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Agnatha, and

Chapter · November 2016

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Antonis Petrou Charitos Zapitis AP Marine Env.Consultancy Ltd & Enalia Physis Environmental Research Centre University of Derby

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CHAPTER 26: AGNATHA, CHONDRICHTHYES AND OSTEICHTHYES

FISHES

Antonis Petrou and Charitos Zapitis

1. INTRODUCTION The classification of is not straightforward since they do not form a natural scientific grouping like the other classes, i.e. the , reptiles, birds and . Indeed, fishes can be considered by exclusion to be that are not (see Chapter 25: Introduction to Vertebrates). They are aquatic, - bearing, ectothermic ('cold-blooded') with a distinguishable head and, when present, digit-less limbs. Traditionally, have been arranged into three groups:

➵ Agnatha 1, the jawless fish (Myxini [] and [lampreys]); ➵ Chondrichthyes, the cartilaginous fish (, skates and rays); ➵ Osteichthyes, the bony fish ( [ray-finned fishes] and [lobe-finned fishes]).

This classification is adequate for general purposes, although Agnatha is paraphyletic and includes several groups of extinct jawless fishes.

2. THE EVOLUTION OF FISHES The first proto-fish appeared in the mid- period around 530 MYA. The best known was gracilens (named after Pike Peak in Canada), a -like crea - ture around 5 cm long, with a slender worm-like body and large prominent head. Although technically not vertebrates, they had a primitive , so were 'chor - dates', had the basic vertebrate body plan, and were at the root of the vertebrate line - age. During the period, 490 - 430 MYA, the first indisputable fish appeared.

2.1. Agnatha Unlike modern fishes, they were jawless and toothless, and were traditionally named agnathans (Greek: 'no '). The most primitive lacked and many were covered with thick -like plates, which provided some protection against arthro - pod predators such as the giant sea scorpion. The agnathans flourished and by the end of the period (445 - 420 MYA) reached the peak of their evolution. During the (420 - 360 MYA), competition from and predation by the sea scorpions (some of which grew up to 2 m in length) and the newly emerging jawed fish, resulted in a decline in agnathans from which they never recovered. There are just two orders of extant jawless fish as shown in Fig 1: Myxiniformes () with around 76 and Petromyzontiformes (lampreys) with Epinephelus marginatus around 38 species. These two lineages had already diverged before the (Dusky ). 1The extant agnathans (the hagfish and lampreys), have been placed together in the grouping Antonis Petrou (Greek: 'round-mouthed') (see Fig 1), the taxonomical status of which is contentious.

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622 The Wildlife of Cyprus Chapter 26 - Agnatha

