming a deep cleft with
mantle, anterior fin lobe reaching level halfway between fin
insertion and mantle border or to mantle margin, posterior
borders of fins convex, less pronounced. Length of fin base
about 33% of mantle length.
Arms (figs la—c, 2a—c, 3) short, rounded aborally, flatten
orally. All arms on both sexes with biserial suckers throughout.
Because most suckers are lost, it was impossible to determine
the sucker ring dentition and if the enlargement of suckers
exists. In males, arm III is the longest and thickest, followed by
arm Il or arm IV. Right arm I of male (figs Id—e, 3) with a flat,
fleshy, tape-like long process, appeared to be modified sucker
stalks of proximal third sucker of dorsal series and proximal
fourth sucker of ventral series fused together throughout their
length. Length of the process reaches to almost the level of
arm tip, thickened along proximal portion ending in a blunt tip.
No sucker on the tape-like process of holotype, but a remnant
of a tiny sucker on the process of paratype 2 (fig. 3). Distal to
the tape-like process, 19 suckers in two series to arm tip on
holotype (23 suckers in paratype 2); no hook-like structures on
the arm. Left arm I in males and both arms I in females with
28-30 suckers in two series, with no peculiar or unusual
development or modification. Arm II with 27-35 suckers in
both sexes with no noticeable special development in either
sex. In males, arms III thickened proximally, slightly tapers
distally to about half of arm length then abruptly tapers
distally; swollen proximal part with no suckers except several
remnants of suckers; distal part strongly curled towards mouth,
with 17-19 suckers. In females, arm III similar to arm II with
2] suckers. Arm IV of both sexes with 26-31 suckers. In
males, aboral keel and swimming membrane absent on arms
I-III, well developed along whole length of arms IV. In
females, aboral keel and swimming membrane present on
distal half of arms I and II, and almost whole length of arms
III and IV. Webs shallow between all arms except between
arms III and IV (web D), which half encloses base of tentacles
in both sexes, web E non-existent.
Tentacles weak, longer than arms. Club (fig. 4a) slightly
expanded, minute carpal suckers in 4 series, minute manal
suckers in 6 series, those on dorsal 2 series larger than the
remaining suckers, numerous minute suckers in 8 series on
dactylus. Sucker ring dentition of largest club sucker finely
toothed around entire minute circle.
Gills with 15—20 lamellae per demibranch, plus a terminal
lamella. A pair of dumbbell-shaped, yellowish photophores,
Opaque with both ends swollen on both sides of ink sac (figs
lg, 4b).
Upper beak (fig. 4d) rostrum slightly curved; jaw angle
obtuse; wing long, shoulder (cutting edge) serrated; rostrum
dark brown to black, hood, shoulder and dorsal part of lateral
wall light brown, posterior part of hood, most of lateral wall
unpigmented, transparent.
4 C.C. Lu & T. Okutani
lable 2. Dextrasepiola taenia n.gen. & n.sp. measurement, counts and indices
S ;»ecimen number i 2»
female
22.06, D.20.2; C:OU. ^, G.94./, 0.599. Se
NidGI AO G «—-. ———
Note: Maturity 4, subadult — sexual characters well distinguished but gonads and accessory organs not completely developed; Maturity 5, adult — sexually mature with
spermatophores in Needham’s sac in males; ovaries, nidamental and oviducal glands fully developed and ripe, eggs sometimes in oviducts, in females; DML, dorsal
mantle length (mm) — measured from anterior-most point of mantle to posterior end of mantle; VMLI, ventral mantle length index — ventral mantle length, measured
from anterior border of mantle at ventral midline, to apex of mantle, expressed as a percentage of DML; HLI, head length index — dorsal length of head measured from
point of fusion of dorsal arms to anterior tip of dorsal mantle expressed as a percentage of DML; HWI, head width index — greatest width of head at level of eyes
expressed as a percentage of DML; NCI, nuchal commissure width index — width of nuchal commissure expressed as a percentage of width of mantle at the position of
nuchal commissure; MWI, mantle width index — mantle width at mantle opening expressed as a percentage of DML; FuLI, funnel length index — length of funnel from
anterior funnel opening to posterior border of funnel measured along ventral midline expressed as a percentage of DML; FuWI, funnel width index — width of funnel at
junction of funnel and head just anterior to funnel locking cartilages expressed as a percentage of DML; FFul, free funnel index — length of funnel from the anterior
opening to the point of dorsal attachment to head expressed as a percentage of funnel length; FLI, fin length index — greatest length of a single fin expressed as a
percentage of DML; FBLI, fin base length index — length of fin attachment to mantle expressed as a percentage of DML; FWI, fin width index — greatest width of a
single fin expressed as a percentage of DML; A LI, arm I length index — length of arm I expressed as a percentage of DML; A,LI, arm II length index — length of arm II
expressed as a percentage of DML; A,LI, arm ri length index — length of arm III expressed as a percentage of DML; A LI, arm IV length index — length of arm IV
expressed as a percentage of DML; A. LI, right arm I length index - length of right arm I expressed as a percentage of DML (only in males); TtLI, tentacle length index
— total length of tentacular stalk and club e expressed as a percentage of DML; CILI, club length index — length of club, measured from proximal-most basal sucker to
distal tip of club expressed as a percentage of DML; CIRC, club row count — number of longitudinal rows of suckers across width of club; GLI, gill length index —
length of gill expressed as a percentage of DML; GWI, gill width index — greatest width of gill expressed as a percentage of DML; GiLC, gill lamellae count — number
of lamellae on outer demibranch, excluding terminal lamella; A SC, sucker counts of arm I — total number of suckers or sucker stalks on arm I; A,SC, sucker counts of
arm II — total number of suckers or sucker stalks on arm II; A, SC, sucker counts of arm III — total number of suckers or sucker stalks on arm III; Á SC, sucker counts of
arm IV — total number of suckers or sucker stalks on arm IV; A, SC, sucker counts of right arm I — total number of suckers or sucker stalks on right arm I (only in
males); CISI, club sucker index — diameter of largest club sucker expressed as a percentage of DML; A SI, arm I sucker index — diameter of largest sucker on arm I
expressed as a percentage of DML; ASI, arm II sucker index — diameter of largest sucker on arm I] expressed as a percentage of DML; A,SI, arm III sucker index —
diameter of largest sucker on arm III expressed as a percentage of DML; A 5I, arm IV sucker index — diameter of largest sucker on arm IV expressed as a percentage of
DML; SpLI, spermatophore length index — length of spermatophore expressed as a percentage of DML; SpWI, spermatophore width index — greatest width of
spermatophore expressed as a percentage of spermatophore length; SpRI, sperm reservoir index — length of sperm reservoir expressed as a percentage of spermatophore
length; CBI, cement body index — length of cement body expressed as a percentage of spermatophore length; NidGI, nidamental gland index — length of nidamental
gland expressed as a percentage of DML.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia 5
Lower beak (figs 4e, f) wide; rostrum with blunt tip; jaw
edge rough, slightly serrated, jaw angle indistinct; blunt tooth
on shoulder; no notch in hood; lateral wall without fold or ridge,
roughly elongate rhomboidal with lower edge slightly concave,
corner faintly produced; rostrum and hood light brown,
posterior part of lateral wall and wings transparent. Radula (fig.
4c) seven series, each row with seven unicuspid teeth.
Gladius absent.
Spermatophores (figs 5a, b) small, five spermatophores from
the holotype approximately 2-2.5 mm long (spermatophore
length index 24-30), greatest width approximately 0.08—0.1 mm
(spermatophore width index 3.4—4.8), sperm reservoir about
0.4—0.5mm (sperm reservoir index 19-22), structure complex,
with spiral appearance in ejaculatory apparatus and the sperm
mass, cement body 0.72—0.78 mm long (cement body index
30-36), connects to sperm reservoir by a narrow neck, oral end
of cement body elongated funnel shaped.
Bursa copulatrix (figs 5c-f ) open type, large, roughly ear-
shaped, longer than wide; anteriorly extends medially towards
midline, running antero-laterally just below right funnel
locking cartilage, reaching posterior end of mantle cavity.
Longitudinal opening of bursa close to mantle ventral midline,
Figure 1. Dextrasepiola taenia n. gen. & n. sp., male: a, dorsal view, holotype (MOV F80458); b, ventral view (MOV F80458); c, lateral view (MOV
F80458); d, hectocotylised arm (right arm I; MOV F80458); e, diagram of the hectocotylised arm showing tape-like modification, paratype 2 (MOV
F74469; drawn by T. Okutanı); f, inside mantle cavity, showing photophores (p) and other organs (MOV F80458); e, diagram of opened funnel and
inside mantle cavity showing dorsal funnel organ (dfo), ventral funnel organ (vfo) and photophores (p; MOV F80458; drawn by T. Okutani).
6 C.C. Lu & T. Okutani
Figure 2. Dextrasepiola taenia n. gen. & n. sp., female, paratype 1 (MOV F91359):
showing sucker arrangements of female.
E-
Figure 3. Dextrasepiola taenia n. gen. & n. sp. male, paratype 2 (MOV F74469). Oral view of arms (grids in background: 1 mm x 1 mm): a, right arm
I, arrow points to remnant of a tiny sucker; b, right arm II; c, right arm III; d, right arm IV; e, left arm I; f, left arm II; g, left arm III; h, left arm IV.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia T
running along long axis of bursa. Mature females with large
nidamental gland, (nidamental gland index 36.6-54.4).
Alcohol-preserved specimens brown in colour, dorsal
mantle surface lighter than ventral surface. Dark chocolate-
brown chromatophores scattered over brown-coloured
background on both dorsal and ventral surfaces of head and
mantle, and along aboral surface of all arms. Surfaces of fins
devoid of chromatophores and pigmented spots. Skin smooth,
lacking sculpture or papillae.
Etymology: Species epithet taenia from Latin taenia meaning
tape-like. The name denotes the tape-like structure in the
copulatory apparatus on the hectocotylised arm of the males.
Distribution: Only known from Moreton Bay, Queensland,
Australia (fig. 13).
Remarks: This is the only known species in the genus. Due to the
poor preservation of the specimens, nearly all suckers are lost or
are without sucker rings. The description of sucker ring dentition
and spermatophores must wait until better materials are available.
The poor state of preservation resulted in distorted
morphology of the specimens studied. This most certainly
contributes to the wide range of the morphometric indices
listed in Table 2.
Amutatiola n. gen.
Diagnosis: Small Sepiolinae with fins rounded with large
anterior lobe, which do not reach the anterior mantle margin;
fin length about 40—66% mantle length. Suckers biseriate on all
arms. Tentacular club suckers in 4-8 longitudinal series.
Nuchal commissure moderately wide, not reaching over the
ocular globes, about 38-59% of mantle width. A pair of
dumbbell-shaped or elongated kidney-shaped photophores on
ventral surface of ink sac. Gladius absent. Ventral mantle
margin slightly sinuate, without any deep funnel indentation.
No arm in mature males hectocotylised. Some arm suckers in
mature males grossly enlarged. Female bursa copulatrix closed
type, pouch-like, opening at level of base of left gill.
Type Species: Amutatiola macroventosa n. gen., n. sp. by
monotypy.
Etymology: Generic name Amutatiola 1s derived from Greek a
meaning without or absent plus Latin mutatus meaning
changed or altered; the ending -ola is the diminutive suffix of
sepiola meaning a small cuttlefish. The name denotes the
unique feature among the Sepiolinae of having no arm
hectocotylised 1n males.
b, inside mantle cavity: photophores (p), left gill (lg), male, holotype (MOV F80458); c, radula (paratype 1, MOV F91359); d, lateral view of upper
beak (MOV F91361); e, lateral view of lower beak (MOV F91361); f, top view of lower beak (MOV F91361).
C.C. Lu & T. Okutani
Figure 5. Dextrasepiola taenia n. gen. & n. sp. (grids in background: 1 mm x 1 mm): a, spermatophore from the holotype (MOV F80458); b,
close-up of cement body; c, bursa copulatrix, paratype 1 (MOV F31359; bc: bursa copulatrix; rg: right gill; rsf: right side of funnel); d, bursa
copulatrix, juvenile (MOV F31360; bc: bursa copulatrıx; Iflc: left funnel locking cartilage; rflc: right funnel locking cartilage; rg: right gill); e,
bursa copulatrix, paratype 3 (MOV F31361; bc: bursa copulatrix; Isf: left side of funnel; ms: mantle septum; rsf: right side of funnel); f, close-up
of a portion of e, arrow points to opening of bursa copulatrix.
Amutatiola macroventosa n. gen., n. sp.
Figures 6—13; Tables 3, 4
Material examined: Holotype: MOV F80081: south-east Tasmania,
42° 38.1'S, 148° 12.4' E, trawl depth 36-42 m, bottom depth 86-90 m,
collected by CSIRO, FRV Soela SO1/85/124, 27 Feb 1985, 0331 hr,
Rectangular Midwater Trawl with 8 m^ mouth area, 1 male, 9.8 mm
mantle length, mature (specimen 71).
Paratype 1: MOV F275293: Great Australian Bight, 32° 43' S,
126° 00' E-32° 45' S, 125° 59' E, 40-170 m, collected by CSIRO, FRV
Soela SO3/80/32, 13 May 1980, 0100 hr, IYGPT, 1 female, 12.2 mm
mantle length, mature (specimen #11).
Paratype 2: WAM 3091-83: west side of Irwin Reef, Port Denison,
Western Australia, 29° 16'S, 114° 55' E; 7-8 m, collected by N. Sinclair,
4 Apr 1983, Rotenone Station, | male, 9.5 mm mantle length, mature
(specimen #3).
Paratype 3: MOV F80083: south-east Tasmania, 42° 39.7' S,
148° 12.1' E, trawl depth 5-10 m, bottom depth 90-95 m, collected by
CSIRO, FRV Soela SO1/85/104, 15 Feb 1985, 2034 hr, Rectangular
Midwater Trawl with 8 m* mouth area, | male, 9.6 mm mantle length,
mature (specimen #2).
Paratype 4: MOV F275294: Great Australian Bight, 33° 30' S,
131° 50.0' E-33° 30'S, 131° 53' E, 200-144 m, collected by FRV Soela
SO3/80/1, 8 May 1980, 1 female, 12.8 mm mantle length, mature
(specimen #9).
Paratype 5: MOV F158244: Luck Bay, western point off beach,
Cape Le Grand National Park, Western Australia, 33° 59' S, 122° 13' E,
5 m, active over algae, collected by D. Rawlins, J. Finn and M. Norman,
26 April 1998, 1915 hr, hand net, 1 male, 8.8 mm mantle length, mature
(specimen #4).
Paratype 6: MOV F275296: Luck Bay, western point off beach,
Cape Le Grand National Park, Western Australia, 33° 59' S, 122? 13' E,
5 m, active over algae, collected by D. Rawlins, J. Finn and M. Norman,
26 April 1998, 1915 hr, hand net, 1 female, 6.2 mm mantle length,
subadult (specimen #17).
Paratype 7: MOV F275295: Luck Bay, western point off beach,
Cape Le Grand National Park, Western Australia, 33° 59' S, 122° 13' E,
5m, active over algae, collect. by D. Rawlins, J. Finn and M. Norman, 26
April 1998, 1915 hr, hand net, 1 male, 8.1 mm mantle length, mature
(specimen #5).
Paratype 8: MOV F91362: Luck Bay, western point off beach, Cape
Le Grand National Park, Western Australia, 33° 59' S, 122? 13' E, 5 m,
active over algae, collected by D. Rawlins, J. Finn and M. Norman, 26
April 1998, 1915 hr, hand net, 1 male, 6.9 mm mantle length, mature
(specimen 77).
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) trom Australia 9
Table 3. Amutatiola macroventosa n.gen. & n.sp. measurement, counts and indices of male specimens
Amutatiola macroventosa
a er a ooy
eaa pops apei M
Ce n —
Station.
Maud SS
PML (mm) — P8 —X— — "— ——
"——T— — —— — wes Mos —
1. — — as — — — d — 9 — — H3 —— 3498 —Àx — HB ——
86L— — — 8 — — — 31888 — — 88 — — HE 383 E — 5 ———
=
Note: Maturity 4, subadult — sexual characters well distinguished but gonads and accessory organs not completely developed; Maturity 5, adult — sexually mature with
spermatophores in Needham’s sac in males; ovaries, nidamental and oviducal glands fully developed and ripe, eggs sometimes in oviducts, in females; DML, dorsal
mantle length (mm) — measured from anterior-most point of mantle to posterior end of mantle; VMLI, ventral mantle length index — ventral mantle length, measured
from anterior border of mantle at ventral midline, to apex of mantle, expressed as a percentage of DML; HLI, head length index — dorsal length of head measured from
point of fusion of dorsal arms to anterior tip of dorsal mantle expressed as a percentage of DML; HWI, head width index — greatest width of head at level of eyes
expressed as a percentage of DML; NCI, nuchal commissure width index — width of nuchal commissure expressed as a percentage of width of mantle at the position of
nuchal commissure; MWI, mantle width index — mantle width at mantle opening expressed as a percentage of DML; FuLI, funnel length index — length of funnel from
anterior funnel opening to posterior border of funnel measured along ventral midline expressed as a percentage of DML; FuWI, funnel width index — width of funnel at
junction of funnel and head just anterior to funnel locking cartilages expressed as a percentage of DML; FFul, free funnel index — length of funnel from the anterior
opening to the point of dorsal attachment to head expressed as a percentage of funnel length; FLI, fin length index — greatest length of a single fin expressed as a
percentage of DML; FBLI, fin base length index — length of fin attachment to mantle expressed as a percentage of DML; FWI, fin width index — greatest width of a
single fin expressed as a percentage of DML; A LI, arm I length index - length of arm I expressed as a percentage of DML: A „LI, arm II length index — length of arm II
expressed as a percentage of DML; A,LI, arm IIT le length index - length of arm Ill expressed as a percentage of DML; A LI, arm IV length index — length of arm I'V
expressed as a percentage of DML; A. LI, right arm I length index — length of right arm I expressed as a percentage of DML (only in males); TtLI, tentacle length index
— total length of tentacular stalk and club e expressed as a percentage of DML; CILI, club length index — length of club, measured from proximal-most basal sucker to
distal tip of club expressed as a percentage of DML; CIRC, club row count — number of longitudinal rows of suckers across width of club; GLI, gill length index —
length of gill expressed as a percentage of DML; GWI, gill width index — greatest width of gill expressed as a percentage of DML; GiLC, gill lamellae count — number
of lamellae on outer demibranch, excluding terminal lamella; A, SC, sucker counts of arm I — total number of suckers or sucker stalks on arm I; A, SC, sucker counts of
arm II — total number of suckers or sucker stalks on arm II; A, SC, sucker counts of arm III — total number of suckers or sucker stalks on arm III; Á SC, sucker counts of
arm IV - total number of suckers or sucker stalks on arm IV: A, SC, sucker counts of right arm I — total number of suckers or sucker stalks on right arm I (only in
males); CISI, club sucker index — diameter of largest club sucker expressed as a percentage of DML; A SI, arm I sucker index — diameter of largest sucker on arm I
expressed as a percentage of DML; ASI, arm II sucker index — diameter of largest sucker on arm II expressed as a percentage of DML; ASI, arm III sucker index —
diameter of largest sucker on arm III expressed as a percentage of DML; A SI, arm IV sucker index — diameter of largest sucker on arm IV expressed as a percentage of
DML; SpLI, spermatophore length index — length of spermatophore expressed as a percentage of DML; SpWI, spermatophore width index — greatest width of
spermatophore expressed as a percentage of spermatophore length; SpRI, sperm reservoir index — length of sperm reservoir expressed as a percentage of spermatophore
length; CBI, cement body index — length of cement body expressed as a percentage of spermatophore length.
10 C.C. Lu & T. Okutani
Table 4. Amutatiola macroventosa n.gen. & n.sp. measurement, counts and indices of female specimens
MOV
F80087
Specimennumber [8 9 tt — |
Type status | [Paratype4 |- [Paratype] |- ———|- b ———
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5.2 94
37
33.6
63
54.6 66
67.2 7.6
ALE [| — — |742 — |675 (697 [882 05 196 |68 ———
63 |
Armfomla | 12341. 1324=1. 2341. (3241. 132=41. 324-1. 32-4.
dui — EB E B ee
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3.5
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NidGl 1399 j415 — 506 ja 1513 72 44.8
Note: Maturity 3, juvenile — young specimen in which some sexual characters are distinguished; Maturity 4, subadult — sexual characters well distinguished but gonads
and accessory organs not completely developed; Maturity 5, adult — sexually mature with spermatophores in Needham's sac in males; ovaries, nidamental and oviducal
elands fully developed and ripe, eggs sometimes in oviducts, in females; DML, dorsal mantle length (mm) — measured from anterior-most point of mantle to posterior
end of mantle; VMLI, ventral mantle length index — ventral mantle length, measured from anterior border of mantle at ventral midline, to apex of mantle, expressed as a
percentage of DML; HLI, head length index — dorsal length of head measured from point of fusion of dorsal arms to anterior tip of dorsal mantle expressed as a
percentage of DML; HWI, head width index — greatest width of head at level of eyes expressed as a percentage of DML; NCI, nuchal commissure width index — width
of nuchal commissure expressed as a percentage of width of mantle at the position of nuchal commissure; MWI, mantle width index — mantle width at mantle opening
expressed as a percentage of DML; FuLI, funnel length index — length of funnel from anterior funnel opening to posterior border of funnel measured along ventral
midline expressed as a percentage of DML; FuWI, funnel width index — width of funnel at junction of funnel and head just anterior to funnel locking cartilages
expressed as a percentage of DML; FFul, free funnel index — length of funnel from the anterior opening to the point of dorsal attachment to head expressed as a
percentage of funnel length; FLI, fin length index — greatest length of a single fin expressed as a percentage of DML; FBLI, fin base length index — length of fin
attachment to mantle expressed as a percentage of DML; FWI, fin width index — greatest width of a single fin expressed as a percentage of DML; A LI, arm I length
index — length of arm I expressed as a percentage of DML; A,LI, arm II length index — length of arm II expressed as a percentage of DML; A.LI, arm III length index
— length of arm Ill expressed as a percentage of DML; A LI, arm IV length index — length of arm IV expressed as a percentage of DML; TtLI, tentacle length
index — total length of tentacular stalk and club expressed as a percentage of DML; CILI, club length index — length of club, measured from proximal-most basal sucker
to distal tip of club expressed as a percentage of DML; CIRC, club row count — number of longitudinal rows of suckers across width of club; GLI, gill length index —
length of gill expressed as a percentage of DML; GWI, gill width index — greatest width of gill expressed as a percentage of DML; GiLC, gill lamellae count — number
of lamellae on outer demibranch, excluding terminal lamella; A, SC, sucker counts of arm I — total number of suckers or sucker stalks on arm I; A,SC, sucker counts of
arm II — total number of suckers or sucker stalks on arm II; ASC, sucker counts of arm III — total number of suckers or sucker stalks on arm III; A „>C, sucker counts of
arm IV — total number of suckers or sucker stalks on arm IV; CISI, club sucker index — diameter of largest club sucker expressed as a percentage of DML; A SI, arm I
sucker index — diameter of largest sucker on arm I expressed as a percentage of DML; A, SI, arm II sucker index — diameter of largest sucker on arm II expressed as a
percentage of DML; A,SI, arm III sucker index — diameter of largest sucker on arm III expressed as a percentage of DML; A,SI, arm IV sucker index — diameter of
largest sucker on arm IV expressed as a percentage of DML; NidGI, nidamental gland index — length of nidamental gland expressed as a percentage of DML.
Mio |A AID W | Oo
O^ | oo On| N o IS
QN
Wim yo | 2 | -- o3 [O2 |
DD W —
[
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia 11
Other material: MOV F80087: Franklin Island, South Australia,
32° 27'S, 133° 40'E, sub-light, 4 Apr 1953, 1 male, 8.0 mm mantle length,
mature; 1 female, 9.2 mm mantle length, subadult (specimens #6, 16).
MOV F80085: Great Australian Bight, 32° 43'S, 126° 00' E-32° 45'S,
125° 59' E, 40-170 m, collected by CSIRO, FRV Soela SO3/80/32, 13
May 1980, 0100 hr, IYGPT, 1 male, 6.6 mm mantle length, subadult; 9
males, 7.7-9.6 mm mantle length, mature; 3 females, 4.7-6.5 mm mantle
length, juveniles.
MOV F80086: Great Australian Bight, 33° S, 126° E, 48-50 m,
collected by CSIRO, FRV Soela S03/80/33, 13 May 1980, IYGPT, 1
male, 7.1 mm mantle length, subadult; 2 males, 8.5-9.8 mm mantle
length, mature; 1 female, 12.2 mm mantle length, mature.
MOV F80084: Great Australian Bight, 33° 22' S, 125° 27 E —
33° 23' S, 125° 27' E, 64 m, collected by CSIRO, FRV Courageous
Q47/51, 7 Apr 1979, 2 females, 10.3-13.2 mm mantle length, mature
(specimens #8, 15).
MOV F77100: Bass Strait, 60 km west of Cape Frankland, Flinders
Island, 39° 53'S, 147° 03' E, trawl depth 20-60 m, bottom depth 66-68
m, collected by CSIRO, FRV Soela SO1/82/1, 16 Jan 1982, 1 female, 5.6
mm mantle length, juvenile; 4 females, 10.6-12.6 mm mantle length,
mature (specimens #10, 12, 13, 14; four larger specimens).
MOV F80082: Great Australian Bight, 33° 30" S,
b
131° 50.0' E-33? 30' S, 131° 53' E, 200-144 m, collected by FRV Soela
S03/80/1,8 May 1980, 9 males, 4.8—7.6 mm mantle length, immature; 3
females, 5.4-6 mm mantle length, immature.
MOV F80080: Great Australian Bight, 32° 22'S, 131° 19' E-32° 17'S,
131° 18' E, 54-68 m, collected by CSIRO, FRV Soela SO3/81/41,6 Aug
1981,2130 hr, IYGPT, 2 females, 10.4-13.3 mm mantle length, mature.
MOV F158291: Port Victoria Jetty, South Australia, 34° 29' 45” S,
137° 28' 53” E, collected by J. Finn and Mark D. Norman, 21 May 1998,
] female, 10.9 mm mantle length, mature.
Description: Mantle (figs 6a—c, 7, 8a—b) short dome-shaped with
blunt posterior end, slightly longer than wide, muscular, studded
by large chromatophores, fused with head dorsally. Anterior
ventral mantle margin (figs 6b, 7b, 8b) shallowly concave with
blunt lateral projections at position of mantle—funnel connectives,
reaching level of posterior margin of eye lens. Head slightly
narrower than mantle, head length about 50% of mantle length.
Nuchal commissure moderately wide, width 38.6-50.5% of
mantle width at commissure in males, 43-59% of mantle width at
commissure in females, commissure does not reach beyond
medial borders of bulbous eyes.
C
Figure 6. Amutatiola macroventosa n. gen. & n. sp. male, holotype (MOV F80081): a, dorsal view; b, ventral view; c, lateral view; d, inside mantle
cavity showing funnel locking cartilages, partial photophores; e, diagram of photophores (p; drawn by T. Okutani); f, arm crown showing arm
sucker arrangement; g, diagram of arm crown showing sucker arrangement (drawn by T. Okutani); h, diagram of arm IV showing sucker
arrangement (drawn by T. Okutan1).
C.C. Lu & T. Okutani
Figure 7. Amutatiola macroventosa n. gen. & n. sp. male, paratype 7 (MOV F275295; grids in background: 1 mm x 1 mm): a, dorsal view; b,
ventral view; c, arm crown showing arm sucker arrangement; d, lateral view.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia 13
a d
Figure 8. Amutatiola macroventosa n. gen. & n. sp.: a, dorsal view, female, paratype 1 (MOV F275293); b, ventral view, female, paratype 1 (MOV
F275293); c, arm crown showing arm sucker arrangement, female, paratype 1 (MOV F275293); d, diagram of funnel locking cartilage, female,
paratype 1 (MOV F275293; drawn by T. Okutani); e, diagram of funnel organs, male, paratype 3 (MOV F80083; drawn by T. Okutani).
Figure 9. Amutatiola macroventosa n. gen. & n. sp. male, paratype 5 (MOV F158244). Oral views of arms (grids in background: 1 mm x | mm):
a, right arm I; b, right arm II; c, right arm III; d, right arm IV; e, left arm I; f, left arm II; g, left arm III; h, left arm IV.
14
Head almost entirely occupied by a pair of large bulbous
eyes with elliptical eye lid. Cornea membrane protecting eye
attached to skin of head along dorsal margin. Olfactory papilla
located behind posterior corner of eye orbit, ventral
photosensitive vesicle not found. Funnel (figs 6b, d, 7b, d, 8b)
long and slender, lacking pigmentation, reaching the level
beyond anterior eye margin, and free from head for 55—80% of
funnel length. Funnel connects to head by an oblique muscle
band extending from beneath anterior end of funnel locking
cartilage to ventro-posterior corner of eye.
Dorsal funnel organ (fig. 8e) broad V-shaped with a blunt
papilla at the apex. A prominent funnel retractor muscle
connected funnel near base with ventro-posterior periphery of
eye. Ventral elements of funnel organ (fig. 8e) tear drop-shaped
pad, narrower anteriorly. Funnel valve well developed on dorsal
roof, tongue shaped, located well behind funnel aperture.
Funnel locking cartilage (fig. 8d) elongated oval with
simple, slightly curved depression in middle, mantle locking
cartilage long, low ridge.
Fins (figs 6a—c, 7a—b, d, 8a—b) circular in outline, attach to
mantle at mid-point of mantle, meeting mantle smoothly
posteriorly, anterior border of fins project forward prominently
forming a deep cleft with mantle, border not reaching level of
mantle margin, posterior borders of fins convex, less
C.C. Lu & T. Okutani
pronounced. Length of fin base about 33% of mantle length.
Arms moderately long, rounded aborally, flattened orally,
lacking aboral keel and protective membranes. Arm formula
inconsistent, in males arm I or II usually longest, arm III or
IV usually shortest; in females, arm II or III longest, arm I or
IV shortest. Arm suckers biserial with strong sexual
dimorphism in sucker sizes and sucker ring dentitions. Web
moderately pronounced between arms III and IV. No
heteromorphism exists in morphology of right and left arms
in males, specifically in the dorsal arms. Webs shallow
between all arms except arms IV where no web exists, web D
encloses base of tentacles in both sexes.
In males (figs 6f—h, 7, 9), arm I with up to 28 suckers, some
suckers in proximal portion of arm enlarged: first to sixth
proximal suckers on both dorsal and ventral series enormously
enlarged, the third sucker on ventral series the largest, enlarged
suckers on ventral series generally larger than dorsal series.
Diameter of enlarged suckers exceed arm width. Arm II with up
to 32 suckers, of which proximal second to seventh on ventral
series slightly enlarged; no enlargement of suckers on dorsal
series, suckers gradually reduced in size from proximal to distal
end of arm II. Arm III with approximately 27 suckers, first to
fourth proximal suckers on ventral series enormously enlarged
(1.6 mm in diameter); no enlargement of suckers on dorsal
Figure 10. Amutatiola macroventosa n. gen. & n. sp. Sucker ring dentition (grids 1n background: 1 mm x 1 mm): a, left arm IV of male, paratype
5 (F158244) showing normal (non-enlarged) suckers near arm tip; b, right arm III of male, paratype 5 (F158244) showing sucker ring of enlarged
suckers; c, d, normal sucker of left arm II of male, paratype 5 (F158244); e, left arm I of male, paratype 5 (F158244) showing enlarged suckers;
f,an enlarged sucker of e showing sucker ring; g, dorsal view of female (F275296) showing non-enlargement of suckers; h, ventral view of female
(F275296) showing non-enlargement of suckers; 1, arm crown of female (F275296) showing non-enlargement of suckers; J, close-up of a portion
of an arm of 1 showing sucker ring dentition.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia 15
series. Arm IV with approximately 29 normal suckers, no
enlargement. Chitinous sucker rings of normal suckers almost at
the same level as muscular rim of suckers, sucker ring margin
minutely denticulated (figs 10a, c, d). Chitinous sucker rings of
enlarged suckers (figs 10b, e, f) extended aborally beyond level
of muscular sucker rims, chitinous ring often covered by thin,
Opaque membrane, sucker ring divided into 2 parts, distal
33-50% of sucker ring long, semicircular shape, proximal
portion of sucker ring lower than distal portion, crescent shaped.
Females with more suckers than the corresponding arm in
males. Arm I with up to 34 suckers, arm II with up to 45
suckers, arm III with up to 38 suckers and arm IV to 38
suckers. No enlarged sucker in females (figs 8c, 108-1).
Chitinous sucker rings (figs 101—]) similar to the non-enlarged
suckers in males, sucker rings entire, with minutely
denticulated margin.
Tentacle weak, longer than arms. Club slightly expanded,
curled due to the presence of the dorsal web, small carpal
suckers in 4 series, manal and dactyl suckers small, numerous
in approximately 6—8 series, carpal and manal suckers much
larger than dactylus suckers, in the central part of club,
particularly those on dorsal 2 series slightly larger than others,
suckers slightly diminish in size towards club margins and
distally (fig. 11a). Sucker ring dentition of largest club sucker
finely toothed around entire minute circle.
Gills with 16-21 lamellae per demibranch, plus a terminal
A
Figure 11. Amutatiola macroventosa n.gen. & n.sp. (grids in background: 1 mm x | mm): a, right tentacular club, male, paratype 5 (MOV F158244);
b, photophores (p), male, paratype 5 (MOV F158244); c, lateral view of upper beak, female, paratype 1 (MOV F275293); d, lateral view of lower beak,
female, paratype 1 (MOV F275293); e, top view of lower beak, female, paratype 1 (MOV F275293); f, radula, female, paratype 1 (MOV F275293).
es
16
lamella. A pair of yellowish photophores (figs 6e, 11b), opaque
with both ends swollen, dumbbell-shaped or elongated
kidney-shaped, on both sides of ink sac.
Upper beak (fig. 11c) rostrum slightly curved with pointed
rostral tip; jaw angle obtuse; wing long, shoulder (cutting
edge) nearly straight; lateral wall deep; rostrum dark brown to
black, hood, shoulder and dorsal part of lateral wall light
brown, posterior part of hood, most of lateral wall
unpigmented, transparent.
Lower beak (figs 11d, e) wide; rostral tip blunt; jaw edge
almost smooth, jaw angle indistinct; blunt tooth on shoulder;
no notch in hood; lateral wall without fold or ridge, roughly
elongate rhomboidal with lower edge concave, corner faintly
produced; rostrum and hood light brown in colour, wings and
posterior part of lateral wall transparent. Radula (fig. 11f) 7
series, each row with 7 unicuspid teeth.
Gladius absent.
Spermatophores (fig. 12a) small, 8 spermatophores from 5
individuals approximately 2.6—4.1mm long (spermatophore
length index 29.3—42.7), greatest width approximately 0.17—
0.25 mm (spermatophore width index 5.4—7), sperm mass
moderately long, sperm reservoir about 0.97—1.5 mm (sperm
reservoir index 27.9—43.2), structure simple, no obvious
ornamental appearance, cement body approximately 0.39-
0.75 mm long, connects to sperm reservoir by a narrow neck,
oral end of cement body elongated funnel-shaped (cement
e
p
ES
Figure 12. Amutatiola macroventosa n. gen. & n. sp.: a, spermatophore, holotype (MOV F80081) Spermatophore length, 3.55 mm (insets: 1,
C.C. Lu & T. Okutani
body index 11-22), appearance of ejaculatory apparatus plain,
without spiral appearance of the preceding species.
Bursa copulatrix closed type, pouch like (Bello, 2020; fig.
12b), opening at level of base of left gill, running dorsally, on a
mature female (Paratype 1) anterior end width approximately
2 mm with slit-like opening approximately 1.1 mm long, length
of pouch approximately 2.3 mm, some spermatangia visible at
opening of bursa. Mature and maturing females with large
nidamental gland, (nidamental gland index 37.2—56.6).
Alcohol-preserved specimens brown in colour, dorsal
mantle surface slightly darker than ventral surface. Dark
blackish brown chromatophores scattered over brown-
coloured background on both dorsal and ventral surfaces of
head and mantle, and along aboral surface of all arms.
surfaces of fins devoid of chromatophores and pigmented
spots, except a semicircular patch of brown chromatophores
along fin insertion on dorsal side of fin. Skin smooth, lacking
sculpture or papillae.
Etymology: Species epithet macroventosa is derived from
Greek makros meaning large plus Latin ventosa meaning
suction cup or sucker. The name denotes the presence of the
ereatly enlarged suckers on some arms of mature males.
Distribution: Flinders Island, Bass Strait, and south-eastern
Tasmania to South Australia and the Great Australian Bight to
Port Denison, Western Australia (fig. 13).
close-up of oral cap; 11, close-up of cement body); b, Bursa copulatrix, paratype 1 (MOV F275293; black scale: 1 mm/division) (Iflc: left funnel
locking cartilage; lg: left gill); arrow points to the bursa copulatrix entrance.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) from Australia 17
Remarks: Apart from Dextrasepiola taenia n. sp. described
earlier in this paper, all known species of the subfamily
Sepiolinae have the left dorsal arm of maturing and mature
males hectocotylised and sucker stalks of some suckers in the
copulatory apparatus of the hetocotylus modified into horn-
like, hook-like, papillae or laminae (Bello, 2020; Naef, 1912a,
b; 1923; Nesis, 1982). The present species is unique in having
enormously enlarged suckers and lacking a hectocotylised arm
bearing highly modified sucker stalks in males. There are some
species of Sepiolinae that carry enlarged suckers in
hectocotylised arms (Bello, 2020; Naef, 1923), but none is so
pronounced as in this species. The female has normal-sized
suckers but more of them. In addition to sucker size, strong
sexual dimorphism exists ın the dentition of the sucker ring: the
re) wu
dentition of the sucker ring of enlarged suckers in males
conspicuously differs from that of the females and the non-
enlarged suckers in males.
The structure of the cement body and the sperm mass of the
spermatophore of this species is simple 1n appearance, with no
obvious ornamentation, as seen in the preceding species or in
Sepietta oweniana (d’Orbigny in Férussac and d’Orbigny, 1841;
cf. figs 5a, b, 12a; Oresland and Oxby, 2021, figs 59-61).
As in the preceding species, the ranges of the
morphometric indices are wide (Tables 3—4). This 1s certainly
due to the range of the state of specimens prior to and during
fixation and preservation. With such a wide range of values, it
is of dubious value to use them to delineate a specific index
for the species.
Porl Moresby
©
Australia
— p ~ «| »
Penh
©
Newcastle
Sydney
n4.
Adelaide
C Canberra
T
IOHIA
: »
Melbourne
9
g
e
Figure 13. Distributional map of Dextrasepiola taenia n. gen. & n. sp. (blue) and Amutatiola macroventosa n. gen. & n. sp. (red).
C.C. Lu & T. Okutani
18
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Discussion
Bello (2020) revised the subfamily Sepiolinae based on
analyses of the structures of the hectocotylised arms, and split
the old Sepiola Leach, 1817, into four genera: Sepiola Leach,
1817, Adinaefiola Bello, 2020, Boletzkyola Bello, 2020, and
Lusepiola Bello, 2020. The genus Euprymna Steenstrup, 1887,
was split into two genera: Euprymna and Eumandya Bello,
2020. According to Bello’s (2020) revision, Sepiolinae includes
nine genera, of which seven genera — Sepiola, Adinaefiola,
Boletzkyola, Inioteuthis, Lusepiola, Rondeletiola and Sepietta
— have a tripartite hectocotylus, and the remaining two genera
— Euprymna and Eumandya — are characterised by the
hectocotylus (left arm I) with sucker-stalks in its distalmost
portion modified into a palisade-like structure.
Both new genera, Dextrasepiola and Amutatiola, share
some basic features with the other members of Sepiolinae:
mantle fused dorsally with head, biseriate arm suckers, bilobed
kidney- or dumbbell-shaped visceral light organ (the latter two
features are deemed plesiomorphic character states by Bello,
2020), reduced arm web, ventral mantle does not produce into a
ventral shield (unlike in Heteroteuthinae) and small body size.
Dextrasepiola n. gen. also shares the same type of
hectocotylisation with the tripartite-hectocotylus genera, but
it conspicuously differs from the other because of its unique
peculiarity that the right dorsal arm of the male is modified
into a copulatory arm instead of the left one. In this genus, the
copulatory apparatus is also unique in being formed by two
suckerless stalks modified and fused into a tape-like structure.
Amutatiola n. gen. 1s also unique among Sepiolinae in lacking
an evident hectocotylised dorsal arm in male and instead
having some arm suckers enormously enlarged. Table 5 shows
the comparison of features of all Il genera of Sepiolinae.
In a recent report, Sanchez et al. (2021) concluded that the
subfamily Sepiolinae is split into Indo—Pacific and Atlantic—
Mediterranean lineages, the former characterised by a closed
type bursa copulatrix, and the latter by an open type bursa. The
discovery of the Dextrasepiola taenia n. gen. and n. sp., (1.e. a
sepioline genus and species with an open type bursa copulatrix)
is not consistent with that systematic-biogeographic pattern.
With Amutatiola macroventosa n. gen. and n. sp., the lack
of a normal sepioline hectocotylus in males is a unique feature
which co-occurs with the enlargement of several suckers 1n
the male arms. The enlargement of some suckers 1s also found
in the mature males of several species of Euprymna (Norman
and Lu, 1997) as well as in some Rossiinae and
Heteroteuthinae (Nesis, 1982). Contrary to Dextrasepiola
taenia n. gen. and n. sp., females of Amutatiola macroventosa
n. gen. and n. sp. display a closed bursa copulatrix, which
parallels the other Indo-Pacific sepioline genera and species.
Based on the present results and descriptions of the new
sepioline genera and species, the definition of the subfamily
sepiolinae should be widened to include Dextrasepiola and
Amutatiola. This can be done by amending the character state
of Hectocotylisation for Sepiolinae in Table | to "left or right
Gn Dextrasepiola) dorsal arm hectocotylised or no
hectocotylisation (in Amutatiola)” and by adding the character
“bursa copulatrix" as "bursa copulatrix in female on left side of
C.C. Lu & T. Okutani
mantle cavity in all Sepiolinae except Dextrasepiola (on right
side)”. Subfamilies Heteroteuthinae and Rossinae do not have
well-differentiated bursa copulatrix, spermatophores are
implanted on the body or on a patch of wrinkled tissue near the
opening of the oviduct (Hoving, et al., 2008, 2009; Naef, 1923).
Acknowledgements
We are indebted to Giambattista Bello, Mola di Barı, Italy, for
his critical review of the manuscript and constructive
suggestions. Discussions with him are most helpful and greatly
appreciated. We are also grateful for the suggestions from the
anonymous reviewers. We also wish to thank Wensung Chung,
Visual Ecology Lab., Sensory Neurobiology Group, Queensland
Brain Institute, University of Queensland, Australia, for his
help in assembling the figures and in photographing figs 1, 2, 8
as well as in constructing fig. 13.
References
Bello, G. 2013. Description of a new sepioline species, Sepiola
bursadhaesa n. sp. (Cephalopoda: Sepiolidae), from the Catalan
Sea, with remarks and identification key for the Sepiola atlantica
group. Scientia Marina TT. 489-499.
Bello, G. 2020. Evolution of the hectocotylus in Sepiolinae
(Cephalopoda: Sepiolidae) and description of four new genera.
European Journal of Taxonomy 655: 1-53
Bello, G. and Salman, A. 2015. Description of a new sepioline species,
Sepiola boletzkyi n.sp. (Cephalopoda: Sepiolidae), from the
Aegean Sea. European Journal of Taxonomy 144: 1-12.
Hei] A. de, and Goud J. 2010. Sepiola tridens spec. nov., an overlooked
species (Cephalopoda: Sepiolidae) living in the North Sea and
north-eastern Atlantic Ocean. Basteria 74: 51—62.
Hoving, H.J.T., Laptikhovsky, V., Piatkowski, U., and Onsoy, B. 2008.
Reproduction in Heteroteuthis dispar (Rüppell, 1844) (Mollusca:
Cephalopoda): a sepiolid reproductive adaptation to an oceanic
lifestyle. Marine Biology 154: 219-230.
Hoving, H.J.T., Nauwelaerts, B., Van Genne, B., Stamhuis, E.J., and
Zumholz, K. 2009. Spermatophore implantation in Rossia
moelleri Steenstrup, 1856 (Sepiolidae; Cephalopoda). Journal of
Experimental Marine Biology and Ecology 372: 75-81.
Lu, C.C. 2001. Cephalopoda. Pp.129-308 in: Wells, A. and Houston,
H.W.K. (eds) Zoological Catalogue of Australia. Vol. 17.2.
Mollusca: Aplacophora, Polyplacophora, Scaphopoda,
Cephalopoda. CSIRO Publishing: Melbourne. x11, 353 pp.
Lu, C.C., and Boucher-Rodoni, R. 2006. A new genus and species of
sepiolid squid from the waters around Tonga in the central South
Pacific (Mollusca: Cephalopoda: Sepiolidae). Zootaxa 1310: 37-51.
Lu, C.C., and Dunning, M.C. 1998. Subclass Coleoidea Bather, 1888. Pp.
499-563 in: Beesley, P.L., Ross, G.J.B., and Wells, A. (eds)
Mollusca: The Southern Synthesis. Fauna of Australia. Volume 5.
CSIRO Publishing: Melbourne. Part A. xvi, 563 pp.
Lu, C.C., and Phillips, J.U. 1985. An annotated checklist of the
Cephalopoda from Australian waters. Occasional Papers of the
Museum of Victoria 2: 21-36.
Lu, C.C., and Roper, C.F.E. 1979. Cephalopods from deepwater dumpsite
106 (Western Atlantic): vertical distribution and seasonal
abundance. Smithsonian Contributions to Zoology 288: 1-36.
Naef, A. 1912a. Teuthologische Notizen. 3. Die Arten du Gattungen
Sepiola und Sepietta. Zoologischer Anzeiger 39: 262-271.
Naef, A. 1912b. Teuthologische Notizen. 7. Zur Morphologie und
Systematik der Sepiola- und Sepietta- Arten. Zoologischer
Anzeiger 40: 78-85.
Two new genera and species of sepioline squids (Cephalopoda: Sepiolidae) trom Australia 23
Naef, A. 1923. Die Cephalopoden, Systematik. Fauna und Flora des
Golfes von Neapel 35: 1-863. Israel Program for Scientific
Translation: Jerusalem [English translation].
Nesis, K.N. 1982. Cephalopods of the world. English Translation from
Russian. Levitov, B.S. (Transl.), Burgess, L.A. (ed.) (1987). T.F.H.
Publications: Neptune city. 351 pp.
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Biological Association of the United Kingdom TT. 1109-1137.
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for bobtail squids. Divers and Scientists West Coast Sweden,
Guide No.1. 122 pp.
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descriptions of cephalopod species. Memoirs of the National
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Memoirs of Museum Victoria 81: 25-41 (2022) Published 2022
1447-2554 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2022.81.02
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their
significance
FEARGHUS R. MCSWEENEY FGS!”, JEFF SHIMETA’, JOHN S'. J.S. BUCKERIDGE FGS!”
! Earth and Oceanic Systems Group, RMIT University, GPO Box 2476, Melbourne, VIC 3001, Australia
* Centre for Environmental Sustainability and Remediation, School of Science, RMIT University, Melbourne, Australia
> Museums Victoria, VIC 3001, Australia
* Correspondence: Fearghus McSweeney, tidal75@gmail.com
McSweeney, F.R., Shimeta, J., and Buckeridge, J.S. 2022. Lower Devonian Zosterophyllum-like plants from central
Three specimens belonging to Zosterophyllaceae are described. Two of these possess bilateral symmetry and are
the first to be described with this arrangement from the Lower Devonian of Victoria. One of these specimens is similar to
Zosterophyllum fertile, and the other cf. Zosterophyllum sp. A. ıs unusual in possessing vascularised long stalks. The third
specimen described cf. Zosterophyllum sp. B. from Ghin Ghin Road, near Yea possesses a small spike and has sporangia
that appear vertically elliptical and similar to some South China taxa. All the specimens are significantly different to
Abstract
Victoria, Australia, and their significance. Memoirs of Museum Victoria 81: 25—41.
previous zosterophyll taxa described from Victoria.
Keywords Victoria, Baragwanathia flora, Platyzosterophyllum, zosterophyll, systematics.
Introduction
The Wilson Creek Shale Formation of Victoria is notable for
providing a link between the lithologies of the Mount Easton
and Darraweit Guim Provinces (fig. 1) of the Melbourne Zone
(VandenBerg, 1988: 114) and for being the type formation of
Baragwanathia longifolia Lang and Cookson, 1935.
Furthermore, it provides a useful approximate marker
delineating the Silurian from Devonian rocks, as it has been
dated as mid Pragian—Emsian (Carey and Bolger, 1995; Mawson
and Talent, 1994). This paper examines three zosterophylls, two
of which come from the Wilson Creek Shale and are the first to
be described with bilateral symmetry in Victoria and are the
first to be described from this formation. The third specimen is
from the base of the Humevale Formation near Yea.
The first zosterophyll described from Victoria was
Zosterophyllum australianum Lang and Cookson, 1930, from
the Centennial Beds at North Road Quarry near Walhalla, now
considered to be part of the Norton Gully Sandstone Formation
(Douglas and Jell, 1985: 161). Lang and Cooksons’ (1930: 148)
work on Z. australianum confirmed an earlier assumption
(Lang, 1927: 449-450) that the reinform appendages of
Zosterophyllum were sporangia, with the presence of in situ
spores. Additionally, Cookson (1935: pl. 10, figs 9-15) described
Z. australianum from Mount Pleasant and Halls Flat Roads
near Alexandra, also considered part of the Norton Gully
Sandstone (VandenBerg, 1975, 1988). Hao and Wang (2000)
reassigned Cookson’s (1935) Z. australianum specimens to Z.
ramosum, primarily based on different — sporangial
morphological characteristics. The Zosterophyllum ramosum
type locality occurs near Zhichang village in the Posongchong
Formation, Yunnan, China (Hao and Wang, 2000).
Tims (1980), in her unpublished thesis, described and gave
informal manuscript names to three zosterophylls in Victoria:
Zosterophyllum minutum from Boola Formation at Tyers,
Pluricaulis biformis from the Yea Formation at Limestone
Road and Chamaecaulon tylosus from the Wilson Creek
Shale on Frenchmans Spur Track near Matlock.
Zosterophyllum minutum was subsequently redescribed and
assigned to Gippslandites minutus McSweeney et al., 2020.
Additionally, McSweeney et al. (2020) described another
zosterophyll, Parazosterophyllum timsiae, from the base of
the Humevale formation on Ghin Ghin Road, Yea.
Stratigraphic setting
Humevale Siltstone. The Humevale Formation was proposed by
Williams (1964: 277) as a sequence of primarily siltstones about
5 km south-west of Tommy’s Hut at Humevale and is found
throughout much of the Darraweit Guim Province (fig. 1) of the
Melbourne Zone (Edwards et al., 1997: fig. 6; VandenBerg,
1988: table 4.1). Williams (1964) delineated the upper and lower
parts of the Humevale Formation with two sandstone units: the
Clonbinane and Flowerdale Members, respectively. Above the
26
Clonbinane Member near the base of the Humevale Formation,
Williams (1964: 276) added the Mount Phillipa Member, a
sandstone unit noted for numerous Nucleospira sp. and
rhynchonellids. Garratt and Wright (1988: 650) subdivided the
Humevale Formation based on brachiopod faunal successions,
with Notoparmella plentiensis Garratt, 1980, above the base of
the Humevale Formation in the Whittlesea area, and the
appearance of Boucotia janaea Garratt, 1980, near the top of
the sequence. However, Talent et al. (2001: 151) suggested
b. janaea maybe B. australis Gill, 1942. Edwards et al. (1997:
22) considered the Clonbinane and Mount Phillipa Members to
be part of the Dargile Formation, and the Flowerdale Member to
be part of the Norton Gully Sandstone, while retaining the
Humevale Siltstone. At Ghin Ghin Road, the fossil exposure P4
(loc 1 1n Garratt, 1978) according to Rickards and Garratt (1990:
fig. 2) 1s at the base of the Humevale Formation, and Pridoli,
upper Silurian; thus, slightly younger than the exposure at
Limestone Road, south-east of Yea, which they consider to be
Ludlovian, with both interpretations based on graptolites. This
Ludlovian—Pragian, upper Silurian age for the exposures in the
Yea area by Garratt (1978), Rickards and Garratt (1990), and
Rickards (2000) is contentious, primarily due to the relatively
advanced plant architectures for this time and the similarity of
some of the flora to younger exposures elsewhere in Victoria,
such as the Wilson Creek Shale (Banks, 1980; Cleal and
Thomas, 1999; Edwards et al. 1979 Hueber, 1983, 1992).
Edwards et al. (1997: 22) placed much of the lithology of Yea,
including both Limestone Road and Ghin Ghin Road exposures,
in the lithologically variable Norton Gully Sandstone.
VandenBerg (pers. comm. June 2021) mapped the Yea area in
2008 (unpublished) confirming much of Edwards et al. (1997),
and found Yea to be predominantly Norton Gully Sandstone
with a narrow band of the Wilson Creek Shale near the Yea
anticline. In this paper, we retain the Humevale Siltstone,
because this new information on the Yea lithology needs to be
published. However, we find Edwards et al. (1997) and
VandenBerg’s argument cogent and so use a conservative
interpretation of the age of the exposure at Ghin Ghin Road,
giving it a Pragian—Emsian, Lower Devonian.
Wilson Creek Shale. The Wilson Creek Shale occurs in both
the Darraweit Guim and Mount Easton provinces of the
Melbourne Zone (fig. 1) and part of the Jordan River Group
(VandenBerg, 1973, 1988). Lithologically, the Wilson Creek
shale consists of black shale that changes from cream to grey
on weathering and is interpreted as representing once deep
anoxic quiescent marine conditions (Edwards et al., 1997;
VandenBerg et al., 2006). Sandstone beds are prevalent towards
the top of the formation and are construed as representing the
srowing influence of turbidity currents late in the sequence
(Edwards et al., 1997). Siltstone occurs primarily in the basal
and upper portions of the formation, and limestone is found to
interfinger with the Coopers Creek Formation at Jacobs Creek,
north of Tyers (Edwards et al., 1997; VandenBerg, 1988;
VandenBerg et al., 2006). The Wilson Creek Shale conformably
overlies the Humevale Siltstones, Eldon Sandstone, Whitelaw
siltstone and Boola Formation, and conformably underlies the
Norton Gully Sandstone, Easts Lookout Siltstone (fig. 2),
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
Marshall Creek Member and Yeringberg Formation, albeit
large scale strike faulting is common according to VandenBerg
(1988) and VandenBerg et al. (2006). Lang and Cookson (1935)
described — in addition to Baragwanathia longifolia — Yarravia
oblonga, Y. subsphaerica and Hostinella sp. as occurring in the
Wilson Creek Shale. Tims (1980) described numerous informal
species in her thesis as occurring in the Wilson Creek Shale,
especially at Frenchmans Spur Track and Coles Clearing on
the Thomson River. Following her thesis, she described two
rhyniophytoids, Salopella australis and S. caespitosa, and the
trimerophyte Dawsonites subarcuatus as occurring in the
Wilson Creek Shale at Frenchmans Spur Track, about 10 km
west of Matlock (Tims and Chambers, 1984). Recently,
Salopella laidae was described from Limestone Road by
McSweeney et al. 2021c.
In the first half of the twentieth century, the lithological
sequence that would be later assigned to the Wilson Creek
Shale (Edwards et al., 1997; Thomas, 1953; VandenBerg,
1975, 2006) was believed to be early Ludlow, upper Silurian,
based on Elles's assignation (Lang and Cookson, 1935: 422).
Jaeger (1966) reassigned the graptolites to the Lower
Devonian based on the occurrence of the monograptids
Uncinatograptus thomasi (=Monograptus thomasi thomasi)
Jaeger, 1966, throughout the formation, and Neomonograptus
notoaequabilis (=M. aequabilis notoaequabilis) Jaeger and
Stein, 1969, in the upper half of the formation (Lenz, 2013).
Neomonograptus notoaequabilis extends Into the overlying
Norton Gully Sandstone according to VandenBerg et al.
(2006). Furthermore, both Eognathodus sulcatus sulcatus
and Polynathus dehiscens occur in the Coopers Creek
Limestone that inter-fingers the Wilson Creek Shale at Tyers,
indicating a mid Pragian—Emsian age (Carey and Bolger,
1995; Mawson and Talent, 1994).
Material and methods
The specimens cf. Zosterophyllum sp. A. (NMV P256740) and
cf. Z. fertile (NMV P50040) were found by J. D. Tims on
Frenchmans Spur Track near Matlock in the Wilson Creek Shale
in the 1970s, with the former referred to in Tims (1980: fig.
4.4.19) indirectly, whereby she described three Hedeia
(=Yarravia) sp., which occur on the same plane. Tims (1980)
noted the specimen cf. Zosterophyllum sp. A. was loaned to H. P.
Banks but made no reference to the associated zosterophyll. The
specimen cf. Zosterophyllum sp. A. (McSweeney et al. 2021a)
was given to the lead author by Prof. Dianne Edwards of Cardiff
University in March 2019. The specimen cf. Z. fertile was
originally assigned to Zosterophyllum sp. by Tims (1980: 91).
The final specimen cf. Zosterophyllum sp. B. (NMV P256742)
was found as float by the lead author at location P4 on Ghin Ghin
Road, near Yea, on 8 August 2012, at location P4 on Ghin Ghin
Road, about 8 km north north-west of Yea Township (37° 13° S,
145° 38° E). P4 is equivalent to loc. | in Garratt (1978: fig. 2).
Dégagement had been undertaken on cf. Zosterophyllum
sp. A. to the right of the spike on the part. Further dégagement
as per Fairon-Demaret et al. (1999) was carried out by the
lead author to better expose sporangia two, four and five, and
around the basal region of the fertile axis, and to expose part
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance
of the axis near the base and apex of the specimen. Samples
taken from sporangia two and four of cf. Zosterophyllum sp.
A. did not reveal any preserved spores, and samples of the
basal axial region of the fertile axis did not yield any
anatomical features when viewed under a low vacuum on a
FEI Quanta 200 ESEM. Degagement was performed on the
distal and proximal parts of the spike of cf. Z. fertile by Tims
(1980: 91) to expose sporangia in those areas, but no further
dégagement was carried out because the specimen was fragile.
Dégagement of cf. Zosterophyllum sp. B. was undertaken by
the lead author around the spike, but no additional parts of the
zosterophyll were found. A veneer of iron-oxide coating the
surfaces was removed and mounted onto a stub and examined
under a low vacuum on a FEI Quanta 200 ESEM. No spores
or internal anatomy were uncovered.
Photographs of cf. Zosterophyllum sp. A., cf. Z. sp. B. and
cf. Z. fertile were taken using a AxioCamMRc5 camera
Victoria
= j |
_ |
i I
,
Ma "
c JE
| Wilsons Promontory
Bass Strait
0 50 100
km
Tasmania’
Clarke Island@
px N
21
attached to a Zeiss microscope and a Leica M205 C microscope
with Leica Application Suite software version 3.8.0. Images
were Z-stacked to improve depth of field using Adobe
Photoshop CC 2017. ImageJ software was used to take
morphological measurements. Rodney Start of Museums
Victoria took images of cf. Zosterophyllum sp. A. on 5
December 2019 using a Canon EOS 5D Mark III camera and
cross-polarised light circular filter with flash strip-lights to
enhance contrast. All photographs had contrast enhanced using
Adobe Photoshop CC 2017 and were arranged using Adobe
Illustrator CC 2017.
Institutional abbreviation
NMV P, Museum Victoria Palaeontology Collection,
Melbourne, Australia
Australia =
| 500 km
Darraweit Guim Province
. Mount Easton Province
Flinders
= Island
=)
* Fossil exposures
Figure 1. Location of Matlock in the Mount Easton Province and Yea in the Darraweit Guim Province of the Melbourne Zone in Victoria,
Australia. Source: after Moore et al. (1998: fig. 2).
28
Systematic palaeontology
Division Tracheophyta
Class Zosterophyllopsida Hao and Xue, 2013
Order Zosterophyllales Banks, 1968
Family Zosterophyllaceae Banks, 1968
Remarks. Initially, Zosterophyllum were separated into two
groups, Platy-zosterophyllum and Eu-zosterophyllum, by Croft
and Lang (1942: 145), but Hueber (1972: 121) split
Zosterophyllum into two subgenera, Platyzosterophyllum and
Zosterophyllum. The — distinguishing character of
Platyzosterophyllum is the arrangement of sporangia in row(s)
of one, two or three along the fertile axes, while the sporangia
of Zosterophyllum are arranged helically (Croft and Lang,
1942; Edward, 1975; Gensel, 1982; Hueber, 1972). Furthermore,
Platyzosterophyllum often possess circinate vernation (Croft
and Lang, 1942; Edward, 1975; Gensel, 1982; Hueber, 1972).
