PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 65, No. 1, pp. 1-39, 25 figs., Appendix 
September 28, 2018 
A Synopsis of the Eastern and Central Atlantic Combers of the 
Genus Serranus (Teleostei: Scorpaeniformes: Serranidae) 
Tomio Iwamoto 1 and Peter Wirtz 2 
1 Department of Ichthyology, California Academy of Sciences, 55 Music Concourse Drive, 
San Francisco, CA 94118 U.S.A. e-mail', tiwamoto@calacademy.org. 
2 Centro de Ciencias do Mar, Campus de Gambelas, PT 8005-139 Faro, Portugal. 
Combers comprise a group of mostly small, shallow-water, tropical to subtropical 
reef fishes of the genus Serranus, subfamily Serraninae. Only a few of the 32 
currently known species attain a size exceeding 25 cm TL and few live at depths 
exceeding 100 m. Three-quarters of the species are found in the Atlantic, with many 
fewer in the eastern Pacific (6 spp.) and western Indian Ocean (2 spp.). The highest 
diversity (14 spp.) is in the western Atlantic, with the eastern Atlantic second (10 
spp.). No species is found in more than one of these four areas. This synopsis provides 
a supplement to the treatment of the group by Heemstra and Anderson (2016) in 
FAO Species Identification Guide for Fishery Purpose. We include three additional 
species: S. pulcher Wirtz and Iwamoto, 2016, and two new species herein described: 
S. drewesi Iwamoto, and S. inexpectatus Wirtz and Iwamoto. Serranus drewesi, 
known only from the holotype taken by spear in Sao Tome, appears most similar to 
S. hepatus but is distinguishable by it cycloid scales, naked interorbital space, 
pigmentation pattern on the fins and body, gill-raker count of the first arch, and 
several other characteristics. The type specimens for Serranus inexpectatus are from 
Angola and Gabon, but we suspect that it occurs elsewhere off West Africa. The type 
specimen from Gabon is that reported by Poll (1954) as Paracentropristis heterurus; 
the excellent illustration in Poll’s work has been used erroneously by others as 
representative of S. heterurus. DNA evidence suggests that S. inexpectatus is sister to 
a clade that includes S. heterurus and S. pulcher. The three are closely similar but 
each can be distinguished by a combination of pigmentation pattern and meristic 
and morphometric characteristics. Descriptions, photographs, and a key to the 
species are provided for the eastern Atlantic species 
The genus Serranus Cuvier, 1817 has long been a catch-basket for a variety of small to medi¬ 
um-sized fishes in the subfamily Serraninae, family Serranidae. They are frequently referred to as 
Combers or Dwarf sea basses. Most species are colorful and easily kept in aquariums, thus their 
popularity among aquarists and divers. Some species are important components of artisanal fish¬ 
eries. Nelson (2006:346) listed 11 genera as examples of the subfamily, although Meisler (1987), 
in a comprehensive unpublished revision of the serranines, provided a classification, based on a 
cladistic analysis, that included only 10 genera, excluding two of the genera Nelson listed and 
adding Mentiperca Gill, 1862. Robins and Starck (1961) in their revision of western Atlantic 
Serranus recognized only two subgenera, Serranus and Paracentropristis Klunzinger, 1884, but 
Meisler (1987) recognized four subgenera: Serranus, Paracentropristis, Dules Cuvier, 1829, and 
Prionodes Jenyns, 1842. 
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