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Heaviside's dolphin
Heaviside's dolphin off Lüderitz, Namibia
Size compared to an average human
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Infraorder: Cetacea
Family: Delphinidae
Genus: Cephalorhynchus
Species:
C. heavisidii
Binomial name
Cephalorhynchus heavisidii
Gray, 1828
Heaviside's dolphin range

Heaviside's dolphin (Cephalorhynchus heavisidii),[2] is one of four species in the genus Cephalorhynchus. The small cetacean is endemic to the Benguela ecosystem along the southwest coast of Africa.[3][4]

Taxonomy and evolution

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Nomenclature

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Early in the 19th century, a specimen was caught off the Cape of Good Hope and brought to the United Kingdom by a Captain Haviside of the British East India Company. Zoologist John Edward Gray, who described the species in his Spicilegia Zoologica,[2] misidentified Haviside as the surgeon John Heaviside, whom was known for his own biological collections at the time.[5] "Heaviside's dolphin" is the recognised common name. Other common names include "Haviside's dolphin" and "Benguela dolphin" the latter especially in Namibia.

The genus name Cephalorhynchus comes from the Greek kephale for ‘head’ and rhynchos for ‘beak’. For the species name "heavisidii" see the above description.

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The genus Cephalorhynchus includes the Chilean dolphin (C. eutropia) and the two sub-species of both the Hector's dolphin (Hector’s dolphin, C. hectori hectori and Māui dolphin, C. h. maui), and the Commerson's dolphin (South American Commerson’s dolphin, C. commersonii commersonii and Kerguelen Islands Commerson’s dolphin, C. c. kerguelenensis). All are located in cool, temperate waters in the Southern Hemisphere.[6]

Genetic studies suggest that the Cephalorhynchus dolphins originated from a single common ancestor in southern Africa, from which Heaviside's dolphins are the basal species.[7] Radiation around the southern hemisphere following the Antarctic Circumpolar Current (otherwise known as the West Wind Drift), first to New Zealand and then to South America, is thought to have led to the subsequent speciation within the genus.[7]

Description

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Morphology

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Heaviside's dolphins off Walvis Bay, Namibia

Heaviside's are small and stocky with adults reaching a maximum length and weight of 1.7m and 75kg respectively.[3] The head is cone shaped with a blunt beak. The dorsal fin is triangular in shape and centred in the middle of the back. The dolphin has a distinct black, grey and white body pattern, and is not easily confused with any other species in its range.[3] The head and thorax are coloured light grey with darker patches around the eye. The dorsal fin, fluke and dorsal cape are a dark grey to bluish black with a band that extends forward from the dorsal fin to the blowhole. The underbelly is white, with bands that extend onto the lower rear of the body forming a trident shape. Small white patches are located just behind the pectoral fins and a single white patch extends between these fins on the chest. Sexual dimorphism is minimal, however variation in the shape of the white patch covering the genital slit is distinct between genders. In males, the patch ends in a point, but in females widens out to cover the mammary slits.[3]

Life history

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Information on reproduction is limited for Heaviside's dolphins, however they are thought to be comparable to Hector's and Commerson's dolphins.[6] Females and males reach sexual maturity approximately between 5–9 years. Mating is thought to occur year-round, however individual females may only produce calves every 2–4 years. Gestation time is unknown. Maximum known lifespan is based on the oldest recorded individual at 26 years old.[5]

Group size

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Typically occurs in small groups of 2–3, but numbers of 1-10 are frequent and large aggregations of 100 individuals or more are known to form in high density areas.[5] Nursery groups (groups exclusively containing adult females and calves) are not formed in this species.[8]

Predation

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Levels of predation are unknown, however killer whales (Orcinus orca) are known predators and there is evidence of shark attack from body scars.[9]

Distribution

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Geographic range

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The species is strongly associated with the cool waters of the Benguela Ecosystem. Although the southern limit of the range is defined as Cape Point, the real southern limit beyond which sightings are extremely rare is Hout Bay, some 40km to the north (a considerable distance for a species which shows very high site fidelity to quite small spatial scales when nearshore). The species occurs more or less continuously for approx. 2500km to the north of this along the South African coast, through Namibia and into southern Angola where the northern boundary for the species remains poorly defined. Several dolphins have been sighted or accidentally caught by fishing vessels north of the Angola-Namibia border, [10][11][12][13] but no sightings were reported during a series of coastal scientific surveys at Tombua which is roughly 170km into Angola but well south of defined northern boundary of the Benguela Ecosystem (REFERENCE). The northern boundary of the Benguela current shifts north and south seasonally and as Heaviside’s dolphins appear closely linked to it’s cool waters, their northern range limit may shift along with water conditions.

