Haemulidae

Blue-striped grunts Haemulon sciurus, copyright Bernard Dupont.

Belongs within: Percomorpharia.

The sweetest of lips
Published 21 August 2014

In an earlier post on this site, I referred to a fish of the family Lethrinidae being known by the name of “sweetlips”. However, as is usually the way with fish vernacular names, there is more than one family of fishes to which this name can be applied. ‘Sweetlips’ is also the vernacular name for fishes in the Plectorhinchinae.

Oblique-banded sweetlips Plectorhinchus lineatus, copyright Richard Ling.

The Plectorhinchinae is most commonly treated as a subfamily within the family Haemulidae, the grunts (some sources will place ‘Plectorhynchidae’ as a separate family, and no, that wasn’t a typo: read on). Plectorhinchines are distinguished from other subfamily of haemulids, the Haemulinae, by characters including a longer dorsal fin and the presence of at least four prominent lateral line pores under the chin (Johnson 1980). The name ‘sweetlips’ refers to the prominent lips of mature individuals of two of the genera of plectorhinchines, Plectorhinchus and Diagramma, which are often a distinct colour from the rest of the head. Members of the third genus, Parapristipoma, have the lips not quite so prominent, and are commonly referred to as ‘grunts’ like the remaining haemulids.

African striped grunts Parapristipoma octolineatum, copyright Juan Cuetos.

The plectorhinchines are found around tropical reefs in the Indo-West Pacific and East Atlantic, with a single species, the rubberlip grunt Plectorhinchus mediterraneus, being found in the in the Mediterranean and Black Seas. No plectorhinchines are found on either side of the Americas. They are nocturnal predators of benthic invertebrates, emerging at night from the secluded crevices and overhangs where they spend the day. Most are medium-sized fish, though the painted sweetlips Diagramma pictum can get up to 90 cm. They are popular with fishers; Smith (1962) referred to them as “among the best if not the best eating fishes of the reef-haunting species”. Many species can go through significant changes in coloration as they mature: spotted juveniles may become unicoloured adults, or blotchy babies may mature into stripes. The differences are great enough that juveniles and adults have often been mistaken for separate species.

Juvenile oriental sweetlips Plectorhinchus vittatus, copyright Jan Messersmith. The adult form of this species resembles the oblique-banded sweetlips pictured above.

But failure to associate parents with their children is not the only way in which this group has been dogged by confusing taxonomy. The name of the type genus has been variously spelled Plectorhinchus or Plectorhynchus, with the family name varying accordingly (it seems that ‘Plectorhinchus‘ is the correct spelling). A surprising number of sources (e.g. Tavera et al. 2012) seem to have it both ways, with the genus being called Plectorhinchus but the higher taxon being called Plectorhynchinae (R. van der Laan et al. confirm the correct family-name spelling). Meanwhile, Smith (1962) argued for the use of the name Gaterin in place of Plectorhinchus, and called the family Gaterinidae. And if you have any interest in the vagaries of taxonomy, settle in: this is going to be a whole thing.

The name ‘Gaterin‘ dates from what is usually known as Forsskål’s (1775) Descriptiones animalium, which Fricke (2008) argued should be attributed to Niebuhr (see, right from the first sentence it’s confusing). Peter Simon Forsskål and Carsten Niebuhr were members of a Danish scientific expedition in 1761 to 1763 to the Red Sea (though Forsskål himself was Swedish, but that’s another story). Forsskål was the expedition’s naturalist, while Niebuhr was there as a geographer. The history of the expedition, and of the composition of Descriptiones animalium, has been summarised by Fricke (2008). The expedition was particularly ill-fated; of six original members, Niebuhr was the only one to make it back to Denmark alive. After returning to Denmark, Niebuhr started preparing Forsskål’s notes for publication. However, he found this no easy task. Forsskål had not prepared a single manuscript, but made notes on various scraps of paper; in the end, Niebuhr suspected that many of these scraps had gone missing. As an engineer, Niebuhr knew little Latin and even less biology, so he obtained the services of an academic adviser. The identity of this adviser was not divulged in the final publication by Niebuhr himself, but he has since been identified as the Danish naturalist Johann Christian Fabricius. The relationship between Niebuhr and Fabricius was not entirely positive (Niebuhr later stated that his adviser on Descriptiones animalium had been a ‘strange fellow’), and Fabricius does not seem to have spent any more time on the Forsskål notes than he absolutely had to. As a result, the final publication that emerged was partly Forsskål, partly Niebuhr, partly Fabricius, and all dog’s breakfast.