The specimens cf. Zosterophyllum sp. A. and cf. Z. fertile
described herein possess bilateral symmetry, but lack enough
characteristics to unequivocally assign to Platyzosterophyllum.
Oaks Creek
L. Devonian
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
cf. Zosterophyllum fertile Leclercq, 1942
Figure 3a—c
Material examined. NMV P50040.1 and P50040.2, part and
counterpart, respectively.
Locality. Occurs on a road cutting on Frenchmans Spur Track,
approximately midway between Big River Road to the north-
north-west, Warburton Road to the east and Frenchmans Spur
Track, ~10 km west of Matlock, central Victoria.
Horizon and age. Wilson Creek Shale Formation, middle
Pragian—Emsian, Lower Devonian (Carey and Bolger, 1995;
Mawson and Talent, 1994).
Description. The specimen consists of part and counterpart of
one partial spike with basal and apical regions missing and
preserved as a carbonised compression. The axis is ~1.0 mm
wide and 10.0 mm long and unbranched, and the spike up to 6.8
mm wide. Eight sporangia are borne alternately in two rows.
The sporangia are attached by recurved stalks; some are
perpendicular to ~60° to the fertile axis and curve sharply
distally to at most 90°. The convex margins of some of the
Lithological units
Wilson Creek Shale
Norton Gully Sandstone
Whitelaw Siltstone
Easts Lookout Siltstone
Associated units:
Monty's Hut Fmn
White Star Sandstone
Dnnm Marshal Creek Member
Dip Coopers Creek Limestone
416 Ma
Sihr Sju — Wurutwun Fmn.
Silurian Dlo Boola Fmn
Sjn Murderers Hill Siltstone
Sir serpentine Creek Sandstone
Sid Donnellys Creek Siltstone
Sl Lazarini Siltstone
Sis Sinclair Valley Sandstone
Sim McAdam Sandstone
Oxe Mount Easton Shale
444 Ma
Ordovician
488 Ma
Fossil site
*r Frenchman's Spur outcrop
Figure 2. Location of the Wilson Creek Shale outcrop on Frenchmans Spur Track, 10 km west of Matlock. Geological map showing the location
of Frenchmans Spur outcrop at the star, north of Springs Creek. Source: Map after Willman et al. (2006).
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 29
sporangia possess a thickened/darker/border, here interpreted
as likely pertaining to dehiscence (fig. 3, sporangia 6, 7). An
adaxially orientated basal lobe occurs on some sporangia (fig.
3b, central arrows).
Remarks. The sporangia are in two rows with no clear
demarcation between the attachment of the stalk and the basal
region of the sporangia. The specimen bears some resemblance
to Edwards’ (1969a: fig. Ib) South Wales Z. ct. fertile. Edwards’
(1969a) specimens are from the Old Red Sandstone Brecon
Beacons and Llanover Quarries, South Wales (Pragian—Emsian
Lower Devonian age) and were placed in Z. cf. fertile, because
the original diagnosis was based on only a single specimen
from Belgium (Leclercq, 1942) such that further specimens
from both Wales and Belgium were thought necessary to
determine specific affinity with confidence. Wellman et al.
(2000) recorded the earliest occurrence of Z. cf. fertile from the
Anglo-Welsh Basin as being mid-Lochkovian (Lower
Devonian). The fructification of Edwards’ (1969a) specimens,
while incomplete, were 1.0 cm high and 3.0 mm wide, which is
comparable in height but over two times smaller than the
Victorian specimen, which reached 6.8 mm wide. Edwards’
(1972) Z. fertile, also based on an incomplete spike but with a
significant proportion preserved, was 7.2 cm high and 3.0 mm
wide, while Wellman et al.’s (2000) Z. cf. fertile reached 2.0 cm
in height and 3.5 mm wide. Axial width for Leclercq’s (1942)
holotype measured 1.0-1.5 mm wide and conforms to the
Victorian specimen. However, the Victorian specimen’s axis,
where visible, was found to be ~1.0 mm wide. The widest axial
width for Z. fertile was found by Edwards (1972: 78) for a
specimen from the Lower Old Red Sandstone of Forfar,
Scotland, with an axial width of 3.0 mm, decreasing only
slightly to 2.8 mm wide, while the specimens of Z. cf. fertile
from Brecon Beacons Quarry, according to Edwards (1969a:
924), showed greater range, 0.8—2.5 mm wide.
of SS ea qun
The stalks for Leclercg (1942), Edwards (1969a, 1972) and
Wellman et al. (2000) range between 0.3-0.5 mm wide and
1.0-1.8 mm long. The Victorian cf. Z. fertile stalks are
broadly similar, 0.5—0.7 mm wide and up to ~2.0 mm long.
The sporangial shape in face view for the Victorian
specimen is reniform (fig. 3, sporangium 8), similar to Wellman
et al. (2000), while Leclercq (1942) described it as elongate—
reniform, and Edwards (1969a) for Z. cf. fertile described it as
irregular. The sporangia examined by Leclercq (1942), Edwards
(1969a, 1972) and Wellman et al. (2000) were all in the range of
2.0-2.5 mm wide and 1.6-3.1 mm high. The sporangial
dimensions of cf. Z. fertile are difficult to ascertain due to their
poor preservation.
Edwards (1969a: 924) found the longest fructification of Z.
cf. fertile. It had eight sporangia but lacked an apical region. Due
to the lack of a complete spike herein and poor preservation
resulting in equivocal characters, the Victorian specimen was
placed in cf. Z. fertile, it being conceivable that given better
preservation the plant might be placed it outside the defining
characteristics of Zosterophyllum.
cf. Zosterophyllum sp. A
Gen. et sp. indet.
Figures 4-9
Material examined. NMV P256740.1 and P256740.2, part and
counterpart, respectively.
Locality. Frenchmans Spur Track, ~10 km west of Matlock,
central Victoria.
Horizon and age. Wilson Creek Shale, middle Pragian—
Emsian, L. Devonian (Carey and Bolger, 1995; Mawson and
Talent, 1994).
Figure 3,01%, here, from the Wilson Creek Shale Formation on Frenchmans Spur, 10 km west of Matlock: a, part (NMV P50040.1); b, line
drawing of partial spike. Abx = abaxial valve, Adx = adaxial valve, Q = ?poorly preserved sporangium, P = stalk, which is normal to the fertile
axis before turning upwards. Central arrows pointing to lobes; c, counterpart (NMV P50040.2). Note. Counterpart image reversed to be in the
same orientation as the part.
30
Description. The specimen consists of a longitudinally elongate
lax spike with its apical region missing. The basal half of the
spike contains about a third of the total sporangia 1n two rows
(figs 4, 5a), and distally, the sporangia are more closely arranged
(?helically) but the insertion points are not clear. The naked
fertile axis is unbranched, 1.3-2.6 mm wide, curving basally,
the spike slightly decreases in width acropetally. The lax spike
is 10 mm wide and up to at least 45 mm long, consisting of 20
sporangia arranged on long vascularised stalks up to 2.0 mm
long and 1.0-1.3 mm wide, at acute angles 15°-45° to the
vertical, before the stalks reorientate towards the apex of the
spike just beneath each sporangium. There is very little vertical
overlap of sporangia. Some fine protuberances and depressions
emanating from the vascular trace (fig. 6) are interpreted here
as representing insertions of further stalks. The junction
between sporangium and stalk is unknown. The sporangia are
circular to reniform in face view, 0.95—3.7 mm wide and 0.5-
S11 Oy
N
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
2.3 mm high, with weakly developed lobes (fig. 7) and a narrow
border visible on distal margin of some sporangia, such as
sporangia two, 10 (figs 7, 8) and 12, are 0.13-0.15 mm wide and
is interpreted as likely pertaining to dehiscence. Sporangia one
and two are longitudinally elliptical and are interpreted to be
infolded, such that half the abaxial valve is visible (fig. 8).
Vascular trace 0.34—1.3 mm in the fertile axis, 0.17-0.21 mm
on the stalks. There are two sporangia 1n close proximity to the
spike, but they are clearly orientated at an angle to indicate they
may come from another axis ın their vicinity (fig. 5a).
The vascular trace is conspicuous in that it is preferentially
preserved compared with cortical tissue (fig. 4), the cortex being
preserved as a grey film in the surrounding matrix. The stalk of
sporangium one (fig. 8) ıs inserted almost perpendicular to the
fertile axis and is bent such that most of the stalk is parallel to
the fertile axis before curving upwards, just beneath the basal
region of the sporangium. Several poorly preserved axes lie
Figure 4. cf. Zosterophyllum sp. A. from the Wilson Creek Shale Formation on Frenchmans Spur, 10 km west of Matlock: a, line drawing, dotted
lines are from faint remains of compression. S=Sporangium; Q=Poorly preserved sporangium or sporangium likely not belonging to same spike;
b, part NMV P256740.1. Arrow at stalk, which has been pushed across fertile axis; c, counterpart, NMV P256740.2 with part of basal region of
spike missing. Note. Counterpart image reversed to be in the same orientation as the part.
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 31
beneath the spike but are too poorly preserved and lack direct
connection to warrant further consideration. However, the
subtending axis to the spike aligns with an axis 4 mm wide (figs.
8, 9) and is suggestive of derivation from the same spike. This
suggests the linear aerial extent of cover of the plant was at least
45 mm wide. Furthermore, the horizontal orientation of this axis
to the spike is suggestive of a rhizomatous system, but it remains
equivocal due to the absence of reticulum axes and H- and
K-branching (sensu Hao et al., 2010: fig. 3; Walton, 1964: fig. 1).
To the right of the apical region of the spike on the part, there
are at least four axes that do not possess any attached sporangia,
and beneath these axes towards the middle of the spike, a poorly
preserved axis is visible with two sporangia (fig. 9) not directly
attached to it but with their sporangial stalks orientated towards
it, suggesting it was once attached. The alignment of these axes
with the spike may indicate a tuft habit, but without clear
evidence of additional spikes, its habit remains inexplicit.
Remarks. The description is based on one specimen — 45 mm
high, part and counterpart with one spike (fig. 4) with sporangia
laxly arranged on vascularised long stalks — preserved as a fine
film of carbonaceous material lacking anatomy. The specimen
occurs with three specimens of Yarravia sp. Lang and Cookson,
1935, on the same plane (McSweeney et al. 2021a: fig. 5a-d).
The limits of the fertile axis and stalks are defined by grey film
on each side of a much darker vascular trace. The fine slender
nature of the darkened linear structures below the stalks are
too narrow to support a sporangium, such that it seems
parsimonious for the original widths of the axes to be defined
by these ghosted grey areas. Lele and Walton (1961: 471), when
describing axes prepared from acetate transfers, found the
xylem to appear as a preferentially preserved dark bands (and
to be about one sixth the axial width) and noted the vascular
traces were often displaced from their central position. This,
they postulated, was likely due to decay of the cortex prior to
burial during early digenesis. This would help explain the
convoluted nature of the vascular trace herein (figs. 4-6),
indicating the structure of the axes had already started to break
down before becoming fully fossilised.
The specimen possesses depressions and protuberances
along parts of its fertile axis, which is especially noticeable
midway along the spike (fig. 6). These are interpreted as likely
insertion points for some axes of sporangia and follows
Edwards’ (1975: 255) interpretation of a similar feature on Z.
myretonianum. Xue (2009: 507), in describing Z.
minorstachyum, suggested that small conical protuberances
along the axes may reflect parasitism. This possibility was
considered, but the irregularities on the vascular trace are
primarily depressions in areas noticeably lacking sporangia,
and in some cases appear to be the basal-most attachment of
the stalk to the fertile axis’ vascular trace. Additionally, we did
not consider areas lacking in sporangia to be indicative of a
deciduous spike, as seen with Z. deciduum from the Emsian,
Lower Devonian of Belgium (Gerrienne, 1988). While it is
plausible that once the more mature proximal sporangia has
dehisced and subsequently abscised, plants would be better
served by losing some sporangia in this region to concentrate
energy on immature sporangia in the distal region of the spike.
However, the specimen still possesses large proximal sporangia
and only some sporangia appear to be missing, suggesting that
= p > i J ;
> i h 2 E > + J
, "3 d AME
_ F . J a “a u
Figure 5. cf. Zosterophyllum sp. A. (part, NMV P256740.1): a, two isolated sporangia at arrows (pre-dégagement) with basal region of both
sporangia orientated away from the spike; b—d, on the reverse of the slab, isolated sporangia with much similar dimensions and weakly developed
sporangial lobes. Image a taken by Rodney Start © Museums Victoria.
32
they may have been lost, most likely as a result of excision due
to the biostratinomy phase (Jackson, 2010: 5) of fossilisation.
The absence of a junction at the axial—sporangial interface does
not mean it never existed because it may have been destroyed
during fossilisation. When examining Llanover specimens of
Zosterophyllum from the Old Red Sandstone of South Wales,
Edwards (1969a: 924) found organs could be superimposed and
amalgamated into the surrounding tissue during preservation,
resulting in them been indistinguishable.
The specimen described herein is atypical in comparison
with most Zosterophyllums because of paucity of folded
sporangia seen in lateral view with only two proximal sporangia
so preserved. Furthermore, the sporangia rarely overlap each
other, with one instance occurring in the proximal region of the
spike where sporangium two has been pushed onto the basal
region of sporangium three (fig. 4b, arrow at stalk of sporangium
two) and distally for sporangia 19 and 20 (fig. 4).
Comparison with other taxa. The sporangia of the specimen
are borne alternatively in two rows on opposite sides of the
axis, akin to Platyzosterophyllum, and so the specimen was
compared to Platyzosterophyllum first. However, some
Platyzosterophyllum possess sporangia emanating from two
rows on one side of the axis, such as Z. cf. fertile in Wellman
et al. (2000: 181) and are noticeably more compact. The stalks
of Z. fertile are perpendicular to the fertile axis, before sharply
turning towards the apex, such that they are borne in an upright
to slightly recumbent position (Wellman et al. 2000: 181). This
characteristic of recurved stalks perpendicular to the fertile
axis 1s also seen in Z. spectabile Schweitzer, 1979, according
to Gensel (1982: 662). However, the specimen clearly differs
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
from these taxa because the stalks are orientated at acute
angles of 15°—45° without any noticeable change in orientation,
other than immediately below each sporangium, where they
sharply reorientate upright and parallel to the fertile axis (figs
5a, 6, 7a). Furthermore, the sporangia of Z. fertile are oblate
(Wellman et al. 2000: 183), being almost linear along the
margins, while the specimen’s sporangia are rounded to
reniform. The dimensions of both taxa also differ slightly,
with Z. fertile possessing stalks that are much narrower than
the 1.0-1.2 mm width for the specimen, with Z. fertile at most
reaching 0.5 mm wide (Edwards, 1972) but generally
(including for Z. cf. fertile) 0.3—0.4 mm wide (Edwards, 1969a;
Leclercq, 1942; Wellman et al. 2000). The sporangial
dimensions for Z. fertile are, in part, similar to the specimen,
with the sporangia of Z. fertile up to 2.3 mm wide (Edwards,
1972), and for Z. cf. fertile specimens the sporangial dimensions
were 2.0-2.3 mm wide (Edwards, 1969a; Leclercq, 1942;
Wellman et al., 2000). The specimen’s sporangial widths vary
more greatly on the same spike and range between 0.95-3.7
mm wide, suggesting the plant was not mature. In comparison
with the Welsh specimen, Z. llanoveranum, sporangia are
arranged in 1—2 alternative rows but differs from the specimen
with sporangia borne close together and in the distal region of
the spike, sometimes helically arranged (Edwards, 1969b).
This could not be confirmed here because the stalk insertion
points are lacking.
Edwards (1975: 263) cautioned against the use of the
arrangement of the sporangia on the spike as a definitive
characteristic with which to delineate species. Edwards noted
bilateral symmetry basally in the spike with the distal part
al
line drawing of medial region of the spike seen in B; b, the cortex (Ctx) is
represented by a tincture of light grey in comparison to a darker coloured vascular trace (Vt). Some perturbations and depressions (*) likely
represent additional stalks. Note small proximal sporangia (at arrow) in a central position on the fertile axis; c, distal part of spike — lower arrow
at stalk that has been pushed across the fertile axis. At the upper arrow sporangium with no stalk, partially behind another sporangium. Smaller
sporangia more centrally located suggests distal part of spike may be helically inserted. Images taken by Rodney Start © Museums Victoria.
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 33
helically arranged in some specimens of Z. myretonianum made similar observations, adding that the difference may
and attributed it to the compression of widely spaced spirally also be due to different ontogenetic stages of individual
arranged sporangia, giving this misleading appearance spikes, and Gensel (1982) noted for Z. divaricatum, sporangia
(Edwards, 1975: 261). Furthermore, Gerrienne (1988: 328) bending and twisting of sporangia to one side.
n b, d, respectively. The sporangia have
weakly developed lobes (Lb), and a lack of a clear junction between the stalk and sporangium. A fine border (Br) is visible only alone distal
margins.
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
sear
FII dui
PERPTT LLL iiti
Figure 8. cf. Zosterophyllum sp. A. (part, NMV P256740.1): a, proximal region of spike; b, interpretative line drawing. Sporangia one and two
(S1-2) appear infolded, with both abaxial (Abx) and partial adaxial (Adx) valves visible in sporangia and two perturbations and depressions (*)
possibly representative of additional stalks that were not preserved. QI, shadowing of possible sporangium, and upper arrow shows change in
orientation of stalk beneath sporangium four.
SOSA, Coco a ER ces
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curving into ?rhizomatous region (arrow 1) and possibly extending out towards axes (arrows 2 and 3). Isolated sporangium (S) in same orientation
as sporangia in spike. Image taken by Rodney Start © Museums Victoria.
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 35
In comparison with species within the subgenus
Zosterophyllum with reniform sporangia, the specimen is
closest to Z. bifurcatum Li and Cai, 1977; Z. deciduum; Z.
myretonianum Lang, 1927; Z. ramosum; Z. rhenanum; Z.
yunnanicum Hsü, 1966; and Z. shengfengense, all of which
have sporangia in approximately the same size range.
Zosterophyllum myretonianum is one of the best studied
Zosterophyllum to date (Edwards, 1975; Lang, 1927; Lele and
Walton, 1961). Edwards (1969: 261) noted when examining Z.
myretonianum from Aberlemno, Scotland, that they possessed
spikes with different levels of sporangial packing, such that
the specimens could be divided into compact, intermediate
and laxly arranged spikes. The sporangial-stalk interface of
Z. myretonianum, according to Edwards (1975), possesses a
dome-like region at the point of insertion on the sporangium
of some of the specimens, which produces its reniform shape.
It is noticeable that in Z. myretonianum, despite different
stages in development, the orientation of the sporangial stalk
remains largely constant, with the sporangial stalk inserted at
almost 90° (Edwards, 1975) to the fertile axis before curving
upwards immediately with the sporangium held erect. This
clearly differs from the specimen where the sporangial stalks
extend from the fertile axis, curving upwards only just
beneath the sporangium and in some cases attached to the
sporangium at an angle, thus producing a splayed appearance.
Zosterophyllum bifurcatum possess well-developed lobes and
much narrower stalks than the specimen reaching up to 0.6
mm wide according to Li and Cai (1977) and Hao and Xue
(2013). Zosterophyllum rhenanum also possess well-
developed sporangial lobes and has a noticeable junction
between sporangium and stalk, and a large border of 0.6 mm
(Hao and Xue, 2013; Schweitzer, 1979). Zosterophyllum
deciduum has weakly developed sporangial lobes (Gerrienne,
1988: 322), similar to the specimen, but the sporangial stalks
were wide (0.4—0.75 mm) near the fertile axis and narrow
(0.1-0.3 mm) near the sporangium (Gerrienne, 1988: 320),
with the contact between the sporangium and subtending
stalk producing a clear junction with no evidence of widening
beneath the sporangia (Gerrienne, 1988: 331). These
characteristics are at odds with what is observed with the
specimen where sporangial stalks remain parallel in width
before widening into the base of the sporangium. Furthermore,
Z. deciduum bifurcates both below and within its fertile parts
(Gerrienne, 1988).
In comparison with Zosterophyllum from the South China
plate with similar sporangial dimension, Zosterophyllum
shengfengense from the Lochkovian, Lower Devonian of
Xitun Formation, Yunnan, China, differs from the specimen
in not possessing any sporangial basal lobes, and shorter
stalks, O0.5—0.8 mm wide and 0.8-1.6 mm long (Hao et al.,
2010; Hao and Xue, 2013). Furthermore, Zosterophyllum
shengfengense (Hao et al., 2010: 222), like Z. myretonianum
(Lele and Walton, 1961: 471), possess tubercles proximally on
the plant, unlike the specimen (Hao et al., 2010: fig. 2a).
Zosterophyllum yunnanicum from the Xujiachong Formation,
Yunnan, possess crowded spikes with up to 50 sporangia
circular to elliptical in face view, dehiscence zone up to 0.5
mm wide, stalks 0.3-0.9 mm wide and 0.6-3.0 mm long
inserted an acute angle to the fertile axis and widening into
the bases of sporangia (Edwards et al., 2015: 223). The stalks
emanate perpendicular to the spike, based on Edwards et al.
(2015: pl. 4, figs 1, 2) and immediately reorientate producing
30°-40° to the fertile axis (Wang, 2007: 528). This
reorientation of the stalks near the fertile axis differs
significantly from the specimen, where the stalks reorientate
only just beneath each sporangium. Furthermore, Z.
yunnanicum produces a dome-like structure at the stalk-
sporangium interface (Edwards et al., 2015).
Comparison with known Victorian zosterophyll taxa. Only four
zosterophylls have thus far been described from Victoria. These
include Z. australianum Lang and Cookson, 1930; Z. ramosum
Hao and Wang, 2000; Parazosterophyllum timsiae McSweeney
et al., 2020; and Gippslandites minutus McSweeney et al., 2020.
Both Z. australianum and Z. ramosum occur in the Norton
Gully Sandstone Formation of Victoria and are younger than
the specimen, which is currently only known from the
underlying Wilson Creek Shale. Zosterophyllum australianum
occurs at North Road Quarry, Walhalla, Victoria, and Yunnan
(Posongchong Formation), China (Hao and Xue 2013; Lang and
Cookson 1930). Zosterophyllum australianum possess
sporangia that are noticeably larger than the specimen and are
longitudinally elliptical or fan-shaped, 2.8—8.0 mm wide and
2.2-5.0 mm high, with short stalks inserted on the fertile axis at
90° (Hao and Xue, 2013; Lang and Cookson, 1930).
Zosterophyllum ramosum occur at Mount Pleasant and Halls
Flat Road, Alexandra (Cookson, 1935; Hao and Wang, 2000).
Mount Pleasant Road is the type locality of Yarravia (Hedeia)
corymbosa Cookson, 1935, and cf. Baragwanathia longifolia,
cf. Yarravia oblonga, ct. Hostinella and Pachytheca sp. have
been found by Cookson (1935) to occur with Z. ramosum
(McSweeney et al., 202la, b). Zosterophyllum ramosum was
originally called Z. australianum by Cookson (1935: pl. 10, figs
9-12), but was later reinterpreted by Hao and Wang (2000: 31)
to be a new species Z. ramosum, which also occurs in Yunnan
(Posongchong Formation), China. Zosterophyllum ramosum
possess circular to reniform sporangia similar to the specimen,
but the sporangia are larger, being 1.6—6.0 mm wide and 1.9—5.5
mm high, on stalks up to 5.0 mm inserted on the fertile axis at
15°-35° (Hao and Wang, 2000; Hao and Xue, 2013). Both Z.
ramosum and Z. australianum, according to Hao and Xue
(2013: fig. 6.5), possess apple-shaped Za-type sporangium with
extended thickened margins, a character not found in the
specimen. Parazosterophyllum timsiae is from Ghin Ghin
Road, Yea, in the base of the Humevale Formation and based on
Rickards & Garratt (1990) Pridoli, upper Silurian—Pragian,
Lower Devonian, and may be either coeval or older than the
specimen and differ significantly from the specimen with its
spike terminating lateral branch (McSweeney et al., 2020).
Gippslandites minutus is from an outcrop of the Boola formation
(Lochkovian—Pragian, L. Devonian) near Boola Quarry, Tyers,
Victoria (Tims, 1980; McSweeney et al., 2000). The Boola
formation is slightly older than the Wilson Creek Shale, which
overlies the Boola formation at Coopers Creek according to
Edwards et al. (1997: 39). Gippslandites minutus differs from
the specimen because its sporangia are much smaller, 0.6—2.6
36
mm wide and 0.3-1.9 mm high, and differ significantly from
Zosterophyllum spp. with anisovalvate sporangia (McSweeney
et al., 2020).
The defining characteristic of the specimen is primarily the
angle of insertion of the vascularised stalks and no overlap
between vertical adjacent sporangia. As noted by Edwards
(1975: 264), the most useful characters in species delimitation
within Zosterophyllum are stalk and sporangial characters. It is
clear that the specimen differs from zosterophylls from Victoria
primarily on sporangial morphology and symmetry. As the
sporangial stalks were likely longer in life when turgid and prior
to degradation resulting in convoluted vascular trace, the lack of
vertical overlap of sporangia and vascularisation of the stalks,
and clear demarcation of insertion points on the fertile axis
means the specimen cannot be readily put into the subgenus
Platyzosterophyllum, and 1s thus assigned to cf. Zosterophyllum
sp. A. until better material becomes available to allow for further
assessment of its phylogenetic and taxonomic position.
Dr +
E d
PEN d
Titi
UEM s^ "ao a
FR 7!
|
rc
E
Bi
-
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
cf. Zosterophyllum sp. b.
Gen. et sp. indet.
Figures 10, 11
Material. NMV P256742.1 and P256742.2 (P4-5 field note identifier),
part and counterpart, respectively.
Locality. P4 is equivalent to Loc. | in Garratt (1978: fig. 2), and
occurs on Ghin Ghin Road, 8 km northwest of Yea township,
central Victoria.
Horizon and age. Humevale Siltstone, Pragian—Emsian, Lower
Devonian (Edwards et al., 1997; Garratt, 1978; Rickards, 2000;
Rickards and Garratt, 1990; VandenBerg et al. 2000; VandenBerg
pers. comm. June 2021).
Description. Single specimen, comprising compact spike, with
part and counterpart preserved in semi-relief as an iron oxide
coated impression and cast, only gross morphological features are
visible (fig. 10). The spike measures 21 mm high and 10 mm
2mm
Figure 10. cf. Zosterophyllum sp. B, from the Humevale Siltstone, Ghin Ghin Road, Yea: a, part (NMV P NMV P256742.1); b, counterpart (NMV
P NMV P256742.2) respectively; c, line drawing, with SI-9 (sporangia one to nine). Note: Part image reversed to be in the same orientation as
the counterpart. Note. Arrow in part, points to sporangium 4, which is obscured in the counterpart.
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 37
oo ^
p «500 um .
hd
Av mmi
ae
U
Figure 11. cf. Zosterophyllum sp. B. NMV P256742.1: a, c, (line drawing) sporangium 3 (counterpart) in-folded, with stalk attachment widening out
in basal region of sporangium. Poorly defined dehiscence zone (zone) and line; b, sporangium one (counterpart) with border visible; d, basal oval
structure (part), with what appears to be a much smaller spike emanating from it (at arrow), and a poorly preserved axis beneath the oval structure.
38
wide, with at least nine sporangia arranged helically, becoming
more compact distally. The fertile naked axis measures 1.5-3.75
mm wide, narrowing distally before terminating in sporangium
nine. Sporangia in face view (1.2—3.0 mm wide and 3.9-4.3 mm
long) occur in the distal part of the spike and are circular to
oblong. Proximally, sporangia one to three are in lateral view,
being longitudinally elliptical and infolded. Sporangia are inserted
at an angle of ~20°. Proximally on the spike there is an elliptical
(lateral view) junction at the point of attachment (face view) on
the abaxial valve between the stalk and the sporangium. The
stalks are decurrent, narrowing slightly before widening distally
at the base of the sporangium. The sporangia possess a dehiscence
zone along their entire distal margins, with a border 0.1—0.2 mm
wide gradually tapering proximally towards the stalk attachment.
Remarks. There is a notable disparity in size between the much
larger proximal sporangia (S1—5) and distal sporangia (S6-9),
suggesting the spike may be immature. Sporangia one-three are
in a lateral position and are infolded, a common feature seen
especially in laterally placed sporangia of Zosterophyllum. It
suggests that the sporangia may have originally had relatively flat
bodies, as proposed by Lang (1927) when explaining this feature
in Zosterophyllum myretonianum. The points of attachment of
the stalk to the sporangia are visible on sporangia one and three
(Figs lla, c), with the stalks widening into the base of the
sporangia. The attachment is elliptical, and it is possible that the
raised regions defining the elliptical region, which appear
partially raised, may reflect the splitting of the attachment at the
base of the sporangium, possibly feeding each of the valve.
However, the preservation 1s too poor and this remains equivocal.
The attachment occurs on the adaxial side of the valve, but the
demarcations between both valves are faint or absent on the
sporangia. Most of the distal sporangia show the stalk centred
beneath the sporangium, with a gradual widening of the stalk into
the valve. Sporangium seven, which is in face-view, appears to
show the widening of the distal part of the stalk on the lower part
of a valve, producing a sub-circular zone of attachment.
Sporangium five, while poorly preserved, is in side view and
ereatly compressed laterally with the projecting edge of the
border visible in the apex, and a fine stalk 1s present 1n the basal
region. The adaxial valve only appears marginally darker than
the surrounding matrix and is convex, while the upper valve is
flatter and delimited by its 1ron-oxide colouration. Sporangia four
and six also appear to show a slight separation at the distal-most
region of the valves. This may be a result of compression as the
sporangia are small relative to the proximal sporangia, suggesting
they were not fully mature.
The dehiscence line can be seen along the distal margins of
the proximal sporangia, visible down to the stalk attachment
(fig. llc) but is only visible on some of the distal sporangia.
There is a dehiscence zone beside the dehiscence line no wider
than 0.2 mm and is clearest on the proximal sporangia.
The proximal part of the fertile axis expands into an oval-
shaped body (fig. 11d) that measures 3.74 mm wide and 5.80
mm high. The point where the axis starts increasing in
diameter was taken as the start of this structure because there
is no other way to differentiate it from the axis. The oval body
is similar to the corm-like structure found basally on
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
Horneophyton lignieri Barghoorn and Darrah (= Hornea
lignieri Kidston and Lang, 1920). However, because no
anatomy is preserved, its nature remains equivocal and may
be a quirk of preservation. A small, fine, faint linear structure
(fig. 11d) appears to emanate from the oval body, but remains
equivocal because sampling did not reveal any organic
remains and high magnification did not reveal any
morphological characters.
The specimen fits into the class Zosterophyllopsida based on
the presence of naked axes, cauline sporangia made up of two
valves that dehisce along their distal margins, and vascularized
stalks (Croft and Lang, 1942). The Zosterophyllopsida includes
two orders, Zosterophyllales and Gosslingiales, and because this
specimen possessed a terminal sporangium, it has been assigned
to order Zosterophyllales. Numerous characteristics are notably
absent, such as H- or K-branching and circinate vernation.
Because the sporangia were not in rows, the specimen was
excluded from the subgenus Platyzosterophyllum and tentatively
placed into the subgenus Zosterophyllum. However, it was
difficult to determine the true morphological outline of the
sporangia, with some of the distal sporangia appearing slightly
longer than wider. Furthermore, the absence of sporangia
proximally in face view added to the uncertainty of whether the
sporangia were slightly vertically elongate. Distally, the
sporangia are poorly preserved with no unequivocal demarcation
of both the valves and junctions between the valves and their
subtending stalks. For the subgenus Zosterophyllum, the
sporangia are reinform, fan-shaped and isovalved, and are
excluded if anisovalvate or vertically longer than wide according
to Edwards et al. (2016) and Edwards and Li (2018a).
Additionally, the presence of an oval region at the proximal end
of the fertile axis added further doubt to its true assignation, and
so the specimen was placed into cf. Zosterophyllum sp.
cf. Zosterophyllum sp. B differs from known Zosterophyllum
spp. from Victoria (Z. australianum and Z. ramosum), despite
the limited characters available for comparison. Zosterophyllum
australianum possess horizontally elliptical sporangia with
large, thickened margins (0.4-1.1 mm wide) and sporangia far
larger than cf. Zosterophyllum sp. B., reaching up to 8.0 mm
wide and 5.0 mm high (Hao and Xue, 2013; Lang and Cookson,
1930). Additionally, the stalks of Z. australianum are inserted at
an angle of ~90°, in contrast to ~20°° in cf. Zosterophyllum sp.
B. Zosterophyllum ramosum, like Z. australianum, possess much
larger sporangia, reaching up to 6.0 mm wide and 5.5 mm high,
and are circular to reinform in shape (Hao and Wang, 2000).
In comparison with other zosterophylls with vertically
elongate sporangia outside the subgenus Zosterophyllum, cf.
Zosterophyllum sp. B. appears distinct, despite the paucity of
available characters with which to compare. For example,
Guangnania cuneata Wang and Hao, 2002, possess anisovalvate
upright sporangia, but differ from cf. Zosterophyllum sp. in the
spike not being compact, the sporangia being much longer than
wide (3.9-6.2 mm high and 1.5-19 mm wide) and not
possessing a terminal sporangium. Yunia dichotoma Hao and
Beck, 1991, from the Zhichang section of the Posongchong
Formation, differs from cf. Zosterophyllum sp. B in the
morphology of the sporangia, which are elongate-elliptical to
ovoid, but the sporangia did not form spikes (Hao and Beck,
Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance 39
1991). Additionally, the axes of Y. dichotoma possess small
spines (Hao and Beck, 1991). Huia recurvata Geng, 1985, of the
Posongchong Formation, Yunnan, China, produced sporangia
that were ovate to ovoid 2D to 3D (Hao and Xue, 2013: 70). Huia
recurvata differs in having adaxially reflected closely arranged
sporangia as opposed to the crowded arrangement seen 1n cf.
Zosterophyllum sp. B., with sporangia 3-5 mm wide and 6-10
mm high (Hao and Xue, 2013).
Primarily, because the specimen lacks unequivocal evidence
as to the characteristics of the sporangia, such as whether the
sporangia are longer than wide, it has been placed in cf.
Zosterophyllum sp. B.
Discussion
The term sporangial stalk is used with regards to cf.
Zosterophyllum sp. A. for convenience, similar to Edwards
(2006: 96), but it is possible they represent lateral branches with
sessile sporangia, as proposed by Hueber (1992: 480) in his
examination of zosterophyll sporangial and based on his
hypothesis that some cooksonioids were the progenitor of this
eroup. The presence of a vascular trace in the sporangial stalk of
cf. Zosterophyllum sp. A. adds credence to Hueber’s hypothesis
(category 3) and has been observed with Z. divaricatum (Gensel,
1982: 654), Z. myretonianum (Edwards, 1975: 262; Lang, 1927:
450), and according to Niklas and Banks (1990: 278), Z.
rhenanum and Z. longhaushanense. Niklas and Banks (1990:
277) proposed two patterns of fertile growth within zosterophylls
governed by apical meristematic growth and speculated the
difference in symmetry for Z. myretonianum, Z. fertile and Z.
llanoveranum was due to uncanalised growth and that the
nature of the apices (terminate and non-terminate) was the key
to determining the type of meristematic growth. Interestingly
Niklas and Banks (1990: 278), in discussing bilateral symmetry,
postulated that the stalks were not the same as fertile axes and
the lateral cluster of initials producing the stalks and sporangia
would likely result in limited to no vascularisation of the stalk.
This limited vascularisation of lateral axes appears to occur
with cf. Zosterophyllum sp. A., whereby the apical initials of
lateral axes were limited to just a single axis before terminating
in a sporangium.
Both cf. Zosterophyllum sp. A. and cf. Z. fertile possess
sporangia arranged in rows on their spikes, and this 1s the first
tme this arrangement has been described for Victorian
Devonian flora. The zosterophyll assemblages from China,
particularly those from the Pragian Posongchong Formation of
the South China Plate, have predominantly been of sporangia
inserted helically into the spike (Hao & Xue, 2013). Only three
cases of sporangia occurring in rows in spikes have been
recorded: Zosterophyllum longhuashanense Li and Cai, 1977;
Distichophytum sp., and Amplectosporangium (Oricilla)
unilaerale Edwards and Li, 2018 (Edwards and Li, 2018; Hao
and Xue, 2013). Another, Ornicephalum (Zosterophyllum)
sichuanense Edwards and Li (2018: 100), superficially appears
to show two-rowed sporangia. The geographical location of
Victoria during the Pragian—Emsian, Lower Devonian,
approximately 30° south of the equator, far from both the South
China Plate and other known Zosterophyllum localities, suggests
two of the specimens described herein may be new taxa. The
palaeocontinental positioning of Victoria and the South China
Plate during the Lower Devonian is based on the work of
Mitchell et al. (2012) and Torsvik and Cocks (2019), and is
summarised in McSweeney et al. (2020). Australia and South
China have only two species in common thus far — Z.
australianum and Z. ramosum — and the specimens described
herein further add to the hypothesis of limited floristic exchange,
as espoused by Xue et al. (2018: 98).
Conclusion
This study has confirmed the first occurrence of bilateral
symmetry in Victoria. Despite the dearth of fossil evidence,
three additional forms — cf. Zosterophyllum sp. A, cf. Z. sp. B.
and cf. Z. fertile — indicate greater diversity in the region
during the Silurio—Devonian than previously known.
However, forms with better preservation are required to
determine their taxonomic positions with certainty.
Acknowledgments
We thank Prof. Dianne Edwards of Cardiff University and Dr
Jackie D. Tims for generously giving us the specimen cf.
Zosterophyllum sp. A to describe and Prof. D. Edwards for her
feedback; Rodney Start and Tim Ziegler for the photography
and organising the use of a Leica microscope respectively, at
Melbourne Museum and RMIT University’s Microscopy and
Microanalysis Facility; Dr Muthu Pannirselvam at RMIT
University, Melbourne; Paul Ter for assisting the lead author ın
viewing the Wilson Creek Shale outcrop on Frenchmans Spur
Track in October 2018; staff at University of Melbourne’s
Baillieu Library for access to J. D. Tims’ PhD thesis; Dr
Michael Garratt for his assistance and advice; and Mrs Peg
Lade in allowing access to her property on Ghin Ghin Road,
the location of P4. This work is part of a Ph.D. thesis by the
lead author, and was supported by an Australian Government
Research Training Program Scholarship.
Disclosure Statement
No potential conflict of interest was reported by the authors.
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Memoirs of Museum Victoria 81: 43-53 (2022) Published 2022
1447-2554 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2022.81.03
Taungurungia gen. nov., from the Lower Devonian of Yea, central Victoria, Australia
FEARGHUS R. McSwEENEY FGS'", JEFF SHIMETA’, JOHN S". J.S. BUCKERIDGE FGS'?
! Barth and Oceanic Systems Group, RMIT University, GPO Box 2476, Melbourne, VIC 3001, Australia
* School of Science, RMIT University, Melbourne, VIC 3001, Australia
> Museums Victoria, VIC 3001, Australia
* To whom correspondence should be addressed. Email: tidal’75@gmail.com
McSweeney, F.R., Shimeta, J. and Buckeridge, J.S. 2022. Taungurungia gen. nov., from the Lower Devonian of Yea,
This paper records a new genus Taungurungia, which is the first new taxon with emergences to be described from
the Lower Devonian of Victoria. The fossil is preserved primarily as a compression and impression, and lacks internal
anatomy. The fossil extends our knowledge of known variations within early land plants, with most characteristics, such as
emergences and H- or K-branching, redolent of affinities with the zosterophylls. However, having a large ovate terminal
sporangium, the fossil adds to taxa that in some cases have been provisonally allied to the zosterophylls with elongate
Abstract
central Victoria, Australia. Memoirs of Museum Victoria 81: 43-53.
sporangia; this further demonstrates the need for reassessment of the Zosterophyllopsida.
Keywords embryophytes, zosterophyll, emergences, Lower Devonian, Victoria
Introduction
Banks (1968, 1975) recognised three subdivisions from the
Psilophytales: the Rhyniophytina, Zosterophyllophytina and
Trimerophytina. Subsequently, these have been raised to
higher taxononic ranks by many workers (Hao and Xue, 2013;
Kenrick and Crane, 1997). Herein, we examine a new form
similar to Zosterophyllopsida sensu Hao and Xue (2013).
Zosterophyllopsida have sporangia that have a width to height
ratio >] and are reniform, globose or rounded in face view
(Banks, 1968, 1975). However, some rare exceptions have
been noted where the sporangia are longer than wide
(Edwards et al., 2016; Edwards and L1, 2018; Hao and Beck,
1991; Hao and Xue, 2013; Wang and Hao, 2002). The
specimen described herein augments taxa with longer-than-
wide sporangia and 1s significant because little is known about
these aberrations and their taxonomic significance. Four
Zosterophyllopsida species are known from Victoria:
Zosterophyllum australanium Lang and Cookson, 1930; Z.
ramosum Hao and Wang, 2000; Parazosterophyllum timsiae
McSweeney et al, 2020; and Gippslandites minutus
McSweeney et al., 2020. But none of these Zosterophyllopsida
resemble the specimen descibed below. Furthermore, axes
with emergences are rare in Devonian Victorian flora and
have only been described by Cookson (1935) and McSweeney
et al. (2021) from the Norton Gully Sandstone Formation of
Alexandra; axes resembling Psilophyon were noted by
Williams (1964: 285) at location A82 on Kerridale Road,
Homewood, near Yea.
Locality, stratigraphy and age
Location F103 (sensu Garratt, 1980) occurs in the Norton
Gully Sandstone Formation according to Edwards et al. (1997,
1998) and VandenBerg et al. (2000). Garratt (1980: 590) did
not record any fossils at F103. The Norton Gully Formation is
found throughout the Mount Easton Province and parts of the
Darraweit Guim Province (fig. 1) according to Edwards et al.
(1997: fig. 6; Vanden Berg et al. 2000: fig. 2.106). The exposure
is on the northern side of the Goulburn Valley Highway (fig.
2) and consists of a sequence of siltstones that are 1ndicative
of quiescent conditions during deposition. The Norton Gully
sandstone Formation is considered to be Emsian age by
Morand and Fanning (2006) and, based on the occurrence of
Uncinatograptus thomasi (Jaeger, 1966), is considered to be
Pragian and possibly Emsian age by Lenz (2013). Therefore, a
late Pragian—Emsian, (Lower Devonian) age is given herein to
account for these minor disparities.
Material and methods
NMV P257028.1 (part) and NMV P257028.2 (counterpart)
were found in situ in January 2015 by the lead author with
Michael Garratt. These buff yellow fine-grained siltstones
easily split along bedding planes, with the fossil preserved as
an iron-stained mould and cast with some carbonaceous film.
The specimen was photographed (figs 3a, b, 4a, 5, 6, 7a) by
Rod Start of Museums Victoria on 17 March 2020, using cross-
polarised light (circular filter) with a Canon 5DSR camera fitted
44
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
Australia
New South Wales
u E =
Victoria 3:
"
; EL
EN i
' Si
Melbourne N
t
Lf Wilsons Promontory
Darraweit Guim Province
‚King
[Jisian Bass Strait M Mount Easton Province
Ysland * Fossil location (Yea)
We Island
$2"
Tasmania
ES»
W
)_ Guba Riger
Figure 2. Goulburn Valley Highway fossil location F103 (Garratt 1980): a, F103 located about 5 km west of Yea; b, part of F103 exposure (37°
I? 30" S; 145° 21’ 56" E) with arrow at location where Taungurungia garrattii gen. et sp. nov. was found in situ. Map after Garratt (1978) and
positioning of Strathbogie Batholith after VandenBerg (1997).
Taungurungia gen. nov., trom the Lower Devonian of Yea, central Victoria, Australia 49
with an EF 100 mm f/2.8L macro lens. Flash strip-lights were
used to enhance contrast. The remaining images were taken
using an AxioCamMRcS5 camera attached to a Zeiss SteREO
Discovery V8 stereomicroscope. Images were Z-stacked to
improve depth of field using Adobe Photoshop CC 2017. ImageJ
software (Image J1.52d, Wayne Rasband, USA, http://1image].
nih.gov/ij) was used to take measurements. Extensive
dégagement (primarily following Fairon-Demaret et al., 1999)
was undertaken by the lead author, mainly on NMV P257028.1,
with a sample removed from an uncovered axis for analysis in
low-vacuum mode on a FEI Quanta-200 scanning electron
microscope at the RMIT Microscopy and Microanalysis
Facility. No anatomical information was obtained. Dégagement
of the distal region of one of the daughter axes on the counterpart
(NMV P257028.2) revealed terminal small fusiform bodies
along the axis (fig. 7a pre-degagement; fig. 7b post-degagement).
Degagement above the large terminal sporangium (fig. 3 pre-
dégagement; fig. 4d post-dégagement) revealed small fusiform
bodies near an axis that was poorly preserved. This dégagement
was undertaken by gently loosening the overlying matrix above
the spike by gently tapping the steel needle on the matrix to
remove grains and work down to the fossil. Degagement of
proximally curved axis revealed additional minute fusiform
bodies (fig. 7c, d post-degagement).
Institutional abbreviations
NMV P, Museums Victoria Palaeontology Collection,
Melbourne, Australia. The specimen NMV P257028.1 and
NMV P257028.2, part and counterpart respectively, are housed
in the Palaeontological section, Museums Victoria, Melbourne.
Systematic palaeobotany
Class. /nsertae sedis
Genus. Taungurungia McSweeney, Shimeta and Buckeridge
cen. nov.
Diagnosis: lerete axes with deltoid to elongate emergences.
Fertile axes terminate in single large sessile ovate sporangium
partially sunken into the axis with a curved junction between
the axis and sporangium. Branching anisotomous and ?H- or
K-branching in basal regions. Small fusiform to elliptical
bodies (buds/sterile sporangia) occur on axes including the
distal regions and on slender lateral axes, some with short
stalks.
Etymology: Named for the indigenous Taungurung people.
Type species: Taungurungia garrattii McSweeney, Shimeta
and Buckeridge gen. et sp. nov.
Shimeta and
Taungurungia garrattiti McSweeney,
Buckeridge, sp. nov.
Figures 3—10
Species diagnosis: As for genus. Plant at least 185 mm high with
axes at least 7.8 mm wide. Emergences up to 1.5 mm wide and
3.4mm long. Mature axes terminate in a single ovate sporangium
at least 7.5 mm wide and at least 15.5 mm long. Small fusiform
to elliptical bodies (buds/sterile sporangia) 0.5 mm wide and |
mm long, some with short stalks occurring on all axes.
Etymology: Named for Michael Garratt in recognition of his
work in Palaeozoic palaeontology, especially on brachiopods.
Holotype: NMV P257028.1 (part; F103-1p) and NMV P257028.2
(counterpart; F103-1cp), part and counterpart respectively.
Locality: The exposure occurs on the Goulburn Valley
Highway (B340) 5 km west of Yea and was designated F103 by
Garratt (1980). 37° 12' 30" S; 145° 21' 56" E.
Stratigraphy and age: Norton Gully Sandstone Formation, late
Pragian—Emsian, Lower Devonian.
Taungurungia garrattii sp. nov.
Description (based on one specimen part and counterpart): The
specimen consists of six parent axes that are 2.4—7.5 mm wide
and taper gently acropetally; three of these axes are poorly
preserved. The axes are oxidised and golden to yellow in
coloration, with a vascular trace evident in parts of the axes,
most notably proximally measuring 1.6 mm wide. Three of the
larger axes terminate in a single sessile large ovate sporangium
(fig. 3a). One is poorly preserved with only part of its apical
region preserved (fig. 3b). The largest sporangium is 7.5 mm
wide and 15.5 mm long and is ovate, reaching its maximum
width 6.4 mm up from its base (figs 3, 4a, b). The sporangium
does not possess a stalk, appears sessile and is partially
embedded in the axis with a curved junction. Another large
sporangium occurs low on the specimen (figs 3, 5) and is 14.7
mm long and 5.6 mm wide and is missing part of its cast basally.
This sporangium sits with part of its basal region embedded into
its subtending axis, again with a curved junction. The distal
region of this sporangium is more elongate. These two sporangia
have two fractures running transversely at an oblique angle.
There is no evidence of a bounded region along these fractures
to indicate they are related to dehiscence. Additionally, there 1s
no evidence of a marginal rim preserved on any of the sporangia.
Emergences are 0.6-1.5 mm wide and 1.3-3.4 mm long,
varying in morphology according to length (fig. 6) and more
occur on the largest axis than the other axes. The smaller
emergences are deltoid, and the larger emergences are elongate
and perpendicular to the parent axes. One elongate emergence
extends perpendicular to the parent axis for about 0.3 mm
before re-orientating at about 45° to the axis (fig. 6d). One of
the deltoid emergences has a fine vascular trace 0.07 mm wide
along its length (fig. 6b).
Fusiform bodies (figs 7, 8) up to 0.5 mm wide and | mm
long occur on the axes, most occurring distally but in two
instances are found to occur in discrete areas of no more than 8
mm long and 2 mm wide on the sides of the main parent axes
(fig. 8a, b). Degagement of the distal region of one of the
daughter axes revealed fine fusiform bodies with short stalks
(fig. 7a, b). Additionally, similar fusiform bodies were found to
occur on the narrow lateral axes that emanate from the main
parent axes at almost right angles (fig. 7e), and on the counterpart,
faint impressions of numerous fusiform bodies were found on
the large central terminal sporangium (fig. 4c). Dégagement
46
above the main large terminal sporangium revealed more of
these fusiform bodies extending beyond the large sporangium
(figs 4d, 9). There is no connection between the small fusiform
bodies and the large terminal sporangium.
Three types of branching are evident. The first is where a
parent axis branches anisotomously and is only seen once,
occurring 33 mm from its apex producing a daughter axis 2.9
mm wide that tapers to 1.5 mm wide (figs 3, 7a). The daughter
axis and distal region of the parent axis both appear lax. The
second type of branching occurs along the sides of the parent
axes and consists of narrow lateral axes usually branching
perpendicular from the parent axis. These narrow lateral axes are
about 0.2 mm wide and up to 2—3 mm long and are sparsely
distributed on the parent axes; some occur just beneath the large
terminal sporangium of the central axis (fig. 7e). None of these
narrow lateral axes branch, and they possess two to three
fusiform bodies interpreted as dormant buds or sterile sporangia,
one of which is always terminal. One large example of a lateral
axis occurs proximally off one of the parent axes and is 15 mm
long, varying in width from 0.3—1.1 mm, curving orthotropically
with a terminal fusiform body 0.24 mm wide and 0.53 mm long
(fig. 7c, d). A third type of branching occurs basally on the
central axis with an oblique axial extension forming an
approximate K-branch (fig. 3a), with some smaller protuberances,
possibly remnants of fusiform bodies (fig. 8c), close to each
other. The acroscopic part of this vegetation is similar in
morphology to the distal regions of the two large terminal
sporangia. A portion of axis was extracted from a poorly exposed
axis (fig. 3) within the matrix, revealing longitudinal structures
possibly indicative of the cell’s original orientation (fig. 10).
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
Comments: The location of sporangia has been used by many
workers (Edwards et al., 1989; Gensel 1992; Kenrick and
Edwards, 1988; Niklas and Banks, 1990) to differentiate
zosterophylls into either the Gosslingiales or Zosterophyllales.
Zosterophyllales have both lateral and terminal sporangia.
However, the presence of emergences suggests affinities with
the Gosslingiales; but Gosslingiales lacks terminal sporangia
(Hao and Xue, 2013). Additionally, the sporangia of
Taungurangia garrattii are not reinform or globose as in
zosterophylls, which precludes assignation to Zosterophyllum
(Edwards et al., 2016; Gensel, 1992: 455). Instead, the terminal
sporangia are noticeably elongate and lack a thickened zone
bounding a dehiscence line as seen in most zosterophylls (Hao
and Xue, 2013). The apparent limited branching and single
terminal elongate sporangium suggests rhyniophytes affinities,
but while branching appears limited, numerous slender axes
emanate from the parent axes. Based on the primary
characteristics — notably the presence of emergences, K- or
H-branching, conspicuous large terminal elongate sporangium
and noticeably sparse branching — we provisionally place
Taungurangia garrattii in the Class incertae sedis.
We believe the small fusiform bodies may represent buds
or sterile vestigial sporangia (figs 7, 8). This is a similar
conundrum to the original consideration of what were
originally believed to be sporangia along the axes of Sawdonia
ornata (Dawson) Hueber but were later found to be buds/
arrested apices (Gensel and Berry, 2016: 619; Hueber, 1992:
fig. 3). We found no fusiform bodies of intermediary size with
the large terminal sporangia; instead, they are all broadly the
Same size. Furthermore, most of the fusiform bodies have
""T———I
m
L si
20mm C
Figure 3. Taungurungia garrattii gen. et sp. nov., holotype: a, part specimen (NMV P257028.1) with lower arrow at K-branching. Terminal ovate
sporangia at S1 and 52. At Ax is an axis that goes into the matrix. K- or H-branching at lower arrow; b, counterpart (NMV P257028.2). At upper
arrow 53 is poorly preserved sporangium. Lower arrow points the axis curving upwards which is visible in fig. 7c, d close-up after dégagment.
The counterpart image is reversed to be in the same orientation as the part; c, line-drawing without small bud/sterile sporangial bodies included.
Images a and b were taken by Rodney Start © Museums Victoria.
Taungurungia gen. nov., from the Lower Devonian of Yea, central Victoria, Australia 4f
Figure 4. Taungurungia garrattii gen. et sp. nov., holotype, part specimen NMV P257028.1: a, ovate sporangium pre-dégagement. Left arrow is
at apex of poorly preserved sporangium. Right arrows are at two lateral axes close to the base of the sporangium. Close-up of lower axis in fig.
7e; b, post-dégagement, reveals sporangium preserved in relief; c, faint impressions at arrows of fusiform bodies (buds/sterile sporangia) on
counterpart. Lower arrow shows a fusiform body attached directly to the axis at the junction with the sporangium; d, dégagement above the
sporanium revealed numerous small fusiform bodies (buds/sterile sporangia) at arrows similar to c. Tip of sporangium disintegrated during
dégagement, positioning highlighted with dotted line. Image a taken by Rodney Start O Museums Victoria.
Figure 5. Taungurungia garrattii gen. et sp. nov., holotype, part
specimen NMV P257028.1. Large sporangium low down on the
specimen. The apex is slightly curved (upper arrow) and more elongate
then the sporangium in fig. 4. Branching visible (lower arrow). Note:
the sporangium appears partly embedded in its subtending axis. Image
taken by Rodney Start © Museums Victoria.
: Dar"; Es -È
p ; 1 * 1
CAR ue E
MER. Le WAS
[s
A 3); | -
r te aon it. „al U _ » y «o
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
narrow subtending axes that clearly differ from the large
sporangia, which are sessile. If these fusiform bodies were to
grow to a similar size as the terminal sporangia, they would
likely have caused the plant to become unstable because they
occur on narrow lateral axes and at the end of daughter axes.
The oblique region of extended vegetation (?K- or
H-branching) seen basally is indicative of creeping vegetation,
such as that seen with Discalis longistipia Hao (1989: 159) and
other zosterophylls (Walton, 1964). Branching frequency and
pattern from the emergences could not be determined. The lax
appearance of the daughter axes (fig. 7a) could be interpreted as
being almost recurved, a similar pattern of growth as circinate
axes (Lyon and Edwards, 1991: 327) associated with
indeterminate growth (Niklas and Banks, 1990). However, we
consider this lax appearance to be due to partial wilting because
there is no evidence of recurved growth on the smaller axes,
which would be expected for this characteristic.
Comparison to other taxa
The specimen described herein has some similarities to
Halleophyton zhichangense Li and Edwards, 1997, from the
Zhichang section of the Posongchong Formation, Yunnan,
China, in that rhomboidal depressions (Li and Edwards, 1997:
fig. 5) are found on the axes and bear a passing resemblance to
the depressions found on the sporangium mould (fig. 4c).
However, H. zhichangense has much smaller sporangia, which
are ellipsoidal and globose, 1.9-3.3 mm long and 1.7—3.1 wide
(Li and Edwards, 1997).
Elongate-ovoid sporangia are characters of rhynialeans, but
the presence of emergences and ?K- or H-branching suggests
affinities with the zosterophylls. Elongate sporangia occurring on
taxa with presumed affinities to the zosterophylls broadly similar
to T. garrattii comprises six species (Table 1) that lived on the
south China plate. Five of these (Guangnania cuneata Wang and
Hao, 2002, Guangnania minor Edwards et al., 2016, Sichuania
uskielloides Edwards and Li, 2018, Baoyinia sichuanensis
Edwards and Li, 2018 and Yunia Guangnania Hao and Xue,
Figure 6. Taungurungia garrattii gen. et sp. nov. holotype, part specimen NMV P257028.1: a-d, ontogenetic changes in morphology and orientation
of emergences. In image b, arrow at possible vascular trace extending out to tip. Images were taken by Rodney Start © Museums Victoria.
Taungurungia gen. nov., from the Lower Devonian of Yea, central Victoria, Australia 49
2013) have naked axes and therefore 7. garrattii can be easily
distinguished from them. Yunia dichotoma Hao and Beck, 1991,
does have emergences in the form of sparely distributed spines
and therefore differs from Taungurungia garrattii. Additionally,
Y. dichotoma differs from T. garrattii because 1t has a marginal
rim on the sporangia; this character is also found with Ramoferis
amalia Hao and Xue, 2011, and Baoyinia sichuanensis (Edwards
and Li, 2018). Ramoferis amalia also differs from T. garrattii in
having ovoid to pear-shaped sporangia (Hao and Xue, 2011).
The sporangia of Y. dichotoma differ from those of T.
garrattii in that they are elliptical to ovoid with rounded
apices (Hao and Beck, 1991). The sporangia of these six taxa
are also noticeably much smaller than T. garrattii. The largest
sporangia G. cuneata, 2.8 mm wide and 9.6 mm long
(Edwards et al., 2016; Wang and Hao, 2002), is much smaller
than the largest sporangium of T. garrattii, which is 7.5 mm
wide and 15.5 mm long. Additionally, the largest axial width
for these six taxa (Table 1) is 5.0 mm for Y. dichotoma (Hao
Table 1. Zosterophyllopsida sensu Hao and Xue (2013) with longer than wide sporangia in comparison to Taungurungia garrattii gen. et sp. nov.
Age; formation;
tvpe location
Pragian, Lower Devonian;
Posongchong Formation;
Daliantang, Yunnan, China
Guangnania
cuneata Wang
and Hao, 2002
Loch-Pragian, Lower
Devonian; Pingyipu
Group, Yanmenba section;
North Sichuan, China
G. minor
Edwards et al.,
2016
Loch-Pragian, Lower
Devonian; Pingyipu
eroup, Yanmenba section;
North Sichuan, China
Sichuania
uskielloides
Edwards and Li,
2018
Baoyinia
sichuanensis
Edwards and Li,
2018
Yunia dichotoma
Hao and Beck,
1991
Loch-Pragian, Lower W: 2-3
Devonian; Pingyipu
eroup, Yanmenba section;
North Sichuan, China
Pragian, Lower Devonian;
Posongchong Formation,
Zhichang section; Yunnan,
China
Pragian, Lower Devonian;
Posongchong Formation;
Diliantang, Yunnan, China
Y. Guangnania
Hao and Xue,
2013
Late Pragian—Emsian,
Lower Devonian;
Taungurungia
garraíti gen. et
SPs tows Norton Gully formation;
Yea, central Victoria,
Australia
Note: All dimensions are in mm; L, length; W, width
Emergences
type; width
(basally) and
length
None Elongate-cuneate | Sporangia helically
W: 12-19 arranged in terminal
spikes
L: 1.755.2
None Elliptical-oval
W: 2.8-4.8
L: 4.0-6.0
None
(spike axis)
Axial spines
W: 0.5-1.4
L: 0.6-1.5
None Elongate-
elliptical
Deltoid-elongate
W: 0.6-1.5
L: 1.33.4
Comment Source(s)
Sporangia
(face view)
Elongate-cuneate | Found also in the
Xujiachong Formation.
Lose spikes, stalks at least
Wang and Hao
(2002);
Edwards et al.
(2016: table 3)
5.2 mm long.
