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<document id="8FBB0D9B8F75690EED4568F073ED55E6" ENCODING="UTF-8" ID-GBIF-Dataset="16d12cf2-d1a3-4705-bc83-2611b7feebe8" ModsDocID="z01451p001" checkinTime="1247402697176" checkinUser="thomas" docAuthor="Brett A. Human" docDate="2007" docId="5CD1B824073C29C41B8D7BD0E2EC323A" docLanguage="en" docName="2007_Human_gg1.xml" docOrigin="Zootaxa 1451" docSource="http://www.zoobank.org/urn:lsid:zoobank.org:pub:1FEC783D-01D3-4458-9EE6-B499AF81A83F" docTitle="Haploblepharus Garman 1913" docType="treatment" docVersion="13" lastPageNumber="7" masterDocId="334F8DC5E1FDEBDBA56006F5B0238504" masterDocTitle="A taxonomic revision of the catshark genus Haploblepharus Garman 1913 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae)." masterLastPageNumber="40" masterPageNumber="1" pageNumber="4" updateTime="1701327483459" updateUser="ExternalLinkService">
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<mods:title id="FE1D0FE3366A61371924BE13269BB521">A taxonomic revision of the catshark genus Haploblepharus Garman 1913 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae).</mods:title>
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<mods:namePart id="FA78F6EB1E53651E288BF19A10470BE8">Brett A. Human</mods:namePart>
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<mods:date id="CBE6592ED7DA5D25E4E19CB6911A9D1B">2007</mods:date>
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<treatment id="5CD1B824073C29C41B8D7BD0E2EC323A" ID-DOI="http://doi.org/10.5281/zenodo.6248465" ID-GBIF-Taxon="100117388" ID-Zenodo-Dep="6248465" LSID="urn:lsid:plazi:treatment:5CD1B824073C29C41B8D7BD0E2EC323A" httpUri="http://treatment.plazi.org/id/5CD1B824073C29C41B8D7BD0E2EC323A" lastPageNumber="7" pageNumber="4">
<subSubSection id="C14404739864C096714EFC7EB8A7E53B" type="nomenclature">
<paragraph id="397C23CB52EABD0F28EDBB54FA4DD731" pageNumber="4">
<taxonomicName id="7BE1D4E27835CFFCC8A7BAB78DF23D17" ID-CoL="4TDF" ID-ENA="1003956" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="genus">Haploblepharus Garman 1913</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="D0D94C9D13A00385F985C0ECD56695F9" type="reference_group">
<paragraph id="7406B163CB8766DBD08FBCF2ACE30ECA" pageNumber="4">
<taxonomicName id="0AC150C566D9457D1FC45C35B8B6D057" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="genus">Haploblepharus Garman, 1913</taxonomicName>
: 101.
</paragraph>
</subSubSection>
<subSubSection id="E4C3CC750C15F71023BBAE10A6DAE443" type="nomenclature">
<paragraph id="FC02BB9B7EBD94C922FAF22A497F9E23" pageNumber="4">
Type species.
<taxonomicName id="E8C47D71E03E216FD403EE9CAF90B5A1" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyllium" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="species" species="edwardsii">Scyllium edwardsii Voigt 1832</taxonomicName>
(=
<taxonomicName id="3EE73A3FB94CF0EF9F196AFAA02A7EC0" LSID-ZBK="urn:lsid:zoobank.org:act:51BDDD26-53A8-415C-A2D4-122F4ADAAE68" class="Elasmobranchii" family="Squalidae" genus="Squalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Squalus edwardsii Schinz 1822:504" order="Squaliformes" pageNumber="4" phylum="Chordata" rank="species" species="edwardsii">Squalus edwardsii Schinz, 1822</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="09564FA550AE5C8110ADAD231DFEED2A" type="diagnosis">
<paragraph id="58BFA53694881066118914E8D7B74F98" pageNumber="4">
Diagnosis. See Garman (1913) and Compagno (1984b, 1988) for a comprehensive diagnosis of
<taxonomicName id="1A70A66E1B098039B9B1338FAA44F049" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
.