The name ‘Gaterin‘ is listed by Forsskål/Niebuhr/Fabricius as one of the sub-divisions of the genus Sciaena, and Smith’s (1962) revival of the name was based on the assumption that Forsskål intended these subdivisions to represent what we would now call subgenera. As such, Gaterin published in 1775 would clearly be an earlier name than Plectorhinchus published in 1802. Smith further supported this interpretation by pointing out that two names listed by ‘Forsskål’ as subdivisions of Chaetodon, Acanthurus and Abudefduf, had since been widely accepted as names for separate fish genera. There were no grounds, he claimed, for taking Abudefduf as valid but refusing Gaterin.

As it happens, Forsskål probably never intended either Gaterin or Abudefduf to represent generic names of any kind. It seems that his notes had used local Arabic names to refer to taxa to which he had not yet supplied formal Latin names. When Fabricius compiled these notes, he simply used the Arabic names as formal names, probably because he just didn’t care. When ‘Forsskål’ referred to ‘Gaterin’ in his introductory paragraph for Sciaena, he was probably referring to the individual species known in Arabia as gaterin rather than any formal group. ‘Abudefduf’ may have been similarly inadvertent, but long usage as a genus name means that it should probably be retained whatever its original status. No such argument can be marshalled in favour of ‘Gaterin’, whose usage in place of Plectorhinchus has been minimal.