Nonterminate
Edwards et al.
(2016: table 3)
Edwards and
Li (2018)
Sporangia laterally
flattened; stalks up to 1.5
mm wide. Spike lax
Edwards and
Li (2018)
Stalks up to 2 mm wide.
Sporangia isovalvate
occurring in clusters; no
vegetative parts
Hao and Beck
(1991); Hao
and Xue
(2013: 119)
Elongate-
elliptical to
ovoid
Peripheral border; axes
sparsely covered in small
spines
Hao and Xue
(2013: 122)
Peripheral border; stalks
<l mm long; sporangia
scatted spirally; isotomous
branching
No stalk evident on large | Herein
sporangia; fusiform and
elliptical bud/sterile
sporangia c.
W: 0.5
L: 1.0
W: 5.3-7.5
L: 14.4-15.5
50 F.R. McSweeney, J. Shimeta & J.S. Buckeridge
and Beck, 1991), and the longest specimen is for G. minor, The emergences on 7. garrattii are broadly similar in
attaining a height of at least 155 mm (Edwards and Li, 2018). morphology to those of Crenaticaulis Banks and Davis, 1969,
In both cases, T. garrattii is noticeably larger, with its axes up and in some respects to those of Forania Jensen and Gensel,
to at least 7.5 mm wide and 185 mm long. 2013, in that they are deltoid and elongate-triangular (Banks
P
(
Figure 7. Taungurungia garrattii gen. et sp. nov. holotype, fusiform bodies (buds/sterile sporangia). Some of these are highlighted with enlarged
dotted outlines near them: a, anisotomous branching with terminal partial fusiform body (upper arrow). Emergence or branching (lower arrow);
b, post-dégagement, several fusiform bodies with short subtending axes are visible along the edge of the axis. Distally the axis narrows and
terminates in an elongate rounded body (Arrow on right); c, curved axis displaying negative geotropism. Close-up (rectangular region) is in (d);
d, distal part of curved axis possesses minute fusiform bodies (at arrows). An insert highlights the morphology. Evidence of determinate growth
can be seen as the axis 1s terminated by one of these fusiform bodies; e, fine axis emanating about 4 mm on the same axis subtending the large
central terminal sporangium, with arrow at small terminal body. All images are from the counterpart (NMV P257028.2). Image a was taken by
Rodney Start © Museums Victoria. Images b-e were taken with a unidirectional light source.
Taungurungia gen. nov., from the Lower Devonian of Yea, central Victoria, Australia 51
Figure 8. Taungurungia garrattii gen. et sp. nov. holotype, fusiform to elongate rounded (buds/sterile sporangia) bodies with some highlighted
with enlarged dotted outlines beside them: a, b, on the side of axes in carbonised matter; c, faint axis (Ax) with ?fusiform bodies; d, close-up of
fusiform bodies (arrows) on one of the large parent axes. No clear dehiscence lines are present. A short narrow stalk is subtending the sporangium
on the left. Images a, b and d are from the counterpart (NMV P257028.2).
RA i — ua Ww
E
Figure 9. Taungurungia garrattii gen. et sp. nov. holotype, part specimen, NMV P257028.1 close-up of fusiform to elongate rounded (buds/sterile
sporangia) body: a, immediate area above and right of the main terminal sporangium with some poorly preserved casts of fusiform bodies evident
at the arrows. An ovate body (arrow c) pre-dégagement; b, the same ovate body post-dégagement. Its shape may represent partial compression
because it 1s slightly dipping down into the matrix from its subtending axis. At arrow evidence of possible dehiscence line.
02
and Davis, 1969; Jensen and Gensel, 2013). However,
Crenaticaulis emergences are biseriate, continuous and in 1—2
rows per side, while the emergences of Forania are biseriate
and discontinuous (Jensen and Gensel, 2013: table 1). With 7.
garrattii, the emergences appear scattered and change
significantly with maturity.
Discussion
Hueber (1992: 479) noted zosterophylls as having sporangia
that were either reinform or globose, and that this morphology
remained the same throughout their history while exhibiting
variations in axial morphology. Thenceforth, several genera
such as Guangnania, Baoyinia and Yunia from the South
China Plate have been described as having noticeably elongate
sporangia, but with characters indicative of affinities with the
zosterophylls. This suggests that zosterophylls exhibited far
more variation in sporangial morphology, or that these taxa
may represent a separate group(s), as suggested by Hao and Xue
(2013: 119) for Guangnania and Yunia. This paper describes
the first known occurrence of elongate sporangia in association
with zosterophyll characters in Australia. However, it cannot be
assumed they have a common ancestor because the elongate
sporangia may be an analogous structure brought about by
homoplasy. Anatomical evidence in the form of xylem anatomy
would be required to better determine their suprageneric
positioning. Unfortunately, xylem structures have only been
found in one taxon, namely Yunia dichotoma, which differs
from zosterophylls (and lycophytes) in showing centrarch
maturation rather than exarch, and similar in having G-type
Figure 10. Taungurungia garrattii gen. et sp. nov. holotype, counterpart,
P257028.2. Micrograph of an axis (part specimen) with longitudinal
structures, possibly indicative of cell orientation. Taken using an FEI
Quanta 200 scanning electron microscope in low vacuum mode.
F.R. McSweeney, J. Shimeta & J.S. Buckeridge
tracheidal structure (Hao and Beck, 1991), which is similar to
Huia gracilis Wang and Hao, 2001. However, Huia gracilis 1s
considered by Wang and Hao (2001) as a questionable
rhyniopsid based on stalk length and its ovate sporangia,
despite the present of a loose spike and K- or H-branching,
while Kenrick and Crane (1997: 172) consider H. gracilis a
basal lycopsid. Differing rates of evolutionary change within
Zosterophyllopsida and Rhyniopsida may have also played a
role in certain characters evolving to a point outside their
respective classes while maintaining other characters for
extended periods, resulting 1n modular evolution.
Conclusions
In this paper we describe the first taxon from Victoria with
emergences and an unusually large terminal elongate
sporangium. We did not assign T. garrattii to Zosterophyllopsida
because anatomical information for sporangial dehiscence,
xylem type and spike is required to better determine its
suprageneric positioning.
Acknowledgements
We thank Museums Victoria, including Isobel Morphy-Walsh
and the Vertebrate Palaeontology manager Tim Ziegler, for
facilitating access to their collection and for organising the use
of the M205 C Leica microscope to photograph specimens.
Additionally, Rod Start of Museums Victoria for additional
photography. The Taungurung Land and Waters Council,
Taungurung elders and Matthew Burns for their consideration
of the genus name and support. Dr Michael Garratt for pointing
out the fossil locality. RMIT University Melbourne, RMIT
Microscopy and Microanalysis Facility and both Dr Muthu
Pannirselvam and Dr Sindra Summoogum Utchanah (technical
officer and coordinator, respectively). The peer reviewers’
insightful feedback. This work was supported by an Australian
Government Research Training Program Scholarship.
Disclosure statement
No potential conflict of interest was reported by the authors.
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Memoirs of Museum Victoria 81: 55-122 (2022) Published 2022
1447-2554 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2022.81.04
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking
morphological characters with molecular lineages
PuiLLiP J. SUTER!”, JULIA H. Mynott! AND MEGAN CRUMP!
! Centre for Freshwater Ecosystems, Department of Ecology, Environment and Evolution, La Trobe University, Albury—
Wodonga Campus, Victoria, Australia
* To whom correspondence should be addressed. Email: p.suter@latrobe.edu.au
Suter, P.J., Mynott, J.H. and Crump, M. 2022. New species of Paratya (Decapoda: Atyidae) from Australian inland
waters — linking morphological characters with molecular lineages. Memoirs of Museum Victoria 81: 55-122.
The taxonomic history of the atyid shrimp Paratya in Australia has been one of confusion due to the high
morphological variability in material collected from its wide range of distribution. Early research concluded that all
material should be considered a single species, P. australiensis Kemp, pending an acceptable revision. After morphological
examination of material throughout the known distribution, others concluded that only a single species occurred in
Australia. Molecular studies have recognised at least 10 distinct lineages. In the current study, fresh material was collected,
and molecular sequencing was undertaken from a single leg from each specimen. Having confirmed the 10 lineages, the
specimens were dissected for morphological examination. These lineages are recognised as distinct species and
morphological descriptions are provided for seven new species: Paratya walkeri n. sp., P. spinosa n. sp., P. williamsi n. sp.,
P. whitemae n. sp., P. strathbogiensis n.sp., P. gariwerdensis n. sp. and P. rouxi n. sp. A new combination, P. arrostra Riek,
is raised from sub-species to species, P. tasmaniensis Riek is reinstated and P. australiensis Kemp is redescribed. A key
Abstract
based on morphology 1s included.
Keywords Taxonomy, glass shrimp, CO/, morphological variation
Introduction
The freshwater Atyid shrimps in the genus Paratya Miers,
1882 (Miers, 1882), are widespread throughout streams 1n the
eastern Pacific with P. borealis Marin, 2018, in the Russian far
east (Marin, 2018); P. marteni Roux, 1925, in the Lesser Sunda
Islands, Indonesia (Chace, 1997); P. boninensis Satake and Cai,
2005, P. improvisa Kemp, 1917, and P. compressa in Japan
(Marin, 2018; Page et al., 2005a; Satake and Cai, 2005); P.
norfolkensis Kemp, 1917, ın Norfolk Island (Page et al., 2005a);
P. caledonica, P. cf intermedia Roux, 1926, and P. cf typa
Roux, 1926, in New Caledonia (Page et al. 2005a) P.
curvirostris (Heller, 1862) in New Zealand (Page et al., 2005a);
and P. australiensis Kemp, 1917, in Australia (Kemp, 1917;
Williams, 1981; Williams and Smith, 1979). Using
mitochondrial sequences of COI and 16S ribosomal DNA, Page
et al. (2005a) examined the dispersal of Paratya throughout the
South Pacific and hypothesised that these shrimp dispersed via
oceanic currents and using the amphidromous life cycle of
some of the species.
The Australian shrimps of the genus Paratya are
widespread throughout eastern Australia, from Queensland to
South Australia and Tasmania (Cook et al., 2006; Williams,
1977, Williams and Smith, 1979; fig. 1). The first species of
Paratya from Australia, Paratya australiensis, was described
by Kemp (1917).
Since first being described, the taxonomic history of
Paratya in Australia has been one of confusion due to the
high morphological variability in material collected from a
wide range of locations. Two authors (Calman, 1926; Roux,
1926) recognised that animals in their collections were not
typical in appearance to P. australiensis, and Calman (1926)
suggested they may represent a distinct local race based on
the examination of rostral characters.
Roux (1926) compared specimens from populations from
the Horton River near Bingara, North Yanco near Narrandera,
Jamberoo on the southern coast of New South Wales, the Nepean
River, Parramatta and Marrickville in Sydney, and Middle
Harbour in Port Jackson. Importantly, he noted two forms from
North Yanco: one with a long rostrum with two postorbital
spines and 5-8 ventral rostral spines, and a second with a short
rostrum with no postorbital spines and 1—3 ventral rostral spines.
Roux (1926) noted that he could not separate samples into
regional races, even though the North Yanco sample suggests
two taxa because samples from the Sydney area also had
specimens with a short rostrum but with postorbital spines. Riek
(1953) recognised five taxa and described two new species and
two subspecies Paratya australiensis, P. australiensis arrostra,
06
P. atacta, P. atacta adynata and P. tasmaniensis. Walker (1973),
in an unpublished honours thesis, suggested all Riek's taxa were
conspecific. Williams (1977) considered Riek's revision to be
“inadequate and cannot be accepted as a serious taxonomic
statement" (p. 403) and "pending an acceptable revision, all
Australian forms of Paratya are regarded as belonging to a
single species, P. australiensis” (p. 403). In a subsequent paper
by Williams and Smith (1979), all Riek's taxa were formally
synonymised with Paratya australiensis Kemp.
Williams and Smith (1979) re-described P. australiensis and
designated a neotype from Riek's material from close to the
type location of "Clyde near Sydney", because Kemp's (1917)
original type material no longer exists (Williams and Smith,
1979, p. 817). The variability of morphology was documented by
these authors (Smith and Williams, 1980; Williams and Smith,
1979) from examination. of specimens from populations
throughout the known geographical distribution and from a
single population in Cardinia Creek near Melbourne, Victoria.
They concluded that only a single highly variable species
(Paratya australiensis) occurred in Australia.
A series of papers on the genetic characteristics of Paratya
(Baker et al., 2004; Cook et al., 2007; Hancock et al., 1998;
Page et al., 2005b) culminated in the paper by Cook et al.
(2006), which demonstrated nine distinct lineages over the
geographical range of Paratya. The nine lineages consisted of
widespread lineages (4, 6 and 8), lineages from only a single
river (3, 5 and 7) and lineages from geographically adjacent
rivers (1, 2 and 9). A further lineage was discovered in south-
west Victoria by McClusky (2007). Cook et al. (2006)
suggested that there may be defining morphological differences
present among the lineages.
This studys aims were to link morphological and
molecular data to find distinguishing morphological
characters that enable the identification of the separate
lineages of “Paratya australiensis” and to revise the
taxonomy of Paratya in Australia.
Specifically, the aims of this study were to:
e use molecular data to assign individual specimens to
lineages
e discover morphological diagnostic characters for females
and males separately to allow for the morphological
identification of the lineages and eliminate any sexual
dimorphism of character expression
e test the hypotheses that where molecular differences are
present, there are corresponding morphological
differences present.
Methods
Study site locations: Study site locations were selected based on
information provided by Cook et al. (2006) relating to the
location of individual lineages and based on specimens kindly
provided by J. Devine from Sydney Water, B. Cook
(Queensland), C. Madden (South Australia), J. Conallın (New
South Wales), B. Mos and personal collections. Collectors are
identified by their initials: B. Cook (BC), Sydney Water (SW),
Environment Protection Authority, Victoria (Vic EPA), C.
Madden (CM), J. Conallin (JC), J. Mynott (JM), M. Crump
P.J. Suter, J.H. Mynott & M. Crump
(MC), J. Hawking (JH), B. Mos (BM), T. Walker (TW), T.
Curmi (TC), S. Oeding (SO), L. Shuveral (LS), B. Kroll (BKr), J.
Webb (JW), D. Black (DB), A. Clements (AC), P. S. Lake (PSL),
B. Knott (BK) and P. Suter (PS).
Morphology: Terminology of anatomical structures is after
Raabe and Raabe (2008). Specimens were examined under a
stereo dissecting microscope. The appendages from one side of
the body only were dissected with the pereiopods, pleopods,
telson, antennae, stylocerite, scaphocerite and mouthparts all
mounted on slides using Euparal for further examination.
Measurements were obtained using an eyepiece graticule
calibrated with a 5 mm micro-ruler to determine scale.
standard measurements, as illustrated by von Rintelen
and Cai (2009), were used throughout. Carapace length was
measured from the anterior margin of the carapace to the
posterior of the eye orbit; rostral length was measured from
the eye orbit to the apex of the rostrum, rostral depth was the
maximum depth; length of ventral spine row was from the
posterior base of the posterior spine to the anterior base of the
most anterior spine; stylocerite and scaphocerite lengths were
from the base of each structure to the apex; pereiopod and
mouthpart measurements were the maximum length of each
segment made on the external margins; width measurements
were the maximum width of the segment; total pereiopod
length was by addition of length of each segment; telson
length was from the base to the apex but not including the
terminal spines and the width was maximum width at base.
Measurements were compared directly and after being
corrected for body size using the carapace length to compensate
for variation related to the size (and age) of an individual. Only
mature females were used in the analysis due to the presence of
known sexual dimorphisms and the limited number of mature
males in the collections. Females were determined by the
presence of the broad thelycum located on the 8th thoracic
segment; the short lanceolate endopod of the first pleopod,
which is similar in shape to the exopod; and the absence of the
appendix masculina on pleopod 2. All measurements are
reported in the descriptions as the holotype character measure
with the range of expression (i.e. maximum and minimum) in
parentheses.
Segment ratios for pereiopods were for all segments and
exopod length with the carpus length, with the carpus length in
parentheses. The antennal peduncle measurement ratios were
all compared with the apical segment length (3as). The
maxilliped 2 comparisons were the ratios of the apical, mid
and basal segments and exopod lengths to the basal segment
length with the basal segment length given in parentheses.
Segment ratios for maxilliped 3 were the ratios of the apical,
mid and basal segments and exopod lengths to the mid segment
length with the mid segment length given in parentheses.
All maps were prepared using Cartographica. (www.
macgis.com).
Laboratory methods: Genetic techniques were used to assess
the relationships within the species Paratya australiensis and
to determine taxonomic resolution of the previously identified
Paratya lineages in Cook et al. (2006) and McClusky (2007).
Total genomic DNA was extracted using two methods.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. Of
b)
Figure 1. Maps: a, distribution of Paratya in Australia. Sources include data from Cook et al. (2006); Baker et al. (2004), Cook (2006), Hurwood
et al. (2003), McCluskey (2007) and data from this study; b, distribution of Paratya specimens analysed in this study. Maps created in
Cartographica.
08
The first DNA extraction method was a standardised
proteinase-K/Chelex solution following the protocol of Webb
and Suter (2010). Leg tissue was removed from each specimen
and placed in 100 pL of Chelex solution (5% Chelex
(weight:volume), 0.2% SDS, 10 mM Tris pH 8 and 0.5 mM
EDTA) and 10 uL of 20 mg/mL proteinase-K. Specimens
were incubated overnight in solution at 55 °C, removed and
centrifuged at 1500 rpm for 5 minutes before being placed in
a thermocycler for 5 minutes at 95 °C to deactivate the
proteinase-K. The extracted DNA was then diluted 1:5 with
IX TE 20 uL DNA extraction: 80 uL TE).
The second DNA extraction method was a Qiagen DNeasy
blood and tissue kit following standard protocols (Qiagen
Handbook, 2006 wwwl.giagen.com/literature/). A region of
the cytochrome c oxidase subunit 1 gene (COI) was amplified
using Folmer primers (HCO2198 and LCOI490 (Folmer et al.,
1994). All primers were MI3-tailed to facilitate sequencing.
Polymerase chain reaction conditions for the CO/ fragment
used the following protocol: 60 seconds at 94 °C; 5 cycles of
60 seconds at 94 °C, 90 seconds at 45 °C and 90 seconds at
72 °C; 35 cycles of 60 seconds at 94 °C, 60 seconds at 50 °C
and 60 seconds at 72 °C; and a final cycle of 4 minutes at 72
°C. Polymerase chain reaction preparations of 40 uL were
made either with: (1) 4 uL buffer reagent, 2 uL 50 mM MgCl,
0.8 uL of each primer, 0.1 uL Platenium taq polymerase
(Invitrogen, Melbourne), | uL of DNA template and 13.3 uL
of ddH,O, or with (11) 20 uL Taq mastermix (Qiagen), 1 uL
DNA template, 0.8 uL of each primer and 17.4 uL of RNA-
free water (Qiagen). Polymerase chain reaction products were
sent to Macrogen Inc. (Seoul, Republic of Korea) for
purification and sequencing.
Sequence data from previous studies were downloaded
from GenBank (Baker et al., 2004; Cook et al., 2006;
Hurwood et al., 2003; McClusky 2007) to form a backbone
for lineages (Supplementary Table 1). Outgroup sequences
were downloaded from GenBank for Caridina and other
Paratya species. Data generated in this study were assembled
in DnaBaser version 2.91.5 (Heracle BioSoft SRL, Romania,
www.DnaBaser.com) with mismatches, if present, assessed
visually. Alignments were generated using MUSCLE (Edgar,
2004) in MEGA version 10.1.8 (Kumar et al., 2018) and
translated to protein sequences to check for stop codons. All
sequences were trimmed to 434 base pairs to match the
sequence length of GenBank data. All new sequences from
this study have been deposited on GenBank with the
identifiers OL420759-OL420929.
Phylogenetic analysis: Genetic analyses were performed in
MEGA version 10.1.8 with base composition for the 383
sequences (in-group only) showing an AT bias overall and
particularly at the third codon position (overall: A = 25.5%, T =
34.2%, C = 21% and G = 19.3% (AT = 59.7%); third codon
position AT = 77.8%). A test of homogeneity showed that
overall, the sequences were predominantly homogenous with
the first codon position showing an equal amount of
homogeneity and heterogeneity between the sequences. The
estimated transition:transversion ratio was R = 3.12. A
maximum likelihood test of model selection returned T92+G+I
P.J. Suter, J.H. Mynott & M. Crump
as the best model, with T92+G as the second best. Neighbour-
Joining analysis was performed in MEGA version 10.1.8 on
397 sequences using the Tamura 3-parameter model with
gamma = 0.81, rates among sites homogeneous and 2000
bootstrap pseudo-replicates.
Distances were also assessed in MEGA Version 10.1.8
using the un-corrected p-distance with default assumptions.
Results
Phylogenetic analysis: Genetic data from previous studies
were used to form a backbone for previously identified
lineages (Baker et al., 2004; Cook et al., 2006; Cook et al.,
2002; McClusky, 2007). The ingroup for the analyses
included 383 sequences with the backbone comprising 198
sequences (Supplementary Table 1). Ten lineages had been
observed in previous studies, 185 sequences generated in this
study were analysed against the lO-lineage backbone to
assess morphological variation among lineages. Molecular
data (fig. 2) showed monophyletic support for all 10 lineages,
with sequence data from this study associated with each
lineage. Inter- and intra-specific variations were consistent
with the values recorded by Cook et al. (2006). Species
boundaries were determined by molecular divergences
following Costa et al. (2007) and Hebert et al. (2003) and the
examination of morphological differences between lineages.
The combined approach supports the designation of the
lineages as species.
Paratya australiensis and P. gariwerdensis show lower
support values (fig. 2) due to the presence of sub-clades that
are predominantly composed of a small set of sequences from
the backbone set. Distance data further reflects the variation,
with P. gariwerdensis showing a maximum intraspecific
distance of 4.8% (Table 1) and inter-specific variation ranging
from 4.6-9./%. Sequences generated in this study for P.
gariwerdensis aligned in well-supported clades (bootstrap
values 84-96%). Paratya australiensis shows less divergence
distance between sequences, with a maximum intra-specific
distance of 2.8%, which is marginally higher than the
maximum intra-specific distance for Decapoda (2.57%; Costa
et al., 2007), and as stated by Hebert et al. (2003), intra-
specific variation is rarely greater than 2%.
Paratya arrostra, P. whitemae and P. tasmaniensis also
showed high intra-specific variation with maxima of 4.1—4.8%
(Table 1). These three species have widespread distributions.
Paratya arrostra comprised five well-supported sub-clades
(fig. 3, supplementary fig. 1). Sub-clade E contained the bulk
of the sequence data and most of the sequences generated in
this study. Sub-clade E showed very little sequence
divergence, despite associated specimens occurring over a
large geographic area from South Australia, Victoria, New
South Wales and Queensland. Sub-clade A included three
sequences from this study that were located from the Clarence
River catchment near Coffs Harbour, New South Wales, to the
Richmond River catchment near Ballina, New South Wales.
Subclade B included two sequences from this study from
northern Queensland at Tinaroo. Sub-clades C (n = 2) and D
(n = 4) comprised haplotypes from Cook et al. (2006).
New species of Paratya (Decapoda: Atyidae) trom Australian inland waters - linking morphological characters with molecular lineages. 59
Paratya tasmaniensis
Paratya gariwerdensis, n. Sp.
Paratya strathbogiensis, n. sp.
B Paratya williamsi, n. sp.
Paratya spinosa, n. sp.
J| Paratya walkeri, n. sp.
Paratya australiensis
Paratya rouxi, n. sp.
Paratya arrostra
99
Paratya whitemae, n. sp.
99 Paratya howensis
sa | Paratya norfalkensis
99
96 Paratya cf caledonica
99 m Paratya outgroup
Paratya cf intermedia
Paratya curvirostris
76
Paratya compressa
93 T
Caridina
re
0.05
Figure 2. Neighbour-joining analysis of Paratya using Tamura-3-parameter, gamma distribution shape parameter of 0.81, homogenous pattern
among lineages and 2000 bootstrap pseudoreplicates. Bootstrap values >72% displayed. Sequence data from Cook et al. (2006); Baker et al.
(2004), Cook (2006), Hurwood et al. (2003), McCluskey (2007) and this study.
60 P.J. Suter, J.H. Mynott & M. Crump
Paratya arrostra
0.005
Figure 3. Sub-tree of Paratya arrostra Riek, 1953, showing supported subclades. Themajority of material collected across a wide geographical
area in this study grouped within a single clade (E), which 1s equivalent to Lineage 4B in Cook et al. (2006). Previous haplotypes from Cook et
al. (2006) form the other sub-clades, but no specific geographical information is known for these sequences. Sub-clades A and B contain some
specimens from this study.
Paratya tasmaniensis
85
92
C'aridina
0.005
Figure 4. Sub-tree of Paratya tasmaniensis Riek, 1953, showing supported sub-clades. Sub-clade A predominantly has material from Tasmania
but with a single specimen collected at Hamilton, Victoria. Sub-clade B contains specimens from the Glenelg River catchment, south-west
Victoria through to the Hastings River in New South Wales. Sub-clade E contains sequences from Cook (2006) from the Strathbogie area, and
sub-clades C and D are from Cook et al. (2006) and McCluskey (2007).
61
New species of Paratya (Decapoda: Atyidae) trom Australian inland waters - linking morphological characters with molecular lineages.
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Table 1. Divergence distances for the Paratya species recognised in this study and linked to lineages from previous studies. Inter— and intra—
specific pairwise p-distances for the CO] fragment from MEGA version 10.1.8
62
Paratya tasmaniensis comprised five well-supported sub-
clades (fig. 4, supplementary fig. 2). Sub-clade A and B had
sequences from this study associated with the backbone material.
Sub-clade C (n = 2) and D (n = 4) comprised haplotypes from
McClusky (2007) and Cook et al. (2006). Sub-clade E comprised
sequences from Cook (2006) with specimens collected from the
Granite Creeks area of the Strathbogie Ranges, Victoria.
Paratya whitemae comprised one well-supported and
three poorly supported sub-clades (Supplementary fig. 3). The
well-supported sub-clade contained only sequences from the
backbone data (Cook et al., 2006; Hurwood et al., 2003;
McClusky, 2007). The bulk of sequences for this species
showed very little divergence and comprised one sub-clade,
the other sub-clades showed more divergence within the clade.
Paratya walkeri, P. spinosa and P. australiensis showed
low minimum inter-specific variation between each other but
were well supported in the neighbour-joining analysis and
morphologically. Paratya spinosa had 3% divergence from
both P. australiensis and P. walkeri. However, P. australiensis
and P. walkeri had a minimum inter-specific divergence of
2.5% (Table 1). This is consistent with the inter-specific
variation within the Atyidae where Chen et al. (2020) observed
inter-specific variation as low as 3.3% (3.3-33%) in Caridina
from China, while Christodoulou et al. (2012) reported 5.9%
(5.9-28./%) variation in Atyaephyra and Shih et al. (2019)
reported 2.17% (2.17-53%) variation in Neocaridina.
Systematics
Genus Paratya Miers, 1882
Paratya australiensis Kemp 1917
Diagnosis: Rostrum longer or shorter than carapace, usually
slender and pointed; dorsally armed with 11-34 teeth of which
0—4 are postorbital spines; ventrally with 1-14 large serrations;
dorsal edge straight or very slightly concave.
Eyes well developed, darkly pigmented.
Carapace with supraorbital spine large and distinct,
antennal spine smaller; pterygostomian spine indistinct, but
pterygostomium angle quite acute; hepatic spine absent.
Antenna | length about half body length. Peduncle with
numerous finely setose spines in row near lateral, ventral and
distal margins and along medial edge; lateral distal angle of
first segment with prominent acute process or stylocerite that
reaches to distal border of peduncle segment.
Antenna 2 longer than body. Peduncle first segment without
setae, overlapping second segment dorsally, with prominent
tooth at outer distal angle; second segment with short row of
setae dorsally; third segment with group of setae at inner distal
angle. Scaphocerite with regular row of setose spines on inner
and distal margins; outer margin extending to a sharp point
overreached by lamella. Flagellum long and slender.
Maxillipeds | exopod flagellum distinct, well developed
and with numerous long setose spines on all margins,
approximately half the length of the caridean lobe; caridean
lobe broad with numerous short setose spines on outer margins
and a few on body of lobe; epipodite small. Maxilliped 2
exopod long and narrow, several setose spines of various
P.J. Suter, J.H. Mynott & M. Crump
lengths near tip and basally. Epipodite with podobranch.
Maxilliped 3 basal segment curved, apical segment with
large terminal claw, medial distal margin with broad teeth-
like spines, largest 1n basal third, outer margin with broad
tooth-like spines longest in basal third; several transverse
spine rows near base; mid and basal segments with several
short simple spines. Exopod long and narrow, with several
long setose spines near tip and several short setose spines near
base. Epipodite with basal conical projection.
Pereiopods 1—5 all possessing an exopodite, only pereiopods
1-4 with epipodite. Pereiopods 1 and 2 with propodus and
dactylus forming chelae each with a terminal tuft of setae.
Dactylus of pereiopod 3 and 4 with a prominent terminal claw
and strong spines on medial margin; dactylus of pereiopod 5
with prominent terminal claw and very regular, comb-like row
of numerous small spines on medial margin.
Pleopod | male with endopod about half length of exopod,
narrowly ovate at base, usually excavated distally with
numerous long setose spines laterally and medial spines.
Pleopod 2 appendix masculina present in males, absent in
females. Appendix interna long and narrow about one-fifth of
length of endopod and exopod, distal margin with long setae.
Peduncle with short and long spines.
lelson long and tapers towards posterior, dorsal surface
with 2—3 pairs of strong sub-marginal teeth-like spines,
posterior margin with | pair of spine-like teeth and 6-14 long
strong terminal spines.
Paratya australiensis Male neotype
This species was fully described by Williams and Smith (1979)
and the description given below adds a morphotype from the
Shoalhaven R and the morphometric characters for direct
comparisons of all species recognised in this revision.
Paratya australiensis Kemp
Figure 5
Paratya australiensis Kemp (1917); primary type material no
longer exists
P. australiensis Riek (1953)
P.australiensis Williams and Smith (1979); neotype male selected
from material named by Riek (1953) AM P28693. Neotype examined
by MC
Lineage A (Baker et al., 2004)
Lineage 1 (Cook et al., 2006)
Numerous authors referred to P. australiensis 1n both taxonomic
(Calman, 1926; Kemp, 1917; Roux, 1926; Smith and Williams, 1980;
Williams, 1977; Williams and Smith, 1979) and ecological studies
(Baker et al., 2004; Balcombe et al., 2007; Boulton, 2003; Bunn and
Hughes, 1997; Chessman and Robinson, 1987; Hancock, 1998;
Hancock and Bunn, 1997; Hancock and Bunn, 1999. Hancock et al.,
1998; Hart et al., 1991; Hladyz et al., 2012; Hughes et al., 2003; Hughes
et al., 1995; Hurwood et al., 2003; Kefford et al., 2004; Marchant et al.,
1999; Marchant et al., 1984; Metzeling, 1993; Piola et al., 2008; Reidet
al., 2008; Richardson and Humphries, 2010; Smith and Williams,
1980; Walsh and Mitchell, 1995; Williams, 1977), but these ecological
studies recognised only a single species from the taxonomic decision
by Williams and Smith (1979). Based on current knowledge, these
identifications should be revisited.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 63
c
-
Figure 5. Paratya australiensis: a—e, P. australiensis Kemp; f-j, P. australiensis Shoalhaven morphotype; a, rostrum; b, left mandible incisors; c,
right mandible incisors; d, first pereiopod; e, second pereiopod; f, rostrum; g, left mandible incisors; h, right mandible incisors; 1, first pereiopod;
j, second pereiopod. Scale lines 0.2 mm.
64
Material Examined: New South Wales: Neotype male AM P28693
Seven Hills, Sydney: Hawkesbury R at Powells Lane, —33.569 S,
150.745 E, 9 March 2011 (SW); Hawkesbury R at North Richmond,
—33.589 S, 150.715 E, 9 March 2011 (SW); Hawksbury R at
Wilberforce, —336018 S, 150.8245 E, 21 April 2011 (SW) Nepean R at
Penrith Weir, —33.746 S, 150.682 E, 14 April 2011 (SW); Warragamba
R upstream of Nepean R confluence, —33.8589 S, 150.611 E, 18 April
2011 (SW); Bedford Ck, Blue Mountains, —33.75 S, 150.4474 E, May
2011 (SW) MC32; Bungonia Ck at Bungonia, —34.8528 S, 149.9437 E,
no date (SW), Shoalhaven R at Hillview, —35.1826 S, 149.9541 E, no
date (SW); Boro Ck at Marlowe upper Shoalhaven R, —35.3426 S,
14977386 E, no date (SW); Hacking R at McKell Ave, 34.1524 S,
151.0284 E,5 May 2011 (SW); South Ck at Richmond Rd, —33.6775 S,
150.8121 E, 30 May 2011 (SW).
Williams and Smith (1979) fully illustrated P. australiensis, and
these have not been duplicated here. Some comparative characters of
the two morphotypes are provided (fig. 5).
Diagnosis: P. australiensis differs from all other species by the
following combination of characters: rostrum long, extending
beyond antennular peduncle and scaphocerite in Sydney
streams, dorsal edge very slightly concave, dorsally armed with
16-28 teeth, 1—3 postorbital spines, ventrally with 4—7 large
serrations over a length of 0.60—1.5 mm, distal half of ventral
edge more or less straight; left mandible with 4—5 teeth
separated by finely ridged notch from a less distinct apical
tooth; right mandible with 4 teeth in 1n 2 separate incisor
processes; scaphognathite of maxilla 2 rounded apically almost
extending to apex of upper endite; maxilliped 1 with exopod
flagellum distinct, well developed and with numerous long
setose spines on all margins, approximately four-fifths length
of caridean lobe; exopod of maxilliped 2 1.70-2.77 times
longer than endopod, epipodite with podobranchs extending
Just to base of third segment of endopodite; maxilliped 3 with
medial distal margin of apical segment of endopod with 9—11
broad teeth-like spines, outer margin with 2—4 broad teeth-like
spines plus 2—4 smaller spines, exopod long and narrow, tip
over-reaching distal end of basal endopod segment; pereiopod
] with long carpus and long slender chelae, exopod extending
to mid carpus-base of propodus; pereiopod 2 with exopod
extending to apex of merus; dactylus of pereiopod 3 with
prominent terminal claw and 7-11 strong spines on medial
margin, exopod extends to mid merus; dactylus of pereiopod 4
prominent terminal claw and 6-11 spines on medial margin,
exopod extends to mid-apical third of merus; dactylus of
pereiopod 5 with prominent terminal claw and very regular
comb-like row of 55—80 small spines on medial margin, exopod
extends to mid merus.
Shoalhaven R morphotype: Rostrum extends beyond peduncle,
but not beyond scaphocerite, dorsally armed with 16—17 spines,
O postorbital spines, ventrally with 2-5 spines over a length of
0.60-1.5 mm.
Carapace: length 4.1-8.0 mm.
Rostrum (Williams and Smith, 1979; fig. 1a): long, 4.0—5.6
mm, extending beyond both antennular peduncle and
scaphocerite in specimens from the Sydney area (fig. 5a) but
slightly shorter and not extending beyond the scaphocerite in
Specimens from the upper Shoalhaven system (fig. 5f), rostral
length 0.62-1.24 times length of carapace, dorsal edge very
P.J. Suter, J.H. Mynott & M. Crump
slightly concave, with slight upwards curve, moderately
slender to broad and pointed, rostrum 6.75—9.33 times longer
than wide; dorsally armed with 16-25 teeth, ratio of rostral
spines to rostral length is 3.15—4.90 with 1-3 post orbital
spines in the Sydney morphotype (fig. 5a) or O post orbital
spines in the Shoalhaven morphotype (fig. 5f); ventrally with
2-6 large serrations over a length of 0.6-1.5 mm all anterior
to widest point; distal half of ventral edge more or less
straight, ratio of ventral rostral spines length to rostral length
= 0.13-0.28 and 3.20-8.00 times more dorsal spines than
ventral spines; rostral length 0.92-1.5 times length of
scaphocerite.
Antenna | (Williams and Smith, 1979; fig. 1, b, c) peduncle
3.40-4.54 mm long, not quite reaching distal tip of
scaphocerite, length 0.90—1.07 times as long as scaphocerite.
Stylocerite 1.40—2.80 mm long, length 6.89-13.33 longer than
width, 0.33-0.43 times carapace length, reaching beyond
distal border of peduncle segment, almost to end of acute
process on the lateral distal angle of the first segment (fig. 5f).
Antenna 2 (Williams and Smith, 1979; fig. Id) second
segment of peduncle 1.00-1.94 mm long, 0.29—0.44 times
length of scaphocerite, 2.53-3.80 times longer than wide.
Scaphocerite 3.40—4.72 mm long, 0.60—0.85 times carapace
length, 2.76—4.25 times as long as wide.
Mouthparts (Williams and Smith, 1979; figs le-1, 2a-c).
Left mandible (fig. 5b, g) with 4—5 teeth separated by finely
ridged notch from a less distinct apical tooth; spine row
immediately below incisor process of 6-10 rugose spines
(lifting spines); spine row above molar process of
approximately over 20 sparsely setose spines. Right mandible
(fig. Sc, h) with 4 teeth separated into 2 incisor groups of 2
teeth, apical and third teeth largest, teeth 2 and 4 shorter;
spine row immediately below teeth with 8—10 spines, each
finely setose basally; spine row above molar process. Molar
process ridged.
Maxilla palps, with | long, setose terminal spine and 1—2
simple sub-terminal ones, inner distal angle may be slightly
acute.
Maxilla 2 scaphognathite rounded apically almost
extending to apex of upper endite. Palps small, terminal parts
narrow with 1-2 setose spines.
Maxilliped 1 palp with broad base, short narrow distal
lobe, several long setose spines on distal margins. Exopod
flagellum distinct, well developed and with numerous long
setose spines on all margins, approximately half to four-fifths
the length of the caridean lobe.
Maxilliped 2 endopod length 0.68—1.32 mm; exopod long
and narrow, length 1.73-2.80 mm, exopod 1.70-2.77 times
longer than endopod. Epipodite with podobranchs extending
just to base of third segment of endopodite.
Maxilliped 3 endopod length 3.07-7.54 mm, 1.70—2.77
times longer than exopod; with 3 distal segments of similar
length; basal segment curved; apical segment with large
terminal claw, medial distal margin with 9—11 broad teeth-
like spines, largest in basal third, outer margin with 2—4 broad
teeth-like spines and 2-4 smaller spines, longest in basal
third. Exopod long and narrow, length 1.80-3.62 mm, tip
over-reaching distal end of basal endopod segment.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 69
Thoracic appendages (Williams and Smith, 1979; figs 2d,
e, 3a—c). Pereiopod 1 (fig. 5d, 1). 3.45—5.49 mm long, 0.68—0.91
times length of carapace. Chelae short to long and slender,
1.09-1.93 mm long, propodus 2.55—3.46 times as long as wide,
1.88—2.21 times longer than dactylus; palm length 1.58-1.91
times width and 1.00-1.38 times dactylus length (fig. 5d, 1).
Carpus long, 1.47-3.08 times longer than greatest width.
segment ratios compared with carpus length 0.54-0.78 :
1.04—1.71 : 1.00 (0.92-1.36) mm : 0.99-1.45 : 0.35-0.84 :
2.21-3.27. Exopod extending to mid carpus—base of propodus.
Pereiopod 2 (fig. 5e, J). length 5.04-9.58 mm, 1.00-1.47
times carapace length. Chelae long and slender 1.11-1.76 mm
long, approximately two-thirds carpus length, 3.59—4.14 times
as long as wide, palm length 1.71-1.98 times width and 0.82—
1.31 times dactylus length (fig. 5e, J). Propodus length 1.61—
2.07 times longer than dactylus. Carpus 5.76-8.16 times as
long as greatest width, slightly broader distally, distal margin
with small excavation. Segment ratios 0.30-0.38 : 0.56-0.72 :
1.00 (1.73-2.84) mm : 0.58-0.96 : 0.34-1.00 : 0.83-1.49.
Exopod extending to apex of merus.
Pereiopod 3 slightly longer than pereiopod 2 and more
slender, length 5.7-10.08 mm, 1.19-1.60 times carapace length.
Dactylus with prominent terminal claw and 7-11 strong spines
on medial margin. Propodus length 3.23-4.56 times longer
than dactylus, length 9.81—14.3 times as long as wide with
12-15 spines on inner margin. Merus with 1 strong spine on
medial margin and | near ventral distal margin. Segment
ratios 0.38—0.54 : 1.62-1.79 : 1.00 (1.07-1.88) mm: 1.60-2.11 :
0.50—0.77 : 1.60—1.74. Exopod extends to mid merus.
Pereiopod 4 similar to pereiopod 3, 6.09—9.60 mm long,
1.20-1.52 times carapace length. Dactylus with prominent
terminal claw and 6-11 spines on medial margin. Propodus
length 3.92-4.94 times longer than dactylus, length 10.74—
16.50 longer than wide, with 11—17 spines on medial margin;
merus with | or 2 strong spines on medial margin and | near
ventral distal margin. Segment ratios 0.37—0.44 : 1.67—1.90) :
1.00 (1.16-1.80) mm : 1.64-2.01 : 0.54-0.82 : 1.18-1.87.
Exopod extends to mid-apical third of merus.
Pereiopod 5 similar length to pereiopods 3 and 4, 5.85—9.79
mm long, 1.21-1.57 times carapace length. Dactylus with
prominent terminal claw and very regular comb-like row of
55—80 small spines on medial margin. Propodus length 2.83—
3.60 times longer than dactylus, length 9.57-16.54 times longer
than wide with 8—16 medial teeth. Carpus approximately half
propodus length with 1 large spine near distal margin. Merus
with | strong medial spine and 0—1 distal spines; ischium less
than half length of propodus; segment ratios 0.52—0.74 : 1.83—
2.09 : 1.00 (1.15—1.85) mm : 1.33-1.90 : 0.58-0.99 : 1.22-1.69.
Exopod extends to mid merus.
Abdomen (Williams and Smith, 1979; figs 3d-f, 4a-c).
Pleopods peduncle of first pleopod short, 0.23-0.33 times
length of carapace, 2.62-3.33 times width, exopod 1.05-1.6
times peduncle length, endopod 0.42—0.83 times peduncle
length; second pleopod peduncle short, 0.29—0.44 times length
of carapace, 2.31—3.5 times width, exopod 1.00-1.58 times
peduncle length, endopod slightly shorter 1.00—1.55 times
peduncle length. Length of first peduncle 0.93-1.38 times
length of second peduncle length.
Telson length 3.00—4.48 mm, 0.55-0.73 times carapace
length, 2.35—3.28 times as long as greatest width, and tapering
distally. Dorsal surface with 2 pairs of strong submarginal
teeth-like spines. Posterior margin convex with | pair of teeth-
like spines outermost, 6 long, strong setose spines, and 2 short,
simple spines.
Uropods approximately equal to telson length.
Males (Williams and Smith, 1979; fig. 3d—f). Smaller than
females, carapace length 5.25 mm; endopod of pleopod 1
strongly excavated with 12 short external spines on medial
margin and 17 long setae on inner margin.
Comments: Lineage | specimens are consistent with the neotype
of P. australiensis described and illustrated by Williams and
smith (1979) from Seven Hills near Sydney. Kemp (1917)
described Paratya australiensis from Clyde, Sydney, but, as the
type was missing, Williams and Smith (1979) attempted to find
material from the type locality, but were unsuccessful. They
selected a specimen from Riek's (1953) collection from the
seven Hills site as the neotype. Both locations (Seven Hills and
Clyde) are in the Parramatta R system and are only 10 km apart
(Williams and Smith, 1979). The material examined in this
study was from adjacent catchments. P. australiensis appears
restricted to coastal catchments in the Sydney area and in the
Shoalhaven catchment. P. australiensis does not have an
overlapping distribution with P. walkeri, P. spinosa, P.
strathbogiensis and P. gariwerdensis but it does overlap with P.
arrostra, P. williamsi, P. whitemae and P. tasmaniensis. P.
whitemae has been found at the same locality as P. australiensis,
and both occur in the Hawksbury R and Nepean R.
P. australiensis can be clearly distinguished from the
species possessing a short rostrum (P. arrostra and P. rouxi)
by the rostrum extending beyond the peduncle compared with
the rostrum that does not extend beyond the peduncle, and
characters listed 1n Table 2.
P. australiensis can be distinguished from all the long
rostrum species by the combination of characters listed in
Table 3. Main characters include the number of dorsal rostral
spines; the number of ventral rostral spine; palm length to
width of second cheliped; carpus of pereiopod 1 long; the
length of the lower rostral spine row; number of long terminal
spines of telson; scaphognathite of maxilla 2 long almost
extending to end of endite; right mandible with 2 pairs of
incisors (Table 3). The presence of two morphotypes of P.
australiensis 1s similar to the observations made on another
Australian atyid shrimp, Australatya, by Choy et al. (2019).
Paratya walkeri n. sp.
Figures 6-8
http://zoobank.org/urn:lsid:zoobank.org:act: EO3B305A-OEDD-
4A BD-92E8-566A E860E9EO
Lineage 2 (Cook et al., 2006)
Type material: Holotype New South Wales. Dingo Ck, —30.3103 S,
152.9822 E, 24 May 2015 (BM) PS9. Body in ethanol and antennae,
mouthparts, pereiopods and abdominal structures dissected, mounted
on 2 slides. Accession Ref. PS9. Australian Museum (AM) Ref No.
P.105600.001.
P.J. Suter, J.H. Mynott & M. Crump
66
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Table 2. Characters that distinguish all Paratya species with a long rostrum which extends beyond the antennal peduncle
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 6
~
Table 3. Characters that distinguish all Paratya species with a short rostrum that does not extend beyond the antennal peduncle
Rostrum shorter than carapace +
Number of dorsal rostral spines —
Number of post-orbital spines
Number of ventral rostral spines
elle
2 =
elige
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88
C10
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>
ge
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pu
=
=
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3
Length of ventral rostral spine row «0.4 20.4
(mm)
Extends just to basal third of process | Extends almost to end of process | Extends almost to end of process
].2-1.4 1.1-1.8
12
0.67—0.71 0.44—0.60
Antenna 1 stylocerite and
peduncle process
First pereiopod carpus length
First pereiopod chelae shape
1
First pereiopod Propodus length/ .
Telson length/carapace length
Paired incisors, each with 2 large teeth | Single incisor of 4—5 large teeth |Single incisor with 4—5 large teet
Truncated, shorter than endite Rounded apically, not truncated,
almost reaching apex of endite
Inland Murray—Darling catchment, NSW| Widespread, Vic, NSW, Qld, SA
Right mandible Incisors
illa 2 scaphognathite
Vix o C
za > 2 d
ay mi; = =
oz 3 yn
= G
e de
=
Note: L9, 4B and 4C refer to lineages of Cook et al. (2006).
Paratypes: New South Wales. 5 specimens in ethanol same data as
holotype; Accession Ref. PS23, PS8 Genbank Registration OL420929;
body in ethanol and other structures dissected, mounted on 2 slides
each; AM Ref No. P.105601; 3 specimens ın alcohol.
: Dingo Ck, —30.3103 S,
Diagnosis: P. walkeri differs from all other species by the
following combination of characters: rostrum long, extending
beyond both antennular peduncle and scaphocerite, dorsal edge
very slightly concave, dorsally armed with 24-30 teeth, 2—4
postorbital spines, ventrally with 14-16 large serrations over a
length of 2.75—4.00 mm, extending from just posterior of greatest
mandible with 4 teeth in a single incisor process; scaphognathite
of maxilla 2 truncated distally not extending to apex of upper
endite; maxilliped 1 with exopod flagellum distinct, well
developed and with numerous long setose spines on all margins,
approximately two-thirds length of caridean lobe; exopod of
maxilliped 2 1.09—2.13 times longer than endopod, maxilliped 3
with medial distal margin of apical segment of endopod with
6—12 broad teeth-like spines, outer margin with 2 broad teeth-
like spines, exopod long and narrow, tip over-reaching distal end
of basal endopod segment; pereiopod 1 with long carpus and
long, robust chelae, exopod extending to mid to apex of carpus;
pereiopod 2 with exopod extending to mid-apex of merus;
dactylus of pereiopod 3 with prominent terminal claw and 7-10
strong spines on medial margin, exopod extends to mid merus;
dactylus of pereiopod 4 prominent terminal claw and 7—8 spines
1
+-
2-3
and separate from rostral spines
Lo de 4 S —— ———
Paratya rouxi (L9) Paratya arrostra (AB) Paratya arrostra (AC)
4.90-5.30 55-63
11-19 19-25 16-19
0-1 spine displaced posteriorly
4—5
>0.4
Long, length 7.0-9.3 times width Long, length 6.2—8.8 times width | Short, length 5.2—5.9 times width
Broad/robust
1.42.1
0.44-0.60
7-13 (usually 11-12) 7-13 (usually 11-12)
-p
Rounded apically not truncated,
almost reaching apex of endite
South-east Qld
on medial margin, exopod extends to apical third of merus;
dactylus of pereiopod 5 with prominent terminal claw and very
regular comb-like row of 53—60 small spines on medial margin,
propodus with 8-11 medial spines; exopod extends to basal third
to mid third of merus. Telson with 9-10 long terminal spines.
Carapace length 6.25 mm (5.75-6.25).
Rostrum long, 6.3 mm (5.75—6.60), extending beyond both
antennular peduncle and scaphocerite (fig. 6a), rostral length
1.08 (0.95-1.15) times longer than carapace, dorsal edge
curved upwards to tip, broad and pointed, rostral length 9.00
(6.76-9.00) times greater than width; dorsally armed with 29
(24-30) teeth, ratio of rostral spines to rostral length is 4.60
(4.17—4.60); 4 (2—4) postorbital spines; ventrally with 16 (14—
16) large serrations (fig. 6a) over a length of 4.00 mm (2.75—
4.00), 2 (1-2) spines posterior to widest point; ratio of ventral
spine length to rostral length is 0.63 (0.42—0.63) and 1.81
(1.60-2.14), more dorsal spines than ventral spines; rostral
length 1.42 (1.25-1.42) times length of scaphocerite.
Antenna | peduncle 3.88 (3.40—4.70) mm long, not
reaching distal tip of scaphocerite (fig. 6b), 0.87 (0.81—0.91)
times length of scaphocerite. Stylocerite 2.88 (2.18—2.88) mm
long, length 11.50 (7.25—11.50) longer than wide, 0.46 (0.36—
distal angle of first segment.
Antenna 2 (fig. 6c) apical segment of peduncle 1.55 (1.55—
1.60) mm long, 0.35 (0.34—0.35) length of scaphocerite, 2.70
(2.56-2.70) longer than wide. Scaphocerite 4.45 (4.45—4.70)
mm long, 0.71 (0.71-0.82) times length of carapace, 3.30
(3.24—3.68) longer than wide.
68 P.J. Suter, J.H. Mynott & M. Crump
Pads
Figure 6. Paratya walkeri sp. nov.: a, head region and rostrum; b, antenna 1 peduncle and stylocerite; c, scaphocerite; d, left mandible; e, enlarged
incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; 1, maxilla 2; jJ, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages.
Figure 7. Paratya walkeri sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus. Scale lines 0.2 mm.
69
70 P.J. Suter, J.H. Mynott & M. Crump
Figure 8. Paratya walkeri sp. nov.: a, pereiopod 4; b, dactylus 4; c, pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g, pleopod 1 of
female. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 11
Mouthparts. Left mandible (fig. 6d, e) with 3 teeth
separated by a notch from 2 less distinct apical teeth; spine
row immediately below incisor process of 6 rugose spines
(lifting spines); spine row above molar process of
approximately over 20 sparsely setose spines. Right mandible
(fig. Of, g) with 4 teeth in a single incisor process with tooth 2
largest; spine row immediately below teeth with 7 lifting
spines; spine row above molar process. Molar process ridged.
Maxilla 1 (fig. 6h) as for P. australiensis.
Maxilla 2 scaphognathite truncated distally (fig. 61), not
extending to apex of upper endite.
Maxilliped 1 (fig. 6j) palp with broad base, short narrow
distal lobe, and several long setose spines on distal margins.
Exopod flagellum distinct, well developed, longer than P.
australiensis with numerous long setose spines on all margins,
approximately Two-thirds to equal length of caridean lobe.
Maxilliped 2 (fig. 6k) endopod 1.14 (1.11-1.30) mm long,
exopod long and narrow 2.35 (1.43—2.38) mm long, exopod 2.03
(1.09-2.13) times longer than endopod. Epipodite with
podobranch.
Maxilliped 3 (fig. 7a) endopod length 7.38 (7.05—7.58)
mm, 2.68—2.69 times longer than exopod, with 3 distal
segments of similar length; basal segment curved, apical
segment with large terminal claw, medial distal margin with 7
(6-12) broad teeth-like spines, largest 1 in basal third, outer
margin with | apical tooth-like spine and | paired sub-apical
setae. Epipodite with basal conical projection.
Thoracic appendages. Pereiopod 1 (fig. 7b) 5.20 (5.12-
5.28) mm long, 0.83 (0.853—0.92) times carapace length.
Chelae long and broad or short and broad, 1.75 (1.75-1.85)
mm long, propodus 3.04 (2.64-3.04) times longer than wide,
2.00 (1.85-2.18) times longer than dactylus, 1.30 (1.30—1.44)
times longer than carpus; palm length 1.48 (1.48—1.65) times
width and 1.03 (1.03-1.32) dactylus length. Carpus long 2.45
(2.00-2.45) times as long as greatest width, broadening
distally, distal margin excavate. Segment ratios 0.65 (0.65—
0.78) : 1.30 (1.30-1.44) : 1.00 (1.35 [1.25-1.35] mm) : 1.15
(1.15—1.24) : 0.41 (0.38—0.42) : 2.50 (2.50-2.68). Exopod
extending to mid to apex of carpus.
Pereiopod 2 (fig. 7c) 7.45 (7.15—7.48) mm long, 1.19 (1.18—
1.30) times carapace length. Chelae long and slender 1.70
(1.58-1.68) mm long, nearly two-thirds length of carpus, 2.96
(2.96—3.53) times longer than wide, palm length 1.61 (1.50—
1.61) times width and 1.12 (0.70—1.12) times dactylus length.
Propodus 2.06 (1.56—2.06) times longer than dactylus. Carpus
7.69 (6.79-7.69) times as long as greatest width, slightly
broader distally, distal margin with small excavation. Segment
ratios 0.33 (0.33-0.43) : 0.68 (0.66—0.68) : 1.00 (2.50 [2.38—
2.50] mm) : 0.80 (0.78—0.82) : 0.49 (0.49—0.57) : 1.36 (1.30—
1.41). Exopod extending to apex of merus to base of carpus.
Pereiopod 3 (fig. 7d, e) distinctly longer than pereiopod 2
and more slender 8.65 (8.55—9.70) mm long, 1.38 (1.38—1.69)
times carapace length. Dactylus with prominent terminal claw
and 7 (7—10) strong spines on medial margin (fig. 7e).
Propodus length 5.26 (4.28—5.33) times longer than dactylus,
length 15.13 (13.78-15.13) times longer than wide with 9
(7—10) spines on inner margin. Merus with | strong spines on
medial margin and | near ventral distal margin; segment
ratios 0.33 (0.32—0.40) : 0.68 (0.66—0.68) : 1.00 (1.65 [1.63—
1.80] mm) : 0.80 (0.78—0.82) : 0.49 (0.50—0.57 : 1.36 (1.30—
1.41). Exopod extends to mid merus.
Pereiopod 4 (fig. 8a, b) similar to pereiopod 3, 9.35 (9.18—
9.35) mm long, 1.50 (1.50-1.60) times carapace length.
Dactylus with prominent terminal claw and 7 (7—8) spines on
medial margin (fig. 8b). Propodus length 4.44 (3.97—4.44)
times longer than dactylus, length 13.33 (11.90—14.40) times
longer than wide, with 11 (10-13) spines on medial margin;
merus with only | strong spine on medial margin and | near
ventral distal margin. Segment ratios 0.35 (0.39—0.49) : 1.83
(1.70—2.00) : 1.00 (1.75 [1.63-1.75] mm) : 1.85 (1.85-2.15) :
0.56 (0.51—0.62 ): 1.70 (1.67—1.75). Exopod extends to middle
third of merus.
Pereiopod 5 (fig. 8c, d) similar length to pereiopods 3 and 4,
9.15 (9.08-9.28) mm long, 1.46 (1.46—1.61) times longer than
carapace. Dactylus with prominent terminal claw and very
regular, comb-like row of 53 (53-62) small spines on medial
margin (fig. Sd). Propodus length 4.25 (3.68—4.25) times longer
than dactylus, length 17.00 (15.00-17.00) times longer than
wide with 8 (8—11) long medial teeth and setae on external
margin. Carpus with | large spines near distal margin. Merus
with | strong medial spine and | distal spine; segment ratios
0.47 (0.47—0.60) : 2.00 (2.00-2.19) : 1.00 (1.70 [1.63—1.70] mm)
: 1.76 (1.72-1.82) : 0.62 (0.61—0.69) : 1.61 (1.43-1.61). Exopod
extends to mid merus (basal third to mid third of merus).
Abdomen. Pleopod peduncle of first pleopod short, 0.26
(0.26—0.37) times length of carapace length, 2.36 (2.36—4.50)
times width, exopod 0.73 (0.73—1.40) times peduncle length,
endopod 0.70 times peduncle length (fig. 8g); second pleopod
peduncle short, 0.36 (0.29-0.36) times length of carapace,
2.64 (1.67-2.64) times width, exopod 1.29 (1.29-1.54) times
peduncle length, endopod slightly shorter 1.13 (1.13—1.46)
times peduncle length.
Telson (fig. Se, f) length 4.00 (4.00—4.25) mm, 0.64
(0.64—0.74) times carapace length, 3.64 (2.86-3.64) times as
long as greatest width, tapering distally. Dorsal surface with 2
pairs of strong submarginal teeth-like spines. Posterior
margin convex with | pair of teeth-like spines outermost, 7
(7—8) long, strong setose spines (fig. 8f).
Uropods approximately equal to telson length.
Males unknown.
Etymology: Named after Dr Terry Walker whose study of
Paratya in Tasmania (Walker, 1973) initiated subsequent
taxonomic and morphometric studies (Smith and Williams,
1980; Williams, 1977; Williams and Smith, 1979) and who has
encouraged this morphological study.
Comments: Paratya walkeri may be confused with P.
australiensis, P. arrostra and P. whitemae due to the very long
rostrum that is slightly concave. P. walkeri can be differentiated
from all the long rostrum species by the combination of
characters in Table 2, including the number of dorsal rostral
spines (24-30), number of post orbital spines (2-4), number of
ventral spines (14-16) over a length of 2.75—4.00 mm; dactylus 4
with 7-8 medial spines; 1—2 ventral rostral spines posterior to
maximum rostrum width.
12
Distribution: P. walkeri 1s restricted to the northern coastal
rivers of New South Wales in the Tweed R and Clarence R
catchments. Cook et al. (2006) recorded this species only in
the Tweed R; we recorded it in the Manning R system but not
at the Tweed R site. It is possible that P. walkeri and P. spinosa
may coexist.
Paratya spinosa n. sp.
Figures 9-11
http://zoobank.org/urn:lsid:zoobank.org:act:2EBBAS31-3594-
4725-A47F-530A A42CC609
Lineage 3 (Cook et al., 2006)
Type Material: Holotype New South Wales. Korrumbyn Ck,
Tributary of Tweed R, Mt Warning, —28.4 S, 153.3 E, no date (BC).
Body in ethanol and antennae, mouthparts, pereiopods and abdominal
structures dissected, mounted on 2 slides. Accession Ref. MC11. AM
Ref No. P.105602.001.
Paratypes: New South Wales. Details as for holotype, Accession
Ref. MCI7 Genbank Registration OL420818, MCI3 bodies in
ethanol and other structures dissected, mounted on 2 slides each
and 2 whole specimens.
Material Examined: New South Wales: Korrumbyn Ck, Tributary of
Tweed R, Mt Warning, —28.4 S, 153.3 E, no date (BC).
