Systematic revision of the eastern Queensland land snail genus Figuladra Köhler and Bouchet, 2020 based on mitochondrial phylogenetics and comparative morpho-anatomy (Eupulmonata, Camaenidae)

ABSTRACT

The current study revises the systematic taxonomy of Figuladra, a genus of camaenid land snails endemic to eastern Queensland, based on the investigation of many recently collected ethanol-preserved samples as well as historic museum material, including types. We employed comparative analyses to investigate the variation in morphological features (shell, penial anatomy) and performed phylogenetic analyses of mitochondrial 16S rDNA sequences to test species hypotheses. Phylogenetic analyses recovered three principal mitochondrial clades, one occurring in mid-east Queensland, one in south-east Queensland, and another in the upland areas of south-east Queensland. We recognise altogether 15 accepted species, including four that are newly described herein (i.e., Figuladra finlaysoni sp. nov., F. robertirwini sp. nov., F. bromileyorum sp. nov. and F. vidulus sp. nov.). The status of 11 previously described species has been assessed and revised diagnoses are provided for each of these. Species of the south-east Queensland clade revealed comparatively low levels of genetic and morphological differentiation and may represent examples of more recent, conceivably ongoing, speciation. Figuladra species have allopatric distributions with their ranges being separated by altitudinal or riverine barriers. Particularly the St Lawrence Gap dry corridor has been identified as a major biogeographic barrier separating two principal clades of Figuladra.

KEYWORDS:

• Biogeography
• gastropoda
• new species
• phylogeny
• systematics
• taxonomy

Introduction

Queensland has a significant number of large to very large, striped land snails colloquially often referred to as ‘hadroid’ snails (Hugall and Stanisic 2011), which belong to the family Camaenidae. Figuladra Köhler & Bouchet, 2020 is one of the lesser known ′hadroid′ genera inhabiting predominantly dry rainforests and vine thickets in coastal and subcoastal lowland areas as well as the adjacent escarpments of the Great Dividing Range. The distribution of Figuladra ranges from north of Gympie in south-eastern Queensland to south of the O′Connell River in mid-eastern Queensland, extending approximately between the latitudes of 20°S to 26°S and longitudes of 148°E to 152°E (Stanisic et al. 2010, Figure 1). This range transverses the St Lawrence Gap, a tract of low-lying open woodland and savannah between St Lawrence and Kunwarara, mid-eastern Queensland, which represents a major biogeographical barrier for various terrestrial biota (Bryant and Krosch 2016, fig. 5). Species of Figuladra are characterised by large tawny yellow to brown and, in general, spirally banded shells that range in size from approximately 20 to 33 mm in height and 23 to 50 mm in diameter (Stanisic et al. 2022).

Figure 1. Current distribution of Figuladra species including candidate species in mid-eastern Queensland (MEQ) and south-eastern Queensland (SEQ).

Historically, Figuladra species have been assigned to various genera. Iredale (1937) listed 13 species under Varohadra Iredale, 1933 which is not an available name (nomen nudum, see Köhler and Bouchet 2020). Smith (1992) reduced the number of recognised species to six, which were placed in Sphaerospira Mörch, 1867. More recently, Stanisic et al. (2010) listed 13 accepted species (including three newly described) under Figuladra Iredale, 1933. However, Köhler and Bouchet (2020) considered that Iredale′s original name was unavailable due to the lack of a diagnosis (Art. 13.1 of the Code; ICZN 2012) and formally validated the name Figuladra while maintaining the treatment of Stanisic et al. (2010). Stanisic and Stanisic (2020) re-examined the types and reviewed the type localities of all nominal species included in Figuladra, thereby setting the nomenclatural foundations for the present work.

More recently, Stanisic et al. (2022) excluded two species previously assigned to Figuladra, namely F. mattea (Iredale, 1933) and F. volgiola (Iredale, 1933), placing each of them into a new monotypic genus, Euryladra and Brigaladra.

In an integrated taxonomic approach, the present study employs comparative morpho-anatomy and mitochondrial phylogenetics to produce a comprehensive revision of Figuladra.

Abbreviations

General: fig. – figure; figs – figures; max – maximum; RC – dry collection; SEM – scanning electron microscopy; SC – spirit collection; spec – specimen(s); var – variation.

Institutions: AM – Australian Museum, Sydney; NEWHM:HMNT – Great North Museum: Hancock, Newcastle, United Kingdom; NHMUK – Natural History Museum, United Kingdom; QM – Queensland Museum, Brisbane; ZMB – Museum für Naturkunde, Leibniz-Institut für Evolutions und Biodiversitätsforschung, Germany.

Geography: alt – altitude; Ck – Creek; Hts – Heights; Id – Island; MEQ – Mid-eastern Queensland; Mt – Mountain; Mts – Mountains; NP – National Park; NEQ – North-eastern Queensland; NW – Northwest; NNW – North-northwest; R – River; Ra – Range; SCQ – Southern-central Queensland; SEQ – South-eastern Queensland; SF – State Forest; X′ing – Crossing.

Habitat data: cnvf – complex notophyll vine forest; mvf – microphyll vine forest; nvf – notophyll vine forest; sevt – semi-evergreen vine thicket.

Molecular phylogenetics: 16S – 16S ribosomal RNA gene; BI – Bayesian Inference; ML – maximum likelihood; MOTUs – molecular operational taxonomic units.

Shell features: D – shell diameter; H – shell height; H/D – ratio of shell height to shell diameter; W – number of whorls.

Genital anatomy: BC – bursa copulatrix; DG – prostate; E – epiphallus; EF – epiphallic flagellum; EP – epiphallic pore; GD – hermaphroditic duct; GG – albumen gland; P – penis; PPL – penial pilasters; PRM – penial retractor muscle; PS – penial sheath; PV – penial verge; PVR – ratio of penial length to vaginal length; SS – bursa stalk; UT – uterus; UV – free oviduct; V – vagina; VD – vas deferens; Y – atrium.

Materials and methods

Specimens and sampling

Examined specimens comprised both dry shells and ethanol preserved samples deposited in the Queensland Museum, Brisbane, the Australian Museum, Sydney and the British Museum of Natural History, London. Individual specimens are identified by their registration number and respective institutional prefix (QMMO – Queensland Museum; AMS C – Australian Museum, Sydney; NHMUK – British Museum of Natural History). Field trips to specific locations in eastern Queensland known to be Figuladra habitat were conducted to collect animal tissue for dissection and genetic analysis. Live specimens collected were transported from the field to the laboratory, drowned in water, usually overnight, then fixed in 95% ethanol and subsequently stored in 70% ethanol for use in anatomical and molecular studies.

Molecular methods

Specimens, DNA extraction, amplification and sequence determination

A mitochondrial DNA sequence data set was created to estimate the amounts of genetic divergence among the Figuladra species and investigate their phylogenetic relationships. This dataset contained partial ribosomal DNA sequences (16S rDNA) of all taxa including four outgroup species. We included additional GenBank sequences published by Hugall and Stanisic (2011) and Stanisic et al. (2022) to enhance the sampling of our analysis (Table 1).

Table 1. Specimens, nomenclature and GenBank accessions used in the 16S mtDNA analyses.

Taxon	Voucher ID	GenBank	Sampling Location	Source
Camaenid BL 45	QMMO87124	OQ676310	Wadallah Ck, MEQ	Present study
F. appendiculata	QMMO86924	OQ676312	Mt Jim Crow, SEQ	Present study
F. appendiculata	QMMO87109	OQ676313	Lakes Ck, SEQ	Present study
F. appendiculata	QMMO87110	OQ676314	Keppel Sands, MEQ	Present study
F. appendiculata	QMMO87114	OQ676311	Mulambin Beach, MEQ	Present study
F. appendiculata	QMMO87119	OQ676317	Lamamoor, MEQ	Present study
F. appendiculata	QMMO87120	OQ676316	Great Keppel I, MEQ	Present study
F. appendiculata	QMMO86833	OL839884	Mt Archer SEQ	L. Stanisic et al. 2022
F. aureedensis	QMMO86835	OQ676318	Johannsen’s Cave, SEQ	Present study
F. aureedensis	QMMO87121	OQ676319	Johannsen’s Cave, SEQ	Present study
F. barneyae	QMMO87097	OQ676320	Nebo, MEQ	Present study
F. barneyae	QMMO87440	OQ676322	Greenhill, MEQ	Present study
F. barneyae	QMMO87127	OL839885	Connors Hump MEQ	L. Stanisic et al. 2022
F. barneyae	QMMO87435	OL839886	St Lawrence, MEQ	L. Stanisic et al. 2022
F. bayensis	QMMO86654	OQ676323	Mt Mudlo SEQ	Present study
F. bayensis	QMMO86659	OL839887	Mt Biggenden SEQ	L. Stanisic et al. 2022
F. bayensis	QMMO86928	OL839888	Mt Mudlo SEQ	L. Stanisic et al. 2022
F. bromileyorum	QMMO86677	OQ676324	Dundowran, SEQ	Present study
F. bromileyorum	QMMO86887	OQ676325	Dundowran, SEQ	Present study
F. bromileyorum	QMMO87064	OQ676326	Aramara SF, SEQ	Present study
F. challisi	QMMO86828	OL839889	Keswick Id, MEQ	L. Stanisic et al. 2022
F. challisi	QMMO86829	OL839890	St Bees Id, MEQ	L. Stanisic et al. 2022
F. finlaysoni	QMMO55753	GQ851068	Seaforth, MEQ	Hugall and Stanisic, 2011
F. finlaysoni	QMMO66973	OQ676327	Mt Ossa, MEQ	Present study
F. finlaysoni	QMMO87133	OQ676328	Oxford-Sarina Rd, MEQ	Present study
F. finlaysoni	QMMO87456	OL839881	Seaforth, MEQ	L. Stanisic et al. 2022
F. incei	QMMO87122	OQ676329	Stanage Bay, MEQ	Present study
F. incei	QMMO87064	OL839891	Percy Id, MEQ	L. Stanisic et al. 2022
F. incei	QMMO87064	OL839891	Percy Id, MEQ	L. Stanisic et al. 2022
F. incei	QMMO87064	OL839891	Percy Id, MEQ	L. Stanisic et al. 2022
F. incei	QMMO87084	OL839892	Percy Id, MEQ	L. Stanisic et al. 2022
F. lessoni	QMMO72163	OQ676330	Mercy Hill, SEQ	Present study
F. lessoni	QMMO72198	OQ676331	Mt Larcom, SEQ	Present study
F. lessoni	QMMO72292	OQ676332	Maurice Hill, SEQ	Present study
F. lessoni	QMMO79347	OQ676333	Bracewell, SEQ	Present study
F. lessoni	QMMO85827	OQ676334	Curtis Id, SEQ	Present study
F. lessoni	QMMO85828	OQ676335	Curtis Id, SEQ	Present study
F. lessoni	QMMO86803	OQ676336	Henickes Rd, SEQ	Present study
F. lessoni	QMMO86839	OQ676337	Targinnie, SEQ	Present study
F. lessoni	QMMO87104	OQ676338	Boyne Id, SEQ	Present study
F. lessoni	QMMO87105	OL839893	Tannum Sands, SEQ	L. Stanisic et al. 2022
F. lessoni	QMMO87142	OQ676339	Ambrose, SEQ	Present study
F. lessoni	QMMO87144	OQ676340	Henickes Rd, SEQ	Present study
F. lessoni	QMMO87148	OQ676360	Old Ten Chain Rd, SEQ	Present study
F. lessoni	QMMO87155	OQ676341	Mt Coulston, SEQ	Present study
F. lessoni	QMMO87159	OQ676342	Bororen, SEQ	Present study
F. lessoni	QMMO87432	OQ676343	Targinnie, SEQ	Present study
F. muirorum	QMMO87137	OL839904	Marlborough, MEQ	L. Stanisic et al. 2022
F. muirorum	QMMO87392	OQ676344	Boomer Ra, MEQ	Present study
F. muirorum	QMMO87431	OQ676345	Mt Slopeaway, MEQ	Present study
F. narelleae	QMMO86831	OL839905	Bouldercombe Gorge, SEQ	L. Stanisic et al. 2022
F. narelleae	QMMO87428	OQ676346	Dan Dan Scrub, SEQ	Present study
F. pallida	QMMO87117	OQ676347	Polka Rd, Byfield, MEQ	Present study
F. reducta	QMMO86669	OQ676348	Goodnight Scrub, SEQ	Present study
F. reducta	QMMO86689	OQ676349	Kroombit Tops, SEQ	Present study
F. reducta	QMMO86815	OL839906	Mt Perry, SEQ	L. Stanisic et al. 2022
F. reducta	QMMO87102	OQ676350	Ubobo, SEQ	Present study
F. robertirwini		GQ851073	Bulburin SF, SEQ	Hugall and Stanisic, 2011
F. robertirwini	QMMO72190	OQ676351	Gwynne Ra, SEQ	Present study
F. robertirwini	QMMO72209	OQ676352	Eurimbula NP, SEQ	Present study
F. robertirwini	QMMO72282	OQ676353	SSE Calliope, SEQ	Present study
F. robertirwini	QMMO72317	OQ676354	Miriam Vale, SEQ	Present study
F. robertirwini	QMMO86795	OQ676355	Turkey Beach, SEQ	Present study
F. robertirwini	QMMO86888	OQ676356	Round Head, SEQ	Present study
F. robertirwini	QMMO86921	OQ676357	Eurimbula NP, SEQ	Present study
F. robertirwini	QMMO87100	OQ676359	Colosseum Ck, SEQ	Present study
F. robertirwini	QMMO87152	OQ676361	Rodd′s Peninsula, SEQ	Present study
F. robertirwini	QMMO87153	OQ676362	Turkey Beach, SEQ	Present study
F. robertirwini	QMMO87160	OQ676363	Granite Ck Rd, SEQ	Present study
F. robertirwini	QMMO87163	OQ676364	Kolan R X’ing, SEQ	Present study
F. robertirwini	QMMO87163 (2)	OQ676365	Kolan R X’ing, SEQ	Present study
F. vidulus	QMMO87062	OQ676358	Watalgan, SEQ	Present study
F. vidulus	QMMO87075	OQ676367	Coonarr, SEQ	Present study
F. vidulus	QMMO87076	OQ676368	Mon Repos, SEQ	Present study
F. vidulus	QMMO87078	OQ676369	Woodgate, SEQ	Present study
F. vidulus	QMMO87079	OL839882	Bagara SEQ	L. Stanisic et al. 2022
F. vidulus	QMMO87663	OQ676366	The Hummock, SEQ	Present study
Outgroups
Monteithosites heliostracum		GQ851007.1	Bakers Blue Mt, NEQ	Hugall and Stanisic, 2011
Billordia nicoletteae	QMMO78854	GQ851018.1	Lotus Ck, MEQ	Hugall and Stanisic, 2011
Brigaladra volgiola		GQ851328	Broadsound Range, SEQ	Hugall and Stanisic, 2011
B. volgiola	QMMO78118	OL839910	Peak Ra, SCQ	L. Stanisic et al. 2022
B. volgiola	QMMO78170	OL839907	Mt McDonald, SCQ	L. Stanisic et al. 2022
B. volgiola	QMMO78170	OL839911	Mt McDonald, SCQ (2)	L. Stanisic et al. 2022
B. volgiola	QMMO87092	OL839908	Duaringa, SCQ	L. Stanisic et al. 2022
B. volgiola	QMMO87092	OL839912	Duaringa, SCQ	L. Stanisic et al. 2022
B. volgiola	QMMO87096	OL839913	Duaringa, SCQ	L. Stanisic et al. 2022
B. volgiola	QMMO87135	OL839909	Langdale Hill, MEQ	L. Stanisic et al. 2022
Euryladra. mattea		GQ851106	Taroom, SCQ	Hugall and Stanisic, 2011
E. mattea	QMMO86640	OL839895	South Bend, SCQ	L. Stanisic et al. 2022
E. mattea	QMMO86643	OL839894	Injune, SCQ	L. Stanisic et al. 2022
E. mattea	QMMO86799	OL839897	Targinnie SEQ	L. Stanisic et al. 2022
E. mattea	QMMO86812	OL839898	Dulacca SCQ	L. Stanisic et al. 2022
E. mattea	QMMO86908	OL839899	Isla Gorge SCQ	L. Stanisic et al. 2022
E. mattea	QMMO86916	OL839900	Henickes Rd, SEQ	L. Stanisic et al. 2022
E. mattea	QMMO87090	OL839901	Moura, SEQ	L. Stanisic et al. 2022
E. mattea	QMMO87138	OL839902	Horrigans Rd, SEQ	L. Stanisic et al. 2022
E. mattea	QMMO87140	OL839903	Fire Creek, SEQ	L. Stanisic et al. 2022
E. mattea	QMMO87234	OL839901	Pointy Vernon, SEQ	L. Stanisic et al. 2022

Note: NA, not available. (Final names are used).

Total genomic DNA was isolated from the small slices of foot muscle of 101 specimens. To avoid inhibition due to the mucopolysaccharides in snail slime, tissue was extracted, placed in 95% ethanol for two weeks and the ethanol changed weekly and at any time it discoloured. For DNA extraction we used the DNeasy tissue kit (QIAGEN, Venlo, The Netherlands) for animal tissue with the Spin Column protocol as per manufacturer′s instructions. Fragments of the 16S ribosomal RNA were amplified by PCR using the primer pairs 16 SCs1 (Chiba 1999) and 16Sbd1 (Sutcharit et al. 2007). The amplified PCR fragments were approximately 770 bp long.

The PCR cycle programs comprised an initial denaturation step at 99°C for 30 s, followed by 35 cycles, each with a denaturation step of 30 s at 98°C, an annealing step of 30 s at 48°C, and an extension step of 30 s at 72°C. The final cycle had a 10 min sequence extension at 72°C. The PCR products were sent for purification and commercial sequencing at Macrogen Inc (Geumchum-Gu, Korea).

Sequence alignment and genetic data analysis

Forward and reverse reads were combined into contigs, manually checked and corrected, where necessary, using Geneious ver. 9.1.8 (Kearse et al. 2012). Sequences were aligned using the MAFFT v7.308 (Katoh and Standley 2013) plug-in in Geneious with default settings. We used Guidance2 (Sela et al. 2015) to identify and remove unreliably aligned regions in the 16S alignment by employing default settings.

Phylogenetic analysis

Phylogenetic relationships were estimated by employing a maximum likelihood-based method of tree reconstruction as well as Bayesian Inference (BI). A maximum likelihood phylogeny was reconstructed by using IQ-TREE 1.6 (Nguyen et al. 2015) using the web W-IQ-TREE interface available at http://iqtree.cibiv.univie.ac. at/ (Trifinopoulos et al. 2016). We employed the integrated ModelFinder (Kalyaanamoorthy et al. 2017) to identify and implement the best-fit model of sequence evolution. Nodal support was inferred by performing 10,000 ultra-fast bootstrap replicates (Minh et al. 2013).

Bayesian posterior probabilities were estimated by running a 1,000,000 generation Metropolis-coupled Markov chain Monte Carlo (4 chains, one heated, sampling rate 1000 generations) with a burn-in of 100, 000 as implemented by the MrBayes (Ronquist et al. 2012) plug-in in Geneious 9.1.8.

Mean genetic p-distances within and between groups were calculated using MEGA6 (Tamura et al. 2013).

All new sequences produced here have been deposited in GenBank under the accession numbers: OQ676310 to OQ676369 (Table 1).

Morphological methods

Shell morphology

Shells were photographed from three different perspectives, apertural view (viewed from aperture with shell parallel to the coiling axis), top view (viewed from above, perpendicular to the coiling axis), and umbilical view (viewed from below, perpendicular to the coiling axis), to show general shell shape.

Shell characters investigated included shape, size (height and diameter), whorl count, and colour patterns of both shell and lip (Figure 2). Standard definitions for conchological characters are used here (Solem and van Bruggen 1984). Shell dimensions (height, diameter) were measured using callipers with a precision of 0.1 mm. Based on shell diameter we classified shells into three arbitrary size classes: small (< 20 mm), large (20–40 mm), very large (> 40 mm). Whorl counts were made to the nearest ¹/₈ whorl (Figure 2). The sculpture on the teleoconch was examined by scanning electron microscopy using a TM-1000 Tabletop Scanning Electron Microscope located at the Queensland Museum, Brisbane. Selected specimens were cleaned by gently brushing in warm soapy water followed by ultrasonication. Specimens were mounted on sticky tabs, gold sputter-coated and imaged under high vacuum.

Figure 2. Primary shell measurements and method of whorl count. SD – shell diameter; SH – shell height.

Genital morphology

Where possible, at least five representatives of each species including both historical museum and freshly collected specimens were dissected in order to confirm constancy of reproductive structures. Specimens dissected were collected at different times of the year to ensure that genital anatomy description was not affected by seasonal variation. The reproductive system was extracted, pinned to a black latex base using very fine ′Austerlitz′ entomological pins, and photographed.

Radula

Radulae and jaws were extracted from preserved specimens by macerating the buccal cavity in 95% NaOH solution overnight followed by rinsing in distilled water and 70% ethanol. Radulae were then transferred to a small vial of 70% ethanol and cleaned by immersing in an ultrasonic cleaner for 15–20 s. The cleaned radulae were then mounted on aluminium stubs, coated with gold and studied with a scanning electron microscope.

Operational criteria for species delimitation

Our operational criterion for the delimitation of species was to test whether candidate species were phenotypically and genotypically distinct from each other (Sites and Marshall 2004) using two stages.

Grouping stage: Specimens were initially grouped using phylogenetic analyses of mitochondrial gene sequences. These analyses resulted in phylogenetic trees which were compared and used as guide trees.

Our initial groups were specimens that formed tight sequence clusters in the mtDNA trees. In this project, we use a threshold measure of genetic divergence as a secondary guide to the grouping. O’Neill et al. (2014) consistently found <4% divergence in Quistrachia Iredale, 1939 species, particularly where geographical distribution was restricted. Given this background and a lower threshold evident using 16S in the above studies generally ranging from 2% to 4%, a minimum divergence value of 2.5% in 16S was used to test the likely status of morphospecies.

Hypothesis testing: We assessed the variation in reproductive anatomy among representatives and whether these groupings represented anatomically coherent sets, distinguishable from each other. For the purposes of this study, pilasters, pustules and longitudinal thickenings of the penial chamber were examined in three zones, an apical zone connecting to the epiphallus, a central zone and a basal zone opening to the atrium. The species threshold limit was set when several major observable differences were evident in the pustules, pilasters and penial wall patterns between each group.

Patterns of shell morphology were examined in detail and evaluated as additional evidence. These details were used to support anatomical homogeneity in the differentiation of the groups.

Where genetic groupings and morphological evaluations aligned, we considered the groups to be independent species and assigned taxonomic names to them.

Results

Mitochondrial phylogenetics

The final dataset contained sequences of 76 ingroup specimens and 22 sequences of outgroup representatives, which were used to root the trees (Table 1). The outgroup was composed of several closely related hadroid species with distributions that overlapped with or abutted that of Figuladra. The final 16S alignment had a total length of 770 base pairs. The best ML phylogram and the Bayesian consensus tree had largely consistent topologies that differed only in minor details (Figure 3).

Figure 3. Reconstruction of phylogenetic relationships of Figuladra Köhler & Bouchet, 2020 from eastern Queensland based on phylogenetic analyses of 16S sequences with Monteithosites heliostracum Stanisic, 1996, Billordia nicoletteae Stanisic, 2010, Euryladra L. Stanisic, 2022 and Brigaladra L. Stanisic, 2022 as outgroups. Values above the nodes represent bootstrap values, numbers below the nodes represent posterior probabilities. Individual samples are named according to their identification. Numbers after taxon names correspond to catalogue numbers or GenBank accession numbers (see Table 1).

In both trees, Figuladra formed a major clade. Within this clade, both analyses recovered three well-differentiated (in terms of basal branch lengths) and well-supported (in terms of statistical support) minor clades: one containing samples from the upland areas of SEQ, a second comprising northern samples from MEQ, and a third consisting of samples from the coastal areas of SEQ. Each of these three clades contained several generally well-supported subclades (Figure 3). These subclades were treated as initial species hypotheses, or molecular operational taxonomic units (MOTUs), which were subjected to further taxonomic scrutiny through assessments of their genetic divergence (p-distances) and comparative analyses of their morpho-anatomy.