Systematic surveys have dedicated effort to describing the distribution in southern Africa South Africa[8] and current research efforts focus on local populations in Walvis Bay and Lüderitz, Namibia (Martin x2 and de rock). These locations are also popular hotspots for watching these dolphins in addition to Table Bay (Cape Town) and and Britannia Bay in South Africa. Sightings are common from land and there are several dolphin watching tour companies by which Heaviside’s dolphins can be seen by boat.[14]

Recent genetic research has demonstrated evidence of population structure across the range, indicating two metapopulations (north and south) with limited genetic exchange.[15] This pattern of fragmentation is a common feature amongst the species in the genus Cephalorhynchus most prevalent in the Hector's dolphin, which displays genetic isolation over very short distances.[16]

Habitat preferences

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Heaviside’s dolphins typically remain nearshore in the mornings where they typically socialise and rest. When nearshore their distribution patterns are remarkably predictable within and between years, with the animals showing highly consistent use of aggregation sites at the exposed western tips of most bays throughout South Africa and Namibia, but they are rarely seen in the protected shallows of these bays. Outside of bays, they show relatively high densities along exposed sandy beaches, but these may be a secondary choice after a preference for areas where there is a high abundance of their main prey item; juvenile hake (Merluccius capensis) in adjacent offshore waters. Most commonly sighted within sea surface temperatures of 9-15°C, depths less than 100m. [8]

Behaviour

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Heaviside's dolphins are energetic and social animals, especially when nearshore in the mornings. Behaviour when offshore tends to be less playful and may include a rest phase when moving offshore to feed (Elwen et al. 2006, 2009). They are attracted to boats and frequently bow-ride.[17] Individuals can also be seen surfing in coastal waves. Iconic vertical leaps clear the water before re-entering headfirst with almost no splash.[5] Mating typically occurs in social groups of 3-7 individuals which remain in a small area exhibiting extensive rolling, touching and position changes with frequent leaps by one or pairs of animals which potentially serve a competitive function.

Diet and Foraging

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Prey items consist of mostly demersal fish and cephalopod species, predominantly juvenile hake (Merluccius capensis) and octopus, however pelagic species such as juvenile goby (Sufflogobius bibarbatus) and horse mackerel (“Trachurus trachurus capensis”) are also consumed, especially in Namibia.[18] Foraging occurs mostly at the seabed, in shallow depths. Feeding nearshore is rarely observed.[19]

Movement patterns

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A diurnal movement pattern is present in South Africa whereby the dolphins move offshore in the afternoon to feed on prey rising vertically to the surface at night.[20] Movement inshore to rest and socialise occurs in the morning.[17] However, the pattern is different in Lüderitz and Walvis Bay, Namibia where the movement is less pronounced and dolphins appear to stay inshore during the night, which is likely associated with foraging on different prey.[21]

Home range and site fidelity

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Heaviside's have small home ranges of 50–80km as measured using satellite telemetry over 2-3 months and photographic re-sightings over up to 3 years. Some individuals have been re-sighted at the same location for up to 10 years.[22]

Dive time and depth

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There has been limited research into Heaviside's diving behaviour, however a study of two dolphins fitted with satellite tags was undertaken in South Africa in 1997.[23] The maximum dive depth recorded was 147 meters, however the majority of dives were less than 50 meters.[23] Dive duration were predominantly less than 2 minutes with most dives between 0 and 1 minutes.[23]

Sympatry with other delphinids

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Whilst typically found further from shore, dusky dolphins (Lagenorhynchus obscurus) are found throughout the range and occasionally both dolphin species are sighted in mixed groups.[8] Where both species overlap in prey selection, Heaviside's dolphins take larger prey items, potentially because they are outcompeted by the larger dusky dolphins for their preferred, smaller sized prey.[24] In central Namibia (especially Walvis Bay) Heaviside’s dolphins overlap with a small population of fewer than 100 common bottlenose dolphins (Tursiops truncatus).[25] The bottlenose dolphin population use only water less 15m depth while in this area Heaviside’s dolphins are almost always encountered in water deeper than 20m (Predicting habitat use paper?) suggesting some form of competitive exclusion[26] although direct interactions have rarely been observered and none were agonistic.

