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http://orcid.org/0000-0003-0014-5560
ABSTRACT
ABSTRACT
The brown algal genus Zonaria (Dictyotales: Dictyotaceae) is widely distributed in temperate waters. Two species, Zonaria turneriana J.Agardh and Zonaria aureomarginata J.A. Phillips & W.A. Nelson have previously been recorded from mainland New Zealand, and a third species, Zonaria diesingiana, has been recorded from the Kermadec Islands. As part of an ongoing study of the order Dictyotales in New Zealand, we have identified an undescribed species of Zonaria present on Stewart Island in southern New Zealand. The new species, described here as Zonaria cryptica, is very similar in external morphology to Z. turneriana, also found on Stewart Island, but differs both genetically and anatomically from Z. turneriana and from other previously described species in the genus. We present a description of the new species based on morphological and anatomical observations, as well as on DNA sequence data from three plastid markers: psaA, psbA and rbcL. Our analysis also indicates the presence of some biogeographic structuring within Z. turneriana and Z. aureomarginata.
Introduction
The genus Zonaria C. Agardh (Dictyotales: Dictyotaceae) was established two centuries ago (Agardh 1817), and currently there are 17 species accepted taxonomically, distributed in the Mediterranean Sea and in temperate regions of the Atlantic and Pacific Oceans (Guiry and Guiry 2017). Species of Zonaria have thalli that are generally flabellate, or lacerate as a result of apical splitting, with growth from a terminal marginal row. The genus Zonaria is defined by both vegetative and reproductive characters (thalli composed of cells regularly aligned in both transverse and longitudinal rows, with cortical cells generally paired above each medullary cell in transverse section; sporangial sori with paraphyses, octosporangia that lack a basal stalk cell; the presence of cortical cells in indusiate sporangial and oogonial sori; antheridial sori slightly raised above the thallus surface) (Papenfuss 1977; Womersley 1987; Phillips et al. 1994; Phillips 1997; Phillips and Clayton 1997).
Phillips and Nelson (1998) traced the recorded history of Zonaria in New Zealand from the earliest reports of Hooker and Harvey (1845), including a summary of the use of synonyms and misapplied names. They lectotypified Zonaria turneriana J.Agardh, and described a new species based on its distinctive anatomy, Zonaria aureomarginata J.A. Phillips & W.A. Nelson. Zonaria aureomarginata was found to possess 10–12 rows of medullary cells that vary greatly in size and that are not always aligned in regular rows, unlike any other species previously described in the genus. In addition, in Z. aureomarginata there were found to be one to four cortical cells above each medullary cell, in contrast to Zonaria turneriana which possesses six to eight medullary cell rows with cells that are very regular in size and shape, and arranged in a very orderly array, and one to two cortical cells per medullary cell. A third species, Zonaria diesingiana J. Agardh, has since been reported (Harper et al. 2012; Nelson and Dalen 2015) from the Kermadec Islands, based on examination of anatomy and morphology. Zonaria diesingiana has two to six medullary cell rows.
Research underway on the order Dictyotales in New Zealand has revealed the presence of taxa that have not been recorded for the region (Harper et al. 2012), including some undescribed species. This paper reports on recent work on the genus Zonaria in New Zealand, presenting phylogenetic analyses based on molecular sequence data that have enabled new understanding of the distribution of species in the region, and the recognition of a new species collected from Stewart Island in southern New Zealand.
Materials and methods
The samples on which this work is based are the result of extensive collection efforts around the New Zealand region (29–53.5°S) with a particular focus on material collected in the past 20 years. Voucher material has been deposited in the Herbarium of the Museum of New Zealand Te Papa Tongarewa (WELT, Thiers 2017). Collection locations, voucher numbers and GenBank accession numbers for specimens included in the phylogenetic analyses are given in .
Table 1. Locations, Voucher numbers and GenBank Accession numbers of sequences in the phylogenetic analysis.