Diagnosis: P. spinosa differs from all other species by the
following combination of characters: rostrum long, extending
beyond both antennular peduncle and scaphocerite, dorsal edge
straight, dorsally armed with 25-30 teeth, 3 postorbital spines,
ventrally with 9-10 large serrations over a length of 1.65—3.00
mm, all forward of greatest depth; distal half of ventral edge
straight; left mandible with 4 teeth separated by finely ridged
notch from a less distinct apical tooth; right mandible with 4-5
teeth in a single incisor process with all teeth large;
scaphognathite of maxilla 2 rounded apically extending to apex
of upper endite; maxilliped | with exopod flagellum distinct,
well developed and with numerous long setose spines on all
margins, approximately half length of caridean lobe; exopod of
maxilliped 2 1.06-1.20 times longer than endopod; maxilliped
3 with medial distal margin of apical segment of endopod with
8-13 broad teeth-like spines, outer margin with 1—2 broad teeth-
like spines, exopod long and narrow, tip over-reaching distal end
of basal endopod segment; pereiopod | with long carpus and
short and broad chelae; pereiopod 2 with exopod extending to
apex of merus; dactylus of pereiopod 3 with prominent terminal
claw and 7-9 strong spines on medial margin, exopod extends to
apex of merus; dactylus of pereiopod with a prominent terminal
claw and 6-12 spines on medial margin, exopod extends to mid
merus; dactylus of pereiopod 5 with prominent terminal claw
and very regular comb-like row of 44-60 small spines on
medial margin, exopod extends to basal third of merus.
Carapace length 6.20 (5.70—6.20) mm.
Rostrum (fig. 9a) long 6.20 (5.70-6.05) mm, extending
beyond both antennular peduncle and scaphocerite, rostral
length 0.81 (0.81—1.16) times longer than carapace, dorsal edge
straight, moderately slender and pointed; rostral length, 6.25
(6.25—8.25) greater than width; dorsally armed with 28 (28—
30) teeth, ratio of rostral spines and rostral length is 5.60
P.J. Suter, J.H. Mynott & M. Crump
(4.24—5.60), with 1-3 postorbital spines (fig. 9a); ventrally with
9 (8-10) large serrations over a length of 1.65 (1.65—3.0) mm, 1
or 2 spines posterior of greatest depth, distal half of ventral
edge straight, ratio of ventral spine length and rostral length is
0.33 (0.33-0.45) with 3.11 (3.00-3.11) times more dorsal
spines than ventral spines; rostral length 1.11 (1.11-1.83) times
length of scaphocerite.
Antenna 1 (fig. 9b) peduncle 4.45 (3.78—4.56) mm long,
not quite reaching distal tip of scaphocerite, length 0.99
(0.59—1.27) times length of scaphocerite. Stylocerite 2.53
(2.2-2.53) mm long, length 7.21 (7.21-9.17) times longer than
width, 0.41 (0.39-0.41) times carapace length, reaching
beyond distal border of peduncle segment almost to end of
broad acute process on distal angle of first segment.
Antenna 2 (fig. 9c) second segment of peduncle 1.50
(1.25-1.68) mm long, 0.33 (0.29-0.4/) times length of
scaphocerite, 2.50 (2.38-3.23) times longer than wide
scaphocerite 4.50 (3.60-4.50) mm long, 0.73 (0.63-0.73)
times carapace length, 2.90 (2.57-2.93) times as long as wide.
Mouthparts. Left mandible (fig. 9d, e) with 4 teeth separated
by finely ridged notch from a less distinct apical tooth; spine
row immediately below incisor process of 10 rugose spines
(lifting spines); spine row above molar process of approximately
over 20 sparsely setose spines. Right mandible (fig. 9f, g) with
4-5 teeth 1n a single incisor process, all teeth large and of equal
length; spine row immediately below teeth with 9 spines each
finely setose basally; spine row above molar process. Molar
process ridged.
Maxilla 1 (fig. 9h) palps short, truncate, with | long, setose
terminal spine and | small, simple subterminal one, inner
distal angle rounded.
Maxilla 2 palps small, but longer than P. australiensis,
terminal parts narrow and with 1 sub-apical setose spine.
Scaphognathite rounded apically, extending to apex of upper
endite (fig. 91).
Maxilliped | (fig. 9J) palp with broad base, short narrow
distal lobe, and several long setose spines on distal margins.
Exopod flagellum distinct, well developed and with numerous
long setose spines on all margins, approximately half length of
caridean lobe.
Maxilliped 2 (fig. 9k) endopod 1.19 (1.19-1.26) mm long;
exopod long and narrow, length 1.43 (1.33—1.43) mm, exopod
1.06-1.20 times longer than endopod. Epipodite with
podobranch.
Maxilliped 3 (fig. 10a) endopod length 4.03 (3.62--6.33)
mm, 2.41 (2.41-2.54) times longer than exopod; with 3 distal
segments of similar length; basal segment curved, apical
segment with large terminal claw, medial distal margin with 9
(8-13) broad teeth-like spines, largest ın basal third, outer
margin with | apical broad tooth-like spines. Exopod long
and narrow 1.67 (1.43—2.64) mm long, tip extends beyond
distal end of basal endopod segment.
Thoracic appendages. Pereiopod 1 (fig. 10b) 4.98 (4.68—
4.98) mm long, 0.80 (0.50-0.52) times carapace length.
Chelae short and broad (fig. 10b), 1.65 (1.60-1.68) mm long,
propodus 2.87 (2.68—2.87) times as long as wide, 1.83 (1.83—
1.97) times longer than dactylus 1.29 (1.29-1.43) times longer
than carpus; palm length 1.47 (1.44—1.79) palm width and
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 13
Figure 9. Paratya spinosa sp. nov.: a, head region and rostrum; b, antenna | peduncle and stylocerite; c, scaphocerite; d, left mandible; e, enlarged
incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; 1, maxilla 2; j}, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
74 P.J. Suter, J.H. Mynott & M. Crump
Figure 10. Paratya spinosa sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 15
Figure 11. Paratya spinosa sp. nov.: a, pereiopod 4; b, dactylus 4; c, pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g, pleopod | of
female; h, pleopod 1 of male; 1, endopod 1 of male. Scale lines 0.2 mm.
16
0.94 (0.93-1.06) times dactylus length. Carpus long 2.32
(2.27-2.50) times longer than greatest width, broadening
distally, distal margin excavate. Segment ratios 0.71 (0.68—
0.74) : 1.29 (1.29-1.43) : 1.00 (1.30 [1.12-1.28] mm) : 1.20
(1.18—1.30) : 0.41 (0.41-0.45) : 2.55 (—). Exopod extending to
mid carpus.
Pereiopod 2 (fig. 10c) 7.05 (6.71—7.05) mm long, 1.14
(1.11-1.18) times carapace length. Chelae long and slender
1.55 (1.50-1.55) mm long (fig. 10c), 3.10 (3.00—3.19) times
longer than wide, 0.65 (0.65—0.69) times carpus length; palm
length 1.80 (1.45-1.82) times longer than palm width, 0.77
(0.53—0.77) times length of dactylus. Propodus length 1.77
(1.53—1.77) times longer than dactylus. Carpus 7.38 (5.60—
7.38) times longer than greatest width, slightly broader
distally with small excavation. Segment ratios 0.36 (0.36—
0.45) : 0.65 (0.65—0.69) : 1.00 (2.40 [2.18—2.40] mm) : 0.80
(0.80) : 0.49 (0.49—0.59) : 1.41 (1.41—1.43). Exopod extending
to apex of merus.
Pereiopod 3 (fig. 10d, e) distinctly longer than pereiopod 2
and more slender 9.08 (7.48—9.08) mm long, 1.46 (1.31-1.46)
times carapace length, dactylus with prominent terminal claw
and 9 (7-9) strong spines on medial margin (fig. 10e).
Propodus length 4.56 (4.56—4.80) times longer than dactylus,
length 12.30 (10.67-12.30) times longer than wide with 8
(6—8) spines on inner margin, outer margin lacks spines.
Carpus with 1 large subapical spine. Merus with 1 strong
Spine on medial margin and | near ventral distal margin;
segment ratios 0.44 (0.34—0.44) : 1.98 (1.64-1.98) : 1.00 (1.55
[1.55-1.56] mm) : 2.35 (1.58-2.35) : 0.52 (0.52-0.57) : 1.65
(—). Exopod extends to mid merus.
Pereiopod 4 (fig. 11a, b) similar length to pereiopod 3, 9.08
(7.68—9.20) mm long, 1.46 (1.35—1.52) times carapace length.
Dactylus with prominent terminal claw and 7 (7—8) spines on
medial margin (fig. 11b). Propodus length 4.96 (4.11—4.96)
times longer than dactylus, 12.90 (9.74—13.33) longer than
wide, with 10 (6-12) spines on medial margin, outer margin
without spines; carpus with large subapical spine; merus with
| strong spine on medial margin and | near ventral distal
margin. Segment ratios 0.40 (0.37—0.40) : 1.98 (1.50-1.98) :
1.00 (1.63 [1.63-1.65] mm) : 2.09 (1.69—2.20) : 0.51 (0.49—
0.56) : 1.71 (1.71—1.82). Exopod extends to mid merus.
Pereiopod 5 (fig. 11c, d). Slightly longer than pereiopods 3
and 4, 9.65 (8.45-9.65) mm long, 1.56 (1.48-1.56) times
carapace length. Dactylus with prominent terminal claw and
very regular, comb-like row of numerous (44—55) small
spines on medial margin (fig. 11d). Propodus length 3.92 (—)
times longer than dactylus, length 14.50 (12.6—14.50) times
longer than wide with 7 (5—11) long medial teeth, external
margin lacking spines. Carpus without large spines near distal
margin. Merus with | strong medial spine and lacking a distal
spine; segment ratios 0.55 (—) : 2.16 (1.56-2.16) : 1.00 (1.68
[1.50—1.81] mm) : 1.85 (1.57—2.23) : 0.75 (0.54—0.75) : 1.67
(1.21—1.92). Exopod extends to mid merus.
Abdomen. Pleopods peduncle of first pleopod short, 0.40
(0.33—0.40) times length of carapace, 4.17 (3.08—4.17) times
longer than wide, exopod 1.14 (1.14—1.43) times peduncle
length, endopod (fig. 11g), (0.53—0.58) times peduncle length;
second pleopod peduncle short, 0.43 (0.37—0.43) times length
P.J. Suter, J.H. Mynott & M. Crump
of carapace, 2.94 (2.81-3.00) times longer than wide, exopod
1.13 (1.13-1.29) times peduncle length, endopod slightly
shorter 1.06 (1.06—1.22) times peduncle length. Length of first
peduncle 1.06 (1.06—1.13) times length of second peduncle.
Telson (fig. lle, f) length 4.41 (3.7-4.41) mm, 0.71 (0.63—
0.71) times carapace length, 5.14 (2.98—3.14) times longer than
ereatest width and tapering distally. Dorsal surface with 1—2
pairs of strong submarginal teeth-like spines. Posterior
margin convex with | pair of teeth-like spines outermost, 7
(6-10) long, strong setose spines (fig. 111).
Uropods approximately equal to telson length.
Males smaller than females, carapace length 4.5 mm; with
endopod of pleopod | strongly excavate apically with 12
external spines on medial margin and 15 long setae on inner
margin (fig. 111).
Etymology: Spinosa after the very spiny rostrum with 28—30
dorsal spines and 9—10 ventral spines.
Comments: P. spinosa 1s most similar to P. tasmaniensis but
may co-occur with P. walkeri. P. spinosa can be distinguished
from all other long rostrum species by the combination of
characters listed in Table 2 including the number of dorsal
rostral spines (28-30); number of ventral rostral spines (9-10);
right mandible incisors with 4-5 teeth; shape of scaphognathite
of maxilla 2.
P. spinosa has a restricted distribution in streams of the
northern coastal streams in New South Wales (Tweed R
catchment) while P. tasmaniensis 1s widespread in Tasmania,
coastal Victoria, New South Wales and Queensland and in the
Murray-Darling catchment in New South Wales and South
Australia. Cook et al. (2006) recorded lineage 3 in the
Clarence R system but we recorded P. spinosa only in the
Tweed R system. It is possible that both P. spinosa and P.
walkeri may coexist in the Tweed R.
Paratya arrostra (Riek), 1953 comb. nov.
Figures 12-14
Paratya australiensis arrostra Riek, 1953; in part = rostrum mid
length (fig. 12a)
Terrors Ck, Dayboro, Queensland. Types examined by MC.
Paratya atacta Riek 1953; in part = rostrum very long, comb. Nov.
(fig. 12c)
Upper Nerang R, southern Queensland
Paratya atacta adynata Riek 1953; in part = rostrum mid length,
comb. Nov. (fig. 12b). Small creek ın upper reaches of Middle Harbour,
Sydney, New South Wales
P. australiensis Williams and Smith (1979); neotype male selected
from material named by Riek (1953), AM P28693.
Paratya australiensis Gan et al. (2016); determination of the
mitogenome of Paratya australiensis.
Lineage 4 (Cook et al., 2006)
Lineage C (McClusky, 2007)
Material Examined: Victoria: Hughes Ck at Hughes Ck Rd,
—37.0075 S, 145.3212 E, 28 September 2011 (PS, JM, MC); King
Parrot Ck at Flowerdale, —37.2953 S, 145.2905 E, 28 September 2011
(PS, JM, MC); Goulburn R past Loch Gary at flood markers,
—36.2411 S, 145.2866 E, 28 September 2011 (JM, MC); Yea R at
Glenburn, —37.4239 S, 145.4210 E, 28 September (PS, JM, MC);
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. ff
in
In,
=
: = =
; NOS
a
=
Sn
LY
pL
en
Figure 12. Paratya arrostra Riek: a, head region and rostrum of lineage 4 long; b, head region and rostrum of lineage 4 short; c, head region and
rostrum of lineage 4c; d, antenna | peduncle and stylocerite; e, scaphocerite; f, maxilla 1; g, left mandible; h, enlarged incisors; 1, right mandible;
j, enlarged incisors; k, maxilla 2; 1, maxilliped 1; m, maxilliped 2. Scale lines 0.2 mm.
18 P.J. Suter, J.H. Mynott & M. Crump
|
we
=.
re
Figure 13. Paratya arrostra Riek: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3. Scale lines 0.2 mm
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 19
Figure 14. Paratya arrostra Riek: a, pereiopod 4; b, dactylus 4; c, pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g, pleopod 1 of
female; h, endopod 1 of male. Scale lines 0.2 mm.
80
Wimmera R downstream of Dimboola Weir, —36.4557 S, 142.0167 E,
6 March 2012 (Vic EPA); Ovens R near Wangaratta, —36.3371 S,
146.3191 E, August 2010 (JM, MC, JM).
New South Wales: Wakool Reserve, —35.4963 S, 144.4541 E, June
2011 (JC); Bagnall’s Lagoon, Albury, —36.070 S, 146.854 E, April
2011 (PS); Murray R below Lake Hume, —36.0998 S, 147.0228 E, 12
August 2010 (JW, MC, JM); Hawksbury R at Windsor Bridge,
—33.6023 S, 150.8233 E, 10 November 2011 (SW); Hawksbury R at
Wilberforce, —33.6020 S, 150.8241 E, 10 September 2011 (SW);
Hawksbury R at Sackville Ferry, -33.5003 S, 150.8746 E, 19
September 2020 (SW); Hawksbury R North Richmond, —33.5684 S,
150.7485 E, 9 March 2011 (SW); South Ck Richmond Rd, —33.6775 S,
150.8121 E, 30 March 2011 (SW); McCarrs Ck, -33.6410 S,
151.2780 E, 13 September 2011 (SW); Nepean R at Sharpes Weir,
—34.0384 S, 150.6793 E, 15 September 2011 (SW); Nepean R at Grove
Rd, —34.0415 S, 150.6964 E,2 April 2011 (SW); Nepean R at Wallacia
Bridge, —33.8655 5, 150.6374 E, 18 April 2011 (SW); Nepean R at
Maldon Weir, —34.2034 S, 150.6301 E,5 April 2011 (SW); Lachlan R,
Newell Highway at Forbes, —33.3956 S, 147.9903 E, 3 November 2011
(PS, JM, MC); Lachlan at Cargellico —32.2033 S, 146.3589 E, Jun
1999 (PS, TC); Lachlan R at Glenmore, —33.4413 S, 145.5377 S, July
1999, (PS, TC); Lachlan R at Goolagong, —33.6060 S, 148.4324 E, July
1999 (PS, TC); Lachlan R at Condobolin, —33.0915 S, 147.1476 E, July
1999 (PS, TC); Macquarie R at Dubbo, —32.2470 S, 148.5990 E, 23
september 2010 (PS, MC); Murrumbidgee R at Wagga Wagga;
—35.1041 S, 147.3751 E, 11 November 2003 (PS); Billabong Ck at
Coree, —35.3556 S, 145.5041 E, 24 June 2001 (PS, LS); Billabong Ck
at Moulamein, —35.0913 S, 144.0334 E, 2 April 2001 (PS, LS);
Billabong Ck at Urana, —35.3598 S, 146.0942 E, 14 May 2001 (PS,
LS); Billabong Ck at Walbundrie, —35.6971 S, 146.7253 E, 14 May
2001 (PS, LS); Billabong Ck at Wanganella, —35.2124 S, 144.3150 E,
2 April 2001 (PS, LS); Way Way Ck, —30.7680 S, 152.9427 E, 24 May
2016 (BM); Maguire Ck —28.8367 S, 153.3364 E Jun 2020 (SO);
Maguire Ck -28.8367 S, 153.3364 E (BM); Tucki Tucki Ck,
—28.8225 S, 153.3362 S, June 2020 (SO); Pinebrush Ck, —30.1306 S,
153.1328 E (BM); Small creek in upper reaches of Middle Harbour,
Sydney, type locality for P. atacta adynata Riek 1953; Wattamollaa
Ck on Clinton Park Rd Kangaroo Valley, 534.7371 E, 150.5929 S, 27
september 2017 (PS, JM, JH); Kangaroo R at Hampden Bridge,
—3472772, S, 150.5218 E, 26 September 2017 (PS, JM, JH); Stream on
Broger Rd Shoalhaven catchment, —34.7105 S, 150.6827 E, 27
september 2017 (PS, JM, JH); Stream on Jarretts Rd — Upper
Kangaroo Valley Rd, —34.7036 S, 150.5880 E, 27 September 2017 (PS,
JM,JH); Brogers Ck in Shoalhaven catchment, —34.7105 S, 150.6827 E,
27 September 2017 (PS, JM, JH); Orara R at Nana Glen, —30.1328 S,
153.0077 E, no date, (BK); Brogo R at Brogo, —36.5402 S, 149.8265 E,
10 March 1999; Williams R at Coreei Bridge Dungog, —32.3968 S,
151.7631 E, 30 October 2011 (PS, JM, MC); The Falls at Forest Falls
Retreat Johns R, —31.709 S, 152.6612 E, 31 October 2011 (MC, JM,
PS); The Cascades at Forest Falls Retreat Johns R, —31.70 S,152.655 E,
31 October 2011 (MC, JM, PS); Lake Yarrunga at Bendeela recreation
area —34.7398 S, 150.4705 E, 27 September 2017 (PS, JM, JH).
South Australia: Brenda Park wetland south of Morgan,
—34.0818 S, 139.6743 E, 8 November 2011 (CM).
Queensland: Terrors Ck, Dayboro, type locality for P.
australiensis arrostra Riek 1953; Upper Nerang R, southern
Queensland, type locality for P. atacta Riek 1953; Kilcoy Ck, upper
Brisbane R, —26.94 S, 152.568 E, no date (BC); Boar Pocket Ck,
Tinaroo, —17.1708 S, 145.6447 E, 20 October 2017 (BM, BKr).
Diagnosis: P. arrostra differs from all other species by the
following combination of characters: rostrum variable, long,
extending beyond antennular peduncle or extending beyond
both the antennular peduncle and scaphocerite, or short not
P.J. Suter, J.H. Mynott & M. Crump
extending beyond the peduncle or intermediate extending
beyond peduncle but not the scaphocerite; dorsal edge slightly
concave or straight, dorsally armed with 22—34 teeth, 2—3
postorbital spines, ventrally with 3—11 large serrations over a
length of 0.40-2.20mm; distal half of ventral edge straight or
curved; left mandible with 2—5 (usually 4—5) teeth separated by
a smooth angular notch from a distinct apical tooth; right
mandible with 4 teeth in 2 separate incisor processes;
scaphognathite of maxilla 2 rounded apically extending to apex
of upper endite; maxilliped 1 with exopod flagellum distinct,
well developed and with numerous long setose spines on all
margins, over half length of caridean lobe; exopod of maxilliped
2 2.18—3.45 times longer than endopod, epipodite with long
podobranchs extending just to basal third of third segment of
endopodite; maxilliped 3 with medial distal margin of apical
segment of endopod with 6-8 broad teeth-like spines, outer
margin with 2 broad teeth-like spines, exopod long and narrow,
tip over-reaching distal end of basal endopod segment;
pereiopod | with long carpus and long slender chelae, exopod
extending to mid to apex of carpus; pereiopod 2 with exopod
extending to apex of merus or base of carpus; dactylus of
pereiopod 3 with prominent terminal claw and 9-11 strong
spines on medial margin, exopod extends to mid merus to base
of carpus; dactylus of pereiopod 4 prominent terminal claw and
8-12 spines on medial margin, exopod extends to mid merus;
dactylus of pereiopod 5 with prominent terminal claw and very
regular comb-like row of 70—90 small spines on medial margin,
exopod extends to mid merus.
Morphotypes of P. arrostra: P. arrostra specimens with a
very short rostrum not extending beyond the second segment
of the antennular peduncle and only to mid scaphocerite
(Lineage 4C) can be distinguished from all other species of
Paratya by this short rostrum, dorsal edge straight and curved
down at end, dorsally armed with 16-19 spines, O-1
postorbital spines, with postorbital separated from other
rostral spines, ventrally with 4—5 large serrations over a length
of 0.60-1.2 mm, extending from posterior of greatest depth;
distal half of ventral edge straight (Table 3).
P. arrostra with the shorter rostrum which does not extend
beyond the scaphocerite (lineage 4B) can be distinguished
from all other species of Paratya by the following combination
of characters: 3—7 ventral spines on rostrum extend over a
length of less than 1.80 mm; rostral length approximately
equal to scaphocerite length 0.73—1.24; exopod of pereiopod
1, 2 and 3 extending to mid merus to base of carpus.
P. arrostra specimens with the longer rostrum character
that extends beyond the end of the scaphocerite (Lineage 4B,
4E) differs from all other species of Paratya by the following
combination of characters: 4—11 ventral spines extending over
a length of 0.6-2.2 mm; rostral length 1.14—-1.31 times longer
than scaphocerite length; carpus of pereiopod | short; chelae
of pereiopod | long and slender (Table 3).
Carapace length 5.10-7.00 mm.
Rostrum variable length either (1, lineages 4B, 4E) long,
extending beyond antennular peduncle and to or beyond the
end of the scaophocerite (fig. 12a), dorsally slightly concave,
moderately slender; length 5.7-7.0 mm, 1.04-1.08 times
length of carapace; dorsally armed with 23-34 teeth, ratio of
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 01
rostral spines to length 4.6—6.67; 2-3 postorbital eye spines
(fig. 12a), ventrally with 4-11 large serrations over a length of
1.10-2.2 mm, extending from posterior of, or from, greatest
depth (fig. 12a), distal half of ventral edge straight or curved,
ratio of ventral spines to rostral length is 0.11—0.34; rostral
length 7.86—8.83 times depth, length 1.14—1.31 times length of
scaphocerite or (11 lineage 4B) rostrum short, length 2.50—5.30
mm, rostrum not extending beyond antennular peduncle (fig.
12b); 0.63—0.91 times length of carapace, dorsal edge straight
and may be angled downwards (fig. 12b); dorsally armed with
22—29 teeth, ratio of rostral spines to length 4.34—7.60, 2—3
postorbital eye spines (fig. 12b), ventrally with 3—7 large
serrations over a length of 0.4—1.8mm all forward of greatest
depth (fig. 12b), distal half of ventral edge straight; rostral
length/depth 6.33-8.53, length 0.73-1.24 times length of
scaphocerite or (111 lineage 4C) rostrum very short 3.10—3.50
mm, not extending beyond the second segment of the
antennular peduncle (fig. 12c) and only to half scaphocerite,
rostral length 0.49—0.60 times length of carapace, shape broad
and pointed, dorsal edge straight and curved down at end;
dorsally armed with 16-19 teeth (fig. 12c), ratio of dorsal
spines to length is 5.15—5.43 and 3.40—4.00 times more spines
than ventral spines; 0—1 postorbital eye spines, when present
spine distinctly posterior and separated from other rostral
spines (fig. 12c); ventrally with 4—5 large serrations over a
length of 0.60—1.20 mm, extending from posterior of greatest
depth, ratio of ventral spines to rostral length 1s 0.18—0.34;
distal half of ventral edge straight; rostral length/depth, 5.17—
6.60; length 0.80—0.90 times length of scaphocerite.
Antenna | (fig. 12d) peduncle not quite reaching distal tip
of scaphocerite, but similar length to scaphocerite itself,
0.42-1.06 times as long as scaphocerite; lateral distal angle of
first segment with prominent blunt process at outer distal
margin with small acute tooth on outer margin of segment but
may be absent in some variations. Stylocerite 1.54—2.48 mm
long, length 7.21-9.17 longer than width, 0.39—0.4] times
carapace length, reaching beyond distal border of peduncle
segment (fig. 12d) almost to end of acute process on distal
angle of first segment (fig. 12d).
Antenna 2 (fig. 12e) second segment length 1.25—1.68mm
long, 0.23—0.34 times length of scaphocerite, 2.00—3.36 longer
than width. Scaphocerite 3.70—5.10 mm long, 0.64—0.94 times
carapace length and 2.73—4.00 times as long as wide.
Mouthparts. Left mandible (fig. 12g, h) with 2—5 (usually
4—5) teeth separated by smooth angular notch from a distinct
acute apical tooth; spine row immediately below incisor
process of 8—10 rugose spines (lifting spines); spine row
above molar process of approximately over 20 sparsely setose
spines. Right mandible (fig. 121, j) with 4 teeth in 2 separate
incisor processes with first and third teeth largest and second
and fourth smaller; spine row immediately below teeth with
8-10 spines each finely setose basally; spine row above molar
process. Molar process ridged.
Maxilla | as for P. australiensis (fig. 12f).
Maxilla 2 as for P. australiensis (fig. 12k).
Maxilliped 1 as for P. australiensis (fig. 121).
Maxilliped 2 (fig. 12m) endopod length 0.44—1.11 mm;
exopod long and narrow, length 2.07—2.69 mm, exopod 2.18—
3.45 longer than endopod. Epipodite with long podobranch
extending to basal third of third segment of endopodite.
Maxilliped 3 (fig. 13a) endopod length 5.72-7.23 mm,
2.20—2.75 times longer than exopod, with 3 distal segments of
similar length; basal segment curved, apical segment with
large terminal claw, medial distal margin with 6-5 broad
teeth-like spines, largest 2 or 3 1n basal half, outer margin
with 2 long teeth-like spines ın apical third. Exopod long and
narrow 2.33-3.07 mm, tip over-reaching distal end of basal
endopod segment.
Thoracic appendages. Pereiopod 1 (fig. 13b) length 3.51—
5.23 mm, 0.66—0.89 times carapace length. Chelae short and
slender (fig. 13b), 1.14—2.08 mm long, propodus 2.50-3.47
times as long as wide, 1.72—3.06 times longer than dactylus,
1.08-1.84 times longer than carpus; palm length 1.13—1.90
times palm width and 0.78—2.09 times dactylus length. Carpus
long, 1.95-2.76 times as long as greatest width, broadening
distally, distal margin excavate. Merus approximately one-
third longer than carpus, parallel-sided. Ischium about one-
quarter length of merus. Segment ratios 0.59—0.75 : 1.08—1.84
: 1.00 (1.01-1.40) mm : 1.14-1.84 : 0.40—0.50 : 1.65-3.16.
Exopod extending to mid-apex of carpus.
Pereiopod 2 (fig. 13c) length 5.49—7.69 mm, 0.99-1.99
times carapace length. Chelae long and slender (fig. 13c)
1.15-1.63 mm long, half to two-thirds length of carpus, 2.87—
4.20 times as long as wide; palm length 1.13—2.08 times
longer than wide and 1.05-2.00 longer than dactylus.
Propodus length 1.43-1.98 times longer than dactylus. Carpus
6.20—8.81 times as long as greatest width, slightly broader
distally, distal margin with small excavation. Merus shorter
than carpus, parallel-sided. Ischium about half as long as
merus. Segment ratios 0.28—0.43 : 0.50—0.62 : 1.00 (2.29—
2.72) mm : 0.73-0.88 : 0.30-0.46 : 1.14-1.22. Exopod
extending to apex of merus to base of carpus.
Pereiopod 3 (fig. 13d, e) distinctly longer than pereiopod 2
and more slender 7.27-9.24 mm long, 1.28-1.57 times
carapace length. Dactylus with prominent terminal claw and
9—]l strong spines on medial margin (fig. 13e). Propodus
length 3.58—4.80 times longer than dactylus, length 11.61—
15.77 times longer than wide with 6-13 spines on inner
margin. Merus longer than propodus with 1—5 strong spines
(usually 2) on medial margin and 1 near ventral distal margin;
ischium approximately one-quarter to one-third length of
propodus; segment ratios 0.36—0.51 : 1.60-1.88 : 1.00 (1.39—
1.75) mm : 1.25-2.05 : 0.46—0.86 : 1.32-1.70. Exopod extends
to mid-merus to base of carpus.
Pereiopod 4 (fig. 14a, b) similar to pereiopod 3, 7.69—9.81
mm long, 1.29-1.67 times carapace length. Dactylus with
prominent terminal claw and 8—12 spines on medial margin
(fig. 14b). Propodus length 3.05-5.00 times longer than
dactylus, length 11.11-13.44 times longer than wide, with
11-16 spines on medial margin; merus with 1-3 strong spine
on medial margin and | near ventral distal margin. Segment
ratios 0.37—0.52 : 1.59-1.79 : 1.00 (1.49-1.93) mm : 1.83-2.38
: 0.50—0.68 : 1.35-1.50. Exopod extends to mid merus.
Pereiopod 5 (fig. 14c, d) similar length to pereiopods 3 and
4, 7.85—9.]9 mm long, 1.37-1.56 times carapace length.
Dactylus with prominent terminal claw and very regular,
82
comb-like row of numerous (70—91) small spines on medial
margin (fig. 14d). Propodus length 2.97-3.91 times longer
than dactylus, length 11.94-18.62 times as long as wide with
9-13 long medial spines and external margin without spines.
Carpus approximately half propodus length without any large
spines near distal margin. Merus similar length to propodus,
with 1 strong medial spine and 1 distal spine; ischium one-
third length of propodus; segment ratios 0.46—0.65 : 1.79-1.97
: 1.00 (1.52-1.80) mm : 1.54-1.97 : 0.47—0.66 : 1.29-1.61.
Exopod extends to mid to apical third of merus.
Abdomen. Pleopods peduncle of first pleopod short,
0.26—0.37 times length of carapace length, 2.40—3.45 times
width, exopod 1.29-1.81 times peduncle length, endopod
0.63—0.69 times peduncle length (fig. 14g); second pleopod
peduncle short, 0.31-0.47 times length of carapace, 2.47—3.67
times width, exopod 1.09-1.35 times peduncle length,
endopod slightly shorter 0.91—1.27 times peduncle length.
Length of first peduncle 1.18—1.28 times length of second
peduncle. Peduncle of pleopod 5 0.20-0.28 times length of
carapace, 1.68-2.31 times width; exopod length 1.57-1.93
times peduncle length; endopod 1.22-1.57 times peduncle
length; exopod length 1.14—1.29 times endopod length.
Telson (fig. 14e, f) length 3.10—4.10 mm, 0.51—0.76 times
carapace length, 2.36—3.75 times as long as greatest width,
and tapering distally, Dorsal surface with 0—2 pairs of strong
submarginal teeth-like spines; posterior margin convex with 1
pair of teeth-like spines outermost, 7-13 (usually 11—12) long,
strong setose spines (fig. 141).
Uropods approximately equal to telson length, exopod
1.05-1.33 times telson length, length 2.84—3.19 times width;
endopod 0.98-1.29 times telson length, length 3.28—4.25
times width.
Males smaller than females, carapace length 4.36 mm;
endopod of first pleopod strongly excavated apically with
9-10 external spines and 10—14 long setae on inner margin
(fig. 14h).
Comments: This 1s the most variable species of Paratya showing
distinct rostral characteristics from rostrum shorter than the
peduncle, rostrum longer than peduncle but not extending
beyond the scaphocerite and rostrum long extending beyond the
scaphocerite. Riek (1953) observed these distinct groups and
described them as species or subspecies based on morphological
character expression only. Williams and Smith (1979) considered
all the taxa described by Riek were synonyms of P. australiensis.
With the development of molecular techniques, it is now possible
to recognise that these variants are all a single taxon and the
taxon P. australiensis arrostra Riek is here raised to species
level. Genetically, all Lineage 4 specimens have low intraspecific
variation, and although Lineage 4C can be reliably identified
morphologically (Table 3), it is slightly more difficult to reliably
define lineages 4A, 4B, 4D and 4E (Cook et al., 2006)
morphologically (Table 2) on the limited material we have been
able to analyse. The presence of different morphotypes in this
species 1s similar to the observations by Choy et al. (2019) in the
Australian atyid shrimp Australatya.
P. arrostra 1s widely distributed through the Murray—
Darling Basin, south-eastern coastal streams in Victoria and
P.J. Suter, J.H. Mynott & M. Crump
New South Wales, north-eastern New South Wales coastal
streams, and south-eastern and northern Queensland (fig.
32a). Lineage 4A has been recorded from northern coastal
streams in Queensland but we do not have any specimens of
this lineage. Lineage 4C is restricted to south-eastern
Queensland in the catchments of the Maroochy R, Mary R
and Brisbane R. We have recorded P. arrostra to occur at sites
in South Australia with P. rouxi and P. tasmaniensis; with P.
rouxi, P. whitemae and P. strathbogiensis in Murray—Darling
Basin rivers and with P. whitemae in coastal rivers.
Gan et al. (2016) defined the complete mitogenome of a
species designated as P. australiensis from the Lodden R at
Baringhup, Victoria. The mitochondrial genome is 15,990
base pairs in length (GenBank accession number: KM978917)
and has 37 mitochondrial genes (13 protein-coding genes, 2
rRNAs and 22 tRNAs) and a non-coding region of 1006 base
pairs (Gan et al. (2016). This genome (strain APRI2) was
analysed with the total GENBANK Paratya data
(Supplementary Table 1) and the species used by Gan et al.
(2016) was embedded with P. arrostra.
Paratya williamsi n. sp.
Figures 15-17
http://zoobank.org/urn:lsid:zoobank.org:act: F299988B-DFO9-
4765-AO01 E-B28875339C5D
Lineage 5 (Cook et al., 2006)
Type Material: Holotype New South Wales. Kangaroo R
Hampden Bridge, —34.7272 S, 150.5218 E, 26 September 2017 (PS,
JM, JH). Body in ethanol and antennae, mouthparts, pereiopods and
abdominal structures dissected, mounted on 2 slides. AM Ref No.
P.105603.001; Accession Ref. PS103, Genbank Registration
OL420884.
Paratypes: New South Wales. Kangaroo R Hampden Bridge,
—34 77272 S, 150.5218 E, 26 September 2017 (PS, JM, JH) Accession
Ref. PS98, AM Ref No. P.105.603, Kangaroo Valley R on Gerringong
Ck Rd, —34.6868 S, 150.6013E, Accession Ref. PS99, AM Ref No.
P.105603, 27 September 2017 (PS, JM, JH), bodies 1n ethanol and
other structures dissected, mounted on 2 slides each.
Material Examined: New South Wales: Hampden Bridge, 34.7272 S,
150.5218 E, 26 September 2017 (PS, JM, JH); Kangaroo Valley R on
Gerringong Ck Rd, —34.6868 S, 150.6013 E, 27 September 2017 (PS,
JM, JH).
Diagnosis: Paratya williamsi differs from all other species by
the following combination of characters: rostrum long, 4.50—
5.20 mm, extending beyond antennular peduncle and to end of
scaphocerite, rostral length 0.84-1.03 times longer than
carapace, dorsal edge curved upwards to tip, narrow and
pointed, rostrum 7.16-9.60 times longer than wide, dorsally
armed with 21-27 teeth, ratio of rostral spines to rostral length
is 4.38—5.38, with 2-3 postorbital spines; ventrally with 1-5
short serrations over a length of 0.10-2.00 mm, 1l spine
posterior to greatest depth, distal half of ventral edge straight,
ratio of ventral spine length to rostral length 1s 0.21—1.00 and
4.40—21.00 more dorsal spines than ventral spines; rostral
length 1.15—1.34 times length of scaphocerite. Left mandible
with 4—5 teeth separated by a notch from 3 less distinct apical
New species of Paratya (Decapoda: Atyidae) from Australian inland waters - linking morphological characters with molecular lineages. 83
b
Figure 15. Paratya williamsi sp. nov.: a, head region and rostrum; b, antenna | peduncle and stylocerite; c, scaphocerite; d, left mandible; e,
enlarged incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; i, maxilla 2; j, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
04 P.J. Suter, J.H. Mynott & M. Crump
Figure 16. Paratya williamsi sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 85
c
Figure 17. Paratya williamsi sp. nov.: a, pereiopod 4; b, dactylus 4; c, pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g, pleopod 1
of female; h, pleopod 2 of female; 1, pleopod 1 of male; j, endopod 1 of male. Scale lines 0.2 mm.
86
teeth; spine row immediately below incisor process of 4—6
rugose spines (lifting spines); right mandible with 4 teeth in a
single incisor process with apical and fourth teeth largest
separated by 2 smaller teeth. Maxilla 2 with scaphognathite
truncated distally, not extending to apex of upper endite. Chelae
of pereiopod | short and broad, 1.40-1.51 mm long, propodus
2.28-3.02 times as long as wide, 1.57—1.97 times longer than
dactylus, 1.27-1.44 times longer than carpus; palm length
1.36-1./0 times longer than wide and 1.00-1.27 times longer
than dactylus length. Carpus very short 2.10—2.33 times as long
as greatest width. Dactylus of pereiopod 3 with prominent
terminal claw and 5-7 strong spines on medial margin.
Dactylus of pereiopod 4 with prominent terminal claw and 5—6
spines on medial margin.
Carapace length 5.40 (4.85—6.00) mm.
Rostrum long, 5.40 (4.80—5.20) mm, extending beyond
antennular peduncle and to end of scaphocerite (fig. 15a),
rostral length 0.96 (0.84—1.03) times longer than carapace,
dorsal edge curved upwards to tip, narrow and pointed,
rostrum 8.00 (7.16—9.60) times longer than wide, dorsally
armed with 24 (21—27) teeth, ratio of rostral spines to rostral
length is 4.61 (4.38—5.38), with 2 (2—3) postorbital spines (fig.
15a); ventrally with 3 (1—5) short serrations over a length of
1.50 (0.10—2.00) mm, 1 spine at greatest depth (fig. 15a), distal
half of ventral edge straight, ratio of ventral spine length to
rostral length 1s 0.29 (0.21—1.00) and 8.00 (3.50—21.00) more
dorsal spines than ventral spines; rostral length 1.27 (1.15—
1.34) times length of scaphocerite.
Antenna | (fig. 15b) peduncle 2.75 (2.25-2.75) mm long,
not reaching distal tip of scaphocerite, 0.90 (0.90—1.00) times
length of scaphocerite. Stylocerite 2.25 (2.05-2.30) mm long,
length 7.50 (6.83-9.20) longer than wide, 0.42 (0.38—0.45)
times carapace length, reaching beyond distal border of
peduncle segment but not to end of broad acute process on
distal angle of first segment (fig. 15b).
Antenna 2 (fig. 15c) second segment of peduncle 1.50
(1.26—1.50) mm long, 0.37 (0.29—0.37) length of scaphocerite,
2.73 (2.29—2.73) longer than wide. Scaphocerite 4.10 (3.75—
4.35) mm long, 0.76 (0.73-0.82) times length of carapace,
3.28 (3.13-3.48) longer than wide.
Mouthparts. Left mandible (fig. 15d, e) with 4—5 teeth
separated by a notch from 3 less distinct apical teeth; spine
row immediately below incisor process of 4—6 rugose spines
(lifting spines); spine row above molar process of
approximately over 20 sparsely setose spines. Right mandible
(fig. I5f, g) with 4 teeth in a single incisor process with apical
and third teeth largest with second and fourth teeth smaller;
spine row immediately below teeth with 8-11 lifting spines;
Spine row above molar process. Molar process ridged.
Maxilla 1I as for P. australiensis (fig. 15h).
Maxilla 2 (fig. 151) scaphognathite truncated distally, not
extending to apex of upper endite; palps small, terminal parts
narrow and with | sub—apical setose spine.
Maxilliped 1 as for P. australiensis (fig. 15j).
Maxilliped 2 (fig. 15k) endopod length 0.88 (0.88—1.11)
mm; exopod long and narrow 2.15 (2.00—2.50) mm, exopod
2.44 (2.27-2.46) times longer than endopodite. Epipodite
with podobranch.
P.J. Suter, J.H. Mynott & M. Crump
Maxilliped 3 (fig. 16a) endopod length 6.10 (5.68—6.27)
mm, 2.71 (2.32—2.71) times longer than exopod; with 3 distal
segments of similar length; basal segment curved, apical
segment with large terminal claw, medial distal margin with 8
(6-8) broad teeth-like spines, largest | in basal third, outer
margin with 7 teeth-like spines. Exopod long and narrow, 2.25
(2.25-2.10) mm long, tip reaching basal third of mid segment,
with several long setose spines near tip and several short setose
spines near base. Epipodite with basal conical projection.
Thoracic appendages. Pereiopod | (fig. 16b) 4.65 (4.65—
4.73) mm long, 0.86 (0.78-0.98) times carapace length. Chelae
short and broad (fig. 16b), 1.40 (1.40—1.51) mm long, propodus
2.54 (2.28-3.02) times as long as wide, 1.87 (1.87-1.97) times
longer than dactylus, 1.27 (1.27-1.44) times longer than
carpus; palm length 1.36 (1.36-1.70) times longer than wide
and equal to dactylus length (1.00-1.27). Carpus very short
2.20 (2.10—2.33) times as long as greatest width, broadening
distally, distal margin excavate. Segment ratios 0.68 (0.68—
0.76) : 1.27 (1.27-1.44) : 1.00 (1.10 [1.05-1.10] mm) : 1.50
(1.43-1.52) : 0.45 (0.45—0.57) : 1.27 (1.27—2.71). Exopod
extending to mid merus (mid merus to mid carpus).
Pereiopod 2 (fig. 16c) 6.20 (5.80—7.50) mm long, 1.15
(1.15—1.25) times carapace length. Chelae long and slender
(fig. 16c), 1.50 (1.35-1.50) mm long, nearly two-thirds length
of carpus, 2.33 (2.25-3.33) times longer than wide, palm
length 1.56 (1.50—1.56) times longer than width and 0.82
(0.52—1.20) times dactylus length. Propodus length 1.76
(1.76—1.80) times longer than dactylus. Carpus 5.71 (5.14—
8.33) times as long as greatest width, slightly broader distally,
distal margin with small excavation. Segment ratios 0.43
(0.34—0.43) : 0.75 (0.60—0.75) : 1.00 (2.00 [1.80-2.50] mm) :
0.68 (0.68—0.80) : 0.68 (0.60—0.75) : 1.50 (1.24—1.50). Exopod
extending to apex of merus to base of carpus.
Pereiopod 3 (fig. 16d, e) distinctly longer than pereiopod 2
and more slender 8.45 (7.65—9.15) mm long, 1.56 (1.53-1.58)
times carapace length. Dactylus with prominent terminal claw
and 5 (5-7) strong spines on medial margin (fig. 16e). Propodus
length 4.58 (4.46-4.58) times longer than dactylus, length
11.00 (10.00-14.50) times longer than wide with 13 (11-13)
spines on inner margin. Merus with 1-3 strong spines on
medial margin and | near ventral distal margin; segment ratios
0.40 (0.37—0.40) : 1.83 (1.67—1.83) : 1.00 (1.50 [1.50-1.65]
mm) : 2.13 (2.03-2.13) : 0.67 (0.40—0.67) : 1.53 (1.53-1.88).
Exopod extends to mid merus (mid merus to apex of merus).
Pereiopod 4 (fig. 17a, b) similar to pereiopod 3, 7.80 (7.45—
9.63) mm long, 1.44 (1.34-1.54) times carapace length.
Dactylus with prominent terminal claw and 5 (5-6) spines on
medial margin (fig. 17b). Propodus length 5.40 (4.75—5.40)
times longer than dactylus, length 10.80 (10.00-11.40) times
longer than wide, with 11 (8—13) spines on inner margin; merus
with 0—2 strong spines on medial margin and 1 near ventral
distal margin. Segment ratios 0.36 (0.36—0.40) : 1.93 (1.79—
1.93) : 1.00 (1.40 [1.40-1.50] mm) : 1.93 (1.80-2.04) : 0.71
(0.50—0.71) : 1.71 (1.61—1.71). Exopod extends to mid merus.
Pereiopod 5 (fig. 17c, d) slightly shorter than pereiopods 4,
7.65 (7.20—8.30) mm long, 1.42 (1.38—1.48) times longer than
carapace. Dactylus with prominent terminal claw and very
regular, comb-like row of 50 (49—66) spines on medial margin
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. o
(fig. 17d). Propodus length 4.07 (3.93—4.07) times longer than
dactylus, length 14.25 (11.00—14.25) times longer than wide
with 7 (7—10) long inner teeth and setae on external margin.
Carpus with | large spines near distal margin. Merus with |
strong medial spine and | distal spine; segment ratios 0.50
(0.47—0.56) : 2.04 (1.91—2.20) : 1.00 (1.40 [1.25—1.60| mm) :
1.71 (1.50—1.92) : 0.71 (0.64—0.78) : 1.43 (1.31—1.43). Exopod
extends to base to third of merus.
Abdomen. Pleopods peduncle of first pleopod short 1.90
(1.50—2.35) mm, 0.35 (0.31-0.39) times length of carapace
length, 2.92 (2.92—5.85) times width, exopod 1.08 (1.08-1.28)
times peduncle length, endopod 0.60 (0.45-0.80) times
peduncle length (fig. 17g); second pleopod peduncle short,
0.42 (0.36—0.45) times length of carapace, 2.50 (2.33-2.81)
times width, exopod 1.16 (—) times peduncle length, endopod
slightly shorter 1.04 (-) times peduncle length (fig. 17h).
Telson (fig. 17e, f) length 3.50 (3.50-4.00) mm, 0.65
(0.67—0.72) times carapace length, 3.04 (2.81-3.17) times as
long as greatest width, tapering distally. Dorsal surface with 2
pairs of strong submarginal teeth-like spines. Posterior margin
convex with | pair of teeth-like spines outermost, 6 (6—8) long,
strong terminal setose spines (fig. 17T).
Uropods approximately equal to telson length.
Male smaller than female, carapace length 4.7mm;
endopod of pleopod 1 strongly excavated and strongly curved
(fig. 171, j) with 11 short spines on outer margin and 12 long
setae on inner margin (fig. 17).
Etymology: The specific epithet is in honour of the late Prof. W.
D. Williams who encouraged and inspired a generation of
Australian limnologists and who provided one of us (PS) a unique
opportunity for post-graduate study at Adelaide University.
Comments: P. williamsi has an overlap of distribution with P.
australiensis and P. arrostra. Characters that distinguish these
species from P. williamsi are listed in Table 2.
P. williamsi may also be confused with P. whitemae and
P. tasmaniensis, both of which are widespread and may have
distributions that overlap, but they can be distinguished by the
following combination of characters: dactylus of pereiopod 3
with 5-7 teeth; dactylus of pereiopod 4 with 5-6 teeth;
dactylus of pereiopod 5 with a comb of 49—74 spines; first
cheliped palm length 1.00-1.27 times dactylus length; 1-5
ventral rostral spines over a length of 0.10-2.00 mm (Table 2).
P. williamsi is restricted to the upper Kangaroo Valley in
the Shoalhaven R catchment south of Sydney and can be
found co-existing with P. whitemae and P. tasmaniensis.
Paratya whitemae n. sp.
Figures 18— 20
http://zoobank.org/urn:lsid:zoobank.org:act:24D43065-1D31-
42BC-A846-7BFB163778AB
Lineage 6 (Cook et al., 2006)
Lineage A (McClusky, 2007)
Type Material: Holotype New South Wales. The Falls at Forest
Falls Retreat Johns R, —31.709 S, 152.6612 E, 31 October 2011 (MC,
JM, PS); Body in ethanol and antennae, mouthparts, pereiopods and
abdominal structures dissected, mounted on 2 slides. Accession Ref.
MC108, AM Ref No. P.105604.
Paratypes: New South Wales. The Falls at Forest Falls Retreat
Johns R, —31.709 S, 152.6612 E,31 October 2011 (MC, JM, PS) Male,
Accession Ref MC110, Genbank Registration OL420801; Bagnall's
Lagoon, Albury, —36.070 S, 146.854 E, April 2011 Accession Ref.
MC95, 97, 99 (PS) Genbank Registration OL420871, OL420872,
OL420874; Wakool Reserve, —35.4963 S, 144.4541 E, June 2011 (JC);
Nepean R at Maldon Weir, —33.7414 S, 150.6846 E, 5 April 2011
Accession Ref. MC37 (SW), Genbank Registration OL420834;
Nepean R at Macquarie Grove Rd, —34.0414 S, 150.6953 E, 2 April
2011 Accession Ref. MC34 (SW), Genbank Registration OL420834;
Bedford Ck —33.75 S, 150.447 E Accession Ref. MC31 (SW), Genbank
Registration OL420830; Hawkesbury R at Wilberforce, —33.5702 S,
150.8382 E, 21 April 2011 Accession Ref. MC24 (SW), Genbank
Registration OL420824; O’Hares Ck near George R, -34.095 S,
150.835 E, 20 April 2011 Accession Ref. MC21-23 (SW), Genbank
Registration OL420821—OL420823; bodies in ethanol and other
Structures dissected, mounted on 2 slides each.
Material Examined: New South Wales: Bagnall’s Lagoon, Albury,
—36.070 S, 146.854 E, April 2011 (PS); Murray R below Lake Hume,
—36.0998 S, 147.0228 E, 12 Aug 2010 (JW, MC, JM); Wakool Reserve,
—35.496 S, 144.454 E, June 2011 (JC); Nepean R at Maldon Weir,
—33.7414 S, 150.6846 E, 5 April 2011 (SW); Nepean R at Macquarie
Grove Rd, —34.0414 S, 150.6953 E, 2 April 2011 (SW); Bedford Ck,
—33.75 S, 150.447 E, 4 May 2011 (SW); Hawkesbury R at Wilberforce,
—33.5702 S, 150.8382 E, 21 April 2011 (SW); O’ Hares Ck near George
R, -34.095 S, 150.835 E, 20 April 2011 (SW); Woolgoolga Ck,
-30.1306 S, 153.1378 E, (BM), Way Way Ck, —30.7681 S, 153.1378 E,
(BM); Nambucca Ck, —30.6408 S, 152.8558 E, (BM); Bellinger R,
—30.4261 S, 152.7794 E, (BM); The Falls at Forest Falls Retreat Johns
R, -31.709 S, 152.6612 E, 31 October 2011 (MC, JM, PS); Jerrys Ck
near Forest Falls Retreat Johns R, —31.7146 S, 152.6625 E, 30 October
2011 (MC, JM, PS); The Cascades at Forest Falls Retreat Johns R,
—31.70 S,152.655 E, 31 October 2011 (MC, JM, PS); Williams R at
Cooreei Bridge Dungog, —32.3968 S, 151.7631 E, 29 October 2011
(MC, JM, PS); Trimble Ck in Shoalhaven catchment, —34.6847 E,
150.5252 S, 26 September 2017 (PS, JM. JH); Small Ck on Kangaroo
Valley Rd, Kangaroo Valley, —34.7229 S, 150.5293 5, 27 September
2017 (PS, JM, JH); Lake Yarrunga at Bendeela Recreation Area,
—34.7398 S, 150.4705 E, 27 September 2017 (PS, JM, JH); Manning R
at Wingham Brush, —31.8706 S, 152.3825 E, 16 September 2016 (BM);
Kangaroo R Hampden Bridge, —34.7272 S, 150.5218 E, 27 September
2017 (PS, JM. JH); Lachlan R at Glenmore, —33.4413 S, 145.5377 S,
July 1999, (PS, TC); Lachlan at Cargellico —32.2033 5, 146.3589 E,
June 1999, (PS, TC); Orara R at Nana Glen, —30.1328 S, 153.0077 E,
no date, (BK); Hawksbury R at Sackville Ferry, —33.5003 S,
150.8746 S, 19 September 2020 (SW); Stream on Gerrigong Ck Rd,
Upper Kangaroo Valley, —34.6870 S, 150.6000 E, 27 September 2017
(PS, JM. JH); Dingo Ck, —30.3103 S, 152.9822 E, 24 May 2015 (BM);
Ellenborough R at Ellenborough Falls, —31.6113 S, 152.2925 E, 31
October 2011 (PS, JM,MC); Blaxland Ck on Armidale Rd, —28.8997 S,
152.7864 E, 8 December 2011 (JW, DB).
Diagnosis: P. whitemae differs from all other species by the
following combination of characters: rostrum long, extending
beyond both antennular peduncle and scaphocerite, dorsal edge
very slightly concave and curved upwards, dorsally armed with
20—34 teeth, 1—4 postorbital spines, ventrally with 4—11 large
serrations over a length of 1.30—2.80 mm, extending from just
posterior to greatest depth; distal half of ventral edge straight;
left mandible with 4—5 teeth separated by finely ridged U-shaped
notch from a short blunt/acute apical tooth; right mandible with
O0 P.J. Suter, J.H. Mynott & M. Crump
Figure 18. Paratya whitemae sp. nov.: a, head region and rostrum; b, antenna 1 peduncle and stylocerite; c, scaphocerite; d, left mandible; e,
enlarged incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; i, maxilla 2; J, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 89
Figure 19. Paratya whitemae sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3; f, pereiopod 4; g, dactylus 4.
Scale lines 0.2 mm.
90 P.J. Suter, J.H. Mynott & M. Crump
Figure 20. Paratya whitemae n. sp.: a, pereiopod 5; b, dactylus 5; c, telson; d, telson terminal spines; e, pleopod 1 of female; f, pleopod 1 of male;
g, endopod 1 of male. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 91
3—4 teeth in a single incisor process; exopod of maxilliped 2
1.63-2.96 times longer than endopod; maxilliped 3 with medial
distal margin of apical segment of endopod with 7-11 broad
teeth-like spines, outer margin with 1-3 broad teeth-like spines,
exopod long and narrow, tip over-reaching distal end of basal
endopod segment; pereiopod | with long carpus and short and
broad chelae, exopod extending to base-apex of carpus;
pereiopod 2 with exopod extending to mid merus to base of
carpus; dactylus of pereiopod 3 with prominent terminal claw
and 7—11 strong spines on medial margin, exopod extends to
mid merus; dactylus of pereiopod 4 prominent terminal claw
and 5-11 spines on medial margin, exopod extends to mid
merus; dactylus of pereiopod 5 with prominent terminal claw
and very regular comb-like row of 81-94 small spines on medial
margin, exopod extends to basal third of merus.
Carapace length 5.7 (4.5—6.6) mm.
Rostrum long 6.82 (4.55-6.82) mm, extending beyond the
antennular peduncle and scaphocerite (fig. 15a), rostral length
longer than carapace 1.2 (0.83-1.26) times length of carapace,
shape long and slender with dorsal edge curved upwards, pointed,
rostrum 8.52 (5.53-9.57) longer than rostral width; dorsally
armed with 33 (20—34) teeth, ratio of number of dorsal spines to
length is 4.84 (3.68—6.15), 2 (1—4) postorbital eye spines (fig.
18a); ventrally with 6 (4—14) large spines over a length of 1.88
(1.30-2.80) mm, with 1—2 spines posterior to greatest width (fig.
15a), distal half of ventral edge straight; ratio of ventral spine row
length to rostral length is 0.28 (0.23-0.46) and 4.57 (2.43—5.80)
times more dorsal spines than ventral spines; rostral length 1.38
(1.04-1.77) times length of scaphocerite.
Antenna | (fig. 18b) peduncle short 3.05 (3.05-5.28) mm,
not reaching distal tip of scaphocerite, length 0.62 (0.62—1.12)
times as long as scaphocerite. Stylocerite 1.97 (1.60—2.64)
mm long, length 0.35 (0.32-0.51) times carapace length,
reaching beyond distal border of peduncle segment and
middle to just beyond process on first segment (fig. 18b).
Antenna 2 (fig. 18c) second segment of peduncle 1.03
(1.03-1.80) mm, 0.21 (0.21—0.45) times length of scaphocerite,
length 2.50 (2.35-3.75) times width. Scaphocerite 4.94 (3.10—
5.20) mm long, 0.87 (0.69—0.96) times carapace length, 2.93
(2.93—3.85) times as long as wide,
Mouthparts. Left mandible (fig. 18d, e) with 4—5 teeth
separated by slightly ridged shallow U-shaped notch from a
short blunt/acute apical tooth; spine row 1mmediately below
incisor process of 7—9 rugose spines (lifting spines); spine
row above molar process of approximately over 40 sparsely
setose spines. Right mandible (fig. 151, g) with 3-4 robust
teeth in incisor process with 2 central teeth larger than apical
and inner teeth; spine row immediately below teeth with 8—12
spines each finely setose basally; spine row above molar
process. Molar process ridged.
Maxilla 1 as for P. australiensis (fig. 18h).
Maxilla 2 as for P. australiensis (fig. 181).
Maxilliped 1 as for P. australiensis (fig. 18}).
Maxilliped 2 (fig. 18k) endopod 1.40 (0.87—1.40) mm long;
exopod long and narrow, length 2.56 (1.73-2.87) mm, exopod
1.82 (1.63—2.96) times longer than endopod. Epipodite with
long podobranch extending to basal third of third segment of
endopodite.
Maxilliped 3 (fig. 19a) endopod length 7.0 (4.85—8.3) mm,
3.38 (2.32-3.38) times longer than exopod; with 3 distal
segments of similar length; basal segment curved, apical
segment with large terminal claw, inner margin with 9 (7-11)
broad teeth-like spines, largest 2—4 in basal half, outer margin
with | (1-3) long teeth-like spines near terminal spine and a
single spine on outer margin; several transverse spine rows
near base; mid and basal segments with several short simple
setae. Exopod 2.07 (1.63-2.85) mm long, narrow, tip over-
reaching distal end of basal endopod segment.
Thoracic appendages. Pereiopod 1 (fig. 19b) short, 5.85
(3.56—5.85 mm), 1.03 (0.73-1.03) times carapace length. Chelae
short and broad (fig. 19b), 1.6 (1.09—1.67) mm long, 2.59 (2.48—
3.84) times as long as wide, 2.0 (1.82—2.48) times longer than
dactylus; palm length 1.55 (1.55-2.25) longer than palm width
and 1.22 (0.82-1.22) times dactylus length. Carpus short, 2.88
(2.10—2.88) times longer than greatest width. Segment ratios
0.61 (0.55-0.68) : 1.22 (1.14—1.49) : 1.00 (1.08 [0.87—1.40] mm)
: 1.60 (1.11-1.60) : 0.64 (0.30-0.64) : — (2.26-2.60). Exopod
extending to base-apex of carpus.
Pereiopod 2 longer than pereiopod 1, 7.23 (5.41—7.83) mm
long, 1.28 (1.11-1.47) times carapace length. Chelae long and
slender (fig. 19c), 1.60 (1.20-1.80) mm long, half to two-thirds
length of carpus, 3.47 (3.43—4.33) times as long as wide, palm
length 2.04 (1.5-2.50) times palm width and 0.87 (0.86-1.29)
times dactylus length. Propodus 1.48 (1.39—1.91) times longer
than dactylus. Carpus 6.54—7.48 times as long as greatest width,
slightly broader distally, distal margin with small excavation.
Merus shorter than carpus, parallel-sided. Ischium about half as
long as merus. Segment ratios 0.39 (0.29—0.43) : 0.58 (0.56—0.70)
: 1.00 (1.80-2.83) mm : 0.77 (0.66-0.82) : 0.29 (0.36-0.56) : —
(1.06-1.33). Exopod extending to mid of merus to base of carpus.
Pereiopod 3 (fig. 19d, e) distinctly longer than pereiopod 2
and more slender 10.11 (7.12-10.21) mm long, 1.78 (1.38—1.78)
times carapace length. Dactylus with prominent terminal claw
and 10 (7—11) strong spines on medial margin (fig. 19e).