The mean genetic p-distances between different MOTUs ranged from 2.9% to 14.1%, while the mean distances between sequences of the same MOTU ranged from 0.9% to 3.6% (Table 3). Hence, there was minimal overlap between the mean within-group and mean between-group distances in 16S. This overlap was restricted to the low mean distances between the MOTUs F. robertirwini sp. nov. and F. bromileyorum sp. nov. (Tables 2 and 3).

Table 2. Mean pairwise distances (%) in 16S within and between candidate species as recognised on the tree shown in Figure 3.

Species	MEQ clade					SEQ upland clade		SEQ clade
	BL45	challisi	finlaysoni	barneyae	incei	bayensis	narelleae	appendiculata	aureedensis	lessoni	muirorum	pallida	reducta	robertirwini	bromileyoryum	vidulus
BL 45
challisi	12.8	0.9
finlaysoni	11.4	5.3	1.1
barneyae	12.6	4.5	5.4	3.6
incei	12.2	4.6	6.3	4.7	0.8
bayensis	13.4	14.1	13.1	13.2	12.5	2.0
narelleae	13.3	12.9	11.8	12.2	11.4	6.4	2.9
appendiculata	5.0	12.2	11.5	12.5	12.0	11.8	12.6	0.4
aureedensis	4.6	12.8	11.6	12.4	11.4	11.7	12.5	4.3	0.1
lessoni	5.1	12.7	11.7	12.5	11.7	12.6	13.0	4.4	3.0	1.2
muirorum	5.1	13.3	12.0	13.0	12.6	12.2	12.6	4.4	4.7	5.1	2.4
pallida	2.3	12.8	11.3	12.4	12.2	13.2	13.2	4.5	4.3	4.1	4.5	n/c
reducta	5.3	12.6	11.7	12.5	11.7	12.4	12.5	4.0	3.6	3.2	5.4	4.2	3.3
robertirwini	5.8	12.5	12.1	12.8	12.4	12.5	12.8	4.1	4.8	5.2	5.6	5.0	4.4	3.2
bromileyorum	5.4	12.8	11.6	12.8	12.0	12.7	12.6	3.5	4.1	4.5	5.1	4.3	3.5	3.4	1.7
vidulus	5.7	13.1	12.0	13.1	12.4	12.4	12.2	3.2	4.1	4.7	5.1	4.4	3.8	3.5	2.9	0.7

Note: Shaded cells show mean within-group p-distances.

Table 3. Comparative overview of major penial chamber characters supporting the hypothetical species units (PVR = Penial/vaginal ratio).

Species	Apical chamber			Central chamber		Basal chamber		PVR
	Pustules	Pilaster	Medial thickening	Longitudinal thickenings	Pilasters	Longitudinal ridges	Pilasters
SEQ upland clade:
bayensis	prominent, wavy	raised	tongue-like pustules	tongue-like pustules	smaller pustules, large longitudinal ridge	thin, corrugated	random pattern, irregularly shaped	2.1
narelleae	prominent, wavy	absent	enlarged tongue-like pustules	tongue-like pustules	large longitudinal ridge, several short ridgelets	short, thin	tongue-like pustules	3.0
SEQ clade:
appendiculata	elongate, crenellated	weak ridge	tongue-like pustules	1 thick, several thinner	absent	two, thick, corrugated	absent	2.1
aureedensis	small rectangular	weak, raised	absent	1 thick, several thinner	tongue-like pustules	thin	numerous, bold, irregularly shaped	2.2
lessoni	diamond-shaped	raised	tongue-like pustules	2 thick	absent	thin	numerous small	2.3
muirorum	rectangular	raised	tongue-like pustules	several	absent	thin		2.3
pallida	rectangular pustules	weak, raised	weak tongue-like	1 thick	absent	thick	short rectangular ridgelets	2.8
reducta	rectangular	raised	prominent tongue-like pustules	1 thick, several thinner	short row, tongue-like, scattered knob-like pustules	several thin
		numerous short ridgelets
robertirwini	diamond-shaped to rectangular	raised	tongue-like pustules	2 thick, several thinner	absent	thin	extensive fine	2.4
bromileyorum	crenellated rectangular	raised	prominent tongue-like pustules	1 thick, several thinner	longitudinal row of large tongue-like pustules	thin	elongate ridgelets	2.3
vidulus	elongate, rectangular	raised	tongue-like pustules	2 thick, several thinner	few tongue-like pustules	thick	scattered tiny pustules	2.5
MEQ clade:
barneyae	elongate, crenellated	absent	tongue-like pustules	1 thick	bold, spade-like pustules	several	scattered, small	2.3
finlaysoni	rectangular	absent	tongue-like pustules	1 thick, 3-4 thinner	spade-like pustules	several	scattered, small	2.3
challisi	crenellated	absent	tongue-like pustules	2 -3 thick	spade-like pustules	thin	random, irregularly shaped	2.2
incei	crenellated	absent	absent	1 thick, 2 thinner	absent	several short	numerous random, irregularly shaped, tiny	2.4

Note: Taxa are arranged according to clades and lineages within the guide tree. Comparative differences are illustrated in Supplementary Figure S1.

Patterns of anatomical variation

For each of the 15 MOTUs delineated on the phylogenetic tree (Figure 3), at least five specimens were dissected, which were sampled from across the extent of the distribution of each MOTU to test if the observed variation in reproductive anatomy revealed patterns consistent with MOTU membership. Generally, we found minimal variation in the configuration of the complete genitalia between all dissected MOTUs, including the general anatomy of the penis, vagina, oviduct and bursa copulatrix. We consider these characteristics to be rather conserved among Figuladra species (for details refer to the anatomical description under the genus). In contrast, the greatest deal of anatomical variation was observed in the penial anatomy. Moreover, this variation was also consistent with the phylogenetic relationships as shown in Figure 3, and all MOTUs were found to be characterised by a combination of typical features that include the configuration of pustules and pilasters in the three penial chambers (Figure 4, Supplementary Figure S1, Table 4).

Figure 4. General features of the genitalia of Figuladra species. A, Close-up of muscular tissue at the epiphallus-penis junction, F. lessoni QMMO86813, Mt Larcom, SEQ; B, Epiphallic flagellum, F. narelleae QMMO86831, Bouldercombe Gorge, SEQ; C, Rectangular pustules forming a central pilaster, F. lessoni, QMMO60270, Boyne Id, SEQ; D, Crenellated pustules forming a central pilaster, F. appendiculata, Mt Dick, QMMO4173. E, Tongue-like pilasters, F. narelleae, QMMO64057, Hourigan, SEQ; F, Large spade-like pilasters at lower end of apical region, F. barneyae, QMMO70436, Connors Ra, MEQ.

Table 4. Shell measurements for Figuladra species.

	Species	No	Height	Diameter	Whorl Count	H/D ratio
SEQ upland	Figuladra bayensis	23	23.8–37.2	35.4–50.0	5.875–6.250	0.672–0.775
	Mean		28.2	40.4	6.00	0.699
	Standard deviation		±0.4	±0.4	±0.2686	±0.0577
	Figuladra narelleae	52	22.7–30.8	33.4–40.8	5.750–6.375	0.616–0.807
	Mean		26.5	38.0	6.000	0.697
	Standard deviation		±2.1	±2.0	±0.1844	±0.0451
SEQ	Figuladra appendiculata	76	17.6–30.9	22.3–41.2	5.875–6.500	0.717–0.825
	Mean		26.4	34.3	6.250	0.777
	Standard deviation		±3.4	±4.2	±0.1965	±0.0366
	Figuladra aureedensis	44	26.1–28.1	34.7–35.8	6.000–6.250	0.749–0.806
	Mean		26.9	35.1	6.125	0.770
	Standard deviation		±0. 9	±0. 5	±0. 1197	±0.0249
	Figuladra bromileyorum	76	24.6–37. 5	30.2–40.6	6.250–7.000	0.737–0.933
	Mean		29.9	35.5	6.500	0.841
	Standard deviation		±0. 4	±0. 4	±0.1529	±0.0141
	Figuladra lessoni	84	23.8–31.3	31.2–38.4	6.000–6.500	0.714–0.873
	Mean		27.6	34.8	6.250	0.790
	Standard deviation		±1.7	±1.7	±0.1894	±0.0389
	Figuladra muirorum	58	24.5–27.9	30.7–32.2	5.625–6.125	0.773–0.871
	Mean		25.9	31.6	6.000	0.821
	Standard deviation		±1.329	±0. 765	±0.2183	±0.0385
	Figuladra pallida	42	27.4–34.2	33.5–40.8	6.500–7.125	0.811–0.928
	Mean		31.1	36.4	6.625	0.863
	Standard deviation		±1.9	±2.4	±0.1964	±0.0437
	Figuladra reducta	39	21.6–27.9	30.8–36.2	5.250–6.250	0.677–0.757
	Mean		25.0	34.3	6.000	0.730
	Standard deviation		±2.1	±2.3	±0.3496	±0.0328
	Figuladra robertirwini	73	23.6–36.8	30.9–43.8	6.250–7.000	0.639–0.974
	Mean		28.8	35.4	6.500	0.813
	Standard deviation		±3.4	±2.1	±0.2356	±0.0575
	Figuladra vidulus	30	25.6–29.5	24.9–39.4	6.125–6.625	0.733–0.833
	Mean		26.6	31.5	6.250	0.801
	Standard deviation		±1.4	±1.9	±0.1892	±0.0395
MEQ	Figuladra incei	61	20.3–32.0	34.7–50.0	5.750–6.625	0.714–0.962
	Mean		25.5	39.7	6.000	0.809
	Standard deviation		±1.4	±1.9	±0.1892	±0.0395
	Figuladra barneyae	91	20.4–33.4	26.5–39.5	5.875–6.750	0.743–0.863
	Mean		27.6	34.8	6.375	0.816
	Standard deviation		±4.4	±4.8	±0.2649	±0.0308
	Figuladra finlaysoni	222	23.8–31.3	24.6–35.2	5.625–6.625	0.682–0.854
	Mean		27.41	30.93	6.000	0.765
	Standard deviation		±2.6627	±2.9565	±0.2263	±0.0398
	Figuladra challisi	32	22.36–23.52	30.43–33.62	5.875–6.125	0.692–0.749
	Mean		22.6	31.6	6.000	0.722
	Standard deviation		±0.4	±0.4	±0.1529	±0.0141

Note: [Range, mean and standard deviation. Height and diameter measurements in mm].

Genitalia (Part A of each of Figures 5–19). The penis was typically elongate and weakly tapered but slightly expanded apically in both SEQ upland species, F. bayensis (Brazier, 18676) and F. narelleae Stanisic & Potter, 2010. A thin penial sheath that envelops the entire penis was present. The epiphallus consisted of two arms demarcated by the insertion of the penial retractor muscle and entered the penis apically through a simple pore adjacent to a knob-like verge (not visible in all illustrated dissections). The ascending arm of the epiphallus, emanating from the vas deferens, was usually shorter and thinner than the descending arm and usually swollen medially. The descending arm of the epiphallus was typically straight, thick, muscular and bound to the head of the penis by a bundle of muscle fibres (Figure 4A). The vas deferens was typically long, thin and attached to the penial sheath by connective tissue (disconnected in all illustrated dissections). A short, thin vestigial epiphallic flagellum was present and inserted at the epiphallus-vas deferens junction (Figure 4B). The almost indiscernible flagellum was tightly bound to the vas deferens.

Figure 5. Genital details of Figuladra bayensis (Brazier, 1875). A, Genitalia; B, Penis interior; C, Apical penial chamber showing tongue-like pustules forming a weak apical pilaster; A–C, QMMO86659, Mt Biggenden, SEQ. Scale bars = 10 mm.

Figure 6. Genital details of Figuladra bromileyorum sp. nov. A, Genitalia; B, Penis interior; C, Apical penial chamber showing verge and the apical raised pilaster of tongue-like pustules. A–C, QMMO86887, Dundowran, SEQ. Scale bars = 10 mm.

Figure 7. Genital details of Figuladra reducta (Iredale, 1937). A, Genitalia; B, Penis interior; C, Apical penial chamber showing rectangular pustules and a medial row of tongue-like pustules forming an apical raised pilaster. A, QMMO86815, Mt Perry, SEQ; B, QMMO86669, Goodnight Scrub, SEQ; C, QMMO77154, Kroombit Tops, SEQ. Scale bars = 10 mm.

Figure 8. Genital details of Figuladra vidulus sp. nov. A, Genitalia; B, Penis interior; C, Apical penial chamber showing a medial row of tongue-like pustules forming an apical raised pilaster. A, B, QMMO39795, The Hummock, SEQ; C, QMMO34327, Norval Park, SEQ. Scale bars = 10 mm.

Figure 9. Genital details of Figuladra robertirwini sp. nov. A, Genitalia; B, Penis interior; C, Apical penial chamber showing a medial row of tongue-like pustules forming an apical raised pilaster and two central longitudinal thickenings. A, QMMO86888, Round Hill Head, SEQ; B, QMMO39791, Town of 1770, SEQ; C, QMMO34975, Eurimbula NP, SEQ. Scale bars = 10 mm.

Figure 10. Genital details of Figuladra lessoni (Pfeiffer, 1846). A, Genitalia; B, Penis interior; C, Apical penial chamber showing a medial row of tongue-like pustules formed into a weak medial ridge. A–C, QMMO60270, Boyne Id. SEQ. Scale bars = 10 mm.

Figure 11. Genital details of Figuladra narelleae Stanisic & Potter, 2010. A, Genitalia; B, Penis interior; C, Apical penial chamber showing longitudinal rows of broad tongue-like pustules. A–C, QMMO86831, Bouldercombe Gorge, SEQ. Scale bars = 10 mm.

Figure 12. Genital details of Figuladra appendiculata (Reeve, 1854). A, Genitalia; B, Penis interior; C, Apical penial chamber showing crenellated pustules arranged in a V-shaped pattern and medial row of tongue-like pustules forming a weak medial ridge. A, B, QMMO7283, Mt Archer, MEQ; C, QMMO4173, Mt Dick, MEQ. Scale bars = 10 mm.

Figure 13. Genital details of Figuladra aureedensis (Brazier, 1872) comb. nov. A, Genitalia; B, Penis interior; C, Apical penial chamber showing crowded, small rectangular pustules arranged in a V-shaped pattern, gathered medially to form a weak raised pilaster. A, B, QMMO34259, Olsen′s Caves, MEQ; C, QMMO43382, Johannsen′s Cave, MEQ. Scale bars = 10 mm.

Figure 14. Genital details of Figuladra pallida (Hedley & Musson, 1892). A, Genitalia; B, Penis interior; C, Apical penial chamber showing crowded, small rectangular pustules arranged in a V-shaped pattern forming a weak medial ridge. A–C, QMMO71220, Polka Ck, Byfield, MEQ. Scale bars = 10 mm.

Figure 15. Genital details of Figuladra muirorum Stanisic, 2010. A, Genitalia; B, Penis interior; C, Apical penial chamber showing a medial row of tongue-like pustules forming a raised medial pilaster. A–C, QMMO76503, Boomer Ra, MEQ. Scale bars = 10 mm.

Figure 16. Genital details of Figuladra incei (Pfeiffer, 1846). A, Genitalia; B, Penis interior; C, Central penial chamber showing two prominent longitudinal ridges. A–C, QMMO39340, Middle Percy Id, MEQ. Scale bars = 10 mm.

Figure 17. Genital details of Figuladra barneyae. A, Genitalia; B, Penis interior; C, Apical central penial chamber showing central ridge and longitudinal row of bold spade-like pustules. A–C, QMMO70436, Connors Hump, MEQ. Scale bars = 10 mm.

Figure 18. Genital details of Figuladra finlaysoni sp. nov. A, Genitalia; B, Penis interior; C, Central penial chamber showing series of longitudinal ridges and longitudinal row of spade-like pustules. A–B, QMMO12864, Seaforth, MEQ; C, QMMO76990, Sarina, MEQ. Scale bars = 10 mm.

Figure 19. Genital details of Figuladra challisi. A, Genitalia; B, Penis interior; C, Apical half of penial chamber showing prominent longitudinal ridges and spade-like pustules. A–C, QMMO86829, St Bees Id, MEQ. Scale bars = 10 mm.

The vagina was typically shorter than the penis. Proportional lengths of the vagina and penis varied between species with all species having the penis at least double the length of the vagina. The greatest proportional difference was seen in F. narelleae where the penis was three times the length of the vagina. (Table S2). The bursa copulatrix stalk was slender and terminated in a large ovate head (not visible in dissection in all illustrations) that was located at the base of the albumen gland. The atrium, free oviduct, prostate, uterus and hermaphroditic duct were typically camaenid and without unusual features. The ovotestis was not dissected from the digestive gland and consequently not illustrated in some figures of dissections.

Penis interior (Figure 4C–F). Differences were observed and noted in three distinctive regions of the penial chamber: apical, central and lowermost (basal). The apical third typically features a short knob-like verge (sometimes retracted into the epiphallus and not shown in all illustrations) and crowded, rectangular to crenellated rectangular pustules often arranged in a V-shaped pattern. In species south of the St Lawrence Gap dry corridor [F. vidulus sp. nov., F. bromileyorum sp. nov., F. reducta (Iredale, 1937), F. robertirwini sp. nov., F. muirorum Stanisic, 2010, F. aureedensis (Brazier, 1872), medial tongue-like pustules formed a raised pilaster or prominent central ridge [F. lesson (Pfeiffer, 1846), F. appendiculata (Reeve, 1854), F. pallida (Hedley & Musson, 1892), Figure 4F]. In two species the apical pustules were spaced, bold and arranged in rows with the medial row of more pronounced tongue-like pustules sometimes forming a pilaster [F. bayensis (Brazier, 1876); Figure 5C] or not (F. narelleae: Figure 4E). In these two species, the rows of pustules descend to the central and basal regions of the penis (Figure 5B, Figure 11B). Species north of the corridor [F. incei (Pfeiffer, 1846), F. barneyae J. Stanisic 2010, F. finlaysoni sp. nov., F. challisi (Cox, 1873)] had crowded apical pustules but no prominent pilaster (Figures 16B–19B).

In the central area of the penis one or two thick, longitudinal ridge-like thickenings were present and accompanied by several shorter, thinner ridges arranged in a chevron pattern, and a row of tongue-like pustules. Three of the species north of the St Lawrence Gap (F. barneyae, F. finlaysoni sp. nov., F. challisi) had an added row of bold or fleshy spade-like pustules in the central area of the penial chamber (Figure 17C, Figure 18C, Figure 19C).

The basal third of the penial chamber varied marginally between species and typically consisted of thinner ridges arranged in a chevron pattern, with crowded pustules. This pustular zone occupied an extensive area of the basal penial chamber in F. incei (Figure 16B). Below this pustular zone thick (Figure 12B) or thin longitudinal thickenings (most species) enter the atrium.

Radulae and jaw (Figure 20). The radula had a central rachidian tooth present with numerous laterals and marginals arranged in transverse rows. The lateral teeth were unicuspid, gradually becoming bicuspid to tricuspid and marginals tricuspid to multicuspid. The jaw was odontognathous with approximately 11 regular, very strong ribs.

Figure 20. Scanning electron micrographs showing jaws, central and lateral parts of radula. A-C, Figuladra bayensis QMMO86659, Mt Biggenden, SEQ; D-F, Euryladra mattea QMMO86640, Arcadia Valley, SCQ. Scale bars = 100 um.

Patterns of shell variation

Size and shape of shell. The total range in shell diameter (Figure 21A) for adult Figuladra measured was 22.3–50 mm (n = 1014, Tables 4 and 6, Supplementary Table S1). The smallest diameter was recorded in F. appendiculata from Great Keppel Id, SEQ. The greatest shell diameter was observed in F. bayensis from the semi-evergreen vine thicket on limestone at Mt Biggenden, SEQ (maximum observed diameter 50 mm). However, smaller F. bayensis specimens occurred in the microphyll vine forests on volcanic substrate at Mt Mudlo, SEQ (mean maximum observed diameter 36 mm). Least variability in mean shell diameter was observed in F. aureedensis (var. 2 mm, n = 44) from the restricted and insular environments of the limestone outcrops, north of Rockhampton, SEQ and in most F. appendiculata (var. 19 mm, n = 77) populations from scattered and varied localities north of Rockhampton, including the Keppel Islands, MEQ. Other species to show little variation in mean shell diameter were the island-based F. challisi (var. 3 mm, n = 37) and F. muirorum (var. 4 mm, n = 44) from the Marlborough area, MEQ. The MEQ species (F. barneyae, F. incei, F. finlaysoni sp. nov., F. bayensis) showed a tendency to have greatly varied shell diameters.

Figure 21. Box and whisker plots showing comparison between shell features of species in the Figuladra clade. A, Shell diameter. B, Shell height. C, Height/Diameter ratio. D, Whorl count. Images A, B: measurements in cm.

The total range of adult shell height was 17–37 mm which is marginally less than the variation in the range of shell diameter (Table 4). The smallest recorded shell height was seen in F. finlaysoni sp. nov. (17 mm) and the largest in F. bromileyorum sp. nov. (37 mm). A greater number of species tended to show a greater variation in mean shell height than in mean shell diameter. These comprised species with relatively larger geographic ranges where a range of microhabitats could be exploited for living space such as F. robertirwini sp. nov. (var. 13 mm, n = 69), F. bromileyorum sp. nov. (var. 12 mm, n = 87), F. barneyae (var. 13 mm, n = 84) and F. appendiculata (var. 13 mm, n = 77). Least variation in mean shell height was observed in the geographically circumscribed F. challisi (var. 1 mm, n = 37) and F. aureedensis (var. 1 mm, n = 44).

The mean H/D ratio in adult Figuladra ranged from 0.697 in F. narelleae from Dan Dan Scrub-Mt Morgan area, SEQ to 0.87 in F. pallida from the Byfield area, north of Rockhampton, SEQ (Figure 21C). The highest individual H/D ratio (0.974) was recorded from a specimen of Figuladra robertirwini from Rosedale, SEQ. The H/D ratio can be affected by several factors, most notably spire protrusion, whorl count and coiling pattern (tightly or loosely). In the case of Figuladra, high whorl counts (above 6.25) correlate with high H/D ratios (F. barneyae, F. bromileyorum sp. nov., F. pallida). These three species are also distinguished by their strongly elevated spires that are tightly rather than evenly coiled. However, apart from these examples, there does not appear to be any overarching trend between coiling pattern, whorl count, spire protrusion and the H/D ratio (Table 5).

Table 5. Comparative overview of shell characters in Figuladra species (large: 17–40 mm; very large: > 40 mm).

Species	Size (diameter)	Shell shape	Spire -domed	Periphery	Colouration
SEQ upland:
F. bayensis	very large	depressedly globose	low-moderate	rounded	banded
F. narelleae	very large	depressedly globose	low	rounded	banded
SEQ:
F. bromileyorum	large	globose	moderate -high	rounded to subangulate	monochrome dark brown;banded
F. reducta	large	depressedly globose	low domed	rounded	banded
F. vidulus	large	globose	low-high	rounded to subangulate	monochrome dark brown;banded
F. robertirwini	large	globose	elevated	subangulate; body whorl strongly angulate	monochrome dark brown;bandedpale based
F. lessoni	large	globose	low – moderately	rounded to subangulate	monochrome chestnut brown;banded
F. appendiculata	large to very large	globose	low-elevated	rounded to subangulate	banded;dark;pale
F. aureedensis	large	globose	moderate	subangulate	banded;dark
F. pallida	large to very large	globose	moderate – high conical	subangulate; body whorl strongly angulate	pale – straw yellow;banded
F. muirorum	large	globose	moderate	subangulate	monochrome dark red-brown;banded
MEQ:
F. incei	large	globose	moderate	rounded to subangulate	monochrome dark brown;banded
F. barneyae	large	globose	elevated	subangulate; body whorl strongly angulate	banded
F. finlaysoni	large	globose	low	rounded	banded
F. challisi	large	globose	low	rounded	banded

Note: See Table 4 for comparative diameter measures.

Body whorl contour tended to be evenly rounded in depressedly globose species where the spire is low and domed (F. challisi, F. reducta, F. bayensis, F. narelleae). The majority of species had globose shells in which the spire was more elevated and exhibited both rounded and weakly angulate ( =  subangulate) shell peripheries (F. bromileyorum sp. nov., F. vidulus sp. nov., F. incei, F. aureedensis, F. lessoni, F. appendiculata, F. barneyae, F. finlaysoni sp. nov.). Exceptions were seen in the high-spired individuals in species such as F. pallida, F. bromileyorum sp. nov. and F. barneyae in which the body whorl becomes strongly angulate. High-spired individuals of F. pallida assumed an almost trochoidal appearance. Juveniles of all species had sharply angulate peripheries.