Vocalisations

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As is the case with all species in the genus, Heaviside's produce narrowband high-frequency (NBHF) echolocation clicks (centred around 125–130 kHz), and do not whistle.[27] This adaptation is theorised to allow acoustic crypsis from eavesdropping predators, as the sounds produced are outside of the detectable frequencies of killer whales.[28] Although NBHF clicks are limited in acoustic range, they have a better resolution for small targets and are thought to provide a foraging advantage in the often cluttered, nearshore environment in which these species occur.[29] Heaviside's also produce a second click type, of lower frequency and broader bandwith, that is within the hearing range of orcas.[30] These calls are produced most frequently in large groups engaging in social behaviour. It is likely that the dolphins use these calls when socialising away from predator threat and switch to high frequency clicks when foraging and travelling.[31]

Population status

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A total abundance has not been calculated, however regional estimates exist in well studied locations. An estimate of 6,345 for the region between Table bay and Lamberts bay, South Africa represents the southernmost populations in the species range.[17] Local population estimates for Walvis bay and Lüderitz are 508 and 494 respectively.[4] A visual and acoustic line-transect ship survey estimated an average of 1594 dolphins in the Namibian Islands’ Marine Protected Area (NIMPA) which spans 400 km of coastline along southern Namibia.[26] Population trends are unknown however, high mitochondrial DNA diversity suggests a relatively stable population,[6] and there is no evidence of population decline.

Threats

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Heaviside’s dolphins are exposed to a variety of anthropogenic threats, including fisheries bycatch related mortality and illegal directed catch (Elwen and Gopal, 2018; Alfaro-Shigueto et al., 2019), changes in prey availability (Coetzee et al., 2008; Roux et al., 2013) and marine eco-tourism (Leeney, 2014).

Bycatch and hunting

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Whilst Heaviside’s dolphins are afforded full legal protection from hunting, direct mortality from fisheries bycatch is considered an ongoing threat.[4] Recently developed mid water trawls for horse mackerel (“Trachurus capensis”) are considered an emerging threat.[4] Directed catch has historically been associated with meat being used for human consumption.[32]

Climate change

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Heaviside's dolphins have been identified as a species vulnerable to the effects of climate change.[33] Warming Sea Surface Temperatures are likely to be problematic for Heaviside’s dolphins and all species in the Cephalorhynchus genus given their preference for cool waters. These species are likely to decline as there is a lack of shallow habitat toward the pole for them to utilise more favourable temperatures.

Heaviside’s dolphins are listed amongst the cetacean species most vulnerable to climate change (Macleod, 2009) as they are limited to a distribution range that includes both suitable shelf habitat and cool water temperatures (Best, 2007). The Benguela Current is the only eastern-boundary current bordered by warm-water currents at both its northern and southern limits. Consequently, small increases in water temperature could result in all shelf waters in this area becoming too warm for both species, and populations are expected to decline rather than shift poleward due to a lack of shelf habitat south of their existing ranges (Macleod, 2009).

Boat interactions

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Heaviside's are exposed to boat traffic and marine tourism At least one vessel strike has been observed[34] and negative effects have been demonstrated for other coastal cetacean species. marine eco-tourism (Leeney, 2014).

Conservation status

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Heaviside’s dolphins are listed by the International Union for Conservation of Nature (IUCN) as ‘Near Threatened’ [1]. Prior to 2018, the species was listed as ‘Data Deficient’ however, research has improved knowledge on the species, leading to the re-classification. Despite this, the overall population trend remains unknown,[4] and there are many aspects of the species biology that remain to be studied.

