This article appeared in a journal published by Elsevier. The attached copy is furnished to the author for internal non-commercial research and education use, including for instruction at the authors institution and sharing with colleagues. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright Author's personal copy Zoologischer Anzeiger 250 (2011) 407–448 Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology Joelle C.Y. Laia,∗ , Jose Christopher E. Mendozaa , Danièle Guinotb , Paul F. Clarkc , Peter K.L. Nga a Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Singapore Muséum national d’Histoire naturelle, Département Milieux et peuplements aquatiques, Case Postale 53, 61 rue Buffon, 752315 Paris cedex 05, France c Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom b Received 30 May 2011; received in revised form 4 July 2011; accepted 5 July 2011 Abstract Currently, 13 subfamilies are recognised in the brachyuran family Xanthidae: Actaeinae, Antrocarcininae, Chlorodiellinae, Cymoinae, Etisinae, Euxanthinae, Kraussiinae, Liomerinae, Polydectinae, Speocarcininae, Xanthinae, Zalasiinae and Zosiminae. This classification has been based on shared adult features like a transversely ovate carapace, well defined dorsal carapace regions, usually with lateral dentition, stout chelipeds and relatively short ambulatory legs. Such characters are now considered to be convergent. Consequently a number of higher xanthid taxa may be artifical and not monophyletic. A broad sample of 147 xanthid species representing 75 out of 124 genera from all 13 xanthid subfamilies were sampled in a multi-gene analysis. Four markers (three mitochondrial and one nuclear) were used and yielded a tree with ca. 30 xanthid clades. Monophyletic support was demonstrated for the Antrocarcininae (although substantially redefined), Cymoinae, and Polydectinae. Almost every other subfamily was para- or polyphyletic. Furthermore, the two other families of the Xanthoidea, Pseudorhombilidae and Panopeidae, were found nested within the Xanthidae. The molecular results were consistent with phylogenetic relationships implied by a suite of novel and/or neglected “ventral” adult characters including sternal characters, position of genital openings and morphology of the first zoea, instead of “dorsal” characters traditionally used to infer xanthid relationships. © 2011 Elsevier GmbH. All rights reserved. Keywords: Systematics; Brachyura: Decapoda; Xanthoidea; Xanthidae; Subfamilies; Revision; Selected genera and species; Multi-gene analysis; Adult characters; First stage zoeal morphology; Phylogeny 1. Introduction ∗ Corresponding author. Tel.: +65 65162969. E-mail addresses: dbslcyj@nus.edu.sg (J.C.Y. Lai), dbsjose@nus.edu.sg (J.C.E. Mendoza), guinot@mnhn.fr (D. Guinot), p.clark@nhm.ac.uk (P.F. Clark), peterng@nus.edu.sg (P.K.L. Ng). 0044-5231/$ – see front matter © 2011 Elsevier GmbH. All rights reserved. doi:10.1016/j.jcz.2011.07.002 Xanthid crabs are common and speciose in macrofaunal communities of reef habitats worldwide. It is the most species-rich family within the Brachyura, with 13 subfamilies, 124 genera and 639 species from a total of ca. 1300 genera and 7000 species (updated from Ng et al., 2008). The taxonomy and systematics of the Xanthidae, changed Author's personal copy 408 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 substantially during the latter half of the last century. Balss (1957) regarded the Xanthidae as a homogenous group divided into four subfamilies that included the pilumnids, carpiliids and menippids, while Glaessner (1969), on the basis of fossil and recent forms, partitioned the Brachyrhyncha into three superfamilies, one of which was the Xanthoidea. Raoul Serène also worked extensively on xanthid taxonomy (e.g., Serène, 1962, 1971; Serène and Nguyen, 1960; Serène and Umali, 1972; Serène and Vadon, 1981). Most of his ideas were eventually synthesised into the classic study on Red Sea and western Indian Ocean xanthoids (Serène, 1984). Xanthids have traditionally been classified on the basis of “dorsal” characters such as the carapace shape and nature of the lateral dentition, cheliped and ambulatory leg characters. However, according to Guinot (1967a,b,c, 1968a,b,c, 1969a,b,c,d, 1971, 1976, 1977a,b, 1978, 1979), these “traditional” characters were subject to substantial convergence. Moreover, she considered that the Xanthidae was likely a polyphyletic assemblage and proposed a radical reclassification, based on ventral features such as sternal morphology and position of genital openings, characters believed to better reflect phylogenetic affinities (Guinot, 1977a,b, 1978, 1979). Her new classification (Guinot, 1978) proposed a superfamily Xanthoidea with eight families: Carpiliidae, Geryonidae, Menippidae, Panopeidae, Pilumnidae, Xanthidae, Platyxanthidae, and Trapeziidae. Later, Martin and Davis (2001) recognised an enlarged Xanthoidea to include the Eumedonidae, Goneplacidae and Pseudorhombilidae. In contrast, Števčić (2005) restricted the Xanthoidea to the Xanthidae, Panopeidae and Speocarcinidae. Karasawa and Schweitzer (2006), however, recognised a different arrangement, with their Xanthoidea containing the Domeciidae, Panopeidae, Pilumnidae, Pseudorhombilidae, Trapeziidae and Xanthidae. Several studies have attempted to integrate groupings based on zoeal morphology and setotaxy with traditional classifications of adult xanthids. The larval data of Rice (1980) did not support the simple divisions of the system of Balss (1957), but neither did it agree well with the proposal of Guinot (1978). Martin (1984) initially endorsed the scheme of Glaessner (1969), but later (Martin et al., 1985) stated that larval morphology appeared to partly support Balss (1957). However, Clark and Galil (1988) suggested that their larval evidence corresponded most closely to the classification of Platyxanthidae and Trapeziidae, as proposed by Guinot (1978). They concluded that careful re-examination of other xanthoid larvae may further substantiate the classification of Guinot (1978). Clark (2007) studied first stage zoeae from 48 species representing 11 subfamilies; but larvae of Antrocarcininae and Zalasiinae were not represented. He found the Xanthidae problematic as his analyses suggested that larval and adult based systematics was incongruent. The analysis of first stage zoeas did not support the existing adult-based classification of the 11 subfamilies studied, in particular the Actaeinae, Cymoinae, Etisinae, Kraussiinae, Liomerinae and Zosiminae. In addition, Clark (2007) questioned the Xanthinae as recognised by Serène (1984) and most modern workers by suggesting that it was not a monophyletic taxon. His study also did not support the Etisinae and Chlorodiellinae (as Chlorodiinae). These two subfamilies were included in a group together with the first stage zoeae represented by Xantho hydrophilus, the type species of Xantho, which is the type genus of the Xanthidae. Finally, he found that on the basis of first stage zoeal antennal types, the xanthids could be divided into five groups that did not correspond to any of the adult-based classifications. Recently, Ng et al. (2008) substantially redefined and restricted the Xanthoidea to three families: Xanthidae, Panopeidae and Pseudorhombilidae. The two latter families are small compared with the Xanthidae, with 25 genera and 94 species and 8 genera and 14 species, respectively. Within the Xanthidae, Ng et al. (2008) recognised 13 subfamilies: Actaeinae, Antrocarcininae, Chlorodiellinae, Cymoinae, Etisinae, Euxanthinae, Kraussiinae, Liomerinae, Polydectinae, Speocarcininae, Xanthinae, Zalasiinae and Zosiminae. As discussed by Ng et al. (2008), their classification was established primarily on adult morphology and they strongly suggested that a number of taxa were probably based on convergent characters with some groups being artificial and/or polyphyletic. Serène (1984), Ng (1993b), Ng and Chia (1994) and Ng and Chen (2004) discussed at length the adult characters used to distinguish these subfamilies. In contrast, the diagnosis of the Speocarcininae by Števčić (2005) was scant and he refered the reader to Serène (1984), Ng (1993a) and Ng and Chia (1994) for further details. While the core genera of some subfamilies share distinctive suites of characters, such as those assigned to the Cymoinae and Polydectinae, other peripheral genera often have less consistent character suites and seem to have been almost arbitrarily assigned to their higher taxa. For example, the Etisinae and Chlorodiellinae share spoon-tipped fingers and sometimes the dactylo-propodal locks on their legs, but whether this phylogenetically unites them, or represents convergence, as the current subfamily recognition implies, needs further investigation. As Ng et al. (2008) discussed, subfamilies such as the Euxanthinae, Actaeinae, Liomerinae, Xanthinae and Zosiminae are difficult to diagnose, as there are many “exceptions”. In fact, a precise diagnosis of the Xanthidae has proven to be extremely difficult. Even the utility of the male first and second gonopods is limited, as the intergeneric variation appears to be too substantial to find clear group patterns. The purpose of the present study is to obtain molecular data from selected xanthid taxa comprising key representatives of all 13 xanthoid subfamilies. They will be used to test the current hypothesis of xanthid systematics and classification as proposed by Ng et al. (2008) and, in conjunction with support from adult morphology and zoeal characters, outline parameters for a revised classification. Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 2. Material and methods 2.1. Taxon sampling A broad sample of taxa representing 147 species (75 out of 124 genera) from 13 Xanthidae subfamilies was sequenced. They were primarily obtained from specimens in the collections of the Raffles Museum of Biodiversity Research, Singapore (ZRC), Natural History Museum, London (NHM), Florida Museum of Natural History, University of Florida, Gainesville (UF), Muséum national d’Histoire naturelle, Paris (MNHN), Ryukyu University Museum, Fujukan (FLMNH), University of the Ryukyus, Okinawa (RUMF-ZC), Philippine National Museum, Manila (NMCR), and National Museum of Science and Technology, Tokyo (NMST). The tissue samples also included three new genera which will be fully described in separate publications. Additional sequence data for 11 xanthoid taxa were obtained from sequences deposited in GenBank. For outgroups, species from three other families were chosen, Benthochascon hemingi (Portunidae), Menippe rumphii (Menippidae), Trapezia cymodoce, and Quadrella coronata (Trapeziidae). The latter two families were included because they were previously associated with the Xanthidae. See Table 1 for list of taxa for DNA, collection sources, and accession numbers. 2.2. DNA extraction, amplification, and sequencing Muscle tissue was excised from appendages or egg masses of alcohol-preserved specimens. DNA was extracted using the DNeasy Tissue and blood kit (QIAGEN Inc., Valencia, CA) following the protocol of the manufacturer. Concentration was quantified using a nanodrop ND 1000 spectrophotometer (Nanodrop Technologies). Molecular characters for phylogenetic inference were obtained from four (three mitochondrial and one nuclear) markers, namely the mitochondrial 12S and 16S ribosomal subunits, the protein coding gene Cytochrome Oxidase I and histone H3 (hereafter referred to as 12S, 16S, COI and H3), respectively, using primers previously published in Thoma et al. (2009) (12S), Lai et al. (2009) (16S) and Colgan et al. (1998) (H3). COI is commonly used for interspecific phylogeny and barcoding studies while the other three, being slightly more conserved were used to examine relationships at the inter-generic and subfamily level. 18S ribosomal subunit data was obtained during study but excluded from analyses as it was uninformative. The Folmer region of COI was amplified using primers designed by Brent Thoma and kindly shared for the present study (see Thoma et al., 2009; Felder and Thoma, 2010). Following an initial denaturation and final extension phase of 94 ◦ C for four minutes and 72 ◦ C for 10 min, respectively, all markers were amplified in 34 standard Polymerase Chain Reaction (PCR) cycles consisting of 409 94 ◦ C for 30 s (denaturation), 45–52 ◦ C for 30 s (annealing; COI [45–50 ◦ C], 16S and H3 [47 ◦ C]), 12 s [52 ◦ C]) and 72 ◦ C for 60 s (extension) run on G-Storm GS1 (Gene Technologies Ltd, Essex, UK) and PTC-100 (MJ Research, Watertown, MA, USA) thermal cyclers. Amplifications were conducted in 25 ␮l volumes containing 2.5–3 mM MgCl2 , 0.025 mM of each dNTP, 12.5 pmol of each primer, 0.2 units of GoTaq DNA polymerase (Promega, Madison, WI, USA), 5 ␮l 5× GoTaq buffer and 1–100 ng of whole genomic DNA. Amplicons were visualized by gel electrophoresis on a 1% agarose gel and successfully amplified PCR products were purified using either Millipore PCR cleanup filter plates (Millipore Corporation, Danvers, MA, USA) or Agencourt AMPure system (Agencourt, Beverly, MA, USA) before being subjected to cycle sequencing using the ABI PRISM® Dye terminator kit containing AmpliTaq and BigDye (version 3) dye terminator. Although the majority of ingroup taxa were sequenced for all target loci, this was not possible for some samples due to repeated failure in amplification and sequencing attempts. Each sequencing reaction comprised 5–8 ng of PCR product, 1 ␮l BigDye, 0.5 ␮l 5× BigDye sequencing buffer, 0.4 ␮l sequencing primer (2 pmol/␮l) and topped up to 5 ␮l with sterile Milli-Q water. Cycle sequencing parameters followed manufacturer’s protocol and all extension products were purified using CleanSEQ dyeterminator removal system (Agencourt) before being read on a 3100 capillary sequencer (Applied Biosystems, Foster City, CA, USA). Sequences were read from both forward and reverse directions and combined in Sequencher v.4.8 (Gene Codes Corporation, Ann Arbor, MI, USA) to eliminate errors and reduce ambiguity. 