Research Article |
Corresponding author: Graham Short ( gshort@calacademy.org ) Academic editor: David Morgan
© 2020 Graham Short, Andrew Trevor-Jones.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Short G, Trevor-Jones A (2020) Stigmatopora harastii, a new species of pipefish in facultative associations with finger sponges and red algae from New South Wales, Australia (Teleostei, Syngnathidae). ZooKeys 994: 105-123. https://doi.org/10.3897/zookeys.994.57160
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A new species of pipefish, Stigmatopora harastii sp. nov., is described based on the male holotype and two female paratypes, 136.3–145.5 mm SL, collected from red algae (sp.?) at 12 meters depth in Botany Bay, New South Wales (NSW), Australia. The new taxon shares morphological synapomorphies with the previously described members of Stigmatopora, including principle body ridges, fin placement, slender tail, and absence of a caudal fin. It is morphologically and meristically similar to Stigmatopora nigra, including snout length and shape, dorsal-fin origin on 6th–7th trunk ring, and lateral trunk ridge terminating on the first tail ring. Stigmatopora harastii sp. nov. is distinguished from its congeners, however, by characters of the head and first trunk ring, distinct sexual dimorphic markings on sides and venter of anterior trunk rings, and red background coloration in life. The new taxon can be further differentiated by genetic divergence in the mitochondrial COI gene (uncorrected p-distances of 9.8%, 10.1%, 10.7%, and 14.6%, from S. argus, S. macropterygia, S. narinosa, and S. nigra, respectively). The type locality is characterised by semi-exposed deep-water sandy areas interspersed with boulders, flat reefs, and an absence of seagrass beds, in which S. harastii has been observed living in facultative associations with a finger sponge and red algae at depths of 10–25 meters, compared to the shallow coastal and estuarine habitats preferred by the fucoid algae and seagrass-associating members of Stigmatopora. Stigmatopora harastii sp. nov. represents the fourth species of Stigmatopora recorded in temperate southern Australia.
Botany Bay, COI, cryptobenthic, ichthyology, Jervis Bay, marine fish, morphology, South Pacific, Sydney, systematics, taxonomy
The Syngnathidae contains over 300 species within 57 genera of predominantly small-bodied and cryptic marine fishes (
The present paper describes a new species of Stigmatopora from NSW, Australia that was first reported by underwater photographers (Oceantrek Diving Resort) in Jervis Bay in 2002, and subsequently observed in Shellharbour and Botany Bay. Stigmatopora harastii occurrs in semi-exposed habitats consistent for the Sydney Basin bioregion (
Type specimens (AMS I. 49510-001, holotype, male; AMS I.47267, paratypes, two females) are deposited in the collections of the Australian Museum (AM).
Counts and measurements to the nearest 0.1 mm were taken from high resolution digital images of specimens using ImageJ (Rasband et al. 1997). Head and body measurements and morphometrics follow Short et al. (2018). External morphological characters were documented using a dissecting microscope and analyses of high-resolution digital images. Georeferenced locations for the type specimens of S. harastii use dWGS84 datum and were captured on GPS units.
DNA extraction, primers and PCR conditions, sequence alignment, and analysis of COI sequence data were performed following protocols described in
Holotype : AMS I.49510-001, male, 145.5 mm SL, collected from a scuba dive area locally referred to as “The Steps”, Kamay Botany Bay National Park, Kurnell, Botany Bay, NSW, Australia, 34°00'07.9"S, 151°13'41.4"E, 13.5 m depth, 18 June 2020, by A. Trevor-Jones and D. Harasti.
Paratypes : AMS I.47267, two females, 130.7 mm and 135.2 mm SL, collected from a scuba dive area known as “The Steps”, Kamay Botany Bay National Park, Kurnell, Botany Bay, NSW, Australia, 34°00'07.9"S, 151°13'41.4"E, 12 m depth, 9 June 2017, by D. Harasti, R. Rodrigues, and A. Trevor-Jones.