And I can think of no better response to all that than the expression of this painted sweetlips Diagramma pictum. Copyright John Natoli.
Systematics of Haemulidae
<==Haemulidae [Gaterinidae, Plectorhynchidae, Pomadasyidae]
    |--OrthopristisND13
    |    |--O. chrysoptera (Linné 1766)D83
    |    |--O. ruber (Cuvier 1830)KC-L22
    |    `--O. scapularis Fowler 1916F16
    `--+--AnisotremusND13
       |    |--A. surinamensisND13
       |    `--A. virginicusF15
       `--HaemulonND13
            |  i. s.: H. albumB02
            |         H. elegansH04
            |         H. flavolineatum [incl. H. xanthopterum Günther 1859]F16
            |         H. macrostomaF15
            |         H. parraF15
            |--+--H. aurolineatumND13
            |  `--H. vittatumND13
            `--+--H. plumieriiND13
               `--H. sciurusND13
Haemulidae incertae sedis:
  BrachydeuterusEA03
    |--B. auritusEA03
    `--B. corvinaeformisF16
  PomadasysM58
    |--P. andamanensis McKay & Satapoomin 1994M01
    |--P. argenteus (Forsskål 1775)M01 [incl. Lutjanus hasta Bloch 1790M58, Po. hastaM01, Pristipoma hastaM58]
    |--P. argyreus (Valenciennes in Cuvier & Valenciennes 1833)M01 [=Pristipoma argyreumM58]
    |--P. auritus (Cuvier in Cuvier & Valenciennes 1830)M01
    |--P. furcatum (Bloch & Schneider 1801)M01 [=Pristipoma furcatumB80, Rhonciscus furcatusM01]
    |--P. jubeliniEA03
    |--P. kaakan (Cuvier in Cuvier & Valenciennes 1830)M01
    |--P. maculatus (Bloch 1793) [=Anthias maculatus, Pristipoma maculatum]M58
    |--P. nigerJR10
    |--P. opercularis Playfair in Playfair & Günther 1867M01
    |--P. perotaei (Cuvier 1830)LD09
    `--P. trifasciatus Fowler 1937M01
  Gaterin Forskål 1775 (see below for synonymy)S62
    |--*G. gaterinus (Forskal 1775) (see below for synonymy)S62
    |--G. ceylonensis Smith 1956S62
    |--G. chubbi (Regan 1919) [=Diagramma chubbi, Plectorhinchus chubbi, Pluchus chubbi]S62
    |--‘Sciaena’ faetala Forskal 1775 (n. d.) [=Diagramma faetala, Plectorhynchus faetala]S62
    |--G. gaterinoides Smith 1962S62
    |--G. giganteus (Gunther 1879) [=Diagramma giganteum]S62
    |--G. griseus (C & V 1830) (see below for synonymy)S62
    |--G. harrawayi Smith 1952 [=G. (Leitectus) harrawayi]S62
    |--G. macrolepis (Boulenger 1899)S62 [=Plectorhinchus macrolepisEA03]
    |--‘Diagramma’ microlepidotum Peters 1866S62
    |--G. nigrus (Cuvier & Valenciennes 1830)S62 (see below for synonymy)
    |--G. obscurus (Gunther 1871) [=Diagramma obscurum]S62
    |--G. papuensis (Macleay 1883) [=Diagramma papuense]S62
    |--G. paulavi (Steindachner 1895)MT73 [=Plectorhynchus paulayiS62; incl. G. baileyi Smith 1953S62]
    |--G. pictus (Tortonese 1935) (see below for synonymy)S63
    |--G. picus (C & V 1830) [=Diagramma pica; incl. D. picoides Peters 1866]S62
    |--G. plagiodesmus (Fowler 1935) (see below for synonymy)S62
    |--G. playfairi (Pellegrin 1914) (see below for synonymy)S62
    |--G. punctatissimus (Playfair 1867) [=Diagramma punctatissimum]S62
    |--G. radja (Bleeker 1853) [=Diagramma radja]S62
    |--G. rayi Menon & Talwar 1973MT73
    `--G. sordidus (Klunzinger 1870)S62 (see below for synonymy)
  Diagramma Oken 1817 [incl. Spilotichthys Fowler 1904]S62
    |--D. labiosum Macleay 1883M01
    |--D. pictum (Thunberg 1792)M01 (see below for synonymy)
    `--D. poecilopterum C & V 1830S62
  Parapristipoma trilineatum Thunb 1793 (see below for synonymy)S62
  Plectorhinchus Lacepede 1802M02, S62 [=Plectorhynchus Berthold 1827S62]
    |--P. albovittatus (Rüppell 1838)M01 (see below for synonymy)
    |--P. chaetodonoides Lacépède 1801M01 (see below for synonymy)
    |--P. chrysotaenia (Bleeker 1855)M01 (see below for synonymy)
    |--P. cinctus (Temminck & Schegel 1843)M01 [=Diagramma cinctumS62, Gaterin cinctusS62]
    |--P. flavomaculatus (Cuvier in Cuvier & Valenciennes 1830)M01 (see below for synonymy)
    |--P. gibbosus Lacepède 1802M01 (see below for synonymy)
    |--P. lessonii (Cuvier in Cuvier & Valenciennes 1830)M01
    |--P. lineatus (Linnaeus 1758)M01 (see below for synonymy)
    |--P. ordinalis (Scott 1962)M01 [=Gaterin ordinalisS62]
    |--P. pictus (Tortonese 1936)M01 (see below for synonymy)
    |--P. picus (Cuvier in Cuvier & Valenciennes 1830) [incl. P. punctatissimus (Playfair 1867)]M01
    |--P. polytaenia (Bleeker 1852)M01 (see below for synonymy)
    |--P. schotaf (Forsskål 1775)M01 (see below for synonymy)
    `--P. vittatus (Linnaeus 1758)M01 (see below for synonymy)
  ParakuhliaC01
  Isacia remensis (Leriche 1908)P93
  Pomadasyidarum gullentopsi Nolf 1978P93
  XenichthysR13
  XenocysR13
  XenistiusR13
  BoridiaR13
  Conodon nobilisR13, F15
Nomina nuda: Diagramma amabile Kent 1893S62
             Diagramma amicium Kent 1893S62

