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Recent Advances and Conflicts in Concepts of Anomuran Phylogeny (Crustacea: Malacostraca)

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Recent Advances and Conflicts in Concepts of Anomuran Phylogeny (Crustacea: Malacostraca) Arthropod Systematics II Phylogeny 119 67 (2) ■■ 119-135 ) Museum fur Tierkunde Dresden, elSSN 1864-8312, 25.8.2009 Recent Advances and Conflicts in Concepts of Anomuran Phylogeny (Crustacea: Malacostraca) 1 2 RAFAEL LEMAITRE * & PATSY A. MCLAUGHLIN 1 Smithsonian Institution, National Museum of Natural History, Department of Invertebrate Zoology, P.O. Box 37012, Washington, D.C. 20013-7012, U.S.A. [[email protected]] 2 Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, Washington, 98221-4042, U.S.A. [[email protected]] * Corresponding author Received26.ii.2009, accepted22.v.2009. Published online at www.arthropod-systematics.de on 25.viii.2009. > Abstract In the debate over phylogenetic relationships within the Decapoda that has raged for more than a century, the Anomura has been the source of many "conflicts", including disagreements over which taxa belong in this morphologically diverse infraorder, and even what name is appropriate (Anomura or Anomala). The Anomura currently includes 17 families, 222 genera, and about 2,469 species, although 54% of the genera and 43% of the species are paguroids. A number of studies have summarized the traditional as well as recent concepts of the infraorder that were based on morphology. This review addresses modern studies on systematics of this group over the last two decades that have been based on molecular as well as morphological data, and which have continued to add controversy to concepts of anomuran phylogeny. The landmark study by C.W. Cunningham and co-workers (published in 'Nature' in 1992), proclaiming that molecular data confirmed the traditional hypothesis on the evolution of king crabs from hermit crabs, was the catalyst for several studies on anomuran evolution that followed, and is the starting point of this review. Modern studies are divided as follows, and discussed: 1) morphological, larval and molecular phylogenies exclusively of the Paguroidea and/or Anomura; 2) spermatologically de- rived phylogenies; 3) information from the fossil record; 4) phylogenetic assessments of anomuran taxa included in general decapod analyses; and 5) auxiliary information pertaining to the Paguroidea in general, and Pylochelidae in particular. These studies have made useful contributions to understanding the "big picture" of anomuran relationships but they also have limi- tations. It is concluded that the Anomura remains today as much a source of discord as it was a century ago, and "conflicts" in analyses will continue to cloud the landscape until more basic, complete information is gathered for all members of this intriguing and varied infraorder. > Key words Anomura, phylogenetic relationships, modern concepts, conflicts, review. 1. Introduction In the debate over phylogenetic relationships within mouelle, 1819, Hippoidea Latreille, 1825, Lomisoi- the Decapoda Latreille, 1802 that has continued for dea Bouvier, 1894 (as Lomoidea) (not 1895 as cited more than a century, the Anomura has been the source by MARTIN & DAVIS 2001), and Paguroidea Latreille, of many of the "conflicts." Not only have disagree- 1802 and bear the name Anomura MacLeay, 1838 in ments flared over which taxa belong in this infraorder, preference to older, but less frequently used, Anomala but even what name is appropriate. MCLAUGHLIN & Latreille, 1816. Both the name and the hierarchy with- HOLTHUIS (1985) reviewed the historical aspects and in the infraorder have yet to gain complete acceptance. ultimately concluded that this major taxon should And, as TAVARES (2003) pointed out, the lingering un- be comprised of the superfamilies Galatheoidea Sa- certainties about the 'primitive crabs' remain a per- 120 LEMAITRE & MCLAUGHLIN: Anomuran phylogeny manent threat to the higher classification of both the & ABELE (1986). MCLAUGHLIN (1983a) removed the Anomura and Brachyura. We begin this review by ini- monotypic Lomisidae Bouvier, 1894 (as Lomidae) tially accepting the Anomura as proposed by MARTIN from the Paguroidea, elevating it to its own super- & DAVIS (2001). Recent advances in several avenues family. MCLAUGHLIN (1983b) suppressed the super- of systematics, however, have resulted in continuing family Coenobitoidea Dana, 1851, uniting the six controversies regarding our concepts of anomuran then recognized hermit crab families, Pylochelidae phylogeny. Although our focus is on modern conflicts Bate, 1888, Coenobitidae Dana, 1851, Diogenidae and concepts, a brief summary of the more prominent Ortmann, 1892, Paguridae Latreille, 1802, Parapa- classification schemes used in the past to define the guridae Smith, 1882 and Lithodidae Samouelle, Anomura provides an informative