Propodus length 4.28 (3.82—4.69) times longer than dactylus,
length, 19.52 (12.10—19.52) times longer than wide with
11(10—18) spines on inner margin. Merus with | strong spine
on medial margin and | near ventral distal margin; segment
ratios 0.42 (0.37—0.49) : 1.82 (1.61—1.88) : 1.00 (1.91 [1.27—
2.00] mm) : 1.91 (1.74—2.23) : 0.64 (0.45—0.64) : — (1.54—1.75).
Exopod extends to mid to apex of merus.
Pereiopod 4 (fig. 19f, g) similar length to pereiopod 3, —
(6.67-9.63) mm long, — (1.45-1.62) times carapace length.
Dactylus with prominent terminal claw and — (8—11) spines on
medial margin (fig. 19g). Propodus — (3.58—5.00) times longer
than dactylus; length — (12.37-16.00) times longer than wide,
with — (11—16) spines on inner margin; merus with 1—2 strong
spine on medial margin and | near ventral distal margin.
Segment ratios — (0.22—0.52) : — (1.56-2.10) : 1.00 ( - [1.17—
1.92] mm): — (1.77-2.23) : — (0.49-0.71) : — (1.25-1.77).
Exopod extends to mid merus.
Pereiopod 5 (fig. 20a, b) similar length to pereiopod 4, 9.16
(6.57—9.73) mm long, 1.62 (1.24-1.75) times carapace length.
Dactylus with prominent terminal claw and very regular,
comb-like row of numerous 72 (72-94) small spines on medial
margin (fig. 20b). Propodus 4.09 (2.78—4.22) times longer than
92
dactylus, length 14.08 (10.42-18.93) times longer than wide
with 14 (10-14) long medial teeth, 2 distally and external
margin without teeth. Carpus without any large spines near
distal margin. Merus with | strong medial spine and | distal
spine; segment ratios 0.45 (0.45—0.71) : 1.85 (1.84—2.41) : 1.00
(1.88 [1.08—1.88] mm) : 1.64 (1.56-1.79) : — (0.52-0.89) : 1.25
(1.07—1.36). Exopod extends to basal to mid third of merus.
Abdomen. Pleopods peduncle of first pleopod short, 1.64
mm, 0.28 (0.30-0.38) times length of carapace, 1.82 (1.82—
3.08) times width, exopod 1.83 (1.06-1.83) times peduncle
length, endopod 0.97 (0.42—0.97) times peduncle length (fig.
20e); second pleopod peduncle short, — (0.33-0.58) times
length of carapace, — (2.50—3.88) times width, exopod —
(0.91—1.35) times peduncle length, endopod slightly shorter —
(0.53—1.20) times peduncle length. Length of first peduncle —
(1.07—1.74) times length of second peduncle.
Telson (fig. 20c, d) length 3.67 (3.20-4.40) mm, 0.65
(0.58—0.83) times carapace length, 3.00 (2.83—3.66) times
longer than greatest width, tapering distally. Dorsal surface
with 2 pairs of strong submarginal teeth-like spines. Posterior
margin convex with | pair of teeth-like spines outermost, 4
(4—10) long strong setose spines (fig. 20d).
Uropods approximately equal to telson length.
Male smaller than females, carapace length 4.9—6.0 mm;
endopod of first pleopod strongly excavated apically with 5-12
external spines and 11-15 long setae on inner margin (fig. 20f, g)
Etymology: Named in honour of the late Dr Mary E. White
(AM), an Australian paleobotanist and whose environmental
publications are inspirational and whose generosity to us while
staying at The Falls Forest Retreat (New South Wales; type
locality) will be remembered always.
Comments: P. whitemae may be confused by other widespread
species and is found in the same areas as P. australiensis, P.
arrostra and P. tasmaniensis. It can be distinguished from all
other long rostrum species by the carpus of pereiopod 1 which
is long with a short, robust chelae; the rostrum 1s concave with
ventral rostral spines extending from posterior to the greatest
width, extending over a length of 1.30—2.80 mm (Table 2).
Paratya whitemae 1s a widespread species in the coastal
streams of Victoria, New South Wales and 1n south-eastern
Queensland and in the Murray R in the Murray-Darling
Basin (fig. 32b) and may co-exist with P. australiensis, P.
arrostra, P. williamsi, P. rouxi and P. tasmaniensis at various
locations throughout its range.
Paratya strathbogiensis n. sp.
Figures 21—23
http://zoobank.org/urn:lsid:zoobank.org:act:6B825 EFF-D5BB-
407E-9CA9-CAA2BB6F92A0
Lineage 7 (Cook et al., 2006)
Type Material: Holotype Victoria. King Parrot Ck at Flowerdale,
—37.2953 S, 145.2905 E, 28 September 2011 (PS, JM, MC). Body in
ethanol and antennae, mouthparts, pereiopods and abdominal structures
dissected, mounted on 2 slides. Accession Ref. MC49. Museum of
Victoria Ref No NMV J75163. Genbank Registration OL420843.
P.J. Suter, J.H. Mynott & M. Crump
Paratypes: Victoria. 1 male and 3 females, King Parrot Ck at
Flowerdale, —37.2953 S, 145.2905 E, 28 September 2011 Accession
Ref. MC40, 43, 46, 47 (PS, JM, MC) NMV J75164—J75167, Genbank
Registration OL420836, OL420839, OL420846, OL420847; 2 males
King Parrot Ck above Goulburn R confluence, —37.0075 S, 145.3212 E,
28 September 2011 Accession Ref. MC53—54 (PS, JM, MC) NMV
J75168—J75169, Genbank Registration OL420845—OL 4208460; bodies
in ethanol and other structures dissected, mounted on 2 slides each.
Material Examined: Victoria: King Parrot Ck at Flowerdale,
—37.2953 S, 145.2905 E, 28 September 2011 (PS, JM, MC); King
Parrot Ck above Goulburn R confluence, —37.0075 S, 145.3212 E, 28
September 2011 (PS, JM, MC).
Diagnosis: P. strathbogiensis differs from all other species by
the following combination of characters: rostrum long, extending
beyond both antennular peduncle and scaphocerite, dorsal edge
straight, dorsally armed with 21—24 teeth, 1—2 postorbital spines,
ventrally with 4-6 large serrations over a length of 1.30-1.80
mm, all forward of greatest depth; distal half of ventral edge
straight; left mandible with 4 large teeth separated by a ridged
straight ridged notch from a blunt apical tooth; right mandible
with 4 teeth in 2 separate incisor processes with first, third and
fourth teeth large; scaphognathite of maxilla 2 truncated apically
and square at distal margin not extending to apex of upper endite;
exopod of maxilliped 2 1.88—2.34 times longer than endopod,
epipodite with long podobranchs extending to basal third of third
segment of endopodite; maxilliped 3 with medial distal margin
of apical segment of endopod with 8—9 broad teeth-like spines,
outer margin with 2 broad teeth-like spines, exopod long and
narrow, tip over-reaching distal end of basal endopod segment;
pereiopod 1 with short carpus, chelae short-long and broad,
exopod extending to base-mid carpus; pereiopod 2 with exopod
extending to mid merus; dactylus of pereiopod 3 with prominent
terminal claw and 8-13 strong spines on medial margin, exopod
extends to mid merus; dactylus of pereiopod 4 prominent
terminal claw and 9-12 spines on medial margin, exopod extends
to apical third of merus; dactylus of pereiopod 5 with prominent
terminal claw and very regular comb-like row of 64—80 small
spines on medial margin, exopod extends to mid merus.
Carapace length 6.3 (5.6—6.5) mm.
Rostrum long 5.90 (5.30—6.00) mm, extending beyond the
antennular peduncle and scaphocerite (fig. 21a), rostral length is
0.94 (0.82—1 00) times length of carapace, shape long and slender
with straight dorsal edge, pointed (fig. 21a); rostrum 8.43 (7.50—
9.33) times longer than wide; dorsally armed with 21 (21—24)
spines, ratio of number of dorsal spines to length is 3.56 (3.56—
4.29) with 2 (1—2) postorbital eye spines; ventrally with 6 (4—6)
large serrations over a length of 1.80 (1.30-1.80) mm, all spines
anterior of greatest width; distal half of ventral edge straight,
ratio of ventral spine length to rostral length is 0.31 (0.23-0.31),
with 3.50 (3.50-6.00) more dorsal spines than ventral spines;
rostral length 1.48 (1.19-1.51) times length of scaphocerite.
Antenna | (fig. 21b) peduncle 4.48 (3.80—4.68) mm long,
not reaching distal tip of scaphocerite, length 1.12 (0.83—1.12)
times scaphocerite length. Stylocerite 2.20 (2.04-2.24) mm
long, length 9.17 (6.88—9.33) times longer than width, 0.35
(0.33-0.36) times carapace length, reaching beyond distal
border of peduncle segment extending to middle of process on
distal angle of first segment (fig. 21b).
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 93
co
LLIL——
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——
SE *
SE
nn "
=
=
—
Figure 21. Paratya strathbogiensis sp. nov.: a, head region and rostrum; b, antenna | peduncle and stylocerite; c, scaphocerite; d, left mandible; e,
enlarged incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; 1, maxilla 2; j, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
94 P.J. Suter, J.H. Mynott & M. Crump
Figure 22. Paratya strathbogiensis sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3; f, pereiopod 4; g, dactylus
4; h, pereiopod 5; 1, dactylus 5. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 95
d
Figure 23. Paratya strathbogiensis sp. nov.: a, telson; b, telson terminal spines; c, pleopod 1 of female; d, pleopod 1 of male; e, endopod 1 of male.
Scale lines 0.2 mm.
96
Antenna 2 (fig. 21c) second segment of peduncle 1.40
(1.20-1.52) mm long, 0.35 (0.25-0.38) times length of
scaphocerite, 2.92 (2.53-3.18) times longer than wide.
Scaphocerite 4.00 (3.70-4.80) mm long, 0.63 (0.63-0.75)
times carapace length, 2.67 (2.47-2.87) times as long as wide.
Mouthpart. Left mandible (fig. 21d, e) with 4 large teeth
separated from a blunt apical tooth by a ridged almost straight
notch. Spine row with 7 spines, each finely setose, spine row
above molar process of approximately over 40 sparsely setose
spines. Right mandible (fig. 211, g) with 4 large teeth in 2
separate incisor processes consisting of one large apical tooth
and | small tooth and two larger separated inner teeth, Apical,
third and fourth teeth large. Spine row immediately below
teeth with 10 spines each finely setose, spine row above molar
process. Molar process ridged.
Maxilla 1 (fig. 21h) as for P. australiensis.
Maxilla 2 (fig. 211) scaphognathite truncated apically and
squared off at distal margin, not extending to apex of upper
endite (fig. 211).
Maxilliped 1 (fig. 21j) as for P. australiensis.
Maxilliped 2 (fig. 21k) endopod length 1.09 (1.01-1.16)
mm; exopod long and narrow, length 2.13 (2.00—2.40) mm,
1.95 (1.88-2.34) times longer than endopod. Epipodite with
long podobranch extending to basal third of third segment of
endopodite.
Maxilliped 3 (fig. 22a) endopod length 6.67 (6.35—7.36)
mm, 2.77 (2.52-2.80) times longer than exopod; with 3 distal
segments of similar length; basal segment curved; apical
segment with large terminal claw, medial distal margin with 7
(7—9) broad teeth-like spines, largest 3 in basal half, outer margin
with 1 long tooth-like spine near terminal spine and 1 long spine
in proximal third. Exopod 2.40 (2.27-2.67) mm long, narrow, tip
over-reaching distal end of basal endopod segment (fig. 22a).
Thoracic appendages. Pereiopod 1 (fig. 22b) short, 5.09
(3.09-5.44) mm long, 0.81 (0.48—0.84) times carapace length.
Chelae short to long and broad (fig. 22b), 1.60 (1.48-1.83) mm
long, propodus 3.00 (2.86—3.26) times as long as wide, 1.76
(1.76—2.36) times longer than dactylus, 1.19—1.31 times longer
than carpus; palm length 2.08 (1.42—2.14) times palm width
and 1.59 (1.27-1.69) times dactylus length. Carpus short, 2.88
(2.30—3.21) times longer than greatest width, broadening
distally. Segment ratios 0.74 (0.54—0.74) : 1.31 (1.19-1.31) :
1.00 (1.22 [1.13-1.47] mm) : 1.41 (1.04—1.41) : 0.43 (0.42—0.52)
: 2.28 (2.00—2.28). Exopod extending to base-mid carpus.
Pereiopod 2 (fig. 22c) longer than pereiopod 1, 7.17 (6.855—
7.89) mm long, 1.14 (1.11—1.22) times carapace length. Chelae
long and slender (fig. 22c), 1.63 (1.47-1.76) mm long, 3.59
(3.57—4.06) times as long as wide, palm length 2.09 (2.00—
2.22) times longer than palm width, 1.15 (0.91-1.15) times
length of dactylus. Propodus 1.74 (1.67-1.81) times longer than
dactylus. Carpus 6.28 (6.28-8.00) times as long as greatest
width, slightly broader distally, distal margin with small
excavation. Segment ratios 0.38 (0.35—0.42) : 0.67 (0.61—0.72) :
1.00 (2.42 [2.37-2.67] mm) : 0.84 (0.80—0.85) : 0.44 (0.35—
0.59) : — (1.08). Exopod extending to mid merus (fig. 22c).
Pereiopod 3 (fig. 22d, e) distinctly longer than pereiopod 2
and more slender 8.67 (8.47—9.40) mm long, 1.38 (1.35-1.51)
times carapace length. Dactylus with prominent terminal claw
P.J. Suter, J.H. Mynott & M. Crump
and 13 (8-13) strong spines on medial margin (fig. 22e).
Propodus 4.60 (4.50—4.93) times longer than dactylus, length
14.38 (12.77-14.86) times longer than wide with 14 (8-14)
spines on inner margin. Merus with | strong spine on medial
margin and | near ventral distal margin. Segment ratios 0.38
(0.35—0.39) : 1.77 (1.63-1.89) : 1.00 (1.73 [1.57-1.88] mm) :
1.69 (1.69-2.03) : 0.54 (0.50—0.55) : 1.46 (1.43-1.46). Exopod
extends to mid merus.
Pereiopod 4 (fig. 221, g) similar to pereiopod 3, 9.12
(8.59—9.36) mm long, 1.45 (1.37-1.45) times carapace length.
Dactylus with prominent terminal claw and 12 (9—12) spines
on medial margin (fig. 22f, g). Propodus 3.78 (3.78—4.04)
times longer than dactylus, length 11.94 (11.89-14.37) times
longer than wide, with 16 spines on medial margin; merus
with | strong spine on medial margin and 1 near ventral distal
margin. Segment ratios 0.44 (0.41—0.44) : 1.68 (1.66-1.69) :
1.00 (1.80 [1.80—1.81] mm) : 1.96 (1.73—1.96) : 0.42 (0.42—0.52)
: — (1.62). Exopod extends to mid-apex of merus.
Pereiopod 5 (fig. 22h, 1) similar length to pereiopod 4, 9.00
(7.80—9.00) mm long, 1.43 (1.20—1.43) times carapace length.
Dactylus with prominent terminal claw and very regular,
comb-like row of numerous 64 (64—80) small spines on
medial margin (fig. 221). Propodus 3.11 (3.04-3.51) times
longer than dactylus, length 12.00 (11.86—14.47) times longer
than wide with 11 (10-11) long medial teeth and external
margin without teeth. Carpus without any large spines near
distal margin. Merus with 1 strong medial spine and | distal
spine. Segment ratios 0.58 (0.49-0.63) : 1.82 (1.72-2.05) :
1.00 (1.76 [1.60—1.82] mm) : 1.69 (1.53-1.78) : 0.61 (0.54—0.61)
: 1.06 (1.06-1.50). Exopod extends to mid merus.
Abdomen. Pleopods peduncle of first pleopod short, 0.29
(0.13—0.32) times length of carapace, 2.64 (1.21—3.00) times
width, exopod 1.41 (1.31-3.35) times peduncle length,
endopod 0.78 (0.44—0.78) times peduncle length (fig. 23c);
second pleopod peduncle short, 0.37 (0.34—0.38) times length
of carapace, 2.94 (2.50—2.94) times width, exopod 1.11 (1.11—
1.51) times peduncle length, endopod slightly shorter 1.06
(1.06—1.33) times peduncle length. Length of first peduncle
1.27 (1.11—2.53) times length of second peduncle.
lelson (fig. 23a, b) length 3.80 (3.30—4.20) mm, 0.60
(0.57—0.66) times carapace length, 3.17 (2.80—3.17) times
longer than greatest width, tapering distally. Dorsal surface
with 2 pairs of strong submarginal teeth-like spines. Posterior
margin convex with | pair of teeth-like spines outermost, 10
(8—12) long, strong setose spines (fig. 23b).
Uropods slightly longer than telson length.
Male smaller than females, carapace length 4.42 mm;
endopod of first pleopod curved, not strongly excavated with
10 external spines and 16 spines on inner margin (fig. 23d, e).
Etymology: After the Strathbogie Range area in Victoria,
where this species occurs and was first recognised as a distinct
lineage by Cook (2006).
Comments: P. strathbogiensis shares the characteristics of a
long straight rostrum extending beyond the scaphocerite with
P. spinosa, P. whitemae and P. tasmaniensis. It differs from all
other species by having a long slender palm of the chelae of
pereiopod 2 (22.00 times width); rostrum with 21-24 dorsal
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 97
spines of which 1—2 are postorbital spines and 4—6 ventral
spines over a length of 1.30-1.80 mm; stylocerite extends to the
middle of the process on the apex of the antenular segment 1;
scaphognathite of maxilla 2 truncated (Table 2).
P. strathbogiensis 1s restricted to the upper Goulburn R in
the Strathbogie Ranges central Victoria and overlaps with P.
tasmaniensis (Cook, 2006) and P. arrostra.
Paratya tasmaniensis Riek, 1953
Figures 24—26
Paratya tasmaniensis Riek, 1953 (fig. 24a); type examined by MC.
Paratya australiensis Williams and Smith, 1979
Lineage 8 (Cook et al., 2006)
Lineage B (McClusky, 2007)
lype Locality: Small stream at Kingston, Tasmania, just above
the tidal zone, 10 January 1947 (Browns R).
Material Examined: Tasmania: Coal R near Campania, —42.6887 S,
147.4359 E, 11 July 2011 (PS); George R on Billabong Bay Rd near St
Helens, —41.3137 S, 148.2656 E, 9 July 2011 (PS); Elizabeth R at
Campbelltown, —41.9332 S, 147.4934 E, 13 July 2011 (PS); Browns R
at Kingston, —42.9659 S, 147.3117 E, 12 July 2011 (PS); Swamp off
Five Mile Rd, Flinders Island, 39.9053 S, 147.9746 E, 1 November
1973 (TW, PSL, BK).
Victoria: Wimmera R downstream of Dimboola Weir, —36.4557 S,
142.0167 E, 6 March 2012 (Vic EPA); Glenelg R at Ford Reserve,
—37.2472 S, 141.8458 S, 11 July 2017 (BM); Hamilton Lake boat ramp,
—37.7327 S, 142.0399 E, 2 February 2018 (PS); Glenelg R, —37.9289 S,
141.2782 E, 1 November 2017 (AC); Glenelg R near Kanagulk,
—37.1497 S, 141.8637 E, 1 November 2017 (AC).
New South Wales Stream on Gerrigong Ck Rd, Upper Kangaroo
Valley, —34.6870 S, 150.6000 E, 27 September 2017 (PS, JM. JH);
Kangaroo R, Hampden Bridge, 34.7272 S, 150.5218 E, 27 September
2017 (PS, JM. JH); Maguire Ck, —28.0837 S, 153.3364 E 26 May 2015
(BM); Hastings R off Oxley Highway, —31.4647 S, 152.6278 E, 1
November 2011 (PS, JM, MC).
South Australia. Brenda Park wetland south of Morgan,
—34.0818 S, 139.6743 E, 8 November 2011 (CM).
Diagnosis: P. tasmaniensis differs from all other species by the
following combination of characters: rostrum long, extending
beyond both antennular peduncle and scaphocerite, dorsal edge
straight or very fine curve, dorsally armed with 22-29 spines,
2—4 postorbital spines, ventrally with 4—13 large serrations over
a length of 2.10—3.40 mm, all forward of greatest depth; distal
half of ventral edge straight; left mandible with 4 teeth separated
by smooth notch then 4 short ridges at base of a less distinct
apical tooth; right mandible with 4 teeth in a single incisor
process with large third tooth; scaphognathite of maxilla 2
truncated and square apically extending to two-thirds length of
upper endite; maxilliped | with exopod flagellum distinct, well
developed and with numerous long setose spines on all margins,
over half length of caridean lobe; exopod of maxilliped 2 1.57-
2.50 times longer than endopod, epipodite with long podobranchs
extending just to base of third segment of endopodite; maxilliped
3 with medial distal margin of apical segment of endopod with
8-11 broad teeth-like spines, outer margin with 3-9 teeth-like
spines, exopod long and narrow, tip over-reaching distal end of
basal endopod segment; Pereiopod 1 with short and broad to
long and slender chelae, carpus short to long and exopod
extending to mid-apex of carpus; pereiopod 2 with exopod
extending to mid merus; dactylus of pereiopod 3 with prominent
terminal claw and 9-11 strong spines on medial margin, exopod
extends to mid merus; dactylus of pereiopod 4 prominent
terminal claw and 8-12 spines on medial margin, exopod extends
to mid merus; dactylus of pereiopod 5 with prominent terminal
claw and very regular comb-like row of 70-85 small spines on
medial margin, exopod extends to basal to mid merus.
Carapace length 5.40—7.50 mm.
Rostrum long 5.65-7.10 mm, extending beyond the
antennular peduncle and well beyond the scaphocerite (fig.
24a), rostral length 0.88—1.26 times carapace length, shape
long and slender, usually straight, pointed; rostral length
6.90-8.50 times greater than width; dorsally armed with
22-29 teeth, ratio of dorsal spines to length 1s 3.33—4.60, 2—4
postorbital spines (fig. 24a); ventrally with 4—13 large spines
over a length of 2.10-3.40 mm, at to anterior to point of
ereatest width (fig. 24a), distal half of ventral edge straight,
ratio of ventral spine length to rostral length 1s 0.33—0.48 with
2.00—6.50 more dorsal spines than ventral spines; rostral
length 1.15—1.69 times length of scaphocerite.
Antenna 1 (fig. 24b) peduncle short 4.04—5.52 mm long, not
reaching distal tip of scaphocerite, 0.78—1.31 times scaphocerite
length. Stylocerite 2.20-2.84 mm long, length 6.55-12.00
longer than wide, 0.33—0.41 times carapace length, reaching
beyond distal border of peduncle segment extending almost to
apex or just beyond distal angle process (fig. 24b).
Antenna 2 (fig. 24c) second segment 1.20-1.68 mm long,
length 1.88—2.83 times width and 0.23—0.57 times length of
scaphocerite. Scaphocerite 4.0—5.4 mm long, 2.86—4.33 times
as long as wide and 0.56—0.85 times carapace length.
Mouthparts. Left mandible (fig. 24d, e) with a blunt apical
tooth and 4 short ridges at its base, separated from incisors by
part smooth and part ridged U-shaped notch. Incisors with 2
large teeth and 2 slightly smaller robust teeth at base. Spine
row with 10 spines, each finely setose. Right mandible (fig.
24f, g) with incisors with 4 large teeth in a single incisor
process, with third tooth larger than other 3. Spine row
immediately below teeth with 9 spines each finely setose,
Spine row above molar process. Molar process ridged.
Maxilla 1 (fig. 24h) as for P. australiensis.
Maxilla 2 (fig. 241) scaphognathite truncated and square
apically extending to approximately two-thirds length of
upper endite.
Maxilliped 1 (fig. 24j) as for P. australiensis.
Maxilliped 2 (fig. 24k) endopod 0.97-1.17 mm long,
exopod long and narrow, length 1.55—2.93 mm, exopod 1.57—
2.50 longer than endopod. Epipodite with long podobranch
extending to basal third of third segment of endopodite.
Maxilliped 3 (fig. 25a) endopod 5.41-8.32 mm long,
2.34-3.12 times longer than exopod; with 3 distal segments of
similar length; basal segment curved, apical segment with
large terminal claw, medial distal margin with 8—11 broad
teeth-like spines, largest 1—2 1n basal half, outer margin with
1-2 long teeth-like spines; middle segment with 2-3 medial
spines and 0—4 spines on outer margin. Exopod long and
narrow 2.6/-3.33 mm, 0.32-0.43 times length of endopod,
extends to base of mid segment.
98 P.J. Suter, J.H. Mynott & M. Crump
Figure 24. Paratya tasmaniensis Riek: a, head region and rostrum; b, antenna | peduncle and stylocerite; c, scaphocerite; d, left mandible; e,
enlarged incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; 1, maxilla 2; J, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 99
Figure 25. Paratya tasmaniensis Riek: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus. Scale lines 0.2 mm.
100
Figure 26. Paratya tasmaniensis Riek: a, pereiopod 4; b, dactylus 4; c,
pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g,
pleopod 1 of female; h, pleopod 1 of male; 1, endopod 1 of male. Scale
lines 0.2 mm.
P.J. Suter, J.H. Mynott & M. Crump
Thoracic appendages. Pereiopod 1 (fig. 25b) short, 4.87-5.68
mm long, 0.7/4—0.84 times carapace length. Chelae short and
broad to long and slender (fig. 25b), 1.49-1.87 mm long, 2.95—
3.43 times as long as wide, |.75—2.22 times longer than dactylus,
1.25-1.41 times longer than carpus; palm length 1.60-1.83
longer than palm width, 1.33-1.67 times dactylus length. Carpus
short to long, 2.00-2.87 times as long as greatest width,
broadening distally, distal margin excavate. Segment ratios
0.45-0.78 : 1.25-1.41 : 1.00 (1.00-2.67) mm : 1.17-1.80 : 0.44—
0.53 : 2.51—2.68. Exopod extending to mid-apex of carpus.
Pereiopod 2 (fig. 25c) longer than pereiopod 1, 6.76-8.17 mm
long, 0.91—1.29 times carapace length. Chelae long and slender
(fig. 25c), 1.47-1.81 mm long, 3.06—3.93 times as long as wide,
half to two-thirds length of carpus; palm length 1.80--3.35 longer
than palm width, 0.80—1.35 times length of dactylus. Propodus
1.57—2.06 times dactylus length. Carpus long 5.79—7.22 times as
long as greatest width, slightly broader distally. Segment ratios
0.31—0.40 : 0.59-0.71 : 1.00 (2.16-2.81) mm : 0.75-1.15 : 0.34—
0.50 : 0.90—1.38. Exopod extending to mid merus.
Pereiopod 3 (fig. 25d, e) distinctly longer than pereiopod 2
and more slender, length 8.43-10.05 mm, 1.32-1.71 times
carapace length. Dactylus with prominent terminal claw and
9-1] strong spines on medial margin (fig. 25e). Propodus
3.8/7—4.81 times longer than dactylus, length 11.36-14.17
tmes longer than wide with 11—18 spines on inner margin and
3 transverse spines apically. Merus with ] strong spine on
medial margin and | near ventral distal margin. Segment
ratios 0.34—0.46 : 1.48—1.77 : 1.00 (1.66-2.03) mm : 1.63-2.06
: 0.44—0.63 : 1.40—1.76. Exopod extends to mid merus.
Pereiopod 4 (fig. 26a, b) similar to pereiopod 3, 8.63-10.47
mm long, 1.29-1.76 times carapace length. Dactylus with
prominent terminal claw and 8-12 spines on medial margin
(fig. 26b). Propodus 3.78—4.80 times longer than dactylus,
length 8.06-15.54 times longer than wide, with 14—19 spines on
medial margin, none on outer margin; merus with 1—2 strong
spines on medial margin and | near ventral distal margin.
Segment ratios 0.32—0.43 : 1.52-1.77 : 1.00 (1.66-2.13) mm :
1.73-2.08 : 0.41—0.66 : 1.55. Exopod extends to mid merus.
Pereiopod 5 (fig. 26c, d) similar length to pereiopod 4,
58.31-10.08. mm long, 1.15-1.68 times carapace length.
Dactylus with prominent terminal claw and very regular,
comb-like row of numerous (70—85) small spines on medial
margin (fig. 26d). Propodus 3.21-4.54 times longer than
dactylus, length 11.80-14.71 times longer than wide with 5-14
long medial teeth and external margin without teeth. Carpus
without any large spines near distal margin. Merus | strong
spine on medial margin and | near ventral distal margin.
Segment ratios 0.41-0.61 : 1.85—2.02 : 1.00 (1.59-1.93) mm :
1.45-1.76 : 0.53-0.70 : 0.99-1.52. Exopod extends to basal to
mid third of merus.
Abdomen. Pleopods peduncle of first pleopod short,
0.27-0.37 times length of carapace, 2.00-3.93 times width,
exopod 1.11-1.78 times peduncle length, endopod 0.55-0.88
times peduncle length (fig. 26g); second pleopod peduncle
short, 0.34—0.42 times length of carapace, 2.25-2.94 times
width, exopod 1.11-1.63 times peduncle length, endopod
slightly shorter 0.91-1.50 times peduncle length. Length of
first peduncle 0.96-1.34 times length of second peduncle.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 101
Telson (fig. 26e, f) length 3.80-4.70 mm, 0.59-0.70 times
carapace length and tapering distally, 2.83—3.79 times as long as
ereatest width. Dorsal surface with 2 pairs of strong submarginal
teeth-like spines. Posterior margin convex with | pair of teeth-
like spines outermost, 6-12 long strong setose spines (fig. 26f).
Uropods approximately equal to telson length.
Males endopod of first pleopod excavated apically with
10-14 short spines on external margin and 16-18 long spines
on inner margin (fig. 26h, 1).
Comments: P. tasmaniensis was described by Riek (1953) from
Browns R at Kingston, Tasmania. It is the only species in
Tasmania with a wide distribution on the eastern, southern and
northern streams, also on the northern west coast streams and
on Flinders Island. It does not occur in lakes of the central
plateau, except at Lakes Crescent and Sorrell at an altitude of
800 m (McClusky, 2007). P. tasmaniensis also occurs on the
mainland coastal and inland streams in South Australia,
Victoria and New South Wales (fig. 32c). It shares the
characteristics of a long straight rostrum extending beyond the
scaphocerite with P. spinosa and P. strathbogiensis and the
distinguishing characters are given in Table 2.
Walker (1973), in his study of Tasmanian Paratya,
described the morphological characteristics of P. tasmaniensis
(not including mouthparts) and described the life history from
the Coal R and a small wetland at Pawleena.
Paratya rouxi n. sp.
Figures 27-29
http://zoobank.org/urn:\sid:zoobank.org:act:8319F351-EA76-
4A F9-9C63-63FOCEBSE292
P. australiensis Roux
Lineage 9 (Cook et al., 2006)
Type Material: Holotype New South Wales. Wakool Reserve,
—35.496 S, 144.454 E, June 2011 (JC). Body in ethanol and antennae,
mouthparts, pereiopods and abdominal structures dissected, mounted
on 2 slides. AM Ref No. P.105605, Accession Ref. MC83, Genbank
Registration OL420861.
Paratypes: New South Wales. Wakool Reserve, —35.4963 S,
144.4541 E, June 2011 Accession Ref. MC672 Genbank Registration
OL420849, PSI-PS2 (JC), Bodies in ethanol and other structures
dissected, mounted on 2 slides each.
Material Examined: As for type material.
Diagnosis: P. rouxi differs from all other species by the following
combination of characters: rostrum very short, extending just
beyond the first segment of the antennular peduncle, rostrum
short and broad with a downward curve, dorsally armed with
11-19 spines, O postorbital spines, ventrally with 1—2 spines over
a length of less than 0.4 mm, all forward of greatest depth; distal
half of ventral edge straight, rostral length 0.57—0.73 times
length of scaphocerite; left mandible with 4 teeth in two groups
separated by smooth U-shaped notch from a distinct apical tooth;
right mandible with 4 teeth in two separate incisor processes;
scaphognathite of maxilla 2 truncated apically with an inner lobe
almost extending to apex of upper endite; maxilliped 1 with
exopod flagellum distinct, well developed and with numerous
long setose spines on all margins, over half length of caridean
lobe; exopod of maxilliped 2 1.9-2.7 times longer than endopod,
epipodite with long podobranchs extending to basal third of third
segment of endopodite; maxilliped 3 with medial distal margin
of apical segment of endopod with 7—10 broad teeth-like spines,
outer margin with 2 long teeth-like spines near terminal spine
and | mid outer spine, exopod long and narrow, tip over-reaching
distal end of basal endopod segment; pereiopod 1 with long
carpus and long slender chelae; pereiopod 2 with exopod
extending to mid merus; dactylus of pereiopod 3 with prominent
terminal claw and 10-12 strong spines on medial margin, exopod
extends to mid merus; dactylus of pereiopod 4 prominent
terminal claw and 8-12 spines on medial margin, exopod extends
to apical third of merus; dactylus of pereiopod 5 with prominent
terminal claw and very regular comb-like row of 70-80 small
spines on medial margin, exopod extends to mid merus.
Carapace length 4.90 (4.90—5.30) mm.
Rostrum very short (fig. 27a), 2.70 (2.10—2.75) mm long,
extending just beyond the first segment of the antennular
peduncle, 0.55 (0.49—0.55) times length of carapace, shape
convex, short and broad with downward curve, pointed; rostrum
6.75 (5.00—6.75) longer than wide; dorsally armed with 19 (11—
19) spines (fig. 27a), ratio of dorsal spine number to length 1s
7.04 (4.61—7.04) without postorbital eye spines; ventrally with 1
(1—2) spines over a length of 0.10 (0.10—0.40) mm, all anterior
to greatest width; distal half of ventral edge straight, ratio of
ventral spines length to rostral length is 0.03 (0.04—0.15) and
19.00 (6.00-19.00) more dorsal spines than ventral spines;
rostral length 0.73 (0.57-0.73) times length of scaphocerite.
Antenna | (fig. 27b) peduncle short, 3.24 (3.24-3.64) mm
long, not reaching distal tip of scaphocerite, 0.88 (0.88—0.98)
times scaphocerite length. Stylocerite 1.80 (1.80—2.03) mm long,
length 11.25 (9.00-11.25) longer than wide, 0.37 (0.34—0.38)
times carapace length, just reaching beyond distal border of
peduncle segment and to base of distal angle process (fig. 24c).
Antenna 2 (fig. 27c) second segment 1.16 (1.00—1.30) mm
long, 0.31 (0.27—0.34) times length of scaphocerite and 2.42
(2.42—2.50) times width. Scaphocerite 3.70 (3.65—3.80) mm
long, 0.76 (0.70—0.76) times carapace length and 3.36 (2.71—
3.36) times as long as wide.
Mouthparts. Left mandible (fig. 27d, e) with incisors of 4
teeth (2 large teeth and 2 robust teeth (one small and one large) at
base), separated from a large acute apical tooth by a smooth
U-shaped notch. Spine row with 9 spines, each finely setose,
spine row above molar process of approximately over 40
sparsely setose spines. Right mandible (fig. 27f, g) with 4 teeth in
2 separate incisor processes with apical and second teeth large.
Spine row immediately below teeth with 12 spines each finely
setose, spine row above molar process. Molar process ridged.
Maxilla 1 (fig. 27h) as for P. australiensis.
Maxilla 2 (fig. 271) scaphognathite apex truncated with an
inner lobe, almost extending to apex of endite.
Maxilliped 1 (fig. 27]) as for P. australiensis.
Maxilliped 2 (fig. 27k) endopod 0.80 (0.76-0.54) mm
long; exopod length 2.07 (1.60—2.07) mm, long and narrow,
exopod 2.58 (1.90-2.71) longer than endopod. Epipodite with
long podobranch extending to basal third of third segment of
endopodite.
102 P.J. Suter, J.H. Mynott & M. Crump
Figure 27. Paratya rouxi sp. nov.: a, head region and rostrum; b, antenna | peduncleand stylocerite; c, scaphocerite; d, left mandible; e, enlarged
incisors; f, right mandible; g, enlarged incisors; h, maxilla 1; 1, maxilla 2; jJ, maxilliped 1; k, maxilliped 2. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters - linking morphological characters with molecular lineages. 103
Figure 28. Paratya rouxi sp. nov.: a, maxilliped 3; b, pereiopod 1; c, pereiopod 2; d, pereiopod 3; e, dactylus 3. Scale lines 0.2 mm.
104 P.J. Suter, J.H. Mynott & M. Crump
Figure 29. Paratya rouxi sp. nov.: a, pereiopod 4; b, dactylus 4; c, pereiopod 5; d, dactylus 5; e, telson; f, telson terminal spines; g, pleopod 1 of
female; h, pleopod 1 of male; 1, endopod 1 of male. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 105
Maxilliped 3 (fig. 28a) endopod 5.36 (5.36—5.95) mm long,
2.51 (2.07-2.51) times longer than exopod; basal segment
curved, apical segment with large terminal claw, medial distal
margin with 8 (8-10) broad teeth-like spines, largest 3 in mid
half and | long spine in basal third, outer margin with 1 long
tooth-like spines near terminal spine and | mid outer spine.
Exopod 2.13 (2.13-2.88) mm long, narrow, tip over-reaching
distal end of basal endopod segment.
Thoracic appendages. Pereiopod 1 (fig. 28b) short, 3.92
(3.92-4.25) mm long, 0.80 (0.74—0.85) times carapace length.
Chelae short and slender (fig. 28b), 1.33 (1.25-1.35) mm long,
propodus 3.85 (3.10-3.85) times as long as wide, propodus
length 2.17 (1.92-2.17) times longer than dactylus; palm length
2.11 (1.90-2.11) times palm width, 1.36 (1.06-1.36) times
dactylus length. Carpus long, 2.33 (2.33-3.00) times longer
than greatest width. Segment ratios 0.66 (0.54—0.66) : 1.43
(1.16—1.43) : 1.00 (0.93 [0.93-1.13] mm) : 1.33 (1.15-1.33) :
0.44 (0.40—0.45) : — . Exopod extending to mid-apex of carpus.
Pereiopod 2 (fig. 28c) longer than pereiopod 1, 6.00
(6.00—6.90) mm long, 1.22 (1.19—1.38) times carapace length.
Chelae long and slender (fig. 28c), 1.33 (1.28—1.33) mm long,
half to two-thirds length of carpus, 4.00 (3.63—4.00) times as
long as wide, palm length 2.38 (2.22-2.38) times palm width,
1.27 (1.25-1.27) times longer than dactylus. Propodus length
1.92 (1.85-2.04) times longer than dactylus. Carpus 6.96
(6.96-9.27) times as long as greatest width, slightly broader
distally, distal margin with small excavation. Segment ratios
0.33 (0.27—0.33) : 0.63 (0.54—0.63) : 1.00 (2.13 [2.13-2.40]
mm) : 0.79 (0.77—0.79) : 0.39 (0.38-0.39) : 0.94 (0.94—1.10).
Exopod extending to mid-apical third of merus.
Pereiopod 3 (fig. 28d, e) distinctly longer than pereiopod 2
and more slender 6.96 (6.48—7.20) mm long, 1.42 (1.22-1.44)
times carapace length. Dactylus with prominent terminal claw
and 10 (10-12) strong spines on medial margin (fig. 25e).
Propodus 3.40 (3.40—4.13) times longer than dactylus, length
12.31 (12.31-12.47) times longer than wide with 7 (7—10)
spines on inner margin. Merus with 2 (1—2) strong spine on
medial margin and | near ventral distal margin. Segment
ratios 0.48 (0.47—0.54) : 1.65 (1.65—2.00) : 1.00 (1.29 [1.13—
1.35] mm) : 2.19 (1.88—2.22) : 0.54 (0.54—0.78) : — (1.75—2.04).
Exopod extends to mid merus.
Pereiopod 4 (fig. 29a, b) similar to pereiopod 3, 6.83
(6.05—6.83) mm long, 1.39 (1.14—1.39) times carapace length.
Dactylus with prominent terminal claw and 8 (8—12) spines on
medial margin (fig. 29b). Propodus 3.36 (3.03-3.58) times
longer than dactylus, length 14.00 (12.57-14.00) times longer
than wide, with 11 (8—11) spines on medial margin and |
apically on outer margin; merus with 2 strong spine on medial
margin and | near ventral distal margin. Segment ratios 0.52
(0.52-0.63) : 1.75 (1.75-2.04) : 1.00 (1.28 [1.07-1.28] mm) :
2.06 (2.00—2.06) : 0.52 (0.52-0.58) : 1.85 (1.85-2.00). Exopod
extends to apical third of merus.
Pereiopod 5 (fig. 29c, d) similar length to pereiopod 4, 7.08
(6.48—7.08) mm long, 1.44 (1.22-1.44) times carapace length.
Dactylus with prominent terminal claw and very regular, comb-
like row of numerous 72 (70—80) small spines on medial margin
(fig. 29d). Propodus 2.90 (2.59-3.00) times longer than dactylus,
length 13.13 (12.94-13.13) times longer than wide, with 8 (8-9)
long medial teeth and external margin without teeth. Carpus
without any large spines near distal margin. Merus with | strong
medial spine and | distal spine; ischium one-third length of
propodus; segment ratios 0.71 (0.67—0.78) : 2.05 (1.93-2.16) :
1.00 (1.28 [1.20—1.43] mm) : 1.67 (1.67—1.83) : 0.81 (0.67—0.81) :
1.61 (1.50—1.67). Exopod extends to mid third of merus.
Abdomen. Pleopods peduncle of first pleopod short, 0.41
(0.29—0.41) times length of carapace length, 2.67 (2.38—3.00)
times width, exopod 1.25 (1.25-1.42) times peduncle length,
endopod — (0.60) times peduncle length (fig. 29g); second
pleopod peduncle short, 0.45 (0.38—0.55) times length of
carapace, 2.93 (1.60—3.93) times width, exopod 1.14 (0.84-
1.23) times peduncle length, endopod slightly shorter —
(0.50-1.13) times peduncle length. Length of first peduncle
1.10 (1.10—1.83) times length of second peduncle.
Telson (fig. 29e, f) length 3.50 (3.50-3.75) mm, 0.71 (0.66—
0.71) times carapace length and tapering distally, 3.24 (2.50—
3.48) times as long as greatest width. Dorsal surface with 2
(2—3) pairs of strong submarginal teeth-like spines. Posterior
margin convex with | pair of teeth-like spines outermost, 12
(8-12) long strong setose terminal spines (fig. 29T).
Uropods approximately equal to telson length.
Male smaller than females, carapace length 3.88 mm;
endopod of pereiopod 1 not strongly excavated with 8 short
spines on outer margin, 2 spines at apex of finger-like
projection and a long spine at base of projection, inner margin
with 10 long setae (fig. 29h, 1).
Etymology: This species 1s named after Dr Jean Roux, who in
1926 clearly recorded the characteristics that are diagnostic for
P. rouxi from North Yanco in New South Wales. Roux also
recognised a second morphological form of Paratya at the
same location, similar to specimens from the Sydney area.
Roux was not prepared to describe a new species based on the
rostral characters, especially as similar morphology was
present in the area of the type for P. australiensis. The second
morphological form was probably P. arrostra.
Comments: P. rouxi can be confused with P. arrostra (AC)
because both have a short rostrum, but P. rouxi lacks post-orbital
spines, has 1—2 ventral rostral spines, telson 0.66-0./0 times
carapace length, rostral length 0.57-0.73 times scaphocerite
length, pereiopod | has a long slender carpus and chelae with
carpus length 2.33-3.00 times longer than wide, pereiopod 2
with dactylus 0.27—0.33 times carpus length and propodus 0.54—
0.63 times carpus length and propodus 3.63—4.00 times longer
than wide, whereas P. arrostra (4C) has 0—1 post-orbital spines,
4—5 ventral spines, telson 0.44—0.60 carapace length, rostral
length 0.80—0.90 times scaphocerite length, pereiopod 1 has a
short carpus and robust, broad cheliped with carpus length 1.71—
1.91 times longer than wide, pereiopod 2 with dactylus 0.45—0.53
times carpus length and propodus 0.74—0.79 times carpus length
and propodus 2.98—3.08 times longer than wide (see Table 3).
The distribution of P. rouxi, which is found in streams in
the Murray Darling Basin in New South Wales, does not
overlap the short rostrum P. arrostra (AC), which occurs ın
the Conondale Ranges south-eastern Queensland and in the St
George district in the upper Condamine R catchment,
Queensland (Calman, 1926).
106
Paratya gariwerdensis n. sp.
Figures 30, 31
http://zoobank.org/urn:|sid:zoobank.org:act:D691687B-83DC-
4E45-81 A B-26BE4F224714
Lineage D McClusky (2007)
Type Material: Holotype Victoria. Stokes R near Dartmoor
—37.8745 S, 141.3014 E, 3 February 2018 (PS). Body in ethanol and
antennae, mouthparts, pereiopods and abdominal structures dissected,
mounted on 2 slides. Accession Ref. PS41 Genbank Registration
OL420914, Museum of Victoria Ref No NMV J75170.
Paratypes: Victoria. Stokes R near Dartmoor -37.87/45 S,
141.3014 E, 3 February 2018 Accession Ref PS40, Genbank
Registration OL420913, PS46 Genbank Registration OL420918 (PS),
NMV J75171-J75172; bodies in ethanol and other structures dissected,
mounted on 2 slides each; Wannon R S of Coleraine —37.6652 S,
141.6632 E, 3 February 2018 Accession Ref. PS42—43 Genbank
Registration OL420915 (PS) NMV J75173—J75174.
Material Examined: Victoria. Stokes R near Dartmoor —37.8745 S,
141.3014 E, 3 February 2018 (PS); Wannon R south of Coleraine
—37.6652 S, 141.6632 E, 3 February 2018 (PS).
Diagnosis: Paratya gariwerdensis differs from all other
species by the following combination of characters: rostrum
long, 4.00—6.25 mm, extending beyond antennular peduncle
and just to end of scaphocerite, rostral length 1.02-1.25 times
longer than carapace, dorsal edge curved downwards to tip,
narrow and pointed; rostral length 8.40—12.50 times greater
than width; dorsally armed with 22-29 teeth, ratio of rostral
spines to rostral length is 4.21-6.67; 1-2 postorbital spines;
ventrally with 7-10 large serrations over a length of 1.30—2.35
mm, 2—3 spines posterior to greatest depth, distal half of ventral
edge straight; ratio of ventral spine length to rostral length is
0.32-0.38 and 2.60-4.00 more dorsal spines than ventral
spines; rostral length 1.27-1.35 times length of scaphocerite.
Antennular peduncle 2.85-3.63 mm long, not reaching distal
tip of scaphocerite, length 0.89-0.94 times length of
scaphocerite. Stylocerite 1.35-2.08 mm long, length 6.60—8.30
times width, 0.38—0.43 times carapace length, reaching beyond
distal border of peduncle segment but not to end of broad acute
process on distal angle of first segment. Right mandible with 4
teeth in a single incisor process with all teeth approximately
equal sized; spine row immediately below teeth with 5-8
lifting spines. Scaphognathite of maxilla 2 truncated distally.
Pereiopod 1, 3.55-4.10 mm long, 0.82—0.99 times carapace
length. Chelae short and broad, 1.10-1.30 mm long, propodus
2.44—3.07 times as long as wide, 1.91-2.19 times longer than
dactylus, 1.21—1.33 times longer than carpus; palm length 1.39—
1.67 times longer than wide and 1.08—1.20 times dactylus
length. Carpus very short, 2.19—2.50 times as long as greatest
width. Pereiopod 2 5.05-5.98 mm long, 1.20-1.40 times
carapace length. Chelae long and slender 1.15—1.30 mm long,
approximately two-thirds length of carpus, 2.61—3.47 times as
long as wide, palm length 1.22-1.73 times longer than width
and 0.54—0.96 times dactylus length. Propodus 1.68-2.04
times longer than dactylus. Carpus 6.27—7.82 times as long as
ereatest width. Pereiopod 3 dactylus with prominent terminal
claw and 6—7 strong spines on medial margin; propodus length
P.J. Suter, J.H. Mynott & M. Crump
3.277—3.84 times longer than dactylus, length 10.29-13.00 times
longer than wide with 11—13 spines on inner margin. Pereiopod
4, 5.40—7.00 mm long, 1.32-1.54 times carapace length:
dactylus with prominent terminal claw and 7-8 spines on
medial margin; propodus length 3.75-3.90 times longer than
dactylus, length 11.43—14.29 times longer than wide, with 10—
ll spines on medial margin. Pereiopod 5, 5.65—7.80 mm long,
1.38-1.57 times longer than carapace; dactylus with prominent
terminal claw and very regular, comb-like row of 44-54 small
spines on medial margin; propodus length 3.00—3.11 times
longer than dactylus, length 11.63—17.40 times longer than wide
with 8—13 long medial teeth and no spines on external margin.
Posterior margin of telson convex with 1 pair of teeth-like
spines outermost, 8—9 long strong terminal setose spines.
Carapace length 4.00 (3.60—4.10) mm.
Rostrum long, 4.40 (4.00—6.25) mm, extending beyond
antennular peduncle and just to end of scaphocerite (fig. 30a),
rostral length 1.10 (1.02-1.25) times longer than carapace,
dorsal edge curved downwards to tip, narrow and pointed;
rostral length 9:78 (8.40—12.50) times greater than width;
dorsally armed with 28 (22—29) teeth, ratio of rostral spines to
rostral length 1s 6.36 (4.21—6.67); 2 (1—2) postorbital spines
(fig. 30a); ventrally with 7 (7—10) large serrations over a length
of 1.50 (1.30-2.35) mm, 23 spines posterior to greatest depth
(fig. 30a), distal half of ventral edge straight; ratio of ventral
spine length to rostral length is 0.33 (0.33-0.38) and 4.00
(2.60—4.00) more dorsal spines than ventral spines; rostral
length 1.35 (1.27-1.35) times length of scaphocerite.
Antenna | (fig. 30b) peduncle 3.05 (2.85-3.63) mm long,
not reaching distal tip of scaphocerite, length 0.94 (0.89—0.94)
times length of scaphocerite. Stylocerite 1.70 (1.35—2.08) mm
long, length 7.56 (6.60—8.30) times width, 0.43 (0.38—0.43)
times carapace length, reaching beyond distal border of
peduncle segment but not to end of broad acute process on
distal angle of first segment (fig. 30b).
Antenna 2 (fig. 30c) second segment of peduncle 0.93
(0.88-1.13) mm long, 0.28 (0.28-0.31) length of scaphocerite
and 2.31 (2.31-3.66) longer than wide. Scaphocerite 3.25
(3.15—3.90) mm long, 2.95 (2.95-3.56) longer than wide, 0.81
(0.78—0.85) times length of carapace.
Mouthparts. Left mandible (fig. 30d, e) with 4 teeth
separated by a ridged notch from 1 less distinct apical tooth;
spine row immediately below incisor process of 5—7 rugose
spines (lifting spines); spine row above molar process of
approximatelyover 20 sparsely setose spines. Right mandible
(fig. 30f, g) with 4 teeth in a single incisor process with all
teeth approximately equal sized; spine row immediately
below teeth with 5-8 lifting spines; spine row above molar
process. Molar process ridged.
Maxilla 1 (fig. 30h) as for P. australiensis.
Maxilla 2 scaphognathite truncated distally, not extending
to apex of upper endite (fig. 301); palps small, terminal parts
narrow and with | sub—apical setose spine.
Maxilliped 1 (fig. 30]) as for P. australiensis.
Maxilliped 2 (fig. 30k) endopod 0.75 (0.73-0.93) mm long;
basal segment length short, exopod long and narrow 1.75
(1.19—1.75) mm long, exopod 2.34 (1.28-2.34) times longer
than endopod. Epipodite with podobranch.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 107
Figure 30. Paratya gariwerdensis n.sp.: a) head region and rostrum; b, antenna ] peduncle and stylocerite; c, scaphocerite; d, left mandible; e,
enlarged incisors; f, right mandible; g, enlarged incisors; h maxilla 1; 1, maxilla 2; J, maxilliped 1; k maxilliped 2; 1, maxilliped 3; m, peraeopod 1; n,
peraeopod 2. Scale lines 0.2mm.
108
Maxilliped 3 (fig. 311) endopod 4.63 (4.20—5.88) mm long,
2.37 (2.37-2.92) times longer than exopod; with 3 distal
segments of similar length; basal segment curved, apical
segment with large terminal claw, medial distal margin with 9
(6-9) broad teeth-like spines, largest 1 or 2 in basal third,
outer margin with 2 (2—4) teeth-like spines plus | apical spine.
Exopod long and narrow, 1.95 (1.55-2.25) mm long, tip
reaching basal third of mid segment.
Thoracic appendages. Pereiopod 1 (fig. 30m) 3.60 (3.55—
4.10) mm long, 0.90 (0.82—0.99) times carapace length. Chelae
short and broad (fig. 31b), 1.10 (1.10—1.30) mm long, propodus
2.44 (2.44—3.07) times as long as wide, 1.91 (1.91—2.19) times
longer than dactylus; palm length 1.39 (1.39—1.67) times longer
than wide and 1.09 (1.08—1.20) times dactylus length. Carpus
very short, 2.19 (2.19—2.50) times longer than greatest width,
broadening distally, distal margin excavate. Segment ratios
0.66 (0.60—0.66) : 1.26 (1.21—1.33) : 1.00 (0.88 [0.88-1.08]
mm) : 1.23 (1.16-1.33) : 0.63 (0.42—0.63) : 2.49 (1.58—2.49).
Exopod extending to mid carpus (apex merus—mid carpus).
P.J. Suter, J.H. Mynott & M. Crump
Pereiopod 2 (fig. 30n) 5.08 (5.05—5.98) mm long, 1.27 (1.20—
1.40) times carapace length. Chelae long and slender (fig. 31c)
1.18 (1.15-1.30) mm long, 2.61 (2.61—3.47) times longer than
wide, palm length 2.00 (1.22-1.73) times longer than width, 0.96
(0.84—0.96) times dactylus length. Propodus length 2.61 (2.61—
3.29) times width, 2.04 (1.68-2.04) times longer than dactylus.
Carpus 6.27 (6.27—7.82) times longer than greatest width,
slightly broader distally, distal margin with small excavation.
Segment ratios 0.33 (0.33-0.37) : 0.68 (0.60-0.68) : 1.00 (1.73
[1.73-2.15] mm) : 0.81 (0.74-0.81) : 0.45 (0.43-0.45) : 1.22
(1.11-1.22). Exopod extending to apex of merus.
Pereiopod 3 (fig. 3la, b) slightly longer than pereiopod 2
and more slender 5.83 (5.53-7.13) mm long, 1.46 (1.35-1.61)
times carapace length. Dactylus with prominent terminal claw
and 7 (6-/) strong spines on medial margin (fig. 3le).
Propodus length 3.27 (3.27-3.84) times longer than dactylus,
length 10.29 (10.29-13.00) times longer than wide with 12
(11-13) spines on inner margin. Merus with | strong spine on
Figure 31. Paratya gariwerdensis sp. nov.: a, pereiopod 3; b, dactylus 3; c, pereiopod 4; d, dactylus 4; e, pereiopod 5; f, dactylus 5; g, telson; h, telson
terminal spines; i, pleopod 1 of female; J, endopod of pleopod 1 of male. Scale lines 0.2 mm.
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 109
c)
Figure 32. Distribution of material examined in this study: a, P. arrostra;
b, P. whitemae; c, P. tasmaniensis. Maps created in Cartographica.
medial margin and | near ventral distal margin. Segment
ratios 0.48 (0.48—0.53) : 1.57 (1.57—1.95) : 1.00 (1.15 [1.00—
1.25] mm) : 1.96 (1.96-2.15) : 0.54 (0.54—0.75) : 1.52 (1.50—
1.92). Exopod extends to mid merus.
Pereiopod 4 (fig. 3lc d) similar to pereiopod 3, 5.68
(5.40-7.00) mm long, 1.42 (1.32—1.54) times carapace length.
Dactylus with prominent terminal claw and 8 (7—8) spines on
medial margin (fig. 31g). Propodus length 3.81 (3.75—3.90)
times longer than dactylus, length 11.43 (11.43—14.29) times
longer than wide, with 11 (10-11) spines on medial margin;
merus with 1 strong spine on medial margin and 1 near
ventral distal margin. Segment ratios 0.53 (0.50—0.53) : 2.00
(1.88—2.00) : 1.00 (1.00-1.25) mm : 2.05 (1.95-2.05) : 0.63
(0.58—0.63) : 1.80 (1.60-1.80). Exopod extends to mid third
of merus.
Pereiopod 5 (fig. 3le f) slightly longer than pereiopods 4,
5.78 (5.65—7.80) mm long, 1.44 (1.38—1.57) times longer than
carapace. Dactylus with prominent terminal claw and very
regular, comb-like row of 44 (44-54) small spines on medial
margin (fig. 311). Propodus length 3.00 (3.00—3.11) times longer
than dactylus, length 11.63 (11.63-17.40) times longer than wide
with 11 (8-13) long medial teeth and no spines on external
margin. Carpus with 1 large spine near distal margin. Merus
with | strong medial spine and | distal spine Segment ratios
0.84 (0.67—0.84) : 2.51 (2.10—2.51) : 1.00 (0.93 [0.93-1.30] mm) :
2.00 (1.77—2.00) : 0.73 (0.65—0.88) : 1.68 (1.35-1.68). Exopod
extends to basal third to mid third of merus.
Abdomen. Pleopods peduncle of first pleopod short 0.28
(0.28—0.35) times length of carapace length, 2.20 (2.20-3.36)
times width, exopod 1.52 (1.27-1.52) times peduncle length,
endopod 0.64 (0.64—0.80) times peduncle length. (fig. 311);
second pleopod peduncle short, 0.34 (0.34—0.40) times length of
carapace, 2.62 (2.62—4.13) times width, exopod 1.27 (1.21—1.36)
times peduncle length, endopod slightly shorter 1.16 (1.08—1.32)
times peduncle length. Length of first peduncle 1.25 (1.00—1.33)
times length of second peduncle.
Telson (fig. 31g h) length 2.80 (2.63-2.80) mm, 0.70 (0.53—
0.70) times carapace length, 3.03 (2.76-3.06) times as long as
ereatest width, and tapering distally. Dorsal surface with 2 pairs
of strong submarginal teeth-like spines. Posterior margin convex
with | pair of teeth-like spines outermost, 9 (6—9) long strong
terminal setose spines (fig. 31h).
Uropods slightly longer than telson.
Males smaller than females, carapace length 4.34 mm;
endopod of first pleopod strongly excavated apically with 8-10
external spines and 19 long setae on inner margin (fig. 31j).
Etymology: Gariwerd 1s the aboriginal name for the Grampians
Mountains, the Grampians (Gariwerd) National Park in south-
western Victoria.
Comments: Paratya gariwerdensis 1s restricted to the south-
western Victoria in streams that drain the Grampians Mountains
where it has been recorded with P. arrostra and P. tasmaniensis.
The long concave rostrum extending beyond the antennular
peduncle is a character shared with P. walkeri, P. spinosa, P.
arrostra, P. williamsi and P. tasmaniensis. Three of these
species (P. walkeri, P. spinosa and P. williamsi) have not been
recorded in Victoria.
110
Paratya gariwerdensis can be distinguished from the
species with a long rostrum by a combination of characters in
Table 2 including: ventral spines cover length of 1.30—2.35 mm;
stylocerite extending to mid process on apex of basal segment
of antennule 1; carapace length of 3.60—4.10 mm; dactylus 3
with 6—7 medial spines; dactylus 4 with 7-8 medial spines;
pereiopod 1 length 0.82—0.99 times carapace length; pereiopod
4 propodus length 1.88—2.00 times longer than carpus 4 length;
dactylus 5 length 0.67—0.84 times longer than carpus 5 length;
propodus 5 length 2.10—2.51 times longer than carpus 5 length;
pereiopod 1 length 3.55—4.10 mm; pereiopod 2 length 5.05—5.98
mm; pereiopod 3-5 less than 7.80 mm; scaphognathite of
maxilla 2 truncated; right mandible with a single group of
incisor teeth, all 4 of similar size
Conclusion
]en species have been recognised in this study because of the
linkage between the molecular and | morphological
characteristics. The aims were all addressed with the genetic
lineages all described with morphological characteristics. It
must be added that, without the benefits of the molecular data,
the conclusion by Williams and Smith (1979) of a single
Species is credible, but Riek’s (1953) revision, although with
some inadequacies as outlined by Williams (1977), was a
serious attempt to address the morphological variability
observed in this genus.