2.3. Alignment MAFFT (Katoh et al., 2002) was used to align sequences for each marker. COI and H3 were aligned using default parameters (FFT-NS-2) and the Q-INS-i strategy employed for 16S and 12S as described by Katoh and Toh (2008). Alignments were then adjusted manually in MacClade 4 (Maddison and Maddison, 2005) before being concatenated into a single dataset. Regions of ambiguous alignment in 16S and 12S sequences were excluded and gaps were treated as “missing” data. 2.4. Phylogenetic analyses Maximum Likelihood (ML) and Bayesian Inference (BI) were used to infer phylogenetic relationships, using a combined data set of all sequences. Exploratory analyses indicated that single gene datasets were not sufficiently informative to yield robust results. MrModeltest version 2.2 (Nylander, 2004) was used for identifying the best-fit model for individual loci partitions and the complete data set based on the Akaike Information Criterion (AIC) for BI. Likelihood tree searches were performed using Randomized Author's personal copy 410 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Table 1. List of terminal taxa used in tree construction with relevant data (Sampling locality, museum catalogue number, and GenBank accession numbers). All specimens were physically examined, except for those marked with an asterisk (*). Taxa Actaeinae Actaea polyacantha (Heller, 1861) Actaea semblatae Guinot, 1976 Actaeodes hirsutissimus (Rüppell, 1830) Actaeodes mutatus Guinot, 1976 Actaeodes tomentosus (H. Milne Edwards, 1834) Epiactaea nodulosa (White, 1848) Gaillardiellus orientalis (Odhner, 1925) Gaillardiellus rueppelli (Krauss, 1843)* Novactaea bella Guinot, 1976 Odhneria echinus (Alcock, 1898)* Paractaea rufopunctata (H. Milne Edwards, 1834) Psaumis cavipes (Dana, 1852) Pseudactaea corallina (Alcock, 1898) Pseudoliomera variolosa (Borradaile, 1902) Serenius kuekenthali (De Man, 1902) Serenius pilosus (A. Milne-Edwards, 1867) Antrocarcininae Cyrtocarcinus truncatus (Rathbun, 1906) Glyptocarcinus lophopus Takeda, 1973 Glyptocarcinus politus Ng & Chia, 1994 Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Japan: Okinawa, Sunabe China: Fujian Province, San Yu Harbour Indonesia: Pulau Bintan ZRC 1999.0261 HM851285 HM798416 HM750945 HM798265 ZRC 2002.0577 HM851286 HM798417 HM750946 HM798266 ZRC 2008.1143 HM851287 HM798418 N/A HM798267 Singapore: Raffles Lighthouse Malaysia: Sabah, Pulau Sapi ZRC 1993.0287–0288 HM851288 HM798419 N/A HM798268 ZRC 2000.1673 HM851289 HM798420 HM750947 HM798269 Singapore: Off Sentosa Island. Singapore: Palau Seringat ZRC 1995.0333 HM851321 HM798450 HM750974 HM798299 ZRC 2000.1196 HM851336 HM798464 HM750987 HM798314 Philippines: Bohol Island ZRC 2010.0162 HM851337 HM798465 HM750988 HM798315 Indonesia: Pulau Bintan Vanuatu: NW coast of Malo Island Guam: Pago Bay ZRC 1998.0981 HM851401 HM798529 HM751044 HM798378 ZRC 2009.1177 HM851402 HM798530 HM751045 HM798379 ZRC 2000.0718 HM851407 HM798535 HM751048 HM798383 ZRC 2010.0157 HM851421 HM798549 N/A HM798395 ZRC 2003.0276 HM851422 HM798550 HM751061 HM798396 Hawaii: Moku Manu Island ZRC 2000.0508 HM851423 HM798551 HM751062 HM798397 Philippines: Panglao Island, Biking. Philippines: Panglao Island, Momo Beach. ZRC 2010.0159 HM851428 HM798556 HM751066 HM798402 ZRC 2010.0158 HM851429 HM798557 HM751067 HM798403 New Caledonia: Ouvéa, Est de Mouly RUMF-ZC-01302 HM851316 HM798445 HM750970 HM798294 Japan: East of Muko-jima NSMT SY-09-04 HM851340 HM798468 HM750991 HM798318 Japan: Kaikata seamount NSMT KT-09-2 KK1-2(1) HM851341 HM798469 HM750992 HM798319 Philippines: Sungcolan Bay, Panglao Island Philippines: Balicasag Island Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 411 Table 1 (Continued) Taxa Chlorodiellinae Chlorodiella barbata (Borradaile, 1900) Chlorodiella corallicola Miyake & Takeda, 1968 Chlorodiella cytherea (Dana, 1852) Chlorodiella laevissima (Dana, 1852) Chlorodiella nigra (Forskål, 1775) Chlorodiella xishaensis Chen & Lan, 1978 Cyclodius ungulatus (H. Milne Edwards, 1834) Garthiella aberrans (Rathbun, 1906) Garthiella sp. nov. Pilodius areolatus (H. Milne Edwards, 1834) Pilodius paumotensis Rathbun, 1907 Cymoinae Cymo melanodactylus Dana, 1852 Cymo quadrilobatus Miers, 1884 Etisinae Etisus anaglyptus H. Milne Edwards, 1834 Etisus dentatus (Herbst, 1785) Etisus electra (Herbst, 1801) Etisus odhneri Takeda, 1971 Etisus splendidus Rathbun, 1906 Etisus utilis Jacquinot, in Jacquinot & Lucas, 1853 Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Australia: Lizard Island UF 17105 HM851301 HM798431 HM750957 HM798279 Philippines: Pamilacan Island ZRC 2010.0296 HM851302 HM798433 N/A HM798281 Philippines: Panglao Island, Alona Beach ZRC 2008.0644 HM851303 HM798434 HM750958 HM798282 Australia: Lizard Island, Yonge Reef UF 17262 HM851304 HM798435 HM750959 HM798283 Oman: Bar Al Hikman Peninsula Philippines: Balicasag Island UF 17948 HM851307 HM798437 HM750961 HM798286 ZRC 2010.0289 HM851308 HM798438 HM750962 HM798287 Philippines: Balicasag Island ZRC 2010.0138 HM851312 HM798441 HM750965 HM798290 Society Islands: Moorea, east of Opunohu Pass Philippines: Balicasag Island Guam: Southwest Orote Peninsula UF 15624 HM851338 HM798466 HM750989 HM798316 ZRC 2010.0142 HM851339 HM798467 HM750990 HM798317 ZRC 2000.0720 HM851415 HM798543 HM751055 HM798390 Indonesia: Anambas Islands, Pulau Jemaja, Telok Tiru ZRC 2003.0559 HM851416 HM798544 HM751056 HM798391 Singapore: Pulau Seringat ZRC 1999.0339 JF930158 FJ548948 HM750967 FJ548931 Guam: Hospital Point ZRC 2001.0755 HM851314 HM798443 HM750968 HM798292 Malaysia: Pulau Tioman, Paya Beach ZRC 1999.0931 HM851322 HM798451 HM750975 HM798300 Guam: Dadi Beach ZRC 2001.0753 HM851324 HM798453 HM750977 HM798302 Hawaii: Oahu, Heeia Kea, Kaneohe Bay Philippines: Pamilacan Island Indonesia: Sulawesi, Bunaken Island Thailand: Phuket, Ao Nam Bay ZRC 2000.0503 HM851325 HM798454 HM750978 HM798303 ZRC 2010.0140 HM851326 HM798455 HM750979 HM798304 ZRC 2008.0526 HM851327 GQ249175 HM750980 GQ249172 ZRC 2002.0586 HM851328 HM798456 HM750981 HM798306 Author's personal copy 412 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Table 1 (Continued) Taxa Etisus aff. villosus Clark & Galil, 1995 Euxanthinae Alainodaeus filipinus Mendoza & Ng, 2008 Cranaothus deforgesi Ng, 1993 Crosnierius carinatus Serène & Vadon, 1981 Danielea noelensis (Ward, 1942) Epistocavea mururoa Davie, 1992 Euxanthus exsculptus (Herbst, 1790) Euxanthus herdmani Laurie, 1906 Euxanthus huonii (Hombron & Jacquinot, 1846) Euxanthus ruali Guinot, 1971 Glyptoxanthus erosus (Stimpson, 1859) Hepatoporus orientalis (Sakai, 1935) Hypocolpus abbotti (Rathbun, 1894) Hypocolpus diverticulatus (Strahl, 1861)* Hypocolpus pararugosus Crosnier, 1997 Ladomedaeus fungillus Manuel-Santos & Ng, 2007 Ladomedaeus serratus (Sakai, 1965) Medaeops granulosus (Haswell, 1882) Medaeops neglectus (Balss, 1922) Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Philippines: Panglao Island, northwest coast ZRC 2008.1480 HM851329 HM798457 N/A HM798307 Philippines: Panglao Island, northwest coast ZRC 2009.0142 HM851290 HM798421 HM750948 HM798270 Philippines: Panglao Island, Arco Point ZRC 2008.1363 HM851310 HM798439 HM750963 HM798288 Philippines: Luzon Island, eastern coast ZRC 2008.1370 HM851311 HM798440 HM750964 HM798289 Maldives: Laamu Atoll ZRC 2007.0757 HM851317 HM798446 N/A HM798295 Philippines: Panglao Island, Looc ZRC 2008.1371 HM637922 GQ249176 HM638032 GQ249173 Malaysia: Pulau Tioman, Paya Beach ZRC 2002.0535 HM851332 HM798460 HM750983 HM798310 Philippines: Panglao Island, Looc Philippines: Panglao Island, Pontod Islet NMCR 27334 HM851333 HM798461 HM750984 HM798311 ZRC 2008.1376 HM851334 HM798462 HM750985 HM798312 Vanuatu: E Aoré Island, Aimbué Bay USA: Georgia ZRC 2009.1178 HM851335 HM798463 HM750986 HM798313 ZRC 1998.9 HM851342 HM798470 HM750993 HM798320 Philippines: Off western coast of Batangas Philippines: Balicasag Island Madagascar: Northwest of Nosy Komba Philippines: Balicasag Island. ZRC 2008.1379 HM851343 HM798471 HM750994 HM798321 NMCR 27335 HM851345 HM798473 HM750996 HM798323 UF 14076 HM851346 HM798474 HM750997 HM798324 ZRC 2008.1389 HM851347 HM798475 HM750998 HM798325 Philippines: Panglao Island ZRC 2010.0130 HM851351 HM798479 HM751001 HM798329 Taiwan: Off Tahsi Port ZRC 2009.0342 HM637923 HM637955 HM638045 HM596617 Singapore: Pulau Seringat ZRC 1999.0856 HM851371 HM798499 HM751019 HM798348 Madagascar: Trois Freres Islet UF 14599 HM851372 HM798500 HM751020 HM798349 Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 413 Table 1 (Continued) Taxa Medaeus danielita Mendoza & Ng, 2010 Medaeus elegans A. Milne-Edwards, 1867 Medaeus ornatus Dana, 1852 Medaeus sp. nov. Miersiella cavifrons Takeda, 1989 Miersiella haswelli (Miers, 1886) Monodaeus couchii (Couch, 1851) Monodaeus cristulatus Guinot & Macpherson, 1988 Monodaeus rectifrons (Crosnier, 1967) Monodaeus aff. tuberculidens (Rathbun, 1911) New genus 2 sp. nov. New genus 3 sp. nov. 1 New genus 3 sp. nov. 2 Olenothus uogi Ng, 2002 Paramedaeus globosus Serène & Vadon, 1981 Paramedaeus aff. simplex Pseudomedaeus distinctus (Rathbun, 1898) Rizalthus anconis Mendoza & Ng, 2008 Visayax osteodictyon Mendoza & Ng, 2008 Kraussinae Kraussia rugulosa (Krauss, 1843)* Palapedia sp. 1* Palapedia sp. 2* Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Society Islands: Moorea, off Sheraton Hotel beach Guam: Apra harbour UF 15469 HM851373 HM798501 HM751021 HM798350 UF 2053 HM851374 HM798502 N/A HM798351 Guam: Apra harbour ZRC 2009.1180 HM851375 HM798503 N/A HM798352 Vanuatu: Aorè island, Port Benier Philippines: Panglao Island, Maribohoc Bay Philippines: Pamilacan Island France: Thalassa Exp., Golfe de Gascogne Namibia: Exp. Benguela VI, dredged from 232 m ZRC 2010.0163 HM851376 HM798504 HM751022 HM798353 ZRC 2008.1396 HM851380 HM798508 HM751026 HM798357 ZRC 2008.1400 HM851381 HM798509 HM751027 HM798358 MNHN B30718 HM851384 HM798512 HM751028 HM798361 MNHN B17498 HM851385 HM798513 HM751029 HM798362 Republic of the Congo: Large de Pointe-Noire MNHN B9580 HM851386 HM798514 HM751030 HM798363 Réunion Island MNHN B16164 HM851387 HM798515 HM751031 HM798364 Philippines: East of Luzon Island Philippines: Balicasag Island Philippines: Balicasag Island Guam ZRC 2010.0164 HM851398 HM798526 HM751041 HM798375 ZRC 2010.0152 HM851399 HM798527 HM751042 HM798376 ZRC 2010.0153 HM851400 HM798528 HM751043 HM798377 ZRC 2002.0176 HM851403 HM798531 HM751046 HM798380 Philippines: Panglao Island, Bolod NMCR 27352 HM851409 HM798537 HM751050 HM798385 Philippines: Panglao Island, Sungcolan Bay USA: Florida, off St. Petersburg ZRC 2008.1408 HM851408 HM798536 HM751049 HM798384 UF 6659 HM851424 HM798552 N/A HM798398 Philippines: Panglao Island, Pontod lagoon ZRC 2008.0215 HM851427 HM798555 N/A HM798401 Philippines: Panglao Island ZRC 2008.0753 HM851432 HM798559 HM751070 HM798405 Taiwan: Hsiao-Liu Kiu ZRC 2009.0335 HM851348 HM798476 HM750999 HM798326 Vanuatu: Tutuba Island Vanuatu: Segond Channel ZRC 2009.1175 ZRC 2009.1176 HM851404 HM851405 HM798532 HM798533 N/A HM751047 HM798381 HM798382 Author's personal copy 414 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Table 1 (Continued) Taxa Liomerinae Liomera bella (Dana, 1852) Liomera cinctimana (White, 1847) Liomera tristis (Dana, 1852) #1 Liomera tristis (Dana, 1852) #2 Liomera tristis (Dana, 1852) #3 Liomera venosa (H. Milne Edwards, 1834) Lipkemera corallina (Takeda & Marumura, 1997) Lipkemera holthuisi Mendoza, 2010 Neoliomera cerasinus Ng, 2002 Neoliomera demani Forest & Guinot, 1961 Neoliomera insularis (Adams & White, 1849) Neoliomera striata Buitendijk, 1941 Menippidae Menippe rumphii (Fabricius, 1798) Panopeidae Glyptoplax smithii A. Milne-Edwards, 1880* Hexapanopeus paulensis Rathbun, 1930* Panopeus herbstii H. Milne Edwards, 1834* Panopeus occidentalis Saussure, 1857* Polydectinae Lybia edmondsoni Takeda & Miyake, 1970 Lybia hatagumoana Sakai, 1961 Lybia leptochelis (Zehntner, 1894) Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 ZRC 1993.646-652 HM851357 HM798485 HM751007 HM798335 ZRC 2000.0730 HM851358 HM798486 HM751008 HM798336 ZRC 2003.0217 HM851359 HM798487 HM751009 HM798337 ZRC 1993.653-656 HM851360 HM798488 N/A HM798338 ZRC 2010.0144 HM851361 HM798489 HM751010 HM798339 ZRC 1999.0383 HM851362 HM798490 HM751011 HM798340 Philippines: Bohol, Maribojoc Bay ZRC 2009.1172 HM851377 HM798505 HM751023 HM798354 Philippines: Panglao Island, north coast. Japan: Ryukyus, Kume-jima ZRC 2009.1171 HM851378 HM798506 HM751024 HM798355 ZRC 2001.2265 HM851390 HM798518 HM751034 HM798367 USA: Hawaii (?), purchased from Singapore aquarium Philippines: Balicasag Island ZRC 2009.1174 HM851391 HM798519 HM751035 HM798368 ZRC 2001.0666 HM851392 HM798520 HM751036 HM798369 Philippines: Balicasag Island ZRC 2008.0641 HM851393 HM798521 HM751037 HM798370 Singapore: Labrador Beach ZRC 2003.0211 HM637946 HM637976 HM638051 HM596626 USA: Southwestern Gulf of Mexico ULLZ 6793 EU863276 EU863342 N/A N/A Panama city: Northern Gulf of Mexico ULLZ 8645 EU863311 EU863377 N/A N/A USA: North Carolina, Masonboro Sound, salt marsh USA: Northern Gulf of Mexico, Panama City ZRC 2001.1015 HM851406 HM798534 JF930161 JF930160 ULLZ 8643 EU863328 EU863394 N/A N/A ZRC 2000.0435 HM851366 HM798494 HM751015 HM798344 ZRC 2010.0147 HM851367 HM798495 HM751016 HM798345 ZRC 2010.0146 HM851368 HM798496 N/A N/A Japan: Okinawa, Motobu-cho, Bise Guam: Apra harbour Malaysia: Pulau Tioman, Paya Beach Japan: Okinawa, Motobu-cho, Bise Philippines: Panglao Island, Momo Beach Indonesia: North coast of Pulau Bintan Hawaii: Mai Palaoa Beach, near Maili Point Philippines: Panglao Island, near San Isidro Philippines: Pamilacan Island Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 415 Table 1 (Continued) Taxa Lybia tesselata (Latreille, in Milbert, 1812) Polydectus cupulifer (Latreille, in Milbert, 1812) Portunidae Benthochascon hemingi Alcock & Anderson, 1899 Pseudorhombilidae Trapezioplax tridentata (A. Milne-Edwards, 1880)* Speocarcininae Speocarcinus lobatus Guinot, 1969* Speocarcinus monotuberculatus Felder & Rabalais, 1986* Trapezidae Quadrella coronata Dana, 1852 Trapezia cymodoce (Herbst, 1801) Xanthinae Cycloxanthops truncatus (De Haan, 1837) Demania cultripes (Alcock, 1898) Demania intermedia Guinot, 1969 Demania scaberrima (Walker, 1887) Eucratodes agassizii A. Milne-Edwards, 1880* Euryxanthops dorsiconvexus Garth & Kim, 1983 Euryxanthops orientalis (Sakai, 1939) Garthiope spinipes (A. Milne-Edwards, 1880)* New genus near Garthiope* Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Guam: Southwest of Orote Peninsula ZRC 2000.0710 HM851369 HM798497 HM751017 HM798346 Hawaii: Magic Island ZRC 2000.0441 HM851420 HM798548 HM751060 HM798394 New Caledonia ZRC 2000.0102 N/A FJ548945 HM750955 FJ548929 USA: Northern Gulf of Mexico. ULLZ 8054 EU863278 EU863344 N/A N/A USA: Northern Gulf of Mexico. ULLZ 7820 EU863341 EU863407 N/A N/A South-western Gulf of Mexico. ULLZ 7562 EU863293 EU863359 N/A N/A Philippines: Balicasag Island Singapore: Southern Islands ZRC 2008.0449 HM851426 FJ548953 HM751065 FJ548936 ZRC 2009.0079 HM851431 FJ548961 HM751069 FJ548941 Philippines: Balicasag Island ZRC 2010.0139 HM851313 HM798442 HM750966 HM798291 Taiwan: Peace Island ZRC 1998.530 HM851318 HM798447 HM750971 HM798296 Philippines: Northwest coast of Panglao Thailand: Phuket, Pichai fishing port ZRC 2009.0187 HM851319 HM798448 HM750973 HM798297 ZRC 2000.0833 HM851320 HM798449 HM750972 HM798298 Northern Gulf of Mexico ULLZ 8400 EU863323 EU863389 N/A N/A Philippines: Bohol/Sulu seas ZRC 2010.0136 HM851330 HM798458 HM750982 HM798308 Philippines: East Luzon, off Aurora Province Southern Gulf of Mexico ZRC 2010.0141 HM851331 HM798459 N/A HM798309 ULLZ 8131 EU863339 EU863405 N/A N/A Northern Gulf of Mexico ULLZ 8183 EU863300 EU863366 N/A N/A Author's personal