<taxonomicName id="AC6FDEFCFFA04ECED2ECFC1F9A240794" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
sharks are slender catsharks, with a long caudal peduncle and fin; distinguished from other southern African scyliorhinids by the expansion and fusion of the anterior nasal lobes into a flap that is united across the ventral midline, extends to the upper labium, and partially overlaps it in some cases, covering the excurrent nasal apertures; small separate symphyseal (basimandibular) cartilage present in lower jaw; sexual dimorphism not apparent. There is a high degree of morphological conservation between species within the genus. The sizes of
<taxonomicName id="E3C27654A194B6B1600DE985E899AD84" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="4" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
species, by sex, and for different maturity stages are shown in Figure 2.
</paragraph>
</subSubSection>
<subSubSection id="79F107A68919402ABDE9378F29D9B796" type="etymology">
<paragraph id="16B8931EF7DD51E333058BB8812FAF7A" pageNumber="5">
Common name. These sharks are normally referred to in the literature as shysharks (Afrikaans - skaamoog), due to their behaviour of covering their eyes with their tails when threatened, however this behaviour is common to
<taxonomicName id="6AE9857AE9F9691902F6AAC7680F4FF6" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
and
<taxonomicName id="118C12F58C6906655A751848A52134D7" class="Elasmobranchii" family="Scyliorhinidae" genus="Poroderma" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Poroderma</taxonomicName>
. Furthermore, the general public, including the angling and SCUBA diving communities, often refer to these sharks as doughnuts, again because of their behaviour when threatened. The term shyshark is used for all members of this genus and the term doughnut is used for both
<taxonomicName id="2C2862866DED9C15A925BCCF583DAAF8" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
and
<taxonomicName id="EF494675EF5E0B1F3E14F01A405CCD57" class="Elasmobranchii" family="Scyliorhinidae" genus="Poroderma" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Poroderma</taxonomicName>
in southern Africa (pers. obs.). Such common names have confounded catch data from both commercial and recreational fishermen, particularly among the latter. In an effort to give the members of this genus common names that will be easily remembered by the general public, Compagno &amp; Human (2003) introduced the name happy sharks, an allusion to their generic name.
</paragraph>
</subSubSection>
<subSubSection id="94367F3EB87B7BA53BE88400A07F5C48" type="discussion">
<paragraph id="64DAC5CE36289E0A39C4AD346887FBB6" pageNumber="5">
Remarks. The fusion of the anterior nasal flaps in
<taxonomicName id="D6E21A126722A19F25E1E6E18703C575" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
is unique to southern African scyliorhinids (not all scyliorhinids as stated by Bass et al., 1975). Elsewhere within the Scyliorhinidae, it occurs in the genera
<taxonomicName id="EB1112D87C276DE43ED92E1488F15CB1" LSID-ZBK="urn:lsid:zoobank.org:act:3328B736-21B0-4CE3-91AE-B13EA2536DA7" class="Elasmobranchii" family="Scyliorhinidae" genus="Atelomycterus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Atelomycterus Garman 1913:100" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Atelomycterus</taxonomicName>
Garman, 1913 (Indo-West Pacific), and
<taxonomicName id="45C52C0B108EEB5B38A831FFEC73EF69" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyliorhinus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Scyliorhinus</taxonomicName>
[in
<taxonomicName id="B5B15034005A68F0E262560CCB77D2FF" LSID-ZBK="urn:lsid:zoobank.org:act:79753A1D-1B94-4E9F-8B3E-B5642279DEF8" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyliorhinus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Scyliorhinus canicula albomaculata Pietschmann 1906:98" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="canicula">S. canicula (Linnaeus, 1758)</taxonomicName>
, Europe and Atlantic north Africa;
