The temperate Australasian genus Papawera Oskars and Malaquias, 2019 (Gastropoda: Cephalaspidea: Haminoeidae), with a redescription of P. zelandiae and P. maugeansis

Figures & data

Figure 1. Bayesian phylogeny depicting relationships and species diversity of the genus Papawera. Hypothesis based on the combined analysis of the mitochondrial COI, 16S rRNA and 12S rRNA and nuclear 28S rRNA and Histone H3 genes. The tree is a composite based on Figures 1 and 2 from Oskars & Malaquias (2019). Values above branches are BI posterior probabilities and below branches are bootstrap values derived by ML (Oskars & Malaquias, 2019). Images of Lamprohaminoea cymbalum, Mozambique (TH64), courtesy of Y. Tibiriçá; Bakawan rotundata, Singapore (496), courtesy of K. Jensen; Smaragdinella cf. sieboldi, Mozambique (TH55), courtesy of Y. Tibiriçá; Haminoea alfredensis, South Africa, NHMUK 20070315 (174), courtesy of G. Branch and C. Griffiths

Figure 2. Female reproductive system. (a) Papawera zelandiae, Hokianga, Northland, New Zealand, (AM 79176, H = 11 mm). (b) Papawera maugeansis, Curlewis Reef, Clifton Springs, Victoria, Australia (ZMBN 125459 ex NVM F194630, H = 5.7 mm). Abbreviations: agl, albumen gland. am, ampulla. amg, anterior mucous gland. gd, gameloytic duct. ggl, gametolytic gland. md, medial duct. ot, ovotestis. pmgl1, posterior mucous gland lobe 1, pmgl2, posterior mucous gland lobe 2. rm, retractor muscle. smg, external seminal groove. v, vestibule. Scale bars: a, b = 0.5 mm

Table 1. List of specimens used in the ABGD analyses including voucher and GenBank accession numbers

Species	DNA extract code	Locality	Voucher No.	COI
Papawera maugeansis (Burn 1966a)	TH9	Shoreham Beach, Victoria, Australia	NMV F 209129	MK473519
Papawera maugeansis (Burn 1966a)	TH10	Curlewis Reef, near Clifton Springs, Victoria, Australia	ZMBN 125458 (ex NVM F 112,423)	MK473517
Papawera maugeansis (Burn 1966a)	TH33	Inverloch, Town Area, Victoria Australia	ZMBN 125459 (ex NVM F 194630)	MK473518
Papawera zelandiae (Gray 1843)	456	Waitemata harbour, north end of Eastern Beach, New Zealand	MA 119920	MK473515
Papawera zelandiae (Gray 1843)	458	Waitemata harbour, north end of Eastern Beach, New Zealand	MA 119920	MK473516

Table 2. COI inter- and intra-specific uncorrected p-distances (%) calculated in MEGA 7

Specimens	456	458	TH10	TH33		Intra-specific p-dist
Papawera zelandiae 456	-
Papawera zelandiae 458	0.3	-			P. zelandiae	0.3
Papawera maugeansis TH10	13.8	13.8	-
Papawera maugeansis TH33	13.5	13.5	0.9	-
Papawera maugeansis TH9	13.6	13.6	1.1	0.2	P. maugeansis	0.2–1.1

Figure 3. Live animals of Papawera zelandiae (Gray, 1843) (a–c), Papawera maugeansis (Burn, 1966a) (d–h). (a) Blockhouse Bay, Auckland, New Zealand (D. Crisp, CC BY-NC, iNaturalist.org). (b) Ambury Farm, Mangere Bridge, Auckland, New Zealand, courtesy of J. Geux. (c) Mahurangi West, North Island, New Zealand (D. Wilson, CC BY-NC, iNaturalist.org). (d) Hamers Haven, courtesy of J. Hales. (e) Inverloch, Victoria, Australia, courtesy of J. Hales. (f) Australia, Victoria, Westernport, Shoreham, (AMS c.105854, H = 11 mm), courtesy of G. Millen. (g) Victoria, Australia, courtesy of L. Altoff. (h) Cape Paterson, Inverloch, Victoria, Australia, courtesy of L. Altoff

