Charles Darwin University The biology and ecology of tropical marine sardines and herrings in Indo-West Pacific fisheries a review Hunnam, Kimberley Published in: Reviews in Fish Biology and Fisheries DOI: 10.1007/s11160-021-09649-9 10.1007/s11160-021-09649-9 Published: 10/04/2021 Document Version Publisher's PDF, also known as Version of record Link to publication Citation for published version (APA): Hunnam, K. (2021). The biology and ecology of tropical marine sardines and herrings in Indo-West Pacific fisheries: a review. Reviews in Fish Biology and Fisheries, 1-36. https://doi.org/10.1007/s11160-021-09649-9, https://doi.org/10.1007/s11160-021-09649-9 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 06. Oct. 2021 Rev Fish Biol Fisheries https://doi.org/10.1007/s11160-021-09649-9 (0123456789().,-volV)( 0123456789().,-volV) REVIEWS The biology and ecology of tropical marine sardines and herrings in Indo-West Pacific fisheries: a review Kimberley Hunnam Received: 24 August 2020 / Accepted: 23 February 2021 Ó The Author(s) 2021 Abstract Tropical sardines and herrings (Family associations and population dynamics. Most research Clupeidae) form important fisheries resources in the on tropical clupeid biology and ecology in the Indo- Indo-West Pacific region. However knowledge of their West Pacific is concentrated on the Round-bodied biology and ecology is largely scattered across diverse Sardinellas (S. longiceps and S. lemuru) from a few localised studies. In addition, their challenging taxon- major fishery locations; other species-groups have omy due to numerous, morphologically similar been studied sporadically across their distributions. species, coupled with data collected from dispersed, Characteristics such as maximum size, life span, size at often small-scale and mixed species fisheries, creates first maturity and some habitat associations are gener- difficulties for comparing species-level research and ally similar within species-groups and differ between fisheries statistics across the region. This review groups. However differences within the same reported considers four main groups of tropical marine clupeid species highlight the importance of continued work to species important for Indo-West Pacific fisheries— improve taxonomic identification. There remain sub- Sardinella (subg. Sardinella) species (‘Round-bodied stantial gaps in knowledge on all species-groups. Sardinellas’), Sardinella (subg. Clupeonia) species Regular critical review of specific research topics, such (‘Flat-bodied Sardinellas’), Amblygaster species as reproductive parameters and population dynamics, (‘Tropical Pilchards’), and Herklotsichthys and Es- would assist to standardise methods and terminology cualosa species (‘Tropical Herrings’)—and collates used, enable consolidation and comparison of findings, and synthesises existing knowledge on early life identify local research agenda, and help build and history, reproduction, trophic biology, habitat improve research on these important tropical species. Keywords Clupeidae Á Sardinella Á Amblygaster Á Supplementary Information The online version contains Herklotsichthys Á Escualosa Á Indo-West Pacific Á supplementary material available at https://doi.org/10.1007/ Fisheries s11160-021-09649-9. K. Hunnam (&) Research Institute for the Environment and Livelihoods, Charles Darwin University, Darwin, NT, Australia Introduction e-mail: [email protected] Clupeoid fishes—sardines, herrings, anchovies and K. Hunnam Research School of Biology, The Australian National their relatives (order Clupeiformes, suborder University, Canberra, ACT, Australia 123 Rev Fish Biol Fisheries Clupeoidei)—form the basis of some of the world’s Fig. 1 a Reported clupeoid (order Clupeiformes, suborder c largest fisheries in both temperate and tropical regions Clupeoidei) marine capture production by family in the IWP (FAO fishing areas 51, 57 and 71) from 1950 to 2017, and (FAO 2018). While most research efforts have focused contribution to global marine clupeoid landings and total IWP on the large single species stocks in productive marine landings. Data from FAO (2019a) with reported temperate and upwelling regions (e.g. Checkley ‘Clupeidae’ reclassified to exclude families Dussumieriidae et al. 2017; Ganias 2014), the importance of these (round herrings, Dussumieria spp.) and Spratelloididae (sprats, Spratelloides spp.) as per recent taxonomic revisions (references fisheries in tropical areas has steadily increased. Over in text). b Average annual IWP clupeoid landings from 2008 to 2 million tonnes of tropical and subtropical sardines 2017 by family (left); and within the Clupeidae family (marine from the Sardinella genus are now caught annually, species only) based on the species-groups used in this review making this group the fourth largest contributor to (right). Data from FAO (2019a) with some disaggregation of ‘Clupeoidei (not specified)’ and ‘Sardinella spp.’ landings global marine capture production in 2016 (FAO 2018). based on species compositions reported in national statistics for Clupeoid production in the largest tropical region, Philippines (BFAR 2008–2017), India (CMFRI 2008–2017), the Indo-West Pacific (IWP),1 has steadily increased Malaysia (DOF 2008–2017) and Australia (Mobsby and over the past 60 years (Fig. 1a). In the last decade Koduah 2017). ‘Round-bodied Sardinellas’ comprise S. longi- ceps (dark blue) and S. lemuru (light blue). ‘Flat-bodied (2008–2017) this region accounted for around half Sardinellas’ comprise species reported as S. gibbosa (dark Sardinella production and 13% of marine clupeoid green); ‘other sardines’, i.e. non-S. longiceps, reported to production globally. In contrast, production in tem- include S. gibbosa, S. fimbriata, S. albella, S. dayi, S. sindensis perate and upwelling areas has remained fairly con- and S. melanura, as well as Amblygaster spp. (Bennet et al. 1992) (mid-green); and predominantly ‘S. fimbriata’, recently stant albeit with high annual variability (FAO 2018; redescribed in the Philippines as S. pacifica (Hata and Moto- FAO 2019a). Tropical clupeoids are caught by both mura 2019b) (light green). ‘Tropical Pilchards’ comprise small-scale and larger-scale fisheries, which provide separately reported A. sirm. ‘Tropical Herrings’ comprise livelihoods for hundreds of thousands of fishers and separately reported Herklotsichthys quadrimaculatus. Landings of Escualosa spp. and marine Gizzard Shads (Nematalosa and post-harvest workers (e.g. FAO 2018; Kripa et al. Anodontostoma spp.) are not separately reported. ‘Sardinops 2018; Rola et al. 2018). Many tropical clupeoid (temperate)’ comprises landings of temperate species Sardinops fisheries are predominantly for human consumption, sagax from cooler parts of FAO fishing areas 57 (i.e. south providing a relatively cheap and affordable source of Australia) and 51 (i.e. South Africa) and are not considered further in this review. For further details, refer to Online protein and micronutrients (e.g. Roeger et al. 2016; Resource 1 Rola et al. 2018). Furthermore, the population of the IWP region is projected to increase around 29% over the next 30 years to reach over 3.8 billion people by numerous, particularly on the most abundantly caught 2050 (United Nations 2019).2 Ensuring the sustain- Sardinella species. However most are localised stud- ability of these fisheries into the future is therefore of ies with few recent attempts to compare and/or critical importance. synthesise existing knowledge across the region. Knowledge of fish biology and ecology is part of Previous reviews of tropical clupeoid biology and the basic information required for fisheries manage- ecology are useful but dated (Longhurst 1971); ment (Sadovy de Mitcheson 2009). Research efforts predominantly focussed on the smaller, short-lived on the biology and ecology of tropical clupeoids are sprats and anchovies (Dalzell 1993); were limited to comparison of a few species on a particular aspect of biology (Milton et al. 1994b); or mainly considered 1 The IWP encompasses the region from the south-east tropical only fisheries landings fluctuations and management coast of Africa, east along the southern continental margin of the (Kripa et al. 2019). Similarly, tropical species have India-Eurasia landmass, to the tropical South China Sea, received little attention in recent reviews on general Hawaii, Micronesia, Polynesia and north-eastern Australia (as clupeoid biology and ecology (e.g. Ganias 2014). per Lavoue´ et al. 2013). For analysis of fisheries production statistics, FAO fishing areas 51, 57 and 71 have been used as per The IWP region has the highest diversity of clupeoid Whitehead (1985), which are all considered tropical areas (FAO species (Lavoue´ et al. 2014; Whitehead 1985) with 2018). representatives from all six currently recognised