Stigmatopora nigra AMS I.42611-009, Botany Bay, NSW, Australia, 03 Feb 2003, K. Parkinson; Stigmatopra narinosa, SAMA F10190, holotype, 150 mm SL, South Australia, Edithburg Pool, 35°05'S, 137°45'E, 31 Dec 2003. Published data was obtained for S. argus, S. macropterygia, S. narinosa, and S. nigra from
Stigmatopora harastii differs from its congeners by the following combination of morphological characters: median ridge, distinct, low, present on dorsum of head and first trunk ring starting from the posterior third of the frontal, over the supraoccipital, to the anterior and posterior nuchal plates; opercular ridge prominent, complete, not angled dorsad; lateromedial ridge, distinct, low, present between opercle and pectoral fin base; dorsal-fin origin on 6th–7th trunk rings, subdorsal rings 19–20 (12 trunk rings + 7 or 8 tail rings); lateral trunk ridge ends on first tail ring. Colouration: red background colour; dorsum of snout with large, irregular pale white spots; sides of head and anterior trunk rings with large, irregular pale white spots or with diffuse pale white stripe; venter of first trunk ring with distinct red elongated spots in longitudinal row, almost forming a stripe, on midline present in male (AMS I. 49510-001); venter of anterior trunk rings pale red with a large cluster of distinct red spots extending posteriad from second trunk ring in male (AMS I. 49510-001), few scattered small red spots in females (AMS I.1.47267).
General body shape as in Figs
Selected counts and morphometric measurements for Stigmatopora harastii. Abbreviations: SnD (snout depth), SnL (snout length), HL (head length), TrL (trunk length), TaL (tail length), SL (standard length).
Voucher | AMS I. 49510-001 | AMS I.47267-001 | AMS I.47267-002 |
---|---|---|---|
Type | holotype | paratype | paratype |
Gender | male | female | female |
Trunk rings | 18 | 18 | 18 |
Tail rings | 68 | 70 | 71 |
Subdorsal rings | 19 | 20 | 20 |
Dorsal-fin origin | 7th trunk ring | 6th trunk ring | 7th trunk ring |
Dorsal-fin rays | 45 | 43 | 43 |
Pectoral-fin rays | 18 | 13 | 13 |
SL (mm) | 145.5 | 136.3 | 138.2 |
SnD/SnL | 7.6 | 7.4 | 7.2 |
SnL/HL | 63.2 | 65.5 | 64.5 |
SnL/TrL | 36.1 | 39.3 | 35.8 |
HL/TrL | 57.1 | 60.0 | 55.6 |
HL/SL | 14.0 | 15.1 | 15.5 |
Trl/SL | 24.5 | 25.2 | 27.9 |
Tal/SL | 61.9 | 60.4 | 56.7 |
In life, S. harastii exhibits red background colouration with unique colour patterns: dorsum of snout with large, irregular pale white spots; sides of head and anterior trunk rings with large, irregular pale white or red spots or with diffuse pale white markings; sexual dimorphic markings with venter of first trunk ring exhibiting distinct red elongated spots in longitudinal row on midline, almost forming a stripe, present in males (Figs
Stigmatopora harastii in situ, AMS I. 49510-001, holotype, male A (right individual) B (left individual); The Steps, Kurnell, Botany Bay, NSW, Australia, 13.5 meters depth, 18 June 2020. The male holotype was photographed with a paired female individual, which was not collected. Note the large cluster of distinct red spots extending posteriad on venter of anterior trunk rings in the male (photographs: Andrew Trevor-Jones).
This species is named after David Harasti, one of the first to recognize S. harastii as being a new species, for recognition of his efforts towards conservation of Syngnathidae in Australia, and for being an aficionado extraordinaire of his beloved genus Stigmatopora. David has stated he counts green pipefish to fall asleep. Harasti’s Pipefish and the Red Wide-bodied Pipefish are proposed here as the common names for S. harastii.
Stigmatopora harastii is currently known to occur in central NSW, Australia from only three localities, including Botany Bay, Shellharbour, and Jervis Bay (Fig.
Single individuals or male-female pairs of S. harastii were observed to be closely associated with the red algae; however, they were infrequently detected and present in only one isolated clump of red algae among all the other clumps in the close vicinity. Multiple dives by the second author between the collection of the type specimens at The Steps in the area where the type specimens were collected, as well as at other nearby large areas with red algae did not consistently detect the presence of S. harastii. Most times individuals were absent or only a single individual was found, possibly suggesting fluctuations in the abundance of the red algae with which S. harastii associates. Individuals were oriented vertically or at an angle and extremely well-camouflaged within the red algae, the distal third of their tails clasped around single fronds. The habitat was subject to strong surge in which individuals of S. harastii and the red algae together were observed swaying in unison with the surge (https://vimeo.com/229093467). The second author has observed on occasion active individuals swimming from one clump of red algae to another or feeding actively just outside the red algae. Feeding behaviour appears to be similar to other Stigmatopora species, with individuals darting out from the cover of their alga to capture food such as small copepods and shrimp. Individuals have been also observed nearby between the Steps and the dive site locally referred to as the Leap, one of which was associated with another species of red algae appearing to be of the genus Gracilaria Greville, 1830 (Fig.