prelude to our dis- 1819, under the superfamily Paguroidea. In contrast, cussion (see Appendix). MARTIN & ABELE (1986) found the families Lomisi- A historical background of the taxa that have been dae (as Lomidae) and Lithodidae (Hapalogastrinae at one time or another included in the Anomura or and Lithodinae) to be sister group to the remaining Anomala was provided by MCLAUGHLIN et al. (2007), Paguroidea. However, it was the startling proclama- although they did not mention, for the most part, the tion "From a hermit to a king" on the cover of the various characters used to propose the different clas- February 6, 1992 issue of the journal 'Nature' that sifications. To define their classifications, early carci- propelled paguroid phylogeny into the arena of major nologists used characters such as: pleon type, whether scientific interest. In an article in that issue entitled it was straight and well developed or folded under the "Evolution of king crabs from hermit crab ancestors", cephalothorax (LATREILLE 1816) or intermediate be- CUNNINGHAM et al. (1992) alleged that their molecu- tween straight and folded (H. MILNE EDWARDS 1837), lar data proved not only the traditional hypothesis mouthpart morphology (DE HAAN 1841), or degree of that hermit crabs (Paguridae) gave rise to king crabs cephalization (DANA 1853). Based on mode of life, (Lithodidae) through carcinization and peramorpho- BOAS (1880) placed the Anomala in his Reptantia or sis (ALBERCH et al. 1979), but that the relationship "creepers". BORRADAILE (1906) derived the Anomala was so close that king crabs were nested in the com- from the Thalassinidea based on six characters that in mon hermit crab genus Pagurus Fabricius, 1775 (see his view separated these two groups from other deca- Fig. 1). CUNNINGHAM et al.'s (1992) data also disas- pods: reduction of the pleon, free carapace not fused sociated the diogenid species Clibanarius vittatus to the epistome, presence of a movable antennal scale, (Bosc, 1802) and an unidentified species of the ter- shape of endopod of first maxilliped, presence of 'ap- restrial coenobitid genus, Coenobita Latreille, 1829 pendices intern ae' on pleopods, and third pair of max- from the other paguroids. Interestingly, in a similar illipeds not covering the mouth. BOUVIER (1940) used: molecular study, SPEARS et al. (1993) found the dro- separation of epistome from the carapace and lack of miid Hypoconcha arcuata Stimpson, 1858 more its clear fusion to the first pleonal somite; presence of closely related to C. vittatus than to a second dromi- 'linea anomurica'; reduced or non-existent rostrum; id, Cryptodromiopsis antillensis (Stimpson, 1858), narrow endopod of the first maxilliped; epipods re- raising not only the question of the relationship of duced or absent on first and third maxillipeds, absent the primitive crabs to the Anomura, but the accu- on second maxilliped; reduced fifth pereopod termi- racy of mitochondrial DNA as an indicator of phylo- nating in chela or subchela; and larval characteristics. genetic relationships, at least when too few taxa were Larval characters were also used by GURNEY (1942) utilized in the analyses. CUNNINGHAM et al. (1992) and MACDONALD et al. (1957). The classifications used strengthened their molecularly based hypothesis by WATERMAN & CHACE (1960) and BOWMAN & ABELE with implications of morphological, larval and fossil (1982) in two major treatises on crustacean biology support. This landmark study was the catalyst for were each based on up-to-date advances at the time, the numerous analyses of phylogenetic relationships whereas DE SAINT LAURENT'S (1979) was based on her within the Anomura and among reptant decapods phyletic re-evaluation of the Decapoda as a whole, and that have followed. Although several avenues sub- SCHRAM'S (1986) incorporated fossil evidence. sequently have been explored in some detail, the resulting phylogenies rarely have been congruent. We will first review the methods and results of post- CUNNINGHAM et al. (1992) phylogenetic studies that Modern concepts have focused directly on the Anomura in general and the Paguroidea in particular, and then consider the conflicts that have arisen when only few exemplars The first challenges to the traditional classification of have been taken as representative of one or more of the Anomura as given by BOWMAN & ABELE (1982) the anomuran superfamilies during the examination were those of MCLAUGHLIN (1983a,b) and MARTIN of decapod relationships. Arthropod Systematics & Phylogeny 67 (2) 121 2.1. Morphological, larval and molecular will describe shortly, the fossil record now provides phylogenies exclusively of the Paguroidea some information that was not available to MCLAUGH- LIN & LEMAITRE (1997). and/or Anomura Although their emphasis was on the Paguroidea and the relationship between lithodids and pagurids, The first
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