copy 416 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Table 1 (Continued) Taxa Lachnopodus bidentatus (A. Milne-Edwards, 1867) Lachnopodus subacutus (Stimpson, 1858) Leptodius exaratus (H. Milne Edwards, 1834) Leptodius nigromaculatus Serène, 1962 Leptodius nudipes (Dana, 1852) Liagore rubromaculata (De Haan, 1835) #1 Liagore rubromaculata (De Haan, 1835) #2 Macromedaeus crassimanus (A. Milne-Edwards, 1867) Metaxanthops acutus Serène, 1984 Micropanope sculptipes Stimpson, 1871* Nanocassiope alcocki (Rathbun, 1902) Neoxanthias michelae Serène & Vadon, 1981 Neoxanthops lineatus (A. Milne-Edwards, 1867) Neoxanthops quadrilobatus (Sakai, 1939) Paraxanthias elegans (Stimpson, 1858) Paraxanthias notatus (Dana, 1852) Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Papua New Guinea: Laing Island ZRC 1995.0411 HM851349 HM798477 HM751000 HM798327 Philippines: Balicasag Island ZRC 2003.0282 HM851350 HM798478 N/A HM798328 Indonesia: Natuna, Pulau Salor ZRC 2003.0549 HM851352 HM798480 HM751002 HM798330 Singapore: Raffles Lighthouse ZRC 2008.0652 HM851353 HM798481 HM751003 HM798331 Malaysia: Sabah, Pulau Sapi Japan: Okinawa, Ishigawa Port ZRC 2000.1675 HM851354 HM798482 HM751004 HM798332 ZRC 2005.0013 HM851355 HM798483 HM751005 HM798333 Philippines: Bohol Island, Cortes ZRC 2010.0143 HM851356 HM798484 HM751006 HM798334 Philippines: Balicasag Island ZRC 2003.0369 HM851370 HM798498 HM751018 HM798347 Philippines: Bohol Island, Bolod ZRC 2010.0148 HM851379 HM798507 HM751025 HM798356 Southeastern Gulf of Mexico ULLZ 6603 EU863338 EU863404 N/A N/A Philippines: Southwestern Luzon, off western coast of Batangas Thailand: Phuket, Pichai fishing port ZRC 2010.0149 HM851389 HM798517 HM751033 HM798366 ZRC 1999.0516 HM851394 HM798522 HM751038 HM798371 Singapore: Sentosa Island ZRC 1987.0531 HM851395 HM798523 HM751039 HM798372 Taiwan: Beibu Wan. ZRC 1999.0022 HM851396 HM798524 N/A HM798373 Philippines: Bohol Island, Baclayon ZRC 2010.0155 HM851411 HM798539 HM751052 HM798386 Japan: Okinawa, Sunabe, Chutan ZRC 1999.0260 HM851412 HM798540 N/A HM798387 Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 417 Table 1 (Continued) Taxa Paraxanthias pachydactylus (A. Milne-Edwards, 1867) Paraxanthodes obtusidens (Sakai, 1965) Xanthias canaliculatus Rathbun, 1906* Xanthias lamarckii (H. Milne Edwards, 1834) Xanthias latifrons (De Man, 1887) Xanthias punctatus (H. Milne Edwards, 1834) Xanthias teres Davie, 1997 Xantho hydrophilus (Herbst, 1790) Xantho pilipes A. Milne-Edwards, 1867* Zalasiinae Banareia nobilii (Odhner, 1925) Calvactaea tumida Ward, 1933 Zalasius dromiaeformis (De Haan, 1839) Zalasius sakaii Balss, 1938 Zosiminae Atergatis floridus (Linnaeus, 1767) Atergatis integerrimus (Lamarck, 1818) Atergatis interruptus Takeda & Marumura, 1997 Atergatis obtusus A. Milne-Edwards, 1865 Atergatopsis aff. amoyensis Atergatopsis aff. obesa Atergatopsis germaini A. Milne-Edwards, 1865 Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Philippines: Bohol Island, Baclayon ZRC 2010.0154 HM851413 HM798541 HM751053 HM798388 Philippines: Off eastern Luzon Island ZRC 2008.1410 HM851414 HM798542 HM751054 HM798389 South Africa: Sodwana Bay ULLZ 4381 EU863316 EU863382 N/A N/A Philippines: Panglao Island, Sungcolan Inlet ZRC 2010.0160 HM851433 HM798560 HM751071 HM798406 Guam: Tepungan Channel Philippines: Balicasag Island ZRC 2000.0728 HM851434 HM798561 HM751072 HM798407 ZRC 2001.0309 HM851435 HM798562 HM751073 HM798408 Philippines: Bohol Island, Maribojoc Bay England: Plymouth ZRC 2010.0161 HM851436 HM798563 HM751074 HM798409 ZRC 2011.0492 HM851437 HM798564 HM751075 HM798410 Ireland: Flaggy Shore NHM 2010.477 HM637925 HM637956 HM638061 HM596616 Philippines: Panglao Island, Bolod Philippines: Panglao Island, Bolod Taiwan ZRC 2010.0131 HM851299 HM798429 HM750954 HM798277 ZRC 2010.0137 HM851300 HM798430 HM750956 HM798278 ZRC 2008.0680 HM851438 HM798565 HM751076 HM798412 Taiwan: Kaoshiung county, Mitou ZRC 1997.0399 HM851439 HM798566 HM751077 HM798413 Singapore: Pulau Seringat Singapore: Beting Bronok Reef ZRC 2009.0373 HM851291 HM798422 HM750949 JF930159 ZRC 2007.0252 HM851292 HM798423 HM750950 HM798271 Philippines: Northwest coast of Panglao Island ZRC 2008.1471 HM637924 FJ548944 HM638024 FJ548928 Philippines: Panglao, Suncolon Inlet ZRC 2010.0132 HM851293 HM798424 HM750951 HM798272 Philippines: Bohol Island, Maribojoc Bay Philippines: Balicasag Island Taiwan: Keelung, Ho-Ping Island ZRC 2010.0133 HM851295 HM798426 N/A HM798274 ZRC 2010.0135 HM851296 HM798427 N/A HM798275 ZRC 2009.0298 HM851297 N/A HM750952 N/A Author's personal copy 418 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Table 1 (Continued) Taxa Atergatopsis granulata A. Milne-Edwards, 1865 Lophozozymus anaglyptus (Heller, 1861) Lophozozymus pictor (Fabricius, 1798) Lophozozymus edwardsi (Odhner, 1925) New genus 1 sp. nov. Paratergatis longimanus Sakai, 1965 Platypodia eydouxi (A. Milne-Edwards, 1865) Platypodia pseudogranulosa Serène, 1984 Platypodia tomentosa (De Man, 1902) Pulcratis reticulatus Ng & Huang, 1997 Zosimus aeneus (Linnaeus, 1758) Zosimus hawaiiensis (Rathbun, 1906) Collected from Catalogue number GenBank accession numbers 12S 16S COI H3 Philippines: Panglao Island, Catarman ZRC 2010.0134 HM851298 HM798428 HM750953 HM798276 Philippines: Pamilacan Island ZRC 2010.0145 HM851363 HM798491 HM751012 HM798341 Singapore: Raffles Lighthouse ZRC 2009.0592 HM851364 HM798492 HM751013 HM798342 Indonesia: Natuna islands, inlet off Pulau Laut Philippines: Pamilacan Island. Taiwan: I-lan county, Tai-chi Port ZRC 2003.0553 HM851365 HM798493 HM751014 HM798343 ZRC 2010.0151 HM851397 HM798525 HM751040 HM798374 ZRC 1998.0047 HM851410 HM798538 HM751051 N/A Hawaii: Mai Palaoa Beach, near Maili Point ZRC 2000.0440 HM851417 HM798545 HM751057 HM798392 Singapore: Cyrene Reef ZRC 2008.0492 HM851418 HM798546 HM751058 HM798393 Philippines: Balicasag Island ZRC 2010.0156 HM851419 HM798547 HM751059 N/A Taiwan: Ping-tung County ZRC 1997.0402 HM851425 HM798553 HM751064 HM798399 Taiwan: Ping-tung County, Heng Chun Peninsula Hawaii: North of Oahu ZRC 1998.0388 HM851440 HM798567 HM751078 HM798414 ZRC 2000.0515 HM851441 HM798568 HM751079 HM798415 Accelerated Maximum Likelihood (RAxML) ver. 7.2.6 (Stamatakis, 2006; Stamatakis et al., 2008) as implemented on the CIPRES portal (http://www.phylo.org/sub sections/portal/) (Miller et al., 2009) with the dataset partitioned according to locus. An estimate of the proportion of invariant sites is not implemented in RAxML because it is sensitive to the number and divergences of the sequences included in the data (see RAxML manual; Stamatakis, 2006). A rapid bootstrap (BS) analysis was performed with 100 replications and use to search for the best scoring ML tree using the GTRCAT model. After implementing the BS analysis, every fifth BS tree was used as a starting point to search for the ML tree using a specified model of sequence evolution, saving 10 best-scoring ML trees (fast ML searches). Finally, RAxML calculated more accurate likelihood scores (slow ML searches) for those 10 trees and placed BS proportions on the best-scoring ML tree. Gene partitioned BI analyses were carried out using MrBayes version 3.1.2 (Huelsenbeck and Ronquist, 2003). Four Metropolis coupled Monte Carlo Markov chains (3 heated and 1 cold) were run for three million generations with trees sampled every 1000 generations (3000 trees sampled). The first 750,000 generations (750 trees) were discarded as “burn-in” and the posterior probabilities were estimated for the remaining sampled generations. Two independent Bayesian runs initiated from random starting trees were performed for each data set, and the log-likelihood values and posterior probabilities were checked to warrant that chains had reached stationarity before conclusion. Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 2.5. Adult morphology For the adults, most taxa included in the molecular tree were examined in detail. The morphological terminology used follows that of Guinot (1979), Serène (1984) and Ng et al. (2008). Abbreviations used in the text include: G1, first male pleopod, or first gonopod; G2, second male pleopod, or second gonopod; mxp1–mxp3, first to third maxillipeds; P1–P5, first to fifth pereiopods with P1 as chelipeds. The thoracic somites and sternites are numbered from 1 to 8 with 1 being anterior and 8, posterior. The thoracic sternal sutures are referred to by the number of the two sternites that are involved, and are numbered from 1/2 to 7/8. The term “coaptation” (from the Latin cum, with; aptare, fit), has been in use for a long time in medical language and refers to the setting of a fracture or luxation. It was revived by the French biologist Lucien Cuénot (1921, 1925) to name, in arthropods, specifically insects, the fitting between two different and originally independent parts of the body, with a more or less pronounced engagement. Many types of coaptation were developed in almost all brachyuran crabs to retain their abdomen beneath the cephalothorax. The more well known is the “bouton-pression” or press-button (Cuénot, 1941). Guinot and Bouchard (1998) in their paper on the different holding mechanisms for the abdomen of brachyuran crabs, also discussed the different types or schemes of coaptation observed in brachyurans. In the present work, the term “coaptation” refers to the close fitting of certain surfaces of the carapace (e.g., subhepatic regions, posterolateral regions) with corresponding surfaces on the pereiopods (e.g., inner surfaces of merus, carpus, propodus and dactylus of P1), and of a pereiopod with other adjacent pereiopods, when they are folded towards the carapace, as in a crab’s resting position (see also Serène, 1984; Mendoza et al., 2009b). When the corresponding surfaces of associated appendages and/or carapace fit tightly with each other, the condition is referred to as “perfect coaptation”. Only a few species exhibit “perfect coaptation”, for instance, Hypocolpus perfectus Guinot-Dumortier, 1960 (Fig. 6a–c). More species show “near-perfect” coaptation (Fig. 6d–f). Even so, near-perfect coaptation is rare in the Xanthidae and can only be seen in two groups: Actaeodes and members of the Euxanthinae sensu stricto. For the rest of the Brachyura, perfect and near-perfect coaptation can also be found in members of the Calappidae, Aethridae and Parthenopidae. In the Discussion, adult morphology for each major clade is treated first, followed by the larval morphology. A list of all adult material examined, is presented in Appendix A. 2.6. First stage zoeas First stage zoeae from 48 species representing 11 of the 13 xanthid subfamilies and 5 panopeids were examined (Appendix B). Morphological terminology follows Clark et al. (1998) and Clark (2007). The sequence of zoeal 419 descriptions was based on the malacostracan somite plan and described from anterior to posterior. Setal armature of appendages was described from proximal to distal segments and in order of endopod to exopod (Clark et al., 1998). The first stage zoeas were described and fully illustrated, except for the mandible, because the only significant character of this appendage is the appearance of the palp in later zoeal stages and it is not present in the stage examined in the present study. The long antennular aesthetascs and the long plumose natatory setae of the first and second maxillipeds were drawn truncated. The first stage zoeas were dissected on glass slides in polyvinyl lactophenol (Gray and Weiss, 1950) under a Leica MZ 16 binocular microscope and the appendages allowed to clear for 24 h before examination. Coverslips were sealed with clear nail varnish. Appendages were drawn using a Leica DMR HC microscope with differential interference contrast (DIC) and a camera lucida. Only the first stage zoea of Xantho hydrophilus was described and illustrated in full, as this is the type genus of the Xanthidae. In order to ensure consistency and standardisation of setal interpretation, described zoeas from other published sources were not used. 3. Results 3.1. Sequence data and molecular analyses Following alignment, curation and concatenation, the final dataset (for 164 terminal taxa) for four markers consisted of 1725 positions. MrModeltest selected the models GTR + I + G (General Time Reversible plus Invariant sites plus Gamma distributed model) for the loci 16S and COI, and HKY + I + G (Hasegawa-Kishino-Yano plus Invariant sites plus Gamma distributed model) for 12S and H3. Bayesian and Likelihood analyses of the combined data produced similar, well-resolved, phylogenetic hypotheses (Fig. 1), with minor differences at nodes with low branch support (<50%). Clades were mostly recovered at the subfamily level, and some at the generic level, with high ML bootstrap (BS) and BI posterior probability (PP) support. However, some clades did not corroborate the view of xanthid classification as recognised by Serène (1984) or Ng et al. (2008). Almost every subfamily was poly- or paraphyletic with the exception of Cymoinae, Polydectinae, and Antrocarcininae. Furthermore, the other families of Xanthoidea (Pseudorhombilidae, Panopeidae) were nested within the Xanthidae. 3.1.1. Actaeinae Actaeinae was represented by 11 genera in the present study, forming three major clades: Act 1 comprising Actaea, Epiactaea, Novactaea, Serenius, with support of 100 (BS) and 1 (PP), Act 2 comprising Gaillardiellus, Paractaea, Pseudoliomera (100/1) and Act 3, comprising Actaeodes (99/1). Three actaeine genera were nested within other subfamilies: Author's personal copy 420 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 1. Bayesian Inference (BI) phylogeny inferred from combined 12S, 16S rRNA, COI and H3 sequences. Numbers above and below branches indicate BI posterior probability and Maximum Likelihood (ML) bootstrap support, respectively. “–” represents values below 0.5 or 50. “” indicates shared clades recovered from BI and ML analyses while “O” indicates taxa with data obtained from Thoma et al. (2009). Subfamily annotations (in parentheses) follow the classification of Ng et al. (2008). Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 1. (Continued ). 421 Author's personal copy 422 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Psaumis within Euxanthinae (Eux 1), Odhneria within Xanthinae (Xan 1) and Pseudactaea within Zalasiinae (Zal). 3.1.2. Antrocarcininae Represented by Cyrtocarcinus and Antrocarcinus, both of which were nested within one of three euxanthine clades (Eux 3) with high support (95/1). 3.1.3. Chlorodiellinae Represented by four genera and clustered into two distant clades: Chl 1, comprising Chlorodiella, Cyclodius, Pilodius (92/1), and Chl 2 with Garthiella (100/1). 3.1.4. Cymoinae Cymoinae was represented by Cymo quadrilobatus and C. melanodactylus, and both taxa were recovered in the same clade (Cym) (with high support under both ML and BI, 100/1) and allied with Polydectinae (Pol), although this was only strongly supported under ML (<50/0.91). Cymoinae was the only monogeneric subfamily. 