Ray-finned fishes are today by far the largest and most diverse vertebrate group with around 30,000 named species, representing around 50% of all the vertebrate Species Common name Sources of data species. They became predominant during the (360 MYA). They are Anguilla anguilla European Zogaris et al ., 2012; Segurado et al ., 2014 divided into two subclasses: the more primitive (Greek: 'cartilaginous Dicentrarchus labrax European Seabass Zogaris et al ., 2012; Segurado et al ., 2014 bone') fishes have cartilaginous with some ossification and comprise around cephalus Flathead Grey Zogaris et al ., 2012; Segurado et al ., 2014 40 extant species of , , and reedfish. All other species are in Liza aurata 1 Zogaris et al ., 2012; Segurado et al ., 2014 the (Greek: 'new fins'). Neopterygii are further divided into two infraclasses: Liza ramada Grey Mullet Zogaris et al ., 2012; Segurado et al ., 2014 the primitive (Greek: 'whole/all bone') fishes with lightly ossified cartilaginous Table 1: List of native fish species in Cyprus. skeletons, comprising just eight extant species of the and and the Teleostei 1Only one specimen was collected and identified from Xeros river in 2012 (Zogaris et al ., 2012). (Greek: teleos = 'complete/perfect', osteon = 'bone'). Teleostei are distinguished by their bony skeletons, their homocercal tail fins and the bones of the upper jaw, which are completely moveable allowing the jaw to be protruded outwards when the mouth is opened. Although fishes, with around 30,000 species, now account for around 95% of all fish species, they started their spectacular radiation only during the late Cretaceous (144 - 65 MYA). Today, lobe-finned fishes are represented by just eight species: two species of and six species of . However, with the first dating back around 420 MY, they were more diverse than the ray-finned fishes during the Devonian and Carboniferous Periods. Around 390 MYA the sarcopterygians split into two major lineages: the and the Dipnotetrapodomorpha (also known as rhipidis - tians). The coelacanths (Greek: koilos = 'hollow'; akantha = 'spine') never left the sea and the peak of their success was during the late Devonian and Carboniferous from around 385 to 300 MYA. They were thought to have disappeared during the K-Pg 65.5 MYA, but in 1938 a specimen was discovered in the waters around South Africa and in 1997 a second specimen was discovered in an Indonesian fish-market. In contrast, the rhipidistians, the ancestors of which probably lived in Fig 2: Anguilla anguilla (European eel) photographed in the pool at the Baths of Aphrodite coastal waters, moved into freshwater where they split into two further in February 2015. Francis Plowman subgroups: the Dipnoi (Greek: dis = 'double; pnoe = 'breathing') (lungfish) and the east Atlantic. During that process, the larvae undergo the transformation of the Tetrapodomorphs (proto-tetrapods).The Dipnoi are represented today by just six different stages of their life-cycle. This is the only freshwater eel species found in Cyprus extant species of lungfish, the best known of which is the Queensland Lungfish. As and is distributed around the island (found in 14 sites among 170 surveyed (Zogaris, described in Chapter 27, the tetrapodomorphs were pivotal in evolving a second verte - 2014)). brate body plan that would give rise to the amphibians and later the reptiles, birds in Cyprus face severe man-made threats, especially from the damming of and mammals. rivers, which blocks off migration routes and destroys many aquatic habitats below the dams; this can result in mass deaths among the eel population. Since 2010, their 3. THE FISHES OF CYPRUS global conservation status has been considered as ‘Critically Endangered’ according 3.1. to the IUCN. 3.1.1. Native species The Mediterranean Toothcarp or Killifish ( Aphanius fasciatus ), is a fish that has been Only five species of native fish inhabit freshwater ecosystems in Cyprus. Although recently discovered in Akrotiri lake. This species is distributed in all countries of the seabass and mullet may primarily be thought of as sea-going species, they also enter Mediterranean region with the exception of the Iberian Peninsula, and its conservation inland brackish and fresh waters, such as , rivers and lakes. status is described as of ‘Least Concern’. Its reintroduction to new sites has been The European eel, Anguilla anguilla , is a catadromous fish (i.e. it spends most of its discussed as a conservation measure to assist in the protection of this species on the life in , then migrates to the sea to breed) widespread in the north-east island (Zogaris, 2014). There is an interest in ecological restoration and A. fasciatus Atlantic and in the . Adults migrate to the Sargasso Sea in could be used instead of alien species for the biological control of mosquitoes the western Atlantic to , and the larvae are transported via the currents back to (Zogaris, 2014).

624 625 The Wildlife of Cyprus Chapter 26 - Agnatha

3.1.2. Introduced species The vast majority (20 species) of freshwater fishes in Cyprus have been introduced. Table 2 gives a list of the introduced freshwater species along with known information about the origin and time of introduction.