Diagramma pictum (Thunberg 1792)M01 [=Perca pictaS62, Plectorhinchus pictusS62, Spilotichthys pictusM58; incl. Diagramma balteatum C & V 1830S62, D. blochii Playfair 1866S62, D. centurio Cuvier 1830S62, D. cinerascens Ruppell 1828S62, Anthias diagramma Bloch 1792 non Perca diagramma Linnaeus 1758S62, *D. diagrammaS62, Plectorhinchus diagrammusS62, Plectorhynchus diagrammusS62, D. moaiab Thiollière 1857 (n. n.)S62, D. pertusum Playfair 1866S62, D. punctatum Rüppell 1830M01, D. radjabau Lacepede 1802S62, D. thunbergi C & V 1830S62]

Gaterin Forskal 1775 [incl. Euelatichthys Fowler 1904, Leitectus Smith 1952, Pluchus Smith 1949, Pseudopristipoma Sauvage 1880]S62

*Gaterin gaterinus (Forskal 1775) [=Sciaena gaterina, Diagramma gaterina, D. gaterinus, D. gatherina; incl. S. abumgaterin Forskal 1775]S62

Gaterin griseus (C & V 1830) [=Diagramma griseum, Plectorhinchus griseus, Plectorhynchus griseus; incl. Gaterin sivalingami Smith 1956]S62

Gaterin nigrus (Cuvier & Valenciennes 1830)S62 [=Euelatichthys nigerM58, Plectorhinchus nigerM58, Pristipoma nigrumM58, Pseudopristipoma nigrusS62; incl. Diagramma affine Gunther 1859S62, D. alta Day 1869S62, D. crassilabre Alleyne & MacLeay 1876S62, D. crassispinum Ruppell 1835S62, Euelatichthys crassispinusJR10, Plectorhynchus crassispinaM58, D. gibbosum Guichenot 1853S62, Pseudopristipoma leucurum C & V 1833S62, D. leucurumS62, Pristipoma leucurumS62]

Gaterin pictus (Tortonese 1935) [=Hapalogenys pictus; incl. Plectorhinchus cinctus punctatus Fang 1942 non Diagramma punctatum Ruppell 1828]S63

Gaterin plagiodesmus (Fowler 1935) [=Plectorhynchus plagiodesmus, Diagramma plagiodesmus, Pseudopristipoma plagiodesmus, Ps. plagiostomus (l. c.); incl. Ps. obscurus Fourmanoir 1957]S62

Gaterin playfairi (Pellegrin 1914) [=Diagramma griseum var. playfairi; incl. G. batata Smith 1952, D. batata]S62

Gaterin sordidus (Klunzinger 1870)S62 [=Diagramma sordidumS62, Plectorhinchus sordidusG75; incl. D. erythrostoma Bliss 1883S62, D. umbrinum Klunzinger 1870S62]

Parapristipoma trilineatum Thunb 1793 [incl. Plectorhinchus aporognathus Regan 1905, Diagramma japonicum Blkr 1852, P. ocyurus Jordan & Evermann 1903]S62

Plectorhinchus albovittatus (Rüppell 1838)M01 [=Diagramma albovittatumS62, Gaterin albovittatusS62, Plectorhynchus albovittatusS62; incl. Plectorhinchus giganteus (Günther 1879)M01, Plectorhinchus harrawayi (Smith 1952)M01, Plectorhinchus obscurum (Günther 1872)M01]

Plectorhinchus chaetodonoides Lacépède 1801M01 [=Gaterin chaetodonoidesS62; incl. Plectorhynchus doanei Seale 1910S62, Diagramma montrouzieri Thiollière 1857M58, D. pardalis Macleay 1884M58, Plectorhinchus plectorhynchus Shaw 1803S62]