3.1.5. Etisinae Etisinae was represented by one genus and seven species. Five species formed a core Etisinae clade (Eti 2) (86/0.98) and two species, Etisus odhneri (Eti 1) and E. electra (Eti 3), were basal to a Chlorodielline clade (Chl 1) and xanthine clade (Xan 3), respectively. The association of E. odhneri with Chlorodiellinae was not statistically supported. 3.1.6. Euxanthinae Seventeen genera clustered into two major euxanthine clades: Eux 1 (57/0.98) comprising Euxanthus, Hepatoporus, Hypocolpus, Olenothus, Rizalthus, Visayax and an actaeine, Psaumis; and Eux 3 (60/0.53) contained Alainodaeus, Crosnierus, Cranaothus, Danielea, Epistocavea, Ladomedaeus, Medaeops, Miersiella, Monodaeus, Paramedaeus and two new genera. Eux 3 also included two xanthine genera (Paraxanthodes and Euryxanthops), a xanthine species, Xanthias teres, and the subfamily Antrocarcininae. Glyptoxanthus, represented by G. erosus (Eux 2), was sister to the Zalasiinae (50/0.98) (Zal). Pseudomedaeus, represented by P. distinctus, was the sister group to the xanthine genus, Cycloxanthops (Xan 5). Of the four Medaeus species, three were nested within a larger xanthine clade (Xan 10) (only supported in BI) with Lachnopodus, Paraxanthias and Metaxanthops species. Medaeus danielita was sister to the chlorodielline genus, Garthiella (clade Chl 2). 3.1.7. Kraussiinae The two kraussiine genera did not group together: Palapedia was in Kra 1 (100/1) and Kraussia (Kra 2) sister to the Etisus (Eti 2) clade (51/0.89). 3.1.8. Liomerinae Liomerinae was represented by three genera, which formed two clades: Lio 1 consisting of Liomera and Lipkemera (92/1) and Lio 2, Neoliomera (100/1). The zosimine Paratergatis and a related new genus were nested within Lio 1 with high support (94/0.99). 3.1.9. Polydectinae This subfamily was recovered as monophyletic (Pol) with high support (99/1) although the four Lybia species included in this study were split into two terminal clades, each with two species. Polydectus was sister to L. hatagumoana and L. leptochelis. 3.1.10. Panopeidae Two Speocarcinus species, representing the Speocarcininae, were nested in a larger clade comprising mostly Atlantic xanthoids. A pseudorhombilid, Trapezioplax tridentata (Pse) was sister to the Speocarcininae (Spe) (94/1) and together they were sister to a xanthine clade (Xan 7) containing Eucratodes and Micropanope (71/0.96). The Spe + Pse + Xan 7 clade, is sister to a monophyletic panopeid (Pan) group (96/1) represented by three genera. Panopeidae is nested within the Xanthidae, being sister to the Pseudorhombilidae + Speocarcininae + Xanthinae clade with good support under BI (62/0.94). Sister to this clade (Spe + Pse + Xan 7 clade) is a new genus near Garthiope (Xan 8) (B. Thoma and D. Felder, pers. comm.), followed by another xanthine, Garthiope spinipes (Xan 6). 3.1.11. Xanthinae Xanthinae was represented by 18 genera, most of which were paraphyletic and widely dispersed throughout the tree (Xan 1–10): Xan 1 included taxa from other subfamilies: Pulcratis reticulatus (Zosiminae) and Odhneria echinus (Actaeinae). Xanthias was not found to be associated with any other xanthine clade; X. punctatus, X. canaliculatus and X. lamarckii were associated with a zosimine clade (Zos 1), X. teres was nested within an euxanthine clade (Eux 3) and X. latifrons (Xan 2) was sister to an actaeine + euxanthine + liomerine + xanthine clade (but with low support under ML). Xantho (the type genus of the Xanthidae), represented by X. hydrophilus and X. pilipes, was not allied with other xanthine genera but was observed to be a sister to a number of other subfamily clades (Xan 4). Nanocassiope (Xan 9) was not closely related to other xanthines and was in a unique monophyletic lineage. 3.1.12. Zalasiinae Zalasiinae was represented by three genera and appears to be monophyletic (81/1). Included in this clade (Zal) was the actaeine, Pseudactaea corallina, as sister to Zalasius with moderate statistical support (71/0.86). 3.1.13. Zosiminae Represented by seven genera and clustered into two major clades (Zos 1 and Zos 2): each sister to an actaeine clade (Act 1 and Act 2). All genera in Zos 1 and Zos 2, except for Zosimus, appeared to be para- or polyphyletic, with Atergatopsis and Lophozozymus present in both clades. Pulcratis Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 423 Fig. 2. Xantho hydrophilus (Herbst, 1790): first stage zoea; (a) anterior view of carapace; (b) antennule; (c) antenna; (d) maxillule; (e) maxilla; (f) telson. Scale bar = 0.1 mm. and Paratergatis were nested within a xanthine (Xan 1) and liomerine (Lio 1) clade, respectively. 3.2. Zoea 3.2.1. Zoeal description Xantho hydrophilus (Herbst, 1790). Xantho incisus—Lebour, 1928: 530–531, Pl. II Fig. 3, Pl. XI Figs. 5–9 (prezoea, zoea I–IV, megalopa, crab 1); Ingle, 1983: 970–973, Figs. 7–12 (zoea I–IV, megalopa); Ingle, 1991: 237–238, Figs. 1.3d, 1.13p, 1.23d, 1.29d, 1.32e, 1.35a, 1.36j; 1.38f, 2.32, Pl. 6g (zoea I–IV). Xantho incisus granulicarpus—Bourdillon-Casanova, 1960: 171 (zoea I–IV). Carapace (Fig. 2a): dorsal spine long, curved distally, longer than rostral spine; rostral spine shorter than antennal protopod, without distal spinulation; lateral spines straight, without spinulation on dorsal margin; anterodorsal setae absent; 1 pair of posterodorsal setae; ventral margin without setae; eyes sessile. Antennule (Fig. 2b): uniramous, endopod absent; exopod unsegmented with 4 (2 broad, 2 slender) terminal aesthetascs of unequal length plus 1 terminal seta. Antenna (Fig. 2c): protopodal process distally multispinulate, longer than rostral spine; endopod spine present; exopod ca. 4% length of protopod with 2 unequal terminal setae. Mandible: palp absent. Maxillule (Fig. 2d): coxal endite with 7 setae; basial endite with 5 setal processes and 2 small setal buds; endopod 2segmented, proximal segment with 1 seta; distal segment with 6 (2 subterminal, 4 terminal) setae; exopod seta absent. Maxilla (Fig. 2e): coxal endite bilobed with 4 + 4 setae; basial endite bilobed with 5 + 4 setae; endopod bilobed, with 3 + 5 (2 subterminal, 3 terminal) setae; exopod (scaphognathite) margin with 4 setae and 1 long distal stout process. First Maxilliped (Fig. 3a): coxa with 1 setae; basis with 10 setae arranged 2,2,3,3; endopod 5-segmented with 3,2,1,2,5 (1 subterminal, 4 terminal) setae, respectively; exopod 2segmented, distal segment with 4 long terminal plumose natatory setae. Second Maxilliped (Fig. 3b): coxa without setae; basis with 4 setae arranged 1,1,1,1; endopod 3-segmented, with 1,1,6 (3 subterminal, 3 terminal) setae, respectively; exopod 2-segmented, distal segment with 4 long terminal plumose natatory setae. Third Maxilliped: absent. Pereiopods: absent. Abdomen (Fig. 3c and d): 5 somites; somite 2 with 1 pair of dorsolateral processes directed anteriorly; somite 3 with 1 pair of dorsolateral processes directed ventrally; somites 1 and 2 each with rounded posterolateral processes and 3–5 each with short posterolateral spinous processes; somite 1 without setae; somites 2–5 each with 1 pair of posterodorsal setae; pleopod buds absent. Telson (Figs. 2f, 3c and d): each fork long, gradually curved distally and not spinulate with 1 large and 1 smaller lateral spine plus 1 large dorsomedial spine; posterior margin with 3 pairs of stout spinulate setae. 3.2.2. First stage zoeal characters Setal morphology is conservative for a number of xanthid characters and these generally provide little phylogenetic information. These characters include the presence of Author's personal copy 424 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 3. Xantho hydrophilus (Herbst, 1790): first stage zoea; (a) first maxilliped; (b) second maxilliped; (c) dorsal view of abdomen; (d) lateral view of abdomen. Scale bar = 0.1 mm. lateral and dorsal carapace spines (Fig. 2a), antennule (Fig. 2b), maxillule (Fig. 2d), maxilla (Fig. 2e), mxp1 (Fig. 3a), mxp2 (Fig. 3b) with the exception of the distal endopod segment, the setation on abdominal somites 2–5 (Fig. 3c and d) and setation of the telson margin (Figs. 2f, 3c and d). Only the characters that differ from the above description will be discussed with reference to support of clades in the DNA analysis and these include the spinulation of the rostral spine (Fig. 4a–d), presence of setae on the ventral margin of the carapace (Fig. 4a–d), the setation of the antennal endopod (Fig. 4e–l), the antennal exopod (Fig. 4e–l) and its distal setation, the presence or absence of seta 3.5 in the distal segment of the second maxilliped (Fig. 5a and b) and the spinulation of the telson (Fig. 5c–f). 4. Discussion 4.1. Systematic and taxonomic implications The topology of the present phylogenetic tree questions the Xanthoidea as currently understood and outlined in Ng et al. (2008). It also indicates that the current classification of the Xanthidae based on “dorsal” adult morphology is artificial, with these traditional characters subject to substantial convergence. Therefore, a radical redefinition and classification of Xanthidae systematics is necessary. While Xanthoidea is considered monophyletic, the assignment and composition of various families are unsatisfactory. For example, results from genetic analyses suggest that the Panopeidae and Pseudorhombilidae are nested within the Xanthidae. Consequently, should these taxa be subsumed as subfamilies, or should some of the xanthid subfamily-groupings be raised to familial status? If the Panopeidae and Pseudorhombilidae are recognised as such, then clades Xan 10 and Chl 2, should perhaps also be regarded as distinct families. In addition, almost every xanthid subfamily and a number of genera (as currently understood) were shown to be para- or polyphyletic. A broad overview of a new xanthid classification based on molecular data with support from known adult and larval morphology is provided below, with additional comments on some genera. 4.2. Mapping of morphological characters 4.2.1. Actaeinae The diagnostic characters of the Actaeinae have been ambiguous because most species superficially conform to a general plan that includes an ovate carapace with a short front, distinct regions, many granules and/or spines arranged in complex ornamentations, short ambulatory legs and relatively short and stout chelipeds (Odhner, 1925; Guinot, 1976; Serène, 1984). Currently, 20 genera and 107 species are assigned to this subfamily (updated from Ng et al., 2008). For the present study, nine genera and 15 species were analysed, and these taxa clustered into three clades suggesting the actaeines are polyphyletic. “Actaeine” genera such as Odhneria, Psaumis and Pseudactaea are nested in clades outside the subfamily and present morphological examinations confirm they are not true actaeines (see discussion for Xan 1, Eux 1, Zal). Act 1: Actaea, Epiactaea, Novactaea, Serenius. This clade represents the Actaeinae s. str., containing the type genus, Actaea. The adult characters supporting this clade include a sparsely setose dorsal carapace surface, a relatively narrow carapace, a moderately produced front, grooves and canals absent from the subhepatic and pterygostomian regions, minimal coaptation of all pereiopods (as seen in most Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 425 Fig. 4. Carapace lateral and rostral spines without well defined spinulation: (a) Gaillardiellus orientalis (Odhner, 1925); (b) Lophozozymus pictor (Fabricius, 1798); (c) Epiactaea nodulosa (White, 1848); (d) Novactaea bella Guinot, 1976. Antennal morphology of xanthid first stage zoea: (e) Acantholobulus bermudensis (Benedict & Rathbun, 1891); (f) Gaillardiellus orientalis (Odhner, 1925); (g) Actaeodes hirsutissimus (Rüppell, 1830); (h) Actaeodes mutates Guinot, 1976; (i) Gaillardiellus orientalis (Odhner, 1925); (j) Lophozozymus pictor (Fabricius, 1798); (k) Epiactaea nodulosa (White, 1848); (l) Novactaea bella Guinot, 1976. xanthids), a dactylo-propodal lock present on ambulatory legs (Fig. 6m and n), the thoracic sternite 4 without transverse or oblique sulci, the G1 moderate in length (Fig. 9g), and the press-button located anteriorly on sternite 5, close to sternal suture 4/5 (Fig. 7g) (see also Guinot, 1976: figs. 41A, B, 42C; Serène, 1984: figs. 54A, B). First stage zoeas are known for Epiactaea nodulosa and Novactaea bella. These zoeas hatch at an advanced stage of development (Clark, 2001, 2005, 2009; Clark and Ng, 2004a,b). A suite of characters are associated with this abbreviated development including the carapace with a pair of anterodorsal setae, a row of setae on the ventral margin of carapace, endopod bud present on antennule and antenna (Fig. 4k–l), presence of a biramous mxp3, pereiopods present with a bilobed chela, and abdominal somites 2–5 with pleopods devoid of endopods. All of these characters are absent in xanthid first stage zoeas (in taxa with four zoeal stages), as in the example illustrated here for Xantho hydrophilus (Figs. 2 and 3). Whether these characters are diagnostic for the whole group or artificial due to early expression associated with abbreviated development can only be determined when more zoeas from this group become known. Other zoeal characters for Novactaea and Epiactaea include an extremely long dorsal carapace spine which is curved distally, without spinulation and just longer than the rostral spine (Fig. 4c, d); a rostral spine equal in length to the antennal protopod and without distal spinulation (Fig. 4c, d); an antennal exopod ca. 8.7% length of the protopod, possessing 3 (1 subterminal, 2 terminal) setae (Fig. 4k–l); antennal protopodal process without distal spinulation, equal in length to the rostral spine and terminally swollen (Fig. 4c, d, k, l); abdominal somite 1 with 1 medial seta and long telson forks. Author's personal copy 426 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 5. Setation of second maxilliped endopod: (a) Acantholobulus bermudensis; (b) Actaeodes tomentosus. Telson spine armature from residual to reduced and lost: (c) Gaillardiellus orientalis (Odhner, 1925); (d) Lophozozymus pictor (Fabricius, 1798); (e) Epiactaea nodulosa (White, 1848); (f) Novactaea bella Guinot, 1976. The first stage zoeas of E. nodulosa can be distinguished from those of N. bella by possessing a dorsal carapace spine terminally swollen (vs. pointed), lateral carapace spines terminally swollen (vs. pointed), and a telson with 3 minute spines (Fig. 5e) (vs. absent; Fig. 5f). Zoeas representing the sister group comprising A. polyacantha and S. pilosus are not known. Act 2: Gaillardiellus, Paractaea, Pseudoliomera. The adult characters that diagnose this clade include minimal coaptation of pereiopods with antero- and postero-lateral margins of carapace, complete transverse sulcus