Biology and Ecology of Groupers

Fabiana Cézar Félix-Hackradt, Carlos Werner Hackradt, José Antonio García-Charton, Fabiana Cézar Félix-Hackradt, Carlos Werner Hackradt, José Antonio García-Charton

• Publisher: CRC Press
• Year: 2022
• ISBN:
  9781351645157
• Edition: 1
• Topic: Biological Sciences
• Subtopic: Zoology
• Index: Biological Sciences

Groupers are fascinating charismatic fishes commonly found in reef habitats around the world that sustain a global multimillionaire fishery. They are animportant top predator species and therefore of paramount ecological importance for reef systems' resilience. The book, Biology and Ecology of Groupers is an up-to-date review of the main bio-ecological topics involving worldwide groupers species oriented to the academic community and managers. It covers aspects of classification and phylogenetic relationships, geographical distribution, and life history related characteristics (Section I), including the major threats of groupers populations, case studies of successful management and comments about the future of groupers in our changing world (Section II).

Section 1 Biology and Ecology of Groupers

Chapter 1.1 Classification of Groupers

Ka Yan Ma^1,# and Matthew T. Craig^2,*

¹ Simon F. S. Li Marine Science Laboratory, School of Life Sciences, The Chinese University of Hong Kong, Hong Kong SAR

² National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Southwest Fisheries Science Center, 8901 La Jolla Shores Drive, La Jolla, CA 92037, USA

^# Current affiliation: School of Ecology, Sun Yat-sen University, Shenzhen, China.

* Corresponding author: [email protected]

Introduction

Biological classification aims to define groups of organisms. As the broader field of systematics has evolved, classification has developed into a system that is based on the evolutionary relationships among organisms. It is generally accepted that a natural classification is one that groups together all the descendants of a common ancestor into a taxon, a condition that is termed “monophyletic” (Hennig 1965). Organisms in a taxon share a common genetic inheritance, which may be expressed in a variety of observable and measurable characters that taxonomists use to develop classification schemes. Traditionally, characters used in fish classification are morphological, such as lateral-line scale count, fin shape and position, and various morphometric ratios. However, similar morphological characters may evolve independently in distantly related lineages due to convergent evolution, and classification based on these analogous characters would create unnatural, non-monophyletic taxonomic groups. Additionally, selection plays a large role in shaping the morphology of an organism, which further complicates evolutionary histories derived from these data. Molecular data have emerged as valuable tools that, along with morphology, can increase our understanding of the evolutionary relationships among lineages, thus enhancing our ability to create robust classification systems. In some cases, these data are thought to be relatively free of strong selective pressures and may not experience such frequent instances of convergent evolution.

Groupers (Epinephelinae sensu Craig and Hastings (2007)) are a species-rich group of marine reef fish whose classification has undergone many changes and remains controversial across various levels of the taxonomic hierarchy from family to species. Historically, their classification was based on morphology. In the past decade or so, a number of molecular phylogenetic studies, using a variety of markers and covering many taxonomic levels, have contributed significantly to clarifying the evolutionary relationships among groupers and creating a more coherent classification (e.g., Maggio et al. 2005, Ding et al. 2006, Craig and Hastings 2007, Craig et al. 2011, Zhuang et al. 2013, Schoelinck et al. 2014, Ma et al. 2016, Qu et al. 2017, Ma and Craig 2018). This chapter discusses the chronological development of grouper classification from the morphology-era to the molecular-era, from family to genera, and to species. We discuss the most comprehensive grouper phylogeny developed to date (Ma and Craig 2018) and its implications for taxonomy. By consolidating both morphological and molecular information, we present a revised grouper classification scheme that reflects evolutionary relationships of this diverse group of marine fish.

The Family for Groupers: Serranidae or Epinephelidae

Groupers have long been recognized as part of the large and diverse family Serranidae, which has traditionally been used as a convenient pigeon-hole for lower percoid species with ambiguous affinity and has received extensive systematic treatments (Fig. 1). In the first attempt to coherently organise the Serranidae, Jordan and Eigenmann (1890) defined six subfamilies: Anthiinae (the anthias), Epinephelinae (the groupers), Grammistinae (the soapfishes and podges), Latinae (the late perches), Percichthyinae (the temperate perches), and Serraninae (the basses) (Fig. 1a). Jordan (1923) elevated the groupers to family level (Epinephelidae), thus restricting the Serranidae to the subfamilies Anthiinae and Serraninae, and treated the monotypic genus Niphon as its own family, Niphonidae (Fig. 1b). The first cladistically-based classification of the family was by Gosline (1966), in which Serranidae was restricted to Jordan and Eigenmann’s (1890) Anthiinae, Epinephelinae, and Serraninae (Fig. 1c). Kendall (1976,1979) added to Serranidae the subfamily Grammistinae (treated as family Grammistidae by Gosline (1966)) based on the similar number and orientation of predorsal bones to those of the Epinephelinae (sensu Jordan and Eigenmann (1890)) (Fig. 1d). Later, Johnson (1983) agreed with Gosline’s (1966) three-subfamily scheme, but also followed Kendall (1976, 1979) in placing members of Grammistinae into the family Serranidae (Fig. 1e). Johnson (1983) defined a monophyletic subfamily Epinephelinae based on the absence of an autogenous distal radial on the first dorsal-fin pterygiophore, and in doing so he included Kendall’s Grammistinae as well as the enigmatic species Niphon spinosus as part of Epinephelinae. Johnson (1983, 1988) further divided his Epinephelinae into five tribes: Diploprionini (soapfishes), Gramminstini (soapfishes), Liopropromini (basslets), Niphonini (Niphon spinosus), and Epinephelini (groupers) (Fig. 1e). Baldwin and Johnson (1993) cladistically demonstrated the monophyly of Johnson’s Epinephelinae and analyzed the relationships among its tribes: Niphon was hypothesized to be most distantly related to all other epinehelines, followed by Epinephelini, and then Diploprionini, whereas Grammistini and Liopropromini, were thought to be closely related sister groups (Fig. 1f). By contrast, Heemstra and Randall (1993) proposed an alternative classification which restricted Epinephelinae to Johnson’s (1983, 1988) Epinephelini, elevated the genus Niphon to subfamily Niphoninae, and united Diploprionini, Gramministini, and Liopropromini to subfamily Grammistinae (Fig. 1g). The classification schemes proposed by Johnson (1983, 1988), Baldwin and Johnson (1993), and Heemstra and Randall (1993), though substantially different, are all used today, causing considerable confusion (Fi g. 1).

Fig. 1 Classification schemes of the family Serranidae propo...