153 ANNOTATED Checklist of the World's Marine Lobsters
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THE RAFFLES BULLETIN OF ZOOLOGY 2010 Supplement No. 23: 153–181 Date of Publication: 31 Oct.2010 © National University of Singapore ANNOTATED CHECKLIST OF THE WORLD’s MARINE LOBSTERS (CRUSTACEA: DECAPODA: ASTACIDEA, GLYPHEIDEA, ACHELATA, POLYCHELIDA) Tin-Yam Chan Institute of Marine Biology, National Taiwan Ocean University, Keelung 20224, Taiwan, R.O.C. Email:[email protected] ABSTRACT. – Marine lobsters are defined as consisting of four infraorders of decapod crustaceans: Astacidea, Glypheidea, Achelata and Polychelida. Together they form the suborder Macrura Reptantia. A checklist of the currently recognized six families, 55 genera and 248 species (with four subspecies) of living marine lobsters is provided, together with their synonyms in recent literature and information on the type locality of the valid taxa. Notes on alternative taxonomies and justifications for the choice of taxonomy are given. Although Caroli Linnaeus himself described the first marine lobster in 1758, the discovery rate of marine lobsters remains high to this day. KEY WORDS. – Crustacea, Decapoda, lobsters, marine, checklist, taxonomy. INTRODUCTION al., 2005). These results also showed that the relationships of the superfamilies and families of marine lobsters are mostly Commercialy, lobsters are generally the most highly prized different from the previously well-established scheme of crustaceans in all parts of the world. The taxonomy of marine Holthuis (1991). However, these phylogenetic studies have lobsters has remained fairly stable over many years and some yielded significantly contrasting results (see Patek et al., 2006, recent authors, such as Burukovsky (1983) and Phillips et al. Tsang et al., 2008; Bracken et al., 2009; Toon et al., 2009). (1980) had complied lists of all the valid species of extant The latest and by far the most robust phylogenetic analysis marine lobsters known in the world, at that time. In the (Tsang et al., 2008) utilises newly developed molecular monumental work of Holthuis (1991), a detailed account was markers, concluding that lobsters are indeed a monophyletic provided for almost all the living marine lobster species up to group, if the thalassinideans (or some of the thalassinideans) 1991. The traditional concept of marine lobsters includes the are excluded. Whilst Scholtz & Richter (1995) argued that superfamilies Nephropoidea (clawed lobsters), Palinuroidea thalassinideans (as Thalassinida) may be more closely (spiny and slipper lobsters), Eryonoidea (blind lobsters) and allied to the freshwater crayfishes than the clawed lobsters the living fossil Glypheoidea contained within the suborder (homarids), recent work has clearly demonstrated a sister Macrura Reptantia. The above three works did not provide relationship between the clawed lobsters and the freshwater any listing for Polychelidae (the only living family of blind crayfishes (Crandall et al., 2000; Porter et al., 2005; Tsang lobsters) because of the extremely confusing taxonomy of this et al., 2008; Toon et al., 2009). Thus, the suborder Macrura group. Furthermore, the reef lobsters genus Enoplometopus Reptantia is revived containing the lobsters and freshwater was not considered to be lobsters by Phillips et al. (1980). crayfishes (see Tsang et al., 2008). Thalassinidea (or now Holthuis (1983) considered Enoplometopus to be axiids Axiidea and Gebiidea, see De Grave et al., 2009) is excluded (Infraorder Thalassinidea or now Axiidea, see De Grave et from this suborder, partly because they are traditionally not al., 2009) rather than nephropoids. Holthuis’s (1991) world considered as true lobsters (e.g. the squat lobsters Galatheidae marine lobster catalog included Thalassinidea among the and Chirostylidae are always not considered as true lobsters), lobsters but no account was given to Enoplometopus. and partly because the most robust phylogeny study (Tsang et al., 2008) strongly suggested that only some clades of Recent advances in morphological and molecular phylogeny thalassinideans are allied to the “traditional” lobsters (i.e. studies have impacted fundamentally on our understanding Thalassinidea is a polyphyletic group, also see Robles et of the evolutionary relationships of marine lobsters and other al., 2009). Decapoda. Most recent analyses suggest that marine lobsters do not comprise a monophyletic group (Scholtz & Richter, This work presents an updated list of all currently considered 1995; Schram, 2001; Dixon et al., 2003; Amati et al., 2004; valid species of living marine lobsters known in the world. Schram & Dixon, 2004; Ahyong & O’Meally, 2004; Porter et The diversity of marine lobsters is not high as compared 153 Chan: Marine lobster checklist to other decapod crustacean