Mean whorl counts showed little variation between species, ranging from 5.875 in F. reducta to 6.675 in F. pallida and generally were within 0.750 of a whorl for all of the 15 species (Figure 21D). Greatest whorl numbers were seen in specimens of F. pallida (7.125), F. bromileyorum sp. nov. (7.000) and F. robertirwini sp. nov. (7.000), species with the largest shells. Smallest whorl counts were recorded in F. reducta (5.250) and F. finlaysoni sp. nov. (5.500). Within-species whorl counts varied by from 0.025 (F. aureedensis) to 1.215 (F. finlaysoni sp. nov.) with most species showing a variability of between three quarters (0.75) and one and an eighth (1.125) whorls.

Aperture, lip and umbilical features. The aperture in all species was ovately lunate and slanted laterally away from the vertical axis of the shell so that most of the opening is located to the side rather than underneath the shell spire. This was most pronounced in F. pallida. The lip was thickened and weakly reflected with an expanded columellar edge occluding some or all of the umbilicus in most species. In F. pallida, the lip is flared and slightly twisted. The umbilicus was very small (some F. pallida), or narrowly open and reduced to a lateral crack by the reflection of the columellar lip edge (most species). The extent of umbilical coverage may vary from half the diameter of the shell to only a hairline crack visible. In some species, the umbilicus was totally closed by the columellar callus (F. narelleae, F. finlaysoni sp. nov.).

Shell colour and banding pattern. Shell colour and spiral banding patterns were extremely variable in the 15 species studied, especially within species. In many cases, differently coloured and spirally banded individuals of a species were often found living together under the same rock or log. In others, unicoloured or spirally banded shells were specific to a particular locality. Most Figuladra species exhibited a combination of dark brown forms; a diverse array of spirally banded forms with few to many brown spiral bands and varying intensities of brown suffusion behind the lip and on the spire; and less commonly, pale yellow forms with few narrow brown spiral bands or lacking banding altogether. A small number of species showed consistency in shell colour and banding pattern. Species with primarily banded shells that also included darker forms comprised Figuladra bromileyorum, sp. nov., F. vidulus sp. nov., F. incei, F. appendiculata, F. barneyae and F. finlaysoni sp. nov.; species with primarily dark shells that also included banded forms comprised F. lessoni and F. muirorum; species with a mixture of banded and dark forms comprised F. aureedensis and F. pallida; and species with strictly banded forms comprised F. bayensis, F. narelleae and F. challisi.

The accompanying images in Figures 22–36 illustrate the more commonly encountered forms exhibited by the 15 Figuladra species which are discussed below. However, the following is a summary only and does not account for the many subtle variations that individual species exhibit.

Figure 22. Shell variation in Figuladra bayensis (Brazier, 1875). A, AMS C.17959, Helix (Hadra) bayensis Brazier, 1875, Wide Bay ( =  Mt Biggenden), SEQ, holotype; B, QMMO42777, Mt Biggenden, SEQ; C, QMMO86654, Mt Mudlo, SEQ. Scale bars = 10 mm or as indicated. Image A: Australian Museum.

Figure 23. Shell variation in Figuladra bromileyorum sp. nov. A, QMMO87485, Dundowran, SEQ, holotype; B, QMMO86848, White Cliffs, Fraser Id, SEQ; C, QMMO86761, Poona, SEQ; D, QMMO39247, Urangan, SEQ. Scale bars = 10 mm or as indicated. Image A: Queensland Museum.

Figure 24. Shell variation in Figuladra reducta (Iredale, 1937). A, AMS C.100653, Varohadra bayensis reducta Iredale, 1937, Tenningering ( =  Mt Perry), SEQ, holotype; B, QMMO86815, Mt Perry, SEQ; C, QMMO21618, Goodnight Scrub, SEQ. Scale bars = 10 mm. D, QMMO86689, Kroombit tops, SEQ; Image A: Australian Museum.

Figure 25. Shell variation in Figuladra vidulus sp. nov. A, QMMO87486, The Hummock, SEQ, holotype; B, QMMO54580, Norval Park, SEQ; C, QMMO12857, Woodgate, SEQ. Scale bars = 10 mm or as indicated. Image A: Queensland Museum.

Figure 26. Shell variation in Figuladra robertirwini sp. nov. A, QMMO87484, Pine Ck, SSE Turkey Beach, SEQ, holotype; B, QMMO86857, Pine Ck, SSE Turkey Beach, SEQ, paratype; C, QMMO72317, Miriam Vale, SEQ; D, QMMO34975, Middle Ck, Eurimbula NP, SEQ. Scale bars as indicated. Image A: Queensland Museum.

Figure 27. Shell variation in Figuladra lessoni (Pfeiffer, 1846). A, ZMB/Moll 260025a, Helix lessoni Pfeiffer, 1846, Port Curtis, SEQ, lectotype; B, Helix curtisiana NEWHM:HANM.1.3531, Boyne Id, SEQ, holotype; C, QMMO72292, Maurice Hill, Gladstone, SEQ; D, QMMO72177, Mt Larcom, SEQ; E, QMMO72207, Farmers Point, Facing Id, SEQ. Scale bars = 10 mm. Image A: ZMB; B: HMNT.

Figure 28. Shell variation in Figuladra narelleae. A, QMMO31294, holotype, Gavial Ck, SEQ; B, QMMO23330, Dan Dan Scrub, SEQ; C, QMMO64057, Horrigans Rd, Raglan, SEQ. Scale bars = 10 mm. Image A: Queensland Museum.

Figure 29. Shell variation in Figuladra appendiculata (Reeve, 1854). A, NHMUK1977033, Helix appendiculata Reeve, 1854, Australia ( =  Berserker Ra, MEQ), holotype; B, QMMO64098, Yeppoon, MEQ; C, QMMO29409, Koongal, MEQ; D, QMMO86924, Mt Jim Crow, MEQ; E, QMMO5004, North Keppel Id, MEQ; F, AMS C.17600, Helix (Camaena) praetermissi Cox, 1868, Cape Direction, NEQ (error = Keppel Id, MEQ), holotype. Scale bars = 10 mm or as indicated. Image A: NHMUK; Image F: Australian Museum.

Figure 30. Shell variation in Figuladra aureedensis (Brazier, 1872) comb. nov. A, AMS C.17604, Helix aureedensis (Brazier, 1872) Aureed Id, Torres Strait (error = Johannsen’s Cave, MEQ), holotype; B–C, QMMO4332, Johannsen’s Cave, MEQ (two forms: banded and dark from same population). Scale bars = 10 mm. Image A: Australian Museum.

Figure 31. Shell variation in Figuladra pallida (Hedley & Musson, 1892). A, AMS C.170704, Helix rockhamptonensis pallida Hedley & Musson, 1892, Rockhampton ( =  Byfield, MEQ), holotype; B, QMMO69927, Byfield, MEQ; C, QMMO36665, Shoalwater Bay Army Training Base, MEQ; D, QMMO71220, Polka Ck, Byfield, MEQ; E, QMMO79052, Jungle Track, Byfield, MEQ. Scale bars = 10 mm or as indicated. Image A: Australian Museum.

Figure 32. Shell variation in Figuladra muirorum Stanisic, 2010. A, QMMO78851, Boomer Ra, MEQ, holotype; B, QMMO74733, Marlborough, MEQ; C, QMMO54428, Glenprairie, MEQ. Scale bars = 10 mm. Image A: Queensland Museum.

Figure 33. Shell variation in Figuladra incei (Pfeiffer, 1846). A, NHMUK20200001, Helix incei Pfeiffer, 1846, Percy Ids, MEQ, lectotype; B, QMMO64867, South Percy Id, MEQ; C, QMMO34275, Stanage Bay, MEQ; D, QMMO76956, Hexham Id, MEQ (two forms: dark with yellow circum-umbilical patch and all yellow). Scale bars = 10 mm. Image A: NHMUK.

Figure 34. Shell variation in Figuladra barneyae. A, QMMO20061, Connors Hump, MEQ, holotype; B, QMMO34539, Greenhill, MEQ; C, QMMO36237, Beautrel Creek, Connors Ra, MEQ; D, QMMO86880, Collaroy Station, MEQ. Scale bars = 10 mm. Image A: Queensland Museum.

Figure 35. Shell variation in Figuladra finlaysoni sp. nov. A, QMMO87487, Halliday Bay, MEQ, holotype; B, QMMO31636, Hazelbrook, Pine Valley, MEQ; C, QMMO54312, Pine Mt, Nebo, MEQ; D, QMMO35782, Finlayson′s Point, Seaforth, MEQ. Scale bars = 10 mm. Image A: Queensland Museum.

Figure 36. Shell variation in Figuladra challisi. A, AMS C.583621, Helix challisi (Cox, 1873), Keswick Id, MEQ, lectotype; B, QMMO86828, Keswick Id, MEQ; C, QMMO86829, St Bees Id, MEQ. Scale bars = 10 mm or as indicated. Image A: Australian Museum.

The basic shell colours of spirally banded forms were tawny yellow and to a lesser extent off-white. To this may be added two prominent brown spiral bands (a supra-peripheral and a subsutural band) plus a variable number of narrower brown spiral bands, a brown suffusion behind the lip and a columellar brown colour patch of variable extent as seen in the shells of F. bayensis (Figure 22), F. reducta (Figure 24) and F. narelleae (Figure 28). This pattern also occurred in some F. robertirwini sp. nov. (Turkey Beach: Figure 26A, B) and F. appendiculata (Berserker Range: Figure 29A, C). Shells of F. bromileyorum sp. nov. (Dundowran: Figure 23B) and F. vidulus sp. nov. (Norval Park: Figure 25B) were strongly banded with a brown suffusion on the last half of the body whorl only. In some chiefly spirally banded species, a brown suffusion may envelop the entire body whorl and spire as seen in F. bromileyorum sp. nov. from Poona (Figure 23C), F. vidulus sp. nov. from Wartburg (Figure 25C), F. robertirwini sp. nov. from Eurimbula (Figure 26D), F. incei from Stanage Bay (Figure 33C) and F. barneyae from Greenhill (Figure 34B). The shells of F. incei from the type locality of Percy Island exhibited both light coloured, weakly banded forms and more darkly coloured forms (Figure 33A, B). In contrast, shells of F. challisi from Keswick and St Bees Islands are always strongly banded but lacking the brown suffusion behind the lip with the columellar colour patch considerably reduced in size (Figure 36). F. finlaysoni sp. nov. from Finlayson Point (Figure 35D) exhibited a spirally banded shell similar to F. challisi in a species with a shell that is usually variously suffused with brown throughout its range. Species with predominantly brown shells such as F. muirorum and F. lessoni usually had some degree of brown spiral banding visible on a paler spire as did the brown shells of chiefly spirally banded species (Figures 27 and 28). A yellow circum-umbilical patch is a feature of the brown shell form of F. aureedensis from the Caves (Figure 30A, C) but is also seen in F. appendiculata from the Keppel Sands-Yeppoon area including Mt Jim Crow (Figure 29D). Shells of F. barneyae from Beautrel Creek (Figure 34C) displayed a similarly coloured circum-umbilical patch. Pale yellow forms with minimal brown banding were present in F. aureedensis from the Johannsen’s Cave (Figure 30B), F. appendiculata from the Keppel Sands-Yeppoon area (Figure 29B) and F. pallida from Byfield (Figure 31A). All yellow or white shells without either banding or brown suffusion, colloquially termed ‘albino’ forms, are known to occur in several species.

The large variation in colour and banding patterns within species and the considerable overlap in these features between species makes identification from shell alone difficult, if not impossible. In many instances, locality may play a key role in species determination.

Shell sculpture. SEM investigation of both protoconch and teleoconch microsculpture revealed similarities between all Figuladra species (Figure 37). The protoconch on all species had weak, crowded, curved radial ridges. The teleoconch microsculpture consisted of fine, irregularly disposed radial, periostracal threads.

Figure 37. Scanning electron micrographs. A, Protoconch of Figuladra lessoni QMMO12196, Boyne Id, SEQ; B-F, Shell microsculpture. B, F. finlaysoni sp. nov. QMMO14795, Slade Pt, Mackay, MEQ; C, F. incei QMMO6973, Stanage Bay, MEQ; D, F. aureedensis QMMO34244, Johannsen′s Cave, MEQ; E, F. appendiculata QMMO29409, Koongal, MEQ; F, F. bayensis, QMMO11967, Mt Biggenden, SEQ. Scale bars: A as indicated; B-F = 500 µm.

Systematics

Figuladra Köhler & Bouchet, 2020

Varohadra Iredale, 1933: 45 (nomen nudum).

Figuladra Iredale, 1933: 45 (nomen nudum); Smith, 1992: 154 (nomen nudum); Stanisic et al., 2010: 470 (nomen nudum).

Figuladra Köhler & Bouchet, 2020: 4.

Type species. Helix incei L. Pfeiffer, 1846, by original designation.

Diagnosis.

External morphology: Shell large to very large, locally quite variable, maximum observed adult diameter 50 mm, whorl count of adults ranging from 5.5–7.0, the extremes correlating with shell diameter parameters. Shell depressedly globose to globose. Spire generally moderately to strongly elevated. Sutures weakly impressed. Apical sculpture of weak, curved radial ridges. Post apical sculpture of fine periostracal threads. Whorls of spire loosely to tightly coiled, normally angulated to rounded or a combination of both; juveniles always strongly angulate. Body whorl descending rapidly. Lip reflected, white with an expanded columella edge covering all or some of the umbilicus in most species. Aperture ovately lunate and slanted laterally away from the vertical axis of the shell. Umbilicus very small or narrowly open and reduced to a lateral crack by the reflection of the columella lip edge or totally closed. Basic shell colour tawny yellow, lighter in some species. Shell colour and spiral banding pattern extremely variable especially within species. Brown suffusion behind lip extending to last half of the body whorl and surrounding umbilicus. Species generally exhibit four colour and banding forms: primarily banded shells that also include darker forms, primarily dark shells that include banded forms, species with a mixture of both and strictly banded forms. Animal dark grey with a pinkish tinge and a pink mantle. Head wart located between superior tentacles.

Reproductive System: Penis (P) long to very long (1.2–5 cm), tapered and slightly expanded apically; penial sheath (PS) present; penial retractor muscle (PRM) inserting at the junction of the two arms of the epiphallus (E); epiphallus with a thin, slightly swollen ascending arm and thick, muscular descending arm entering penis through a simple pore; externally, a bundle of muscle fibres (MF) connecting the descending arm of the epiphallus to the head of the penis; vas deferens (VD) thin and attached to penial sheath by connective tissue; a short, thin, vestigial epiphallic flagellum (EF) present, situated near the epiphallus-vas deferens junction, tightly bound to the vas deferens (detached in dissections); vagina (V) short, approximately half the length of the penis; atrium (Y) simple; free oviduct (UV) similar in length to the vagina; bursa copulatrix (BC) simple with a slender stalk (SS) and a large ovate head situated at the base of the albumen gland (GG); prostate (DG), uterus (UT) and hermaphroditic duct (GD) without unusual features.

Penis interior. Apically with a knob-like verge, epiphallus entering through a simple pore. Penis with three chambers; apically, interior wall with variable configuration of pustules and pilasters at the inner penial wall; centrally, with large longitudinal ridges and several short ridgelets arranged in a chevron pattern; basally with small, irregularly shaped pustules, several thin, longitudinal ridges or corrugated thick longitudinal ridges entering the atrium. For more detail see penial chamber descriptions under each species.

Range. Eastern Queensland from north of Gympie, SEQ to south of the O′Connell River, MEQ, including off-lying islands from the South Cumberland Island group to Fraser Island, and west to the Great Dividing Range.

Remarks

Among eastern Australian camaenids, Figuladra is distinguished by its almost smooth microsculpture and the pinkish mantle. Genital anatomy separates Figuladra from other hadroid snail genera by having a band of muscular tissue connecting the head of the penis to the epiphallus. This feature is not present in any other hadroid genus. Figuladra is also separated from Euryladra and Brigaladra by the penial-vaginal ratio in which the vagina is considerably shorter than the penis.

SEQ species (south to north)

Figuladra bayensis (Brazier, 1875)

Helix (Hadra) bayensis Brazier, 1875: 2.

Varohadra bayensis.— Iredale, 1937: 34, pl. 3, Figure 13.

Sphaerospira appendiculata .— (Reeve, 1854): Smith, 1992: 154 (in part)

Figuladra bayensis.— Stanisic et al. 2010: 470, sp. 750, text figure

Taxonomic note. Figuladra bayensis was first figured by Cox (1868: 55, pl. 18, Figure 1) as a large, thin, numerously and finely banded variant of ′Helix incei′ from the Wide Bay district, SEQ before being described as a separate species by Brazier (1875).

Holotype. Wide Bay (restricted type locality = Mt Biggenden, SEQ), 25° 54′ S, 152° 6′ E, (AMS C.17595) J. Brazier collection, pre-1900 – original designation. Height of shell 30.22 mm, diameter 41.77 mm, H/D 0.723, whorls 6.25.

Material examined

Type material. See above. Other material (all SEQ). Clifton Ra, 25° 32.92′ S, 152° 28.174′ E: (QMMO86920, 2 RC). Deep Ck, Tony Farrell property 11.8 km west northwest of Biggenden SEQ, 25° 30.16′ S, 151° 55.58′ E: (QMMO86861, 2 RC). Gayndah, NE c. 3.5 km SW of Wetheron, 25° 34′ S, 151° 41′ E: (QMMO76046, 1 SC). Mt Biggenden, 25° 32′ S, 151° 50′ E: (QMMO11967, 27 RC), (QMMO19570, 17 RC), (QMMO42777, 34 RC); 25° 31.989′ S, 151° 58.835′ E: (QMMO86659, 3 SC/4 RC), (QMMO86912, 8 RC); 25° 32′ S, 151° 57′ E: (QMMO7963, 3 SC/32 RC). Mt Mudlo NP, 26° 01.374′ S, 152°14.308′ E: (QMMO86928, 8 SC/3 RC), 26° 04.844′ S, 152° 07.986′ E: (QMMO86654, 4 SC/1 RC). Mudlo Gap N of Kilkivan, 26° 1′ S, 152° 13′ E: (QMMO43556, 21 SC/13 RC). Mt Woowoonga, 25° 26.388′ S, 152° 05.934′ E: (QMMO86918, 5 RC), (QMMO86665, 1 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for additional AM records from AM database.

Description

Shell (Figure 22A–C). Very large, diameter 35.36–50.00 mm (mean 40.36 mm), height 23.75–37.15 mm (mean 28.27 mm), H/D ratio 0.67–0.76 (mean 0.70), whorls 5.87–6.25 (mean 6.00); depressedly globose with a low domed spire; whorls rounded, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus reduced to a chink; animal dark grey to black, head and dorsal area with brown tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 22). Shell tawny yellow with prominent broad subsutural and supra-peripheral bands; numerous narrower dark brown spiral bands continuous on base; narrow dark brown suffusion behind lip and surrounding umbilicus. Based on 23 measured adult specimens (Table S2).

Genital anatomy (Figure 5A). As for genus; vagina approximately half the length of the penis; PVR 1.9–2.3 (mean 2.1).

Penis interior (Figure 5B, C). Apically with knob-like verge (not visible in dissection) and several longitudinally arranged rows of crowded, prominent, wavy pustules (PP). Medial row of prominent tongue-like pustules forming distinct raised pilaster (PT); centrally a row of large tongue-like pustules (PPT), several adjacent rows of smaller pustules; large longitudinal ridge (PPL) and several short ridgelets (PPR) arranged in chevron pattern; basally with small, randomly disposed, irregularly shaped pustules (PPP); several thin, corrugated longitudinal ridges entering atrium. Based on 4 dissected specimens (Supplementary Table S2).

Distribution and habitat. From west of Maryborough to Gayndah in the Wide Bay region, SEQ; microphyll vine forest and semi-evergreen vine thicket, living under rocks and under or inside logs.

Key localities. Mt Biggenden, Mt Woowoonga, Mt Mudlo.

Remarks

In the molecular phylogeny (Figure 3), Figuladra bayensis is the sister species of F. narelleae. F. bayensis is distinguished from other Figuladra species by the molecular analyses that places F. bayensis as a sister group to F. narelleae (p-distance = 6.4%) in a subclade basal to other Figuladra species and separated from other Figuladra species by a p-distance >12% (Table 3). Figuladra bayensis is differentiated from other Figuladra species by penial chamber anatomy comprising several rows of prominent, wavy pustules and a medial row of prominent tongue-like pustules forming a distinct raised pilaster. The rows of prominent pustules are continuous in the central part of the chamber and comprise a row of large tongue-like pustules flanked by rows of smaller pustules. These penial chamber features are distinct from those of other members of the SEQ clade but most similar to those of F. narelleae from the Boyne and Mt Morgan Ranges, SW of Rockhampton, SEQ. Figuladra bayensis differs from F. narelleae by having much bolder apical pustules, a much smaller PVR, and the distinct raised pilaster in the apical chamber which is absent in F. narelleae. Conchologically, F. bayensis differs from F. bromileyorum sp. nov. by combination of larger size, lower spire and prolific banding, which is continuous on base of shell.

Common name. Biggenden banded snail

Figuladra bromileyorum L. Stanisic, sp. nov.

Holotype. Dundowran, Anson's Road, SEQ, 25° 16.192′ S, 152° 46.219′ E, coll. J. Stanisic, L. Stanisic, 14 Mar. 2018, (QMMO87485), under logs/on ground – here designated. Height of shell 37.45 mm, diameter 40.57 mm, H/D 0.923, whorls 6.500.

Paratypes. Same data as holotype (QMMO86677, 4 SC/26 RC). Dundowran Beach, SEQ, 25° 16.133′ S, 152° 46.33′ E, coll. J. Stanisic, L. Stanisic, 3 Jul. 2013, (QMMO86887, 33 SC/5 RC), under bark of tree, littoral rainforest.

Material examined.

Type material. See above. Other material (all SEQ). Poona Conservation Park, 25° 07.04′ S, 152° 34.12′ E: (QMMO86761, 1 RC, QMMO86923, 2 RC). Burrum Heads, Salford Scrub, 25° 11′ S, 152° 36′ E: (QMMO24245, 17 SC/1 RC). Eli Creek Rd, Hervey Bay, 25° 15.87′ S, 152° 48.92′ E: (QMMO86867, 3 RC). Tinnanbar, N of Tin Can Bay, 25° 46′ S, 152° 57′ E: (QMMO16523, 16 SC). Poona Ck, 25° 45.92′ S, 152° 51.88′ E: (QMMO86864, 1 RC). St Mary SF, SW of Maryborough, 25° 44.6′ S, 152° 28.45′ E: (QMMO39860, 1 RC), 25° 43.18′ S, 152° 29.05′ E: (QMMO39857, 11 SC/1 RC), 25° 43.18′ S, 152° 29.05′ E: (QMMO43408, 1 SC), 25° 42.7′ S, 152° 29.03′ E: (QMMO86874, 10 RC), 25° 41.42′ S, 152° 31.67′ E: (QMMO86863, 5 RC), 25° 41.12′ S, 152° 28.26′ E: (QMMO87067, 7 RC). South Aramara SF, 25° 38.52′ S, 152° 19.43′ E: (QMMO87064, 1 RC/2 SC). Ellerslie Rd, c. 39.9 km WSW of Maryborough, 25° 38.38′ S, 152° 19.2′ E: (QMMO86875, 6 RC). Woocoo NP via Ellerslie Rd, c. 38.9 km WSW of Maryborough, 25° 38.32′ S, 152° 19.55′ E: (QMMO86876, 4 RC), Raven Hills c. 3.8 km NNW of Poona, 25° 38.03′ S, 152° 53.6′ E: (QMMO86866, 1 RC). Fraser Id, Central Station, 25° 28.5′ S, 153° 03′ E: (QMMO86847, 5 RC). Fraser Id, Nth White Cliffs, 25° 24′ S, 153° 1′ E: (QMMO59985, 5 SC, QMMO86848, 5 RC). Fraser Id, 25° 15′ S, 153° 10′ E: (QMMO4673, 6 SC, QMMO5152, 1 SC), NW of Maryborough, 25° 23.7′ S, 152° 31.05′ E: (QMMO72357, 1 RC). Hervey Bay, 25° 17′ S, 152° 50′ E: (QMMO4546, 20 RC). Fraser Id, Orchid Beach, 24° 58′ S, 153° 18.98′ E: (QMMO61777, 10 SC, QMMO71562, 6 SC). Urangan c. 12 km SE at boat jetty, Ariadne St, River Heads, 25° 25.78′ S, 152° 55.37′ E: (QMMO87167, 4 RC). Urangan, Hervey Bay, 25° 17′ S, 152° 53′ E: (QMMO39247, 10 SC), 25° 17.23′ S, 152° 54.35′ E: (QMMO86865, 25 RC). Pialba Esplanade, 25° 16.88′ S, 152° 50.78′ E: (QMMO87063, 3 RC/1 SC). Dundowran Beach, Hervey Bay, 25° 16.13′ S, 152° 46.33′ E: (QMMO86862, 9 RC, QMMO86887, 33 SC/5 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for additional AM records from AM database.