3.1.3. Freshwater Fish Farms Only a limited number of freshwater fish are farmed in Cyprus, with rainbow trout, Oncorhynchus mykiss , being most common. During the last few , attention has been drawn to the sturgeon ( baerii ) which, according to the Department of and Marine Research, is currently farmed in three units. Furthermore, the Department runs an unit in Kalopanagiotis community in Nicosia district. The research station keeps a brood stock of the following species: Anguilla anguilla (European Eel), Micropterus salmoides (Largemouth Bass), Ictalurus punctatus (Channel Fig 3: A school of goldfish, photographed in an agricultural tank at Farmakas, April 2015. Eddie John ), Cyprinus carpio (Common ), Carassius auratus (Goldfish), rutilus Species Common name of Comments Sources of data (Common Roach) and Tinca tinca (Tench). introduction Abramis bjoerkna Silver Bream 1972 Probably UK stock Zogaris et al ., 2012; Segurado et al ., 2014 3.2. Marine Fishes of Cyprus Alburnus alburnus Bleak 1972 Probably UK stock Zogaris et al ., 2012; Segurado et al ., 2014 Although marine species are not the focus of this book, we include for complete - ness (and since many of them appear in fishmongers’ shops) a short section on the Aphanius Mediterranean ? Recently discovered Zogaris et al ., 2012; Segurado et al ., 2014 fasciatus Killifish in Akrotiri fish present in waters around Cyprus. Aspius aspius 1 Asp/Rapfen ? Anon, 1998 The Mediterranean is the world's largest enclosed sea with a surface area of 2.5 million km 2. It is connected to the (via the 14 km-wide Strait of Atherina boyeri Big-scale Sand Smelt ? Marine transient Zogaris et al ., 2012; Segurado et al ., 2014 Gibraltar), to the (via the straits of Bosphorus and Dardanelles) and to the Carassius auratus Goldfish ? Zogaris et al ., 2012; Segurado et al ., 2014 Red Sea (via the Suez Canal). The Mediterranean contains around 10 - 12,000 species Carassius carassius 1 Crucian Carp ? Anon, 1998 of fish, 20 - 30% of which are endemic, so it is rich in biodiversity, but it is very low in Carassius cf. gibellio Carp ? Zogaris et al ., 2012; Segurado et al ., 2014 biomass. The reasons for this are: Cyprinus carpio Common Carp 1960s Israeli stock Zogaris et al ., 2012; Segurado et al ., 2014 Gambusia holbrooki Eastern Mosquitofish 1939 Syrian stock Zogaris et al ., 2012; Segurado et al ., 2014 ➵ The oligotrophic (i.e. low in nutrients) waters around Cyprus determine which fish Ictalurus punctatus Channel Catfish 1975 American stock Zogaris et al ., 2012; Segurado et al ., 2014 species are present. Small pelagic (i.e. open sea) fishes are absent from the coastal Lepomis gibbosus Pond Perch recently Zogaris et al ., 2012; Segurado et al ., 2014 waters, because the lack of nutrient-rich inflows of freshwater results in low abun - Micropterus salmoides Largemouth Bass 1971 Probably UK stock Zogaris et al ., 2012; Segurado et al ., 2014 dance of phytoplankton, which is the primary food source of these species. Hence Oncorhynchus mykiss Rainbow Trout 1969 USA stock Zogaris et al ., 2012; Segurado et al ., 2014 the and fisheries present in both Greece and Turkey are absent Oreochromis aureus Blue Tilapia 1976 Israeli stock Zogaris et al ., 2012; Segurado et al ., 2014 from Cyprus. Perca fluviatilis European Perch 1971 UK stock Zogaris et al ., 2012; Segurado et al ., 2014 ➵ The fish caught commercially in Cyprus are mainly benthic (i.e. bottom dwellers) Rutilus rutilus Common Roach 1972 probably UK stock Zogaris et al ., 2012; Segurado et al ., 2014 and benthopelagic species, as well as a few migratory pelagic species. The main methods used in Cyprus are set and gill nets, bottom and floating longlines, Salmo trutta Brown Trout 1948 UK stock Zogaris et al ., 2012; Segurado et al ., 2014 traps, troll and jigged lines. Two bottom trawlers also fish in Cyprus waters. Like Stizostedion Pikeperch / ? Anon, 1998 lucioperca most benthic fisheries in the Mediterranean, Cyprus has a multispecies . This means that rarely is one species targeted, such as in the case of albacore . Tinca tinca 2 Tench ? Anon, 1998 Professional fishing brings in a wide variety of fish species, which makes the Table 2: Introduced freshwater species along with known information about the origin and time management of fisheries much more complex and difficult. of introduction. 1 Considered by Zogaris et al . (2012) as unconfirmed. 2 Observed at Kalopanagiotis Hatchery in 2011 (Zogaris et al. , 2012). The main species, both commercial and non-commercial, present in Cyprus waters are given in Table 3.