Heaviside's dolphin is listed on Appendix II of the Convention on the Conservation of Migratory Species of Wild Animals[35] and is included in the Memorandum of Understanding Concerning the Conservation of the Manatee and Small Cetaceans of Western Africa and Macaronesia. The Memorandum of Understanding was established in 2008 and aims to protect these species at a national, regional and global level.

References

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Citations

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  1. ^ a b Elwen, S.; Gopal, K. (2018), "Heaviside's Dolphin, Cephalorhynchus heavisidii", The IUCN Red List of Threatened Species 2018: e.T4161A50352086, doi:10.2305/IUCN.UK.2018-2.RLTS.T4161A50352086.en
  2. ^ a b Gray, John Edward (1828). Spicilegia Zoologica: Original Figures and Short Systematic Descriptions. Vol. Vol.1. Soho, London: Treüttel, Würtz & Co. {{cite book}}: |volume= has extra text (help)
  3. ^ a b c d Best, Peter B. (2007). Whales and Dolphins of the Southern African Subregion. Cambridge University Press. p. 388. ISBN 978-0-521-89710-5.
  4. ^ a b c d e Gopal, K; Elwen, S; Plön, S (2016). "A conservation assessment of Cephalorhynchus heavisidii. In Child MF, Roxburgh L, Do Linh San E, Raimondo D, Davies-Mostert HT. The Red List of Mammals of South Africa, Swaziland and Lesotho" (PDF). South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa.
  5. ^ a b c d Carwardine, Mark (2020). Handbook of Whales, Dolphins and Porpoises. Bloomsbury Publishing. pp. 359–361. ISBN 978-1-4729-7715-1.
  6. ^ a b c Dawson, S (26 February 2009). "Cephalorhynchus dolphins". In Würsig, B; Thewissen, J.G.M; Kovacs, K.M (eds.). Encyclopedia of Marine Mammals (3rd ed.). London: Academic Press. ISBN 9780080919935.
  7. ^ a b Pichler, F; D. Robineau, R; Goodall, M; Meyer, M; Olivarria, C; Baker, C (2001). "Origin and radiation of Southern Hemisphere coastal dolphins (genus Cephalorhynchus)". Molecular Ecology. 10 (9): 2215–2223. doi:10.1046/j.0962-1083.2001.01360.x. PMID 11555263. S2CID 24368161.
  8. ^ a b c d Elwen, S.H.; Thornton, M.; Reeb, D.; Best, P.B. (2010). "Near-shore distribution of Heaviside's (Cephalorhynchus heavisidii) and dusky dolphins (Lagenorhynchus obscurus) at the southern limit of their range in South Africa". African Zoology. 45 (1): 78–91. doi:10.1080/15627020.2010.11657256. hdl:2263/14283. ISSN 1562-7020. S2CID 219289679.
  9. ^ Best, Peter B.; Abernethy, R. Blake (1994). "Heaviside's dolphin, Cephalorhynchus heavisidii (Gray, 1828)". In Ridgway, Sam H.; Harrison, Richard (eds.). Handbook of Marine Mammals. Vol. Vol. 5 The First Book of Dolphins. Academic Press. ISBN 978-0125885058. {{cite book}}: |volume= has extra text (help)
  10. ^ Morias, Miguel (2012). "Marine mammal sightings off the Angolan coast recorded from the R/V Dr. Fridtjof Nansenin August 2004 and July 2005". In Van Waerebeek, Koen (ed.). Conserving cetaceans and manateesin the western African region (PDF) (CMS Technical Series No. 26 ed.). Bonn, Germany: CMS Secretariat. pp. 26–30.
  11. ^ Weir, C. R (2019). "The Cetaceans (Whales and Dolphins) of Angola". In Huntley, B; Russo, V; Lages, F; Ferrand, N (eds.). Biodiversity of Angola. Cham: Springer. pp. 445–470. doi:10.1007/978-3-030-03083-4_16. ISBN 978-3-030-03082-7.