groups (e.g. 3,268 species in the type lost and original description inadequate to determine caridean shrimps and 6,559 species in crabs, see De Grave its identity, no taxonomic resolution at present is possible et al., 2009). Although the catalog of marine lobsters of (see Holthuis, 1991). These issues are discussed in Holthuis the world by Holthuis (1991) had encompassed almost all (1983, 1991, 2002). The present work makes no attempt to the species known at that time, there have been many new settle them and the most widely used names are adopted. discoveries in the last two decades. Moreover, Galil (2000) made a worldwide revision of Polychelidae and clarified The present list recognizes 248 valid species (with four most of the taxonomic problems of this difficult group. The valid subspecies) of marine lobsters in 6 families and 55 higher classification used here follows the results of the latest genera. Marine lobsters were first described by Linnaeus phylogenetic analyses of lobsters (Chan & Ng, 2008; Tsang in 1758. There was a gradual increase in species only after et al., 2008, 2009; Palero et al., 2009; Bracken et al., 2009; 1800 (Fig. 1). At the end of the nineteen century during Tshudy et al., 2009; Ahyong, 2009) instead of that presented the age of great exploration, the lobsters also had a rapid in Martin & Davis (2001). Such a scheme has been presented increase in their discovery rate similar to that of many other in the recently published classification of living and fossil decapod groups (Dworschak, 2000; Yeo et. al., 2008; De genera of decapod crustaceans in De Grave et al. (2009), Grave et. al., 2008; Baba et al., 2008). The discovery rate except here the suborder Macrura Reptantia is recognized. significantly slowed down between the Two World Wars but Reasons for the present classification scheme are given under after 1950 quickly returned to its pre-war rate. Surprisingly, the remarks in the corresponding higher taxon. for a group of generally large sized animals with high economic value, the number of new species discovered in The spelling of author’s names follows Ng et al. (2008) marine lobsters since then has remained high, even very for the world brachyuran crabs list. An “*” refers to the recently. For example, nearly 11.7% (29 species) of marine type species of the genus. As nearly all groups of marine lobsters were only described in the last decade (i.e. since lobsters have been recently revised, and full synonymies of 2000). From the still very steep discovery curve shown in most species can be found for Arctides (Holthuis, 2006), Fig. 1, no extrapolation for total number of marine lobster Indo-Pacific Scyllarinae (Holthuis, 2002), Polychelidae seems possible. Even to the most common and commercially (Galil, 2000), Enoplometopus (Holthuis, 1983; Poupin, 2003) important genera such as Palinurus and Panulirus, new and the rest of the marine lobsters (Holthuis, 1991), only species have been added in the last few years (Sekiguchi synonyms still used in taxonomic literature after 1960 are & George, 2005; Groeneveld et al., 2006). The discovery given. Taxonomic decisions for synonymy after the major curve of marine lobsters is very similar to that of the squat works of Holthuis (1983, 1991, 2002, 2006) and Galil (2000) lobsters (see Baba et al., 2008). However, squat lobsters are are explained in the remarks of the corresponding higher generally small and of no commercial importance. Recent taxon. If the original name given for a taxon is different employment of molecular tools in separating cryptic and from its current generic allocation and/or usage (or recent very similar species has contributed to the discovery of usage for synonyms), the original name is provided at the more lobster species as in other decapod crustaceans under end of the name in square brackets. Synonyms with spelling this modern trend. Nevertheless, the high discovery rate as or similar to the original name are not repeated in the of lobsters is no doubt more related to the revived large synonymy. For example in the species Stereomastis nana scale expeditions in the Indo-West Pacific (see Richer de (Smith, 1884), it was originally described as Pentacheles Forges & Justine, 2006; Bouchet et al., 2008). It is believed nanus. The generic assignment of this species has been in that many more marine lobsters with novel morphological flux and sometimes the genus Pentacheles was used again diversity (e.g. the new genus living fossil Laurentaeglyphea for this species. However, the synonym Pentacheles nanus neocaledonica (Richer de Forges, 2006) discovered in is not repeated in the synonymy because such a comination of names is already given as the original name in square brackets next to the name currently in use. Sometimes a different suffix was used for this species such