DNA was extracted using the CTAB-Proteinase K method of Zuccarello and Lokhorst (2005). Three plastid markers were amplified using PCR: psaA (primers 870F and 1760R, Yoon et al. 2002, annealing at 41–50 °C), psbA (primers psbAF1 and psbAR2 or psbAR1, Yoon et al. 2002, annealing at 41–50 °C), and rbcL. The rbcL locus was amplified using either primers DJF1 (this study, CTA GCT TTA TTC CGA ATC AC) or DRL2F (Hwang et al. 2005) paired with reverse primer DRL2R (Hwang et al. 2005), annealing at 45 °C. Polymerase chain reaction products were visualised on agarose gels, purified by Exonuclease I/ Alkaline Phosphatase digestion using standard methods, and sequenced at Macrogen Inc. (Seoul, Korea).
Sequences were trimmed and aligned using Geneious V10 (http://www.geneious.com, Kearse et al. 2012). We included in the alignment sequences of other Zonaria specimens available in GenBank, and sequences attributed to other genera within tribe Zonarieae as outgroups, choosing taxa for which sequence data were available for at least two of our markers ().
Phylogenetic analysis
Appropriate models of sequence evolution for the maximum-likelihood analyses were estimated using jModeltest 2 (psaA: GTR + I + Γ; psbA: GTR + Γ; rbcL: GTR + I + Γ; concatenated data set: GTR + I + Γ; Darriba et al. 2012). Maximum-likelihood trees were estimated using PhyML v3.3 (Guindon et al. 2010), using subtree pruning and regrafting branch swapping and assessing support using both the approximate likelihood ratio test (aLRT, Anisimova and Gascuel 2006) and bootstrapping (1000 replicates). Analyses were not partitioned because PhyML does not support partitioning.
For the Bayesian analysis, appropriate partitioning strategies and models of sequence evolution were estimated using Partitionfinder v2.1.1 (Lanfear et al. 2012). Accordingly, the analysis was divided into the following six partitions and models: psaA codon 1 (GTR + G), psbA codon 1 + rbcL codon 2 + psaA codon 2 (GTR + I + G), psaA codon 3 (GTR + I + G), psbA codon 3 + rbcL codon 1 (GTR + I), psbA codon 2 (GTR + I + G) and rbcL codon 3 (GTR + I + G). Four Markov Chain Monte Carlo chains were run for 5 million generations, sampling every 1000 generations, using MrBayes V3.2.6 (Ronquist et al. 2012). Burn-in was assessed through inspection of log-likelihood plots and average parameter values using Tracer V1.6.0 (http://tree.bio.ed.ac.uk/software/tracer/) and confirmed by potential scale reduction factor values calculated in MrBayes. Trees were visualised using FigTree 1.4.0 (http://tree.bio.ed.uk/software/figtree).
Results
The final concatenated data set consisted of a total of 2605 bp (psaA 789 bp, psbA 901 bp, and rbcL 915 bp). Phylogenetic trees derived from each of the single gene data sets were congruent, confirming that concatenation of markers was an appropriate strategy. Results of phylogenetic analyses of the concatenated three-marker data set are shown in Figure 1.
Published online:
28 June 2018Figure 1. Maximum likelihood phylogram estimated from the concatenated psaA, psbA and rbcL data set. Support values are shown on each branch: approximate Likelihood Ratio Test (aLRT) values and maximum likelihood bootstrap (%) above, and Bayesian Posterior Probability (PP) values below. An asterisk indicates support of 1/100/1. Only values greater than 0.8 (aLRT), 70% (bootstrap) and 0.9 (PP) are shown. Support values are only shown if two support methods reach the cut-off value. Scale bar represents substitutions per site.
![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzb20/2018/tnzb20.v056.i03/0028825x.2018.1478310/20180802/images/medium/tnzb_a_1478310_f0001_b.gif"})
Figure 1. Maximum likelihood phylogram estimated from the concatenated psaA, psbA and rbcL data set. Support values are shown on each branch: approximate Likelihood Ratio Test (aLRT) values and maximum likelihood bootstrap (%) above, and Bayesian Posterior Probability (PP) values below. An asterisk indicates support of 1/100/1. Only values greater than 0.8 (aLRT), 70% (bootstrap) and 0.9 (PP) are shown. Support values are only shown if two support methods reach the cut-off value. Scale bar represents substitutions per site.