Plectorhinchus chrysotaenia (Bleeker 1855)M01 [=Diagramma chrysotaeniaS62, Gaterin chrysotaeniaS62; incl. Plectorhinchus celebicus Bleeker 1873M01, Diagramma celebicumM58]

Plectorhinchus flavomaculatus (Cuvier in Cuvier & Valenciennes 1830)M01 [=Diagramma flavomaculatumS62, Gaterin flavomaculatusS62; incl. Gaterin citronellus Smith 1956S62, Diagramma jayakari Boulenger 1887S62, D. kumut Thiollière 1857S62, D. maculatus Fourmanoir 1957S62, D. ornatum Kossman & Rauber 1877S62, Plectorhinchus reticulatus Gunther 1859M01, Diagramma reticulatumS62, D. reticulatusS62, Gaterin reticulatusS62, D. roughleyi Whitley 1930S62, Plectorhinchus saidae Steindachner 1895S62]

Plectorhinchus gibbosus Lacepède 1802M01 [=Diagramma gibbosumS62; incl. Diagramma affine Gunther 1859S62, D. alta Day 1869S62, D. crassilabre Alleyne & MacLeay 1876S62, D. crassispinum Ruppell 1835S62, Euelatichthys crassispinusJR10, Plectorhynchus crassispinaM58, D. gibbosum Guichenot 1853S62, Pseudopristipoma leucurum C & V 1833S62, D. leucurumS62, Pristipoma leucurumS62, Euelatichthys niger Cuvier in Cuvier & Valenciennes 1830M58, M01, Gaterin nigrusS62, Plectorhinchus nigrusM01, Pristipoma nigrumM58, Pseudopristipoma nigrusS62]

Plectorhinchus lineatus (Linnaeus 1758)M01 [=Perca lineataS62, Diagramma lineatusS62, Gaterin lineatusS62, Plectorhinchus lineataS62, Plectorhynchus lineatusS62, Sciaena lineataS62; incl. Diagramma goldmanni Bleeker 1853M58, M01, Gaterin goldmanniS62, Plectorhinchus goldmanniM01, Plectorhynchus goldmanniS62, Diagramma haematochir Bleeker 1854S62, Plectorhynchus haematochirJR10, D. multivittatum MacLeay 1878S62, Plectorhinchus multivittatusH90, D. nitidum Gunther 1859S62, D. vavad Thiollière 1857M58]

Plectorhinchus pictus (Tortonese 1936) non Perca picta Thunberg 1792 (not preoc. if in dif. gen.)M01 [incl. Plectorhinchus fangi Whitley 1951M01, Gaterin fangiS63, P. sinensis Chu, Wu & Join 1977M01]

Plectorhinchus polytaenia (Bleeker 1852)M01 [=Diagramma polytaeniaM58, Gaterin polytaeniaS62; incl. D. ouan Thiollière 1857M58, D. polytaenioides Bleeker 1854S62]

Plectorhinchus schotaf (Forsskål 1775)M01 [=Diagramma schotafS62, Gaterin schotafS62, Plectorhynchus schotafS62, Sciaena schotafS62; incl. Diagramma durbanense Gilchrist & Thompson 1908S62, D. unicolor Macleay 1883S62]

Plectorhinchus vittatus (Linnaeus 1758)M01 [incl. Lutjanus aurantius Lacepede 1802S62, Bodian cuvier Bennett 1829S62, Diagramma cuvieriS62, Plectorhinchus cuvieriM58, Diagramma lessoni Cuvier & Valenciennes 1830S62, Anthias orientalis Bloch 1793M58, M01, Diagramma orientaleS62, Gaterin orientalisS62, Plectorhinchus orientalisM01, Plectorhynchus orientalisS62, Diagramma sebae Bleeker 1859M58, Plectorhynchus sebaeS62, D. sibbaldi Bennett 1832S62]

*Type species of generic name indicated

References

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[C01] Carpenter, K. E. 2001. Kuhliidae Flagtails. In: Carpenter, K. E., & V. H. Niem (eds) FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western Central Pacific vol. 5. Bony fishes part 3 (Menidae to Pomacentridae) pp. 3317–3320. Food and Agriculture Organization of the United Nations: Rome.