<taxonomicName id="1DCF1A6420BB6AF3E198A2C8C35AA8E3" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyliorhinus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="comoroenis">S. comoroenis Compagno, 1988</taxonomicName>
, tropical west Indian Ocean; and
<taxonomicName id="F77C8D5F600570AC5877029D3E713848" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyliorhinus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="garmani">S. garmani (Fowler, 1934)</taxonomicName>
, Indo-West Pacific]. Other sharks in southern African waters that have fused anterior nasal flaps are
<taxonomicName id="E1D23D2512207D7A8B966765849569F4" LSID-ZBK="urn:lsid:zoobank.org:act:6A36EC92-2E72-4403-B606-7B58F4E7569D" class="Elasmobranchii" family="Triakidae" genus="Scylliogaleus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Scylliogaleus quecketti Boulenger 1902:51" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="quecketti">Scylliogaleus quecketti Boulenger, 1902</taxonomicName>
(Triakidae), and
<taxonomicName id="F33527CF6E5F728DDC7DB5CD37991321" class="Elasmobranchii" family="Stegostomatidae" genus="Stegostoma" higherTaxonomySource="CoL" kingdom="Animalia" order="Orectolobiformes" pageNumber="5" phylum="Chordata" rank="species" species="fasciatum">Stegostoma fasciatum (Hermann, 1783)</taxonomicName>
(Stegostomatidae) (Compagno, 1984b, 1988, 2001; Compagno et al., 1989, 2005). Smith (1837) placed
<taxonomicName id="C20CBABB83DB8634ADDD939220B97C75" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
in the genus
<taxonomicName id="59B5C110293865B80678758673F8D207" class="Elasmobranchii" family="Scyliorhinidae" genus="Catulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Catulus</taxonomicName>
(attributed to Willoughby by Smith, 1837) along with
<taxonomicName id="90AD3894ACDDCED507C78A7AEE7CE983" class="Elasmobranchii" family="Scyliorhinidae" genus="Atelomycterus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="marmoratus">Atelomycterus marmoratus (Bennett, 1830)</taxonomicName>
and
<taxonomicName id="74EA49D84AA8ADBD8D4B5FEC74CF0B57" LSID-ZBK="urn:lsid:zoobank.org:act:79753A1D-1B94-4E9F-8B3E-B5642279DEF8" class="Elasmobranchii" family="Scyliorhinidae" genus="Scyliorhinus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Scyliorhinus canicula albomaculata Pietschmann 1906:98" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="canicula">Scyliorhinus canicula (Linnaeus, 1758)</taxonomicName>
, and may have based this grouping on the fused anterior nasal flaps. Garman (1913) described
<taxonomicName id="C366E0B7AC0E0CB789985D931623ABFA" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
as lacking posterior nasal valves, this is incorrect, as the excurrent nasal apertures are hidden by the fused anterior nasal flap and are obvious once this is lifted.
</paragraph>
<paragraph id="9FE8A226F7B4624FF60D74B9997F4248" pageNumber="5">
The presence of a small cartilage at the symphysis of the lower jaw (basimandibular) was first described and figured by Bass et al. (1975) and they suggest that the extra cartilage strengthens the jaw at the symphysis with the compromise of reduced jaw mobility. The presence of the basimandibular is an autapomorphic character of
<taxonomicName id="01F13C5F357A0B166A48A072F9E9A7EC" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
(Compagno, 1988; Human et al., 2006).
</paragraph>
<paragraph id="8BF14E580BFAFC00B04913977677B0E7" pageNumber="5">
<taxonomicName id="C19B63961C7DFA49EA4B31AA1B68E71F" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
sharks are generally common to abundant, and are regularly taken on hook and line by commercial and recreational anglers, and as bycatch in demersal trawls (Compagno, 1988; pers. obs.). As with other scyliorhinids,
<taxonomicName id="D27EC14A825EE5FB1F4FAF7AE91A9710" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
sharks are considered pests. Despite this, Springer (1979) states that this group of sharks are poorly known, and museum specimens outside of South Africa are uncommon.