New Zealand Zonaria specimens were resolved as a monophyletic group divided into three main clades, each of which was supported at aLRT 1.0, bootstrap 100% and Posterior Probability 1.0. Two of these corresponded to Z. aureomarginata and Z. turneriana based on established morphological and anatomical criteria for these taxa.
Within both Z. aureomarginata and Z. turneriana some genetic differentiation was apparent: each was further resolved into two sub-clades. In both species, one sub-clade contained sequences from specimens collected in northern localities, while the second contained sequences from specimens collected over a much wider geographic range.
Specimens identified as Z. aureomarginata collected at Manawatāwhi (Three Kings Islands), were resolved as a sister clade to specimens from the North Island, Chatham Island and the northern South Island of New Zealand (Marlborough Sounds). In the case of Z. turneriana, the northern sub-clade included specimens collected from Manawatāwhi and also from the Bay of Islands (northeastern North Island), and the more widely distributed clade included representatives from collections from Whangarei (northern North Island) through to Stewart Island in the south of New Zealand.
The third New Zealand Zonaria clade consisted of five specimens collected from three sites on Stewart Island, the southernmost of the three main islands of the New Zealand archipelago. This clade was resolved as a sister clade to Z. turneriana, but differed from it by over 26 bp (92% identity, psaA), 16 bp (98%, psbA) and 19 bp (97% identity, rbcL). They could be distinguished from Z. aureomarginata and Z. turneriana on the basis of the number of cell rows in the thallus. We consider that this degree of genetic differentiation coupled with morphological distinctiveness warrants recognition at the species level.
Published online:
28 June 2018![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzb20/2018/tnzb20.v056.i03/0028825x.2018.1478310/20180802/images/medium/tnzb_a_1478310_f0002_c.jpg"})
Figure 2. Holotype of Zonaria cryptica –WELT A033574.
Published online:
28 June 2018Figure 3. Cross section of Isotype (WELT A033573) showing eight cell rows, with two cortical cells per medullary cell. Scale bar = 20 µm.
![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzb20/2018/tnzb20.v056.i03/0028825x.2018.1478310/20180802/images/medium/tnzb_a_1478310_f0003_c.jpg"})
Figure 3. Cross section of Isotype (WELT A033573) showing eight cell rows, with two cortical cells per medullary cell. Scale bar = 20 µm.
Published online:
28 June 2018Figure 4. Cross-section of Holotype showing stupose filaments on upper and lower surfaces of the blade. Scale bar = 100 µm.
![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzb20/2018/tnzb20.v056.i03/0028825x.2018.1478310/20180802/images/medium/tnzb_a_1478310_f0004_c.jpg"})
Figure 4. Cross-section of Holotype showing stupose filaments on upper and lower surfaces of the blade. Scale bar = 100 µm.
Published online:
28 June 2018Figure 5. Cross-section of Holotype showing development of filaments from cortical cells on upper and lower surfaces. Scale bar = 50 µm.
![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzb20/2018/tnzb20.v056.i03/0028825x.2018.1478310/20180802/images/medium/tnzb_a_1478310_f0005_c.jpg"})
Figure 5. Cross-section of Holotype showing development of filaments from cortical cells on upper and lower surfaces. Scale bar = 50 µm.
Zonaria cryptica sp. nov. W.A. Nelson & J.E. Sutherl. Figures 2–5
Diagnosis
Thalli erect with branched fan-shaped fronds (Figure 2). Branches flattened and linear, expanding to rounded, fan-shaped tips to 15 mm across with apical growth from the marginal cell row. Blades with six to eight regularly arranged cell rows in transverse section (Figures 3–5), with fewer cell layers towards the margins and apex. Thalli have four to six layers of medullary cells, and a single cortical row on the adaxial and abaxial surfaces, and one to two cortical cells per medullary cell. Holdfast and lower part of the thallus stupose (Figures 4–5), with filaments continuing up the mid-line of the thallus, more densely on the abaxial surface.