626 627 Chapter 26 - Agnatha Scientific Name Common Name (English) Scientific Name (cont.) Common Name (cont.) Anthias anthias Swallowtail Seaperch Pagellus bogaraveo The fish on sale in Cyprus fish markets is a mix of both local and imported species. Apogon imberbis Cardinal Fish Pagellus erythrinus This is due to low local production and high demand. Various types of fish can be found lacunosus Hardyhead Silverside Pagrus pagrus Common Seabream around the year, complemented by local seasonal catches. Local consumers have capriscus Grey Parexocoetus mento African Sailfin Flyingfish adapted their taste to include various Lessepsian migrants (see 3.2.1) such as siganids Belone belone /Garpike vanicolensis Vanikoro and Fistularia commersonii , whilst some fish species such as albacore tuna, although Blennius ocellaris Butterfly Blenny Phycis phycis Forkbeard Boops boops Bogue/Bream Polyprion americanus Wreckfish caught in large quantities, are not favoured by local consumers and are exported. The Brama brama Atlantic Pomfret Pseudocaranx White Trevally largest fish stock appears to be that of the Picarel ( Spicara smaris ). This, however, poses Caranx crysos Blue Runner clavata /Skate a distinct problem to fishermen in that, since various EU measures have been in place, Centracanthus cirrus Curled Picarel Ruvettus pretiosus Oilfish the healthiest available stock cannot be fished. macrophthalma Red sarda Atlantic Bonito In an attempt to enhance fish stocks, whilst at the same time assisting the recre - Chlorophthalmus agassizi Shortnose Greeneye Sardina pilchardus ational diving industry on the island, the Department of Fisheries and Marine Research Damselfish Sardinella aurita Conger conger Conger Eel Sargocentron rubrum Red Soldier Fish/Redcoat has embarked on a programme of sinking artificial reefs, including ships. These artificial Coris julis Rainbow Sarpa salpa Cow Bream /Salpa/Salema Porgy structures add hard substrate in unproductive areas and thus promote the increase Coryphaena hippurus Common Dolphinfish umbra Brown Meagre of biodiversity, the protection and growth of fisheries and the development of under - Dactylopterus volitans Flying Gurnard Scomber colias Atlantic Chub water attractions. pastinaca Common Scomber scombrus Atlantic Mackerel Marine Aquaculture is highly developed in Cyprus and plays an important role in Dentex dentex Striped Bream/Common Dentex commerson Narrow-barred Dentex gibbosus Pink Dentex Scorpaena scrofa Large-scaled Scorpionfish the fish trade on the island. Marine fish hatcheries and offshore cage facilities are found Dicentrarchus labrax European Seabass canicula Lesser Spotted Dogfish on the south coast. The main marine finfish species produced are Dicentrarchus labrax Annular Seabream dumerili Greater (Seabass) and Sparus aurata (Seabream). Species such as Argyrosomus regius (Meagre), Diplodus cervinus Zebra Seabream cabrilla Comber Siganus luridus (Dusky Spinefoot/Rabbitfish), Seriola dumerili (Greater Amberjack) and Diplodus puntazzo Sharpsnout Seabream Serranus scriba Painted Comber Pagrus japonicas (Japanese Seabream) have also been produced for aquaculture Diplodus sargus White Seabream Siganus luridus Dusky Spinefoot purposes in the past but are not reared on a large scale. Two-banded Seabream Siganus rivulatus Marbled Spinefoot Epinephelus aeneus White Grouper Solea solea Common Sole The whole process from to table-size fish is undertaken locally, and great effort Epinephelus caninus Dogtooth Grouper Sparisoma cretense Parrotfish is made to avoid the use of hormones, antibiotics or other pharmaceutical products. Epinephelus costae Goldblotch Grouper Sparus aurata Gilt-head Seabream Additionally, the isolation of the island from the European continent and areas where Epinephelus marginatus Dusky Grouper Sphyraena sphyraena European Barracuda wild populations of these species are found, make the likelihood of any diseases very Etrumeus teres Round Spicara maena Blotched Picarel remote. Hence, the quality of the two main species of farmed fish (Seabass and Fistularia commersonii Bluespotted Cornetfish Spicara smaris Picarel cemiculus Blackchin Spondyliosoma cantharus Black Seabream Seabream) is very high and has even been recognized with international awards. They niger Squalus acanthias Piked Dogfish/ also constitute a prized export product, with 70% of the fish produced being exported Helicolenus dactylopterus Blackbelly Rosefish Symphodus tinca Peacock Wrasse to major cities in Europe, North America and the Middle East. Silver-cheeked Toadfish Synodus saurus Bluestriped Lizardfish Lichia Leerfish Tetrapturus belone Mediterranean Spearfish 3.2.1. Alien invasive species Lithognathus mormyrus Striped Seabream alalunga Albacore Due to the close proximity of the Suez Canal, the offshore waters of Cyprus are Lophius budegassa Black-bellied Angler Thunnus thynnus Atlantic Bluefin Tuna Macroramphosus scolopax Longspine Spinefish marmorata Marbled Torpedo Ray impacted by an increased presence of species known as Lessepsian Migrants (named European Hake Trachinotus ovatus Derbio after Count Ferdinand de Lesseps, the French diplomat who oversaw the canal's Mugil cephalus Flathead Mullet draco Greater construction). These species are of Indo-Pacific origin and have passed from the Red barbatus Trachurus sp . Horse Mackerel Sea to the Mediterranean through the Suez Canal. The acclimatization of most of these Mullus surmuletus Trigloporus lastoviza Streaked Gurnard species to the warmer, more saline waters of the eastern Mediterranean has occurred helena Black/ Umbrina cirrosa Shi Drum Mycteroperca rubra Mottled Grouper Upeneus moluccensis Goldband in recent years at a much faster rate than previously noted. This may be partly due to aquila Common Uranoscopus scaber Stargazer increased scientific interest in the subject, but the effect of climate change cannot be Naucrates Pilotfish Xiphias gladius completely excluded. Non-native fish species arriving from the Red Sea (or other Oblada melanura Saddled Seabream Xyrichtys novacula Pearly Razorfish sources) are also known as alien invasive species. For the most part, they are thought Axillary Seabream Zeus faber John Dory to affect the ecosystem negatively, by either competing with or preying upon native Table 3: The main fish species, both commercial and non-commercial, present in Cyprus waters. Mediterranean fish species and thus affecting the food chain. (List adapted from Department of Fisheries and Marine Research – 100 most important fish Species of Cyprus Waters.) 629 The Wildlife of Cyprus Chapter 26 - Agnatha