</paragraph>
<paragraph id="6DCB01CE3AEAE3FC435D78A6FE079A83" pageNumber="5">
A factor that may have previously inhibited research on
<taxonomicName id="544CBB6B7AAE74BC107A36F768AB8E67" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
, is the difficulty in identifying these sharks to species, as they are morphologically conserved and most identification keys have used colour patterns to differentiate the species. As demonstrated in the current study, colour pattern is variable to highly variable in some of the
<taxonomicName id="9D94AF7B3538C46B454D9880866160AF" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
species, and the use of colour alone will inevitably lead to misidentification of some specimens. Another problem with identification keys currently available for
<taxonomicName id="E5623ED95C52C1CD1E7DB1A314376353" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
is the use of fin position (first dorsal fin relative to pelvic fin) for species identification, which also proves to be non-diagnostic. The author speculates that some hybridisation may also be occurring between the species, as some specimens are particularly difficult to classify, sharing morphological and/or colour patterns of different
<taxonomicName id="D63620AC9BA8C89A1FC217DD56B05159" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
taxa.
</paragraph>
<paragraph id="C61D987D95F0EC7C81AA926BFD7FF2F9" pageNumber="5">
The juveniles of most of the
<taxonomicName id="5AADA8A8E8723193D90D7DC6A8354DD1" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
species are elusive, with poor representation in collections. The author speculates that this could be due to a period of rapid growth from time of hatching to late adolescence. If juveniles are collected, it should be considered a priority to collect a genetic sample before preserving the specimen, as there is still considerable confusion as to the identity of some juvenile
<taxonomicName id="693BFE207A1B3F9ACD3F3EEC30E18E33" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
specimens. The collection of genetic material may also shed light on the ontogenetic segregation apparent in some species (see Distribution, below). Genetic samples are also needed for adult
<taxonomicName id="883B9284DAEF5C9AF1EB638321A5167C" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="species" species="kistnasamyi">H. kistnasamyi</taxonomicName>
. A genetic study of this genus at the species and population level is needed to resolve these issues.
</paragraph>
<paragraph id="06E3418BB8FDBC3119AC4BCFFA42618C" pageNumber="5">
<taxonomicName id="3FF1CEC1AFC1706DB6537B762A93A54A" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="5" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
sharks occurring in the Eastern Cape and kwaZulu-Natal require a great deal more research in order to determine their basic life history parameters.
</paragraph>
<paragraph id="7CB1189CF6FC893E1BE2EE22E6FC623C" pageNumber="6">
As determined by the current study,
<taxonomicName id="DD3675E3B5FE2F79BB400CF13BD54D37" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
contains four valid taxa. These are
<taxonomicName id="FABE7A640068C86FA99A21B504C0401E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii (Schinz, 1822)</taxonomicName>
,
<taxonomicName id="3957A00BC15C6F19097B6514ACF61EBC" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="pictus">
H. pictus (
<normalizedToken id="FDAC913E1A274FD92A9D046FDD89DCAC" originalValue="Müller">Mueller</normalizedToken>
&amp; Henle, 1838)
</taxonomicName>
,
<taxonomicName id="3C0E1195F958B3CC7564BCA67E9A1153" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus Smith, 1950</taxonomicName>
, and
<taxonomicName id="15AEC1E9EF57C8B37714E6ACB07FC6F8" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="kistnasamyi">H. kistnasamyi Human &amp; Compagno, 2006</taxonomicName>
. Due to the tentative placement of some juvenile specimens in nearly all of the species groups, the species identification key (below) is based on type material and mature specimens (maturity stage 4 or 5).
</paragraph>
</subSubSection>
<subSubSection id="790A3CAD8561726FA3BC12EEA1E871E7" type="etymology">
<paragraph id="FA9247A69C03B310A2CC909367E53CA9" pageNumber="6">
Etymology. Although not specified by Garman (1913), it is assumed that
<taxonomicName id="5A7C17EC9603767A3B6F7D8D3CF71655" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
is derived from the Greek words haplos, meaning single or simple, and blepharon, meaning eyelid, and refers to the simple structure of the lower eyelid that can close against the upper eyelid to protect the eye.