Holotype: New Zealand, Stewart Island, Tikotahi Bay (−47.08689, 168.15186), 31 March 2017, R. D'Archino, (ASO058), WELT A033574; on rock, subtidal, 5 m depth.
Isotypes: WELT A033573 (ASR057), WELT A033577 (ASR074)
Etymology: cryptica in reference to this species being unrecognised because of its morphological similarity to Z. turneriana occurring in the same region.
Morphological and anatomical observations
At present Z. cryptica is known from five collections, three of which were from the same site, and all specimens were collected in March (late summer/autumn). Based on the material available, this species falls within the range of size found in Z. turneriana. Although all specimens collected to date are less than 15 cm high, it is not known whether this is the maximum height attained by this species. The cell arrangement in the medulla is typical of the genus, with very regularly arranged rows and columns of cells, unlike the arrangement found in Z. aureomarginata, in which some medullary cells are deeper or taller than others. All specimens collected to date are sterile. The lower parts of the axes are markedly stupose, a feature also seen in Z. turneriana but which appears to be particularly well developed in Z. cryptica.
Representative specimens
Zonaria cryptica: Stewart Island: Port Adventure, 6.5 m depth in Macrocystis bed, 30 March 2017, R. D'Archino, (ASR 034), WELT A033575; Port Pegasus, North Arm, Albion Inlet, 6 March 2009, 8–12 m, R. D'Archino, (ASJ311), WELT A033576.
Zonaria aureomarginata–northern sub-clade: Manawatāwhi: Great Island, South East Bay, 20 m depth, 19 April 2013, R. D'Archino, (ASN747), WELT A033571; also (ASN748), WELT A033572, and (ASN729), WELT A033647; 14 April 2013, R. D'Archino (ASN551), WELT A033644.
Zonaria aureomarginata–main clade: North Island, Hawke’s Bay, Mangakuri, 23 February 2016, W.A. Nelson, (ASQ021), WELT A033648; North Island, Wellington, Owhiro Bay, 3 September 2007, R. D'Archino, S. Miller, T. Farr, W. Nelson, (ASH284), WELT A024491; North Island, Wellington, Lyall Bay, 19 June 2007, W.A. Nelson & R. D'Archino, (ASH250), WELT A024435; Chatham I., Manukau reef, 31 January 2006, W. Nelson, (ASG202), WELT A024220; South Island, Marlborough Sounds, 23 March 2017, R. D'Archino (MS194), WELT A033649.
Zonaria turneriana–northern sub-clade: Manawatāwhi, Princes Islands, north side of middle island, slope on west side of northern opening of the archway, 18–21 m depth, 10 April 2013, R. D'Archino, (ASN383), WELT A033578; Manawatāwhi 14 April 2013, R. D'Archino, (ASN550), WELT A033643, also R. D'Archino, (ASN552), WELT A033645; Manawatāwhi, 17 April 2013, R. D'Archino (ASN588), WELT A033646, also Great Island, South East Bay, 14.5–19 m depth, 17 April 2013, R. D'Archino, (ASN605), WELT A033579; North Island, Bay of Islands, Te Miko Reef, 08 February 2010, S. Miller & R. D'Archino, (RHO0015), WELT A033650;
Zonaria turneriana–main clade: North Island, Marsden Point, Whangarei, 01 October 2009, R. D'Archino & S. Miller, (SS1138), WELT A031160; North Island, Auckland, Brown’s Bay, 13 December 2012, J.E. Sutherland, (MATS2818), WELT A033580; North Island, Wellington, Princess Bay, 7 October 2007, M. Francis, (ASH442), WELT A024428; South Island, Fiordland, Doubtful Sound, Shelter I, 27 January 2008, C. Hepburn & D. Richards, (ASI055), WELT A033583; South Island, Fiordland, Caswell Sound, Hansard Pt, 3 February 2009, R. D'Archino, (ASJ148), WELT A033642; South Island, Fiordland, entrance to Doubtful Sound, 25 January 2008, C. Hepburn & D. Pritchard, (ASI020), WELT A033581; South Island, Fiordland, Bligh Sound, Postman’s Bluff, 5 February 2009, R. D'Archino, (ASJ156), WELT A033582; South Island, Fiordland, Dusky Sound, Cook Channel, 31 January 2009, R. D'Archino, (ASJ069), WELT A033641; South Island, Fiordland, Doubtful Sound, Shelter I, 27 January 2008, C. Hepburn & D. Richards, (ASI056), WELT A033584; Stewart I, East of Bishop, 1 April 2017, R. D'Archino, (ASR100), WELT A033585.