Description

Shell (Figure 23A–D). Large, diameter 30.20–40.57 mm (mean 35.54 mm), height 24.64–37.45 mm (mean 29.92 mm), H/D ratio 0.74–0.93 (mean 0.84), whorls 6.25–7.00 (mean 6.50); globose with moderately elevated to high spire; whorls rounded to subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak, crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus partly open; animal dark grey, head and dorsal area with grey tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 23). Shell light to dark brown with orange-brown banding and paler spire; prominent dark subsutural band and numerous thinner dark brown bands; brown suffusion behind the lip and surrounding the umbilicus; occasionally with extensive brown suffusion on body whorl; some specimens from Poona and Fraser Island, SEQ are monochrome dark brown without any trace of banding (Figure 23C). Based on 76 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 6A). As for genus; vagina less than half the length of the penis; PVR 1.725–3.257 (mean 2.258).

Penis interior (Figure 6B, C). Apically with a knob-like verge (PV) and crowded crenellated rectangular pustules (PP) arranged in a V-shaped pattern and featuring a medial row of prominent tongue-like pustules forming a raised pilaster (PT); centrally a prominent thick longitudinal ridge (PPL), several thinner longitudinal ridges, and a single longitudinal row of large tongue-like pustules (PPT); basally an extensive zone of elongate ridgelets arranged in a chevron pattern (PPR); thin longitudinal thickenings entering the atrium. Based on 18 dissected specimens (Table S2).

Distribution and habitat. Hervey Bay region south of the Isis River to Inskip Point including Fraser Island, west to Aramara SF, SEQ; in vine thicket living under and inside logs, under rocks and under the bark of trees.

Key localities. Dundowran Beach, Fraser Id, Aramara SF, Pialba and Poona.

Remarks

Figuladra bromileyorum sp. nov. is distinguished from F. robertirwini sp. nov. (p-distance of 3.4%), by several visible differences in the genital anatomy, viz. in the apical chamber the pustules are crenellated as compared to rectangular in F. robertirwini sp. nov., a double longitudinal thickening (single in F. robertirwini sp. nov.) and the row of large tongue like pilasters in the central chamber which are missing in F. robertirwini sp. nov. Figuladra bromileyorum sp. nov. differs from the geographically neighbouring species F. bayensis and is separated by a genetic p-distance of 12.7%. Anatomically, the penis interior of F. bromileyorum sp. nov. differs significantly from that of F. bayensis which features a proliferation of prominent, wavy pustules in both the apical and central parts of the chamber. In F. bromileyorum sp. nov., the apical chamber has small crenellated pustules and a medial row of tongue-like pustules forming a raised pilaster; centrally the chamber has a single longitudinal thickening and only a single longitudinal row of tongue-like pustules; basally the extensive zone of chevron-arranged ridgelets of F. bromileyorum sp. nov. contrasts with irregularly disposed tiny pustules of F. bayensis. Conchologically, the globose shell with an elevated spire and rounded to subangulate whorls of F. bromileyorum sp. nov. differs from that of F. bayensis which has a low-spired, flatter shell with rounded whorls. Figuladra bromileyorum sp. nov. inhabits the coastal and sub-coastal regions of the Maryborough area, in contrast to F. bayensis which mainly inhabits the mountain ranges to the west.

Etymology. Named for the Bromiley family, my father′s family who were early settlers in the nineteenth century in the Hervey Bay area and well known for farming coffee, sugar cane, pineapples and timber.

Preferred common name. Dundowran banded snail

Figuladra reducta (Iredale, 1937)

Varohadra bayensis reducta Iredale, 1937: 34, pl.3, Fig. 15.

Sphaerospira appendiculata.— (Reeve, 1854): Smith, 1992: 154 (in part).

Figuladra reducta.— Stanisic et al., 2010: 472, sp. 748, Fig. in text.

Holotype. Tenningering, near Mt. Perry, W of Bundaberg, SEQ, 25° 15′49" S, 151° 53′19" E, J.C. Cox collection, pre-1912 (AMS C.100653) – original designation. Height of shell 24.50 mm, diameter 34.38 mm, H/D 0.713, whorls 6.000.

Paratypes. Same data as holotype (AMS C.79076, 2 RC).

Material examined.

Type material. See above. Other material (all SEQ).

Tellebang Mt, 24° 59.03′ S, 151° 13.62′ E: (QMMO71571, 5 SC). Yarrol Scrub, 30 km E Monto, 24° 54.28′ S, 151° 20.58′ E: (QMMO38976, 4 SC). Kroombit Tops NP Razorback Rd, 24° 24.95′ S, 150° 53.02′ E: (QMMO86784, 1 RC), Kroombit Tops NP The Barracks, 24.357°S, 150° 0.96′ E: (QMMO86683, 1 RC), 24° 21.42′ S, 150° 57.72′ E: (QMMO86683, 4 SC), Kroombit Tops NP Ironbark Ck, top falls, 24° 22.09′ S, 150° 54.02′ E: (QMMO86773, 1 RC), Kroombit Tops, upper Dry Ck, 24° 21.38′ S, 150° 58.33′ E: (QMMO77154, 1 RC). Ubobo, 7 km W Broom Ck, 24° 23.78′ S, 151° 13.95′ E: (QMMO87102, 2 SC/1 RC). Goodnight Scrub, 25° 16′ S, 151° 55′ E: (QMMO85417, 4 SC/1 RC), 25° 15.110′S, 151° 56.51′E: (QMMO86669, 1 SC/2 RC), 25° 15′ S, 151° 34′ E: (QMMO21618, 20 SC). Limestone Ck SF, W of Childers, 25° 15.08′ S, 151° 55′ E: (QMMO13123, 12 SC/34 RC). Gin Gin, c. 28 km SSW on Gin Gin-Gayndah Rd, 25° 15′ S, 151° 56′ E: (QMMO7966, 18 SC/37 RC) Mt Perry, NE at Scrub Rd, 25° 10.65′ S, 151° 39.84′ E: (QMMO86815, 2 SC/1 RC). Mulgildie Plateau, 25° 01.08′ S, 151° 11.87′ E: (QMMO65136, 28 RC).

Description

Shell (Figure 24A–C). Large, diameter 30.79–36.22 mm (mean 34.26 mm), height 21.58–27.95 mm (mean 25.02 mm), H/D ratio 0.68–0.76 (mean 0.73), whorls 5.25–6.25 (mean 6.00); depressedly globose with a low domed spire; whorls rounded, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, purplish to white; umbilicus narrowly open; animal dark grey, head and dorsal area with dark brown tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 24). Shell tawny yellow with prominent broad, brown subsutural band and narrower supra-peripheral band, and numerous narrower dark brown spiral bands continuous on the base; dark brown suffusion behind lip extending to and surrounding the narrow umbilicus. Based on 39 adult measured shells (Supplementary Table S1).

Genital anatomy (Figure 7A). As for genus; vagina less than half the length of the penis, PVR 2.1–2.8 (mean 2.3).

Penis interior (Figure 7B, C). Apically with rectangular pustules (PP) arranged in a V-shaped pattern and a medial row of prominent tongue-like pustules forming a raised pilaster (PT); centrally a thick longitudinal ridge (PPL), several thinner longitudinal ridges and a short row of tongue-like pustules (PPT) terminating in small, scattered knob-like pustules; basally with an extensive zone of numerous short ridgelets arranged in a chevron pattern (PPR) and scattered small pustules (PPP); several thin, longitudinal thickenings entering the atrium. Based on 12 dissected specimens (Table S2).

Distribution and habitat. Inland, north of the Burnett River in the Many Peaks, Dawes and Burnett Ranges, and the Goodnight Scrub, SEQ; semi-evergreen vine thicket, living under and inside logs and under rocks.

Key localities. Mt Perry, Goodnight Scrub, Monto, Kroombit Tops.

Remarks

Figuladra reducta is separated from its nearest relative on the molecular phylogeny, F. lessoni (p-distance 3.2%), by several differences in the penial architecture viz. the apical chamber has rectangular pustules as compared to diamond-shaped pustules in F. lessoni. The central chamber has one thick longitudinal thickening and F. lessoni has two; F. reducta has a short row of scattered knob-like pustules that are absent in F. lessoni. The uncorrected pairwise distances between F. reducta and F. bromileyorum sp. nov., a geographical neighbour, were low (3.5%) but the species are distinguishable by the shell and genital anatomy. Anatomically, the penis interior of F. reducta differs from that of F. bromileyorum sp. nov. by having a short central row of tongue-like pustules terminating in a series of small, irregularly disposed, knob-like pustules. Conchologically F. reducta is distinguished from F. bromileyorum sp. nov. by having a smaller, flatter shell with rounded whorls and a much lower H/D ratio (Figure 21). Results from molecular analyses (Figure 3) show F. reducta from Kroombit Tops and Ubobo (at the base of Kroombit) within the sub-clade containing F. lessoni. These specimens are included as F. reducta on the basis of broad similarity in shell and anatomical features. However, results from molecular analyses also show that F. reducta from the Goodnight Scrub and the Mt Perry localities are quite distinct from the Kroombit Tops material. Additional field work in the Dawes and Burnett Ranges, SEQ, is required to establish the true status of these populations. F. reducta from the Mt Perry district was originally described as a sub-species of F. bayensis on the basis of its smaller size and purplish-white lip. However, a genetic distance of 12.4% and a distinctly different penial architecture suggests these to be two separate species.

Common Name. Squat Cumberlands banded snail

Figuladra vidulus L. Stanisic, sp. nov.

Holotype. QMMO87486, The Hummock, c. 7 km NE Bundaberg, SEQ, 24° 51′ S, 152° 25′ E, coll. J. Stanisic, D. & N. Potter, 24 May 1990, remnant vine forest, under rocks, – herein designated. Height of shell 28.32 mm, diameter 33.96 mm, H/D 0.833, whorls 6.000.

Paratypes. (all SEQ). Same data as holotype (QMMO39795, 80 SC/155 RC). The Hummock, c. 7 km NE Bundaberg, 24° 51′ S, 152° 25′ E, SEQ, coll. T. Carless, 6 Dec. 1983, (QMMO14695, 24 RC), coll. J. Ferguson, 1990, (QMMO31621, 6 RC), under lantana bushes.

Material examined.

Type material. See above. Other material (all SEQ).

Bargara Caravan Park, 24° 48.49′ S, 152 27.48′ E: (QMMO87079, 5 SC/4 RC). Coonarr, end Davenport Drive, 24° 58.87′ S, 152° 28.90′ E: (QMMO87075, 1 SC/2 RC). Isis River (townsite), 23° 22 ′S, 150° 32′ E: (AMS C.112152, 3 RC). Mon Repos, 24° 48.07′ S, 152° 26.7′ E: (QMMO87076, 8 SC/20 RC). Moore Park, 24° 42.42′ S, 152° 15.7′ E: (QMMO86674, 2 RC). Norval Park, c. 40 km NW of Bundaberg, 24° 37′S, 152° 7′ 15” E: 4, 7 SC/43 RC). The Hummock, 24° 50.63′ S, 152° 25.56′ E: (QMMO86909, 2 SC/8 RC, QMMO87080, 5 SC/12 RC). Woodgate NP, 25° 07.21′ S, 152° 34.35′ E: (QMMO86759, 7 RC), 25° 07.26′ S, 152° 34.42′ E: (QMMO87078, 5 SC/1 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 25A–C). Large, diameter 31.33–35.35 mm (mean 33.17 mm), height 25.61–29.45 mm (mean 26.60 mm), H/D ratio 0.73–0.83 (mean 0.80), whorls 6.13–6.63 (mean 6.25); globose with an elevated, domed spire; whorls rounded to subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus partly open; animal dark grey, head and dorsal area with brown tentacles and pink-tinged foot, pink mantle.

Shell colour and pattern variation (Figure 25). Shell tawny yellow to brown with paler spire and darker brown banding; broad brown subsutural and narrower supra-peripheral bands; brown suffusion behind the lip and surrounding the umbilicus. Monochrome brown specimens occur at some localities (Woodgate, SEQ: Figure 25C). Based on 30 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 8A). As for genus; vagina less than half the length of the penis; PVR 1.5–2.7 (mean 2.5).

Penis interior (Figure 8B, C). Apically with a knob-like verge (not visible in dissection) and crowded elongate, rectangular pustules (PP) arranged in a V-shaped pattern, medially with tongue-like pustules forming a long, raised pilaster (PT); centrally two prominent longitudinal ridges (PPL), several smaller ridges and a few tongue-like pustules (PPT); basally an extensive zone of bold, elongate ridgelets arranged in a chevron pattern (PPR) and scattered, tiny pustules; thin longitudinal thickenings entering the atrium. Based on 15 dissected specimens (Table S2).

Distribution and habitat. Coastal and sub-coastal lowlands of the Bundaberg region between Kolan River and the Isis River, SEQ; in semi-evergreen vine thicket, living under and inside logs and under rocks.

Key localities. The Hummock, Bagara, Norval Park, Woodgate NP.

Remarks

The p-distances between the phylogenetically and geographically neighbouring species, Figuladra vidulus sp. nov., F. bromileyorum sp. nov. and F. reducta species averaged 3%–4%, possibly due to the occurrence of recent speciation, in which case species identity was confirmed by dissection. Anatomically, the penis of F. vidulus sp. nov. differs from that of F. bromileyorum sp. nov. by having two central longitudinal thickenings compared to one in the latter species. Compared to F. bromileyorum sp. nov. and F. reducta, the apical section of the penial chamber of F. vidulus sp. nov. features a series of long, crowded rectangular pustules compared to a series of shorter, discrete pustules in the former two species. Conchologically, F. vidulus sp. nov. is distinguished from F. bromileyorum sp. nov. by having a generally lower spire and smaller H/D ratio (Figure 21). The distribution of F. vidulus sp. nov. is bounded by the distributions of F. bromileyorum sp. nov. in the south, F. reducta in the west and F. robertirwini sp. nov. in the north.

The results of the molecular analysis include specimens from Bulburin SF, Colosseum Creek and Granite Creek in the F. vidulus sp. nov. clade. The penial anatomies of these records accord more with that of F. robertirwini sp. nov. as does their geographical distribution. Hence, these populations are herein included with those of F. robertirwini sp. nov. It is recommended that this phylogenetic anomaly be investigated further using additional genetic markers.

Etymology. From the Latin vidulus = suitcase, noun in apposition referring to a community of these snails that were found surviving in large numbers in an old, abandoned suitcase in the vine thickets of The Hummock, SEQ.

Preferred common name. Bundaberg banded snail.

Figuladra robertirwini L. Stanisic, sp. nov.

Holotype. Turkey Beach, c. 10.1 km SSE at Pine Ck area, SEQ, 24°09.97′ S, 151°41.65′ E, coll. J. Ferguson, 20 Nov. 2006, (QMMO87484), rainforest, under logs – herein designated. Height of shell 32.43 mm, diameter 38.95 mm, H/D 0.832, whorls 6.875.

Paratypes. Same data as holotype (QMMO86857, 3 RC). Turkey Beach, c. 3.6 km SSE at Tanti Road, SEQ, 24°07.12′ S, 151° 39.26′ E, coll. J. Ferguson, 19 Nov. 2006, under logs, (QMMO87858, 8 RC).

Material examined.

Type material. See above. Other material (all SEQ).

Blackman′s Ck, c. 55 km SSE, 24° 27.17′ S, 151° 23.6′ E: (QMMO72289, 72 SC/5 RC). Builyan railway station, ca.1.8 km E, 24° 32′ S, 151° 24′ E: (QMMO10316, 30 SC/5 RC). Dawes NP, W at headwaters of Deception Ck, 24° 32.67′ S, 151° 17.72′ E: (QMMO87004, 1 RC). Bulburin SF, Bobby Ra, 24° 29.75′ S, 151° 30.25′ E: (QMMO38959, 10 SC). Kolan R X’ing, c. 15 km NW of Bundaberg, 24° 48.08′ S, 152° 11′ E: (QMMO54587, 5 SC), 24° 42.7′ S, 151° 41.85′ E: (QMMO87163, 5 SC). Calliope, c. 29 km SSE Bloomfield Ck, 24° 13.75′ S, 151° 21.65′ E: (QMMO72281, 3 SC/6 RC). Calliope, c. 10 km SSE Taragoola Rd, 24° 5.07′ S, 151° 14.4′ E: (QMMO72282, 10 SC).Calliope, Old Ten Chain Rd, c. 4 km SE, 24° 0′ S, 151° 14.7′ E: (QMMO87148, 13 SC/3 RC). Colosseum Ck, 56 km SSE, 24° 24.85′ S, 151° 29.65′ E: (QMMO72164, 3 SC). Colosseum Ck, Blackmans Gap Rd, 24° 24.93′ S, 151° 29.75′ E: (QMMO87100, 1 SC). Colosseum Ck, c. 51 km SSE, 24° 23.65′ S, 151° 27.68′ E: (QMMO72319, 1 SC/2 RC). Deepwater NP, 24° 19.33′ S, 151° 57.73′ E: (QMMO86775, 1 RC). Euleijah Ck, c. 36 km ESE, 24° 28.8′ S, 151° 53.18′ E: (QMMO72310, 6 RC). Eurimbula, 16 km SE of Turkey Beach, 24° 10.22′ S, 151° 46.18′ E: (QMMO72209, 2 SC/1 RC). Eurimbula NP, 24° 11′ S, 151° 50′ E: (QMMO38289, 6 SC/3 RC, QMMO86921, 5 SC, QMMO86810, 22 RC, QMMO86811, RC, QMMO86808, 17 RC, QMMO54571, 43 SC/66 RC). Eurimbula NP, Middle Ck, 24° 8′ S, 151° 46′ E: (QMMO34975, 4 SC). Eurimbula NP, Bustard Beach, 24° 11′ S, 151° 50′ E: (QMMO34977, 5 SC). Granite Ck Forestry Rd c. 12 km from Bruce Highway, 24° 38.05′ S, 151° 34.35′ E: (QMMO87160, 10 SC/9 RC). Granite Creek Crossing, Bruce Hwy, 24° 36.76′ S, 151° 40.09′ E: (QMMO86822, 4 SC). Littabella Conservation Park, 24° 64.2′ S, 151° 95.89′ E: (QMMO87083, 1 RC). Rodd′s Peninsula, c. 5 km E of Turkey Beach, 24° 4.03′ S, 151° 41.57′ E: (QMMO72313, 13 SC), 24° 03.83′ S, 151° 41.42′ E: (QMMO86854, 5 RC), 24° 01.73′ S, 151° 41.88′ E: (QMMO86855, 2 RC), 24° 11′ S, 151° 50′ E: (QMMO86851, 2 RC). Rodds Peninsula, Ethel Rocks Rd, 24° 04.03′ S, 151° 41.45′ E: (QMMO86850, 7 RC). Rodds Peninsula, near NP gate, 24° 5.47′ S, 151° 42.32′ E: (QMMO87149, 2 SC/16 RC). Rodds Peninsula, Table Hill ridge crossing on peninsula track, 24° 01.3′ S, 151° 39.65′ E: (QMMO86853, 2 RC). Rodds Peninsula on W facing slopes of Table Hill, 24° 0.7′ S, 151° 39.7′ E: (QMMO86852, 5 RC). Rosedale, NW nr Lowmead, 24° 30.87′ S, 151° 46.02′ E: (QMMO67832, 1 RC, QMMO67834, 1 RC). Round Hill Head, 24° 09.02′ S, 151° 53.15′ E: (QMMO86888, 3 SC/10 RC). Seventeen Seventy, c. 50 km E Miriam Vale, 24° 9.08′ S, 151° 53′ E: (QMMO39791, 11 SC). Pine Ck, c. 18 km NNE Turkey Beach, 24° 13.55′ S, 151° 41.95′ E: (QMMO72317, 6 SC). Turkey Beach, 24° 05.39′ S, 151° 38.93′ E: (QMMO86914, 4 RC, QMMO87153, 6 SC/21 RC). Turkey Beach, c. 1 km W, 24° 5.39′ S, 151° 38.93′ E: (QMMO86793, 8 RC). Turkey Beach, c. 6 km W, 24° 6.98′ S, 151° 38.08′ E: (QMMO86795, 2 SC/1 RC). Turkey Beach, 8 km SE near Worthington Ck, 24° 7.85′ S, 151° 41.85′ E: (QMMO72210, 5 SC). Wartburg, N of Bundaberg, 24° 30′ S, 151° 55′ E: (QMMO17058, 2 RC). Watalgan SF, 24° 34.8′ S, 152° 0372′ E: (QMMO66599, 2 SC/RC), 24° 39.59′ S, 152° 01.19′ E: (QMMO87062, 11 RC/1 SC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 26A–D). Large, diameter 30.85–43.77 mm (mean 35.35 mm), height 23.61–36.83 mm (mean 28.83 mm), H/D ratio 0.64–0.97 (mean 0.81), whorls 6.25–7.00 (mean 6.50); globose with a low to high domed spire; whorls subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus partly open to closed; animal pinkish grey, head and dorsal area with brown tentacles, pink mantle.

Shell colour and pattern variation (Figure 26). Shell with variable colour forms: tawny yellow with thin brown banding, prominent brown supra-peripheral and subsutural bands generally present, brown suffusion behind the lip and on lower half of body whorl and surrounding umbilicus. Monochrome brown to dark brown; shells at Eurimbula NP, SEQ with paler spire. Shells with a pale base occur at Bustard Head, SEQ. Based on 73 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 9A). As per genus; vagina less than half the length of the penis; PVR 1.9–3.2 (mean 2.4).

Penis interior (Figure 9B, C). Apically with a knob-like verge (PV) and crowded diamond-shaped to rectangular pustules (PP) arranged in a V-shaped pattern, medially with a row of tongue-like pustules forming a raised pilaster (PT); centrally an attenuating, longitudinal row of tongue-like pustules (PPT), two prominent longitudinal thickenings (PPL) and several thinner ridges; basally an extensive zone of fine, longitudinal ridgelets (PPR) arranged in a chevron pattern; thin longitudinal thickenings entering the atrium. Based on 27 dissected specimens (Table S2).

Distribution and habitat. North of the Kolan River to Rodds Peninsula and west to the Many Peaks Range (Bobby Range), SEQ.

Key localities. Turkey Beach area, Rodds Peninsula, Granite Ck, Colosseum Ck, Eurimbula NP, Bulburin SF.

Remarks

Molecular analysis suggests that Figuladra robertirwini sp. nov. and F. vidulus sp. nov. are closely related (p-distance = 3.5%). Anatomically, F. robertirwini sp. nov. is distinguished from F. vidulus sp. nov. by having diamond-shaped pustules in the apical region of the penis as compared to rectangular pustules in the latter. F. robertirwini sp. nov. is further differentiated from F. vidulus sp. nov. by having more developed tongue-like pustules in the central part of the penial chamber and finer ridges and pustules in the lower half of the penial chamber. Conchologically, F. robertirwini sp. nov. differs most notably from both F. vidulus sp. nov. and F. lessoni by having a higher mean whorl count (mean 6.500, n = 73) compared with that of F. vidulus sp. nov. (mean 6.250, n = 30) and F. lessoni (mean 6.375, n = 84). In Eurimbula NP, SEQ, the shells of F. robertirwini sp. nov. from the microphyll vine forest on sand are monochrome dark brown in contrast to those in adjoining coastal dry vine thicket which are banded, with limited degrees of brown suffusion. The specimen from Bulburin SF is an anomaly as it occurs in the clade containing F. vidulus sp. nov. As the locality for this specimen from GenBank cannot be confirmed through checking museum specimens and the sequence was short, only 440 bp, a final decision was determined by morphology matching shell and genital anatomy to that of F. robertirwini sp. nov. Further investigation as to whether this is a cryptic species or a hybrid is warranted.’