between thoracic sternites 3 and 4, ambulatory dactylo-propodal lock absent, abdomen generally broader (vs. narrower in Actaeinae s. str. and Actaeodes), disto-lateral angles of male abdominal somite 6 well produced to accommodate a relatively large press-button, which is close to suture 4/5, and male telson short and broad (see Guinot, 1976: figs. 41D, 42A; Serène, 1984: figs. 54D, E). The first stage zoea is known for Gaillardiellus orientalis (see Ng and Clark, 1994) and the characters indicate that this species has four zoeal stages and not abbreviated as listed for Actaeinae sensu stricto. Gaillardiellus orientalis zoeas differ from those of X. hydrophilus, possessing a relatively long rostral spine (Fig. 4a) (vs. proportionally shorter; Fig. 2a), a long dorsal carapace spine with a swollen tip (Fig. 4a) (vs. proportionally shorter) with a swollen tip (vs. pointed tip; Fig. 2a), lateral carapace spines with swollen tips (Fig. 4a) (vs. pointed; Fig. 2a), the antennal protopod being relatively long (vs. shorter) without distal spinulation (vs. spinulate) and terminally swollen (vs. pointed) (Fig. 4a, f vs. Fig. 2a, c), antennal protopod without endopod spine (vs. present) (Fig. 4f vs. Fig. 2c), and long (vs. proportionally shorter) telson forks with 3 minute spines (Fig. 5c) (vs. 3 prominent spines; Fig. 2f). Gaillardiellus orientalis shares a number of zoeal characters with members of the Actaeinae s. str. (Act 1) including long carapace rostral and dorsal spines, and antennal protopod all without spinulation; carapace dorsal and lateral spines, and antennal protopod terminally swollen; and long telson forks. The first stage zoea is not known for Pseudoliomera variolosa (sampled for molecular analysis), but Clark and Galil (1998) described that of P. speciosa. However, this zoea differs significantly from that of G. orientalis in that the carapace dorsal and rostral spines, and the antennal protopod are relatively short (vs. significantly longer); the carapace dorsal and lateral spines, and the antennal protopod are terminally pointed (vs. swollen); the carapace dorsal spine is distally spinulate (vs. smooth); the carapace lateral spines are dorsally spinulate (vs. smooth); the rostral spine is spinulate (vs. smooth); the antennal protopod with a prominent endopod spine (vs. absent); the antennal exopod is proportionately longer (vs. proportionately shorter); length of telson fork proportionately short (vs. much longer) and telson spinulation prominent (vs. minute). The significance of these differences between P. speciosa and G. orientalis zoeas is uncertain. For example, the first stage zoea of P. speciosa differs only in a few carapace characters from Actaeodes mutatus and A. tomentosus (both assigned to Act 3) by possessing a distally spinulate dorsal spine (vs. smooth), dorsally spinulate Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 427 Fig. 6. Example of perfect coaptation. (a–c) Hypocolpus perfectus Guinot-Dumortier, 1960, MNHN-B6516. Example of near-perfect coaptation, (d–f) Banareia nobilii (Odhner, 1925), ZRC. (g–i) Presence of modified cutting tooth on dactylus of major chela: (g) Medaeops granulosus (Haswell, 1882), ZRC 1965.11.10.46-50; (h) Xantho poressa (Olivi, 1792), ZRC 1988.694-697; (i) Xanthias teres Davie, 1997, ZRC 2001.0689. (j–l) Absence of modified cutting tooth on dactylus of major chela: (g) Medaeops granulosus (Haswell, 1882), ZRC 1965.11.10.46-50; (h) Xantho poressa (Olivi, 1792), ZRC 1988.694-697; (i) Xanthias teres Davie, 1997, ZRC 2001.0689; (j) Demania splendida Laurie, 1906, ZRC 2001.0650, (k) Palapedia roycei (Serène, 1972), ZRC; (l) Lobiactaea lobipes (Odhner, 1925), ZRC. (m–o) Ambulatory dactylo-propodal locking mechanism: (m) Zosimus aeneus (Linnaeus, 1758), ZRC 1995.612; (n) Pilodius pilumnoides (White, 1848), ZRC 1965.11.11.150; (o) Liomera cinctimana (White, 1847), ZRC 2000.0705. Abbreviations: dpl = dactylo-propodal lock; dt = modified dactylar cutting tooth. lateral spines (vs. smooth) and a spinulate anterodorsal region. Only further work will establish if Pseudoliomera is not monophyletic and/or its zoeas are really distinct from those described for G. orientalis by Ng and Clark (1994). Act 3: Actaeodes. The adult characters that diagnose this clade include a densely tomentose carapace, carapace transversely broad, front not well-produced, grooves and canals present on subhepatic region (Guinot, 1976: fig. 38D), near-perfect coaptation of all pereiopods with carapace and with each other (Fig. 6a–f), dactylo-propodal lock on ambu- latory legs present (Fig. 6m,n), transverse and oblique sulci on sternite 4 (Fig. 7a,d,e), G1 relatively longer (Fig. 9o) (see also Serène, 1984: figs. 55B, 78, 79), and press-button touching sternal suture 4/5 (Fig. 7h,i; Guinot, 1976: fig. 41C). Actaeodes superficially resembles members of Actaeinae s. str. externally, but it is the only actaeine genus for which all the pereiopods are tightly and near-perfectly coapted against the carapace, with the retracted ambulatory legs pressing closely against the prominent concavity on the smooth posterolateral surfaces. The same applies to the smooth cheliped palm, Author's personal copy 428 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 7. Male anterior thoracic sterna: (a) Euxanthus huonii (Hombron and Jacquinot, 1846), ZRC 1987.471; (b) Liagore pulchella Ng and Naruse, 2007, ZRC 2007.0642; (c) Liomera cinctimana (White, 1847), ZRC 2000.0705; (d) Banareia nobilii (Odhner, 1925), ZRC; (e) Medaeops granulosus (Haswell, 1882), ZRC 1965.11.10.46-50; (f) Xantho poressa (Olivi, 1792), ZRC 1988.694-697. (g–l) Male sternoabdominal cavities: (g) Banareia nobilii (Odhner, 1925), ZRC; (h) Euxanthus huonii (Hombron and Jacquinot, 1846), ZRC 1987.471; (i) Liomera cinctimana (White, 1847), ZRC 2000.0705, (j) Medaeops granulosus (Haswell, 1882), ZRC 1965.11.10.46-50; (k) Xantho poressa (Olivi, 1792), ZRC 1988.694-697; (l) Neoliomera demani Forest and Guinot, 1961, ZRC 2009.1174. Abbreviations: ab 4–6 = abdominal somites 4–6, respectively; con 1 = coxo-sternal condyle of pereiopod 1; cox 1–4 = coxae of pereiopods 1–4, respectively; ml = median line; pb = sternal press-button; s = slit for receiving the tips of the G1; st 1–7 = thoracic sternites 1–7, respectively. which neatly fits the smooth concavity of the anterolateral carapace margin. Serène (1984: 16) commented upon this unusual apomorphy (see also Guinot, 1976: pl. 15), but was apparently unaware of its phylogenetic significance. First zoeas were obtained for Actaeodes hirsutissimus and A. tomentosus (see Clark and Al-Aidaroos, 1996) together with those of A. mutatus. They possess the normal xanthid zoeal development of four stages (as with Act 2 but unlike Act 1 zoea). Apart from the characters associated with abbrevi- ated development, the zoeas of Actaeodes differed from those of Actaeinae s. str. and G. orientalis (Act 2) by possessing a relatively short (vs. longer) rostral spine with distal spinulation (vs. smooth); a shorter (vs. longer) dorsal carapace spine being terminally pointed (vs. swollen except Novactaea), lateral carapace spines that are terminally pointed (vs. swollen except Novactaea), a shorter (vs. long) antennal protopod distally multispinulate (vs. smooth) (Fig. 4g, h; vs. Fig. 4f, k, l), and proportionately shorter (vs. longer) telson forks. Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 The first stage zoea of Actaea areolata is known but tissues of this species were not sampled for genetic analysis. This zoea differed from those of Actaeodes hirsutissimus, A. mutatus and A. tomentosus by possessing only a distally smooth rostral spine (vs. spinulate); from the antennal exopod of A. hirsutissimus in possessing 3 (1 subterminal, 2 terminal) setae (vs. 1 terminal seta) and from Pseudoliomera speciosa by possessing a smooth dorsal carapace spine (vs. spinulate), smooth lateral carapace spines (vs. dorsally spinulate) and a smooth anterodorsal carapace region (vs. spinulate). Although the position of Actaea areolata requires further clarification, on zoeal evidence it may be assigned to clade Act 3. 4.2.2. Euxanthinae and Antrocarcininae Euxanthinae is one of the largest xanthid subfamilies with 27 genera and 90 species (updated from Ng et al., 2008). Twenty-one genera and 37 species were analysed in the present study, forming two major clades. Genera excluded from these two clades were Medaeus, with three species nested in a xanthine clade (Xan 10) and one species sister to a chlorodielline clade (Chl 2); Glyptoxanthus (Eux 2), sister to the zalasiine clade (Zal); and Pseudomedaeus, allied with Cycloxanthops, a xanthine (Xan 5). The two major clades recovered by the molecular analysis have morphological support. Davie (2002) and Ng et al. (2008) commented on the problems associated with defining the Euxanthinae sensu lato, the main character being a well-defined carapace anterolateral margin that descends towards the buccal cavity (Ng, 2002: fig. 2b; Mendoza and Ng, 2008: figs. 1b, 3c, 5c; Mendoza et al., 2009a: fig. 2b,e). In many genera, however, this character is not clearly defined and is therefore difficult to use. The present results corroborate the observations of Ng et al. (2008) that two broad groups may be differentiated within the Euxanthinae, separated by carapace, pereiopod, and thoracic sternal features. Eux 1 represents Euxanthinae s. str. as the type species, Euxanthus sculptilis Dana, 1851 (=E. huonii (Hombron and Jacquinot, 1846)) is clustered here. Eux 3 forms the other major euxanthine group and also includes Antrocarcininae as well as three xanthine taxa (Euryxanthops, Paraxanthodes and Xanthias teres). The data indicates that Medaeus is not monophyletic and its traditional assignment in Euxanthinae is untenable (see below). Glyptoxanthus is in a unique lineage (Eux 2), far from other euxanthines and closer to Zalasiinae (Zal). It is difficult to draw conclusions on the placement of Pseudomedaeus as the species used in the present study is not the type. More representatives from the two genera are needed in future analyses before any definite conclusion can be made. Eux 1: Euxanthus, Hepatoporus, Hypocolpus, Olenothus, Rizalthus, Visayax (+actaeine Psaumis). The adult characters that diagnose this clade are: anterior portion of anterolateral margins clearly descending to meet the buccal cavity; nearperfect coaptation of pereiopods to carapace (Fig. 6a–c); inner surfaces of cheliped carpus, propodus, and dactylus 429 curved to conform to convex pterygostomian region (best seen from anterodorsal view) and carapace posterolateral margins are concave so as to receive the folded posterior ambulatory legs; subhepatic cavities present in some genera (Fig. 6c) (e.g., Hypocolpus, Hepatoporus); the basal article of antennae is large, L-shaped or trapezoidal and obliquely oriented; the chelipeds are usually symmetrical and sometimes subequal but never markedly unequal; semi-parallel longitudinal grooves are present on the anterior of the male abdominal cavity (for receiving the tips of G1 s) (Fig. 7 h); the press-button is immediately adjacent to sternal suture 4/5 (Fig. 7h); the anterior portion of the exposed sternite 7 is as large as its episternite; the male abdomen is long with the telson reaching to the level above the coxo-sternal condyles of pereiopod 1 (Fig. 7a); the lateral margins of the male abdomen are oblique and straight with base of male abdominal somite 3 not much wider than tip of somite 5; and G2 relatively short, length less than half of G1 (Fig. 9b) (see also Guinot-Dumortier, 1960: pl. 1 figs. 4–6, pl. 2, pl. 5 figs. 28, 29, pl. 11 figs. 60–63; Guinot, 1979: fig. 23; Ng, 2002: figs. 1–4). The first stage zoeal morphology for Eux 1 is only known for Psaumis cavipes. This species has four predicted zoeal stages and differs from X. hydrophilus in having a rostral spine that is terminally spinulate (vs. smooth) and the antennal exopod is proportionately slightly longer (although the significance of this character is not known) with 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal). Eux 2: Glyptoxanthus is in a clade sister to the Zalasiinae (Zal) but an extensive suite of characters (carapace, epistome, pereiopod, male thoracic sternum, male abdomen, G1 and G2) indicate that its morphology has significantly diverged from the clades Eux 2 and Zal. Furthermore, there are no features linking Glyptoxanthus with genera assigned to clades Eux 1 and Eux 3. The zoeal morphology is not known for Glyptoxanthus. Eux 3: Alainodaeus, Crosnierus, Cranaothus, Danielea, Epistocavea, Ladomedaeus, Medaeops, Miersiella, Monodaeus, Paramedaeus + xanthines Euryxanthops, Paraxanthodes and Xanthias and antrocarcinines Cyrtocarcinus and Glyptocarcinus. The characters that define this clade include: the anterolateral margin in three possible states–anterior portion distinctly descending towards the buccal cavity (e.g., Cranaothus, Crosnierius, Danielea, Epistocavea, Medaeops, Paramedaeus, Paraxanthodes), anterior portion not well-defined and terminating at the external orbital angle (e.g., Alainodaeus, Miersiella, Monodaeus) or well-defined throughout and ends at the external orbital margin (e.g., Cyrtocarcinus, Euryxanthops), basal article of antenna simple or rectangular and vertically to obliquely oriented; imperfect coaptation of pereiopods to carapace, chelipeds markedly unequal and never symmetric, dactylus of major chela with a modified cutting tooth (Fig. 6g,i), press-button nearer centre of sternite 5 but never immediately adjacent to sternal suture 4/5 (Fig. 7j), anterior portion of exposed sternite 7 much larger than its episternite (Fig. 8e,f), male abdomen short, telson reaching to level just below coxo-sternal condyles of Author's personal copy 430 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Fig. 8. Shape of thoracic sternite 7: (a) Actaea semblatae Guinot, 1976, ZRC 2002.0577, (b) Zalasius dromiaeformis (De Haan, 1839), ZRC 2001.0383, (c) Liagore pulchella Ng and Naruse, 2007, ZRC 2007.0642, (d) Etisus dentatus (Herbst, 1785), ZRC 2001.0753, (e) Xantho poressa (Olivi, 1792), ZRC 1988.694-697, (f) Medaeus ornatus Dana, 1852, USNM 29523. Exposure of thoracic sternite 8: (g) Liagore pulchella Ng and Naruse, 2007, ZRC 2007.0642, (h) Medaeus ornatus Dana, 1852, USNM 29523, (i) Etisus dentatus (Herbst, 1785), ZRC 2001.0753. Abbreviations: ab 1–5 = abdominal somites 1–5, respectively; cox 4–5 = coxae of pereiopods 4–5, respectively; ep 6–7 = episternites of sternites 6–7, respectively; st 6–7 = thoracic sternites 6–7, respectively. P1 (Fig. 7e), abdominal somite 6 with disto-lateral expansions and concave lateral margins, abdominal somite 6 usually broader than long, lateral margins of male abdomen prominently concave (Fig. 7e), base of male abdominal somite 3 much wider than tip of somite 5, G2 relatively long, and G1 about twice as long as G2 (Fig. 9a) (see also Guinot, 1967a: figs. 20–22, 25; Davie, 1993: figs. 5–7; 1997: figs. 4–8, 12–14, Mendoza et al., 2009a: figs. 1–5). There may be five subclades nested within Eux 3 – the first containing Alainodaeus, Miersiella and Euryxanthops dorsiconvexus; the second: Cranaothus, Danielea, Paramedaeus aff. simplex and Medaeops; the third: Monodaeus; fourth: Crosnierus, Cyrtocarcinus, Epistocavea, Euryxanthops, Glyptocarcinus, Ladomedaeus, Paraxanthodes and Xanthias teres; and lastly the fifth, with Paramedaeus globosus. The first stage zoeas of Medaeops granulosus and Monodaeus couchii are known and predicted to have four zoeal stages. Both have a distally smooth rostral spine and antennal exopod proportionately slightly longer with 3 (one subterminal, 2 terminal) setae (vs. shorter and 2 terminal setae in X. hydrophilus). In Medaeops granulosus, seta 3.5 is absent on the distal segment of the mxp2 giving 5 (2 subterminal, 3 terminal) setae (vs. present, 6: 3 subterminal, 3 terminal in Monodaeus couchii), the terminal endings of the dorsal and lateral carapace spines are swollen (vs. pointed in M. couchii), the two lateral spines on the telson are relatively small (vs. larger in M. couchii) and the posterior lateral pro- cess of the fifth abdominal somite is exceeding long, and swollen terminally (vs. short and pointed in M. couchii). These differences in larval morphology between Medaeops granulosus and Monodaeus couchii appear to support the subclade division for Eux 3. 