Species such as Lagocephalus sceleratus (Puffer Fish/Silver-cheeked Toadfish) and Pterois miles (Common Lionfish), negatively affect native fish species by preying on them. Species such as Siganus rivulatus (Marbled Spinefoot) which has been in the Mediterranean since the 1940s, compete for food resources with species like Sarpa salpa (locally known as Salema Porgy, a species of seabream often served in Mediterranean restaurants, and which can cause hallucinations when eaten). The effect of alien species on commercial fisheries varies: the pufferfish, for example, being highly toxic is of no commercial value, but also negatively influences the catch and causes damage to nets; the spinefoot, on the other hand, is highly commercial.

ACKNOWLEDGEMENTS The authors extend their thanks to Eddie John and Francis Plowman for the provi - sion of photographs.

REFERENCES

Anonymous. 1998. Fishes of the Cyprus Reservoirs : Angler's Guide. Ministry of Agriculture, Natural Re - sources and Environment, Fisheries Department. Press and Information Office, Republic of Cyprus. 18 pp. Gess, R. W., Michael I., Coates, M. I. & Rubidge, B.S. 2006. A from the Devonian period of South Africa. 443 : 981-984. Segurado, P., Zogaris, S., Chatzinikolaou, Y. & Ferreira, M. T. 2014. Fish as indicators of ecological status in Mediterranean island streams: Insights from Cyprus. Jornadas Ibéricas de Ictiologia, pre - sented at Conhecer para preservar, Lisbon. Shimeld, S. M. & Donoghue P. C. J. 2012. Evolutionary crossroads in developmental : cyclostomes (lamprey and hagfish). Development 139 (12): 2091-9. Zogaris, S. 2014. Ichthyological Study for Oroklini Lake. April 2014. BirdLife Cyprus. A Study within the Project LIFE10 NAT/CY/716 ‘Restoration and Management of Oroklini Lake SPA, Larnaca’. Unpublished Final report, 58 pp. Nicosia, Cyprus. Zogaris, S., Chatzinikolaou, Y., Koutsikos, N., Oikonomou, E., Giakoumi, S., Economou, A. N., Vardakas, L., Segurado, P. & Ferreira, M. T. 2012. Assessment of fish assemblages in Cyprus Rivers for the implementation of Directive 2000/60/EC. Specialized Consultancy Services for the Assessment of Fish Assemblages in Cyprus Rivers – Implementation of the Directive 2000/60/EC. Final Report of Second Phase of the Project. Hellenic Center for Marine Research - Institute of Marine Biological Resources and Inland Waters / Instituto Superior de Agronomia, Universidade Técnica de Lisboa. 205 pp.

Fig 4: A scorpion fish (top of page) . The exact Fig 5: Red Soldier Fish ( Sargocentron rubrum ), species is difficult to determine from the picture a common alien species. but is possibly Scorpaena scrofa. Antonis Petrou Antonis Petrou

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