Stigmatopora harastii was also observed at the dive site locally referred to as the Minmi Trench, located at the northern headland of Botany Bay, that consists of a flattish reef with small to large boulders at about 16 meters depth, which then drops off to 22–23 meters depth. A male-female pair was observed associating with a finger sponge appearing to be a member of the family Callyspongiidae Laubenfels, 1936 at 18 meters depth (Fig.
Stigmatopora harastii shares morphological synapomorphies with S. argus, S. macropterygia, S. narinosa, and S. nigra, including principle body ridges, dorsal-fin placement, distally attenuated, slender tail, and absence of caudal fin (
Comparison of morphological characters between S. harastii and other members of Stigmatopora.
S. harastii | S. nigra | S. argus | S. macropterygia | S. narinosa | |
---|---|---|---|---|---|
Data source | This study | Dawson, 1982 | Dawson, 1982 | Dawson, 1982 | Browne & Smith, 2007 |
Trunk rings | 18 | 16–19 | 16–23 | 21–22 | 18 |
Tail rings | 68–71 | 67–79 | 78–91 | 85–92 | 68 |
Dorsal-fin rays | 43–45 | 35–47 | 37–64 | 63–74 | 37–45 |
Pectoral-fin rays | 13–18 | 11–16 | 13–18 | 15–19 | 12–13 |
Dorsal-fin origin | 6th–7th | 5th–7th | 9th–13th | 8th–10th | 5th–7th |
Lateral trunk ridge reaches | 1st tail ring | Anal or 1st tail ring | 8th–20th tail ring | 22nd–35th tail ring | 2nd tail ring |
Snout length | long | long | long | long | medium, laterally flattened, dorsally elevated |
Median dorsal ridge on head and first trunk ring | from frontal to posterior nuchal plate | from frontal to 1st trunk ring | from anterior to posterior nuchal plate | absent | supraoccipital |
Longitudinal ridge between opercle and pectoral-fin base | present | present | absent | absent | absent |
Comparison of the dorsal median ridge present on the head and first trunk ring in: A S. harastii and B S. nigra, AMS I.42611-009. Note the dorsal median ridge (DMR) extending into the first trunk ring in S. nigra versus ending on the posterior nuchal plate in S. harastii. Abbreviations: ANP, anterior nuchal plate; DMR, dorsal median ridge; Fr, frontal; PNP, posterior nuchal plate; SOC, supraoccipital; first TnR, first trunk ring (illustration by Kent Sorgon).
The new species is most easily distinguished from S. nigra by features of the colour pattern in life (Figs
Fucoid algae and seagrass associating members of Stigmatopora in situ A S. nigra, male, Nelson Bay, NSW (photograph: David Harasti) B S. nigra, female, Port Hughes, Gulf St Vincent, South Australia (photograph: David Muirhead) C S. argus, Port Hughes, Gulf St Vincent, South Australia (photograph: Graham Short) D S. narinosa, Port Hughes, Gulf St Vincent, South Australia (photograph: Graham Short) E S. macropterygia, Winstones Cove, North Island, New Zealand (photograph: Nick Shears).
Stigmatopora harastii, S. nigra, and S. narinosa share the presence of large, irregular pale white spots on dorsum of snout (vs. absence of white spots in S. argus and S. macropterygia). Finally, S. harastii and S. nigra share the presence of large, irregular pale white spots with scattering of small red dots, or diffuse pale white stripes, on sides of head and anterior superior trunk rings (vs. absence of colour markings on sides of head and anterior superior trunk rings in S. argus and S. macropterygia). In contrast, Stigmatopora narinosa exhibits a camouflage pattern of brown and white diffuse colouration on sides of head.