Key to female Paratya from Australia
The following key is based on mature females and caution is
required if identifying 1mmature specimens. We have found
that mature males can be identified with this key. Many of the
species will key out in several couplets because we have tried
to incorporate the variation present in each species of
Paratya.
l Rostrum short, not extending beyond antennal peduncle
(gel 2D ZN. . nennen 2
l’ Rostrum long, extending beyond antennal peduncle figs
ab Erba Dd. PAL i us da ESN EA SE HER IE E ET PEE E ER 4
2(1) No post — orbital spines (fig. 27a), rostrum with 1—3
ventral spines over a length of less than 0.4 mm, all
anterior to widest point; right mandible with paired
incisors, each with two large teeth (fig. 27g); stylocerite
extending to basal third of peduncle process (fig. 27b);
scaphognathite of maxilla 2 truncated shorter than endite
South Wales]
De Post-orbital spines present (fig. 12b, c), rostrum with
rostrum with 3—7 ventral spines over a length of 0.4—1.1
mm; right mandible with single incisor of 4—5 large teeth
(fig. 12]); stylocerite extending almost to end of peduncle
process (fig. 12d); scaphognathite of maxilla 2 rounded
apical almost to apex of endite (fig. 12k) 3
3(2)
3"
5’
6(5)
6’
7(6)
10(9)
P.J. Suter, J.H. Mynott & M. Crump
2-3 post orbital spines all contiguous with rostral spines,
3—7 ventral spines over a length of 0.4-1.5mm, all
anterior of widest point (fig. 12b)
P. arrostra (lineage 4B short rostrum)
| post-orbital spine separated posteriorly from other
rostral spines (fig. 12c), 4-5 ventral spines over a length
of 0.6-1.2 mm, with some spines posterior to greatest
width P. arrostra (lineage 4C very short rostrum)
Rostrum long extending to end or beyond scaphocerite
(figs 5a, 6a, 9a, 12a, 18a, 21a, 24a, 30a) o
Rostrum long but not extending beyond scaphocerite
(figs 5b, 12a, 15a) 5
Post-orbital spines absent (fig. 5f); 16-17 dorsal rostral
spines u... P. australiensis (in part) [New South Wales,
Shoalhaven R catchment]
Post-orbital spines present (1-3) (figs 12a, 15a); greater
than 21 dorsal rostral spines 6
*90909248699648300908349608599686000909200€9€0€0000600909060€600980202029000686048002294€00€6608086020224€069299289€
small species, carapace length less than 4.1mm; 2-3
ventral rostral spines posterior to greatest width of
rostrum (fig. 50a); south-western Victoria...
LEE P. gariwerdensis n.sp. [South-western Victoria in
streams draining the Grampians Mts]
Larger species with carapace length greater than 4.8 mm;
maximum of 2 ventral rostral spines posterior to greatest
width of rostrum (fig. 12a), or all spines anterior greatest
width of rostrum (figs 15a, 18a) 7
First pereiopod chelae shape short and slender (fig. 13b);
telson with usually 9-10 long terminal spines (fig. 141)
but may be as few as 5 or as many as 12 spines; dactylus
3 with 8—11 teeth; dactylus 4 with 8-12 teeth |...
P. arrostra (long rostrum)
First pereiopod chelae shape short and broad (figs 16b,
19b); telson with usually 6-8 long terminal spines (fig.
17f); dactylus 3 with 5-8 teeth; dactylus 4 with 5-6
teeth P. williamsi n.sp. [New South Wales, Upper
Kangaroo Valley in the Shoalhaven catchment]
Ventral spines of rostrum all anterior to widest point ..... 9
Ventral spines of rostrum 1-2 posterior to widest point
Rostrum shorter than carapace; ventral rostral spines
extend over a length less than 2.0 mm 10
Rostrum longer than carapace; ventral rostral spines
extend over a length greater than 2.0 mm ....................
. P. tasmaniensis | Tasmania and coastal streams in south-
east Australia]
Ventral rostral spines extend over a length of 0.6-1.5 mm
(fig. 5a); telson with 6 long terminal spines (fig. 4c in
Williams and Smith, 1979); carpus of pereiopod long
(1.47-2.84 mm) (fig. 5d, i) ann
RTT P. australiensis 1n part [New South Wales coastal,
Sydney area south of Newcastle]
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 111
10° Ventral rostral spines extend over a length of 1.3—1.5 mm
(hg. 21a); telson with 6-10 long terminal spines (fig.
23b) P. strathbogiensis n.sp. | Victoria, Strathbogie
Ranges in the upper Goulburn R catchment]
11(8) Rostrum with greater than 14 ventral spines (fig. 6a)........
— P. walkeri n.sp. [New South Wales coastal, Bellinger
and Tweed catchments |
ll’ | Rostrum with «14 ventral spines .......... 12
12(11) First pereiopod chelae shape short and slender (fig. 13b);
telson with usually 9—10 long terminal spines (fig. 141)
but may be as few as 5 or as many as 12 spines ca.
P. arrostra (long rostrum)
12° First pereiopod chelae shape short and broad (figs 16b,
19b); telson with usually 4—10 long terminal spines (figs
| 1f, 23d) 13
13(12) Rostrum length longer than carapace length; Dactylus 4
with 8—11 teeth; dactylus 5 with comb of 72—94 spines;
propodus 2 length 3.0—3.2 times width; 0.53—0.77 length
TIREE N MUS Zr. em mmm en m el en
ooper P. whitemae n.sp. [Widespread mainland Australa,
Victoria coastal and inland Murray—Darling Basin, New
South Wales coastal and inland Murray—Darling Basin,
Queensland coastal, South Australia inland MDB]
13’ Rostrum length shorter than carapace length; dactylus 4
with 7-8 teeth (fig. 11b); dactylus 5 with comb of less
than 60 spines; propodus 2 length 3.4—4.3 times width;
0.86—1.5 times length of dactylus 2 s.
P. spinosa n.sp. [New South Wales, northern
coastal rivers (Tweed R catchment)]
Acknowledgements
We wish to thank all the collectors who have provided
material for this study (see methods), especially Jamie Devine
from Sydney Water, Ben Cook, Tim Page and Ben Mos. We
would also like to thank those who reviewed this manuscript
and provided constructive comments. Although there were
few samples from National Parks, they were collected under
permits 10005961, 10007492,10008606 (Victoria), SL100434
(New South Wales) and WITKO6190909 (Queensland).
Funding for this work was provided by La Trobe University
through honours and research support.
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New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages.
Supplementary material
113
Supplementary Table 1: GenBank sequence data used as a genetic backbone for the morphological examination of lineages. GenBank accession
number, source publication and new species determination for the sequences.
Sequence
EU25 1947 1 McClusky haplotype D9
EU251946 1 McClusky haplotype D8
EU251945 1 McClusky haplotype D7
EU251944 1 McClusky haplotype D6
EU251943 1 McClusky haplotype D5
EU251942 1 McClusky haplotype D4
EU251941 1 McClusky haplotype D3
EU251940 1 McClusky haplotype D2
EU251939 1 McClusky haplotype D12
EU251938 1 McClusky haplotype D11
EU251937 1 McClusky haplotype D10
EU251936 1 McClusky haplotype D1
EU251935 1 McClusky haplotype C7
EU251934 1 McClusky haplotype C6
EU251933 1 McClusky haplotype C5
EU251932 1 McClusky haplotype C4
EU251931 1 McClusky haplotype C3
EU251930 1 McClusky haplotype C2
EU251929 1 McClusky haplotype Cl
EU251928 1 McClusky haplotype B9
EU251927 1 McClusky haplotype B8
EU251926 1 McClusky haplotype B7
EU251925 1 McClusky haplotype B6
EU251924 1 McClusky haplotype B5
EU251923 1 McClusky haplotype B4
EU251922 1 McClusky haplotype B3
EU251921 1 McClusky haplotype B2
EU251920 1 McClusky haplotype B16
EU251919 1 McClusky haplotype B15
EU251918 1 McClusky haplotype B14
EU251917 1 McClusky haplotype B13
EU251916 1 McClusky haplotype B12
EU251915 1 McClusky haplotype B11
EU251914 1 McClusky haplotype B10
EU251913 1 McClusky haplotype B1
EU251912 1 McClusky haplotype A6
EU251911 1 McClusky haplotype A5
EU251910 1 McClusky haplotype A4
EU251909 1 McClusky haplotype A3
EU251908 1 McClusky haplotype A2
EU251907 1 McClusky haplotype Al
EF076817 1 Cook HAPY
EF076816 1 Cook HAPF
EF076815 1 Cook HAPQ
EF076814 1 Cook HAPS
Group
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky D
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Species
determination
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. gariwerdensis
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. arrostra
P. arrostra
P. arrostra
P. arrostra
Publication source
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
114
Sequence
EF076813 1 Cook HAPR
EF076812 1 Cook HAPG
EF076811 1 Cook HAPO
EF076810 1 Cook HAPZ
EF076809 1 Cook HAPE
EF076808 1 Cook HAPC
EF076807 1 Cook HAPOO
EFO76806 1 Cook HAPI
EF076805 1 Cook HAPD
EF076804 1 Cook HAPH
EF076803 1 Cook HAPL
EF076802 1 Cook HAPMM
EF076801 1 Cook HAPM
EF076800 1 Cook HAPP
EF076799 1 Cook HAPB
EF076798 1 Cook HAPA
AY641791 1 Baker haplotype 33
AY 641790 1 Baker haplotype 32
AY 641789 1 Baker haplotype 63
AY 641788 1 Baker haplotype 56
AY 641787 1 Baker haplotype 55
AY 641786 1 Baker haplotype 54
AY 641785 1 Baker haplotype 62
AY 641784 1 Baker haplotype 58
AY 641783 1 Baker haplotype 66
AY 641782 1 Baker haplotype 65
AY 641781 1 Baker haplotype 81
AY 641780 1 Baker haplotype 82
AY 641779 1 Baker haplotype 118
AY 641778 1 Baker haplotype 131
AY 641777 1 Baker haplotype 129
AY 641776 1 Baker haplotype 130
AY 641775 1 Baker haplotype 128
AY 641774 1 Baker haplotype 108
AY 641773 1 Baker haplotype 116
AY 641772 1 Baker haplotype 106
AY 641771 1 Baker haplotype 104
AY 641770 1 Baker haplotype 111
AY 641769 1 Baker haplotype 114
AY 641768 1 Baker haplotype 113
AY 641767 1 Baker haplotype 110
AY 308175 1 Baker isolate PIGCI
AY 308174 1 Baker isolate SHC33
AY 308173 1 Baker isolate BC372
AY 308172 1 Baker isolate TR51
AY308171 1 Baker isolate BC370
AY 308170 1 Baker isolate LY5
AY 308169 1 Baker isolate Cro4
AY 308168 1 Baker isolate NCK3
Group
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Cook 7
Cook 7
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Cook 9
Lineage D Baker/Cook 5
Cook 7
Lineage D Baker/Cook 5
Cook 7
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
McClusky C/Cook 4
Species
determination
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. arrostra
P. arrostra
P. arrostra
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. whitemae
P. whitemae
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. rouxi
Pwilliamsi
P. strathbogiensis
P-williamsi
P. strathbogiensis
P. tasmaniensis
P. arrostra
P. arrostra
P.J. Suter, J.H. Mynott & M. Crump
Publication source
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
McCluskey unpublished thesis 2
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 115
Sequence Group Species Publication source
determination
AY 308167 1 Baker isolate SHA3
AY 308166 1 Baker isolate TWR2
AY308165 1 Baker isolate RY6
AY 308164 1 Baker isolate LY6
AY 308163 1 Baker isolate 3Mol
AY308162 1 Baker isolate SHA4
AY308161 1 Baker isolate KN3
AY 308160 1 Baker isolate GCK5
AY 308159 1 Baker isolate NCK4
AY308158 1 Baker isolate TWRI
AY308157 1 Baker ısolate WR3
AY308156 1 Baker ısolate JCK2
AY308155 1 Baker isolate WRI
AY308154 1 Baker isolate GINI
AY308153 1 Baker ısolate MCK3
AY308152 1 Baker isolate PR1
AY308151 1 Baker isolate RY5
AY308150 1 Baker isolate RY2
AY 308149 1 Baker isolate PR2
AY 308148 1 Baker isolate BC112
AY 308147 1 Baker isolate GIN2
AY 308146 1 Baker isolate BC113
AY 308145 1 Baker isolate BC234
AY 308144 1 Baker isolate MCK1
AY 308143 1 Baker isolate Vck2
AY 308142 1 Baker isolate BC141
AY308141 1 Baker isolate RY 1
AY 308140 1 Baker isolate DIN2
AY 308139 1 Baker isolate 85-3
AY 308138 1 Baker isolate BRLR1
AY 308137 1 Baker isolate 85-12
AY 308136 1 Baker isolate TR55
AY308135 1 Baker isolate WESI
AY 308134 1 Baker isolate Vck3
AY308133 1 Baker isolate 85-13
AY 308132 1 Baker isolate BC52
AY 308131 1 Baker isolate RY3
AY 308130 1 Baker isolate 011-5
AY 308129 1 Baker isolate BRHRI
AY 308128 1 Baker isolate Yan4
AY 308127 1 Baker isolate 032-2
AY 308126 1 Baker isolate 85-6
AY 308125 1 Baker isolate KN5
AY 308124 1 Baker isolate KN1
AY 308123 1 Baker isolate ECK1
AY 308122 1 Baker isolate BR4
AY 308121 1 Baker isolate LNAI
AY 308120 1 Baker isolate SHC32
AY 308119 1 Baker isolate 013-2
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Cook 2
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
Cook 3
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Cook 2
Cook 2
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
McClusky C/Cook 4
Lineage C Baker/Cook 8/McClusky B
Cook 2
McClusky C/Cook 4
Lineage A Baker/Cook 1
Lineage C Baker/Cook 8/McClusky B
Lineage A Baker/Cook 1
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
McClusky C/Cook 4
Lineage A Baker/Cook 1
Lineage C Baker/Cook 8/McClusky B
Cook 2
Lineage B BakerMcClusky A/Cook 6
Lineage C Baker/Cook 8/McClusky B
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
Lineage A Baker/Cook 1
Cook 3
Cook 3
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
P. tasmaniensis
P. tasmaniensis
P-walkeri
P. tasmaniensis
P. arrostra
P. tasmaniensis
P. spinosa
P. tasmaniensis
P. arrostra
P. tasmaniensis
P. arrostra
P. whitemae
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P.walkeri
P-walkeri
P. arrostra
P. tasmaniensis
P. arrostra
P. tasmaniensis
P. tasmaniensis
P. arrostra
P. arrostra
P. tasmaniensis
P.walkeri
P. arrostra
P.australiensis
P. tasmaniensis
P.australiensis
P. tasmaniensis
P. arrostra
P. arrostra
P.australiensis
P. tasmaniensis
P-walkeri
P. whitemae
P. tasmaniensis
P. arrostra
P. whitemae
P.australiensis
P. spinosa
P. spinosa
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
116 P.J. Suter, J.H. Mynott & M. Crump
Sequence Group Species Publication source
AY308118 1 Baker isolate 034-3
AY308117 1 Baker isolate MURI
AY308116 1 Baker isolate KN2
AY308115 1 Baker isolate 032-3
AY308114 1 Baker isolate 013-3
AY308113 1 Baker isolate BR3
AY308112 1 Baker isolate CON4
AY308111 1 Baker isolate PAR4
AY308110 1 Baker isolate DAL2
AY308109 1 Baker isolate S91-2
AY308108 1 Baker isolate TR36
AY308107 1 Baker isolate NCK1
AY308106 1 Baker isolate BC442
AY308105 1 Baker isolate H31
AY308104 1 Baker isolate H30
AY308103 1 Baker isolate H29
AY308102 1 Baker isolate H28
AY308101 1 Baker isolate H27
AY308100 1 Baker isolate H26
AY 308099 1 Baker isolate H25
AY308098 1 Baker isolate H24
AY 308097 1 Baker isolate H23
AY 308096 1 Baker isolate H22
AY 308095 1 Baker isolate H21
AY 308094 1 Baker isolate H20
AY308093 1 Baker isolate H19
AY 308092 1 Baker isolate H18
AY308091 1 Baker isolate H17
AY308090 1 Baker isolate H16
AY 308089 1 Baker isolate H15
AY308088 1 Baker isolate H14
AY 308087 1 Baker isolate H13
AY 308086 1 Baker isolate H12
AY308085 1 Baker isolate Hll
AY 308084 1 Baker isolate H10
AY308083 1 Baker isolate H9
AY308082 1 Baker isolate H8
AY308081 1 Baker isolate H7
AY308080 1 Baker isolate H6
AY308079 1 Baker isolate H5
AY 308078 1 Baker isolate H4
AY 308077 1 Baker isolate H3
AY 308076 1 Baker isolate H2
AY 308075 1 Baker isolate H1
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Cook 3
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage C Baker/Cook 8/McClusky B
Lineage A Baker/Cook 1
Lineage B BakerMcClusky A/Cook 6
Lineage D Baker/Cook 5
Lineage C Baker/Cook 8/McClusky B
Lineage A Baker/Cook 1
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage B BakerMcClusky A/Cook 6
Lineage A Baker/Cook 1
Lineage B BakerMcClusky A/Cook 6
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage C Baker/Cook 8/McClusky B
Lineage C Baker/Cook 8/McClusky B
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage B BakerMcClusky A/Cook 6
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
Lineage A Baker/Cook 1
determination
P. whitemae
P. whitemae
P. spinosa
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. tasmaniensis
P.australiensis
P. whitemae
Pwilliamsi
P. tasmaniensis
P.australiensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P. tasmaniensis
P.australiensis
P.australiensis
P.australiensis
P. whitemae
P.australiensis
P. whitemae
P.australiensis
P.australiensis
P. tasmaniensis
P. tasmaniensis
P.australiensis
P.australiensis
P.australiensis
P.australiensis
P.australiensis
P. whitemae
P.australiensis
P.australiensis
P.australiensis
P.australiensis
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Cook et al. (2006)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
Baker et al. (2004)
AF534904 1 Hurwood Pall
AF534903 1 Hurwood Pal0
AF534902 1 Hurwood Pa9
AF534901 1 Hurwood Pa8
AF534900 1 Hurwood Pa7
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
P. whitemae
P. whitemae
P. whitemae
P. arrostra
P. arrostra
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages.
Sequence
AF534899 1 Hurwood Pa6
AF534898 1 Hurwood Pa5
AF534897 | Hurwood Pa4
AF534896 1 Hurwood Pa3
AF534895 1 Hurwood Pa2
AF534894 | Hurwood Pal
AY 622605 1 Paratya howensis
AY 661487 1 Paratya curvirostris
AY 661488 1 Paratya compressa
AY 661489 1 Paratya compressa
AY 661490 1 Paratya compressa
AY 661491 1 Paratya compressa
AY 661492 1 Paratya norfolkensis
AY 661493 1 Caridina indistincta
AY 661494 1 Caridina cf imitatrix
AY 661495 1 Paratya cf caledonica
AY 661496 1 Paratya cf caledonica
AY 6614981 Paratya cf caledonica
AY 661499 | Paratya cf intermedia
AY 661500 1 Paratya cf intermedia
AY 661501 1 Paratya cf intermedia
01420759 JWA2019
OL420760 JWA2020
OL420761 JWA2023
OL420762 JWA2026
OLA20763 JWA2027
OL420764 JWA2028
OLA20765 JWA2032
OL420766 JWA2033
OL420767 JWA2034
OL420768 JWA2035
OLA20769 JWA2036
OLA20770 JWA2113
OLA20771 JWA2116
OLA20772 JWA2118
OLA20773 JWA2119
OLA20774 JWA2123
OLA20775 JWA2125
OL420776 JWA2126
OL420777 JWA2127
OL420778 JWA2129
OLA20779 JWA2131
OLA20780 JWA2132
OLA20781 JWA2135
OL420782 JWA2136
OLA20783 JWA2137
OL420784 JWA2138
OLA20785 JWA2141
OLA20786 JWA2143
Group
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Caradina
Caradina
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Paratya outgroup
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
McClusky C/Cook 4
Lineage B BakerMcClusky A/Cook 6
Lineage B BakerMcClusky A/Cook 6
McClusky C/Cook 4
Species
determination
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. whitemae
P. whitemae
P. whitemae
P. arrostra
P. whitemae
P. whitemae
P. arrostra
P. whitemae
P. whitemae
P. whitemae
P. whitemae
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. arrostra
P. whitemae
P. arrostra
P. arrostra
P. arrostra
P. whitemae
P. whitemae
P. whitemae
P. arrostra
P. arrostra
P. whitemae
P. whitemae
P. arrostra
Publication source
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
Hurwood et al. (2003)
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
This study
117
118 P.J. Suter, J.H. Mynott & M. Crump
Sequence Group Species Publication source
determination
OL420787 JWA2144 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420788 JWA2184 McClusky C/Cook 4 P. arrostra This study
OL420789 JWA2435 McClusky C/Cook 4 P. arrostra This study
OL420790 JWA2436 McClusky C/Cook 4 P. arrostra This study
OL420791 JWA2437 McClusky C/Cook 4 P. arrostra This study
OL420792 JWA2438 McClusky C/Cook 4 P. arrostra This study
OL420793 JWA2439 McClusky C/Cook 4 P. arrostra This study
OL420794 JWA2440 McClusky C/Cook 4 P. arrostra This study
OL420795 JWA2441 McClusky C/Cook 4 P. arrostra This study
OL420796 MCI McClusky C/Cook 4 P. arrostra This study
OL420797 MC10 McClusky C/Cook 4 P. arrostra This study
OL420798 MC101 McClusky C/Cook 4 P. arrostra This study
OL420799 MC103 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420800 MC109 Lineage B BakerMcClusky A/Cook 6 P. tasmaniensis This study
OL420801 MC110 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420802 MC115 Lineage B BakerMcClusky A/Cook 6 P. tasmaniensis This study
OL420803 MC116 Lineage B BakerMcClusky A/Cook 6 P. tasmaniensis This study
OL420804 MC117 Lineage B BakerMcClusky A/Cook 6 P. tasmaniensis This study
OL420805 MC119 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420806 MC120 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420807 MC121 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420808 MC125 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420809 MC126 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420810 MC130 F Cook 7 P. strathbogiensis This study
OL420811 MC133 McClusky C/Cook 4 P. arrostra This study
OL420812 MC134 McClusky C/Cook 4 P. arrostra This study
OL420813 MC135 Cook 9 P. rouxi This study
OL420814 MC136 Cook 9 P. rouxi This study
OL420815 MC137 Cook 9 P. rouxi This study
OL4208 16 MC138 Cook 9 P. rouxi This study
OL420817 MC14 Cook 3 P. spinosa This study
OL420818 MCI7 Cook 3 P. spinosa This study
OL420819 MC2 McClusky C/Cook 4 P. arrostra This study
OL420820 MC20 McClusky C/Cook 4 P. arrostra This study
OL420821 MC21 F Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420822 MC22 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420823 MC23 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420824 MC24 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420825 MC26 Lineage A Baker/Cook 1 P. australiensis This study
OL420826 MC28 Lineage A Baker/Cook 1 P. australiensis This study
OL420827 MC29 Lineage A Baker/Cook 1 P. australiensis This study
OL420828 MC3 McClusky C/Cook 4 P. arrostra This study
OL420829 MC30 Lineage A Baker/Cook 1 P. australiensis This study
OL420830 MC31 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL42083 1 MC32 Lineage A Baker/Cook | P. australiensis This study
OL420832 MC33 Lineage A Baker/Cook 1 P. australiensis This study
OL420833 MC34 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420834 MC37 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420835 MC4 McClusky C/Cook 4 P. arrostra This study
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 119
Sequence Group Species Publication source
determination
OLA420836 MC40 Cook 7 P. strathbogiensis This study
OLA20837 MC41 McClusky C/Cook 4 P. arrostra This study
OL420838 MC42 Cook 7 P. strathbogiensis This study
OL420839 MC43 Cook 7 P. strathbogiensis This study
OLA420840 MC46 Cook 7 P. strathbogiensis This study
OL420841 MC47 Cook 7 P. strathbogiensis This study
OLA20842 MC48 Cook 7 P. strathbogiensis This study
OL420843 MC49 Cook 7 P. strathbogiensis This study
OL420844 MC52 Cook 7 P. strathbogiensis This study
OL420845 MC53 Cook 7 P. strathbogiensis This study
OL420846 MC54 Cook 7 P. strathbogiensis This study
OLA20847 MC63 McClusky C/Cook 4 P. arrostra This study
OLA420848 MC66 McClusky C/Cook 4 P. arrostra This study
OL420849 MC672 Cook 9 P. rouxi This study
OL420850 MC70 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OLA20851 MC71 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420852 MC72 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420853 MC73 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420854 MC74 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420855 MC75 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420856 MC76 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420857 MC78 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OLA420858 MC79 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420859 MC80 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OLA20860 MC82 McClusky C/Cook 4 P. arrostra This study
OL420861 MC83 Cook 9 P. rouxi This study
OL420862 MC84 Cook 9 P. rouxi This study
OLA420863 MC86 McClusky C/Cook 4 P. arrostra This study
OLA20864 MC87 McClusky C/Cook 4 P. arrostra This study
OLA420865 MC88 McClusky C/Cook 4 P. arrostra This study
OLA20866 MC89 McClusky C/Cook 4 P. arrostra This study
OL420867 MC9 McClusky C/Cook 4 P. arrostra This study
OL420868 MC90 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OLA420869 MC90 FS McClusky C/Cook 4 P. arrostra This study
OL420870 MC91 McClusky C/Cook 4 P. arrostra This study
OL420871 MC95 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420872 MC97 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420873 MC98 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420874 MC99 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OLA20875 PJ1 Lineage A Baker/Cook 1 P. australiensis This study
OLA208706 PJ2 Lineage A Baker/Cook 1 P. australiensis This study
OLA20877 PJ3 Lineage A Baker/Cook 1 P. australiensis This study
OLA20878 PJ4 Lineage A Baker/Cook 1 P. australiensis This study
OL420879 PJ5 McClusky C/Cook 4 P. arrostra This study
OLA20880 PJ6 McClusky C/Cook 4 P. arrostra This study
OLA20881 PS100 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420882 PS101 McClusky C/Cook 4 P. arrostra This study
OL420883 PS102 Lineage D Baker/Cook 5 P. williamsi This study
OL420884 PS103 Lineage D Baker/Cook 5 P. williamsi This study
120 P.J. Suter, J.H. Mynott & M. Crump
Sequence Group Species Publication source
determination
OL420885 PS104 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420886 PS105 Lineage D Baker/Cook 5 P. williamsi This study
OL420887 PS106 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420888 PS107 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420889 PS108 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420890 PS 109 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420891 PS110 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420892 PS111 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420893 PS112 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420894 PS113 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420895 PS114 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420896 PS115 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420897 PS116 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420898 PS117 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420899 PS12 Cook 2 P. walkeri This study
OL420900 PS 14 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420901 PS15 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420902 PS16 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420903 PS18 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420904 PS20 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420905 PS22 McClusky C/Cook 4 P. arrostra This study
OL420906 PS24 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420907 PS25 Lineage A Baker/Cook 1 P. australiensis This study
OL420908 PS26 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420909 PS27 Lineage A Baker/Cook 1 P. australiensis This study
OL420910 PS28 Lineage A Baker/Cook 1 P. australiensis This study
OL420911 PS29 Lineage A Baker/Cook | P. australiensis This study
OL420912 PS4 McClusky C/Cook 4 P. arrostra This study
OL420913 PS40 McClusky D P. gariwerdensis This study
OL420914 PS41 McClusky D P. gariwerdensis This study
OL420915 PS43 McClusky D P. gariwerdensis This study
OL420916 PS44 McClusky D P. gariwerdensis This study
OL420917 PS45 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420918 PS46 McClusky D P. gariwerdensis This study
OL420919 PS47 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420920 PS48 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420921 PS49 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420922 PS50 Lineage C Baker/Cook 8/McClusky B P. tasmaniensis This study
OL420923 PS51 McClusky D P. gariwerdensis This study
OLA420924 PS53 McClusky D P. gariwerdensis This study
OL420925 PS54 McClusky D P. gariwerdensis This study
OL420926 PS55 McClusky D P. gariwerdensis This study
OL420927 PS6 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420928 PS7 Lineage B BakerMcClusky A/Cook 6 P. whitemae This study
OL420929 PS8 Cook 2 P. walkeri This study
New species of Paratya (Decapoda: Atyidae) from Australian inland waters — linking morphological characters with molecular lineages. 121
AY308128 1 Rater actam HCKÀ
| " AYRES | her jicisis MCKJ
wars
e
ANY | Baden meii md
" WSOP 980 ^ Bale ee Ce)
AY300544 N Maker now MC
rer)
" ENT | Geter hie WO
& ke ADC | Pieter tosis WII
m ADAD 1 ner ete Vea?
| AY ROR Ds | Pater aois vend "
= P^
mra
E Av 300154 | Esker mote ONT | é
AY3ORIAT | Bibe wein QU
= A84]T77 | Ricker Pacictyps 179
AY84]775 | Taker fupictype 130
ATBATTTS | aber tapings 128
; ANETTE T ber age 131
AO 1 Murad Pl
AF 334555 \ mures Pal
APSMéMEBM | era Pat
ASTA | haramia Ped
AVES 1 Faber potia Yar
SIDE | Pered Pal
EFQ798!3 ! Cock MEG
= Aral Ti 1 Ramer nginya 118 F
m ww
CUZS 1900 | MoCuabg apictype C7
- MOS
PUSS 1832 1 Wucht Ca
ART 00 1 Beier fapssiype 114
Av308552 | ink nog FF N
— A4 1770 ^ Rue hemes 11*
Arie 9 Hades Peppe 113
CUZ 780" | hut hapicteps C3
AYAJO | uber haciatype 1)
81051834 1 Ar A C^
CPOE | Coon HAPE
warn“
ATIA 9 Bete ie VD!
wore F
me
AWG FT 5 Rakes Fapefype SDi
uw
wen
DEIN 1 er, Magione Ct
Near
PS vo
Piar
msn
Nw
Av"3081 40 ! Dohe chain CINC
nam!
AY308 14) " Bahr mobs PRT
wore res
EFG76iH 4 | Cock MARS
Ata 3 Ta 5 Baker Happs 1354
Nui
Peres 1 Coca PAP
AYE41T72 © Makes hagierype 196
£u | Casa HARA
Nw 2508
EF378817 | Coca Pay
woe F
Ar | Bier inte Mi]
MCINX
EUISHOS | McClusry hapietups C7
PABow! i
Na)”
MINTS
FUSE | McChaiy hapitus CS
Bear | Soom AAR
Neto
| Bar | Cook mu
— POM | Cosa WAT
Avid 1772 * Raker mapenypa TH
b
Supplementary Figure 1: Expanded sub-tree for Paratya arrostra Riek
from fig. 3 in text.
EU?St917 * McChusiey hapiatype B13
| DUIS PbS t MeCkaur, haplotype B1
Mcre
AY 308108. 3 Makar incisko K
KY308126 1 Baker motets TRE
EU731918 1 Weck hapictype Bhd
EU251015 ! MoClusi hapietype B12
KY30BOBS 1 Bohai ieolaie M12
EUZS1972 1 Mery hapiafype EJ
n AY3OBTST 1 Baloo rotae SHA)
| JV uysobta2 1 Baker nomte DM
| | AY308*60 | Baier isciate OTRS
A Y30810*3 1 Baker nolute H27
ANSOBOUS Y Baer solate H21
Wr
A | Baker motae H75
AYO 1 Baker molte H72
AY3QBQUT | Baron! estate M23
AY208129 | Baker octets BAHS
Av SORS | Baker testete H24
EU2514921 1 MeChaioy hagita EC
P10
PISS
aca
ro
Pass
Paar
Pods
LI
| BURY 1824 $ MoClusiry haplotype BS
" | AWG A6 1 Baier isolate TAI
AG 1 Balow mela TWRT
EU251825 1 Minsky Nagiotype 58
ABER j Baar istas LY
P Aachá
AY308170 1 [haber hotste Y
MC120
| AY 3080885 | Maer asiyta HTI
LIC
esin
P5113
, P8118
| PaT
P5114
EISIN 1 MoCusiry Aare B' ] ,
m V AS08120 1 Bale isolate BALA
EU244926 * Melk hapietpee BY
AY841788 1 Baier haplotype 58
e AYÉASETET 1 Passer hapietepe SA
2641784 1 Baker hapiotyse 54
= Ayi Gh T Baker Papiclypa 62
| — DOSS | Cock RAP
t. AY B41783 1 faker haplotype $8
mom
l EUT * Metus haplotype BS
FU231829 1 MoChusiny haplotype BIG
j UFU? WOO 1 Cook HAPF
[T raonta I I
EFUTSA0) 1 Coo MAP,
ADD an 1 Baker mote BCH
N13081458 | Baer melale BC112
EFUTSS08 | Cong HIPC
1 I 841786 1 Baloe haplotype &?
BUCS 1038 1 McCiusiry haplotype B9
EUQS1914 1 MeCiusiry hagietype BIG
1 EFCTOT90 ! Cock HIPS
AYv308142 | Bakar iaceo DENN
AYZENK | Baer incite BC52
— DS 1 Cock HAPOO
EFGTGBO! 1 Cook MAP
—— (0251910 | McClusky Sapictype 815
FONE 1 Cook MAPS
EFDUT;OROZ | Coca HAPAI
EFU7UBO4 1 Coot MAPH
AGATE 1 Baer hapierype 58
4 v
L— (
supplementary Figure 2: Expanded sub-tree for Paratya tasmaniensis
Riek from fig. 4 in text.
122 PJ. Suter, J.H. Mynott & M. Crump
m AO0308127 1 Bator molate O52-2
= £F254004 | Hurmeos Patt
| AY305115 1 Baker aa 052-3
AvY20811B. 3 Baer inaite 094-3
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AFS34807 ! Hurancd Pap
à AYADI 1 Baker molte CAL
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| AY306130 1 Baker kole O1 1-5
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r EUZS 1810 7 ic uni hapiotype Ad
= 1308120 1 Dakar motte HC?
" ll papas
Av S008 1 Baker mola H15
— — EU2819127 1 MecCiosky hagplatypa AB
! AYG64St TE Baker haplotype B2
py P365
| AYSOS154 1 Bater iolate JOCKS
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| AY308123 ! Dakar cite ECHT
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—
supplementary Figure 3: Expanded sub-tree for Paratya whitemae n.sp.
Memoirs of Museum Victoria 81: 123-133 (2022) Published October 2022
ISSN 1447-2554 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/j.mmv.2022.81.05
Articullichirus, a new genus of ghost shrimp (Crustacea: Axiidea: Callichiridae) with
one new species
(http://zoobank.org/urn:Isid:zoobank.org:pub:449E07EF-3824-4B86-87CA-D8433A FOD166)
GARY C.B. Poore!, PETER C. DWORSCHAK? AND KAREEN E. SCHNABEL?
! Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia [gpoore& museum vic.gov.au] (http://zoobank.org/urn:lsid:zoobank.
org:author:c004d784-e842-42b3-bfd3-3174d35918975; https://orcid.org/0000-0002-7414-183X )
? Dritte Zoologische Abteilung, Naturhistorisches Museum, Burgring 7, A-1010 Vienna, Austria [peter.dworschak@nhm-wien.ac.at|
(http://zoobank.org/urn:|sid:zoobank.org:author:4BCD9429-46A F-4B DA-BE4B-439EE6A DC657; https://orcid.org/0000-0003-4705-6426)
* Coasts and Oceans Centre, National Institute of Water & Atmospheric Research, Private Bag 14901 Kilbirnie, Wellington 6241, New
Zealand |kareen.schnabel@niwa.co.nz] (http://zoobank.org/urn:|sid:zoobank.org:author:90CD9ESD-8E26-4E08-8A5F-3263CBE9D6BD;
https://orcid.org/0000-0002-9965 9010)
Abstract
Poore, G.C.B., Dworschak, P.C., and Schnabel, K.E. 2022. Articullichirus,a new genus of ghost shrimp (Crustacea:
Axiidea: Callichiridae) with one new species. Memoirs of Museum Victoria 81: 123—133.
Articullichirus gen. nov., close to Corallianassa Manning, 1987 and Calliapagurops de Saint Laurent, 1973, 1s
diagnosed to include Callianassa articulata Rathbun, 1906 from Hawaii and French Polynesia, Callianassa collaroy
Poore and Griffin, 1979 from southern Australia and Articullichirus chiltoni sp. nov. from northern New Zealand. Previous
records of the two described species from the Indo-West Pacific are reassigned.
Keywords
Introduction
Callichiridae Manning and Felder, 1991 is one of seven families
of callianassid-like Axiidea de Saint Laurent, 1979, most
recently reviewed by Poore et al. (2019), whose classification
was based on their molecular phylogeny (Robles et al., 2020).
Corallianassa Manning, 1987, one of its 17 genera, has proved
problematic. The question of differences between
Corallianassa, Corallichirus Manning, 1992 and Glypturus
Stimpson, 1866 was settled by Komai et al. (2015). Manning
(1987) was uncertain whether to include Callianassa articulata
Rathbun, 1906 in Corallianassa; this question is investigated
here by re-examining this species and two similar others.
During this study, it was realised that Calliapagurops de Saint
Laurent, 1973, a small genus with exceptionally long eyestalks,
Shares several features with Corallianassa. Here, we erect a
new genus for Callianassa articulata and two others, one new.
Material is lodged in Museums Victoria, Melbourne,
Australia (NMV), Canterbury Museum, Christchurch, New
Zealand (CMNZ), Muséum national d'Histoire naturelle,
Paris, France (MNHN) and Naturhistorisches Museum,
Vienna, Austria (NHMW), Bernice P. Bishop Museum,
Honolulu, USA (BPBM), and National Museum of Natural
History, Washington, D.C., USA (USNM). Size is expressed
as carapace length (cl.), including rostrum, in mm. The
Crustacea, Decapoda, Axiidea, Callichiridae, Articullichirus, taxonomy, new genus, new species, Pacific Ocean, taxonomy
diagnosis of the new genus is derived from the same DELTA
database (Dallwitz, 2018) used by Poore et al. (2019).
Family Callichiridae Manning and Felder, 1991
Articullichirus gen. nov.
http://zoobank.org/urn:lsid:zoobank.org:act:EB81E6A6-9CFI1-
4B8B-BF06-3E2440B2A6BA
Type species. Callianassa articulata Rathbun, 1906 by present
designation.
Diagnosis. Anterior branchiostegal lobe sclerotised, well-
produced anteriorly beyond junction with oblique branchiostegal
ridge with which it articulates by means of a virtual condyle;
rostrum spine-like; anterolateral spines prominent. Pleomere |
tergite undivided or with weak transverse step. Antennal
scaphocerite acute. Maxilliped 3 ischium and merus together
ovoid, axial length slightly greater than width at their articulation;
crista dentata of few separate spines proximally and toothed
ridge distally overlapping proximal margin of merus; propodus
about as wide as long, propodus free distal margin nearly
transverse. Male major cheliped merus with 3 proximal sharp
oblique spines, | or more distally along length of lower margin;
without spines on upper margin of merus and propodus or lower
124
margin of carpus. Minor cheliped of male slightly more than half
wide as major, carpus slightly shorter than palm, fingers as long
as palm; merus lower margin with row of spines. Pereopod 3
propodus subpentagonal, with strong broad proximal lobe on
lower margin, lower margin straight. Pereopod 4 subchelate,
fixed finger shorter than dactylus. Male pleopod | of 2 articles.
Male pleopod 2 appendix interna absent. Pleopods 3-5
appendices internae longer than broad, clearly emerging from
margin of endopod. Uropodal endopod with convex anterior
margin, acute-rounded apex, slightly curved posterior margin,
longer than wide; exopod with elevated dorsal plate with row of
setae diverging from row of setae on distal margin. Telson wider
than long, convex-sided, widest near midpoint, posterior margin
concave between rounded posterolateral corners, with transverse
row of spiniform setae at midpoint.
Etymology. Alliteration of articulata, specific name of the
type species, and Callichirus, type genus of the family.
Included species. Callianassa articulata Rathbun, 1906;
Callianassa collaroy Poore and Griffin, 1979; Articullichirus
chiltoni sp. nov.
Remarks. Callianassa articulata Rathbun, 1906 was provisionally
included in Corallianassa by Manning (1987) and Dworschak
(1992) and without qualification by Poore et al. (2019). Poupin
(1998) argued for its inclusion in Cheramus Bate, 1888, a genus
now belonging in another family, Callianassidae Dana, 1852.
Sakai (1999, 2005, 2011) included the species in Glypturus
Stimpson, 1866. Dworschak (1992) noted how the telson, without
a median posterior prominence, and the more operculiform
maxilliped differed from other species of Corallianassa (see
figures here and in Rathbun, 1906; Edmondson, 1944).
Callianassa collaroy Poore and Griffin, 1979 has been variously
included in Glypturus (Sakai, 1988, 2011), Corallianassa (Tudge
et al., 2000; Davie, 2002; Komai et al., 2015; Poore et al., 2019)
and Neocallichirus Sakai, 1988 (Sakai, 1999). It was included in
Komai et al.’s (2015) thorough review of differences between
Glypturus and Corallianassa, but differences between C.
articulata and both these genera were not commented on. The
discovery of new material of C. articulata from French Polynesia,
of C. collaroy in Australia, and of a similar species from New
Zealand, stimulated a reappraisal of specimens from throughout
the Pacific identified as one or the other of these two species.
Several are misidentified and a new genus is warranted.
Articullichirus resembles Corallianassa and Glypturus
(but not Neocallichirus) in the possession of a pair of sharp
anterolateral carapace spines. Articullichirus differs from both
genera in the much broader oval maxilliped 3 (narrow with an
oblique meral margin in Glypturus and Corallianassa); shape
of the telson (evenly convex-sided, with concave posterior
margin in Articullichirus, with semicircular posterior half in
Glypturus, tapering from greatest width anteriorly in
Corallianassa); and an acute scaphocerite (small in Glypturus,
sometimes absent in Corallianassa). In addition,
Articullichirus, like Corallianassa, differs from Glypturus in
the absence of spines along the upper margin of the chelipeds
and in having a horizontal rostrum. Komai et al. (2015) noted
that the combined length of pleomeres | and 2 of at least four
G.C.B. Poore, P.C. Dworschak & K.E. Schnabel
species of Corallianassa 1s greater than or equal to the carapace
length, whereas in G. armatus, Corallianassa intesi (de Saint
Laurent and Le Loeuff, 1979) and Articullichirus collaroy
comb. nov. pleomeres | and 2 are shorter than the carapace.
Calliapagurops differs from all callichirids in having
exceptionally stout antennae, much more prominent than the
antennules; the antennae may be used for suspension feeding
(Dworschak and Wirtz, 2010). Calliapagurops was placed in
its own subfamily Calliapaguropinae by Sakai (1999) but its
relationship to Callichiridae was realised by Ngoc-Ho (2002).
Calliapagurops is notable for exceptionally long eyestalks
with terminal cornea, differing from most callichirids that
have more or less flattened eyestalks with subterminal cornea
and a mesiodistal lobe (Ngoc-Ho, 2002; Dworschak and Wirtz,
2010). While the eyestalks of species of Corallianassa,
Glypturus and Articullichirus are much shorter, the cornea 1s
terminal with only a small mesiodistal lobe. Calliapagurops
and these three genera all have sharp prominent anterolateral
carapace spines that, with the sharp rostrum, have more or less
weakly calcified bases. Species of Articullichirus resemble the
two recognised species of Calliapagurops. The oval telson
with a transverse row of robust setae is similar to that of
Articullichirus, quite different from those of Corallianassa
and Glypturus. The chelipeds of Calliapagurops are generally
similar to those of Articullichirus; the coxae carry a pair of
mesial hooks as in A. collaroy and A. chiltoni, but these are
absent in A. articulatus. The uropodal endopod of
Calliapagurops bears two proximal teeth on the dorsal surface
similar to those seen in most species of Articullichirus, but not
seen in the other two genera. The maxilliped of Calliapagurops
is operculate as in Articullichirus, more so than in the other
two genera. Its merus bears four distal teeth, whereas some
individuals of A. articulatus and all those of A. collaroy have a
single tooth or tubercle; A. chiltoni has a tubercle.
Articullichirus articulatus (Rathbun, 1906) comb. nov.
Figures 1,2
Callianassa articulata Rathbun, 1906: 892, fig. 47.—Manning,
1987: 396.
Callianassa (Callichirus) articulata.—De Man, 1928: 28, 94,
108.—Edmondson, 1944: 54, fig. 9.
Corallianassa articulata.—Dworschak, 1992: 210, fig. 14.—
Tudge et al., 2000: 144.—Komai et al., 2015: 54.—Poore et al., 2019:
136, 144.
Corallianassa collaroy.—Sakai, 1992: 212, fig. 1.
Glypturus articulatus.—Sakai, 2011: 433 (part).
Cheramus articulatus.—Poupin, 1998: 31.
Not Callianassa | articulata.— Chilton,
Articullichirus chiltoni sp. nov.
1911: 551-552 =
Material examined. Hawaii, Oahu, Honolulu, Harbour Entrance, coll.
V. Pietschmann, 1927, NHMW 6621 (ovigerous female, 7.5 mm).
French Polynesia, Tuamotu, Mataiva, Hoa Papino platier, coll. Mario
Monteforte, 1982, MNHN IU-2013-19994 (ZTh1232) (female, 7.3 mm).
Material not examined. Hawaii, Oahu, Kahala, coll. C.H. Edmondson,
1930-1933, BPBM S4669, S4670, S4671, 84672, 84673, S4674 (6
specimens); Kawailoa, coll. C.H. Edmondson, 1921, BPBM S4668;
Hanauma, coll. C.H. Edmondson, 1933, BPBM 84675, S4676 (2); coll.
Banner, 1938, 54677 (1); Waikiki, coll. Simon, 1941, BPBM S4678 (1).
Articullichirus
Nihoa Island, RV Albatross, 48-60 m, USNM 30532 (female,
holotype); Nihoa Island, RV Albatross, 42-48 m, USNM 30995
(female). Oahu Island, Waikiki, coll. C.H. Edmondson, 1921 USNM
78119 (1).
Size. Cl. to 7.5 mm.
Type locality. USA, Hawaii, 42—60 m.
Diagnosis. Maxilliped 3 basis without mesial tooth or with 2
small mesial teeth; merus with or without small tooth on distal
margin. Pereopod | coxa with small mesiodistal process with
terminal setae. Uropodal endopod midlength about 1.3-1.4
times as long as wide, with tapering apex, with 2 dorsal
clusters of setae, with 2 proximal dorsal teeth. Telson 1.4-1.7
times as wide as long, with sinuous posterior margin; with
transverse dorsal row of 2 pairs of spiniform setae.
Description of female from Tuamotu (MNHN IU-2013-19994).
Carapace length 0.28 of total length; cervical groove at 0.8 length
of carapace; dorsal oval well defined; hepatic region with weakly
sclerified line between dorsal oval and linea thalassinica. Rostrum
and anterolateral carapace spines with unsclerified basal region;
rostrum an anteriorly directed, acute spine as long as eyestalk;
anterolateral spines set well back from rostrum, acute, one third
125
as long as rostrum. Anterolateral branchiostegal margın
extending dorsal to linea thalassinica, with small separate
sclerified plate near beginning of linea thalassinica. Pleomere |
with weak transverse groove, with dorsolateral longitudinal setal
row; pleomere 2 about 1.4 times as long as pleomere | tergite;
pleomeres 3—5 scarcely expanded laterally, with dense setose
areas; pleomere 6 1.2 times as long as pleomere 5, pleomere 6
with posterolateral notch, with pair of lobes on posterior margin.
Eyestalks shorter than first article of antenna 1; with
rounded mesiodistal lobe not visible in dorsal view; cornea
globular, distally placed. Antennular peduncle reaching two
thirds along antennal peduncle article 5. Antennal peduncle
with acute scaphocerite. Right maxilliped 3 (left missing) basis
with 2 small mesial teeth; ischium with crista dentata of 13
teeth, larger distally; merus width about 0.7 as long as ischium
and merus together; merus shorter than ischium, with tooth on
free distal margin; carpus articulating distolaterally on merus;
propodus as wide as long, expanded as asymmetrical lobe;
dactylus one third as wide as propodus, 0.7 times as long.
Pereopods 1 unequal, dissimilar; coxae with small
mesiodistal process with terminal setae. Major pereopod 1
(right cheliped) carpus-propodus upper margin 0.5 times
carapace length; ischium lower margin with row of 6 spines,
7
N
Y,
; rT \\\ \
mt NM
d
SU
|
ay
Figure |. Articullichirus articulatus (Rathbun, 1906). Hawaii, NHMW 6621 (ovigerous female, 7.5 mm). a, anterior carapace, eyestalks, antennular
peduncle, antennal peduncle; b, c, telson, uropods (depressed); d, left uropod, telson; e, right uropod, telson. Scale bar = 1 mm.
126 G.C.B. Poore, P.C. Dworschak & K.E. Schnabel
Figure 2. Articullichirus articulatus (Rathbun, 1906). French Polynesia, MNHN IU-2013-19994 (female, 7.3 mm). a, carapace, eyestalks,
pleomeres 1, 2; b, pleomeres 5, 6, telson (depressed); c, d, anterior carapace, eyestalks, antennular peduncle, antennal peduncle; e, pleomere 6,
telson, lateral view; f, telson, dorsal view; g, uropodal endopod, dorsal view; h, uropodal exopod, dorsal view; 1, maxilliped 3, outer view; J, right
maxilliped 3, coxa to merus, mesial view (palp removed); k, proximal articles of left maxilliped 3 and cheliped in situ, ventral oblique view; 1,
left cheliped coxa, ventral view; m, right cheliped, mesial view; n, right cheliped fingers, lateral view; o, right cheliped propodus, dactylus, upper
view; p, left cheliped; q, pereopod 2; r, pereopod 4. Scale bars = 1 mm. cl, bl, coxa, basis of right cheliped; cm, bm, 1m, coxa, basis, ischium of
maxilliped 3. Scale bars = 1 mm.
Articullichirus
larger distally; merus lower margin with 2 proximal teeth
followed by irregularly toothed blade, upper margin evenly
convex; carpus twice as wide as long, with blunt tooth at end of
upper and lower margins; propodus upper margin smooth, with
submarginal carina on mesial face, most visible distally, palm
1.1 times as long as wide, distomesial edge straight; fixed finger
0.45 times upper margin, cutting edge with broad tooth at
midlength; dactylus stout, slightly longer than fixed finger, with
terminal tooth, cutting edge with blunt tooth at midlength; ratio
of dorsal lengths, merus: carpus: propodus - 1 : 0.5 : 1.45.
Minor pereopod 1 (left cheliped) about as long as major;
ischium lower margin with 7 similar spines; merus lower
margin with 3 proximal spines followed by irregularly toothed
blade; carpus 1.1 times as wide as long, with tooth at end of
upper and lower margins; propodus upper margin 1.2 times as
long as greatest width; fixed finger 0.8 times upper margin,
cutting edge lateral, crenellate over distal third; dactylus
overreaching fixed finger, cutting edge smooth; ratio of dorsal
lengths, merus: carpus: propodus - 1 : 0.7 : 1.0.
Uropod endopod 1.5 times as long as wide, anterior margin
convex, without setae; posterior margin slightly concave,
tapering to a narrowly rounded apex; marginal setae confined
to distal and posterodistal margins; upper face with cluster of
setae close to posterior margin, few scattered setae near
anterior margin, with 2 proximal teeth and another at
midlength. Exopod subtriangular, anterior margin straight,
posterior margin concave, width 0.7 times anterior margin,
upper face with proximal tooth; dorsal plate oblique, well
separate from distal margin, armed with imbricating robust
setae, posterodistal angle rounded, armed with imbricating
robust setae, diminishing along posterior margin.
Telson 1.4 times as wide as long, lateral margins strongly
evenly convex, posterior margin concave; with dorsal transverse
row of 2 pairs of spiniform setae separated medially by cluster
of fine setae; posterolateral margin with row of setae.
Distribution. Eastern Indo-Pacific (Hawaii, French Polynesia);
upper shelf.
Remarks. We were unable to examine the holotype at the
USNM but have illustrated others. Articullichirus articulatus
is distinguished from the other two species by its tapering
uropodal endopod, narrower in published figures (Rathbun,
1906; Edmondson, 1944) than in our figures. Rathbun (1906)
and Edmondson (1944: fig. 9d) noted a meral tooth on
maxilliped 3 of specimens from Hawaii, also present on the
female from French Polynesia examined by GCBP, but the
tooth is absent on NHMW 6621 from Hawaii examined by
PCD. Both specimens examined have a submarginal mesial
crest on the major cheliped propodus, two pairs of short
spiniform setae on the telson and a sharp scaphocerite. The
mesial hook on the pereopod | coxa (present in A. collaroy and
A. chiltoni) 1s absent or replaced in the French Polynesian
female by a short fingerlike process.
The description above is based on the specimen reported by
Poupin (1998). We have not examined the individual from
Moorea now in the Senckenburg Museum, Frankfurt, illustrated
by Sakai (1992), which we assume to belong to this species.
127
Articullichirus chiltoni sp. nov.
Figures 3, 4
http://zoobank.org/urn:lsid:zoobank.org:act:5205A319-D237-
4CEE-9ABE-6A037685A386
Callianassa articulata.— Chilton, 1911: 551—552.
? Glypturus articulatus.—Sakai, 1999: 76—78, fig. 15 (Gilbert Is,
Kiribati).
Glypturus collaroy.—Sakai, 2005: 139—141, fig. 29 (New Zealand).
Material examined. Holotype. New Zealand, Kermadec Is, Sunday I.
[now Raoul I.|, rockpool, Captain Bollons, 1907, CMNZ AQ3372
(male, 14.7 mmy).
Diagnosis. Maxilliped 3 basis with mesial spine; merus with
tubercle on distal margin. Pereopod 1 coxa with mesiodistal
hook. Uropodal endopod midlength about 1.5 times as long as
wide, with broadly rounded apex, with submarginal dorsal
cluster of setae midway along posterior margin, with 1 proximal
tooth. Telson 1.4 times as wide as long, with excavate posterior
margin; with transverse dorsal row of 12 or 13 pairs of
contiguous spiniform setae plus 1 or 2 more lateral.
Description of holotype. Carapace length 0.28 of total length;
cervical groove at 0.8 length of carapace; dorsal oval well
defined; hepatic region with weakly sclerified line between
dorsal oval and linea thalassinica. Rostrum and anterolateral
carapace spines with unsclerified basal region; rostrum an
anteriorly directed, acute spine nearly as long as eyestalk;
anterolateral spines acute, set slightly back from rostrum, half as
long as rostrum. Anterolateral branchiostegal lobe margin
extending dorsal to linea thalassinica, with sclerified plate below
linea thalassinica. Pleomere 1 (damaged) with weak transverse
eroove, with dorsolateral longitudinal setal row; pleomere 2
about 1.5 times as long as pleomere | tergite; pleomeres 3-5
scarcely expanded laterally, with dense setose areas; pleomeres 5
and 6 subequal in length, with posterolateral notch.
Eyestalks shorter than first article of antennular peduncle,
without produced mesiodistal apex; cornea globular, distally
placed. Antennular peduncle little shorter than antennal peduncle.
Antennal peduncle with acute scaphocerite. Right maxilliped 3
(left missing) basis with recurved mesial hook; ischium with
distinct crista dentata, teeth diminishing distally; merus width
about 0.8 as long as ischium and merus together; merus slightly
shorter than ischium, with blunt tooth on free distal margin;
carpus articulating distolaterally on merus; propodus slightly
wider than long, expanded as round lobe on lower margin;
dactylus one third as wide as propodus, 0.7 times as long.
Pereopods | unequal, dissimilar; coxae with strong mesial
hook. Major pereopod | (left cheliped) carpus-propodus upper
margin 0.9 times carapace length; ischium lower margin with
row of 6 spines, larger distally; merus lower margin with 3
proximal spines, upper margin strongly convex proximally;
carpus 1.6 times as wide as long, with blunt tooth at end of
upper and lower margins; propodus upper margin smooth,
with submarginal carina on mesial face, palm slightly longer
than wide, distomesial edge convex; fixed finger 0.5 times
upper margin, cutting edge with broad tooth at midlength,
proximally serrated; dactylus stout, slightly longer than fixed
finger, with terminal tooth, cutting edge slightly concave, with
128 G.C.B. Poore, P.C. Dworschak & K.E. Schnabel
= ~
TNT gay
b um
Figure 3. Articullichirus chiltoni sp. nov. New Zealand, CMNZ AQ3372 (holotype male, 14.7 mm). a, carapace, eyestalks, dorsal view; b, pleon,
telson, dorsal view; c, anterior carapace, pterygostomium, eyestalk, antennular peduncle, antennal peduncle, lateral view; d, anterior carapace,
eyestalk, antennular peduncle, antennal peduncle, dorsal view; e, thoracic sternite 7, pereopodal coxae 1—4, detail of right pereopodal coxa 1; f,
maxilliped 3, outer view, detail of tooth on upper meral margin; g, maxilliped 3, inner view; h, major cheliped, left, lateral view; 1, major cheliped,
left,upper view; J, minor cheliped, right, lateral view. Scale bars = 1 mm.
Articullichirus
blunt tooth at midlength; ratio of dorsal lengths, merus: carpus:
propodus - 1 : 0.76: 1.38.
Minor pereopod | (right cheliped) about 0.8 times length of
major; ischium lower margin with 5 spines, larger distally; merus
lower margin with | proximal spine; carpus subtriangular, 1.2
times as wide as long, with tooth at end of upper and lower
margins; propodus upper margin about as long as greatest width;
fixed finger 1.1 times upper margin, cutting edge with 2 teeth in
\
ms
i ^T T
129
distal quarter; dactylus as long as fixed finger, cutting edge smooth;
ratio of dorsal lengths, merus: carpus: propodus — | : 0.85 : 0.92.
Pereopod 2 chelate. Pereopod 3 ischium slightly longer than
wide, about half length of merus; carpus slightly shorter than
merus; propodus 0.7 times length of carpus, lower margin
broadly expanded proximally, densely setose; dactylus simple,
about half length of propodus. Pereopod 4 propodus semichelate,
with spiniform seta at base of finger; dactylus simple.
=
33 _
=z ST
GE a ES
E »
EA )
BZ
TS) = — =—
À
SN
Figure 4. Articullichirus chiltoni sp. nov. New Zealand, CMNZ AQ3372 (holotype male, 14.7 mm). a, left pereopod 2, outer view; b, left pereopod
3, inner view (setae not shown); c, left pereopod 3, outer view; d, left pereopod 4; e, left pereopod 4, detail of distal propodus and dactylus; f,
pleopod 1; g, h, right and left pleopods 2; 1, telson; J, uropodal endopod and exopod. Scale bars = 1 mm.
130
Male pleopod 1 uniramous, biarticulate; article 2 with
small distolateral triangular lobe. Male pleopod 2 biramous;
endopod with obsolete distomesial appendix masculina;
exopod longer than endopod.
Uropod endopod 1.5 times as long as wide, anterior
margin convex, without setae; posterior margin slightly
concave, tapering to a narrowly rounded apex; marginal setae
confined to distal and posterodistal margins; upper face with
cluster of setae close to posterior margin, few scattered setae
near anterior margin, with | proximal tooth. Exopod longer
than endopod, subtriangular, as wide as anterior margin,
upper face with proximal tooth; dorsal plate oblique, well
separate from distal margin, armed with imbricating robust
setae, posterodistal angle rounded, armed with imbricating
robust setae.
Telson 1.4 times as wide as long, lateral margins evenly
convex, posterior margin concave; with dorsal transverse row
of spiniform setae, | or 2 robust setae placed laterally, remote
from row of 12 (right) and 13 (left), separated medially by
cluster of fine setae; posterolateral margin with row of setae.
Etymology. Named after New Zealand carcinologist Charles
Chilton (1860-1929) who first reported this specimen from the
Kermadec Islands.
Distribution. Kermadec Islands, New Zealand and possibly
Kiribati; intertidal.
Remarks. The single specimen collected from the Kermadec
Islands is more similar to Articullichirus collaroy than to A.
articulatus. The new species differs from A collaroy and A.
articulatus as follows: the transverse row of 28 spiniform setae
on the telson exceeds those of the other species (< 9), and the
single or two spiniform setae placed laterally on the outside of
the submedian ridges are absent in both A. collaroy and A.
articulatus. The uropodal endopod has one distinct proximal
spine on the dorsal surface, rather than two distinct spines as in
both other species. Articullichirus chiltoni shares the distinct
curved spine on the maxilliped 3 basis and the hook on the
pereopod | coxa with A. collaroy; these are absent or reduced
in A. articulatus.