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[F16] Fowler, H. W. 1916. The fishes of Trinidad, Grenada, and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 520–548.

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[G75] Grant, E. M. 1975. Guide to Fishes. The Co-ordinator-General’s Department: Brisbane (Australia).

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[H90] Hutchins, J. B. 1990. Fish survey of South Passage, Shark Bay, Western Australia. In: Berry, P. F., S. D. Bradshaw & B. R. Wilson (eds) Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee pp. 263–278. Western Australian Museum.

Johnson, G. D. 1980. The limits and relationships of the Lutjanidae and associated families. Bulletin of the Scripps Institution of Oceanography 24: 1–114.

[JR10] Jordan, D. S., & R. E. Richardson. 1910. Check-list of the species of fishes known from the Philippine archipelago. Bureau of Printing: Manila.

[KC-L22] Kmentová, N., A. J. Cruz-Laufer, A. Pariselle, K. Smeets, T. Artois & M. P. M. Vanhove. 2022. Dactylogyridae 2022: a meta-analysis of phylogenetic studies and generic diagnoses of parasitic flatworms using published genetic and morphological data. International Journal for Parasitology 52: 427–457.

[LD09] Li, B., A. Dettaï, C. Cruaud, A. Couloux, M. Desoutter-Meniger & G. Lecointre. 2009. RNF213, a new nuclear marker for acanthomorph phylogeny. Molecular Phylogenetics and Evolution 50: 345–363.

[M01] McKay, R. J. 2001. Haemulidae (=Pomadasyidae). Grunts (also sweetlips, rubberlips, hotlips, and velvetchins). In: Carpenter, K. E., & V. H. Niem (eds) FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western Central Pacific vol. 5. Bony fishes part 3 (Menidae to Pomacentridae) pp. 2961–2989. Food and Agriculture Organization of the United Nations: Rome.

[MT73] Menon, A. G. K., & P. K. Talwar. 1973. On a new species of the genus Gaterin Forskål, 1775 (Pisces: Gaterinidae) from the Andaman Islands. Journal of the Bombay Natural History Society 69 (3): 658–660.

[M58] Munro, I. S. R. 1958. The fishes of the New Guinea region: A check-list of the fishes of New Guinea incorporating records of species collected by the Fisheries Survey Vessel “Fairwind” during the years 1948 to 1950. Papua and New Guinea Agricultural Journal 10 (4): 97–369 (reprinted: 1958. Territory of Papua and New Guinea Fisheries Bulletin no. 1).

[ND13] Near, T. J., A. Dornburg, R. I. Eytan, B. P. Keck, W. L. Smith, K. L. Kuhn, J. A. Moore, S. A. Price, F. T. Burbrink, M. Friedman & P. C. Wainwright. 2013. Phylogeny and tempo of diversification in the superradiation of spiny-rayed fishes. Proceedings of the National Academy of Sciences of the USA 110 (31): 12738–12743.

[P93] Patterson, C. 1993. Osteichthyes: Teleostei. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 621–656. Chapman & Hall: London.

[R13] Regan, C. T. 1913. The classification of the percoid fishes. Annals and Magazine of Natural History, series 8, 12: 111–145.

[S62] Smith, J. L. B. 1962. The fishes of the family Gaterinidae in the Red Sea and the Indian Ocean, with a resume of all Indo-Pacific species. Ichthyological Bulletin 25: 469–502.

[S63] Smith, J. L. B. 1963. The identity of Hapalogenys pictus Tortonese, 1935. Annals and Magazine of Natural History, series 13, 5: 637, pl. 13.

Tavera, J. J., A. Acero P., E. B. Balart & G. Bernardi. 2012. Molecular phylogeny of grunts (Teleostei, Haemulidae), with an emphasis on the ecology, evolution, and speciation history of New World species. BMC Evolutionary Biology 12: 57. http://www.biomedcentral.com/1471-2148/12/57.

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