1 | Dorsal-fin origin on 5–7th trunk ring, lateral trunk ridge ends on anal, first, or second trunk ring | 2 |
– | Dorsal-fin origin on 8th–13th trunk ring | 5 |
2 | Snout long and slender | 3 |
– | Snout shorter, laterally flattened, and dorsally elevated | 4 |
3 | Red dots on venter of anterior trunk rings (male and female) | Stigmatopora harastii |
– | Black stripes on venter of anterior trunk rings (female) | Stigmatopora nigra |
4 | 2 large transverse pairs of black dots on venter of trunk (female) | Stigmatopora narinosa |
5 | Lateral trunk ridge reaches 8th–23rd tail ring, black spots on dorsum and venter of trunk | Stigmatopora argus |
– | Lateral trunk ridge reaches 22nd–35th tail ring, no distinctive markings on venter of trunk | Stigmatopora macropterygia |
Stigmatopora harastii inhabits semi-exposed bay entrances and ocean embayments in which the underwater terrain is characterised by sandy areas interspersed with boulders and hard flat reefs. Fucoid algae, and seagrass species that occur commonly in NSW coastal and estuarine areas such as Posidonia australis and Zostera capricorni, were absent but were recorded within the shallow areas of the bays or in nearby adjacent bays (
The unique habitat associations of S. harastii with a finger sponge and red algae differ markedly from the fucoid algae and seagrass associating members of Stigmatopora. Stigmatopora argus, S. narinosa, S. nigra, and S. macropterygia inhabit sheltered seagrass and algal beds in bays and estuaries throughout southern Australia and New Zealand, respectively. In NSW, the congenerics S. nigra and S. argus occur in sympatry in seagrass beds and form an abundant component of the inshore ichthyofauna (
Table
Uncorrected genetic distances (p-distances) summary between S. harastii and other members of Stigmatopora based on cytochrome c oxidase I (COI) sequences analysed in this study.
Here we consider S. harastii as a valid species due to its morphological characters of the head, distinct sexual dimorphic markings on the ventral trunk rings, and genetic divergence from its congeners. We have identified a subtle but useful diagnostic morphological character consisting of a median dorsal ridge spanning the frontal, supraoccipital, and anterior and posterior nuchal plates that differentiate S. harastii from the superficially similar S. nigra (Figs
The unique habitat associations of S. harastii with a finger sponge and red algae differ markedly from the fucoid algae and seagrass associating members of Stigmatopora. Even though no close association has been reported between fish and red algae in the literature until this study,
Other species of syngnathids form a component of the inshore ichthyofauna at the type locality where the authors and underwater photographers have recorded the following (1) seahorse species: Hippocampus abdominalis, H. histrix, H. kelloggi, and H. whitei; (2) pipefish species: Festucalex cinctus, Heraldia nocturna, Lissocampus runa, Maroubra perserrata, Notiocampus ruber, and Trachyramphus bioarctatus; (3) pygmy pipehorse species: Idiotropiscis lumnitzeri; and (4) weedy seadragon species: Phyllopteryx taeniolatus. Hippocampus abdominalis, H. kelloggi, and H. whitei have been observed associating with various species of sponges, as well as kelp and large tunicates (Pyura spinifera), and in a wide variety of habitat types at other localities in NSW (
Habitat structural complexity plays an important role in shaping populations, community dynamics, and distribution of seagrass associating pipefish (
We are grateful to many research colleagues and fish enthusiasts who contributed in the field, lab, observations, and congenial discussions: Amanda Hay, Kerryn Parkinson, Sally Reader, and Mark McGrouther, Department of Ichthyology, Australian Museum, for amazing curatorial assistance; Andrew King and Scott Ginn, Australian Centre for Wildlife Genomics, Australian Museum for providing COI sequence data for the paratype (AMS I.47267-001); Merrick Ekins, Queensland Museum (QM), for sponge identification; Michael McFadyen, underwater photographer, for images of the aerial view of the shore dive site The Steps; Duncan Heuer, underwater photographer, for images of S. harastii taken at Minmi Trench; Craig Taylor, underwater photographer, for images of S. harastii taken at the dive site The Gutter, Shellharbour, NSW; Lyn Clairly, Oceantrek Diving Resort, and underwater photographer Sue Newlson, for their observations of S. harastii in Jervis Bay, NSW; David Muirhead, underwater photographer, for images of S. nigra taken at Port Hughes South Australia; Roney Rodriguez, Donna Cassidy, and Cody Sheridan for assistance in location of the type specimens, Kent Sorgon, University of the Philippines Los Baños, for creating the science illustrations of the head morphology of S. harastii and S. nigra; Greg Misner, Aotearoa Hydrogen Alliance, for the generation of the geographic information system (GIS) map of the distribution of S. harastii within Australia. Specimens were collected under NSW Department of Primary Industries permit P01/0059(A)-4.0. This research was supported in part by funding from Aotearoa Hydrogen Alliance, AHA-NZ.Energy.