Sakai (2005) identified and partly illustrated two
ovigerous females of “Glypturus collaroy" from Flax Bush
Bay, New Zealand — these specimens cannot now be found (R.
Webber, pers. comm., 14 Dec 2021). Sakai (2005) specifically
mentioned a spine on the merus of maxilliped 3 and a telson
with a concave posterior margin. These features are consistent
with Articullichirus but the upturned rostrum is enigmatic,
more like that of Glypturus. Sakai (1999) recorded a very
small male (cl. 4.5 mm) from Gilbert Is (Kiribati) as
“Glypturus articulatus”; this specimen (Swedish Museum of
Natural History, SMNH 16226) cannot now be found (S.
Stöhr, pers. comm., 7 Mar 2022). Saka1’s illustrations of the
short excavate telson, uropodal endopod and maxilliped 3
resemble those of A. chiltoni. Minor differences, such as more
teeth on the merus of the major cheliped, could be due to this
specimen being only one third of the size of the holotype
from New Zealand.
G.C.B. Poore, P.C. Dworschak & K.E. Schnabel
Articullichirus collaroy (Poore and Griffin, 1979) comb. nov.
Figure 5
Callianassa collaroy Poore and Griffin, 1979: 260-263, figs 24, 25.
Glypturus collaroy.—Sakai, 1988: 61.—Sakai, 2011: 434 (part).
Corallianassa collaroy.—Tudge et al., 2000: 144.—Davie, 2002: 460.
Neocallichirus collaroy.—Sakai, 1999: 98.
Corallianassa sp. MoV 4965.—Poore et al., 2008: 95, colour figure.
Not Glypturus collaroy.—Sakai, 2005: 139-141, fig. 29 ? =
Articullichirus chiltoni sp. nov.
Not Corallianassa collaroy.—Sakai,
Articullichirus articulatus.
1992: 212, fig. 1 =
Material examined. Australia, Western Australia, Great Australian
Bight, 110 nm (204 km) SW of Eucla, 33° 20'S, 127° 45'E, 260 m, SAM
C17888 (female, 9.3 mm). Off Bald Island, 35.19°S, 118.649°E, 161-169
m (stn SS10/2005/038), NMV J55438 (female, 13.2 mm). Off Two
Rocks, 31.7244°S, 115.244°E, 102 m, NMV J53458 (male, 10.5 mm)
Diagnosis. Maxilliped 3 basis with mesial spine; merus with
tooth on distal margin. Pereopod | coxa with mesiodistal hook.
Uropodal endopod midlength about 1.2 times as long as wide,
with semicircular apex, with 5 dorsal clusters of setae in distal
half, with 2 proximal dorsal teeth. Telson 1.3 times as wide as
long, with excavate posterior margin; with transverse dorsal
row of 5 pairs of spiniform setae.
Supplementary description. Carapace length 0.27 of total length;
cervical groove at 0.9 length of carapace; dorsal oval well
defined; hepatic region with weakly sclerified line between
dorsal oval and linea thalassinica. Rostrum and anterolateral
carapace spines with unsclerified basal region; rostrum an
anteriorly directed, acute spine longer than eyestalk; anterolateral
spines set slightly back from rostrum, acute, half as long as
rostrum. Anterolateral branchiostegal lobe well defined, margin
extending dorsal to linea thalassinica, with sclerified plate below
linea thalassinica. Pleomere | without transverse groove, with
dorsolateral longitudinal setal row followed by cluster of simple
setae; pleomere 2 1.8 times as long as pleomere | tergite;
pleomeres 3-5 scarcely expanded laterally with dense setose
areas; pleomere 6 1.3 times as long as pleomere 5, with
posterolateral notch.
Eyestalk without produced mesiodistal apex; cornea
swollen, distal. Antenna with acute scaphocerite. Antennular
peduncle reaching almost to end of antennal peduncle article 5.
Antennal peduncle with acute scaphocerite. Maxilliped 3 basis
with recurved mesial hook; merus with prominent tooth on free
distal margin.
Pereopods | coxae each with strong mesial hook. Major
pereopod | (cheliped) ischium lower margin spinose; merus
lower margin with 3 proximal spines, upper margin strongly
convex; carpus with tooth at end on upper and lower margins;
propodus upper margin rounded, with submarginal carina on
mesial face, about as long as greatest width, distomesial edge
straight; fixed finger 0.8 times upper margin, cutting edge with
small tooth; dactylus as long as fixed finger, cutting edge with
triangular tooth.
Minor pereopod 1 (cheliped) 0.8 times length of major;
ischium lower margin spinose; merus lower margin with 2
proximal spines; carpus as long as wide; propodus upper margin
as long as greatest width; fixed finger 1.3 times upper margin,
Articullichirus
131
a, b, k-m Wm
c-f, h
Figure 5. Articullichirus collaroy (Poore and Griffin, 1979). Western Australia, NMV J53438 (male, 10.5 mm). a, carapace, eyestalks, pleon,
telson, dorsal view; b, carapace, eyestalk, pleomere 1, lateral view; c, d, anterior carapace, eyestalk, antennular peduncle, antennal peduncle,
lateral and dorsal views; e, pleomere 6, telson, dorsal view; f, right uropod, dorsal view; g, pereopodal coxae 1—4, thoracic sternite 7, ventral view;
h, proximal articles of right cheliped and maxilliped 3 in situ, ventral oblique view; 1, jJ, maxilliped 3, inner and outer views; k, major cheliped
(left), mesial view; |, major cheliped fingers, lateral view; m, minor cheliped (right), mesial view; n, o, right pleopods 1, 2, posterior views. Scale
bars = | mm. cl, coxa of right cheliped; cm, bm, im, coxa, basis, ischium of maxilliped 3. Scale bar = 1 mm.
-—)
eee |
p
5
—
pe]
132
cutting edge smooth; dactylus as long as fixed finger, cutting
edge smooth.
Male pleopod | biarticulate; article 2 with small distolateral
triangular lobe. Male pleopod 2 biramous; endopod with
obsolete distomesial appendix; exopod longer than endopod.
Uropod endopod 1.6 times as long as wide, with distal marginal
setae, upper face with 5 clusters of setae over distal half, with
2 proximal teeth. Exopod about as wide as anterior margin,
with proximal tooth on dorsal face, dorsal plate oblique, well
separate from distal margin, setose, posterodistal angle rounded,
densely setose.
Telson 1.3 times as wide as long, lateral margins strongly
convex, posterior margin concave; with 5 pairs of dorsal
spiniform setae in transverse row, finer setae mesially.
Size. Cl. to 13 mm.
Type locality. Australia, New South Wales, Long Reef,
Collaroy.
Distribution. Temperate Australia (from off Sydney to off
Perth); intertidal—260 m.
Remarks. Poore and Griffin's (1979) figures did not show the
acute scaphocerite or spiniform setae on the telson that
characterise this new genus. This is remedied here.
Sakai’s uses of the species name are in error. His
"Glypturus collaroy" from New Zealand (Sakai, 2005) has an
upturned rostrum, not typical of the genus, but could
otherwise be synonymous with A. chiltoni (see comments
under that species above). His "Corallianassa collaroy" from
French Polynesia (Sakai, 1992) is A. articulatus.
The present identifications extend the distribution of this
species from the intertidal of central NSW to deep water off
southeastern WA, Australia.
Acknowledgements
This study was supported by the National Taxonomy Research
Grant Program (Grant numbers CN216-14 and CBG18-06 to
GCBP) from the Australian Biological Resources Study. KES
was funded by the National Institute of Water and Atmospheric
Research under Coasts and Oceans Research Programme 2
Marine Biological Resources: discovery and definition of the
marine biota of New Zealand (2021-2022 SCI). We thank
Johnathon Ridden and Erna Tidy (Canterbury Museum,
Christchurch, New Zealand) for the loan of the Chilton material,
Rick Webber (Museum of New Zealand Te Papa Tongarewa,
Wellington) and Sabine Stohr (Swedish Museum of Natural
History, Stockholm) for searching for specimens in their
collections, and Laure Corbari (Muséum national d’ Histoire
naturelle, Paris) for a loan.
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Memoirs of Museum Victoria 81: 135-162 (2022) Published 2022
ISSN 1447-2554 (On-line)
httos://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/
DOI https://doi.org/10.24199/|.mmv.2022.81.06
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera:
Halictidae: Halictini: genus Lasioglossum) — *“Wood-Splitting Axe Bees"
http://zoobank.org/urn:|sid:zoobank.org:pub: DDDBDF43-838A-4C99-8F73-38ED597A6C F4
KENNETH L WALKER
(ORCID ID https://orcid.org/0000-0001-8750-2035). Museums Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia.
|kwalker@museum.vic.gov.au]
Abstract
Keywords
Walker, K.L. 2022. Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae:
Halictini: genus Lasioglossum) — "Wood-Splitting Axe Bees". Memoirs of Museum Victoria 81: 135-162.
The Australian Lasioglossum Curtis 1833 subgenus Australictus Michener 1965 is revised.
Of the 11 available names listed by Michener (1965) for the subgenus Australictus, six are placed in synonymy. A
species placed by Michener (1965) in the Lasioglossum subgenus Parasphecodes Smith 1853 is recombined to the subgenus
Australictus, and four other species, placed in Parasphecodes by Michener (1965), are synonymised with this valid,
recombined taxon, and the species name of the taxon is reverted to its original spelling. In addition, a species placed in
Australictus by Michener (1965) is synonymised with a valid species in the Lasioglossum subgenus Chilalictus Michener
1965. These changes provide five valid names for the subgenus Australictus.
New synonymies, recombined names and valid species proposed for Lasioglossum (Australictus) are as follows:
New synonymies — Lasioglossum (Australictus) kurandense (Cockerell 1914) syn. nov., Lasioglossum (Australictus)
nigroscopaceum (Friese 1917) syn. by Cockerell 1929 but listed by Michener (1965: 165) as valid = Lasioglossum
(Australictus) davide (Cockerell 1910a); Lasioglossum (Australictus) insculptum (Cockerell 1918) syn. nov., Lasioglossum
(Australictus) rufitarsum (Rayment 1929) syn. nov. and Lasioglossum (Australictus) fulvofasciae Michener 1965 syn. nov.
= Lasioglossum (Australictus) tertium (Dalla Torre 1896); Lasioglossum (Australictus) franki (Friese 1924) syn. nov. =
Lasioglossum (Chilalictus) orbatum (Smith 1853). New combination and new synonymies — Lasioglossum (Parasphecodes)
lithuscum (Smith 1853) comb. nov. moved to Lasioglossum (Australictus) lithusca; Lasioglossum (Parasphecodes)
adelaidae (Cockerell 1905) syn. nov., Lasioglossum (Parasphecodes) griseipenne (Cockerell 1929) syn. nov., Lasioglossum
(Parasphecodes) stuchilum (Smith 1853) syn. nov., Lasioglossum (Parasphecodes) wellingstoni (Cockerell 1914) syn. nov.
= Lasioglossum (Australictus) lithusca. Valid is Lasioglossum (Australictus) plorator (Cockerell 1910b).
New female subgeneric mandibular characters are added to Michener’s (1965) diagnostics for Australictus —
mandible with elongated and enlarged preapical tooth, reduction in width of basal tooth at apical end and ın dorsal view,
broadening of width at base of mandible. The mandibular modifications, widening of the head basally and enlarged gena
are associated with Australictus behaviour to nest in wood rather than ground nesting in soil as is usual for Halictidae bees.
The shape of the female mandible, especially in dorsal view, resembles a wood-splitting axe — hence the common name
coined here, “Wood-Splitting Axe Bees”. Australictus 1s the first record of wood-nesting bees for Australian Halictidae.
All valid species are redescribed; keys to both sexes, montage diagnostic images and distribution maps are provided
to assist with species identification.
Hymenoptera, Halictidae, Halictini, Lasioglossum, Parasphecodes, Chilalictus, Australictus, nesting behaviour.
Introduction
The world’s bee fauna consists of seven families, Apidae,
Colletidae, Halictidae, Megachilidae, Stenotritidae, Andrenidae
and Melittidae, with the latter two families absent from Australia.
Halictidae is the second largest bee family in the world, with
4,510 species (Ascher and Pickering, 2022), and is found on all
continents except Antarctica (Michener, 2007). Most of this
species richness 1s concentrated in the tribe Halictini, with 2,882
species (Ascher and Pickering, 2022).
Lasioglossum Curtis 1833 is one of the most species-rich
and taxonomically challenging groups of all world bee genera,
with 2,645 named species to date (Ascher and Pickering, 2022).
Lasioglossum bees are distinguished from other Halictini
genera by having the third submarginal (2rs-m) (“strong-veined
Lasioglossum series”) and sometimes the second submarginal
136
cross veins (Irs-m) (“weak-veined Hemihalıctus series’)
weaker than the first submarginal cross vein (Michener 2007,
see especially figs. 66—6a and c).
The Australian component of this most diverse genus within
the Halictidae has 308 available names (ABRS, 2022a; note:
update to ABRS, 2022a includes changes for Callalictus
Michener 1965 — see Walker (2022); and here, Homalictus
Cockerell 1919 is treated as a subgenus of Lasioglossum — see
Danforth, 1999, Danforth et al., 2001, 2003, 2008; Gibbs et al.,
2012, 2013; Zhang et al., 2022). In Australia, Lasioglossum 1s
divided into nine subgenera as follows: Australictus Michener
1965 (11 species); Austrevylaeus Michener 1965 (six species);
Callalictus (three species); Chilalictus Michener 1965 (139
species); Ctenonomia Cameron 1903 (nine species); Glossalictus
Michener 1965 (one species); Homalictus (46 species);
Parasphecodes Smith 1853 (92 species); and Pseudochilalictus
Michener 1965 (one species).
The subgenus Australictus, with 11 available names, is a
striking Lasioglossum subgenus of the Australian bee fauna.
The subgenus occurs from north Queensland (two species),
down coastal New South Wales, across Victoria, widely in
Tasmania (two species), and there are three specimen records
from south-east South Australia (igs. I2A-E). Bees of this
subgenus nest in wood, either excavating their own nests or
modifying the vacated burrows of beetles. This is the first record
of wood-nesting Halictidae in Australia. To accommodate this
wood-nesting behaviour, the mandibles of females have been
modified as described below.
A full morphological diagnosis for the subgenus
Australictus 1s provided. Species can be generally
distinguished from other Australian Lasioglossum subgenera
by the following characters: body length moderately large
(females: 8.32 mm to 10.23 mm; males: 8.01 mm to
10.21 mm); female head widened basally in frontal view, with
lower inner eye width almost equal to or wider than upper
inner eye width (fig. 1A); bidentate mandible of females with
elongated and enlarged preapical tooth (fig. IC) and broadened
at base (fig. IE); female labrum medium process truncate
apically with lateral margins pectinate (fig. 2A); dorsolateral
angles of pronotum rounded (figs. 5A, 6A, 7A, 8A, 9A); body
colour black, one species with metasoma red-brown (figs. 6A,
6B); several species with no body tomentum (figs. 5A—D,
6A-D, 8A—D), several species with various combinations of
yellow/white tomentum on dorsolateral angles of pronotum,
mesoscutum, metanotum and basal band across metasomal
12-I3 (figs. 7A-D, 9A-D); female anterior metatibial spur
finely serrate (fig. 2B); and male genitalia with gonobase
either basally widened (figs. 10A, 10B, 11C, 11D) or narrowed
(figs. IOC—F, 12A, 12B), retrorse lobe either short (figs. 11A,
11C) or elongate (figs. 10A, LOC, 10E) and gonostylus either
short (figs. LOA, LOC, 11A) or absent (figs. LOE, 11C).
The shape of the female mandible, especially in dorsal
view, ıs broad basally and apically pointed (fig. IE),
resembling a wood-splitting axe — hence the common name
suggested here, “Wood-Splitting Axe Bees”.
The Australian Lasioglossum subgenus Australictus 1s
revised. Of the 11 available names listed by Michener (1965) for
the subgenus, six are placed in synonymy. A species placed by
K.L. Walker
Michener (1965) in the Lasioglossum subgenus Parasphecodes
is recombined to the subgenus Australictus and four other
species, placed in Parasphecodes by Michener (1965), are
synonymised with this valid, recombined taxon, and the species
name of the taxon is reverted to its original spelling. In addition,
a species placed in Australictus by Michener (1965) is
synonymised with a valid species in the Lasioglossum subgenus
Chilalictus Michener 1965. These changes provide five valid
names for the subgenus Australictus.
All valid species are redescribed, keys to both sexes,
montage images and distribution maps are provided to assist
with species identifications.
Material and methods
Terminology follows Michener (2007), except the propodeum
is called the metapostnotum (Brothers, 1976; Gibbs et al.,
2013; Gibbs et al., 2022; Engel and Gonzalez, 2022) and the
inner hind tibial spur is called the anterior metatibial spur
(Aguiar and Gibson, 2010). To assist with the descriptions, the
following notes are provided: relative head measurements
were Standardised to a head width of 100 units; absolute
measurements are expressed as minimum-mean-maximum
if multiple specimens were available; body length was
measured from antennal sockets to end of metasoma; forewing
length was measured from the base of the arcuate basal vein
(vein M) to the distal margin of the third submarginal cell
(vein 2rs-m); and, intertegular distance was measured as the
ereatest width between the tegulae. Sculpture nomenclature
follows Harris (1979). Mesoscutum punctation nomenclature
is as follows: dense, interspaces between punctures less than
diameter of a puncture; close, interspaces between punctures
equal to diameter of puncture; open, interspaces between
punctures greater than | x but less than 2 x diameter of
puncture; sparse, interspaces between punctures equal to or
sreater than 2 x diameter of puncture. To standardise the
description of the mesoscutum punctation, the mesoscutum
was subdivided into areas as follows: anteromesial, area along
the leading edge of the mesoscutum and on each side of the
midline; anterolateral, area on the anterior lateral corners of
the mesoscutum; mesial, area between parapsidal lines;
parapsidal, areas between parapsidal line and nearest lateral
margin; laterad of parapsidal lines, area adjacent to outer
margin of the parapsidal lines; mesiad of parapsidal lines,
area adjacent to the inner margin of the parapsidal lines;
posterior, area along the posterior margin of the mesoscutum
(see diagram in Walker, 1995: 258, fig. 3).
Due to the importance of vestiture as diagnostic characters
on the male Lasioglossum genital capsule, in particular on the
conocoxa, retrorse lobe and gonostylus, a decision was made
to photograph dry mounted capsules rather than the usual
method of photography in glycerol. Treatment of the genital
capsule was as usual: specimen placed in humid chamber for
24 hours, removal of genital capsule from the bee using a
hooked no. 3 entomological pin (38 mm x 0.53 mm), 24 hours
in 10% KOH cold, several water washes, then placed on tissue
paper and allowed to dry for several hours. A micro-headless
pin B3 (0.193 mm x 15 mm) was pinned to a piece of foam
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 137
attached to a no. 3 entomological pin so that the blunt end of
the micropin was pointing out. A droplet of water-soluble glue
was placed on the end of the micropin, one side of the
micropin was scraped to reduce the amount of glue, and the
blunt end was brought in contact with the left side basal corner
of the gonobase and the genitalia allowed to dry. While
drying, it is important to keep the glued genital capsule at the
üp of the micropin. To achieve this dried position, the
micropin must be held in a horizontal position with the genital
capsule underneath its tip. If not placed in this drying position,
the genital capsule can move down or around the micropin,
obscuring the gonobase for photography. Ventral and dorsal
montage images were taken against a black background to
highlight diagnostic vestiture and structures. Following
photography, each pinned capsule was attached to the
specimen's pin pith. A decision was made to exclude images
of the seventh and eighth metasomal sternal segments. These
two sterna were similar in shape across all species and offered
no unique diagnostic characters.
Photographs were taken using a Leica M205C microscope
with a Leica DFC500 c-mount camera using LAS Version 3.8
to create image montage stacks and montaged images. Images
were then post-processed and image plates formed in Adobe
Photoshop 6.
Abbreviations used in the text are as follows: AOD,
antennocular distance; AF, antennal flagellomeres; CL, clypeal
length; EW, eye width, in side view; FL, flagellum length; GW,
gena width, in side view; HL, head length; HW, head width; IAD,
interantennal distance; IOD, interocellar distance; LID, lower
interorbital distance; OAD, ocellantennal distance; OOD,
ocellocular distance; SE — South East; SW — scutum width; S1—S5,
metasomal sterna 1-5; SL, scape length; SW, scutum width; T1-T6,
metasomal terga 1-6; UID, upper interorbital distance.
This study was based on examination of just over 1,000
specimens (and several citizen science images posted on
Facebook - links within species descriptions) from the
following collections.
AM - Australian Museum, Sydney, New South Wales; ANIC
— Australian National Insect Collection, CSIRO, Canberra,
Australian Capital Territory; BDBSA - Department for
Environment and Water, SA Fauna, Adelaide, South Australia;
BMNH - Natural History Museum, London, United Kingdom;
Cornell — Cornell University, Ithaca, New York; NMV Museums
Victoria (formerly the National Museum of Victoria), Melbourne,
Victoria; QDPI — Queensland Department of Primary Industries,
Brisbane, Queensland; OAI — Orange Agricultural Institute,
Orange, New South Wales; QM — Queensland Museum, Brisbane,
Queensland; SEM — Snow Entomological Museum, Kansas
University Biodiversity Institute, University of Kansas, Kansas,
United States of America; TMAG — Tasmanian Museum and Art
Gallery Invertebrate Collection, Hobart, Tasmania; TDA -
Tasmanian Department of Agriculture, Hobart, Tasmania; WAM
— Western Australian Museum, Perth, Western Australia.
Keys to both sexes of Lasioglossum (Australictus) species
] Metasoma with yellow/white basal tomentum bands on
T2-T3 (figs. female: 7A, 9A; male: 7C, 9C) 2
- Metasoma without tomentum bands on terga (figs.
female: 5A, 6A, 8A; male: 5C, 6C, 8C)
2 Female with yellow/white tomentum on posterolateral
corners of mesoscutum and on metanotum (fig. 7A); male
with an apical V-patch of tomentum on posterior vertical
posterior surface of metapostnotum (figs. 2C, 70)
L. peraustrale (Cockerell, 1904)
- Female without yellow/white tomentum on mesoscutum or
metanotum (fig. 9A); male without apical V-patch of
tomentum on posterior vertical posterior surface of
metapostnotum (fig. 9C) L. tertium (Dalla Torre, 1896)
3 Metasoma red-brown (figs. female: 6A; male: 6C)
AER bik EN ll MEAP OAT PT bt VRE TLE REI L. lithusca (Smith, 1853)
- . Metasoma black (figs. female: 5A, 8A; male: 5C, 8C) 4
4 Dorsal surface of metapostnotum posterior rim defined by
raised carina, dorsal sculpture ruguloso-striolate (figs.
female: 2E, 5A; male: 5C) L. davide (Cockerell, 1910a)
- Dorsal surface of metapostnotum posterior rim not defined
by carına, rım smooth and elevated, dorsal sculpture
microalveolate (figs. female: 2F, 8A; male: 8C) .......
L. plorator (Cockerell, 1910b)
Taxonomy
Subgeneric diagnosis: body length moderately large (female:
8.32 mm to 10.83 mm; male: 8.01 mm to 10.21 mm), robust,
non-metallic species; in both sexes, body colour black (figs. 5A—
D, 7A-D, 8A—D, 9A-D) except L. lithusca metasoma red-brown
(figs. 6A-D); L. tertium male metasoma either black (figs. 9C—
D) or banded with alternating dark to light red-brown tergal
colours (fig. 2D); mesoscutum and metasomal terga of L. davide
and L. plorator with tinge of deep blue colour (figs. 5A, 8A);
female inner eye widths vary from narrowed to almost parallel
to diverging basally from upper to lower inner eye widths (e.g.
in L. tertium lower inner eye width 1.2 x upper inner eye width),
meaning head widened ventrally (fig. 1A), and gena width
enlarged (fig. 9B), (Australian Lasioglossum subgenera typically
with eyes widths converging basally — e.g. L. (Parasphecodes)
hiltacum (Smith, 1853) lower inner eye width 0.94 x upper inner
eye width (fig. 1B); female bidentate mandible modified by
elongation and enlargement of preapical tooth, reduction in
width of basal tooth, enlargement of height and width at base of
mandible, reduced width of condylar groove (figs. IC, LE) (see
Table 1 for comparison of Australictus (figs. IC, 1E) with
Parasphecodes (figs. 1D, 1F)); labrum medium process apically
truncate, laterally pectinate (fig. 2A); pronotum with rounded
dorsolateral angles (figs. 5A, 6A, 7A, 8A, 9A); wing veins with
second transverse cubital vein (1rs-m) strong, 1* recurrent vein
(Im-cu) entering second submarginal cell or in line with Irs-m
except in L. tertium, where 1* recurrent vein enters third
submarginal cell; female hind basitibial plate rounded to weakly
pointed apically; female anterior metatibial spur finely serrate
(fig. 2B) to smooth; dorsal surface of metapostnotum either
carinate, as defined by raised carina forming raised lip or ridge
(fig. 2E), or acarinate, as defined by crescent-shaped, elevated
138 K.L. Walker
Table 1. Female mandibular morphological differences between Lasioglossum subgenera Australictus and Parasphecodes. (Note: L. (Australictus)
lithusca and L. (Parasphecodes) hiltacum were used for measurements)
Character Australictus
(figs. IC, 1E)
Condylar groove
Width of condylar groove 0.14 x width of base
of mandible between the acetabulum and
Parasphecodes
(figs. ID, 1F)
Preapical tooth length Preapical tooth elongated to almost same length | Preapical tooth shorter than length (0.79 x
(0.93 x length) as basal tooth length) of basal tooth
Preapical tooth width Preapical tooth enlarged and broader (1.14 x Preapical tooth not enlarged and narrower (0.42
width) than width of basal tooth x width) than width of basal tooth
Basal tooth width Width of basal tooth 0.13 x width of base of Width of basal tooth 0.21 x width of base of
mandible mandible
Width of condylar groove 0.18 x width of base
of mandible between the acetabulum and
condyle
condyle
Base of mandible width in side view Base of mandible length between acetabulum Base of mandible length between acetabulum
and condyle at 0.53 x length of mandible and condyle at 0.44 x length of mandible
Base of mandible width in dorsal view Dorsal width at base of mandible 0.47 x length Dorsal width at base of mandible 0.21 x length
of mandible of mandible
medium area distinctly raised above vertical and lateral surfaces
(fig. 2F); in both sexes, metasomal terga of L. peraustrale and L.
tertium with conspicuous basal bands of tomentum on
metasomal T2-T3 (figs. 7A—D, 9A-D), L. davide, L. lithusca
and L. plorator without such metasomal tergal tomentum bands
(figs. 5A-D, 6A—D, 8A-D). Male: some species with lateral hair
tufts on metasomal S3-S5 (figs. 6F, 7F, SF, 9F); genitalia with
sonobase widened basally (figs. 10A, 11C) or narrowed
(figs. 10C, LOE, 11A), gonostylus small (figs. 10A, 10B, 11A) to
absent (figs. LOE, 11C), retrorse lobes small (figs. LIA, 11C) to
enlarged (figs. 10A, 10C, 10E), either not meeting at midline
(figs. LOA, 10E) or overlapping at midline (fig. 10C), either
glabrous (figs. 10A, 11A, 11C) or setose (figs. 10C, LOE).
Notes. The elongated and enlarged preapical mandibular
tooth, the narrowing of basal tooth and condylar groove,
enlargement at the base of the mandible, the widening of the
head basally and the enlargement of the gena are all
adaptations to provide effective wood-splitting and wood-
excavating mandibles for female Australictus to nest in wood
rather than in soil as is usual for Halictidae bees.
Several specimen labels (at least five labels) refer to collected
specimens found nesting in “rotten wood”, and one specimen
had a label observation “from Sirex emergence hole”.
A series of Facebook images by Christopher Robbins for
L. (Australictus) plorator documented the wood excavation
nesting of this species (figs. 3A—F). The same species was also
found using a vacated beetle exit hole as a nesting site in wood
(fig. 4) (https://www.facebook.com/groups/1041684025880609/
search?q=christopher%20robbins).
Christopher Robbins posted these bee observations with
his Facebook images:
Chapple Vale, Victoria 19/04/2020. Found today while
splitting wood. There was no bark on the log though, the crack
in the log they make their tunnels into where the brood cells
were filled with old termite bed from an extinct termite colony.
There were four adults all up and two pupae. They do not seem
to worry about their access point to the log, they used cracks on
the top, sides and underneath, in some cases they had excavated
the chambers into decayed wood, a reddish wood rot, I believe
commonly known as red cube rot, we used to find the larvae of
a small reddish stag beetle in the same substrate.
These images and observations demonstrate
Australictus bees nest in wood.
Danforth et al. (2019) summarised the nest architecture of
bees. The most common nesting strategy used by solitary bees
is ground nesting in soil. An estimated 64% of non-parasitic bee
species nest in underground soil excavations (Cane and Neff,
2011). However, a few bee species excavate nests in wood (both
solid and rotten). Of the 27 known subfamilies of bees
worldwide, wood-nesting occurs primarily in only the
subfamilies Xylocopinae, Lithurginae and Callomelittinae
(Danforth et al., 2019). The tunnels and cells of wood-nesting
bees are entirely constructed using their mandibles, and wood-
nesting bees have modified mandibles that differ from the
mandibles of ground soil-nesting bees.
Most Halictidae are ground soil-nesting bees, and halictid
bees show limited diversity in their mandibular structures
(Danforth et al., 2019). However, wood-nesting halictids have
been recorded in several Halictini genera. Of the 37 genera of
halictine Augochorini that occur in North and South America,
four are known to nest in wood (Augochlora Smith 1853,
Megalopta Smith 1853, Neocorynura Schrottky 1910 and
Xenochlora Engel, Brooks and Yanega 1997 (Tierney et al.,
2008). Wood-nesting halictids have specialised bidentate
mandibles (Michener and Fraser, 1978). The mandible of wood-
excavating halictids is characterised as more stout and more
robust than the mandible shape of ground soil-nesting
Halictidae, with modified apical teeth, in particular an elongated
and enlarged preapical tooth; the mandible is widened at its base
and has groove variations on the outer and inner surfaces
(Michener and Fraser, 1978).
that
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 139
Australictus 1s the first record of wood nesting for any
Australian Halictidae. The bidentate mandible of Australictus
looks similar 1n shape and structure to the mandible of the
wood-nesting Lasioglossum subgenus Eickwortia McGinley
1999, especially the elongated and enlarged preapical tooth
found in both subgenera (for an Eickwortia mandible image, see
Gibbs and Dumesh, 2013, fig. 3). However, Australictus belongs
to the informal "strong-veined" Lasioglossum series, whereas
Eickwortia belongs to the informal “weak-veined” Hemihalictus
series of Lasioglossum, which can be recognised by the weaken
Irs-m and 2rs-m veins in the forewing (Michener, 2007; Gibbs
and Dumesh, 2013).
Species descriptions
Lasioglossum (Australictus) davide (Cockerell)
(Figs. 2E, SA-F, 10A-B, 12A)
Halictus davidis Cockerell 1910a: 234.
Halictus kurandensis Cockerell 1910a: 234. syn. nov.
Halictus nigroscopaceus Friese H. 1917 syn. by Cockerell 1929: 211.
Lasioglossum (Australictus) davide — Michener 1965: 165.
Lasioglossum (Australictus) kurandense — Michener 1965: 165.
Lasioglossum (Australictus) nigroscopaceum — Michener 1965: 165.
Material examined: Holotype of davidis X, Queensland, Kuranda (has
Cairns typed on label and Kuranda hand written), 4.02. Turner BMNH
Hym.17.a.914 (BMNH) (view type data and image at https://data.nhm.
ac.uk/object/f00271eb-7e4e-A41fb-9e8d-2736fa91179f accessed 16
August 2022).
Holotype of kurandensis 5, Queensland, Kuranda (Cairns typed
on label and Kuranda handwritten), 4. 02. Turner BMNH Hym.17.a.956
(BMNH) (view type data and image at https://data.nhm.ac.uk/object/
eleec50b-a2b4-4b89-84ca-6bea23587e8e accessed 16 August 2022).
Syntypes ("TY PUS”) of nigroscopaceus — Queensland: Malanda,
Mjöberg. All three specimens have same label and type number: Am.
Mus. Nat. Hist. Dept. Invert. Zool. No. 26926 (29, 14%). The male
specimen has Cockerell's handwritten label saying "Halictus davidis
CkII" (AMNH).
Other specimens examined: (139, 68): QUEENSLAND: (19) Upper
Mulgrave 19 June 1991, J.H. Barrett, nesting 1n rotten log (QDPI);
(1?) Tully Falls, S.F. 730m, 18 km SSW Ravenshoe, 7 Dec 1987 — 7
Jan 1988, R. Storey & B. Dickinson (QDPI); (1?) Mt Halifax summit,
45 km WNW Townsville, 4 Dec 1990 —8 Jan 1991, A. Graham, pitfall
and intercept traps (QDPI); (12) Mt Halifax summit, 45 km WNW
Townsville, 2 Dec 1990, A. Graham, hand collecting (QDPI); (19) Mt
Lewis nr Mossman, 22 Oct 1984, NW. Rodd (AM); (19, 14) Cairns
District, F.P. Dodd (SAM); (2%) Kuranda, 2 Jan 1953 & 25 Sept 1954,
GB (NMV); (59, 34) Kuranda Black Mountain Forest Road, Jan
2001, K. Walker, on Lophostemon grandifloris subsp. riparius
(NMV); (15) Kuranda, Jan 1952, J.G. Brooks (AM); (14%) Mt Spec
Nat. Pk, 10 km E Paluma, 22 Nov 1988, K. Walker, on Eucalyptus
(NMV).
Floral record: Family visited: 1 (Myrtaceae (2)). Genera visited: 2
(Eucalyptus, (1), Lophostemon (1)).
Flight phenology capture records: Jan (3) Feb (0) Mar (0) Apr (0) May
(0) June (1) July (0) Aug (0) Sept (1) Oct (1) Nov (1) Dec (3).
Diagnosis. Lasioglossum (Australictus) davide 1s most like L.
(Australictus) plorator in body colours. This species can be
distinguished, in both sexes, from other Australictus species
by black body colour with bluish tinge on mesoscutum; the
lack of tomentum on the mesosoma or metasoma; dorsal
surface of metapostnotum posterior margin carinate, defined
by raised posterior and lateral carinae; metapostnotum dorsal
surface sculpture ruguloso-striolate (figs. 2E, 5A-D); male
genitalia with gonobase widened basally (a characteristic
shared only with L. tertium), large retrorse lobes with small,
apically rounded gonostylus (figs. LOA, 10B); and males
lacking any distinctive metasomal sternal vestiture (fig. 5F).
This species is restricted to North Queensland (fig. 12A).
Description of female: (figs. 5A-B, 5E) body length: 9.58—
9.68-10.21 mm (n=10); forewing length: 2.45-2.47—2.49 mm
(n=10); head width: 2.78—2.88—2.95 mm (n=10); intertegular
width: 2.06—2.38-2.40 mm (n-10) Relative head
measurements: HW: 100, HL: 84-85, UID: 54-55, LID: 50-
52, IAD: 08-09, OAD: 21-22, IOD: 08-09, OOD: 14-15, CL:
19-21, GW: 18-20, EW: 20-22, SL: 37-40, FL: 63-65.
Head: (fig. 5E) inner eyes margins weakly narrowed
basally; median frontal carina reaching less than one third
way to median ocellus; clypeus entirely polished and smooth,
anterior half weakly concave medially, sparsely deeply
punctate, punctures rounded to elongate; supraclypeal area
distinctly raised above paraocular area, polished, smooth,
punctures small, round and open with small rounded
punctures; frons sculpture above antennal bases smooth but
dull, microtessellate with small punctures for setae insertion
points, paraocular area smooth and closely punctate.
Mesosoma: (fig. 5A) mesoscutum anterior mesial margin
weakly produced mesoanteriorly, surface smooth with dull,
“oily” sheen, medially openly punctate, laterad of parapsidal
areas closely punctate, parapsidal areas and posterior margin
densely punctate; scutellum 2 x as long as dorsal surface of
metapostnotum, scutellum smooth, with dull sheen, weakly,
openly punctate; dorsal surface of metapostnotum carinate
(fig. 2E), posterior margin with well-defined, raised,
semicircular carina and posterolateral carinae, dorsal surface
transversely ruguloso-striolate medially, striate laterally,
sculpture reaches posterior carina, lateral margins smooth
microalveolate, vertical posterior surface of metapostnotum
defined by lateral carinae meeting dorsal surface
posterolateral carinae; mesepisternum and metepisternum
plicate; first recurrent vein (lm-cu) meeting Irs-m vein or
entering second submarginal cell.
Metasoma and legs: (figs. 5A, 5B) metasomal TI-T5 dull,
smooth, closely to densely punctate with minute punctures;
anterior metatibial spur finely serrate to simple, with no
distinct teeth.
Colour: (figs. 5A, 5B) body and legs black except
mesoscutum and scutellum dark "oily" grey with bluish tinge.
Vestiture: (figs. SA, SB, 5E) sparse, clypeus and
supraclypeal area glabrous, frons with sparse small, black,
erect hair; mesoscutum and scutellum almost glabrous but
with sparse small, black, erect hair; long hair on lateral,
vertical posterior surface of metapostnotum; apical one third
of Tl with long, erect, white hair, remainder of Tl and T2
glabrous, T3 and T4 with some black, adpressed hair apically.
K.L. Walker
140
Figure 1. Lasioglossum (Australictus) tertium A; Lasioglossum (Australictus) lithusca C, E; Lasioglossum (Parasphecodes) hiltacum B, D, F. Head
front: A & B; Mandible outer view: C & D; mandible dorsal view: E & F. All female images.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 141
Figure 2. A, Lasioglossum (Australictus) lithusca female labrum; B, Lasioglossum (Australictus) tertium female anterior metatibial spur; C,
Lasioglossum (Australictus) peraustrale male vertical metapostnotum tomentum; D, Lasioglossum (Australictus) tertium, banded male metasoma
form; E, Lasioglossum (Australictus) davide female dorsal surface of metapostnotum; F, Lasioglossum (Australictus) plorator female dorsal
surface of metapostnotum.
142 K.L. Walker
-
Figure 3. A-F Lasioglossum (Australictus) plorator: A, B, live female; C, wood nesting gallery construction; D, dorsal view of pupa; E, brood
cell; F, lateral view of pupa. All images copyright Christopher Robbins.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 143
ene?
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> = ie
* TU X -
DO "Ll Hm
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Figure 4. Lasioglossum (Australictus) plorator female using a beetle exit hole in wood. Image copyright Christopher Robbins.
Description of male: (figs. SC, SD, SF) body length: 7.07—8.45—
8.79 mm (n=6); forewing length: 1.94—2.48—2.54 mm (n=6);
head width: 1.82—2.42—2.50 mm (n=6); intertegular width: 1.44—
2.15-2.21 mm (n=6). Relative head measurements: HW: 100,
HL: 88—89, UID: 56-57, LID: 42-43, IAD: 12-13, OAD: 23-24,
IOD: 15-16, OOD: 14-15, CL: 26-29, GW: 16-18, EW: 28-29,
SL: 20-23, AF4/AF2+3 (18/14, 20/14) 1.29—1.43, FL: 178—180.
Differs from female as follows: inner eyes converging more
basally; median frontal carina reaching about one quarter to
median ocellus; frons sculpture reticulate across surface to inner
margins of eyes; scape reaches basally level of median ocellus;
clypeus surface shiny weakly microtessellate basally, medium
area rounded not concave, openly punctate, supraclypeal area
protruding above paraocular area, supraclypeal area bulbous,
shining, openly to closely punctate; mesoscutum surface similar
to female in colour but densely punctate medially, openly
punctate laterad of parapsidial areas, densely punctate in
parapsidal areas and anterolaterally; scutellum shiny as in
female but closely punctate; dorsal surface of metapostnotum
same as in female, posteriorly carinate, dorsal surface ruguloso-
striolate; apical two thirds of clypeus pale yellow and metasoma
in some specimen with distinct blue tinge.
Vestiture: frons hair dense, erect, black; paraocular area
hair sparse, white, adpressed; mesoscutum appearing glabrous
but with sparse, short, erect, black hair; metapostnotum lateral
sides with short, white, adpressed hair; apical posterior
vertical posterior surface of metapostnotum glabrous;
metasomal sterna with moderately dense, short erect and
adpressed black setae, no distinct patterns observed (fig. 5F).
Genitalia: (figs. 10A, 10B) gonobase widened basally,
complete ventroapically, gonocoxa wider and longer than
sonobase, with sparse, erect setae dorsoapically, glabrous
ventrally, dorsal inner margins of gonocoxa basally rounded,
apical inner margin not produced continuing contour of
sonostylus, glabrous; retrorse lobes large, overlapping at
midline, membranous, basal inner margins with short setae,
apical inner margins of retrorse lobes glabrous; gonostylus
small (about one third length of gonocoxa) erect, small,
apically rounded, with sparse short setae; penis valves curved
apically, with short dense hair dorsolaterally.
Distribution: (fig. 12A) the species is restricted to north
Queensland between approximately Townsville to Cairns.
Remarks: Cockerell (1929: 211) synonymised Halictus
nigroscopaceus with Halictus davidis; however, Michener
(1965: 165) listed Lasioglossum (Australictus) nigroscopaceum
as a valid species. Cockerell's (1929) synonymy was checked and
accepted here. Due to the restricted distribution of this species,
fewer than 20 specimens were located for this study. The only
significant variation observed was in the body length of males
(7.07 mm to 8.79 mm). Lasioglossum davide and L. tertium are
the only two Australictus species in which the gonobase is
widened basally (figs. 10A, 11C); ın the other three Australictus
species, the gonobase narrows basally (figs. LOA, LOE, 11A).
144 K.L. Walker
Figure 5. A-F Lasioglossum (Australictus) davide: A, dorsal female; B, lateral female; C, dorsal male; D, lateral male; E, female head front; F,
male vestiture on metasomal sterna.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 145
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Figure 6. A-F Lasioglossum (Australictus) lithusca: A, dorsal female; B, lateral female; C, dorsal male; D, lateral male; E, female head front; F,
male vestiture on metasomal sterna.
146 K.L. Walker
Urs
Fil
0.5 mm
ae
Figure 7. A-F Lasioglossum (Australictus) peraustrale: A, dorsal female; B, lateral female; C, dorsal male; D, lateral male; E, female head front;
F, male vestiture on metasomal sterna.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 147
Figure 8. A-F Lasioglossum (Australictus) plorator: A, dorsal female; B, lateral female; C, dorsal male; D, lateral male; E, female head front; F,
male vestiture on metasomal sterna.
148 K.L. Walker
Figure 9. A-F Lasioglossum (Australictus) tertium: A, dorsal female; B, lateral female; C, dorsal male; D, lateral male; E, female head front; F,
male vestiture on metasomal sterna.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 149
Figure 10. Lasioglossum (Australictus) male genital capsules: Lasioglossum davide, A, ventral view, B, dorsal view; Lasioglossum lithusca, C,
ventral view, D, dorsal view; Lasioglossum peraustrale, E, ventral view, F, dorsal view.
150
K.L. Walker
Figure 11. Lasioglossum (Australictus) male genital capsules: Lasioglossum plorator, A, ventral view, B, dorsal view; Lasioglossum tertium, C,
ventral view, D, dorsal view.
Lasioglossum (Australictus) lithusca (Smith)
(Figs. IC, IE, 2A, 6A—F, 10C—D, 12B)
Parasphecodes lithusca Smith 1853: 41.
Parasphecodes adelaidae Cockerell 1905: 297. syn. nov.
Parasphecodes wellingstoni griseipennis Cockerell 1929: 11.syn. nov.
Parasphecodes stuchila Smith 1853: 43. syn. nov.
Parasphecodes wellingstoni Cockerell 1914: 45. syn. nov.
Lasioglossum (Parasphecodes) lithuscum — Michener 1965: 168.
Lasioglossum (Australictus) lithusca comb. nov.
Lasioglossum (Parasphecodes) adelaidae — Michener 1965: 167.
Lasioglossum (Parasphecodes) griseipenne — Michener 1965: 168.
Lasioglossum (Parasphecodes) stuchilum — Michener 1965: 168.
Lasioglossum (Parasphecodes) wellingtoni — Michener 1965: 169.
Material examined: Holotype of lithusca 7, Van Diemen's Land (ie.
Tasmania) F. Sm Coll. 79.22 BMNH Hym.17.a.618 (BMNH) (view
type data and image at https://data.nhm.ac.uk/object/9179e027-f637-
4d01-aff4-8cb0bc82d480 accessed 16 August 2022).
Holotype of adelaidae 9, South Australia: Adelaide BMNH
Hym.17.a.628 (BMNH) (view type data and image at https://data.nhm.
ac.uk/object/646ab614-8703-419a-9f3b-O805dbecac2e accessed 16
August 2022).
Holotype of griseipennis 9, New South Wales: Jenolan, 20 April
W.P. Cockerell (ANIC).
Holotype of stuchila 9, Van Diemen’s Land (ie. Tasmania) F. Sm
Coll. 79.22 BMNH Hym.17.a.613 (BMNH) (view type data and image at
https://data.nhm.ac.uk/object/2fel4c22-bacO-44f2-8f09-1250906cf baf
accessed 16 August 2022).
Holotype of wellingtoni 9, Tasmania, Mt Wellington Jan 15 —
Feb 6 1913, 1,300-2,000 ft R.E. Turner. Four females are mentioned
in the description but one specimen is labelled “type”. BMNH
Hym.17.a.636 (BMNH) (view type data and image at https://data.
nhm.ac.uk/object/3b470052-986f-43f2-a975-7dab4b48al2e accessed
16 August 2022).
Other specimens examined (2399, 418): QUEENSLAND: (19) Mt
Norman area, via Wallangarra, 7-8 Oct 1972, S.R. Monteith (ANIC);
(19) Mt Norman area, via Wallangarra, 2-3 Oct 1971, G.B. Monteith
(QM).
NEW SOUTH WALES & AUSTRALIAN CAPITAL
TERRITORY: (29) 3.19 km SW Condor Ck Bridge, Brindabella Rd, 06
Nov 1992, GJ. Davis (NMV); (29) 1.9 km S Condor Ck, Bridge,
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 191
Brindabella Rd, 17 Oct 1993, GV. Maynard & G.J. Davis, on Davesia sp.
(NMV); (19) Lees Camp, Warks Rd, 10 Dec 1993, G.J. Davis on
Leptospermum (NMV); (39) Bull's Head Repeater Station, 26 Nov
1994, GV. Maynard & G.J. Davis, on Pultenea and Davesia (NMV);
(19) Hill W side Condor Ck Bridge, 06 Nov 1993, GV. Maynard & G.J.
Davis on Davesia mimosoides (NMV); (19) Jink Ck, Blue Mtns, 21 Jan
1982, N.W. Rodd (AM); (12) Mt Boboyan 6 March 1993, G.J. Davis on
Kunzea ericoides (NMV); (4?) New England Nat. Pk, 5 Nov 1981, NW.
Rodd (AM); (19) 8 km S Mt Wilson, Blue Mtns, 10 Oct 1978, N.W. Rodd
(AM); (42) Mt Tomah Blue Mtns, 26-29 Aug 1977 — 2 Oct 1977, N.W.
Rodd (AM); (39) Clyde Mt, 27 Oct 1968, (ANIC); (29) Picadilly Circus,
Brindabella Ra., 24 Nov 1981, J.C. Cardale (ANIC); (19) Blundell’s 15
Oct 1952, Paramonov (ANIC); (12) Blundell’s, 14 Oct 1947, E.F. Riek
(ANIC); (19) Blundell’s, 13 Feb 1950, E.F. Riek (ANIC); (19) Blundell’s,
10 Oct 1930, L.F. Graham (ANIC); (19, 15) Mt Gingera, 6 Feb 1951,
E.F. Riek (ANIC); (39) Winburndale, 12 mi E Bathurst, 6 Dec 1974
(ANIC); (25) Bendora, 6 Feb 1952, E.F. Riek (ANIC); (12) Brindabella,
24 Nov 1931, L.F. Graham (ANIC); (12) Bull’s Head, 26 Feb 1952, L.J.
Nick (ANIC); (12) Mt Tomah, Blue Mtns, 5 Nov 1980, N.W. Rodd (AM);
(309) Picadilly Circus, Brindabella Ra.,1 Nov 1988, R.R. Snelling & J.
Grey #88 (LACM); (2%) Mt Victoria, 20 Oct 1930, A.N. Burns (NMV);
(102) NE Nat. Pk, near Entrance, 04-20 Nov 1960, C.W. Frazier (ANIC);
(1%) New England Nat. Pk, via Ebor, 22-23 Jan 1966, B. Cantrell (QM);
(19) 15 km SW Ebor, 12 Dec 1984, D. Yeates, on Leptospermum
mrytifolium (ANIC); (19, 1:5) Nadgee Reserve, 4 km N Newton’s Beach,
29 Dec 1985, E.A. Sugden (AM); (15) Guthega Pondage, Kosciusko
Nat. Pk, 16 Mar 1983, G.R. Brown & A.E. Westcott by sweeping (OAI).
VICTORIA: (119) Warburton, 19 Dec 1968, E.M. Exley, on
Leptospermum ericoides (QM); (1%) Halls Gap, 1 Nov 1928, GB
(SAM); (19) Black Sands, 16 Oct 1936 (ANIC); (19) Brisbane Ranges,
Oct 1982, P. Bernhardt, on Acacia (NMV); (12) Mason’s Ck, Kinglake,
20 Oct 1976, A.A. Calder (NMV); (39) Gorae West, 7 Oct 1957, A.N.
Burns (NMV); (189) Gorae West, 10 Jan 1950, T. Rayment (ANIC);
(12) Cockatoo, 24 Dec 1950, RD (NMV); (19) Arthur Plain, 4 Feb 1965,
Neboiss (NMV); (19) Ferntree Gully, 1909, F.P. Spry (ANIC); (49)
Grampians, Oct 1928, F.E. Wilson (NMV); (12) Reefton, 9 Feb 1955,
Neboiss (NMV); (19) Brighton (NMV); 239, 31%) Mt Hotham, 18
March & 6 April 2021, K. Walker, on Achillea millefolium (NMV), (29)
Portland, Dec 1979, G. Knerer (NMV); (19) Asgard Swamp, 4 km NE
Victoria, 10 Oct 1981, G.R. Brown (OAD; (19) Mt Drummer, 4 Dec
1956, E.F. Riek (ANIC); (29) Mt Difficult, Grampians, 2600 ft, 2 Jan
1966, B. Cantrell (QM); (19) Mt St Bernard, 4000 ft, 9 Jan 1932, A.N.
Burns (NMV); (39) Flowerdale, 8 Dec 1954, Neboiss (NMV); (19) Mt
Donna Buang, 5 Feb 1955, Neboiss (NMV); (19) Ferntree Gully,
1 Nov 1911, A.D.D (NMV); (15) Crystal Brook, Mt Buffalo, 24 Feb
1955, A.N. Burns (NMV); (19) 5 km W Toolangi, 7 Nov 2000, Thomas
Walker, on Ranunculus lappaceus (NMV ).
TASMANIA: (19) Huon Camping Area, 25 Jan 1983, I.D.
Naumann & J.C. Cardale (ANIC); (39) Mt Wellington, 2,500 ft, Feb
1936, A.J. Turner (QM); (19) Mt Field Nat Pk, 13 Jan 1997, A. Hingston
(NMV); (19) 5 km WNW Mt Alma, 27 Jan 1981, I.D. Naumann & J.C.
Cardale (ANIC); (2?) SW Nat. Pk, (Tyenna) 11 Jan 1997, A. Hingston
(NMV); 29) Tasmania, A. Simson (SAM); (59) 6 mi SW Queenstown,
|] Jan 1969, E.M. Exley on Melaleuca (QM); (4?) 11 mi E Strahan, 1 Jan
1968, E.M. Exley, on Melaleuca (QM); (62) 9 mi E Strahan, 1 Jan 1969,
E.M. Exley, on Bursaria (QM); (29) just N of junction of Waratah &
Murchison Hwys, 31 Dec 1968, E.M. Exley (QM); (1?) nr Nunamarra,
9 Jan 1969, E.M. Exley, on Melaleuca (QM); (49) Marrawah, 30 Jan
1949, E.F. Riek (ANIC); (19) Southport, 18 Jan 1948, E.F. Riek (ANIC);
(1?) Meredith R, 12 mi from Corinna, 6 Jan 1954, T.G. Campbell
(ANIC); (19) Derwent R, 740 m Lake St Clair, 24-28 Jan 1980,
Lawrence & Weir (ANIC); (19) 7 km SWW Derwent Bridge, 16 Jan — 2
Feb 1983, I.D. Naumann & J.C. Cardale, ex pantrap (ANIC); (12) 7 km
S Frodshams Pass, 25 Jan 1983, I.D. Naumann & J.C. Cardale (ANIC);
(19) Lake St Clair, 15 Apr 1955, T. Rayment (ANIC); (19) Lake St Clair,
alt. 2,500 ft, 08 Apr 1956, T. Rayment (ANIC); (1?) Collinsvale Road,
6 Jan 1972, W.F. Calvert (TDA); (19) Scarlet Ck, 3 km W Collingwood
R,8 Jan 1992, G. & A. Daniels (QM); (19, 15) 3.5 km SE Murdunna, 10
Feb 1988, G. & A. Daniels (QM); (12) 22 km N Dunkeld, 6 Nov 1988,
R.R. Snelling & J. Grey (LACM); (8?) Hasting, 15 Jan 1949, E.F. Riek
(ANIC); (19) Hobart, 4 March 1989, K. Walker, on Eucalyptus (NMV);
(12) Melaleuca nr Bathurst Harbour, 9 Nov 1991, I. Naumann & G.
Clarke, heathy sedge land & closed forest margin (ANIC); (19)
Melaleuca nr Bathurst Harbour, 3—7 Dec 1990, I. Naumann, margin
Melaleuca / Leptospermum (ANIC); (29) Celery Top Is., Bathurst
Harbour, 4 Dec 1990, I.D. Naumann (ANIC); (1?) Claytons, Bathurst
Harbour, 6 Dec 1990, I.D. Naumann (ANIC); (19) Railton, 8 Jan 1991,
B.B. Lowery, in rotten log (ANIC); (19) Liffey Valley, May 1980, S.
Fearn, ex alcohol storage (ANIC); 2?) 9 km WSW Derwent Bridge, 21
Jan 1983, I.D. Naumann & J.C. Cardale (ANIC); (29) Mt Wellington, 6
Jan 1918, G.H. Hardy (QM); (4?) Hobart, 10 Dec & 20 Dec 1913, G.H.
Hardy (QM); (19) Lake St Clair, 13 Jan 1937, C. & C. Davis (AM); (19)
Pieman Bridge, 8 Jan 1937, C. & C. Davis (AM); (19) Dover, 1963, from
sirex emergence hole (QM ).
SOUTH AUSTRALIA: (19) Adelaide (BMNH).
Floral record: Families visited: 6 (Asteraceae (2), Fabaceae (4),
Orchidaceae (1), Myrtaceae (11), Pittosporaceae (2), Ranunculaceae
(1)). Genera visited: 10 (Acacia (1), Achillea (2), Brenesia (1), Bursaria
(2), Davesia (2), Kunzea (1), Eucalyptus (3), Leptospermum (3),
Pultenaea (1), Ranunculus (1)).
Flight phenology capture records: Jan (27) Feb (14) Mar (7) Apr (3)
May (1) June (0) July (0) Aug (1) Sept (0) Oct (17) Nov (14) Dec (14).
Diagnosis. Lasioglossum (Australictus) lithusca 1s unlike any
other Australictus species 1n body colour. This species can be
distinguished, in both sexes, from other Australictus species by
red-brown-coloured metasoma contrasting with black
mesosoma; lack of tomentum on mesosoma or metasoma; dorsal
surface of metapostnotum posterior margin acarinate but dorsal
surface elevated above surrounding lateral and vertical surfaces;
metapostnotum dorsal surface sculpture ruguloso-striolate
(figs. 6A-D); male genitalia with gonobase narrowed basally,
large retrorse lobes and small, erect gonostylus (figs. I0C—D);
and metasomal sternal S5 lateral margins with distinct white,
semi-erect hair tufts (fig. 6F). This species occurs in SE Australia,
including Tasmania (fig. 12B).
Description of female: (figs. 6A, 6B, 6E) body length: 9.11—
10.49-10.83 mm (n=10); forewing length: 2.73—2.76—2.78 mm
(n=10); head width: 2.64—2.72—2.78 mm (n=10); intertegular
width: 1.92—2.04—2.06 mm (n=10). Relative head measurements:
HW: 100, HL: 88-90, UID: 56-57, LID: 53-54, IAD: 10-11,
OAD: 22-23, IOD: 14-15, OOD: 14-15, CL: 23-25, GW: 21-
23, EW: 21-23, SL: 39-40, FL: 71-73.
Head: (fig. 6E) inner eyes weakly narrowed basally; median
frontal carina reaching about than one third way to median
ocellus; clypeus smooth, polished with microtellesate sculpture
on basal one third, medially flattened to weakly concave, closely
punctate with large, deep, rounded punctures, medially with
vertical grooves, supraclypeal area distinctly raised above
paraocular area, surface dull, dense microtessellate pattern,
open to sparsely punctate with shallow punctures; frons dull,
close, vertical striae and punctate, paraocular area smooth,
shining closely punctate.
152 K.L. Walker
at
E
Figure 12. Species distribution maps: A, Lasioglossum (Australictus) davide; B, Lasioglossum (Australictus) lithusca; C, Lasioglossum (Australictus)
peraustrale: D, Lasioglossum (Australictus) plorator; E, Lasioglossum (Australictus) tertium; F, Lasioglossum (Chilalictus) orbatum.
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 193
Mesosoma: (fig. 6A) mesoscutum anterior mesial margin
weakly produced mesoanteriorly, surface dull, with minute
microtessellate pattern, medially openly to closely punctate,
laterad of parapsidal areas open to sparsely punctate, in
parapsidal areas densely punctate, densely punctate along
posterior margin; scutellum 2 x longer than dorsal surface of
metapostnotum, scutellum smooth, shining, openly to sparsely
punctate; dorsal surface of metapostnotum acarinate across
posterior margin defined by crescent shaped ridge distinctly
raised above vertical and lateral surfaces, dorsal surface
ruguloso-striolate, sculpture reaches posterior margins, lateral
margins smooth, dull with microalveolate pattern, vertical
posterior surface of metapostnotum defined by lateral carinae;
mesepisternum and metepisternum plicate; first recurrent vein
(Im-cu) entering second submarginal cell.
Metasoma and legs: (figs. 6A-B) metasomal T1- T5 with
dull sheen, appears impunctate, sparsely punctate with minute
punctures; anterior metatibial spur finely serrate with distinct,
small, backwardly pointing teeth.
Colour: (figs. 6A-B) head and mesosoma black, some
specimens apical half of clypeus red-brown, T1-T3 red-brown,
T4-T5 red-brown suffused with black; legs with fore, mid
coxae, trochanters, femora black, remainder red-brown.
Vestiture: (figs. 6A-B, 6E) body sparse, clypeus and
supraclypeal area glabrous, frons with sparse small, black, erect
hair, paraocular area with downwardly pointing white hair,
mesoscutum and scutellum appear glabrous with sparse small,
black, erect hair, long, white hair on lateral, vertical posterior
surface of metapostnotum, metasoma sparse, laterally with short,
erect white hair.
Description of male: (figs. 6C-D) Body length: 8.95—10.08—
10.21 mm (n=10); forewing length: 2.69-2.81-2.98 mm (n=10);
head width: 2.35—2.61—2.69 mm (n=10); intertegular width: 1.92—
2.07-2.16 mm (n=10). Relative head measurements: HW: 100,
HL: 88-89, UID: 56-57, LID: 42-43, IAD: 12-13, OAD: 22-23,
IOD: 16-18, OOD: 13-14, CL: 25-26, GW: 16-18, EW: 28-29,
SL: 22-23, AF4/AF2+3 (24/15, 22/14) 1.57—1.60, FL: 248—252.
Differs from female as follows: upper and lower inner eye
margins converging basally; median frontal carina extending
just beyond antennal bases; frons sculpture densely reticulate
across surface to inner margins of eyes; scape reaches basally
level of median ocellus; clypeus surface shiny with weak
microtessellate pattern basally, medium area weakly concave,
openly punctate with shallow punctures, supraclypeal area
protruding above paraocular area, bulbous, dull, with dense
microtessellate sculpture, closely punctate with minute, shallow
punctures; mesoscutum dull, with dense microtessellate
sculpture, closely to densely punctate, scutellum and metanotum
similar to mesoscutum; dorsal surface of metapostnotum same
as in female, acarinate, dorsal surface defined by raised ridge,
dorsal surface coarsely ruguloso-striolate; colour similar to
female except apical two thirds of clypeus pale yellow, metasoma
colour ranges from TI-I3 red-brown, T4-I6 red-brown
suffused with black, or T1 almost black, T2—T3 red-brown with
median black band, T4-T6 almost black.