  12. ^ Findlay, K. P.; Best, P. B.; Ross, G. J. B.; Cockcroft, V. G. (1992). "The distribution of small odontocete cetaceans off the coasts of South Africa and Namibia". South African Journal of Marine Science. 12 (1): 237–270. doi:10.2989/02577619209504706. ISSN 0257-7615.
  13. ^ Payne, A. I. L; Brink, K. H; Mann, K. H; Hillborn, R (eds.). Benguela Trophic Functioning. South African Journal of Marine Science 12:237–270.
  14. ^ Leeney, Ruth (2014). "Towards sustainability of Marine Wildlife-Watching Tourism in Namibia". Journal of the Namibian Scientific Society. 62: 9–33.
  15. ^ Gopal, K; Karczmarski, L; Tolley, K.A (2019). "Patterns of geographic variation between mitochondrial and nuclear markers in Heaviside's (Benguela) dolphins (Cephalorhynchus heavisidii)". Integrative Zoology. 14 (5): 506–526. doi:10.1111/1749-4877.12380. hdl:2263/74315. PMID 30688009.
  16. ^ Hamner, Rebecca M.; Pichler, Franz B.; Heimeier, Dorothea; Constantine, Rochelle; Baker, C. Scott (2012). "Genetic differentiation and limited gene flow among fragmented populations of New Zealand endemic Hector's and Maui's dolphins". Conservation Genetics. 13 (4): 987–1002. doi:10.1007/s10592-012-0347-9. ISSN 1566-0621. S2CID 17218356.
  17. ^ a b c Elwen, Simon H.; Reeb, Desray; Thornton, Meredith; Best, Peter B. (2009). "A population estimate of Heaviside's dolphins,Cephalorhynchus heavisidii, at the southern end of their range". Marine Mammal Science. 25 (1): 107–124. doi:10.1111/j.1748-7692.2008.00246.x. hdl:2263/10207. ISSN 0824-0469.
  18. ^ Sekiguchi, K.; Klages, N. T. W.; Best, P. B. (1992). "Comparative analysis of the diets of smaller odontocete cetaceans along the coast of southern Africa". South African Journal of Marine Science. 12 (1): 843–861. doi:10.2989/02577619209504746. ISSN 0257-7615.
  19. ^ Elwen, SH; Snyman, L; Leeney, RH (2010). "Report of The Namibian Dolphin Project 2010: Ecology and Conservation of coastal dolphins in Namibia - Submitted to the Ministry of Fisheries and Marine Resources, Namibia" (PDF). {{cite journal}}: Cite journal requires |journal= (help)
  20. ^ Elwen, Simon H.; Best, Peter B.; Reeb, Desray; Thornton, Meredith (2009). "Diurnal Movements and Behaviour of Heaviside's Dolphins,Cephalorhynchus heavisidii, with some Comparative Data for Dusky Dolphins,Lagenothynchus obscutus". South African Journal of Wildlife Research. 39 (2): 143–154. doi:10.3957/056.039.0204. ISSN 0379-4369. S2CID 85572457.
  21. ^ Leeney, R. H; Carslake, D; Elwen, S. H (2011). "Using static acoustic monitoring to describe echolocation behaviour of Heaviside's dolphins (Cephalorhynchus heavisidii) in Namibia". Aquatic Mammals. 37 (2): 151–160. doi:10.1578/AM.37.2.2011.151. hdl:2263/17051.
  22. ^ Elwen, Simon; Meÿer, Michael A.; Best, Peter B.; Kotze, P. G H.; Thornton, Meredith; Swanson, Stephan (2006). "Range and Movements of Female Heaviside's Dolphins (Cephalorhynchus heavisidii), as Determined by Satellite-Linked Telemetry". Journal of Mammalogy. 87 (5): 866–877. doi:10.1644/05-MAMM-A-307R2.1. ISSN 0022-2372.
  23. ^ a b c Davis, RW; David, JHM; Meÿer, MA; Sekiguchi, K; Best, PB; Dassis, M; Rodríguez, DH (2014). "Home range and diving behaviour of Heaviside's dolphins monitored by satellite off the west coast of South Africa". African Journal of Marine Science. 36 (4): 455–466. doi:10.2989/1814232X.2014.973903. hdl:2263/43900. ISSN 1814-232X. S2CID 56296143.