4.2.3. Xanthinae, Speocarcininae, Panopeidae, and Pseudorhombilidae The Xanthinae, as presently understood, is the largest xanthid subfamily with 39 genera and 166 species (updated from Ng et al., 2008). The 15 genera and 29 species of Xanthinae sequenced for this analysis split into several major clades and lineages, confirming the polyphyly of this speciose group. Some genera were nested within other subfamilies, while other “xanthine” clades did not appear to be monophyletic. Of interest is the position of Xantho, the type genus of the subfamily and therefore Xanthinae sensu stricto. Xantho, does not appear to be closely affiliated with other xanthines and forms a separate clade (Xan 4) basal to a number of xanthid subfamily clades. As for the remaining xanthines, the majority of taxa tested grouped into four clades (Xan 1, 3, 4, 9) and their affiliations are supported by morphology. Xan 1: Demania, Neoxanthias, Liagore + zosimine Pulcratis, actaeine Odhneria. The characters that define this clade are the presence of distinct longitudinal rows of dense setae on the dactylus of all four ambulatory legs, a broad male thoracic sternite 7 (Fig. 7b), abdominal somites 1 and Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 431 Fig. 9. Gonopods. Comparative lengths of G1 and G2: (a) Euxanthus ruali Guinot, 1971, ZRC 2009.1178; (b) Alainodaeus rimatara Davie, 1993—(a, b) G1; (a1 , b1 ) G2. (c and d) Prominent crest on lateral margin of G1 (black arrow): (c) Monodaeus tuberculidens (Rathbun, 1911), (d) Crosnierius carinatus Serène and Vadon, 1981. Variation in G1 morphology: (e) Cymo andreossyi (Audouin, 1826), (f) Liomera tristis (Dana, 1852), (g) Actaea savignyi (H. Milne Edwards, 1834), (h) Paratergatis longimanus Sakai, 1965, (i) Chlorodiella cytherea (Dana, 1852), (j) Cycloxanthops truncatus (De Haan, 1837), (k) Leptodius exaratus (H. Milne Edwards, 1834), (l) Etisus laevimanus Randall, 1840, (m) Zozymodes xanthoides (Krauss, 1843), (n) Lybia tesselata (Latreille in Milbert, 1812), (o) Actaeodes tomentosus (H. Milne Edwards, 1834), (p) Nanocassiope alcocki (Rathbun, 1902), (q) Cataleptodius occidentalis (Stimpson, 1871), distal tip, (r) Xantho hydrophilus (Herbst, 1790), distal tip. Scale bars = 1.0 mm. a, a1 , after Davie (1993); c, after Guinot and Macpherson (1988); d, after Serène and Vadon (1981); e-p, after Serène (1984); q, r, after Guinot (1968). 2 broad (Fig. 8g); male thoracic sternite 8 just visible when abdomen is closed (Fig. 8g), episternites 5–7 delimited by a sulcus (Fig. 7b), and coxo-sternal condyle of P5 inserted between abdominal somites 2 and 3 (Fig. 8g). Demania, Liagore and Neoxanthias cluster with the supposed actaeine Odhneria and zosimine Pulcratis. Demania might not be monophyletic, clustering in two groups, with D. cultripes and D. scaberrima forming one group (regarded as Demania s. str. because the type species, D. splendida, is morphologically similar to the two exemplars), while D. intermedia appears to be sister to Odhneria. Zoeas for clade Xan 1 are not known. Author's personal copy 432 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Xan 2: Xanthias latifrons. This clade is only represented by X. latifrons and while not a true Xanthias species, it has several unusual sternal and other features (see also Edmondson, 1962). More Xanthias species and allied genera will need to be examined to confirm their affinities. Zoeas for clade Xan 1 are not known. Xan 3: Leptodius, Neoxanthops, Macromedaeus with the etisine Etisus electra (Eti 3) basal to the group. Morphological examination shows it to be a coherent group with distinct characters which include a relatively shallow sternoabdominal cavity without deep depressions at the level of sternite 4 (Fig. 7i), the male thoracic sternal suture 2/3 is more or less straight, the press-button is medial on sternite 5 equidistant from sutures 4/5 and 5/6, the front is produced when viewed ventrally, mxp3 quadrate, male thoracic episternites not delimited by a sulcus, and male thoracic sternite 7 not completely divided. Also notable about this group is the presence of fully or partially spoon-tipped chelae. First zoeas are known for Leptodius exaratus (see Clark and Paula, 2003), L. sanguineus and Macromedaeus crassimanus, and four zoeal stages are predicted for these taxa. Their first stage zoeas differ from X. hydrophilus (Xanthinae s. str.) in the following features: the rostral spine is terminally spinulate (vs. smooth), the antennal exopod has 1 terminal seta (vs. 2), the antennal exopod is proportionally longer (vs. smooth), the terminal setation of the distal endopod segment in mxp2 is 5 (2 subterminal, 3 terminal) setae with seta 3.5 absent (vs. 6: 3 subterminal, 3 terminal, seta 3.5 present). Xan 3 zoea differs from Lachnopodus subacutus (Xan 10) with respect to possessing an antennal exopod with 1 terminal seta (vs. 3: 1 subterminal, 2 terminal setae in L. subacutus). The first stage zoea of Cataleptodius floridanus, a genus that superficially resembles Leptodius, is known and although this species was not sampled for the genetic analysis, Ng et al. (2008) referred it to the Xanthinae. However, the zoea differs from that of X. hydrophilus in that the rostral spine is distally spinulate (vs. smooth) and the distal endopod segment of mxp2 comprises 5 (2 subterminal, 3 terminal) setae with setae 3.5 absent (vs. 6: 3 subterminal, 3 terminal, with setae 3.5 present). These characters suggest that C. floridanus is not a true xanthine as indicated by the present study. Cataleptodius may not even be closely allied to clade Xan 3 due to its different adult gonopods (Fig. 9q; also see Guinot, 1968a) and unpublished genetic data (B. Thoma and D. Felder, pers. comm.). Xan 4: Xantho. As discussed, a narrower definition of Xanthinae is necessary. The adult morphological characters that diagnose this taxon include the male thoracic sternal suture 2/3 being V-shaped (Fig. 7f), presence of deep depressions in the anterior sterno-abdominal cavity to accommodate the tips of the G1 structure (Fig. 7k), and a press-button that is positioned near or on sternal suture 5/6 (Fig. 7k). First zoeas are known for X. hydrophilus, X. pilipes and X. poressa, all with four zoeal stages (e.g., see Rodrìguez and Martin, 1997; Paula and dos Santos, 2001) and the first zoeal stage appendages for X. hydrophilus are described and illustrated (Figs. 2 and 3) in the present study. The first stage zoeas of these Xantho species differ from those of Xan 3 with respect to the antennal exopod possessing 2 terminal setae (Fig. 2c) (vs. 3: 1 subterminal, 2 terminal setae), the terminal setation of the distal endopod segment in mxp2 is 6 (3 subterminal, 3 terminal) setae, with seta 3.5 present (vs. 5: 2 subterminal, 3 terminal, seta 3.5 absent). The first stage zoea of X. pilipes differs from those of X. hydrophilus and X. poressa in that the rostral spine is spinulate (vs. smooth), the carapace lateral spines are dorsally spinulate (vs. smooth) and the antenna exopod is significantly longer (vs. proportionally smaller). Xantho poressa first zoea is distinct from X. hydrophilus and X. pilipes in that lateral processes are present on abdominal somites 4 and 5 (vs. absent), the posterior margin of abdominal somites 2–5 are spinulate (vs. absent) and the telson forks are spinulate (vs. smooth). Xan 5: Cycloxanthops, Pseudomedaeus. This is a clade of American taxa and few conclusions can be drawn because only two species were tested. Additional species of both genera will need to better test the integrity and/or homogeneity of this clade. Zoeas for clade Xan 5 are unknown. Xan 6–8: These xanthid clades are all part of a distinct group of wholly American/Atlantic taxa. Garthiope spinipes (Xan 6), Eucratodes agassizi, Micropanope sculptipes (Xan 7) and a new genus near Garthiope (Xan 8). Micropanope sculptipes and E. agassizi (Xan 7) are sister taxa while the new genus (Xan 8) is basal to a clade comprising xanthids, speocarcinines, panopeids and pseudorhombilids. The present analyses also indicate that Speocarcinus (Spe) is a sister clade to the Pseudorhombilidae (Pse) (represented by Trapezioplax tridentata) and supports the proposed assignment of the Speocarcininae into the Pseudorhombilidae on the basis of the coxo-sternal condition of the male penis (D. Guinot, M. Tavares and P. Castro, pers. comm.). Therefore, it is more parsimonious to recognise the Pseudorhombilidae as a subfamily within the Xanthidae, with Speocarcininae as a junior synonym. The Panopeidae also form a monophyletic clade (Pan) within this American group, suggesting that it should be recognised as a subfamily within Xanthidae, as Panopeinae. These results corroborate those of Thoma et al. (2009), based on similar data. Xan 7 first stage zoeas were not available for the present study. However, the first zoeas of Garthiope barbadensis, Hexapanopeus paulensis and Panopeus occidentalis are known, and all have a predicted four zoeal stages with distinctive characters. The first zoea of G. barbadensis differs from X. hydrophilus in that the rostral spine is terminally spinulate (vs. smooth), the carapace lateral spines are dorsally spinulate (vs. smooth), the antennal exopod a proportionally long (vs. small), the antennal exopod has 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal) and the second lateral telson spine is small (vs. longer). The first zoea of H. paulensis is distinct in that rostal and dorsal spine is long (vs. much shorter in X. hydrophilus, G. barbadensis, P. occidentalis), the antennal protopod is smooth (vs. multispinulate in X. Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 hydrophilus, G. barbadensis, P. occidentalis), the antennal protopod is relatively long (vs. much shorter in X. hydrophilus, G. barbadensis, P. occidentalis), the antennal exopod is relative minute with 1 terminal seta (vs. relatively large in X. hydrophilus with 2 terminal setae, G. barbadensis with 3: 1 subterminal, 2 terminal setae), the antennal endopod spine is absent (vs. present in X. hydrophilus, G. barbadensis), the mxp2 for which the terminal setation of the distal endopod segment is 5 (2 subterminal, 3 terminal) setae with seta 3.5 absent (Fig. 5a) (vs. 6: 3 subterminal, 3 terminal setae with seta 3.5 present in X. hydrophilus, G. barbadensis), and the telson is without lateral and medial spinulation (vs. present in X. hydrophilus, G. barbadensis). The first zoea of Panopeus occidentalis is distinctive in that the antennal exopod has 1 terminal seta (vs. 2 in X. hydrophilus, 3 in G. barbadensis) and in mxp2, the terminal setation of the distal endopod article is 5 (2 subterminal, 3 terminal) setae with seta 3.5 absent (vs. 6: 3 subterminal, 3 terminal setae with seta 3.5 present in X. hydrophilus, G. barbadensis (Fig. 5a). The zoeas of P. occidentalis and H. paulensis appear to support separate clades for the two genera. Although the first stage zoea of G. barbadensis is distinct, this species was not sampled for DNA analysis and therefore the taxon cannot be attributed to clades Xan 6 or 8. Xan 9: Nanocassiope. Based on the present molecular analyses, Nanocassiope is an isolated clade and its unique position is corroborated by morphological characters (Guinot, 1967a: 355, figs. 8, 9, 12–14; Davie, 1995: 202, 205, figs. 1 and 2). A suite of characters defines this taxon: the chelipeds are strongly unequal, there is a distinct gap between epistome and superior margin of mxp3 merus, the male thoracic sternites are relatively broad, the press-button is closer to sternal sutures 5/6 (Fig. 7j, k), a minute section of thoracic sternite 8 is visible when the abdomen is closed (Fig. 8g, h), the male abdomen is relatively short with the distal tip of the telson posterior to level of coxo-sternal condyle of P1 (Fig. 7d–f), and the G1 is disproportionately large and stout for the relatively small sterno-abdominal cavity, with stout setae distally (Fig. 9p). First zoeas are known for two species, N. granulipes (possibly a junior synonym of N. alcocki (Rathbun, 1902), see Takeda, 1976: 87) and N. melanodactylus, both with four stages (Ko and Clark, 2002, Dornelas et al., 2004). The zoea of N. granulipes is superficially similar to X. hydrophilus but differs with respect to the antennal exopod possessing 1 terminal seta (vs. 2 terminal setae) and the mxp2 for which the terminal setation of the distal endopod segment possesses 5 (2 subterminal, 3 terminal) setae with 3.5 absent (vs. 6: 3 subterminal, 3 terminal with setae 3.5 present) (Ko and Clark, 2002). The larva of N. melanodactylus, the type species of the genus, however, is markedly distinct, differing from N. granulipes in that its carapace lateral spines are dorsally spinulate (vs. smooth), the antennal exopod possessing 3 (1 subterminal, 2 terminal) setae (vs. 1 terminal), the terminal setation of the distal endopod in mxp2, segment is 6 (3 subterminal, 3 terminal) setae with seta 3.5 present (vs. 5: 2 subterminal, 3 433 terminal, seta 3.5 absent) (Dornelas et al., 2004). These zoeal differences between the two species suggest that Nanocassiope is not monophyletic. Xan 10: Lachnopodus, Paraxanthias, Metaxanthops + euxanthine Medaeus. The traditional xanthine taxa Lachnopodus, Paraxanthias, and Metaxanthops clustered with the euxanthine genus Medaeus in a basal clade. Even so, there are few characters to separate this clade from the rest of the Xanthidae although the structures of the G1 and G2 as well as the sterno-abdominal cavity appear to be useful. The morphological and genetic characters that diagnose this clade will need further study in order to understand the relationships of the currently included taxa. Only the first zoea of Lachnopodus subacutus is known and predicted to have four zoeal stages. Its first zoea differs from those of X. hydrophilus in that the antennal exopod possesses 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal) and the mxp2 for which the terminal setation of the distal endopod segment is 5 (2 subterminal, 3 terminal) setae with setae 3.5 being absent (vs. 6: 3 subterminal, 3 terminal, seta 3.5 present). It differs from those of Xan 3 with respect to possessing an antennal exopod with 3 terminal setae (vs. 1). The significance of these differences is apparent, as the alignment of Lachnopodus, Paraxanthias and Metaxanthops with Medaeus has not been previously proposed. 