Zonaria diesingiana: Kermadec Islands, Meyer Islands, 2 June 1992, J. Williamson, WELT A032337.
Discussion
Investigations of Dictyotales based on extensive collections and employing molecular tools have revealed the presence of a species of Zonaria previously unrecognised in New Zealand, and an improved understanding of the distributions of Z. aureomarginata and Z. turneriana within the New Zealand archipelago. Before this study, Zonaria in New Zealand was thought to be represented by these two ‘mainland species’, and a third species, Z. diesingiana, described from Australia and present within the New Zealand region only on the Kermadec Islands/Rangitāhua (Harper et al. 2012).
Morphological species delimitation in Zonaria is challenging: comparative studies on the Australian species of Zonaria have shown that ‘species can only be delimited by differences in thallus morphology’ (Phillips 1997). Both vegetative and reproductive anatomy (sporangial and gametangial structure) are remarkably similar across all species of Zonaria investigated to date (Phillips et al. 1994; Phillips 1997; Phillips and Clayton 1997). Except for the lack of sporangial paraphyses in Zonaria angustata (Kütz.) Papenf., and the irregular medullary cell rows in Z. aureomarginata, anatomical features are uninformative in species discrimination for Australasian species in this genus (Phillips 1997). Securing fertile material may be enabled by future collections from different seasons and also lunar phases, which was effective in locating all reproductive stages in Z. aureomarginata, and also noted for species of Zonaria and Homeostrichus by Phillips and Clayton (1997).
Although both Z. aureomarginata and Z. turneriana show evidence of geographically based differentiation with genetically distinct northern sub-clades, members of these northern sub-clades cannot be distinguished morphologically. We do not consider that the genetic differences between the sub-clades in each species justify recognition of separate species. The Z. aureomarginata northern and mainland New Zealand clades differ by a maximum of 3, 2 and 4 bp at psaA, psbA and rbcL respectively, while the Z. turneriana northern and southern clades differ by a maximum of 4, 8 and 6 bp; in both cases the percentage sequence identity between the clades exceeds 99.5%. The genetic break between taxa on Manawatāwhi and those on the North Island of New Zealand has been observed in other species, with high endemicity seen on these offshore islands. Our study supports the observation of limited gene flow between Manawatāwhi and the North Island (Nelson and D’Archino 2014).
As a result of this study, four species of Zonaria can be recognised in New Zealand waters from north to south, based on the number of cell layers in combination with their distribution: Z. diesingiana (four to eight cell layers) at the Kermadec Islands; Z. aureomarginata (12–14 cell layers), present from Manawatāwhi, North, Chatham to the South Island; Z. turneriana (8–10 cell layers), from Manawatāwhi, North, Chatham, South, and Stewart Islands, and Z. cryptica (six to eight cell layers) from Stewart Island. Sequence data are still required to confirm the identity of Z. diesingiana from the Kermadec Islands. A map of the currently understood distributions of these species within New Zealand is presented in the Supplementary material Figure S1. In central New Zealand where Z. aureomarginata and Z. turneriana overlap in range, these species can be distinguished in most cases by habitat and external morphology, with Z. aureomarginata generally occurring in shallower waters and with distinctively broader and more golden apices. Zonaria cryptica may be present in other southern New Zealand locations, although recent collections in Fiordland (southwestern South Island) did not reveal any material of this new species. It is not clear yet, given the limited number of collections available, whether it will be possible to distinguish Z. cryptica from Z. turneriana in Stewart Island where they are sympatric, by habitat, depth or growth characteristics.