  24. ^ Heinrich, Sonja; Elwen, Simon; Bräger, Stefan (2010). "Patterns of sympatry in Lagenorhynchus and Cephalorhynchus: dolphins in different habitats". In Würsig, Bernd; Wursig, Melany (eds.). The Dusky Dolphin: Master Acrobat Off Different Shores. Academic Press. pp. 313–332. ISBN 978-0-08-092035-1.
  25. ^ Elwen, SH; Leeney, RH; Gridley, T (2 January 2019). "Abundance estimates of an isolated population of common bottlenose dolphins Tursiops truncatus in Walvis Bay, Namibia, 2008–2012". African Journal of Marine Science. 41 (1): 61–70. doi:10.2989/1814232X.2019.1572538.
  26. ^ a b Martin, MJ; Gridley, T; Roux, J-P; Elwen, SH (2020). "First Abundance Estimates of Heaviside's (Cephalorhynchus heavisidii) and Dusky (Lagenorhynchus obscurus) Dolphins Off Namibia Using a Novel Visual and Acoustic Line Transect Survey". Frontiers in Marine Science. 7: 555–659. doi:10.3389/fmars.2020.555659.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  27. ^ Morisaka, Tadamichi; Karczmarski, Leszek; Akamatsu, Tomonari; Sakai, Mai; Dawson, Steve; Thornton, Meredith (2011). "Echolocation signals of Heaviside's dolphins (Cephalorhynchus heavisidii)". The Journal of the Acoustical Society of America. 129 (1): 449–457. doi:10.1121/1.3519401. hdl:10722/140937. ISSN 0001-4966. PMID 21303024.
  28. ^ Morisaka, T.; Connor, R. C. (2007). "Predation by killer whales (Orcinus orca) and the evolution of whistle loss and narrow-band high frequency clicks in odontocetes". Journal of Evolutionary Biology. 20 (4): 1439–1458. doi:10.1111/j.1420-9101.2007.01336.x. ISSN 1010-061X. PMID 17584238. S2CID 23902777.
  29. ^ Kyhn, L. A.; Jensen, F. H.; Beedholm, K.; Tougaard, J.; Hansen, M.; Madsen, P. T. (2010). "Echolocation in sympatric Peale's dolphins (Lagenorhynchus australis) and Commerson's dolphins (Cephalorhynchus commersonii) producing narrow-band high-frequency clicks". Journal of Experimental Biology. 213 (11): 1940–1949. doi:10.1242/jeb.042440. ISSN 0022-0949. PMID 20472781.
  30. ^ Martin, Morgan J.; Gridley, Tess; Elwen, Simon H.; Jensen, Frants H. (2018). "Heaviside's dolphins (Cephalorhynchus heavisidii) relax acoustic crypsis to increase communication range". Proceedings of the Royal Society B: Biological Sciences. 285 (1883): 20181178. doi:10.1098/rspb.2018.1178. ISSN 0962-8452. PMID 30051842.
  31. ^ Martin, Morgan J.; Elwen, Simon H.; Kassanjee, Reshma; Gridley, Tess (2019). "To buzz or burst-pulse? The functional role of Heaviside's dolphin, Cephalorhynchus heavisidii, rapidly pulsed signals". Animal Behaviour. 150: 273–284. doi:10.1016/j.anbehav.2019.01.007. ISSN 0003-3472. S2CID 72334526.
  32. ^ Best, P; Ros, G.J.B (1977). Exploitation of small cetaceans of the coast of Southern Africa. Report to the International Whaling Commission27:494-497
  33. ^ MacLeod, CD (2009). "Global climate change, range changes and potential implications for the conservation of marine cetaceans: a review and synthesis". Endangered Species Research. 7: 125–136. doi:10.3354/esr00197. ISSN 1863-5407.
  34. ^ Elwen, Simon H.; Leeney, Ruth H. (1 December 2010). "Injury and Subsequent Healing of a Propeller Strike Injury to a Heaviside's Dolphin (Cephalorhynchus heavisidii)". Aquatic Mammals. 36 (4): 382–387. doi:10.1578/AM.36.4.2010.382.
  35. ^ "Appendices I and II of the Convention on the Conservation of Migratory Species of Wild Animals (CMS)" (PDF). 5 March 2009. p. 6. Archived from the original (PDF) on 21 February 2012.
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