Etymology. Named for Robert Irwin, conservationist and wildlife photographer, Australia Zoo.

Preferred common name. Robert Irwin′s banded snail

Figuladra lessoni (Pfeiffer, 1846)

Helix lessoni Pfeiffer, 1846: 71, sp. 571; Stanisic and Stanisic, 2020: 142–145, figs 7–9.

Helix rangii.— Lesson, 1830: 305.

Helix gulosa.— Gould, 1846: 17.

Helix curtisiana.— Pfeiffer, 1864: 528; Stanisic and Stanisic, 2020: 145–148, figs 10–12.

Helix seminigra.— Morelet, 1864: 289.

Helix basalis.— Schmeltz, 1869: 135 (nomen nudum).

Helix (Hadra) parsoni.— Cox, 1872: 18–20.

Helix incei var. lessoni.— Hedley and Musson, 1892: 556.

Varohadra lessoni.— Iredale, 1933: 45.

Varohadra curtisiana exedra.— Iredale, 1933: 45.

Sphaerospira incei curtisiana.— Smith, 1992: 156.

Sphaerospira incei lessoni.— Smith, 1992: 156.

Figuladra incei curtisiana.— Stanisic et al., 2010: 472, sp. 751, Figure in text.

Taxonomic note. Iredale (1933: 45) considered it was likely that Helix curtisiana and H. lessoni were identical. Stanisic and Stanisic (2020) provided conclusive evidence as to why these two taxa are one and the same and synonymy is here formally recognised. Figuladra lessoni has often been confused with F. incei in the literature. Molecular analyses clearly differentiate F. lessoni which inhabits the Greater Port Curtis region, SEQ from F. incei which inhabits the Percy Islands and Stanage Bay, MEQ.

Lectotype. ′Australia′ ( =  Port Curtis), Helix lessoni Pfeiffer, 1846. ex. Pfeiffer collection (ZMB/Moll 260025) – subsequent designation. Height of shell 32.55 mm, diameter 26.55 mm, H/D 0.815, whorls 6.5.

Paralectotype. Same data as lectotype (ZMB/Moll 260026).

Material examined.

Type material. See above, Port Curtis, SEQ, Holotype Helix curtisiana Pfeiffer, 1864, Angas collection, (NEWHM:HANM.1.3531). Other material (all SEQ). Ambrose, limestone quarry, c. 1 km S Ambrose railway station, 23° 47′ S, 150° 55′ E: (QMMO43535, 16 SC/5 RC). Machine Ck, Ambrose-Bracewell Rd, 8 km SW Mt Larcom c. 3° 50.32′ S, 150° 55.22′ E: (QMMO87142, 2 SC/4 RC). Henickes Rd, off Ambrose-Bracewell Rd, 23° 49.28′ S, 150° 55.62′ E: (QMMO86803, 1 SC/9 RC, QMMO86915, 6 RC), 23° 49.28′ S, 150° 55.62′ E: (QMMO87144, 2 SC/4 RC). Targinnie, 23° 49.46′ S, 151°06.71′ E: (QMMO86797, 32 SC/1 RC), 23° 47.2′ S, 151° 07.2′ E: (QMMO86798, 4 SC), 23° 47.25′ S, 151° 06.57′ E: (QMMO86839, 3 SC). Targinnie Rd, c .0.5 km past Forest Rd, 23° 47.22′ S, 151° 06.57′ E: (QMMO86807, 2 SC/1 RC). Targinnie at Swan Rd, 23° 45.75′ S, 151° 5.11′ E: (QMMO79918, 9 SC/2 RC). Tannum Sands, Canoe Point, opp. Wild Cattle Id, 23° 56.5′ S, 151° 22.08′ E: (QMMO87105, 1 SC/8 RC). Tannum Sands, Wild Cattle Creek walk, 23° 57.53′ S, 151° 22.97′ E: (QMMO87108, 8 SC). Scrubby Mt, c. 2.5 km SE of Gladstone, 23° 43.8′ S, 151° 2.67′ E: (QMMO74610, 7 RC). O’Connell Ridges, 12 km NNW of Gladstone, 23° 57.03′ S, 151° 16.63′ E: (QMMO72173, 21 RC) Mt Larcom, 23° 47.82′ S, 151° 06.29′ E: (QMMO86813, 1 RC), 23° 47.82′ S, 151° 06.29′ E: (QMMO86938, 2 RC), 23° 45.28′ S, 151° 3.6′ E: (QMMO72198, 1 SC/4 RC). Mt Larcom, Millar Rd, c. 20 km ENE Gladstone, 23° 49.25′ S, 151° 5.58′ E: (QMMO54527, 25 SC/23 RC), 23° 48′ S, 151° 7′ E: (QMMO85356, 1 RC). Mt Larcom, c. 2.1 km W Yarwun to Targinnie Rd, 23° 49.13′ S, 151° 5.6′ E: (QMMO39451, 14 SC/40 RC). Mt Larcom, 11k W Clay Ck, 23° 47.87′ S, 150° 52.38′ E: (QMMO72177, 13 RC). Mt Larcom, 1 km SW Bottle Tree Hill, 23° 48.95′ S, 150° 58.43′ E: (QMMO72211, 5 SC/1 RC). Mt Larcom, S at Scrub Ck, 23° 55.02′ S, 150° 58.43′ E: (QMMO80419, 1 RC). Monduran Ck, c. 30 km NW of Gladstone, 23° 39.62′ S, 151° 3.03′ E: (QMMO72329, 2 SC). Maurice Hill, c. 8.5 km S Gladstone, 23° 55.98′ S, 151° 14.38′ E: (QMMO72292, 3 SC/4 RC). Facing Id, Farmers Point, 10 km E Gladstone, 23° 46.9′ S, 151° 19.53′ E: (QMMO72207, 4 SC/26 RC). Curtis Id, 23° 37′ S, 151° 9′ E: (QMMO85827, 2 RC, QMMO85828, 2 RC). Mercy Hill, 24 km ENE Calliope, 24° 5.82′ S, 151° 24.9′ E: (QMMO72163, 7 SC). Bracewell, 23° 54′ S, 150° 54′ E: (QMMO23563, 18 SC, QMMO79347, 1 SC/1 RC). Boyne Id, 23° 55′ S, 151° 20′ E: (QMMO60270, 1 RC), 23° 54.57′ S, 151° 19.62′ E: (QMMO78028, 5 RC). Boyne Id, Wyndham Pk, 23° 57′ S, 151° 21′ E: (QMMO12196, 1 RC, QMMO87104, 5 SC/8 RC). Bororen, c. 1 km N on Bruce Hwy, 24° 13.91′ S, 152° 33.35′ E: (QMMO86791, 12 SC/6 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 27A–E). Large, diameter 31.18–38.36 mm (mean 34.83 mm), height 25.41–31.30 mm (mean 27.53 mm), H/D ratio 0.71–0.87 (mean 0.79), whorls 6.00–6.63 (mean 6.25); globose with low to moderately elevated, domed spire; whorls rounded to subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of extremely fine periostracal threads; lip reflected, white; umbilicus partly open; animal dark brown, head and dorsal area with brown tentacles and pink-tinged foot, pink mantle.

Shell colour and pattern variation (Figure 27). Variable; more common dark forms with shells dark red-brown with paler spire without orange-brown banding; some dark forms also have faint spiral banding present on the body whorl (Figure 27C). Banded forms have a tawny yellow shell with prominent broad subsutural and supra-peripheral bands and numerous narrower dark brown spiral bands, often darker on the last half of the body whorl (Figure 27D). Based on 84 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 10A). As for genus; vagina less than half the length of the penis; PVR 1.9–3.2 (mean 2.3).

Penis interior (Figure 10B, C). Apically with an extensive area of rectangular to diamond-shaped pustules (PP), arranged in a V-shaped pattern extending to the central part of the chamber and a medial row of raised tongue-like pustules forming a medial ridge (PT); centrally a longitudinal row of tongue-like pustules (PPT) and two prominent longitudinal thickenings (PPL); basally numerous small ridgelets arranged in a chevron pattern (PPR); thin longitudinal thickenings entering the atrium. Based on 21 dissected specimens (Table S2).

Distribution and habitat. Port Curtis region including Gladstone and the nearby islands, Mt Larcom, Bororen, and the more westerly limestone areas of Bracewell and Ambrose, SEQ; in semi-evergreen vine thicket, living under and inside logs and under rocks.

Key localities. Boyne Id, Curtis Id, Facing Id, Mt Larcom, Targinnie, Bracewell, Ambrose.

Remarks

Figuladra lesson is distinguished from F. aureedensis (p-distance 3%) in the same sub-clade by several differences in the penial anatomy viz. a more prominent raised pilaster in the apical chamber, the apical pustules, diamond-shaped in F. lessoni and rectangular in F. aureedensis, tongue like pustules at the lower end of the apical chamber which are absent in F. aureedensis and the absence of tongue-like pustules in the central penial chamber which are present in F. aureedensis. F. lessoni is separated from geographically neighbouring F. robertirwini sp. nov. by associating with a different sub-clade (p-distance of 5.2%). F. lessoni differs most notably from F. robertirwini sp. nov. in elements of the penial chamber. F. lessoni has a much more extensive area of apical pustules that extend well into the central part of the chamber compared with F. robertirwini sp. nov., whereas F. lessoni has a much shorter basal area of pustules than F. robertirwini sp. nov. (compare Figure 9B and Figure 10B). F. lessoni also lacks the formed pilaster of F. robertirwini sp. nov. instead having a medial ridge of tongue-like pustules. Conchologically, F. lessoni and F. robertirwini sp. nov. can be easily confused due to numerous similarly coloured and patterned variants of both species. The two species are essentially geographically separate with F. lessoni occurring north of the Boyne River, SEQ. The distribution of F. lessoni is bisected by the Calliope River, SEQ. To the south of the river the greater Port Curtis area basically supports monochrome brown snails with minimal banding. Localities north of the river (Mt Larcom, Ambrose, Bracewell) support specimens with an increasing amount of banding together with rarely occurring monochrome brown forms at the same locality (Mt Larcom, Targinnie). Further to the north, the Fitzroy River, SEQ geographically separates F. lessoni from F. appendiculata.

Reassigned common name. Port Curtis dark snail.

Note. Figuladra lessoni, previously known as the Yeppoon varicoloured snail, was considered by Stanisic et al. (2010) to occur in the Yeppoon region, SEQ. The species is herein restricted to the greater Port Curtis region and consequently has been assigned the alternative common name of Port Curtis dark snail to reflect the geographic distribution of the species more accurately.

Figuladra narelleae Stanisic & Potter, 2010

Figuladra narelleae Stanisic & Potter, 2010 (in Stanisic et al. 2010): 472, sp. 749, Figure in text.

Holotype. Gavial Ck, via Bouldercombe, W of Rockhampton, SEQ, 23° 36′ S, 150° 29′ E, coll. J. Stanisic, D. Potter, 18 May 1993, (QMMO31294), mvf on creek banks, under logs – original designation. Height of shell 28.64 mm, diameter 37.84 mm, H/D 0.757, whorls 5.875.

Paratypes (all SEQ). Same data as holotype (QMMO78850, 10 SC/12 RC), Mt Morgan, c .7 km E, 23° 38′ S, 150° 29′ E, coll. J. Stanisic, A. Green, 11 Jul. 1980, (QMMO8267, 20 SC/34 RC). Mt Moore, W side, NE Gladstone, 24° 2′ S, 151° 5′ E, coll. J. Stanisic, D. Potter, W.F. Ponder, O. Griffith, 5 Sep. 1982, (QMMO13211, 13 SC/46 RC). Dawson Hwy, c. 6 mls W of Calliope, 23° 59′57" S, 151° 5′58" E, coll. W.F. Ponder, L. Moore, 22 Jul. 1969, (AMS C.75134, 28 SC/1 RC), in scrubby bush on side of road. Dan Dan Scrub, SW of Calliope, 24° 12′ S, 151° 5′ E, coll. J. Stanisic, D. Potter, J. Chaseling, 27 Jun. 1989, mvf/sevt/Araucaria, under logs and rocks, (QMMO23330, 30 SC/34 RC). Dan Dan Scrub, SW of Calliope, 24° 10′ S, 151° 5′ E, coll. J. Stanisic, D. Potter, J. Chaseling, 27 Jun. 1989, (QMMO23391, 15 SC/15 RC), mvf/sevt/eucalypt emergent, under and in logs. Dan Dan Scrub, ca 30 km SW of Calliope, 24° 10′ S, 151° 5′ E, coll. J. Stanisic, D. Potter, W.F. Ponder, O. Griffith 5 Sep. 1982, (QMMO12217, 10 SC/72 RC, AMS C.136547, 20 SC/1 RC), sevt/Araucaria. Crocodile Ck near Bouldercombe Gorge, SW of Rockhampton, 23°35′37"S, 150°28.32′ E, coll. W.F. Ponder, M.V. Tippett, 22 Jul. 1969, (AMS C.75135, 15 RC), dense bush on sides of stony gully.

Material examined.

Type material. See above. Other material (all SEQ). Bouldercombe Ck, 23° 34′ S, 150° 28′ E: (QMMO12853, 5 RC). Boyne Ra, 21 km NNW of Calliope, 24° 10.66′ S, 151° 3.38′ E: (QMMO72193, 4 SC/1 RC), 24° 7.45′ S, 151° 02.83′ E: (QMMO80365, 5 RC). Boyne Ra, upper Neill Ck, 24° 7.37′ S, 151° 02.97′ E: (QMMO80388, 7 RC). Dan Dan Scrub, Boyne Ra, c. 30 km SW Calliope, 24° 10.67′ S, 151° 05.5′ E: (QMMO65155, 12 RC), 24° 10.22′ S, 151° 05.2′ E: (QMMO80332, 10 RC), 24° 9.65′ S, 151° 05.2′ E: (QMMO80352, 7 RC), 24° 08.71′S, 151° 04.56′E, (QMMO87422, 2 SC/4 RC), Tableland Rd, 24°10.38′ S, 151° 05.4′ E: (QMMO87428, 1 SC/1 RC). Dee Ra, northern spur, c. 5.2 km E Mt Morgan, 23° 38′ S, 150° 27′ E: (QMMO21496, 33 SC/1 RC). Dululu, c. 7 km NE on ridge opposite Piebald Mt, 23° 48′ S, 150° 21′ E: (QMMO23410, 5 SC/5 RC). Gavial Ck, via Bouldercombe, 23° 36′ S, 150° 29′ E: (QMMO31294, 1 SC). Hourigan Ck, 2 mls from junction with Ragan Ck, 23° 42′ S, 150° 49' E, (QMMO64057, 1 SC/1 RC). Mt Morgan, 10 km NE Bouldercombe Gorge, 23° 35.57′ S, 150° 28.27′ E: (QMMO72205, 10 SC). Mt Usher, 2 km SW Bouldercombe, 23° 35.6′ S, 150° 28.33′E, (QMMO86831, 7 RC/3 SC). Norton Ck, 38 km NNW of Calliope, 24° 19.48′ S, 151° 21.6′ E: (QMMO72184, 7 SC). Raglan, 23° 43′ S, 150° 49′ E: (QMMO7296, 2 SC). Stag Ck, WSW Biloela, 24° 15.95′ S, 150° 51.05′ E: (QMMO65165, 1 SC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 28A–C). Large to very large, diameter 33.38–40.80 mm (mean 38.04 mm), height 22.69–30.82 mm (mean 26.53 mm), H/D ratio 0.62–0.80 (mean 0.69), whorls 5.75–6.38 (mean 6.00); depressedly globose with a low domed spire; whorls rounded, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded curved radial ridge, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white with purple tinge; umbilicus closed to almost closed; animal dark grey, head and dorsal area with dark grey tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 28). Shell tawny yellow with prominent broad, dark brown subsutural band and a narrower brown supra-peripheral band, together with numerous thin brown spiral bands; brown suffusion behind lip extending onto the columella and surrounding the umbilicus. Based on 52 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 11A). As for genus; vagina about a third the length of the penis, PVR 2.0–3.7 (mean 3.0).

Penis interior (Figure 11B, C). Apically with a knob-like verge (not visible in dissection) and several longitudinally arranged rows of crowded, prominent, wavy pustules (PP) featuring a medial row of enlarged tongue-like pustules; centrally a row of large tongue-like pustules (PPT), several adjacent rows of smaller pustules, a large longitudinal ridge (PPL) and several short ridgelets (PPR) arranged in a chevron pattern; basally with several rows of tongue-like pustules and small, randomly disposed, irregularly shaped pustules below (PPP); several thin, corrugated longitudinal ridges entering the atrium. Based on 11 dissected specimens (Table S2).

Distribution and habitat. Boyne Range, Dee and Mt. Morgan Ranges and environs, west of Rockhampton, SEQ; semi-evergreen vine thicket and lowland rainforest with eucalypt emergents, living under and inside logs and under bark of trees.

Key localities. Bouldercombe Gorge, Dan Dan NP, Boyne Range, Struck Oil.

Remarks

Molecular analysis groups F. narelleae and its sister species F. bayensis in a subclade separate from mainstream Figuladra (p-distance >11.7%). Anatomically, F. narelleae has a penial chamber comprising rows of prominent tongue like-pilasters that extend to the basal third of the chamber. Apically, the medial row of prominent tongue-like pustules differentiates F. narelleae from other Figuladra species. The most similar penial anatomy is seen in F. bayensis from the Wide Bay area, SEQ but that differs from this species primarily by having very bold apical pustules lacking a distinct apical pilaster and a large longitudinal ridge with several short ridgelets in the central chamber. An additional distinguishing feature of the genitalia of F. narelleae is the large PVR showing a penial length three times that of the vagina. Conchologically, F. narelleae differs from neighbouring congeners by having a spirally banded tawny yellow shell with a prominent brown subsutural band and brown suffusion behind the lip. F. narelleae is a species that has often been mis-identified in private and museum collections as F. appendiculata based on shell characters alone (J. Stanisic, pers. obs.).

Common name. Mt Morgan banded snail.

Figuladra appendiculata (Reeve, 1854)

Helix appendiculata Reeve, 1854 (August): 373, pl.193, fig. 1353a, b.

Helix appendiculata.— Pfeiffer, 1854 (April 1855): 149.

Helix (Camaena) praetermissi.— Cox, 1868: p. 111, pl. 20, fig. 13; Stanisic and Stanisic, 2020: 148–149, figs 13, 14.

Helix thatcheri.— Cox, 1870: 170, pl. 16, fig. 2.

Helix lessoni var. appendiculata.— Pilsbry, 1890: 163. pl. 33, figs 71, 72.

Helix (Hadra) yeppoonensis.— Beddome, 1897: 123, figure in text.

Varohadra bernhardi.— Iredale, 1933: 45, pl.3, fig. 7.

Varohadra yeppoonensis.— Iredale, 1937: 31.

Varohadra bernhardi.— Iredale, 1937: 32.

Varohadra appendiculata.— Iredale, 1937: 34.

Varohadra praetermissi. – Iredale, 1937: 35.

Varohadra keppelensis.— Iredale, 1937: 33, pl. 3, fig.8.

Varohadra keppelensis degener.— Iredale, 1937: 33, pl. 3, fig.9.

Sphaerospira appendiculata.— Smith, 1992: 154.

Figuladra appendiculata.— Stanisic et al., 2010: 472, sp. 750, figure in text.

Figuladra incei lessoni.— (Pfeiffer, 1846): Stanisic et al., 2010: 472, sp. 752, figure in text [error].

Figuladra incei incei.— (Pfeiffer, 1846): Stanisic et al., 2010: 474, sp. 753, upper figure in text [error].

Taxonomic note. Reeve′s (1854) figured specimen in the NHMUK was labelled syntype and is herein designated as lectotype. The recently taken photograph of this specimen (Figure 29A) features a considerably faded shell but which on closer inspection still bears all the markings present in Reeve′s original illustration. Helix (Camaena) praetermissi Cox, 1868 (corrected type locality. Keppel Id), Helix (Hadra) yeppoonensis Beddome, 1897 (type locality. Yeppoon), Varohadra bernhardi Iredale, 1933 (type locality. Rockhampton) and Varohadra keppelensis Iredale, 1937 (type locality. North Keppel Id) and Varohadra keppelensis degener Iredale, 1937 (type locality. South Keppel Id) are herein placed in synonymy.

Lectotype. Australia ( =  Berserker Range, SEQ), H. Cuming collection, (NHMUK 1977033) – herein designated. Height of shell 35.11 mm, diameter 26.44 mm, H/D 0.754, whorls 6.375.

Material examined.

Type material. See above. Other material (all SEQ). North Keppel Id, 23° 4′ S, 150° 53′ E: (QMMO5004, 1 SC). Great (South) Keppel Id, 23°11.68′ S, 150°56.05′ E: (AMS C.107796, 9 RC), 23° 10.78′ S, 150°56.13′ E: (QMMO87120, 4 SC/52 RC), 23° 10′ S, 150° 57′ E: (QMMO9975, 1 SC). Cawarral, Rockhampton, Emu Park Rd, 23° 15′ S, 150° 40′ E: (QMMO19564, 32 SC/RC). Emu Park, Ritamanda Rd, 23° 13.85′ S, 150° 48.65′ E: (QMMO87113, 1 SC). Frenchville, foot of Mt. Archer, N. Rockhampton, 23° 20′ S, 150° 34′ E: (QMMO29405, 3 RC), 23° 20′ S, 150° 34′ E: (QMMO29418, 1 RC), 23° 20′ S, 150° 34′ E: (QMMO7224, 3 SC/2 RC), 23° 20′ S, 150° 34′ E: (QMMO7283, 6 SC/1 RC), 23° 20.67′ S, 150° 34.83′ E: (QMMO87469, 1 RC). Keppel Sands CP, 23° 20.32′ S, 150° 46.5′ E: (QMMO87110, 3 SC/2 RC). Koongal, N. Rockhampton, 23° 23′ S, 150° 33′ E: (QMMO29409, 2 RC), 23° 22′ S, 150° 33′ E: (QMMO6796, RC). Lakes Ck, N Rockhampton, 23° 23′ S, 150° 33′ E: (QMMO19864, 6 RC), Rockonia Rd, 23° 22.13′ S, 150° 33.82′ E: (QMMO87109, 7 SC/1 RC). Lamamoor Beach, Pilkington Park, 100m S of Robinson Rd/Scenic Highway intersection, 23° 9.22′ S, 150° 45.82′ E: (QMMO87119, 2 SC/4 RC). Mt Archer, Berserker Ra, 23° 21′ S, 150° 35′ E: (QMMO11663, 11 SC/RC, QMMO1469, 2 RC, QMMO30588, 3 RC), 23° 20′ S, 150° 34′ E: (QMMO11842, 1 RC), 23° 20.87′ S, 150° 34.04′E, (QMMO86833, 6 SC), 23° 20.67′ S, 150° 34.83′ E: (QMMO54512, 1 SC/4 RC). Mt Dick, Berserker Ra, 23° 22′ S, 150° 35′ E: (QMMO4173, 6 SC, QMMO4174, 3 RC). Mt Jim Crow, 23° 13.17′ S, 150° 37.75 E, (QMMO86924, 10 SC/9 RC). Mulambin Beach, Bluff Point NR, 23° 11′ S, 150° 47′ E: (QMMO4620, 4 SC), 23° 11.12′ S, 150° 47.55′ E: (QMMO87114, 7 SC/10 RC). Yeppoon, 23° 3.33′ S, 150° 46.03′ E: (QMMO78009, 10 RC), 23° 7′ S, 150° 44′ E: (QMMO64098, 59 SC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 29A–F). Large to very large, diameter 22.30–41.19 mm (mean 34.33 mm), height 17.57–30.96 mm (mean 26.41 mm), H/D ratio 0.71–0.83 (mean 0.77), whorls 5.88–6.50 (mean 6.25); globose with a low to elevated domed spire; whorls rounded to subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak, crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus almost closed; animal brown to grey, head and dorsal area with brown to grey tentacles, pinkish tail and pink mantle.