| Taxon | Location/Strain/Voucher (WELT) | GenBank Accession No. | |||
|---|---|---|---|---|---|
| Reference | psaA | psbA | rbcL | ||
| Outgroups | |||||
| Lobophora pachyventera | Taiwan | Wang et al. 2013 | DQ866963.1 | DQ866942.1 | DQ866930.1 |
| Lobophora australis | South Australia | Wang et al. 2013 | DQ866958.1 | DQ866944.1 | DQ866924.1 |
| Homoeostrichus formosana | Taiwan | Wang et al. 2013 | DQ866964.1 | DQ866952.1 | DQ866931.1 |
| Homoeostrichus canaliculatus | South Australia | Wang et al. 2013 | DQ866956.1 | DQ866943.1 | DQ866922.1 |
| Padina japonica | Korea (psaA + psbA) and Japan (rbcL) | rbcL Hoshina et al. 2004; psaA and psbA Wang et al. 2013 | AY430319.1 | AY430360.1 | AB096905.1 |
| Padina australis | – | Bittner et al. 2008 | EU579915.1 | – | EU579959.1 |
| Spatoglossum asperum | – | Bittner et al. 2008 | EU579921.1 | – | EU579963.1 |
| Spatoglossum crassum | South Korea | Wang et al. 2013 | AY430314.1 | AY430355.1 | AY430336.1 |
| Dictyopteris polypodioides | – | Bittner et al. 2008 | EU579899.1 | – | EU579932.1 |
| Dictyopteris australis | – | Bittner et al. 2008 | EU579895.1 | – | EU579940.1 |
| Zonaria | |||||
| Zonaria crenata | South Australia | Wang et al. 2013 | DQ866970.1 | DQ866955.1 | DQ866936.1 |
| Z. tournefortii | – | Bittner et al. 2008 | EU579930.1 | – | EU579973.1 |
| Z. subarticulata | – | Bittner et al. 2008 | EU579929.1 | – | EU579972.1 |
| Z. spiralis | – | Bittner et al. 2008 | EU579928.1 | – | EU579971.1 |
| Z. angustata | South Australia | Wang et al. 2013 | DQ866966.1 | DQ866946.1 | DQ866932.1 |
| Z. diesingiana Korea | South Korea, WJ8 | Hwang et al. 2005 | AY422606.1 | AY422644.1 | AY422682.1 |
| Z. diesingiana KU1071 | KU-MACC:KU-1071 | Kawai et al. 2015 | AB899217.1 | AB899260.1 | AB899284.1 |
| New Zealand Zonaria specimens | |||||
| Z. aureomarginata | Three Kings Is Manawatāwhi/ASN748 /A033572 | This study | MG811833.1 | MG807130.1 | MG807160.1 |
| Z. aureomarginata | Three Kings Is Manawatāwhi/ASN747/A033571 | This study | MG807104.1 | MG807129.1 | MG807159.1 |
| Z. aureomarginata | Three Kings Is Manawatāwhi/ASN729/A033647 | This study | MG807103.1 | MG807128.1 | MG807158.1 |
| Z. aureomarginata | Three Kings Is Manawatāwhi/ASN551/A033644 | This study | MG811832.1 | MG811829.1 | MG807154.1 |
| Z. aureomarginata | North I, Wellington, Owhiro Bay/ASH284/A024491 | This study | MG807096.1 | MG807119.1 | MG807148.1 |
| Z. aureomarginata | Chatham I, Manukau reef/ASG202/A024220 | This study | MG807091.1 | MG807117.1 | MG807143.1 |
| Z. aureomarginata | South I, Marlborough Sounds/MS194/A033649 | This study | MG807111.1 | MG807137.1 | MG807167.1 |
| Z. aureomarginata | North I, Wellington, Lyall Bay/ASH250/A024435 | This study | MG807092.1 | MG807118.1 | MG807118.1 |
| Z. aureomarginata | North I, Hawke’s Bay, Mangakuri/ASQ021/A033648 | This study | MG807105.1 | MG807131.1 | MG807131.1 |
| Z. cryptica | Stewart I, Tikotahi Bay/ASR074/A033577 | This study | MG807116.1 | MG807142.1 | MG807172.1 |
| Z. cryptica | Stewart I, Tikotahi Bay/ASR057/A033573 | This study | MG807113.1 | MG807139.1 | MG807169.1 |
| Z. cryptica | Stewart I, Tikotahi Bay/ASR058/A033574 | This study | MG807114.1 | MG807140.1 | MG807170.1 |