4.2.4. Chlorodiellinae Ng et al. (2008) listed eight genera and 47 species for this taxon and four genera and 12 species were analysed. They formed two unrelated clades, suggesting that this subfamily as currently understood is not monophyletic. Chl 1 is considered to be Chlorodiellinae s. str., containing Chlorodiella, the type genus and its type species, C. nigra (Forskål, 1775). Morphological examination revealed a suite of characters that may be used to define the subfamily. Garthiella, together with the euxanthine Medaeus danielita, forms a second chlorodielline clade (Chl 2). There are morphological similarities between Garthiella spp. and M. danielita and this clade is likely to be natural. Chl 1: Chlorodiella, Cyclodius, Pilodius. The characters common to these taxa include front not produced (particularly in a ventral view), merus of mxp3 quadrate, episternites not delimited by a sulcus, thoracic sternites 2 and 3 flat, thoracic sternite 7 not divided by coxo-sternal condyle of P4, which is at a distance from the abdomen, male abdominal somite 3 does not form a locking system with episternite 7, and the tip of the male telson is level with the coxo-sternal condyle of P1. The traditional features of this subfamily listed in Serène (1984), viz. spoon-tipped fingers of the chelipeds and dactylo-propodal locks on the ambulatory legs, are also found in other xanthid groups (e.g., Etisinae). Serène (1984) distinguished Chlorodiellinae from Etisinae primarily by the presence (in the latter) or absence or reduction (in the former) of a strong antero-lateral lobe on the basal article of the antenna, which occludes the orbital hiatus. Author's personal copy 434 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 The first zoeas of Chlorodiella nigra, Cyclodius obscurus (as C. monticulosus), Pilodius areolatus, P. paumotensis and P. pugil are known (Ng and Clark, 2000) and the morphology of the first stage zoea suggests all have four zoeal stages. They are identical and differ from X. hydrophilus only in the possession of minute spines on the dorsal carapace spine (vs. smooth), and a distally spinulate rostral spine (vs. smooth). The antennal exopod in Cyclodius obscurus, P. areolatus, P. paumotensis and P. pugil is distinctly proportionately longer than that of X. hydrophilus. The first zoeas are similar to those of Actaeodes (Act 3) except the small antennal exopod with 2 terminal setae (vs. 1 terminal or 3: 1 subterminal, 2 terminal) and telson with a slightly larger second lateral spine (vs. respectively smaller). Chl 2: Garthiella + euxanthine Medaeus danielita. Medaeus danielita differs from Medaeus s. str. (type species M. ornatus) in having a poorly produced front and a relatively shorter, stouter and less curved G1, and may not be a true Medaeus. These features are also seen in Garthiella, which has been classified under Chlorodiellinae (see Ng et al., 2008). However, Garthiella must be excluded from this subfamily as it does not possess the diagnostic features of the subfamily including the spoon-tipped chelae, the ambulatory dactylo-propodal lock, and the subterminal calcareous spine on the flexor margin of the ambulatory dactyli (Serène, 1984; Mendoza and Manuel-Santos, in press). Furthermore, Chl 2 is positioned “low” in the tree and is well separated from Chlorodiellinae s. str. (Chl 1). Zoeas of Chl 2 are not known. male thoracic sternal and male abdominal characters and is probably monophyletic. The first stage zoeas are known for Etisus anaglyptus and E. utilis, and four zoeal stages are predicted. They differ from X. hydrophilus in that the rostral spine is spinulate (vs. smooth) and the antennal exopod is proportionately slightly longer (vs. smaller). Etisus first stage zoeas may be distinguished from those of Chl 1 in that the carapace dorsal spine is without minute spines (vs. present), in all other respects the zoeas are identical. Eti 3: Etisus electra. This is a distinct clade from Eti 1 and 2, and it is supported by a number of distinct male thoracic sternal and carapace features that may substantiate the recognition of a long-synonymised genus, Etisodes. The type species of Etisodes, Etisus frontalis, is morphologically similar to E. electra (Eti 3) and the two are likely to be congeneric. The first stage zoea for a closely allied species, E. frontalis (not included in the current molecular analyses) is known and four zoeal stages are predicted. It shares the same differences as with the earlier discussed zoeas of E. anaglyptus and E. utilis from X. hydrophilus and chlorodiellines of clade Chl 1. However, E. frontalis differs from E. anaglyptus and E. utilis by possessing a mxp2 for which the terminal setation of the distal endopod segment is 6 (3 subterminal, one of which 3.5 is minute, 3 terminal) setae (vs. 6: 3 subterminal, seta 3.5 is prominent, 3 terminal) setae. If this character is consistent with the first stage zoea of E. electra, this larval evidence may support the validity of Etisodes. 4.2.5. Etisinae Two genera and 25 species are currently assigned to the Etisinae, of which seven species of Etisus were analysed. The results from the present molecular and morphological analyses suggest that this subfamily requires extensive revision (see Kraussiinae and Xanthinae). The main Etisus clade (Eti 2) is considered to be Etisus s. str. as it contains E. dentatus, the type species. Etisus villosus, from its position in the tree may be included in Etisus s. str. (Eti 2) but has a combination of characters that distinguishes it from congeners. Etisus electra is in its own clade (Eti 3) and has a suite of morphological characters that suggest it should be assigned to a different genus. Having examined adult E. electra specimens, numerous characters were found differentiating it from Etisus s. str. On the other hand, Etisus odhneri forms a separate clade (Eti 1) that appears to be basal to the Chlorodiellinae, but statistical support for this relationship is weak. Nonetheless, there are morphological characters justifying the exclusion of E. odhneri from Etisus sensu stricto. Eti 1: Etisus odhneri. This unusual species has a completely different male thoracic sternum as well as other characters that separate it from Etisus s. str. (Eti 2). It may need to be referred to its own genus. Zoea of Etisus odhneri are not known. Eti 2: Etisus anaglyptus, E. dentatus, E. utilis. This is a distinct group united by a strong suite of carapace, antennal, 4.2.6. Kraussiinae Of the three genera and 17 species currently assigned to Kraussiinae, only two genera and three species were sequenced. The results of the present molecular analysis suggest that Kraussiinae is polyphyletic, with Kraussia (Kra 2) affiliated with Etisinae s. str. (Eti 2), and Palapedia (Kra 1) forming its own clade. There is, however, a suite of strong characters that separate Kraussiinae s. str. (Kra 2) and Etisinae s. str. (Eti 2). These characters are so marked that for over a century, Kraussia and its allies were assigned to the Atelecylidae and other families, before Ng (1993a) argued they were highly modified xanthids and proposed a new subfamily for Kraussia, Palapedia and Garthasia. Although Palapedia is superficially similar to Kraussia (see Ng, 1993a), a number of characters have been observed that argue against them being in the same clade so the tree may prove to be correct. However, these three genera share one character absent in all other known xanthids (with the exception of those now in Xan 10 and Chl 2) in that the G1, while long, just reaches suture 4/5 and sternite 4 does not contribute to the sterno-abdominal cavity. In all other xanthine subfamilies (with the exception of those in Xan 10 and Chl 2), the posterior region of sternite 4 contributes to the sterno-abdominal cavity, and the G1 reaches up to this part of sternite 4. This condition in Kraussia, Palapedia and Garthasia may merely be a consequence of their burrowing habits, which has resulted in a relatively elongated carapace and thoracic sternum (Ng, 1993a). Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Kra 1: Palapedia. Although the carapace, pereiopods and thoracic sternum of Palapedia and Kraussia are both superficially similar, the posterior margin of the branchiostegite is markedly serrated in Palapedia (gently sinuous in Kraussia), the pleurites can barely be seen outside the branchiostegite in Palapedia, with only a sliver of pleurite 7 visible (pleurites 5–7 are visible as distinct rectangular plates in Kraussia). These characters are significant and suggest that Palapedia may not be as closely related to Kraussia as had been believed. In addition, Palapedia has pointed fingers on the chelae whereas those of Kraussia are spoon-tipped (Ng, 1993a). The first zoeas of Palapedia integra (see Ko et al., 2004) and P. valentini (see Clark and Ng, 1998) are known and probably have four zoeal stages. Palapedia integra differs from X. hydrophilus in that the rostral spine is terminally spinulate (vs. smooth) and the second lateral telson spine is proportionally much shorter (vs. longer). Palapedia valentini first stage zoea differ from those of P. integra with respect to their carapace dorsal spine being much longer and with some minute spinulations (vs. absent), the antennal exopod with 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal setae), the telson fork with two small lateral spines (vs. both proportionately larger) and with a disitinctly large curved medial dorsal spine on the telson fork (vs. proportionately smaller). Kra 2: Kraussia. The differences with Palapedia have been discussed above; and the genetic markers suggest Kraussia is closer to members of Etisinae s. str. (Eti 2) than to Palapedia. Kraussia and Etisinae s. str. share a number of characters such as a serrated anterolateral margin of the carapace, similar male thoracic sternal structures and chelipeds, and it is not difficult to conceive Kraussia as a “longitudinally stretched” Etisus villosus or E. dentatus. Moreover, while Garthasia differs from Kraussia in the shape of the carapace and chelae, their branchiostegal margin is similar in form, indicating that they may be closely related (see Ng, 1993a: Table 2). Kraussia zoea are not known. 4.2.7. Liomerinae Six genera and 57 species are currently assigned to Liomerinae of which three genera and 12 species were included in the molecular analyses. Members of this subfamily are morphologically difficult to separate from most Actaeinae or Xanthinae sensu lato. Sakai (1976) established this subfamily mainly for taxa with a relatively broad, brightly coloured carapace and the propodal finger of the cheliped usually lacking a proximal or median cutting tooth. According to the present study, the Liomerinae as traditionally understood appears to be paraphyletic with the three genera and 10 species analysed clustering into two separate clades that are well supported statistically and morphologically. Lio 1 is considered to be Liomerinae s. str. as the type genus Liomera s. str. is in this clade. However, a revision of the subfamily is required because the zosimine genus Paratergatis and an undescribed genus are nested together with Liomera and Lipkemera while Lio 2 consists entirely of Neoliomera species. 435 A new subfamily may have to be established to accommodate Neoliomera. Lio 1: Liomera, Lipkemera + zosimine Paratergatis and an undescribed genus. This clade is supported by the following characters: absence of slits in male sternoabdominal cavity (at level of sternite 4) (Fig. 7i) (vs. present in Neoliomera, Fig. 7l), longitudinally narrow thoracic sternite 2 (Fig. 7c) (vs. well defined triangle in Neoliomera), dactylo-propodal lock on ambulatory legs absent (Fig. 6o) (vs. present in Neoliomera), and anterior part of thoracic sternite 1 bifurcated with two spines (vs. not bifurcated in Neoliomera). The analysis also indicates that Liomera is paraphyletic with the type species L. cinctimana basal to the other Liomera species. Liomera cinctimana was compared with specimens of L. tristis, the type species of Carpilodes and can be distinguished by a narrow thoracic sternite 7 (vs. broad), thoracic episternite 7 only slightly covering the P5 coxa (vs. covering a larger part of the coxa), male telson relatively longer (vs. relatively shorter), male abdominal somite 2 narrower (vs. broader) and male thoracic sternites 3 and 4 broader (vs. narrow). These morphological characters support the recognition of Carpilodes, and it includes most of the species placed in Liomera by Serène and Nguyen (1960) and Serène (1984). First zoeas are known for Liomera bella, L. cinctimana, L. laevis, and all probably have four stages. Liomera first stage zoeas are distinguished from those of X. hydrophilus by exhibiting a rostral spine with terminal spinulation (vs. smooth) and a proportionately longer (vs. small) antennal exopod with 3 (1 subterminal and 2 terminal) setae (vs. 2 terminal setae). Similar to what is seen in adult morphology and the current phylogenetic tree (Fig. 1), the zoeas of L. laevis (similar in adult morphology to L. tristis) differ markedly from the other two congeners in that they possess an exceedingly long and spinulate dorsal carapace spine, spinulation on the surface of the anterior carapace, setae on the ventral carapace margin, a long antennal protopod equal in length to the dorsal spine and proximally spinulate, abdominal somites 4 and 5 with lateral processes, abdominal somites 2 and 3 possess a pair of minute dorsal medial spines and the first lateral spine of the telson is markedly longer. Lio 2: Four Neoliomera species were analysed: N. demani, N. cerasinus, N. insularis and N. striata. The presence of supplementary slits/grooves in the male sterno-abdominal cavity (at level of thoracic sternite 4) (Fig. 7l) to accommodate the elaborate setation of the distal end of the G1 s (vs. absent in Liomerinae s. str.) is diagnostic of this group. Other characters include the fused male thoracic sternites 1 and 2 widely triangular and prominent (vs. narrow in Liomerinae s. str.), presence of a dactylo-propodal lock on the ambulatory legs (vs. absent in Liomerinae s. str.) and male thoracic sternite 1 ridge not bifurcated (vs. bifurcated with two spines in Liomerinae s. str.). Neoliomera zoeas are not known. 