</paragraph>
</subSubSection>
<subSubSection id="6CCFE54B54C2CE9B35843C78CC012FF1" type="distribution">
<paragraph id="58A049EF4B0FED38C4606D596C38D5E6" pageNumber="6">
Distribution.
<taxonomicName id="63892B652B6A684E5034C62A7306D28A" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
species are rare to abundant, and are present in southern Namibia and South Africa. They also demonstrate habitat preferences, and
<taxonomicName id="BE4729F6F6C79602B330683800DDC6C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
apparently exploits different niches either side of Cape Agulhas, by occurring at different depths, possibly to avoid competition with other
<taxonomicName id="079FD34D70B50D2D3F598F59EE0125AF" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
in the Eastern Cape.
</paragraph>
<paragraph id="7A6115793E079777220BB7F6BC46C074" pageNumber="6">
<taxonomicName id="FBB4AF4132EB2202FA3E98E7B2BF0F13" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">Haploblepharus edwardsii</taxonomicName>
is abundant close inshore in the Western Cape, certainly west of Cape Agulhas, whereas it appears to prefer deeper water east of Cape Agulhas (see species account below). Smith (1950) commented that he had not known
<taxonomicName id="7E879AC5957179B7B66F8E238868A35A" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
to be caught from the shore (in reference to the Eastern Cape), whereas they are easily observed when diving from the shore in the Western Cape (pers. obs.). The deeper habit of
<taxonomicName id="713AAF82598FE0150F49514742B972E2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
in the Eastern Cape has also been noted by Bass et al. (1975), and Compagno (1984b, 1988). This may serve to avoid competition with
<taxonomicName id="2A83F8969B91C5A033F91CEC2D39E527" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
and
<taxonomicName id="8A7A7C6BDE0536E37D3F97D60EFDD2B3" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="pictus">H. pictus</taxonomicName>
in that area (Compagno 1984b, 1988; current study), however,
<taxonomicName id="44E072ADD010C9023C2B24F4031CE259" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
is not unknown to occur in inshore waters of the Eastern Cape (Buxton et al., 1984; Wallace et al., 1984; current study). The deepest record for
<taxonomicName id="899D258FA14112E0F9937535991BF189" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
verified here is 111 m, and Bass et al. (1975) record this species from 130 m.
<taxonomicName id="006CFB7027D6FFDFBB3B782F224774B4" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">Haploblepharus edwardsii</taxonomicName>
is apparently segregated by depth from
<taxonomicName id="4CBC65CC5E6D892A23AF08A05109FCC8" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
and
<taxonomicName id="8EC8B281FE37A2A081D8F1BB8F2D4EBC" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="pictus">H. pictus</taxonomicName>
in Eastern Cape waters, with
<taxonomicName id="96E3FC6B3A936846253C2A4DB015BA32" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
and
<taxonomicName id="BFF0B855E320D9548BF962BAE3ED9C2D" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="pictus">H. pictus</taxonomicName>
occurring at shallower depths than
<taxonomicName id="A61E9B39714AADD83F7042C310C83ADD" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
, which may serve to reduce the amount of competition between the species there (Compagno, 1984b, 1988; current study).
</paragraph>
<paragraph id="2EC4D6D6C63D13E574A4F42DF6B7677A" pageNumber="6">
An interesting phenomenon that was brought out in the current study was the difficulty encountered in collecting juvenile and adolescent
<taxonomicName id="9684E99E9E5C6160C552D734C3D29335" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
females from False Bay and Table Bay (also noted by Dainty, 2002; A. Bertolini, pers. comm.). Dainty (2002) had noted the difficulty in obtaining any female specimens. Sex ratios, particularly of juveniles and adolescents, are highly biased towards males, therefore it appears that mature females come to False Bay and Table Bay from elsewhere, this requires further investigation.
</paragraph>
<paragraph id="7A947EF3FF22075596B67DD0FD374440" pageNumber="6">
The juveniles of
<taxonomicName id="12411CBB247553F0030A4AD8E1F817EB" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
are scarce (Bass et al., 1975; M. J. Smale, pers. comm.; current study), and there appears to be an unknown habitat that is used by
<taxonomicName id="0DA79B43B279CFF95AB251EEAD02BEC8" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
for egg laying and where juveniles spend that stage of their life history.