| Z. cryptica | Stewart I, Port Adventure/ASR034/A033575 | This study | MG807112.1 | MG807138.1 | MG807168.1 |
| Z. cryptica | Stewart I, Port Pegasus/ASJ311/A033576 | This study | MG811830.1 | MG807121.1 | MG807150.1 |
| Z. turneriana | Three Kings Is Manawatāwhi/ASN383/A033578 | This study | MG807099.1 | MG807123.1 | MG807152.1 |
| Z. turneriana | Three Kings Is Manawatāwhi/ASN550/A033643 | This study | MG811831.1 | MG807124.1 | MG807153.1 |
| Z. turneriana | North I, Bay of Islands, Te Miko Reef/RHO0015/A033650 | This study | MG807109.1 | MG807135.1 | MG807165.1 |
| Z. turneriana | Three Kings Is Manawatāwhi/ASN605/A033579 | This study | MG807102.1 | MG807127.1 | MG807157.1 |
| Z. turneriana | Three Kings Is Manawatāwhi/ASN588/A033646 | This study | MG807101.1 | MG807126.1 | MG807156.1 |
| Z. turneriana | Three Kings Is Manawatāwhi/ASN552/A033645 | This study | MG807100.1 | MG807125.1 | MG807155.1 |
| Z. turneriana | North I, Auckland, Brown’s Bay/MATS2818/A033580 | This study | MG807098.1 | MG807122.1 | MG807122.1 |
| Z. turneriana | North I, Marsden Pt, Whangarei/SS1138/A031160 | This study | MG807110.1 | MG807136.1 | MG807166.1 |
| Z. turneriana | South I, Fiordland, Doubtful Sound/ASI056/A033584 | This study | MG807095.1 | – | MG807147.1 |
| Z. turneriana | Stewart I, East of Bishop/ASR100/A033585 | This study | MG807115.1 | MG807141.1 | MG807171.1 |
| Z. turneriana | North I, Wellington, Princess Bay/ASH442/A024428 | This study | MG807097.1 | MG807120.1 | MG807120.1 |
| Z. turneriana | South I, Fiordland, Doubtful Sound/ASI055/A033583 | This study | MG807094.1 | – | MG807146.1 |
| Z. turneriana | South I, Fiordland, Caswell Sound/ASJ148/A033642 | This study | MG807108.1 | MG807134.1 | MG807164.1 |
| Z. turneriana | South I, Fiordland, Doubtful Sound/ASI020/A033581 | This study | MG807093.1 | – | MG807145.1 |
| Z. turneriana | South I, Fiordland, Bligh Sound/ASJ156/A033582 | This study | MG807107.1 | MG807133.1 | MG807163.1 |
| Z. turneriana | South I, Fiordland, Dusky Sound/ASJ69/A033641 | This study | MG807106.1 | MG807132.1 | MG807162.1 |
Supplemental Material. Map of the currently understood distributions of Zonaria species within New Zealand waters.
Download PNG Image (239 KB)
Related Research Data
Acknowledgements
We acknowledge the support of SSIF funding within NIWA, collections made through a range of projects funded by the Ministry for Primary Industries, collections from Manawatāwhi in conjunction with the Auckland Museum (led by Dr Tom Trnski), collections from Stewart Island, Otago and Fiordland in conjunction with University of Otago Department of Marine Sciences (led by Ass. Prof Chris Hepburn). We thank staff of the Herbarium of the Museum of New Zealand Te Papa Tongarewa for assistance with registration of specimens and providing the image of the Holotype, and Erika Mackay and Kate Neill (NIWA) for preparing the illustrations.
Disclosure statement
No potential conflict of interest was reported by the authors.
ORCID
Wendy A. Nelson http://orcid.org/0000-0003-0014-5560
Judy E. Sutherland http://orcid.org/0000-0002-1722-8218
References
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