Shell colour variation (Figure 29). Highly variable. Banded forms are tawny yellow with a broad brown subsutural band and numerous thinner chestnut bands; dark brown suffusion behind lip extending to the columella and around umbilicus (Figure 29C); shells on Great Keppel ( =  South) Island have an indistinct supra-peripheral band and are often darker on the top half of the shell (Figure 29F); Shells from Mt Jim Crow are dark brown with faint banding visible and a pale yellow circum-umbilical patch (Figure 29D). Pale forms with a basically yellow shell have pale chestnut bands (Figure 29B, E). Based on 76 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 12A). As for genus; penial-vaginal ratio variable but averaging double, PVR 1.4–3.3 (mean 2.1).

Penis interior (Figure 12B, C). Apically with a knob-like verge (not visible in dissection), elongate, crenellated pustules (PP) arranged in a V-shaped pattern, a row of tongue-like pustules forming a weak medial ridge (PT); centrally with a short row of tongue-like pustules (PPT), a thick longitudinal ridge (PPL), and several thinner longitudinal ridgelets (PPR) arranged in a chevron pattern; basally with smaller pustules (PPP); two, thick, corrugated longitudinal ridges (PPC) entering the atrium. Based on 10 dissected specimens (Table S2).

Distribution and habitat. Berserker Range; Yeppoon and environs; Keppel Island group, MEQ; microphyll vine forest (Berserker Range), coastal semi-evergreen vine thicket and palm forest (Yeppoon and environs; Keppel Island group); living in and under logs, in palm fronds and at the base of fig trees.

Key localities. Mt Archer, Frenchville, Mulambin Beach, Yeppoon, Keppel Sands, Keppel Is.

Remarks

Figuladra appendiculata is distinguished from other members of the sub-clade (F. bromileyorum sp. nov., p-distance 3.5%, F. vidulus sp. nov. p-distance 3.2%, and F. robertirwini sp. nov. p-distance 4.1%) by the penial chamber architecture. The apical chamber is distinctive with long, crenellated pilasters in a chevron shape that unlike the three congeners form a very weak ridge (see associated remarks) and the presence of two thick corrugated ridges in the basal chamber not seen in any other SEQ clade species. F. appendiculata is separated from the nearby limestone-restricted F. aureedensis (p-distance of 4.3%) anatomically, with penial chamber features comprising apical crenellated pustules as compared to rectangular pustules of the latter, tongue-like pustules that form a weak ridge and the two basal corrugated thickening previously mentioned. The extensive synonymy which covers a wide range of shell colour and pattern is based on alignment of molecular analysis and genital anatomy. Conchologically, F. appendiculata displays a diverse array of shell colour and banding patterns that have formerly led to a proliferation of names. Shell colour and banding pattern in neighbouring and oft-confused F. narelleae (p-distance 12.6%) and the mainly limestone associated F. aureedensis are strikingly similar to those of some F. appendiculata. The molecular phylogeny (Figure 3) shows an extremely tight clustering of populations from diverse localities despite the highly variable shell colour and patterns comprising the populations sampled.

Common name. Berserker Range banded snail.

Figuladra aureedensis (Brazier, 1872) comb. nov.

Helix (Camaena) aureedensis Brazier, 1872: 640; Stanisic and Stanisic, 2020: 149-151, figs 15, 16.

Helix incei var. aureedensis.— Hedley and Musson, 1892: 556.

Varohadra aureedensis.— Iredale, 1937: 32.

Sphaerospira incei lessoni.— (Pfeiffer, 1846); Smith, 1992: 157 (in part)

Figuladra incei lessoni.— Stanisic et al, 2010: 538 (in part)

Holotype. Aureed Id, Torres Strait (error! reassigned type locality = Johannsen’s Cave, MEQ see Stanisic and Stanisic, 2020: 151), Hargraves collection (AMS C.17604) – original designation. Height of shell 27.54 mm, diameter 39.26 mm, H/D 0.705, whorls 6.25.

Material examined.

Type material. See above. Other material (all MEQ). Johannsen′s Cave, c. 30 km N Rockhampton, 23° 8.38′ S, 150° 28.02′ E: (QMMO21601, 63 SC/24 RC, QMMO34244, 55 SC/24 RC, QMMO43382, 100 SC/111 RC, QMMO64066, 21 SC), 23° 09.45′ S, 150 27.51′ E: (QMMO86835, 8 SC/3 RC, QMMO87468, 2 RC), 23° 9.43′ S, 150 27.52′ E: (QMMO87121, 8 SC/1 RC). Olsen′s Caves, c. 29 km N Rockhampton, 23° 10′ S, 150° 29′ E: (QMMO21586, 84 SC/19 RC), 23° 10′ S, 150° 30′ E: (QMMO34259, 22 SC/20 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 records from the AM database.

Description

Shell (Figure 30A–C). Large, diameter 34.70–35.77 mm (mean 35.07 mm), height 26.11–28.11 mm (mean 26.97 mm), H/D ratio 0.75–0.81 (mean 0.77), whorls 6.00–6.25 (mean 6.13); globose with moderately domed spire, whorls weakly subangulate, body whorl descending rapidly in front, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus partly open; animal brown, head and dorsal area with brown tentacles and pink foot, pink mantle.

Shell colour variation (Figure 30). Shell colour variable. Tawny yellow with narrow spiral chestnut bands, supra-peripheral band prominent; or with dark brown suffusion on upper half of shell with banding visible on first few whorls, supra-peripheral chestnut band visible through suffusion, tawny yellow circum-umbilical patch present. Based on 44 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 13A). As for genus; vagina about half the length of the penis, PVR 1.8–2.9 (mean 2.2); free oviduct (UV) shorter than vagina.

Penis interior (Figure 13B, C). Apically with a knob-like verge (not visible in dissection) and numerous crowded, small rectangular pustules (PP) gathered medially to form a weak pilaster (PT); centrally with a longitudinally arranged series of tongue-like pustules (PPT), a prominent longitudinal ridge (PPL), and several thinner longitudinal ridges; basally with several ridgelets (PPR) arranged in a chevron pattern and numerous, bold, irregularly shaped pustules (PPP); thin longitudinal ridges entering the atrium. Based on 10 dissected specimens (Table S2).

Distribution and habitat. Mt Etna NP and associated limestone habitats, MEQ; semi-evergreen vine thicket, living under rocks and logs.

Key localities. Johannsen′s Cave, Mt Etna Caves, Olsen′s Caves, Cammoo Caves.

Remarks

Figuladra aureedensis comb. nov. is distinguished from the other member of the sub-clade containing F. lessoni (see comparative remarks for F. lessoni). Molecular analysis separates the populations of F. aureedensis from those of the geographically neighbouring F. appendiculata (p-distance = 4.3%) even though the shells of F. aureedensis are very similar to those of the former. Features of the penial chamber readily separate the two species. The numerous small pustules in the apical penial chamber of F. aureedensis contrast with the crenellated pustules arranged in a V-shaped pattern seen in F. appendiculata; the apical pilaster formed by the gathered pustules in F. aureedensis differs from the weak medial ridge formed by tongue-like pustules of F. appendiculata; and the basal penial chamber of F. aureedensis lacks the two thick corrugated ridges of F. appendiculata. Conchologically, F. aureedensis is easily confused with neighbouring F. appendiculata. The shell colour and banding patterns of the two forms of F. aureedensis are also present in populations of F. appendiculata. F. aureedensis is endemic to the limestones of Mt Etna NP and highlights the importance of these geographically rocky outcrops to land snail evolution in the greater landscape.

Preferred common name. Fitzroy Caves banded snail.

Figuladra pallida (Hedley & Musson, 1892)

Hadra rockhamptonensis var. pallida Hedley & Musson, 1892: 556.

Varohadra rockhamptonensis.— (Hedley & Musson, 1892: 556); Iredale 1937: 31 (in part).

Sphaerospira incei incei.— (Pfeiffer, 1846); Smith, 1992: 156 (in part).

Figuladra incei incei.— (Pfeiffer, 1846): Stanisic et al., 2010: 474, sp. 753, lower figure in text [error].

Figuladra pallida.— Stanisic et al., 2010: 474, sp. 754, figure in text.

Holotype. Rockhampton (reassigned type locality herein = Byfield NP, MEQ), coll. C.T. Musson, Sept. 1887 (AMS C.170704) – original designation. Height of shell 30.52 mm, diameter 33.66 mm, H/D 0.907, whorls 6.75.

Material examined.

Type material. See above. Other material (all MEQ). NE Shoalwater Bay at Pinetrees Pt, 22° 20′ S, 150° 38′ E: (QMMO54841, 6 RC). Island Head, 70m N. of Yeppoon, 22° 20′S, 150° 39′E, (QMMO6803, 2 RC). N side of Island Head Ck, 22° 22.56′ S, 150° 38.27′ E: (QMMO86007, 8 RC). Shoalwater Bay, Army Base, 22° 23′ S, 150° 41′ E: (QMMO33665, 2 SC/15 RC). Shoalwater Bay, 22° 45′ S, 150° 21′ E: (QMMO24247, 1 SC). Parnassus Ra, W side, 60m, 22° 45′ S, 150° 24′ E: (QMMO24248, 5 SC), upper Reaches Nob Ck, 22° 51′ S, 150° 39′ E: (QMMO19681, 7 RC/1 SC). Byfield, Polka Rd c. 0.6 km S of Castle Rock Rd, 22° 49.98′ S, 150° 37.57′ E: (QMMO87117, 2 SC/2 RC). Byfield, Polka Ck, c. 15 km NW of Bowenia Forestry Camp, 100m, 22° 50′ S, 150° 40′ E: (QMMO25944, 4 SC), 22° 50′ S, 150° 37.28′ E: (QMMO71220, 29 SC/RC). Byfield NP, 22° 51′ S, 150° 39′ E: (AMS C.51655, 4 RC), 22° 52′ S, 150° 44′ E: (QMMO79052, 1 RC). Byfield SF, Stony Ck, 22° 55′ S, 150° 39′ E: (QMMO31407, 1 RC), upper Stoney Ck camping grounds, N of Rockhampton, 22° 55′ S, 150° 41′ E: (QMMO69927, 3 RC). Byfield SF, Farnborough Rd, 8.5 km S Byfield Rd intersection, 22° 57.25′ S, 150° 40.77′ E: (QMMO87115, 1 RC), c. 40 km N Yeppoon Byfield-Yeppoon Rd, 22° 50.92′ S, 150° 38.92′ E: (QMMO54523, 8 SC/1 RC). See Supplementary Material, Appendix 1 for additional QM material studied.

Description

Shell (Figure 31A–E). Large to very large, diameter 33.54–40.79 mm (mean 36.38 mm), height 27.42–34.15 mm (mean 31.12 mm), H/D ratio 0.81–0.93 (mean 0.86), whorls 6.50–7.13 (mean 6.63); globose with a moderately elevated spire to high conical spire; body whorl descending rapidly in front; whorls subangulate, sutures impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip strongly reflected, white; umbilicus narrowly open to almost closed; animal grey, head and dorsal area with dark tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 31). Highly variable. Shell straw yellow to dark tawny yellow with pink spire, sometimes with indistinct brown spiral bands (Figure 31A); or shell chestnut brown with indistinct brown spiral bands and tawny yellow circum-umbilical patch (Figure 31B); or shell yellow with prominent chestnut supra-peripheral and subsutural spiral bands and pale yellow base (Figure 31C–E). Based on 42 measured shells (Supplementary Table S1).

Genital anatomy (Figure 14A). As for genus; vagina less than half the length of the penis, PVR 1.9–3.9 (mean 2.8).

Penis interior (Figure 14B, C). Apically with crowded rectangular pustules (PP) arranged in a V-shaped pattern, crowded medially to form a weak medial ridge (PT); ridge extending centrally and terminating in row of weak tongue-like pustules (PPT) adjacent to a prominent thick longitudinal ridge (PPL); basally an extensive zone of short rectangular ridgelets arranged in a chevron pattern (PPR) and numerous randomly arranged small pustules; thick longitudinal thickenings entering the atrium. Based on nine dissected specimens (Table S2).

Distribution and habitat. North of Yeppoon in Byfield NP and surrounding areas of the Shoalwater Bay region, Peninsula Range and off-shore islands, MEQ; semi-evergreen vine thicket and riparian rainforest, living under logs and rocks.

Key localities. Parnassus Range, Polka Creek, Byfield SF, MEQ.

Remarks

Molecular analysis separates the Polka Creek Rd, Byfield NP population of Figuladra pallida from other species in the SEQ clade (p-distance = 4–13%, average = 6.2%). The medial ridge in the apical penial chamber of F. pallida is similar to that of F. appendiculata but differs from that of F. aureedensis. The shell shape, colour and banding pattern of F. pallida are variable but separate this species from its congeners. The core distribution of F. pallida appears to be centred on the Parnassus Range. The populations from the Peninsula Range, Island Head and nearby off-shore islands (MEQ) are tentatively included here. These populations need further study dependent on the availability of fresh material for dissection and molecular analysis.

Common name. Pale Parnassus Range snail

Figuladra muirorum Stanisic, 2010

Figuladra muirorum Stanisic, 2010 (in Stanisic et al. 2010): 476, sp. 756, figure in text.

Holotype. Boomer Ra, c. 1.5 km W of Comanche Stn, Gogango – Glenroy Rd, MEQ, 23° 20.05′ S, 149° 51.8′ E, coll. J. Stanisic, D. & N. Potter, 18 May 1993, sevt, under rocks, (QMMO78851) – original designation. Height of shell 26.57 mm, diameter 31.96 mm, H/D 0.831, whorls 5.5.

Paratypes (all MEQ). Same data as holotype (QMMO46318, 41 SC/54 RC). Marlborough, c. 14 km N, Glenprairie Rd, 22° 43′ S, 149° 53′ E, coll. J. Stanisic, D. Potter, 28 Apr. 1990, woodland, under logs and rocks (QMMO34404, 27 RC). Marlborough, c. 6 km N, Glenprairie Rd, 22° 45′ S, 149° 54′ E, coll. J. Stanisic, D. Potter, 29 Apr. 1990, remnant vine thicket, under rocks and logs, (QMMO34409, 15 SC/8 RC). Marlborough, c. 7.5 km SW, old Marlborough-Sarina Rd, 22° 52′ 6" S, 149° 50′ 6" E, coll. J. Stanisic, D. Potter, 16 May 1993, degraded vine thicket/eucalypt under logs, (QMMO46279, 13 SC/21 RC).

Material Examined.

Type material. See above. Other material. (All MEQ) Boomer Ra, at Mongrel Scrub, 23° 12′ S, 149° 46′ E, (QMMO76503, 1 RC). Boomer Ra, at Python Scrub, 23° 12′ S, 149° 44′ E, (QMMO77392, 7 SC). Marlborough, Coorumburra Rd, 285m, 21° 30.31′ S, 149° 07.34′ E, (QMMO87431, 1 SC/2 RC). Marlborough, c. 19 km N on Glenprairie Stn, 22° 39.5′ S, 149° 52.58′ E, (QMMO54428, 1 SC/38 RC). Marlborough, 7.7 km N on Glenprairie Rd, 100m, 22° 45.3′ S, 149° 54.22′ E, (QMMO87137, 1 SC/2 RC). Rockhampton, NW, near Marble Ridges Stn, 23° 11.48′ S, 150° 7.47′ E, (QMMO78030, 34 RC). Rockhampton, NW on Eden-Bann Rd, 23° 4.65′ S, 150° 13.13′ E, (QMMO78092, 8 SC/9 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 32A–C). Large, diameter 30.78–32.27 mm (mean 31.57 mm), height 24.50–27.91 mm (mean 25.95 mm), H/D ratio 0.77–0.87 (mean 0.82), whorls 5.63–6.13 (mean 6.00); globose with a moderately domed spire; whorls subangulate, sutures impressed; body whorl descending rapidly in front; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white to white with a slight purple tinge; umbilicus partly open; animal dark grey to black, head and dorsal area with brown to black tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 32). Shell dark brown with pale apex and indistinct spiral bands on early whorls grading to monochrome dark red-brown on the body whorl; or pale yellow with prominent brown subsutural and supra-peripheral bands, brown suffusion behind the lip extending to the last half of the body whorl, pale umbilicus (Figure 32C). Based on 58 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 15A). As for genus; vagina less than half the length of the penis, PVR 2.2–2.8 (mean 2.3).

Penis interior (Figure 15B, C). Apically with a knob-like verge (not visible in dissection), crowded rectangular pustules arranged in a V-shaped pattern and a medial row of tongue-like pustules forming a prominent raised pilaster (PT); centrally a short, longitudinal row of weak tongue-like pustules (PPT), several, widely spaced, thick longitudinal ridges (PPL) and smaller ridgelets; basally with longitudinal ridgelets arranged in a chevron pattern (PPR); thin longitudinal thickenings entering the atrium. Based on 5 dissected specimens (Table S2).

Distribution. Eden-Bann, Boomer Range to Mt Slopeaway, Broadsound Range and Marlborough environs (including Pine Mts), MEQ; semi-evergreen vine thicket and scattered remnants, living inside hollow trees and under logs.

Key localities. Marble Ridges, Glenprairie Stn, Pine Mountains, Mt Slopeaway, Marlborough, MEQ.

Remarks

Molecular analysis and genital anatomy associate Figuladra muirorum with Figuladra species of the SEQ clade south of the Fitzroy River, MEQ (p-distance 4.4–12.6%, average = 7%) rather than neighbouring species north of the river (p-distance 12–13%, average = 12.7%). Anatomically, the prominent raised apical pilaster distinguishes F. muirorum from F. appendiculata and F. pallida which have weak apical, medial ridges. Conchologically, F. muirorum is readily distinguished from near neighbours, F. aureedensis and F. pallida by its smaller sized and generally monochrome dark red-brown shell. In contrast to its surrounding neighbours, the shell colour and banding pattern are relatively conservative. The main range of F. muirorum appears to be centred on the vine thickets of the Pine Mountains, northeast of Marlborough, MEQ. Populations of the species also occur in scattered vine thicket patches of the Boomer and Broadsound Ranges, MEQ.

Common name. Marlborough dark snail

MEQ species (south to north)

Figuladra incei (Pfeiffer, 1846)

Helix incei Pfeiffer, 1846 (February): 126; Stanisic and Stanisic, 2020: 136–142, figs 1–6.

Helix incei. – Philippi, 1846 (February): 83, pl. 7, fig. 3.

Helix incei var. depressor. – Schmeltz, 1869: 73, (nomen nudum).

Varohadra incei. – Iredale, 1937: 33.

Sphaerospira incei incei. – Smith, 1992: 156.

Figuladra incei incei. – Stanisic et al., 2010: 474, sp. 753, lower figure in text.

Taxonomic note. Figuladra incei was considered a subspecies within Sphaerospira Mörch, 1867 by Smith (1992) along with S. incei lessoni and S. incei curtisiana. Stanisic et al. (2010) followed Smith but reassigned the three subspecies to Figuladra. Figuladra incei is herein raised to full specific status on the basis of distinctive reproductive anatomy and a strongly divergent molecular profile.

Lectotype. Percy Is, MEQ, coll. J.M.R. Ince, Commander RN, (Cuming collection), Feb. 1843, (NHMUK20200001). Height of shell 36.66 mm, diameter 32.0 mm, H/D 0.873, whorls 6.625.

Material examined.

Type material. See above. Other material. (All MEQ) Alligator Bay, 24 km S Stanage Bay, 22° 8.33′ S, 150° 3.75′ E: (QMMO8911, 3 SC/1 RC, QMMO15025, 35 RC, QMMO6973, 47 RC). Cullen Id off Sarina, MEQ, 21° 25′ S, 149° 30′ E: (QMMO17052, 9 RC, QMMO19408, 1 SC/1 RC). Hexham Id, northwestern corner, 22° 01.02′ S, 150° 21.72′ E: (QMMO76956, 3 SC/14 RC). Hunter Id, c. 13 nautical miles N Stanage Bay, 21° 58′ S, 150° 8′ E: (QMMO11479, 1 SC/1 RC). Middle Percy Id, 21° 39′ S, 150° 16′ E: (AMS C.112684, 20 RC). Middle Percy Id, 21° 40′ S, 150° 16′ E: (QMMO39430, 2 SC), 21° 39.59′ S, 150° 15.79′ E: (QMMO87084, 6 SC/1 RC). Percy Id, 21° 42′ S, 150° 20′ E: (QMMO5863, 1 RC). Pine Islet, Percy Is, 21° 39′ S, 150° 13′ E: (QMMO19146, 1 RC, QMMO19147, 1 RC, QMMO19930, 1 RC). South Percy Id, 21° 40′ S, 150° 16′ E: (AMS C.132840, 2 RC). South Percy Id, Central Valley, 21° 45′ S, 150° 19′ E: (QMMO64867, 1 SC). South Percy Id, Northwest Bay, 21° 45′ S, 150° 18′ E: (QMMO64866, 2 RC). Stanage Bay, Schnapper Drive, 22° 06.15′ S, 150° 03.1′ E: (QMMO87122, 4 SC/32 RC). Stanage Bay, 22° 9′ S, 150° 4′ E: (QMMO4271, 2 SC. QMMO4272, 5 RC, QMMO34272, 54 SC/36 RC). Stanage Bay, c. 8 km S, Stanage Bay Rd, 22° 13′ S, 150° 3′ E: (QMMO34275, 19 SC/1 RC). See Supplementary Material, Appendix 1 for additional QM material studied see Supplementary Material, Appendix 2 for records from AM database.

Description

Shell (Figure 33A–D). Large, diameter 24.89–39.37 mm (mean 31.48 mm), height 20.30–32.00 mm (mean 25.47 mm), H/D ratio 0.714–0.962 (mean 0.809), whorls 5.750–6.625 (mean 6.000); globose with moderately elevated spire; whorls rounded to subangulate, body whorl descending rapidly in front; sutures weakly impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip expanded and moderately flared, white; umbilicus narrowly open; animal dark grey, head and dorsal area with dark tentacles, pink mantle.

Shell colour and pattern variation (Figure 33). Shell tawny yellow to light brown with prominent broad subsutural and supra-peripheral bands and numerous narrower dark brown spiral bands; prominent dark brown suffusion on the last half of the body whorl in some specimens (Percy Is, MEQ); or monochrome dark brown shells and little or no banding present (Stanage Bay, MEQ); or either brown with a yellow circum-umbilical patch or monochrome yellow (Hexham Island, MEQ). Based on 61 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 16A). As for genus; vagina less than half the length of the penis, PVR 1.9–5.2 (mean 2.4).

Penis interior (Figure 16B, C). Apically with a knob-like verge (PV) and crowded crenellated rectangular pustules (PP); centrally with one prominent longitudinal ridge (PPL) and two thinner ridges; basally, an extensive zone of short longitudinal ridgelets arranged in a chevron pattern (PPR) and numerous randomly arranged, irregularly shaped tiny pustules; thin longitudinal thickenings entering the atrium. Based on 14 dissected specimens (Table S2).

Distribution and habitat. Percy Islands, Cannibal Island group; Stanage Bay to Alligator Creek on the Torilla Peninsula, MEQ; in coastal semi-evergreen vine thicket, living under and inside logs and under rocks.

Key localities. Middle Percy Id, South Percy Id, Stanage Bay.

Remarks

Results of the molecular analysis place Figuladra incei firmly within the MEQ Figuladra clade (p-distance: F. barneyae = 4.7%, F. finlaysoni sp. nov. = 6.3%, F. challisi = 4.6%). F. incei is readily distinguished from F. barneyae, F. finlaysoni sp. nov. and F. challisi by the lack of large, spade-like pilasters in the central chamber of the penis. An additional distinguishing feature is the extensive zone of basal ridgelets and pustules that occupy almost two-thirds of the penial chamber. The lack of a raised apical pilaster also relates F. incei to the MEQ clade. Shells of F. incei are variable in colour and banding pattern. Percy Island populations comprise large shells that include both banded and dark brown forms whereas the Stanage Bay population comprises smaller shells having only dark brown forms. The Cullen Island population east of Sarina, MEQ is an unusual range extension for the species that needs further investigation.

Preferred Common name. Percy Islands banded snail

Figuladra barneyae Stanisic, 2010

Figuladra barneyae Stanisic, 2010 (in Stanisic et al. 2010): 476, sp. 757, Figure in text.

Holotype. Connors Ra, S end, 15 km W St Lawrence turnoff on Bruce Hwy, MEQ, 22° 22′ S, 149° 19′ E, coll. J. Stanisic, D. Potter, 17 Jun. 1987, mvf, under logs and rocks, (QMMO20061). Height of shell 33.15 mm, diameter 38.09 mm, H/D 0.870, whorls 6.75.