Vestiture: lower half of frons, between antennal bases and
paraocular area with dense, erect, white hair, gena with dense
beard of long, plumose, white hair, mesoscutum with sparse,
short, erect, brown hair, metapostnotum lateral sides with
short, white, adpressed hair, apical posterior vertical posterior
surface of metapostnotum glabrous, metasomal sterna SI-S4
with sparse white, adpressed hair, S5 lateral margins with
distinct white, semi-erect hair tufts (fig. 6F).
Genitalia: (figs. 10C, 10D) gonobase sides narrowed
basally, complete ventroapically, gonobase width almost half
of gonocoxa width, gonocoxa glabrous ventrally and dorsally,
dorsal inner margins of gonocoxa basally rounded, gonocoxa
apical inner margin not produced, continues contours of
sonostylus, glabrous; retrorse lobes large, meeting at midline,
membranous, inwardly pointing, outer margins of retrorse
lobes glabrous, inner margins basally with erect, large setae,
apically with short adpressed, erect setae; gonostylus small
(about one third length of gonocoxa), erect, apically rounded,
with simple, erect hair; penis valves curved apically, with
short dense hair dorsolaterally.
Distribution: (fig. 12B) this species occurs from SE Queensland,
down New South Wales coast, across Victoria, throughout
Tasmania and one South Australian record. It has been recorded
from sea level (Portland 7.23 m) up to high elevations in Victoria
(Mt Hotham 1,861 m) and Tasmania (Mt Wellington 1,271 m).
Remarks: Australian Halictidae species now in the genus
Lasioglossum were originally placed in either Halictus Latreille
1804 or Parasphecodes, and most original species names ended
with “us” or “a”. Michener’s (1965) revision of the Australian bee
fauna recombined these species into the genus Lasioglossum. In
accordance with ICZN rules (Articles 31 and 34), Michener
changed species name endings to agree with the gender of
Lasioglossum and endings with "us" or "a^ were changed to
“um”. Therefore, Michener 1965 changed Parasphecodes
lithusca to Lasioglossum (Parasphecodes) lithuscum.
John Ascher (pers. comm.) argued that some of these
names should not have been changed. John reasoned that 12
of Smith's (1853) new species names were partial anagrams of
the word "Halictus", meaning these species names should be
treated as nouns in apposition rather than adjectival and
therefore should not have been changed in Michener's (1965)
revision. Accordingly, Michener's (1965) L. (Parasphecodes)
lithuscum name has been reverted to Smith's (1853) original
species name of L. (Australictus) lithusca.
Cockerel (1914: 46) commented that Parasphecodes
wellingtoni was close to Parasphecodes lithusca but differed in
leg colour; that colour difference is within the known colour
range. Cockerell (1929: 11) commented that Parasphecodes
wellingtoni griseipennis was “typical” of Paraspheodes
wellingtoni but that species occurred in Tasmania, whereas the
new subspecies “griseipennis” occurred in New South Wales.
Lasioglossum (Australictus) lithusca is now recorded from SE
Queensland across to South Australia, including Tasmania.
several female specimens carried deutonymphal Anoetus
Dujardin 1842 (Sarcoptiformes: Astigmatina: Histiostomatidae)
hypopial mites apically across Tl, though one specimen had
several mites on the right forewing. No acarinarium was present
on T1, but this area has a moderate cover of erect setae. See
Walter et al. (2002) for full discussion of mites on bees.
154
Rayment identified a specimen as this species, his
handwritten label reading “Bolgart WA, 12 Jan 1950”. This is
the only specimen of this species and the entire subgenus
recorded from Western Australia. I consider it to be an
incorrect labelling, because Rayment also collected and
labelled, in his handwriting, specimens of L. lithusca at Gorae
West, Victoria on 10 Jan 1950. I have not included Rayment’s
WA specimens in the “Other Material Examined” or on the
species distribution map.
Although this species was originally described in
Parasphecodes by Smith (1853) and placed ın Lasioglossum
(Parasphecodes) by Michener (1965), it is better placed in
Lasioglossum (Australictus) due to the presence of all the
Australictus subgeneric diagnostic characters, especially the
elongated and enlarged preapical tooth on the mandible, finely
serrate anterior metatibial spur, shape of labrum and male
genitalia characters.
Lasioglossum (Australictus) peraustrale (Cockerell)
(Figs. 2C, 7A—F, IOE, LOF, 12C)
Halictus peraustralis Cockerell 1904: 211.
Halictus odyneroides Rayment 1939: 279. syn. nov.
Lasioglossum (Australictus) peraustrale — Michener 1965: 165.
Lasioglossum (Australictus) odyneroides — Michener 1965: 165.
Material examined: Holotype of peraustralis 9, South Australia.
“bicingulatus var. Smith” F. Sm. Coll. 79.22 BMNH Hym.17.a.693
(BMNH) (view type data and image at: https://data.nhm.ac.uk/object/
c23639db-ald1-40e5-886a-ef654e4d3c8f accessed 16 August 2022).
Holotype of odyneroides 9, New South Wales, White Swamp,
Macpherson Range, May 1939, J. Hardcastle. (ANIC).
Other specimens examined (2199, 473): QUEENSLAND: (119)
Kuranda, Black Mountain Road, 3 Oct 2005, K. Walker, on
Lophostemon grandifloris (NMV); (19) Cairns dist. F.P. Dodd (QM);
(349) Brisbane, 4 Jul 1914, 1 Sept 1914, 24 Sept 1914, 29 Sept 1914, 2
Feb 1918, 13 Mar 1918, 12 Feb 1918, 6 Mar 1918, H. Hacker (QM);
(1?) Mt Moffat Nat Pk, Kenniff Cave, 840 m, 22 Nov 1995, C. Burwell
(QM); (59) Stradbroke Is., 3 Dec 1912, 7 Sept 1914, 17 Sept 1915, H.
Hacker (QM); (19) Birkdale, 13 Mar 1918, H. Hacker (QM); (25)
Cairns district, F.P. Dodd (SAM); (28) Upper Cedar Creek via
Samford, 21 Nov 1965, B. Cantrell (QM); (34) Cedar Ck, 21 Nov
1965, T. Weir (QM); (15) Brisbane, 17 Apr 1955, J. Kerr (NMV); (25)
5 km N Karara, 6 Jan 1984, K. Walker on Eucalyptus (NMV); (15)
Murphy's Creek, 30 Nov 1988, K. Walker, on Eucalyptus (NMV); (2%,
14) Leslie Dam, 13 km W Warwick, 13 Nov 1978, K. Walker, on
Eucalyptus (QM); (15) Broken R, Eungella, 16-17 Nov 1992, 750 m,
Monteith, Thompson, Cook & Janetzki (QM); (12, 35) 3 mi
Cunningham's Gap, 25 Feb 1959, C.D. Michener on Bursaria spinosa
(SEM); (14) Tambourine, 18 Dec 1958, C.D. Michener (SEM); (19)
Tambourine, 1923, W.H. Davidson (QM); (2%) Amiens, 4 Nov 1965,
J.C. Cardale (QM); (19) Severnlea via Stanthorpe, 10 Dec 1980, E.M.
Exley & J. King on Leptospermum (QM); (29) 11 km S Cunningham's
Gap, 12 Nov 1980, J. & C. King, on Bursaria spinosa (QM); (12) Lucas
Heights, 5 Nov 1995, A. Sundholm, on Leptospermum polygalifolium
(QM); (29) Springbrook, 12 Feb 1943, A.J. Turner (QM); (29) 8 mi N
of Landsborough, 28 Oct 1965, J.C. Cardale (QM); (12) Capalaba, 23
Sept 1961, R. Shepard (QM); (2?) Burleigh, 28 Sept 1958, A.N. Burns
(NMV); (19) 13 mi N Stanthorpe, 29 Dec 1958, C.D. Michener (SEM).
NEW SOUTH WALES & AUSTRALIAN CAPITAL
TERRITORY: (239) In rotten log, Bankstown nr Sydney, 30 June 1984,
K.L. Walker
B.J. Day (AM); (339, 15) Bilpin Blue Mtns, 11 Mar 1981, 28 Feb 1986,
10 Sept 1979, 14 Nov 1977, 22 Sept 1977, 19 Apr 1978, 21 Nov 1978,
2 Oct 1978, N.W. Rodd (AM); (19) Kurrajong Heights, 27 Feb 1978
(AM); (19) Woronara, 12 Jan 1982, M.L. Mason (AM); (45) Castle Flat,
Clyde R, 4.5 km W Pigeon House, 8 Jan 1984, L. Hill, ex cluster c. 150
on twigs (AM); (16) Liston, 23 Dec 1969, J.C. Cardale, on Eucalyptus
(ANIC); (19) National Park, 23 Dec 1906, C. Gibbons (AM); (19)
Patonga, 25 Nov 1945 (AM); (339,25) Cheltenham, 22 Oct 1949, 1 Apr
1950 (AM); (19) Approx. 1 km S Kew, 23 Dec 1981, G. & T. Williams,
on Leptospermum blossoms (AM); (15) Brunswick Heads, 12 Jan 1938,
E.F. Riek (ANIC); (15) Hornsby, C. Gibbons (AM); (99, 15) Nadgee
Reserve, 7 km S of Newton’s Beach, 21 Dec 1985, E.A. Sugden (AM);
(13') Gosford, 13 Mar 1932, A.J. Turner (QM); (19) Mountain Lagoon,
Blue Mtns, 23 Nov 1977, NW. Rodd (AM); (49) Sydney, W.W, Froggatt
(ANIC); (19) Pearl Beach, near Woy Woy, 9-11 Dec 1988, M.J. Fletcher
& J.A. MacDonald (OIA); (19) nr Glenbrook Ck, Blue Mtns, 23 Dec
1998, G.R. Brown (OIA); (1?) Lane Cove, 26 Sept 1987, S.G. Hunter
(OIA); (12) Georges River, nr Lugarno, 16 Apr 1941, A. Holmes (AM);
(12) Barrington House via Salisbury, 25-28 May 1963, A. Macqueen
(QM); (39) State Forest Gibraltar Range, 29 Dec 1969, C.W. Frazier
(ANIC); (19) Nelligen, 3 Nov 1949, Cane & Gemmell (ANIC); (19)
Jervis Bay, 7 Nov 1956, E.F. Riek (ANIC); (19) 2.7 km NE Queanbeyan,
670 m, 16 Dec 1979, I.F.B. Common (ANIC); (19) 3 km N Lansdowne,
nr Taree, 3 Jan 1992, G. Williams on Tristaniopsis laurina blossoms
(NMV); (29) Royal National Park, 28 Dec 1970, D.K. McApline (AM);
(19) Black Mountain, 15 Jan 1934, FJ. Gray (ANIC); (29) Wisemans
Ferry, 18 Dec 1927, A.N. Burns (NMV); (19) Buchan, 26 Jan 1937, A.N.
Burns (NMV); (19) Mt Wilson, 4 Jan 1931, A.N. Burns (NMV); (19)
Wattle Flat, W.W. Frogeatt (ANIC).
VICTORIA: (29, 15) N of Lakes Entrance, Colquhoun State
Forest, 21 Feb 1985, 6 Feb 1987, K. Walker, on Eucalyptus (NMV);
(6:5) 9 km N Bruthen, 8 Feb 1992, G. Daniels & C. Burwell (QM);
(39) Coranderrk, 16 Nov 1984, P. Bernhardt on Acacia mearnsii
(NMV); (29) Woori Yallock, 23 Nov 1930, A.N. Burns (NMV); (49)
Tambo Crossing, Jan 1935, F.E. Wilson (NMV).
Floral record: Families visited: 3 (Fabacaea (1), Myrtaceae (10),
Pittosporaceae (2)). Genera visited: 6 (Acacia (1), Bursaria (2),
Eucalyptus (4), Lophostemon (1), Leptospermum (2), Tristaniopsis (1)).
Flight phenology capture records: Jan (27) Feb (14) Mar (7) Apr (3)
May (1) June (0) July (0) Aug (1) Sept (0) Oct (17) Nov (13) Dec (14).
Diagnosis. Lasioglossum (Australictus) peraustrale 1s most like
L. (Australictus) tertium with pronotum and metasomal
tomentous yellow hair bands. This species can be distinguished in
both sexes from other Australictus species by black body colour;
tomentum on pronotum, posterolateral corners of mesoscutum,
metanotum and metasomal bands on T2-T3; dorsal surface of
metapostnotum posterior margin acarinate but elevated above
surrounding lateral and vertical surfaces; metapostnotum dorsal
surface sculpture microalveolate; metapostnotum vertical surface
with V-shaped patch of white tomentum (weak in female and
dense in male) (figs. 2C, 7A—D); male genitalia with gonobase
narrowed basally, large retrorse lobes and gonostylus absent
(absent gonostylus 1s shared with L. tertium only; figs. LOE, 1OF,
11C, 11D); and metasomal sternal S5 lateral margins with weak,
white, semi-erect hair tufts (fig. 7F). This species occurs in North
Queensland but is primarily found in SE Australia, and is absent
from Tasmania (fig. 12C).
Description of female: (hgs. 7A, 7B, 7E) body length: 8.32—
9.59-10.05 mm (n=10); forewing length: 2.35—2.89—3.07 mm
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 155
(n=10); head width: 2.54—2.76—2.83 mm (n=10); intertegular
width: 1.68-1.83-1.92 mm (n=10). Relative head measurements:
HW: 100, HL: 80-81, UID: 58-59, LID: 56-57, IAD: 09-10,
OAD: 21-22, IOD: 12-13, OOD: 16-17, CL: 21-22, GW: 20-
21, EW: 23-25, SL: 41-42, FL: 62-65.
Head: (fig. 7E) distance between inner upper and lower
inner eye margins almost parallel; median frontal carina
reaching about half way to median ocellus; clypeus smooth,
polished with minute microtesellate sculpture, medially
flattened, closely to sparsely punctate with shallow, rounded
punctures, medially several punctures; supraclypeal area
slightly raised above paraocular area, surface with dull sheen,
with weak microtessellate pattern, open to closely punctate
with shallow punctures; frons dull, with dense reticulate
punctures which extend to inner margins of eye, paraocular
area shining, close to densely punctate.
Mesosoma: (fig. 7A) mesoscutum anterior mesial margin
rounded, continuing lateral contour, surface with dull sheen,
with dense punctures, many punctures fused; scutellum 1.35 x
longer than dorsal surface of metapostnotum, scutellum dull,
with densely punctate sculpture; dorsal surface of metapostnotum
acarinate across posterior margin but defined by V-shaped ridge,
distinctly raised above vertical and lateral surfaces, dorsal
surface microalveolate but with several weak rugulose striae
basally, striae not reaching mid-point of dorsal surface, dorsal
surface pattern forming V-shape onto vertical posterior surface,
lateral margins smooth, dull with microalveolate pattern, with
some large, shallow punctures, vertical posterior surface of
metapostnotum microalveolate with carinae extending basal one
third of length; mesepisternum and metepisternum plicate; first
recurrent vein (Im-cu) interstitial with or entering second
submarsinal cell.
Metasoma and legs: (figs. 7A-B) metasomal TI-T5 with
dull sheen, densely punctate across entire surfaces; anterior
metatibial spur finely serrate to smooth.
Colour: (figs. 7A, 7B, 7E) body black except: apical two
thirds of clypeus, lower paraocular area, antennae red-brown,
T4-T5 red-brown, coxa, trochanter dark brown to black,
remainder of legs red-brown; coloured dense patches of vestiture
as follows: dense white vestiture around spiracle, yellow to
orange on pronotum corners, lateral corners of mesoscutum,
mesial of metanotum, dense tomentum basally across T2-T3, T3
tomentum thicker than tomentum on T2, T4-T5 brown; forewing
with brown tinge from submarginal cells to tip of wing.
Vestiture: (figs. 7A, 7B, 7E) body vestiture sparse, clypeus
with sparse simple hair, supraclypeal area almost glabrous,
frons with sparse, small, black, erect hair, paraocular area
with downwardly pointing adpressed, white branched hair,
mesoscutum, scutellum almost glabrous but with tomentum
on posterolateral corners of mesoscutum, metanotum, basally
on T2-T3, sparse small, black, erect hair, long, white hair on
lateral, vertical posterior surface of metapostnotum, except
for tomentum bands, metasomal terga appears glabrous but
with sparse, short, adpressed hair.
Description of male: (figs. 2C, 7C, 7D) body length: 8.01—9.25—
9.42 mm (n=10); forewing length: 2.50—2.54—2.59 mm (n=10);
head width: 2.06—2.33—2.40 mm (n=10); intertegular width: 1.52—
1.85-1.87 mm (n=10). Relative head measurements: HW: 100,
HL: 90-92, UID: 57-58, LID: 42-43, IAD: 16-17, OAD: 23-24,
IOD: 15-16, OOD: 16-17, CL: 26-27, GW: 15-16, EW: 29-30,
SL: 22-23, AFA/AF243 (23/15, 22/14) 1.53—1.57, FL: 215—220.
Differs from female as follows: upper and lower inner eye
margins converging basally; median frontal carina extending
about one third to two thirds to medium ocellus; frons
sculpture densely reticulate-punctate across surface to inner
margins of eyes; scape reaches level of median ocellus;
clypeus surface polished, shiny, weakly convex, openly
punctate with shallow punctures; supraclypeal area weakly
protruding above paraocular area, rounded, polished, closely
punctate with minute, shallow punctures with microalveoate
sculpture; mesoscutum as in female, dull, densely punctate
appearing as coarse reticulate sculpture, scutellum,
metanotum similar to mesoscutum; dorsal surface of
metapostnotum same as in female though reticulation more
extensive, acarinate, dorsal surface defined by raised, rounded
ridge, dorsal surface with microtessellate sculpture but
without weak striae basally; colour similar to female except
apical two thirds of clypeus pale yellow, with similar yellow-
orange vestiture except V-shaped tomentum on upper vertical
posterior surface of metapostnotum (fig. 2C), T3 basal
tomentum much broader than basal tomentum on T2.
Vestiture: lower half of frons, between antennal bases,
paraocular area with dense erect, white hair, gena with
moderate beard of long, plumose, white hair, mesoscutum
appears glabrous but with sparse, short, erect, brown hair,
metapostnotum lateral sides with short, white, adpressed hair,
apical vertical posterior surface of metapostnotum with
V-shaped tomentum of white hair (fig. 2C), metasomal sterna
SI-S4 with sparse white, adpressed hair, S5 lateral margins
with weak white, semi-erect hair tufts (fig. 7F).
Genitalia: (figs. IOE-F) gonobase sides narrowed basally,
complete ventroapically, gonobase width almost half of
eonocoxa width, gonocoxa with single seta dorsolaterally,
remainder glabrous, dorsal inner margins of gonocoxa basally
rounded, gonocoxa apical inner margin not produced, continues
contours of gonostylus, glabrous; retrorse lobes large, meeting at
midline, membranous inwardly pointing, outer margins of
retrorse lobes with small, erect setae, inner margins of retrorse
lobes basally with area of erect, large setae, apically with fine,
short adpressed, some larger erect setae; gonostylus absent but
area with simple, erect hair; penis valves curved apically, with
short dense cover of hair dorsolaterally.
Distribution: (fig. 12C) this species extends from north
Queensland (Cairns), down eastern New South Wales and
across Victoria, but is absent from Tasmania. The type locality
of Halictus peraustralis is South Australia, but no location was
named so the South Australian record has not been mapped.
Remarks: Cockerell’s (1904) description of “Halictus
peraustralis" was unusual as the species description was part of a
dichotomous key to “Halictus specimens in the British Museum”,
with the species description of “Halictus peraustralis” as part of
couplet one. Cockerell noted (1904: 211) that the specimen used
for the species description (Type) was originally labelled by F.
Smith as a variety of “Halictus bicingulatus var Smith’. Note that
156
Michener (1965) placed “bicingulatus” in the Lasioglossum
subgenus Chilalictus, but Cockerell recognised the variety as a
valid species. Rayment (1939) commented that his new species
"odyneroides" belonged to the "bicingulatus" group and that it
was closest to “H. peraustralis”, but he distinguished it as a new
species due to a minor colour variation. This colouration 1s
recognised within L. peraustrale. The colour patterns of L.
peraustrale are best described as mimetic, because they are
replicated in a range of species in Lasioglossum subgenera.
Lasioglossum (Australictus) plorator (Cockerell)
(Figs. 2F, 3A-F, 4, SA-F, 11A, I IB, 12D)
Halictus plorator Cockerell 1910b: 274; Rayment 1953: 29.
Lasioglossum (Australictus) plorator — Michener 1965: 165.
Material examined: Holotype of plorator 9 ‚Victoria Melbourne Aug 1900,
C.F. Turner Collection 1900-7 C.7. 8.00 BMNH Hym.17.a.633 (BMNH)
(view type data and image at https://data.nhm.ac.uk/object/9154124c-377b-
4ea6-8421-90965bbc6284 accessed 16 August 2022).
Other specimens examined (2482, 1598): QUEENSLAND: (19)
Beechmont, 3 Oct 1984, N.W. Rodd (AM); (29) Wyberba National
Park, 8 Jan 1967 Houston, T.F. on Eucalyptus (WAM).
NEW SOUTH WALES & AUSTRALIAN CAPITAL
TERRITORY: (59) Blue Mtns, 26 Sept 1978, N.W. Rodd (AM); (2%,
15) Tianjara Falls, 60 km SW Tomerong, 5 Feb 1988, N.W. Rodd
(AM); (19) Orange, Jan 1934 (ANIC); (29, 15) 3 km S Mt Wilson,
Blue Mtns, 12 Sept 1978, NW. Rodd (AM); (15) Narrow Neck, Blue
Mtns, 27 March 1979, N.W. Rodd (AM); (19) Mt Victoria, Blue Mtns,
29 Dec 1981, NW. Rodd (AM); (25) 6 km NE Bilpin, Blue Mtns, 4
March 1986, NW. Rodd (AM); (39,18) Mt Tomah, Blue Mtns, 9 Feb
1986, NW. Rodd (AM); (19, 14) Mt York, Blue Mtns, 29 Jan 1982,
N.W. Rodd (AM); (19, 15) Haystack Ridge, 26 Feb 1979, N.W. Rodd
(AM); (19,15) 3 km S Mt Wilson, Blue Mtns, 13 Jan 1986, N.W. Rodd
(AM); (19,15) Mt Tomah, 25 Jan 1979, N.M. Rodd (AM); (19, 15)
Nadgee Reserve, 7 km S Newton Beach, 29 Dec 1985, E.A. Sugden,
sweeping Kunzea ercoides (AM); (27) 8 km W Tyalgum, 24 Sept
1983, N.W. Rodd (AM); (22) Mt Kaputar Nat. Park, 1362 m, 5 Dec
1974, 1.F.B. Common & G.E.D. Edwards (ANIC); (42) Dawson’s
Spring, Mt Kaputar Nat. Pk, 30 Nov — 10 Dec 1978, G.R. Brown
(OIA); (19) Armidale, 18 Nov 1959, C.W. Frazier (ANIC); (59) NE
Nat. Pk, 4 Nov 1960, C.W. Frazier (ANIC); (92) Blundell’s, 18 Feb
1931, L.F. Graham (ANIC); (19) Blundell’s, 18 Feb 1950, E.F. Riek
(ANIC); (15) Mt Wilson, 4 Jan 1931, A.N. Burns (NMV); Q9)
Winburndale, 12 mi E Bathurst, 6 Dec 1974 (ANIC); (29) Picadilly
Circus, Brindabella Ra.,1 Nov 1988, R.R. Snelling & J. Grey #88
(LACM); (39) NE Nat Pk, near Entrance, 04-20 Nov 1960, CW.
Frazier (ANIC).
VICTORIA: (39) Gorae West, 1951, 22 Aug 1956 (ANIC); (19)
Packenham, 20 Nov 1936 (ANIC); (29) Grampians, Oct 1928, F.E.
Wilson (ANIC); (1?) Grampians, 20 Oct 1945, A.N. Burns (NMV);
(39) Flowerdale, 15 Dec 1954, A.N. Burns (NMV); (39) Flowerdale, 8
Dec 1954, Neboiss (NMV); (1?) Melbourne, F.P. Spry 1909 (ANIC);
(19) Erica, 20 April 1983, P. Bernhardt (NMV); (7?) Warburton, 19
Dec 1968, E.M. Exley on Leptospermum ericoides (QM); (19)
Warburton, 19 Dec 1968, E.M. Exley on Prostanthera lasianthoe (QM);
(27) Portland, 6 Dec 1974, G. Knerer (NMV); (19) Macedon, 29 Feb
1967 (ANIC); (19, 53) Emerald, 26 July 1936, Rayment (ANIC); (19)
11 km Halls Gap, 21 Oct 1983, I.D. Naumann & J.C. Cardale (ANIC);
(12) Mt Difficult, Grampians, 2 Jan 1966, B. Cantrell (QM); (25)
Anglers Rest, 5 March 1992, K. Walker, on Eucalyptus (NMV); (22,18)
Reefton, 9 Feb 1955, Neboiss (NMV); (172) Croydon (NMV); (29)
K.L. Walker
Lake Hattah, 11 Apr 1920, J.E. Dixon (NMV); (19) San Remo, 17 Oct
1927, A.D.D. (NMV); (39) Hamilton, 14 Sept 1914, G.S. (NMV); (39)
Wombargo Ck, 9 May 1947, G.B. (NMV); (19) Grampians, 19 Oct
1945, G.B. (NMV); (19) High Tap, 7 Aug 1951 (NMV); (19) Lorne, 3
March 1954, F.E. Wilson, burrowing in punk of rotten log (NMV); (29)
Mt Dandenong, 2,000 ft, 21 Dec 1930, A.N. Burns (NMV); (19)
Dromana, 3 Nov 1931, G.B. (NMV); (2?) Warburton, 8 Sept 1959, A.N.
Burns (NMV); (29) Ferntree Gully, 5 Dec 1915, F.P. Spry (NMV); (19)
Nariel, 12 Feb 1963, A.N. Burns (NMV); (349) Cobboboonee State
Forest, 1.3 km E Wright’s Swarm Road, 3 March 1990, W.T. Wcislo ex
Brenesia (SEM); (379) Cobboboonee State Forest, 12-13 March 1990,
W.T. Wcislo, ex Eucalyptus (SEM); (37,95) 22 km N Portland farm, at
NW Border of Cobboboonee State Forest and Glenelg National Forest,
15 Feb 1990, WT. Wcislo ex red Eucalyptus (SEM); (119, 148)
Cobboboonee State Forest, 4 Feb 1997, K. Sparks and C. Mcphee on
Bursaria (NMV); (19) Victoria Valley, 11 Feb 1947, B. Given (NMV);
(19) 715 Lt Hampton Rd Glenlyon, Gayle Osborne, on Achillea
millefolium (1Naturalist https://ww w.inaturalist.org/
observations/68516396); (1?) Wilsons Promontory, 5 Mile Beach Road,
26 Feb 1996, K. Walker, on Eucalyptus (NMV); (39) Mt Hickey,
Tallarook, 14 Nov 1987, P. Carwardine (NMV).
TASMANIA: (19) 3 km SEE Black River, 18 Jan 1983, I.D.
Naumann & J.C. Cardale (ANIC); (29) 11 mi E Strahan, 1 Jan 1969,
E.M. Exley on Melaleuca (QM); (19) 10 mi W Upper Blessington, 29
Dec 1968, E.M. Exley, on Leptospermum (QM); (19) Tullah, 31 Dec
1968, E.M. Exley on Leptospermum (QM); (39) 1 km SSE Gladstone, 29
Jan 1983, I.D. Naumann & J.C. Cardale (ANIC); (19) The Lea, 6 km S
Hobart, 27 Dec 1979, J.C. Cardale (ANIC); (12) Hobart, Lea (SAM);
(29) Sandy Bay, 20 May 1947, A.N. Burns (NMV); (15) 7 km SW
Buckland, 27 Jan 1983, I.D. Naumann & J.C. Cardale (ANIC); (29)
Huon-Picton R Junction, 14 Nov 1972, A. Neboiss (NMV); (219)
Marrawah, 30 Jan 1949, E.F. Riek (ANIC); (19) Mt Claude, 680m, 21
March 1990, L. Hill, on Leptospermum (TDA); (29) Lake Leake,
2000 ft, 27 Feb 1963, I.F.B. Common & M.S. Upton (ANIC); (19)
10 km NNW St. Helens, 14 Jan 1983, I.D. Naumann & J.C. Cardale
(ANIC); (19) 2 km NNE Pioneer, 29 Jan 1983, I.D. Naumann & J.C.
Cardale (ANIC); (2?) Bruny I Lea (SAM); (14%) Hastings, 15 Jan 1949,
E.F. Riek (ANIC); (2?) SW Nat. Park, Tyenna, 11 Jan 1997, A. Hingston,
on Leptospermum lanigerum (TDA); (12) 4 km WSW Maydena, 11 Dec
1981, I.D. Naumann (ANIC); (29) Tyenna, 16 Dec 1917, C.E. Cole,
(SAM); (19) National Park, 21 Jan 1949, E.F. Riek (ANIC); (19, 1105)
Florentine Valley, 20 km W Maydena, 9 Feb 1986, K. Walker, on
Leptospermum (NMV); (19) Ellendale, 30 Jan 1973, R.J. Hardy (TDA);
(19) Mount Wellington, Icehouse Track, 27 Jan 2001, K. Hergstrom
(TDA); (19) Mt Wellington, Hobart, 8 Nov 1996, A. Hingston, on Hakea
lissosperma (TDA); (29) Mt Nelson, 31 Jan 1997, A. Hingston, on
Leptospermum scoparium (NMV); (19) Liffey: Fernery, 4 Dec 1993,
C.P. Spencer & L. Richards (TMAG); (19) 6 mi SW Queenstown, 1 Jan
1969, E.M. Exley, on Melaleuca (QM); (19, 25) 6 km N Zeehan, 6
March 1989, K. Walker, on Eucalyptus (NMV); (19, 15) Cradle
Mountain, 7 March 1989, K. Walker, on Leptospermum rupestre (NMV).
Floral record: Families visited: 4 (Asteraceae (1), Lamiaceae (1),
Myrtaceae (5), Proteaceae (1)). Genera visited: 5 (Achillea (1), Eucalyptus
(6), Hakea (1), Kunzea (1), Leptospermum (7), Prostanthera (1)).
Flight phenology capture records: Jan (18) Feb (14) Mar (7) Apr (3)
May (2) June (0) July (1) Aug (2) Sept (4) Oct (6) Nov (8) Dec (15).
Diagnosis. Lasioglossum (Australictus) plorator is most like L.
(Australictus) davide in body colour. This species can be
distinguished, in both sexes, from other Australictus species by
black body colour with bluish tinge on metasoma; lack of
tomentum on mesosoma or metasoma; dorsal surface of
metapostnotum posterior margin acarinate but surface elevated
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 157
above surrounding lateral, vertical surfaces, metapostnotum
dorsal surface sculpture microalveolate (figs. 2F, 8A—D); male
genitalia with gonobase narrowed basally, moderate sized retrorse
lobes, small, erect gonostylus (figs. 11A, 11B); and metasomal
sterna with erect lateral hair tufts on S4-S5 (fig. 8F). This species
occurs in SE Australia, including Tasmania (fig. 12D).
Description of female: (figs. 2F, SA, 8B) body length: 9.89—
10.12-10.21 mm (n=10); forewing length: 2.97-3.05-3.12 mm
(n=10); head width: 2.64-2.71-2.78 mm (n=10); intertegular
width: 1.87—1.98—2.06 mm (n=10). Relative head measurements:
HW: 100, HL: 82-83, UID: 60-62, LID: 59-60, IAD: 09-10,
OAD: 22-23, IOD: 08-09, OOD: 17-18, CL: 20-21, GW: 20—
21, EW: 20-21, SL: 43-45, FL: 70-72.
Head: (fig. 8E) upper and lower inner eye margins almost
parallel; median frontal carina barely reaching above upper
antennal bases; clypeus polished, smooth on apical half,
remainder with microtessellate sculpture pattern giving dull
sheen, surface weakly convex, openly punctate with shallow
punctures, few irregular, elongate punctures apically,
supraclypeal area distinctly raised above paraocular area,
surface dull with microtessellate sculpture pattern, sparsely
punctate with shallow punctures; frons sculpture above antennal
bases densely reticulate, laterally almost smooth but plicate with
weak, wavy, raised, vertical carinae extending laterally to inner
margins of eyes, apically to just basally level of median ocellus,
paraocular area smooth, shining, weakly plicate.
Mesosoma: (figs. 2F, SA-B) mesoscutum anterior mesial
margin weakly produced mesoanteriorly, surface smooth, with
dull sheen due to weak microtessellate sculpture pattern across
surface, medially open to closely punctate, laterad of parapsidal
areas closely punctate, in parapsidal areas densely punctate,
densely punctate along posterior margin; scutellum 1.6 x length
of dorsal surface of metapostnotum, scutellum with dull sheen,
with microtessellate sculpture pattern, open to closely punctate;
dorsal surface of metapostnotum acarinate across posterior
margin with weak carinae in posterolateral corners, dorsal
surface defined by rounded, raised ridge above vertical, lateral
surfaces, dorsal surface with microalveolate sculpture but with
several weak striae basally, striae not reaching mid-point of
dorsal surface, lateral margins smooth with microalveolate
pattern, vertical posterior surface of metapostnotum defined by
lateral carinae (fig. 2F); mesepisternum and metepisternum
plicate; first recurrent vein (lm-cu) meeting Irs-m vein or
entering second submarginal cell.
Metasoma and legs: metasomal TI-T5 shining, smooth,
sparsely punctate with minute punctures; anterior metatibial
spur finely serrate, teeth barely discernible.
Colour: (figs. 8A, 8B) body black except scapes dark
brown, flagella segments light brown, metasomal tergal
segments black with bluish tinge, legs dark brown.
Vestiture: (figs. 8A, 8B, 8E) body vestiture sparse, clypeus,
supraclypeal area almost glabrous, frons with sparse, small,
black, erect hair, mesoscutum, scutellum almost glabrous but
with sparse, small, black, erect hair; long hair on lateral,
vertical posterior surface of metapostnotum, apical one third
of Tl with some long, erect, hair, remainder of TI, T2
glabrous, T3-I4 with some black, adpressed hair apically.
Description of male: (figs. 8C, 8D) body length: 8.48—9.98—
10.99 mm (n=10); forewing length: 2.64-3.08-3.12 mm (n=10);
head width: 2.26-2.57-2.64 mm (n=10); intertegular width:
1.58—2.07—2.16 mm (n=10). Relative head measurements: HW:
100, HL: 83-85, UID: 60-62, LID: 47-48, IAD: 09-10, OAD:
201, IOD: 15-16, OOD: 16-17, CL: 27-28, GW: 19-20, EW:
28-29, SL: 19-20, AFA/AF243 (21/14, 22/15) 1.46-1.50, FL:
210—214.
Differs from female as follows: upper and lower inner eye
margins converging basally; median frontal carina reaching
about one quarter to median; frons sculpture reticulate across
surface to inside inner margins of eyes; scape almost reaches
level of median ocellus; clypeus smooth, shiny, with weak
microtessellate pattern basally, medium area rounded, open to
closely punctate; supraclypeal area protruding above paraocular
area, surface dull with microtessellate sculpture covering entire
surface, openly punctate; mesoscutum surface dull with
reticulate, densely punctate sculpture, except anterolaterally
with microtessellate sculpture, openly punctate; scutellum
surface dull with reticulate, densely punctate sculpture except
medially openly punctate but dull; dorsal surface of
metapostnotum same as in female except V-shaped, weak striae
reaching dorsolateral margin, posterolaterally weakly plicate;
colour similar to female except apical two thirds of clypeus pale
yellow, fore tibiae light brown and metasoma with distinct blue
tinge; apical margin of T6 yellow.
Vestiture: frons with erect, black hair; between antennal
bases, paraocular area with dense white semi-erect hair, gena
with erect, long beard-like, white hair; mesoscutum with short,
erect, brown hair, metapostnotum lateral sides with sparse, long,
white, erect hair, apical posterior vertical posterior surface of
metapostnotum glabrous; metasomal sterna with short erect,
adpressed black setae, S4—S5 with elongate, erect lateral hair
tufts (fig. SF).
Genitalia: (figs. 11A-B) gonobase sides narrowed basally,
complete ventroapically, gonobase width almost half of
sonocoxa width, gonocoxa glabrous dorsally and ventrally,
dorsal inner margins of gonocoxa basally broadly truncate to
broadly rounded, gonocoxa apical inner margin not produced,
continues contours of gonostylus, glabrous; retrorse lobes
moderate in length, not meeting at midline, membranous,
inwardly pointing, outer margins of retrorse lobes glabrous,
inner margins of retrorse lobes with area of erect, large, inwardly
pointed setae, apically retrorse lobes glabrous; gonostylus
moderate sized (about one third length of gonocoxa), erect,
apically rounded, sparse short setae on inner margin, dense
elongate simple, erect hair on outer margin; penis valves curved
apically, glabrous dorsolaterally.
Distribution: (fig. 12D) this species occurs from SE
Queensland, down eastern New South Wales, across Victoria,
and is widespread in Tasmania.
Remarks: Rayment (1953) described the male of L. plorator,
from Victoria (Gorae West) and labelled the specimen an
Allotype; however, this specimen has no type status.
Lasioglossum plorator is the species found nesting in rotting
wood in the Facebook posts mentioned above (figs. 3A-F, 4).
158
Lasioglossum (Australictus) tertium (Dalla Torre)
(Figs. IA, 2B, 9A-F, 11C, 11D, 12E)
Halictus rufipes Smith 1853: 56. [junior primary homonym of
Halictus rufipes Fabricius, 1793]
Halictus tertius Dalla Torre 1896: 86. [nom. nov. for Halictus
rufipes Smith, 1853]
Parasphecodes insculptus Cockerell 1918: 118. syn nov.
Parasphecodes rufitarsus Rayment 1929: 127. syn nov.
Lasioglossum (Australictus) fulvofasciae Michener 1965: 310. syn
nov.
Lasioglossum (Australictus) tertium — Michener 1965: 165.
Lasioglossum (Australictus) insculptum — Michener 1965: 165.
Lasioglossum (Australictus) rufitarsum — Michener 1965: 165.
Lasioglossum (Australictus) fulvofasciae -Michener 1965: 165.
Material examined: Holotype of rufipes 9, label reads “Melb N.H. F.
Sm. Coll. 79.22” = Australia, Victoria, Melbourne (has additional
label as: Halictus rufipes Sm Type = tertius D.T. Det. Michener 1960)
BMNH Hym.17.a. 2837 (BMNH) (view type data and image at https://
data.nhm.ac.uk/object/68143dfc-dOb8-474d-af44-71aacdecb7706
accessed 16 August 2022).
Holotype of insculptus 9, Queensland, Tamborine Mountain, 28
Dec 1911, H. Hacker (190) Hy/4142 (OM). Note the holotype of
insculptus has a Cockerell handwritten label designating this specimen
as the type. Associated with the Holotype are two specimens with the
same locality data, one female and one male. Neither specimen carries
Cockerell's handwritten label; they are not mentioned in the original
description and have no type status.
Holotype of rufitarsus &, Victoria, Cann River, Nov 1928, J. Clark.
1-118564 (NMV) (view type data at https://collections.museumsvictoria.
com.au/specimens/1018349 accessed 16 August 2022).
Holotype of fulvofasciae 4, Queensland, 3 mi W Cunningham's
Gap, 25 Feb 1959, C.D. Michener, on Bursaria spinosa 1-6910 (QM).
Allotype 2, same data as Holotype except has an additional label of
“375”. T-6911 (QM). The Holotype locality label is printed, while the
allotype locality label 1s handwritten by Michener.
Other specimens examined (239, 513): QUEENSLAND: (19) Mt
Lindesay, 25 Sept 1967, T.F. Houston, 87/450 (WAM); (205) 3 mi W
Cunningham's Gap, 25 Feb 1959, C.D. Michener, on Bursaria spinosa
(SEM); (12) Bald Mt area, 3-4000 ft via Emu Vale, 17-22 May 1980,
G.B. Monteith (QM); (13) 13 mi N Stanthorpe, 29 Dec 1958, C.D. | | l ]
~ so on S5 (fig. OF). This species occurs in SE Australia but is
Michener (SEM); (34) Barney Ck, 7 Mar 1965, S.R. Curtis (QM).
NEW SOUTH WALES: (15) Nadgee State Forest, 7 km S of
Newton's Beach, 7 Dec 1985, E.A. Sugden (AM); (25) Cheltenham,
12 Feb 1950 (AM); (29,95) 6 km NE Bilpin, Blue Mtns, 13 Dec 1984,
28 Feb, 4 Mar 1986, N.W. Rodd (one pair labelled “in cop") (AM);
(14) Haystack Ridge, nr Mt Tomah, 29 Mar 1978, NW. Rodd (AM);
(19) Dunns Swamp, nr Kandos, 13 Nov 1982, N.W. Rodd (AM); (25)
7 km NE Bilpin nr Kurrajong, 17 Mar 1981, NW. Rodd (NMV); (15)
Murray Beach, Jervis Bay, 18 Feb 1987, NW. Rodd (AM); (16)
Narrow Neck, Blue Mtns, 21 Mar 1979, NW. Rodd (AM); (165)
Tianjara Falls, 60 km SW ‘Tomerong, 5 Feb 1988, N.W. Rodd (AM);
(1%) 3 km S Mt Wilson, Blue Mtns, 9 Mar 1978, NW. Rodd (AM);
(1?) Mt White, 23 Sept 1995, A. Sundholm, on Leptospermum (QM);
(1?) Mt Wilson, 4 Jan 1931, A.N. Burns (NMV); (2%) Wisemans
Ferry, 18 Dec 1927, G.B. (NMV); (19) Gibraltar Range, 29 Dec 1969,
C.W. Frazier (ANIC); (19) Dawson's Spring, 1420m, Mt Kaputar Nat.
Park, 1-10 Dec 1987, G.R. Brown (OIA).
VICTORIA: (39, 45) N of Lakes Entrance, Colquhoun State
Forest, 21 Feb 1985, K. Walker, on Eucalyptus (NMV); (13) LaTrobe
Survey, Tanjil Junction (NMV); (15) Wilson’s Promontory, 5 Mile
Beach Road, 26 Feb 1996, K. Walker, on Eucalyptus (NMV); (15)
K.L. Walker
Anglers Rest, 5 March 1992, K. Walker, on Eucalyptus (NMV); (13)
Cobboboonee State Forest, 3 Feb 1997, K. Sparks & C. McPhee, on
Bursaria (NMV); (15) 22 km N Portland, Cobboboonee State Forest,
15 Feb 1990, W.C. Wcislo (SEM); (42) Warburton, 2 Dec 1918, Mar
1920, F.P. Spry (NMV); 29) Lake Hattah, Jan 1920, J. Dixon (NMV);
(1?) Mittagong, 17 Dec 1947, G.M. Goldfinch (NMV); (19) 35 km N
Cann River, 3 Nov 1988, R.R. Snelling & J. Grey (LACM); (19) 8 km
S Cann River, 5 Feb 1987, K. Walker & C. McPhee, on Eucalyptus
(NMV); (19) Freestone Creek Rd, Moornapa, M. Smith (iNaturalist
— https://www.inaturalist.org/observations/26301693 ); (19) 115
Cranbourne-Frankston Rd, Langwarrin (1Naturalist — https://www.
inaturalist.org/observations/25985414 ); (19) Yan Yean, 19 Nov 1999,
Walker & Danforth (Cornell); (19) 42 Reserve Rd, Wonga Park
(Naturalist — https://www.inaturalist.org/observations/25749227 );
(1?) Edgar Track, Montrose, R. Richter (iNaturalist — https://www.
inaturalist.org/observations/25719040 ); (19) Rocklands-Cherrypool
Rd, Rocklands, R. Richter (1Naturalist — https://www.inaturalist.org/
observations/25735753).
SOUTH AUSTRALIA: (12) Nangwarry, 09 Nov 2014, R. Leijs,
on Leptospermum (BDBSA).
Floral record: Families visited: 2 (Myrtaceae (5), Pittosporaceae (2)).
Genera Visited: 3 (Bursaria (2), Eucalyptus (3), Leptospermum (2)).
Flight phenology capture records: Jan (1) Feb (9) Mar (7) Apr (0) May
(1) June (0) July (0) Aug (0) Sept (2) Oct (0) Nov (4) Dec (8).
Diagnosis. Lasioglossum (Australictus) tertium is most like L.
(Australictus) peraustrale with yellow tomentum on pronotum
and metasomal hair bands. This species can be distinguished,
in both sexes, from other Australictus species by: black body
colour (except some males in the SE Queensland area with light
coloured banded metasomal tergal segments (fig. 2D)); dense,
yellow tomentum on pronotum lateral corners, vestiges of
yellow tomentum on posterolateral corners of mesoscutum,
none on metanotum but with metasomal bands on T2-T3;
dorsal surface of metapostnotum posterior margin carinate;
metapostnotum dorsal surface sculpture microalveolate
(figs. 9A-D); male genitalia with gonobase widened basally (a
character shared with L. davide), small retrorse lobes,
sonostylus absent (figs. 10E, IOF); and metasomal sternal S3-
S5 with erect, lateral hair tufts, more so on S4 and even more
absent from Tasmania (fig. 12E).
Description of female: (figs. 1A, 9A, 9B) body length: 8.79-
10.37-10.83 mm (n=10); forewing length: 2.50—2.79—2.83 mm
(n=10); head width: 2.64—3.06—3.17 mm (n=10); intertegular
width: 1.78-1.85-1.87 mm (n=10). Relative head measurements:
HW: 100, HL: 74-75, UID: 55-56, LID: 64-65, IAD: 10-11,
OAD: 20-21, IOD: 11-12, OOD: 16-17, CL: 16-17, GW: 24—
25, EW: 23-24, SL: 41-42, FL: 65-66.
Head: (fig. 1A) upper and lower inner eye margins
diverging basally (head widened basally); median frontal
carina reaching about one third way to median ocellus;
clypeus smooth, polished, medially flattened, open to closely
punctate with small, shallow, rounded punctures except
densely punctate along apical margin; supraclypeal area not
distinctly raised above paraocular area, surface flat, smooth,
polished, open to closely punctate with shallow punctures;
frons smooth, polished, above antennal bases half way to
inner margins of eyes closely to densely punctate, along inner
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 199
eye margins closely to densely punctate wıth minute punctures
with smaller diameters than punctures above antennal bases,
paraocular area smooth, shining, close to densely punctate;
cena enlarged (fig. 9B).
Mesosoma: (figs. 9A, 9B) mesoscutum anterior mesial
margin produced forward over pronotum, surface smooth,
polished with dull sheen, medially open to closely punctate,
laterad of paraspidal lines closely punctate, anterolaterally, in
parapsidal areas densely punctate; scutellum 0.81 x shorter than
length of dorsal surface of metapostnotum, scutellum smooth,
polished, long midline densely punctate, laterally sparse to
openly punctate; dorsal surface of metapostnotum carinate
posterolaterally only, dorsal margin not defined by raised ridge,
dorsal surface covered with microalveolate sculpture but with
several weak striae, striae just reaching mid-point of dorsal
surface, dorsal surface pattern forming rounded V-shape pattern
onto vertical posterior surface, lateral margins smooth, dull with
microalveolate pattern, vertical posterior surface of
metapostnotum with carinae extending to dorsal posterolateral
carinae; mesepisternum and metepisternum plicate; first
recurrent vein (1m-cu) entering third submarginal cell.
Metasoma and legs: (figs. 9A, 9B). metasomal T1—T5 with
dull sheen, densely punctate across entire surfaces; anterior
metatibial spur finely serrate to smooth (fig. 2B).
Colour: (figs. LA, 9A, 9B) frons, supraclypeal area, apical
one third of clypeus, mesoscutum, scutellum black suffused
with dark blue tinge, in some specimens basal two thirds of
clypeus red-brown, in other specimens entire clypeus dark
brown, metanotum, dorsal surface of metapostnotum black
suffused with blue tinge, metasomal TI-T5 black, in some
Specimens terga suffused with brown tinge, in some
specimens 14-14 light red-brown coxa, trochanter, basal two
thirds of femur dark brown, remainder of legs light red-brown;
tegula light red-brown, forewing with brown tinge from
submarginal cells to tip of wing; pronotum dorsolateral
corners, basal margin of T2-T3 with dense yellow tomentum.
Vestiture: (figs. LA, 9A, 9B) body vestiture sparse, frons,
clypeus, supraclypeal area, paraocular area with sparse, erect
hair, pronotum dorsolateral margins with dense tomentum,
mesoscutum scutellum almost glabrous with sparse cover of
small, black, erect hair, in some specimens posterolateral
corners of mesoscutum with weak, yellow tomentum or hair
tufts, in other specimens, perhaps worn, with no tomentum or
hair tufts, lateral surfaces of metapostnotum with long, black
hair; metasoma terga appears glabrous but with dense cover of
short, adpressed hair.
Description of male: (figs. 2D, 9C, 9D) body length: 8.63—9.87—
10.21 mm (n=10); forewing length: 2.31—2.62—2.68 mm (n=10);
head width: 2.11-2.35-2.40 mm (n=10); intertegular width:
1.73-1.87-1.92 mm (n=10). Relative head measurements: HW:
100, HL: 90-92, UID: 57-59, LID: 48-49, IAD: 15-16, OAD:
23-24, IOD: 15-16, OOD: 17-18, CL: 30-31, GW: 20-22, EW:
27-28, SL: 25-26, AFA/AF243 (20/15, 18/14) 1.28-1.33, FL:
187-190.
Differs from female as follows: upper and lower inner eye
margins converging basally; median frontal carina extending
about one third to medium ocellus; frons sculpture densely
reticulate-punctate across surface to inner margins of eyes;
scape reaches level of median ocellus; clypeus surface polished,
shiny, surface weakly convex, close to openly punctate with
shallow punctures; supraclypeal area weakly protruding above
paraocular area, supraclypeal area rounded, dull, with
microtessellate sculpture pattern, closely punctate with minute,
shallow punctures; mesoscutum as in female, anterior margin
produced forward, mesoscutum smooth with dull sheen,
medially closely punctate, anterolaterally, laterad or parapsidal
lines, in parapsidal areas densely punctate, scutellum,
metanotum similar to mesoscutum; dorsal surface of
metapostnotum acarinate, not defined by raised ridge, with
microtessellate sculpture, with weak striae, striae just reaching
mid-point of dorsal surface; colour similar to female except
apical two thirds of clypeus pale yellow, with similar yellow-
orange vestiture on pronotum dorsolateral corners, metasomal
12-13 with basal, yellow tomentum on 13 broader than basal
tomentum on T2, in some specimens metasoma with light
brown colour banding across metasomal terga T1—15 (fig. 2D).
Vestiture: lower half of frons, between antennal bases,
paraocular area with dense, erect, white hair, gena with
moderate beard of long, plumose, white hair, mesoscutum
appear glabrous but with sparse cover of short, erect, brown
hair, metapostnotum lateral sides with basal, short, white,
adpressed hair, metasomal sterna SI-S4 with sparse, white,
adpressed hair, 53—55 with erect, lateral hair tufts, more so on
S4 and even more so on S5 (fig. YF).
Genitalia: (figs. 11C, 11D) gonobase sides widened
basally, complete ventroapically, gonocoxae 1.1 x than width
of gonobase, 1.2 x longer than length of gonobase, gonocoxa
glabrous, dorsal inner margins of gonocoxa basally broadly
truncate to broadly rounded. Gonocoxa apical inner margin
not produced, continue contours of gonostylus, glabrous;
retrorse lobes small, narrow, not meeting at midline,
membranous, retrorse lobes glabrous except for one or two
setae; gonostylus small to almost absent, apically rounded,
with cover of short erect setae; penis valves curved apically,
glabrous dorsolaterally.
Distribution: (fig. 12E) this species extends from SE Queensland
down eastern New South Wales, across Victoria, one record
from SE South Australia, and is absent from Tasmania.
Remarks: Cockerell (1918: 118) commented that Parasphecodes
insculptus was close to P. plorator but noted a colour and
sculpture difference. The metasomal T2-T3 yellow tomentum is
weak on this type specimen; however, the head measurements
are typical for this species. Rayment (1929: 128) commented that
P. rufitarsus was also close to P. plorator but noted colour
differences. Michener (1965: 311) commented that Lasioglossum
fulvofasciae was close to L. peraustrale but differed in
metasomal colour banding on his new species. Michener (1965:
311) primarily used the metasomal colour banding in the male
only to distinguish his new species from L. insculptum. Note that
Michener’s male Holotype of L. fulvofasciae has the metasomal
colour banding, but his allotype female is typical colouration for
L. tertium. Apart from Michener’s type series for L. fulvofasciae
collected at Cunnginham’s Gap, I have located only a few other
specimens with similar banded metasomal colour markings. It
160
appears the distinctive metasomal banding seen on male
specimens of L. fulvofasciae are all from SE Queensland (e.g.
Cunningham’s Gap, Emu Vale, Mt Lindesay and Bald
Mountains), suggesting some kind of colour morph variant in
that area. | have examined a series of male specimens without
the metasomal colour banding also collected by Michener at
Cunningham’s Gap when he collected the type specimens of L.
fulvofasciae. Male genitalia examinations of the Cunningham’s
Gap banded type series specimens and the non-banded males
collected by Michener at Cunningham’s Gap at the same time
showed no differences between these specimens and between
other male L. tertium genitalia examinations. This is the basis for
the L. fulvofasciae synonymy. Of note, the banded metasomal
morph specimens occur about the northernmost known
distribution for this species. The male genitalia of L. tertium
have shared characters with two other Australictus species: the
sonobase widened basally is shared with L. davide (figs. 10A,
10B, 11C, 11D), and the absence of gonostylus is shared with L.
peraustrale (figs. 10E, IOF, 11C, 11D). Note that L. davide, L.
peraustrale and L. tertium are all absent from Tasmania, while
both L. lithusca and L. plorator do occur in Tasmania.
Lasioglossum (Chilalictus) orbatum (Smith)
(Figs. 12F; Walker 1995, pp. 193-194, figs. 132 A-H)
Halictus orbatus Smith 1853: 58-59.
Halictus viridarii Cockerell 1930: 42. syn. by Walker 1995: 193.
Halictus franki Friese H. 1924 syn. nov.; Cockerell 1929: 13.
Lasioglossum (Chilalictus) orbatum — Michener 1965: 177.
Lasioglossum (Chilalictus) viridarii - Michener 1965: 177.
Lasioglossum (Australictus) franki — Michener 1965: 165.
Remarks: in the present study, the TYPUS of Halictus franki
could not be located. ABRS (2022b) lists the type of Halictus
frankias in AMNH. A search of the AMNH Invertebrate Zoology
database for "Halictus franki" returned “Objects 0” result
(AMNH, 2022a) and a search for just "frank1" returned one result
for Paracolletes franki Cockerell 1929 (AMNH 2022b).
The description of Halictus franki lists the T Y PUS sex and
location as “9 Freemantle (sic Fremantle), 20 July 1906 Frank
leg”. This study has determined that for valid Lasioglossum
(Australictus) species, the subgenus occurs only along the east
coast of Australia and a few records in SE South Australia
(figs. I2A-E). If this species does belong to the subgenus
Australictus, it would suggest the TYPUS location label for
Halictus franki in Western Australia (Fremantle) is doubtful.
While searching for the Halictus franki TY PUS specimen
in AMNH, a Friese specimen, missing the metasoma, labelled
as TYPUS was located (this specimen was borrowed and
examined) with Friese’s handwritten label as “Halictus” and
with an unpublished Australian place-based species name. This
specimen has the following labels in descending order: the
location label of “Australia: Sydney, 14.9.06 Frank”; a Friese
handwritten species name label with "1909 Friese det’; a
printed orange TYPUS label; a printed AMNH registration
number of “Am. Mus. Nat. Hist. Dept. Invert. Zool. No. 26905”;
a handwritten Cockerell determination label; and an undated,
typed and handwritten label stating “Lasioglossum [typed]
Australictus [handwritten] det. G.C. Eickwort’.
K.L. Walker
Below the orange Friese TYPUS label is a Cockerell
handwritten label with “Halictus frank1’. Cockerell (1929: 13)
examined this specimen and under the species name “Halictus
franki”, wrote these remarks:
A second specimen, which has lost the abdomen, but is
evidently the same species, is labeled (sic) Sydney, 14.9.06
(Frank). It carries a manuscript name by Friese referring to it as
Australian. I assume that Friese withdrew the latter from
publication, finding it to be identical with H. franki. The hind
spur has short noduliform teeth. The mesothorax is excessively
densely punctured all over.
Cockerell’s (1929) remarks and his handwritten specimen
label of Halictus franki on the unpublished Friese named
specimen provide an insight into the actual species Halictus
franki. Examination of this unpublished Friese TYPUS
specimen confirmed that it is Lasioglossum (Chilalictus)
orbatum. The identification was confirmed by the specimen
having the following subgeneric and species characters.
Subgeneric characters: mandibular preapical tooth is not
enlarged or elongated but of typical Chilalictus length and
shape; anterior metatibial spur is typical of Lasioglossum
(Chilalictus) species (1.e. one large basal tooth followed by a
wavy margin to the apex). Species characters: mesoscutum
punctation is typical of L. (Chilalictus) orbatum; shape of
dorsal surface of the metapostnotum is diagnostic for L.
(Chilalictus) orbatum (1.e. lateral margins of metapostnotum
expanded at level of dorsal surface). Only five species of
Chilalictus have this type of metapostnotum sculpture (Walker,
1995: 193), and the Sydney location is within the known
distribution for L. (Chilalictus) orbatum (fig. 12F). From the
identification of this specimen, I conclude that Halictus franki
is a synonym of Lasioglossum (Chilalictus) orbatum, a species
which is restricted to the east coast of Australia (Walker, 1995,
fig. 12F).
Acknowledgments
I would like to sincerely thank the people and institutions who
allowed me to borrow their material. These include Derek Smith
(AM), Jo Cardale and Ian Naumann (ANIC), Gary Taylor
(BDBSA), George Else (BMNH), Bryan Danforth (Cornell),
Murray Fletcher and Peter Gillespie (OIA), Ross Storey (QDPI),
Geoff Monteith (QM) and Robert Brooks (SEM), Cathy Byrne
(TMAG), Margaret Williams (TDA) and ‘Terry Houston
(WAM). I would also like to especially thank Christopher
Robbins, whose wonderful live images helped to confirm the
wood-nesting behaviour of Australictus bees. Christopher
provided permission for me to reproduce his images.
References
ABRS. 2022a. ABRS 2022. Australian Faunal Directory. Lasioglossum
statistics. Australian Biological Resources Study, Canberra.
https://biodiversity.org.au/afd/taxa/Lasioglossum/statistics
(accessed 31 May 2022)
ABRS. 2022b. A BRS 2022. Australian Faunal Directory. Lasioglossum
(Australictus) franki. Australian Biological Resources Study,
Canberra. _ https://biodiversity.org.au/afd/taxa/36aed2d6-3905-
4e00-aff1-6c5cccdc2819 (accessed 31 May 2022)
Taxonomic revision of the Australian native bee subgenus Australictus (Hymenoptera: Halictidae: Halictini: genus Lasioglossum) 161
Aguiar, A.P., and Gibson, G.A.P. 2010. The spatial complexity in
describing leg surfaces of Hymenoptera (Insecta), the problem
and a proposed solution. Zootaxa 2415: 54-62. https://doi.
org/10.11646/zootaxa.2415.1.5
AMNH. 2022a. Search on Invertebrate Zoology database at AMNH for
Halictus franki returned no results: https://emu-prod.amnh.org/
imulive/iz.html#view=list&id=8685&modules=
ecatalogue&IdeCurrentGenusLocal-Halictus&IdeCurrentSpeciesL
ocal=frankı&SecDepartment_tab=Invertebrate%20Zoology&SecLo
okupRoot=Invertebrate%20Zoology&CatDepartment=Invertebra
te%20Zoology&CatObject lype=Specimen%2FLoté&AdmPublishW
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tab=Invertebrate%20Zoology&SecLookupRoot=Invertebrate%20
Zoology&CatDepartment=Invertebrate%20Zoology&CatObjectIy
pe=Specimen%2FLot&AdmPublishWebNoPassword=Yes
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