</paragraph>
<paragraph id="CE9CA48030C944ED922755475AF79FF8" pageNumber="6">
If the specimens tentatively assigned here as juvenile
<taxonomicName id="F0CAB799142CB8AE3188FE40D8FB107A" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="kistnasamyi">H. kistnasamyi</taxonomicName>
prove to be accurate (see species account below), then there is also ontogenetic segregation of that species with juveniles occurring in the southern part of their range, and adults occurring in the northern part of their range.
</paragraph>
</subSubSection>
<subSubSection id="56066DCBDBBEEBBE9AECADD1C1744B95" type="key">
<paragraph id="2BD5FDA0068A2718BC5F81C70BC1995A" pageNumber="6">
Key to the species of the genus
<taxonomicName id="9F23397CE699669282976BDC53E35032" LSID-ZBK="urn:lsid:zoobank.org:act:87028B01-5D5B-4004-918B-492D3059F90E" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus Garman 1913:101" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="genus">Haploblepharus</taxonomicName>
(based on type material and mature specimens only)
</paragraph>
<paragraph id="3336867696762CDDA15F77A29F199E83" pageNumber="6">1a. Snout acutely rounded, often coming to a point in adults; body slender and compressed to moderately depressed in adults; less than 70 tooth rows total in upper jaw of adults....................................................2</paragraph>
<paragraph id="F0CF889496258E1186AE9C3AC2724970" pageNumber="6">1b. Snout broadly rounded, not coming to a point in adults; body stocky and moderately to greatly depressed in adults; more than 75 tooth rows total in upper jaw of adults..................................................................3</paragraph>
<paragraph id="E27E8146D48E3DD78D7CB6ECD054CD9F" pageNumber="6">
2a. Body moderately depressed; head width at pectoral fin origin 1.31 times or greater than head height at pectoral fin origin; abdomen width 0.95 times or greater than abdomen height.......................
<taxonomicName id="93309D4C2A5EAE1AC5458B8531F38F21" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="edwardsii">H. edwardsii</taxonomicName>
</paragraph>
<paragraph id="83E9AEDE8B588B58E340E79551BE0563" pageNumber="6">
2b. Body weakly depressed to weakly compressed; head width at pectoral fin origin 1.29 times or less than head height at pectoral fin origin; abdomen width 0.90 times or less than abdomen height.............
<taxonomicName id="CDBA1132731D237D324D4578FBC3103B" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="6" phylum="Chordata" rank="species" species="kistnasamyi">H. kistnasamyi</taxonomicName>
</paragraph>
<paragraph id="64E5B24F5269845D96938C3AD904E305" pageNumber="7">
3a. Saddles and/or spotting always present, saddles and spots usually occurring together, saddles and spots obvious when present; 84 or more tooth rows total in upper jaw ....................................................
<taxonomicName id="B23B803E01BF5EB4F9D9C94AAABC32D5" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" order="Carcharhiniformes" pageNumber="7" phylum="Chordata" rank="species" species="pictus">H. pictus</taxonomicName>
</paragraph>
<paragraph id="77A4B6837E90FACAB58901032AA194BC" pageNumber="7">
3b. Saddles or spotting usually not present, saddles and spots rarely occurring together, saddles and spots inconspicuous when present; 83 or less tooth rows total in upper jaw ..........................................
<taxonomicName id="34E6C8F277CFCB9DCF6E7308B5143922" LSID-ZBK="urn:lsid:zoobank.org:act:5B016902-D9E6-47E1-9F91-E23E5AA226C2" class="Elasmobranchii" family="Scyliorhinidae" genus="Haploblepharus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Haploblepharus fuscus Smith 1950:883" order="Carcharhiniformes" pageNumber="7" phylum="Chordata" rank="species" species="fuscus">H. fuscus</taxonomicName>
</paragraph>
</subSubSection>
</treatment>
</document>
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