Paratypes (all MEQ). Same data as holotype (QMMO78819, 12 SC/35 RC). St Lawrence, c. 16.3 km W at Stony Ck, St Lawrence – Barmount Rd, 22° 22′ S, 149° 21′ E, coll. J. Stanisic, D. Potter, 30 Apr. 1990, riparian forest, under logs and rocks, (QMMO34418, 4 RC). Connors Ra, S end, 15 km W St Lawrence turnoff on Bruce Hwy, 22° 22′ S, 149° 19′ E, coll. J. Stanisic, D. Potter, 30 Apr. 1990, mvf, under rocks, (QMMO34422, 10 SC/9 RC). Connors Ra, S end, 15 km W St Lawrence turnoff on Bruce Hwy, 22° 22′ S, 149° 19′ E, mvf, under rocks, coll. J. Stanisic, D. Potter, G. Ingram, C. Eddie, 24 Jul. 1994, (QMMO54325, 14 SC/36 RC).

Material examined.

Type material. See above. Other material (all MEQ). Beautrel Ck, Old Sarina – Marlborough Rd, SE Sarina, 22° 3′ S, 148° 59′ E: (QMMO36237, 16 SC/7 RC). Blue Mts, at Stony Ck, 21° 36.83′ S, 148° 58.92′ E, (QMMO62891, 3 RC). Clairview, 22° 6′ S, 149° 32′ E: (QMMO34532, 3 SC/6 RC, QMMO14780, 2 RC), 22° 7′ S, 149° 32′ E: (QMMO30495, 1 RC, QMMO87437, 1 SC/1 RC, QMMO19970, 1 RC). Clairview Stn, SW, Clairview Ck, 22° 12′ S, 149° 25′ E: (QMMO34522, 4 SC/1 RC). Collaroy Stn ESE of homestead c. 18.5 km SSW of Carmila, 22° 04′ 36.7′ S, 149° 23.43′ E: (QMMO86880, 4 RC). Connors Hump on St Lawrence-Croydon Rd, 235m, 22° 20.98′ S, 149° 20.58′ E: (QMMO87127, 1 SC/1 RC). Connors Ra, 14 km N St Lawrence, Croydon Rd, Collaroy Rd, 22° 16.5′ S, 149° 17.5′ E: (QMMO45985, 1 SC/7 RC). Connors Ra, c. 8.1 km SE Killarney Stn, Rd – Burwood Stn, 22° 24.6′ S, 149° 18.4′ E: (QMMO46262, 18 SC/10 RC). Flock Pigeon Id, E Clairview, 22° 8′ S, 149° 35′ E: (QMMO49807, 7 RC). Girnwood Stn, Connors Ra, S Sarina, 180m, 21° 55.03′ S, 149° 9.25′ E: (QMMO76990, 13 SC/13 RC), 21° 55.73′ S, 149° 10.3′ E: (QMMO76977, 11 RC). Greenhill, E Ilbilbie, SE Sarina, 21° 41′ S, 149° 27′ E: (QMMO34539, 19 SC/RC) Greenhill, c. 1 km NW, Greenhill access Rd, SE Sarina, 21° 41′ S, 149° 26′ E: (QMMO34541, 4 RC, QMMO87440, 1 SC). St Lawrence, Hansens Rd, 184m, 21° 38.06′ S, 149° 27.13′ E: (QMMO87435, 1 SC). St Lawrence, c. 5.9 km E at Hansen′ S Rd, 15m, 22° 20.88′ S, 149° 34.53′ E: (QMMO87125, 1 RC). St Lawrence, c. 5 km E, 22° 20′ S, 149° 34.17′ E: (QMMO54426, 18 SC/4 RC). St Lawrence, c. 20 km W Connors Barmount Rd Ra Summit, 22° 21′ S, 149° 20.75′ E: (QMMO70436, 14 SC/10 RC). See Supplementary Material, Appendix 1 for additional QM material studied. See Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 34A–D). Large, diameter 26.51–39.49 mm (mean 34.77 mm), height 20.42–33.44 mm (mean 27.56 mm), H/D ratio 0.74–0.86 (mean 0.82), whorls 5.86–6.750 (mean 6.38), globose with subangulate whorls and elevated domed spire or globose with rounded whorls and low domed spire; whorls subangulate to rounded; body whorl descending rapidly in front; sutures weakly impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip white; umbilicus narrowly open; animal grey, head and dorsal area with dark tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 34). Variable. Shell tawny yellow with prominent broad subsutural and supra-peripheral bands and numerous narrower dark brown spiral bands continuous on base (Connors Hump, MEQ); or shell with dark brown suffusion and indistinct supra-peripheral chestnut band on body whorl, spire pale with brown banding visible (most populations); or dark brown with yellow circum-umbilical patch (Figure 34C). Based on 91 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 17A). As for genus; vagina less than half the length of the penis, PVR 2.2–2.9 (mean 2.3).

Penis interior (Figure 17B, C). Apically with a knob-like verge (PV) and crowded elongate, crenellated rectangular pustules (PP) arranged in a V-shaped pattern with a pronounced medial row of tongue-like pustules (PPT) terminating centrally in a longitudinal row of large, bold, spade-like pustules (PPS) alongside a large longitudinal ridge (PPL); basally with longitudinal ridgelets arranged in a chevron pattern (PPR) and scattered small pustules; thin longitudinal thickenings entering the atrium. Based on 8 dissected specimens (Table S2).

Distribution and habitat. Connors Range (Connors Hump in the south to the Pine Mountains in the north), east to coastal areas from St Lawrence to Ilbilbie, MEQ; microphyll vine forest on rocky outcrops (upland areas) and coastal semi-evergreen vine thicket; living under rocks and logs.

Key localities. Connors Hump, Pine Mountains area in the northern Connors Range, Clairview, Greenhill, St Lawrence, MEQ.

Remarks

Molecular analysis associates Figuladra barneyae with the northern most MEQ clade species containing congeners F. finlaysoni sp. nov. (p-distance 5.4%), F. incei (p-distance 4.7%) and F. challisi (p-distance 4.6%). Anatomically, F. barneyae is characterised by a relatively long penis and a penial chamber with apical pustules that feature a medial row of pronounced tongue-like pustules that terminate in a row of bold and solid, spade-like pustules. This differs from F. finlaysoni sp. nov. which has much smaller pustules and several large, central, longitudinal thickenings as compared to F. barneyae which has only one. F. barneyae from the microphyll vine forest of Connors Hump is readily distinguished from F. finlaysoni sp. nov. by the larger shell with multiple brown bands, and elevated spire. However, specimens of F. barneyae from other localities in coastal semi-evergreen vine thicket (Greenhill and Clairview, MEQ), are smaller. F. barneyae has a subangulate shell compared to F. finlaysoni sp. nov. which has a rounded shell. The distributions of F. barneyae and F. finlaysoni sp. nov. appear to be allopatric but there is a possible contact zone in the northern Connors Range near Funnel Creek, MEQ. Study of reproductive anatomy will be essential for correct identification in this area.

Common name. Connors Range banded snail

Figuladra finlaysoni L. Stanisic, sp. nov.

Holotype. Haliday Bay, Seaforth, MEQ, 20° 53.6′ S, 148° 59.28′ E, softwood scrub adjacent to the beach, under bark of trees, coll. T. Carless, 25 Jun. 1982. (QMMO87487). Height of shell 26.05 mm, diameter 33.70 mm, H/D 0.773, whorls 6.25.

Paratypes. Same data as holotype, (QMMO87488, 1 RC).

Material examined.

Type material. See above. Other material (all MEQ). Carpet Snake Id, N end of St Helens Beach, 20° 49′ S, 148° 50′ E: (QMMO24371, 4 SC/RC). Seaforth, Finlaysons Point Rd, c. 0.75 km from Seaforth port, 20° 52.75′ S, 148° 57.27′ E, 20 m: (QMMO87456, 1 SC/11 RC), 20° 52.82′ S, 148° 57.27′ E, 50 m: (QMMO87459, 20 RC). Balls Bay, 20° 54.18′ S, 148° 59.75′ E, 20 m: (QMMO87463, 5 RC), 20° 53.35′ S, 148° 50.85′ E, 65 m: (QMMO87478, 1 RC). Seaforth, NW of Mackay, 20° 53.5′ S, 148° 56.5′ E: (QMMO35782, 6 SC/33 RC), 20° 54.17′ S, 148° 58.92′ E: (QMMO55753, 3 SC), 20° 54′ S, 148° 58′ E: (QMMO8909, 2 RC). Seaforth, Finlayson Pt, 20° 52′ S, 148° 57′ E: (QMMO12862, 2 RC). Haliday Bay, 20° 54′ S, 148° 59′ E: (QMMO12864, 11 SC), 20° 53.6′ S, 148° 59.29′ E, 27 m: (QMMO87461, 1 RC). Cape Hillsborough, via Seaforth, NW of Mackay, 20° 54′ S, 149° 3′ E: (QMMO35788, 5 RC). Cape Hillsborough Rd, c. 7 km W Cape Hillsborough, 20° 55′ S, 148° 57′ E: (QMMO11688, 2 SC/3 RC). Mt Ossa, c. 7 km E: Mt Ossa – Seaforth Rd, NW of Mackay, 20° 56′ S, 148° 53′ E: (QMMO35658, 1 RC), 20° 57.28′ S, 148° 49.43′ E: (QMMO66973, 2 SC). Cherry Tree Ck, 1.2 km off Pinnacle Station Road, 21° 12′ S, 148° 42′ E: (QMMO85407, 1 SC/1 RC). Endeavour Ck, upper reaches, Clarke Ra W of Mackay, 21° 15.5′ S, 148° 37.5′ E: (QMMO35794, 3 SC/6 RC). Eton, c. 2 km E on Peak Downs Highway, 21° 17.28′ S, 149° 17.54′ E, 65 m: (QMMO87449, 3 RC). Hay Point, end of Hay Point Rd, 21° 17.28′ S, 149° 17.54′ E, 68 m: (QMMO87451, 2 RC). Hay Pt, S Mackay, 21° 17′ S, 149° 17′ E: (QMMO34865, 2 SC/5 RC). Cut Ck, southwest of Eton on road to Nebo, 21° 18′ S, 148° 56′ E: (QMMO60246, 2 SC Eimeo), 8 m NE Mackay, 21° 2′ S, 149° 10′ E: (QMMO4942, 3 RC). Bailey Islet, off Mackay, 21° 2′ S, 149° 33′ E: (QMMO6773, 1 RC). Pine Valley, Mia Mia, Hazelbrook Stn, 21° 20′ S, 148° 52′ E: (QMMO31636, 4 RC). Mt Chelona, 1.7 km N on Main Hwy, S of Mackay, 21° 23′ S, 149° 13′ E: (QMMO20163, 6 SC/1 RC). Kuttabul, 0.2 km W Kuttabul T′off, Farleigh – Seaforth Rd, 21° 3′ S, 148° 59′ E: (QMMO19917, 1 SC/2 RC). Bells Gap, SW of Sarina, 1 km S on W Funnel Ck banks, 21° 30′ S, 149° 8′ E: (QMMO20227, 1 SC/1 RC). Sarina, SW Plum Tree Ck, upper reaches, 21° 32′ S, 148° 12′ E: (QMMO34857, 3 RC). Hatfields Gap, Koumala-Bolingbroke Rd, 21° 36′ S, 149° 12′ E, 75 m: (QMMO87442, 1 SC, QMMO19983, 2 RC). Mackay, Slade Pt, Ibis St, 21° 4′ S, 149° 13′ E: (QMMO14795, 25 RC). Sarina, ck c. 40 km S, Sarina – Marlborough Rd, 21° 40′ S, 149° E: (QMMO11862, 7 SC/19 RC). Nebo at Nebo Ck,21° 41.6′ S, 148° 41.37′ E, 213 m: (QMMO87097, 9 SC). Dipperu NP, at Pine Mt, c. 20 m SE Nebo, 21° 44.75′ S, 148° 50.58′ E: (QMMO54312, 18 SC/91 RC), 21° 45′ S, 148° 50.38′ E: (QMMO59241, 1 SC/11 RC), 21° 46.5′ S, 148° 50.5′ E: (QMMO36233, 16 SC/26 RC). Strathfield, Oxford-Sarina Rd, c. 8 km E Denison Ck X’ing, 21° 46.3′ S, 148° 50.7′ E, 227 m: (QMMO87133, 4 SC/18 RC). Mackay, 21° 9′ S, 149° 11′ E: (QMMO4160, 2 RC, QMMO4178, 3 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 35A–D). Large, diameter 24.55–35.17 mm (mean 30.93 mm), height 16.92–27.37 mm (mean 23.27 mm), H/D ratio 0.68–0.85 (mean 0.77), whorls 5.64–6.64 (mean 6.00); globose with a low domed spire and rounded whorls; sutures weakly impressed; body whorl descending rapidly in front; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip white; umbilicus narrowly open; animal grey, head and dorsal area with dark tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 35). Shell tawny yellow to light brown with prominent broad subsutural and supra-peripheral bands, numerous narrower dark brown spiral bands continuous on base and a brown suffusion on last half of body whorl (most populations). Shells from Finlayson Point, SEQ lack brown suffusion (Figure 35D). Based on 222 measured adult specimens (Supplementary Table S1).

Genital anatomy (Figure 18A). As for genus; vagina less than half the length of the penis, PVR 1.7–2.9 (mean 2.3).

Penis interior (Figure 18B, C). Apically with a knob-like verge (not visible in dissection) and crowded rectangular pustules (PP) with a medial row of prominent tongue-like pustules terminating centrally in a row of fleshy spade-like pustules (PPS) centrally, a large longitudinal ridge (PPL), and three to four thinner ridges; basally with chevron-oriented, longitudinal ridgelets (PPR) and scattered small pustules; longitudinal thickenings entering the atrium. Based on 15 dissected specimens (Table S2).

Distribution and habitat. From Pine Mt via Nebo in the west, east to the northern part of the Connors Range west of Funnel Creek (Blue Mountains); coastally from south of Sarina to Mackay and north to Seaforth and St Helen′s Beach, MEQ; in coastal semi-evergreen vine thicket and microphyll vine forest; living under timber, in hollow trees, palm fronds and under rocks and the bark of standing trees.

Key localities. Pine Mt via Nebo, Blue Mountains, Eton, Slade Point, Finlayson Point, St Helens Beach.

Remarks

Figuladra finlaysoni sp. nov. is distinguished from the parapatric F. barneyae (p-distance 5.3%). While the molecular phylogeny shows all the populations of F. finlaysoni sp. nov. sampled as distinct from those of F. barneyae, F. finlaysoni sp. nov. has a shorter penis (mean length 1.7 mm) compared with 2.3 mm in F. barneyae. Internally the penial chamber of F. finlaysoni sp. nov. lacks the distinct row of bold and solid spade-like pustules seen in F. barneyae, instead having fleshy spade-like pustules. Centrally, the three to four accessory ridges distinguish this species from F. barneyae. Shells of F. finlaysoni sp. nov. from some localities can be easily confused with those of F. barneyae and knowledge of precise locality and reproductive anatomy will be crucial to correct identification in some cases. F. finlaysoni sp. nov. is the most northerly mainland species of Figuladra with a relatively extensive distribution including microphyll vine forests of the uplands of the northern Connors Range and coastal vine thickets from Koumala to St Helen′s Beach, MEQ.

Etymology. Named for Hector Mackenzie Finlayson, an early settler of the Seaforth area.

Preferred common name. Seaforth banded snail.

Figuladra challisi (Cox, 1873)

Helix (Camaena) challisi Cox, 1873: 565, pl. 48, fig. 3; Stanisic and Stanisic, 2020: 151–154, figs 17–19.

Varohadra challisi.— Iredale, 1937: 33.

Sphaerospira challisi.— Smith, 1992: 154.

Figuladra challisi.— Stanisic et al., 2010: 476, sp. 758, figure in text.

Lectotype. Keswick Id, MEQ, W.H. Hargraves collection, pre-1877 (AMS C.583621). Height of shell 24.80 mm, diameter 31.14 mm, H/D 0.796, whorls 6.5.

Paralectotypes. Same data as lectotype (AMS C.17597, 7 RC).

Material examined.

Type material. See above. Other material (all MEQ). Keswick Id, Cumberland Group, NE of Mackay, 20° 53.37′ S, 149° 45.17′ E: (QMMO70382, 5 SC/20 RC), 20° 53.37′ S, 149° 25.21′ E: (QMMO86827, 4 RC), 20° 55.33′ S, 149° 25.4′ E: (QMMO86828, 4 SC/1 RC, QMMO87074, 4 RC), 20° 55′ S, 149° 25′ E: AMS C.133804, 2 RC). Scawfell Id northern end, 20° 44.59′ S, 149° 45.17′ E: (QMMO70385, 7 RC). St Bees Id, Cumberland Group, 20° 54.89′ S, 149° 25.75′ E: (QMMO86829, 10 SC/2 RC, QMMO87072, 14 RC). See Supplementary Material, Appendix 1 for additional QM material studied; see Supplementary Material, Appendix 2 for AM records from AM database.

Description

Shell (Figure 36A–C). Large, diameter 30.43–33.62 mm (mean 31.63 mm), height 22.36–23.52 mm (mean 22.64 mm), H/D ratio 0.69–0.75 (mean 0.72), whorls 5.88–6.13 (mean 6.00); globose with low domed spire and rounded whorls; body whorl descending rapidly in front; sutures weakly impressed; protoconch with weak crowded, curved radial ridges, teleoconch smooth, microsculpture of fine periostracal threads; lip reflected, white; umbilicus partly closed. animal pinkish-grey, head and dorsal area with pinkish-grey to grey tentacles and pink foot, pink mantle.

Shell colour and pattern variation (Figure 36). Consistent. Tawny yellow to off-white with prominent broad subsutural and peripheral bands and numerous narrower dark brown spiral bands continuous on base; small brown circum-umbilical patch. Based on 32 measured adult specimens (Table S2).

Genital anatomy (Figure 19A). As for genus; vagina less than half the length of the penis, PVR 1.9–2.7 (mean 2.2).

Penis interior (Figure 19B, C). Apically with a knob-like verge (not visible in dissection); crowded crenellated rectangular pustules (PP); centrally with a row of tongue-like pustules (PPT) becoming fleshy spade-like (PPS), two to three prominent longitudinal ridges (PPL); basally with prominent longitudinal ridgelets (PPR) arranged in a chevron pattern and numerous scattered, randomly arranged, irregularly shaped pustules (PPP); thin longitudinal ridges entering the atrium. Based on 5 dissected specimens (Table S2).

Distribution and habitat. Southern Cumberland Islands including Keswick Id, St Bees Id and the northern end of Scawfell Id, MEQ; coastal vine scrub, remnant semi-evergreen vine thicket with eucalypt emergents, living under rocks and logs and in litter.

Remarks

The molecular analyses show the populations of Figuladra challisi from the South Cumberland Island group as distinct from the F. finlaysoni sp. nov. (p-distance 5.3%). The penial chamber of F. challisi has three thick longitudinal thickenings compared to the multiple thickenings in F. finlaysoni sp. nov. The basal chamber of F. challisi features thin longitudinal ridges different from the small, scattered, pilasters of F. finlaysoni sp. nov. Shells of F. challisi are consistently banded in contrast to mainland F. finlaysoni sp. nov. which has similar looking shells at Finlayson Point and shells with brown suffusions at other localities. F. challisi is not liable to be confused with any other Figuladra species in the current study due to its exclusively insular distribution on the South Cumberland Islands, MEQ. On Scawfell Island, MEQ, F. challisi has been found in microphyll vine forest at the northern end of the island. At the southern end of the island, a putative new camaenid species, suspected of belonging to Figuladra, has been found but is only known from a large series of litter collected and faded shells.

Common Name. Squat Cumberlands banded snail

Discussion

Systematics

This revision of Figuladra from south-eastern and mid-eastern Queensland including adjacent islands, based on newly collected specimens as well as historical museum material, revealed a diverse land snail radiation. In the most recent review of the genus, Stanisic et al. (2010) recognised 11 species based mainly on shell morphology, including one with three identified sub-species and a further two which have subsequently been shown to belong to other genera, viz. Euryladra mattea (Iredale, 1933) and Brigaladra volgiola (Iredale, 1933). The large variation in shell colour and banding patterns within species and the considerable overlap in these features between species has made past identification from the shell alone difficult, if not impossible. Accordingly, our investigation implemented an integrated taxonomic approach combining morpho-anatomy and mitochondrial phylogenetics in an effort to more clearly discriminate species within the clade. An additional consideration was geography (including landscape) and the identification of historical barriers to dispersal as well as refugia for isolation. The relationship between geographic and genetic distance was significant, showing a general trend of increasing genetic divergence as geographic distance increased, as might be anticipated given the ecology and environment of the genus. The outcome presented a broad vicariant pattern of north–south replacement of 15 identified species of Figuladra, four of which are newly described (Table 6, Figure 38).

Figure 38. A, Occurrence records of Figuladra species described in this study in eastern Queensland from records including all sequenced individuals in the malacological collection of the Queensland Museum. B, Three major Figuladra clades (see Figure 3). The incongruences in the tree (F. reducta from Kroombit Tops/Ubobo and the Bulburin SF specimen) are shown in red. The final decision for the naming of these incongruencies was determined by dissection. See discussion of Kroombit Tops locality in the ′Remarks′ section for F. reducta. The Bulburin specimen cannot be traced to a museum specimen so that any anomalies in its locality data cannot be confirmed.

Table 6. Summary of all species in Figuladra.

Names used in Stanisic et al. 2010 and putative species in museum collections	Current taxonomy
Figuladra incei curtisiana & Figuladra incei lessoni Camaenid SQ 19	Figuladra lessoni
Figuladra incei incei	Figuladra incei
Camaenid BL 50
Camaenid BL 51
Figuladra bayensis	Figuladra bayensis
Figuladra narelleae	Figuladra narelleae
Figuladra reducta	Figuladra reducta
Figuladra appendiculata Helix praetermissi	Figuladra appendiculata
Figuladra muirorum	Figuladra muirorum
Figuladra pallida	Figuladra pallida
Figuladra challisi Camaenid BL 48	Figuladra challisi
Figuladra barneyae Camaenid BL 46	Figuladra barneyae
Camaenid BL 47	Figuladra finlaysoni sp. nov.
Camaenid SQ 15	Figuladra robertirwini sp. nov.
Camaenid SQ 16	Figuladra bromileyorum sp. nov.
Camaenid SQ 17	Figuladra vidulus sp. nov.
Helix aureedensis	Figuladra aureedensis comb. nov.
Camaenid BL 45	Camaenid BL 45 (Wadallah Creek, MEQ)

In this study, operational criteria for delimiting species were defined and data gained from a variety of sources. Firstly, the guide tree and the genetic distance data were used to delimit groupings of closely related individuals. However, these data did not fully resolve ’species’ in all cases but gave mitochondrial groupings on which to base a broad hypothesis. This hypothesis was then tested by exploring patterns within the morphological data. When genital anatomy data were added to these data, the picture became much clearer, and the hypothesis honed. Observable differences were found in genital anatomy between the taxonomic units identified in the initial hypothesis using the guide tree and were consistent with those groupings (see Table 3). The main features that contributed to this discrimination were identified in the apical and central chambers of the penis. A refined hypothesis was overlaid by the shell morphology which by itself created some confusion due to shell plasticity. By using approaches to detect divergence at both the molecular and morphological levels, congruence between the lines of evidence demonstrated that it was possible to develop a robust Figuladra taxonomy, thus helping to unravel this taxonomic conundrum. Comparing evidence from multiple independent sources improved confidence in species boundaries and suggested that Figuladra species are reproductively isolated and independently evolving units.

Both molecular and morphological data indicate that the camaenid land snails of Figuladra constitute three major lineages in SEQ, SEQ uplands and MEQ with each containing a number of species. Internally, the general form of the reproductive system (genitalia) was deemed uninformative in species delimitation with all species showing a similar structure. The detailed penial chamber architecture has been shown to be species-specific among camaenid species, and showed a level of inter-specific variation in Figuladra spp. consistent with species-level differences seen in some other camaenid groups studied (e.g., Solem 1992b; Stanisic 2018; Stanisic et al. 2022). More significantly, the observed variations align with the molecular structuring shown in the phylogram of Figure 3. The apical third of the chamber in the majority of species in the SEQ clade had crowded pustules and either an apical pilaster (four species) or an apical medial ridge (five species). Two species (SEQ Upland clade), identified as a separate branch on the mitochondrial 16S phylogram, had multiple rows of bold pustules occupying most of the chamber in what is a major departure from the general pattern of apical pustules, central ridges with pustules and basal pustules. Within the MEQ clade, a binding characteristic among three of the species was the presence of bold spade-like pustules in the central part of the chamber.