Figure 4. Papawera zelandiae (Gray, 1843). (a) apertural view of shell (AM 79,176, H = 11 mm; New Zealand). (b) apertural view of shell (NHMUK 20170322, Haminea obesa, holotype; H = 17 mm; New Zealand). (c) apertural view of shell (AMS C.159944, H = 22 mm; New Zealand). (d) apertural view of shell, (AMS c.457260, H = 27 mm; New Zealand). (e) SEM, whole radula (AM 119920, H = 11 mm; New Zealand). (f) SEM, detail of rachidian and first laterals in central part of radula (AM 119920, H = 11 mm; New Zealand). (g) Right lateral view of whole gizzard plate (AM 119920, H = 11 mm; New Zealand). (h) SEM, dorsal surface of whole gizzard plate (AM 119920, H = 11 mm; New Zealand). Scale bars: e, g, h = 100 µm. f = 0.15 µm

Figure 5. Papawera zelandiae (Gray, 1843). (a) SEM, detail of ridges of gizzard plate (AM 119920, H = 11 mm; New Zealand). (b) SEM, Detail of jaws (AM 119920, H = 11 mm; New Zealand). (c) detail of external male reproductive system (AM 119920, H = 11 mm; New Zealand). (d) detail of interior of ventrally opened atrium and fundus (AM 119920, H = 11 mm; New Zealand). Abbreviations: as, atrium sheet. at, atrium. bc, body cavity. fu, fundus. pr, prostate. sd, seminal duct. smg, seminal groove. rm, retractor muscles. Scale bars: a = 20 µm. F = 4 µm. c, d = 0.25 mm

Figure 6. Papawera maugeansis (Burn, 1966a). Detail of left lateral Hancock’s organ, dorsal branches partially covered by cephalic shield (ZMBN 125458, H = 8 mm). Abbreviations: cs, cephalic shield. db, dorsal branch. ho, Hancock’s organ. lo, labial organ. pl, parapodial lobe. vb, ventral branch. vm, visceral mass. Scale bar = 0.3 mm

Figure 7. Papawera maugeansis (Burn, 1966a). (a) adpertural view of holotype (NMV F26134, H = 7 mm; Australia). (b) apertural view of shell (ZMBN 125458, H = 8 mm; Australia). (c) apertural view of shell (AMS c.105854, H = 5 mm; Australia). (d) apertural view of shell, (NHMUK MacAndrew col. 1563, H = 7.5 mm; Australia). (e) SEM, whole radula (AMS c.105854, H = 5 mm; Australia). (f) SEM, detail of rachidian and inner laterals in central part of radula (ZMBN 125458, H = 8 mm; Australia). (g) Right lateral view of whole gizzard plate (ZMBN 125458, H = 8 mm; Australia). (h) SEM, dorsal surface of whole gizzard plate (ZMBN 125458, H = 8 mm; Australia). Scale bars: e = 100 µm. f = 50 µm. g, h = 30 µm

Figure 8. Papawera maugeansis (Burn, 1966a). (a) SEM, detail of ridges of gizzard plate (ZMBN 125458, H = 8 mm). (b) SEM, Detail of jaws (AMS c.105854, H = 5 mm). (c) detail of external male reproductive system (AMS c.105854, H = 5 mm). (d) detail of interior of ventrally opened atrium and fundus (ZMBN 125458, H = 8 mm). Abbreviations: as, atrium sheet. at, atrium. fu, fundus. pr, prostate. sd, seminal duct. smg, seminal groove. rm, retractor muscles. Scale bars: a = 20 µm. b = 2 µm. c = 0.5 mm, d = 0.25 mm

Table 3. Synopsis of the most useful morphological characters for diagnosis of species in the genus Papawera.

Characters/Species	Papawera zelandiae (Gray 1843)	Papawera maugeansis (Burn 1966a)
1. External morphology
Animal colouration	Light grey or brown to completely black	Animal light yellowish-grey or light brown to densely black; orange blotches and white dots maybe present
Cephalic shield	Shallowly bilobed	Deeply bilobed
Parapodial and pallial lobes	Large, meet dorsally	Small, do not meet dorsally
2. Shell
Shape	Rounded, bulbous	Oval-elongate
Shell colour	Whitish; periostracum transparent to yellowish	Whitish transparent; periostracum transparent
3. Radula
Formula	27–22.1.1.1.22–27	7–5.1.1.1.5–7
4. Gizzard plates
Ridges	Rachis present; 8–16 ridges	Pseudo-rachis present; 14–20 ridges
Sculpture	Rods along rachis and inner part of posterior side of ridges	Single row of rods along top of ridges
5. Male reproductive system
Prostate	Oblong, tapering, annulated	Oblong, oval, annulated
Penial sheath/Atrium	Fundus semi-enclosed with glandular epithelium	Fundus open, pocket-like
6. Key References	Rudman 1971a, 1971b	Burn 1966a, 1969, 1974

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