The radulae of Figuladra were not a focal point of the current study. However, the radulae of two species (F. lessoni, F. bayensis) were investigated with SEM and showed no significant differences in tooth structure from the radulae of Euryladra and, Brigaladra (Stanisic et al. 2022).

Similarities were seen in external characteristics. All Figuladra animals have a pink mantle distinguishing Figuladra from other large hadroid genera in south-eastern Queensland (Stanisic et al. 2022). Shell shape discriminated species in SEQ with the upland species possessing flatter shells (depressedly globose with a low domed spire) from those with a globose shell and an elevated spire (Table 5). These species (F. bayensis, F. reducta, F. narelleae) also had rounded rather than subangulate whorls. In the MEQ clade, species with globose shells and a low spire (F. finlaysoni sp. nov., F. challisi) had a rounded shell contour suggesting that the low spire and a rounded whorl is a linked change. However, in general, shell contour was a less useful character in differentiating species with most having subangulate whorls to some extent. Shell variation as the primary basis for classification needs to be treated cautiously, as the external shell is considered particularly susceptible to environmental selection processes (Schander and Sundberg 2001 and references therein; Walker 2018). Conchologically, shell colour and banding pattern were the least informative in identifying species with a similar range of colour and patterns present in most of the Figuladra clade species (Table 5).

While mate recognition appears to play little part in the development of dark or banded shells in Figuladra, camouflage may. The dominant colours in Figuladra shells, though dramatically varied, encompass a subtle palette of yellows and browns arranged in either monochrome or banded forms. These are not only the colours of dead leaves on the ground but also of the Australian forests in which they live. They are all long wave-length colours towards the infra-red end of the spectrum and are less visible to birds that are known to predate snails (Savazzi and Sasaki 2013). Nevertheless pitta birds (Pitta versicolor 25Swainson, 1825) found in eastern coastal Australia are known predators, using ‘anvils’ to break the shells of snails which are one of its primary foods (Rose 1999; Australia Zoo 2023). Anvils can be a rock stuck in the ground, a tree stump, or even a broken bottle. Anvils have been sighted on many occasions in the forests of eastern Queensland (pers. obs).

Geographic distribution and habitat

(Figures 39–41)

Figure 39. Habitat and live snails. A, Coastal vine thicket, Boyne Id, SEQ, habitat of Figuladra lessoni; B, Vine thicket on sand, Eurimbula NP, SEQ, habitat of F. robertirwini sp. nov.; C, Vine thicket, The Hummock, Bundaberg, SEQ, habitat of F. vidulus sp. nov.; D, Vine thicket, Mt Perry, SEQ, habitat of F. reducta; E, Microphyll vine forest, Mt Mudlo, SEQ, habitat of F. bayensis; F, Coastal vine thicket, Urangan, SEQ, habitat of Figuladra bromileyorum sp. nov.

Figure 40. Habitat and live snails. A, Microphyll vine forest Byfield, MEQ, habitat of Figuladra pallida; B, Remnant vine thicket, Glenprairie, MEQ, habitat of F. muirorum; C, Vine thicket on limestone, Johannsen′s Cave, MEQ, habitat of F. aureedensis; D, Coastal vine thicket, Mulambin Beach, SEQ, habitat of F. appendiculata; E, Vine thicket on rocky scree, Bouldercombe Gorge, SEQ, habitat of F. narelleae.

Figure 41. Habitat and live snails. A, Microphyll vine forest, Connors Hump, MEQ, habitat of Figuladra barneyae; B, Vine thicket, St Bees Id, MEQ, habitat of F. challisi; C, Coastal vine thicket, Stanage Bay, MEQ, habitat of F. incei; D, Coastal vine thicket, Finlaysons Point, Seaforth, MEQ, habitat of Figuladra finlaysoni sp. nov.

The mainly lowland and near-coastal species in the southern-most species of the clade (Figuladra reducta, F. bromileyorum sp. nov., F. vidulus sp. nov., F. robertirwini sp. nov.) have contiguous distributions and the latter three form a close-knit group of phenotypically similar species in the semi-evergreen vine thickets and vine forests of the region’s lowlands south of the Boyne River. F. bromileyorum sp. nov. is also present on K’gari (Fraser Id) as well as the neighbouring mainland suggesting a recent connection.

F. reducta occupies araucarian microphyll vine forest in the Dawes and Burnett Ranges and some lowland locations (Goodnight Scrub) to the west of the Burnett River. Among the material studied, molecular data show two distinct reducta lineages, a southern one (Mt Perry, Goodnight Scrub) and a northern one (Kroombit Tops in the Dawes Range) with close association to F. lessoni. In two cases within the SEQ clade, uplands appear to have been significant refugia for the altitudinal isolation of F. bayensis and F. narelleae from lowland and near coastal groups. The distribution of F. bayensis is centred on araucarian microphyll vine forests of Mt Biggenden, Mt Walsh, Mt Mudlo and Mt Woowoonga west of the distribution of F. bromileyorum sp. nov. F. narelleae occupies araucarian microphyll vine forests in the Mt Morgan, Boyne, Dee and Ulam ranges and some adjacent lowland areas (Dan Dan Scrub) south-west of Rockhampton and west of F. lessoni.

To the north of the Boyne River and south of the Fitzroy River F. lessoni occupies vine thickets of the region’s coastal lowlands on either side of the intervening Calliope River as well as the off-lying islands (Curtis, Farmers, Wild Cattle Id). Populations south of the Calliope River have mainly dark brown shells whereas those to the north have lighter coloured and variously banded shells.

To the north of the Fitzroy River, F. appendiculata occupies vine thickets and vine forests of the Berserker Range and the coastal lowlands north to Yeppoon including those on the nearby Keppel Island Group. F. pallida, a species with a similar penial chamber to F. appendiculata (Figure 14) but with a shell that differs significantly from that of F. appendiculata in colour, banding pattern and shape, occupies the Parnassus Range and the environs of the Byfield NP. Currently included here are populations living in dry coastal vine thickets of the Peninsula Range and off-lying islands to the north of Byfield NP. These populations are known only from dead shells and their inclusion is tentative pending the availability of live material for study. An undescribed species, camaenid BL 45 (Figuladra ‘Wadallah Creek’) separates these populations from Byfield F. pallida suggesting that the Peninsula Range populations may also represent a distinct species.

To the west of the former two species and endemic to the semi-evergreen vine thickets of the limestones of Mt Etna NP is F. aureedensis. These limestones are inhabited by other endemic land snails (Stanisic et al. 2010) which highlights the insular nature of such habitats within the woodland landscape.

F. muirorum inhabits semi-evergreen vine thickets north of the Fitzroy River and extends to the Boomer Range north-west of Rockhampton to the Pine Mountains north-west of Marlborough. The geographical separation of this species from the main mass of the SEQ clade north of the Fitzroy R (∼ 200 km) indicates that the relationship is an old one, borne out by the molecular analyses which identify this species as a clearly divergent lineage within the SEQ clade.

Species of the MEQ clade are distributed north of the St Lawrence Gap with F. incei being the southern-most in the Broadsound area, including some off-lying islands, MEQ. F. incei was originally described from the Percy Islands that are 130 km off the coast. As a result of this study, the species is now known to occur on a number of closer in-shore islands (Hunter Id, Marble Id, Avoid Id) and at Stanage Bay on the mainland. The Percy Island shells are relatively large compared with those on the other islands and on the mainland and include both dark and lightly banded forms.

Whereas upland areas in SEQ harbour species that are quite different in their reproductive anatomy from the adjacent lowland species, this is not the case in MEQ. Two species with upland populations also inhabit the lowlands. F. barneyae has a large, banded shell found in the microphyll vine forests of the upland areas of the Connors Range north of the St Lawrence Gap but has a smaller, darker form in the semi-evergreen vine thickets of adjacent coastal areas. Similarly, F. finlaysoni sp. nov. occurs in araucarian microphyll vine forests of Pine Mountain via Nebo, Mt Ossa and the uplands of the northern Connors Range at Mia Mia SF, but also in the coastal vine thickets of the Proserpine to Sarina lowlands from Sarina to St Helens Beach south of the O’Connell River, MEQ.

F. challisi is an island species (Keswick Id, St Bees Id, Scawfell Id) that has no known mainland population. Shells are strongly banded with some very similar to those of F. finlaysoni sp. nov. from Finlayson Point, Seaforth (compare Figure 35D, Figure 36A). Molecular analyses show these island populations as distinct from both F. finlaysoni sp. nov. and F. barneyae.

Biogeography

The current data give an increased understanding of the biogeography of eastern Australian Camaenidae. The camaenids were considered mid-Tertiary immigrants from land masses to Australia’s north by McMichael and Iredale (1959) an opinion reinforced by Solem (1992b) and which now seems to be the prevailing view. McMichael and Iredale (1959) were the first to offer a biogeography of the Australian land snail fauna but were constrained by the crude taxonomy of the time, based solely on shell morphology. As a result, there was no attempt to relate snail evolution at the species level to fine scale landscape and vegetation changes. Based on shell and reproductive anatomy, Solem (1992a and references therein; 1992b) divided the camaenids into two subfamilies with the east coast radiation belonging to the Camaeninae. However, Solem’s interests were mainly with the camaenids in South Australia, Western Australia, and Northern Territory and he provided little detail on the diversification of the eastern fauna. The most recent comprehensive biogeographical study of eastern Australian camaenids was conducted by Hugall (2011) who suggested that these camaenids stemmed from an Oligo-Miocene Laurasian immigration and that ‘relictual endemics indicated that many ancestral lineages were in place before the major decline of the mesic forest realm’ (Hugall 2011: v).

The arrival of the camaenids on the continent in the Miocene occurred when the Australian land mass had split from Gondwanaland and xerification was beginning to exert environmental control over mesic habitats. Along the east coast, the palaeo-environments in which the camaenids must have dispersed and radiated subsequent to their appearance were to undergo drastic change. In eastern Australia, two geological events contributed to the maintenance of mesic communities in the wake of climatic phases that elsewhere resulted in the removal of widespread rainforest flora (Martin 2006). Firstly, episodic uplift of the eastern highlands began in the Palaeocene with major elevation occurring in the Miocene and Pliocene. The eastern highlands ensured orographic rainfall and enabled species to radiate altitudinally. Secondly, there was widespread igneous activity in eastern Australia along and adjacent to the eastern highlands (Wellman and MacDougall 1974). Indirectly, volcanism provided favourable soil-water conditions in a basically dry and infertile continent. The fact that most of the eastern hadroid radiation now occurs in ‘rainforest’ suggests that the persistence of moist refugia to the present day is pivotal to their survival. The main mesic refugia during dry phases were in the moist uplands, and in the lowlands along major drainage lines, areas which were not large, but wet.

Climatic changes in the late Quaternary were similar in amplitude to those experienced in the Miocene and Pliocene but occurred in more rapid succession. Galloway and Kemp (1981) concluded that these changes must have placed considerable stress on montane and coastal environments and that modern biotic communities in these situations are recent assemblages consisting of biota which have survived in isolated refugia. During the last 1.8 million years, there have been at least 17 glacial-interglacial cycles (Williams 1984). These extreme climatic changes have been responsible for the most recent environmental sifting of the rainforest biota including land snails.

The molecular-based phylogeographic study of the hadroid camaenid Sphaerospira by Hugall et al. (2003) was the first to provide an insight into the historical evolution of any land snail lineage in eastern Australia. Combining mtDNA analyses with bioclimatic modelling the study showed that the monophyletic Sphaerospira group formed a phylogenetically nested north–south series of discrete allopatric sister lineages and concluded that these lineages developed via vicariant isolation as rainforest blocks became progressively fragmented and isolated from the mid-Miocene onwards. Particularly relevant to the present study were the findings that the St Lawrence Gap was an effective biogeographic barrier separating MEQ species from those in SEQ and that the Shoalwater/Byfield and Kroombit/Bulburin areas were bioclimatic refuges for the evolution of these snails. Throughout MEQ and SEQ Sphaerospira occurs in sympatry with Figuladra and some of that study’s findings aid the understanding of the evolution of this genus.

Similarly to Sphaerospira, Figuladra exhibits a broad pattern of north–south replacement of species within a distribution ranging from the Kilkivan-Gympie-Maryborough area, SEQ to the O’Connell River, south of Proserpine, MEQ. Unlike Sphaerospira, there is no sympatry and allopatry appears to be the normal pattern of distribution for these snails. The St Lawrence Gap dry corridor (Dick 1964; Bryant & Krosch 2016) separates the SEQ clade (11 species) from the MEQ clade (4 species). This separation is supported by both genetic and anatomical data and once again highlights this corridor as a major influence on the dispersal and evolution of biota along Australia’s east coast. Not unexpectedly, this dry corridor has particular significance for low vagility, moisture-sensitive land snails as the study of Sphaerospira by Hugall et al. (2003) has shown.

Species of Figuladra inhabit dry rainforest (low microphyll vine forest, semi-evergreen vine thickets) on the lowlands and coastal areas and slightly wetter rainforest (araucarian microphyll vine forest) in upland areas of their distribution. The dry vine thicket/vine forest biome within SEQ and MEQ lowlands is an archipelago of scattered environments that would have expanded and contracted many times in response to the rapidly fluctuating climate regime of the Pleistocene. These episodic climatic shifts, from humid moist to cool dry, would have segregated lowland and coastal biotic communities across emerging barriers as the rainforest blocks became more fragmented. Figuladra populations in these situations would have become isolated on more than one occasion and also been provided with opportunities for dispersal and contact. With the overriding trend to more xeric conditions, these isolated populations would gradually develop into incipient species as separation became more environmentally pronounced. Shifts in penial chamber structure, reflecting species-species interactions tending to promote reproductive isolation, would have occurred gradually.

Within the SEQ clade south of the Fitzroy River, rivers can be identified as geographic barriers as they appear to form the distributional limits of the lowland species. The Isis, Kolan, Boyne and Fitzroy river systems have all played major roles. Species ranges abut on different sides of the identified barriers (F. bromileyorum sp. nov./F. vidulus sp. nov.: Isis River; F. vidulus sp. nov./F. robertirwini sp. nov.: Kolan River; F. robertirwini sp. nov./F. lessoni: Boyne River; F. lessoni/F. appendiculata: Fitzroy River). The low genetic diversity between species would suggest that it is likely that the rivers have formed ecotonal barriers over time and that each species has adapted to its habitat on either side of the ecotone (Bull 1991; Hamilton and Johnson 2015).

Reproductive character displacement may arise as an interaction between competition, gene flow and abiotic environments and may shape or stabilise distributional boundaries over time (Case and Taper 2000; Case et al. 2005; McQuillan and Rice 2015). This may be seen in the distribution of F. lessoni and F. robertirwini sp. nov. where no distinct barrier is evident, and a parapatric hybrid zone may be present. Species records were geographically too scattered to define the extent of the possible contact zones.

Separation of lowland populations from upland populations would have been more dramatic than the isolation of lowland populations, as would separation of upland populations from each other. In contrast to the lowland scenario, upland areas of araucarian microphyll vine forests on rocky substrates with higher annual precipitation (rain and cloud drip) would have provided relatively more stable refugial environments for Figuladra species. Species such as F. bayensis and F. narelleae probably have evolved over an equally long period of time in relative altitudinal isolation. This separation from lowland species is reflected in their radically different penial chambers.

The Kroombit Tops area was identified as a bioclimatic (ecological) refuge for the evolution of these snails by Hugall et al. (2003) and is implicated in the upland separation of F. reducta in the Dawes Range.

F. aureedensis is a disjunct species belonging to a relatively uniform fauna but evolved in an insular habitat. Most likely this species was able to survive in isolation in the limestone outcrops of Mt Etna NP during the environmental attrition of the Pleistocene. F. aureedensis is considered to be a topographic isolate that is restricted to the limestones of Mt Etna NP and is now unable to expand into adjacent fire-prone environments. Limestone restricted camaenids with non-limestone associated congeners also inhabit similar outcrops at Greenvale, MCQ, Chillagoe, NEQ and Palmerville, NEQ (Stanisic et al. 2010).

The occurrence of F. muirorum in vine thickets in the Marlborough Plains bioprovince north of the Fitzroy River, and its potential relationship to lowland species south on the Boyne River, is indicative of an ancient distribution of lowland vine thickets that have undergone contraction, fragmentation and even extinction in response to Pleistocene climate changes.

The MEQ clade represents a differing set of evolutionary scenarios to those of the SEQ clade. The overarching picture is still one of climate-induced rainforest fragmentation and isolation but without the dramatic shift in penial chamber structure seen within the SEQ clade. Upland species such as F. barneyae and F. finlaysoni sp. nov. also have lowland populations suggesting a shorter or less complete separation of their upland and lowland biomes during the Pleistocene climatic fluctuations. Both species occur in the uplands of the Connors Range and the adjacent Proserpine-Sarina lowlands. Together with F. challisi, these species share features of the penial chamber (unusual bold spade-like pustules) that indicate their derivation from an ancestral population presumably by climate-induced fragmentation of rainforest habitat. During this fragmentation, F. challisi may have become isolated on islands of the Cumberland Group, MEQ by rises in sea level during the many glacial cycles of the Pleistocene. These cycles were quite dramatic as is evidenced by the estimation of sea levels 200 m lower than present day in the last glacial maximum (15,000–20,000 ybp) (Willmott 2006, 2014). For F. challisi, separation from the mainland during these cycles was sufficient to cause speciation. In contrast F. incei, which occurs on the Percy Islands, also has mainland populations indicating that these and the island populations have recently been connected.

Conservation and threats

Terrestrial biodiversity is being lost at an alarming rate and no taxa are more affected than the invertebrates. This decline is at least partly related to habitat loss due to a number of factors. For example, Figuladra has been impacted by climate induced reduction of habitat and more recently, habitat loss through land clearing and both natural and anthropogenic fire regimes.

Habitat loss due to land clearing for agriculture and farming and more recently mining developments, has significantly affected Figuladra populations. This is likely to continue into the future particularly where urban and related human development is progressing rapidly. The total area of land cleared in Queensland was 2,446,600 hectares from 2010 to 2018 and, of that, 370,900 hectares were forests over 30 years old (Kilvert 2020). In 2018, land clearing laws were introduced, and these were expected to reduce the rate of clearing for pasture. However, Queensland is considered by many to be a deforestation hotspot (DELW 2021; Wilderness Society 2022) and the Queensland Conservation Council reports that the clearing has not reduced but doubled, placing native wildlife at serious threat (Copeman 2022). The Maryborough, Bundaberg regions in SEQ and the Mackay area in MEQ are areas where land clearing for agriculture and urban development has led to small, isolated remnant vine thickets being largely reduced to coastal strips. Where vine thickets have been severely degraded, but fallen timber or rocky outcrops remain, Figuladra snails have shown remarkable resilience (Figure 42) indicating that for the survival of land snails and other invertebrates, small tracts of native vegetation should be maintained in any land clearing regime.

Figure 42. Figuladra appendiculata surviving under rocks in severely degraded vine thicket from unnecessary clearing at the side of Roconia Road, Lakes Creek, SEQ.

Fire is a major threat to the existence of invertebrates through habitat modification and destruction. Low vagility land snails are particularly prone to the effects of fire and even cool burns adversely affect populations. Hot burns destroy land snail microhabitat (Stanisic and Ponder 2004; Ray and Bergey 2014). Dry rainforest and vine thickets have little tolerance to fire. Introduced grasses (e.g., Buffel grass, Cenchrus ciliaris) are a major threat to dry rainforests and vine thickets. These grasses are highly flammable and contribute to the destruction of many fire-sensitive rainforest plants. Fire also causes the dry rainforest to contract in size, until it is no longer a functioning ecosystem (Grimshaw 2017). This extreme sensitivity of dry rainforest and vine thickets to fire, coupled with their restricted nature, indicates they are considered to be communities of great ecological concern. Recently, the effects of anthropogenic fire regimes have been shown to encroach into vine thickets and cause serious damage and a marked delineation between vine thickets and woodland communities, affecting the transition zone between the two (Stanisic and Ponder 2004; McLauchlan et al. 2020).

There is a growing recognition that fire may have evolutionary consequences in animals and in particular, the structure of invertebrate communities (Gill et al. 1999; Thom and Seidl 2016; Hewitt et al. 2016; Pausas and Parr 2018; McLauchlan et al. 2020). Given the influence of fire on dry rainforest and open woodland combined with the aridification of the continent, an emphasis on the role of a managed fire regime for the maintenance of vegetation types and structure across the landscape is critical (Burgess et al. 2015). The survival of low vagility land snails depends on this.

Figuladra aureedensis, endemic to the limestones of Mt Etna NP, was found in extremely large numbers when the vine thickets were in a pristine state in 2019 (Figure 43A). After a bushfire in 2020 (Figure 43B), the devastation of the local snail populations, including F. aureedensis, was significant and visible (Figure 43C). However, live snails were found still thriving in rocky crevices in the limestones. Of greater concern, a follow up visit two years later showed a greatly reduced population that will take years to re-establish.

Figure 43. Johanssen’s Cave, Mt Etna NP, MEQ; A, 2019. B, 2020 after a bush fire. C, Effects of the fire on the population of Figuladra aureedensis.

Most species dealt with in this study occur outside the current conservation reserve system and would require ‘listing’ to gain further protection. The current listing of Brigalow Scrub and other threatened forest communities such as vine thicket, does protect Figuladra species in these susceptible areas.

The question remains as to which anthropogenic pressures are most likely to affect species in the future? For those species restricted to the upland areas of the Great Dividing Range, habitat destruction appears to be a minimal risk for now. For the lowland species, the risk is significantly greater. Fortunately, on the lowlands, populations of some (but not all) species still occur within National Parks and State Forests.

The overall goal of any conservation strategy should be to protect the processes, both ecological and evolutionary, that sustain genetic diversity in the ecosystem. Fundamental to this aim is the preservation of species. From an Australian conservation perspective, low vagility land snails are an important element of biodiversity and should not be overlooked in the conservation of our native forests (Parkyn and Newell 2013).

Concluding remarks and future directions

This revision of the Figuladra radiation should not be seen as an endpoint. Rather. the study opens the door for further research on the group. At the species taxonomy level, there are a number of issues to resolve that were identified by the molecular analyses. These comprise the status of the F. reducta populations in the Dawes and Burnett Ranges, SEQ; the status of the Peninsula Range, MEQ populations currently incorporated with F. pallida; the distribution and description of camaenid BL 45; and the molecular analysis of associated island populations of Figuladra currently only known from shells (Fraser Island/K’gari, Scawfell Id, Hunter Id). Based on unpublished preliminary results, the use of single nucleotide polymorphism (SNP) technology will be useful in identifying possible contact zones between such a tightly bound geographic radiation of species. Such studies will require further fieldwork but should lead to a greater understanding of camaenid evolution in eastern Australia at a time when climate change increasingly threatens the rainforest biome.

Permits

In order to allay any conservation concerns, the majority of specimens were collected in areas outside National Parks and State Forests. Where specimens were taken from within these gazetted jurisdictions they were carried out under Scientific Purposes permit number WlTK16810116.

Acknowledgements

We acknowledge Griffith University, the Queensland Museum and Australian Museum for allowing us to access facilities and collections for this study. We would like to sincerely thank Darryl Potter, collection manager, Queensland Museum for his assistance with the collection and the databae. A special thanks to Alistair Meltzer and Kathryn Radcliffe, for their assistance with the collection of some specimens from the South Cumberland Islands and Percy Island respectively for this project in times when museum fieldwork was unable to be conducted. Thank you also to Geoff Thompson from the Queensland Museum Digital Imaging Unit for providing high resolution images of the new holotypes, and to Kim Maxwell for her cartographic expertise. We thank the reviewers for their constructive comments.

Data availability

All data generated or analysed during this study are included in this article and its Supplementary Information. Sequence information generated in this study are available at GenBank, accession numbers OQ676310–OQ676369.

Disclosure statement

No potential conflict of interest was reported by the author(s).

Additional information

Funding

This research was funded by HDR candidate support funding from Griffith University to LS and CM.