4.2.8. Zosiminae Nine genera and 69 species are currently assigned to this subfamily (Ng et al., 2008), characterised by exhibiting Author's personal copy 436 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 cristate anterolateral margins on the carapace and/or cristate pereiopods. However, the monophyly of Zosiminae has been questioned (see Serène, 1984; Ng et al., 2008) and as such, eight genera and 20 species were selected for inclusion in the present genetic study. These sequenced taxa clustered into two main clades with most genera being paraphyletic; e.g., included species of Atergatopsis form two distinct morphological groups within the two Zosiminae clades. The type species, A. signatus, while not sampled in the molecular study is morphologically similar to A. germaini and A. granulata, but both these species are not observed to be reciprocally monophyletic. The same is observed for Lophozozymus. The separation of L. anaglyptus in Zos 2 from L. pictor and L. edwardsi in Zos 1 suggests that the assignment of the former species from Platypodia to Lophozozymus by Ng et al. (2008) may not be justified. Therefore, the present study demonstrates that the cristate anterolateral margins and/or cristate pereiopods are not reliable subfamilial characters, with taxa previously assigned to the Zosiminae clustering into four clades: Zos 1, 2, Lio 1 and Xan 1. However, finding reliable characters to distinguish between the major zosimine clades proved difficult and those cited here might not be totally satisfactory. Only Zosimus appears to be monophyletic, but this taxon too may become problematical when additional species are analysed. Furthermore, Zos 2 contains the type species Zosimus aeneus and this clade must therefore be considered as Zosiminae sensu stricto. Zos 1: Comprises Atergatopsis aff. amoyensis, A. aff. obesa, Lophozozymus pictor, L. edwardsi + the xanthines Xanthias canaliculatus, X. lamarckii, X. punctatus. Only one character appears to define this cluster, the tip of the telson level with or posterior to (as in X. punctatus) the coxo-sternal condyle of P1. The first zoea of Zos 1 is known only for Lophozozymus pictor which has four stages (Clark and Ng, 1998). The first zoea differ from that of X. hydrophilus in that the carapace lateral spines are minutely spinulate along the dorsal margin, the antennal protopod is smooth (vs. multispinulate), the antennal exopod is proportionately longer (vs. relatively short), the setae of the expod comprises 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal setae) and the two lateral spines on the margin of the telson are minute (vs. relatively distinct). Zos 2: This clade includes the type species Zosimus aeneus (therefore Zosiminae s. str.) and comprises Atergatis obtusus, A. integerrimus, A. interruptus, A. floridus, Atergatopsis germaini, A. granulatus, Lophozozymus anaglyptus, Platypodia, Zosimus. Only the tip of the male telson reaching beyond the level of coxo-sternal condyle of P1 appears to define this clade. Some genera (e.g., Zosimus) also have dactylo-propodal locks on their ambulatory legs (Fig. 6m). The molecular analysis indicates that Atergatopsis A. Milne-Edwards, 1862, is polyphyletic. Atergatopsis s. str. forms a discrete morphological group with the type species, Atergatopsis signata (Adams and White, 1849) (not included in the phylogeny), and the two included species, A. germaini and A. granulata are clustered separate from a second group including Atergatopsis aff. amoyensis and A. aff. obesa. Lophozozymus A. Milne-Edwards, 1863, is also polyphyletic, as L. pictor (senior synonym of Xantho octodentatus H. Milne Edwards, 1834, type species) clusters with L. edwardsi, but is separate from L. anaglyptus. The latter species clusters with Atergatopsis s. str., Atergatis, Platypodia and Zosimus. This suggests that the re-assignment of L. anaglyptus from Platypodia to Lophozozymus by Ng et al. (2008) is unwarranted. However, finding reliable characters that distinguish between the clades Zos 1and Zos 2 proved difficult. First zoeas are known for Atergatis floridus (see Clark and Paula, 2003), Atergatopsis germaini, Platypodia eydouxi (see Clark et al., 2004), P. spectabilis and Zozymodes xanthoides (see Clark and Paula, 2003), all of which are believed to have four zoeal stages. The first stage zoeas of Zos 2 differ from that of Xantho hydrophilus in that the rostal spine is terminally spinulate (vs. smooth), the antennal exopod is proportionately much longer (vs. short) and the antennal exopod has 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal setae). First stage zoeas of Zos 2 differ from L. pictor (Zos 1) in that the carapace lateral spines are smooth (vs. minutely spinulate along its dorsal margin), rostral spine is distally spinulate (vs. smooth), the antennal protopod is multispinulate (vs. smooth) and the 2 lateral spines on the telson are distinct (vs. minute). While no sequence data are available, the zoeas of Atergatis subdentatus, Platypodiella spectabilis and Zozymodes xanthoides compare well with those described for this clade. 4.2.9. Cymoinae Ng et al. (2008) assigned one genus and eight species to this subfamily. Two species were sequenced for the present analysis, and Cymoinae appears to be monophyletic (Cym). Cymoinae is the sister group to the monophyletic Polydectinae (Pol). The suite of adult morphological characters that has been used to define this subfamily by Serène (1984) appears to be diagnostic. First zoeas are known for Cymo lanatopodus and C. melanodactylus, with the characters indicating that the development comprises four zoeal stages. The first zoeas are similar to those of X. hydrophilus except that the rostral spine is terminally spinulate, the antennal exopod is slightly longer (vs. shorter) and has 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal setae). These first stage zoeas are difficult to distinguish from those of Atergatis floridus (see Clark and Paula, 2003), Atergatopsis germaini, Platypodia eydouxi (see Clark et al., 2004), P. spectabilis and Zozymodes xanthoides (see Clark and Paula, 2003). 4.2.10. Polydectinae Two genera and 11 species are currently assigned to this subfamily, with both genera and five species sampled for the present analysis and these clustered together into one clade (Pol), with Cymoinae (Cym) as its sister group. The suite of Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 characters established by Guinot (1976) and Serène (1984) to define the Polydectinae remain diagnostic. However, the relationship between Lybia and Polydectus will need to be further assessed as P. cupulifer is shown to be basal to L. hatagumoana and L. leptochelis while L. edmondsoni and L. tessallata are sister species. The first zoea is known for Lybia plumosa (see Clark and Paula, 2003) and is predicted to have four stages. This zoea is similar to that of X. hydrophilus except that the rostral spine is distally spinulate (vs. smooth), the antennal exopod has 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal) and the mxp2 with the terminal setation of the distal endopod segment is 5 (2 subterminal, 3 terminal) setae with seta 3.5 absent (vs. 6: 3 subterminal, 3 terminal with seta 3.5 present). 4.2.11. Zalasiinae Zalasiinae currently comprises three genera and 23 species of which three genera and four species were analysed. They form a monophyletic group (Zal) with the inclusion of the actaeine, Pseudactaea corallina. A number of diagnostic characters define the subfamily including the cutting edges of the cheliped fingers being shear-like and the tips cross when closed (Guinot, 1976: pl. 6 figs. 5, 6, 10; pl. 7 figs. 9, 10), distinct pereiopod coaptation with the carapace (Fig. 6d–f), thoracic sternites 1 and 2 are large and fused (Fig. 7d), relatively wide slits separating the central portion of epistome from lateral portions (Fig. 6f), and the tips of the G1 lodged in deep slits in the anterior sterno-abdominal cavity and slightly produced beyond the abdomen/telson (Fig. 7d, g) (see also Guinot, 1976: figs. 25, 27, 28). The results of the present analyses corroborate these adult features. Guinot (1976) assigned Banareia and Calvactaea to the subfamily originally established by Serène (1968) for Zalasius only. Traditionally (e.g., Odhner, 1925), Banareia and Calvactaea have been allied with the Actaeinae. Guinot (1976), however, showed that there was a continuity of characters that connects the various Banareia species with Zalasius, arguing that they form a monophyletic assemblage. This hypothesis is confirmed by the present results. In addition, a typical zalasiine character was considered to be the hirsute carapace, usually with long and dense setae covering the entire surface (Banareia and Zalasius) or with short velvety setae as in Calvactaea. Pseudactaea appears to be a miniature “denuded” zalasiine and, other than hirsuteness, possesses all the other diagnostic characters of the subfamily. The first stage zoeas are known for Banareia subglobosa but tissues of this species were not sampled in the molecular analysis. When compared with Xantho hydrophilus first stage zoea, B. subglobosa possesses a proportionately longer antennal expopod (vs. shorter) with 3 (1 subterminal, 2 terminal) setae (vs. 2 terminal), mxp2 with a distal endopod segment comprising 5 (2 subterminal, 3 terminal) setae with seta 3.5 absent (vs. 6: 3 subterminal, 3 terminal with seta 3.5 present) and a minute second lateral spine on the telson fork that could almost be described as a seta. 437 5. Conclusions The present study uses genetic datasets to establish a basephylogenetic tree for the Xanthidae and the results suggest that a major revision of the family is required. It confirms earlier predictions that many (7 out of 13) of the subfamilies are para- or polyphyletic. In addition, they demonstrate that traditional characters based on general appearances of the carapace, legs, and chelipeds are unreliable phylogenetic indicators. Instead, the veracity of the newly identified clades was tested against new adult and larval characters. Relationships implied by non-traditional “ventral” features in adults, such as thoracic sternal and gonopodal characters as originally proposed by Guinot (1967a,b,c, 1968a,b,c, 1969a,b,c,d, 1971, 1976, 1977a,b, 1978, 1979) proved to be more consistent with molecular results than traditional “dorsal” characters. First zoeal characters proved to be selectively valuable in supporting a number of clades. However, the limitation of these data is that the first zoeas were examined for only 48 out the 161 species analysed in the present study. Consequently, the phylogenetic significance of some characters such as size of lateral telson spines, varying between large to minute or absent, is difficult to resolve. In addition, the evolutionary significance of species possessing a lateral carapace which is dorsally spinulate, is unclear. In any event, larval data remain valuable in phylogenetic inference, when deployed in conjunction with adult morphology and molecular data. One fundamental change to the current systematics is the assignment of taxa to the Xanthinae. From our findings, only the Atlantic genus Xantho appears to belong to this subfamily (Xan 4). Furthermore, within this genus, the zoeas of X. hydrophilus, X. pilipes and X. poressa are distinct compared with those of chlorodielline genera which appear to have similar, if not identical setotaxy. Most xanthines clustered into a “Leptodius” group (Xan 3) with others scattered throughout the tree in 10 widely dispersed clades: Xanthias (Xan 2, Zos 1, Eux 3), Demania/Liagore (Xan 1), Garthiope (Xan 6), Nanocassiope (Xan 9), Euryxanthops (Eux 3) and Lachnopodus (Xan 10). More importantly, this segregation of the Xanthinae based on genetic markers is well supported by adult and some zoeal characters. The discovery that Xanthias is polyphyletic comes as no surprise as numerous authors, including Serène (1984), had previously alluded to this. Furthermore, the few Xanthias species analysed here are assigned to three separate clades (Xan 2, Zos 1, Eux 3), and these can be supported by adult morphology. Euryxanthops (Eux 3) too, possesses “xanthine” carapace characters, but has the thoracic sternum, abdomen and gonopods typical of antrocarcinines. This is strongly supported by the genetic data presented in Figure 1. Interestingly, Demania and Liagore (Xan 1) can be accomodated in the Liagorinae Števčić, 2005, although the revised diagnostic characters presented here are completely different from that originally proposed by Števčić (2005) which was based only on “dorsal” adult morphology (see also Ng and Chen, 2004). There is also strong adult and Author's personal copy 438 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 larval support for the isolated lineages (Xan 9) of Nanocassiope and Garthiope (Xan 6); both may eventually warrant subfamily or even family status. Nanocassiope is currently being studied by some of the present authors, while Garthiope is undergoing revision by B. Thoma and D. Felder (pers. comm.). The Actaeinae, Etisinae, Euxanthinae, Kraussiinae, Liomerinae, and Zosiminae are not monophyletic as they have been constituted in Ng et al. (2008) and will require extensive revision. The establishment of new subfamilial taxa will be necessary for at least five clades, comprising Paractaea-Gaillardiellus (Act 2), Actaeodes (Act 3), Neoliomera (Lio 2), Lophozozymus (Zos 1), and Garthiella (Chl 2). The support from adult and larval morphological characters is especially compelling for Act 2, Act 3 and Lio 2. Further, the consistent molecular dichotomy between the two kraussiines, Palapedia (Kra 1) and Kraussia (Kra 2) will also need a systematic review as the thoracic sternal characters suggest that the two genera resemble each other only because both are burrowers in fine reef sand. The complex situation with the Euxanthinae merits comment. Davie (1993, 1997, 2002), Ng (1993b, 2002) and Ng et al. (2008) have suggested that the Euxanthinae may be paraor polyphyletic and may not be a good taxon. The molecular data confirm Euxanthinae s. str. (Eux 1), although this subfamily should be restricted to genera such as Euxanthus and Hypocolpus, and surprisingly, also including one actaeine, Psaumis. However, the majority of other euxanthines cluster in a separate large clade (Eux 3) that also contains members of a small aberrant subfamily, the Antrocarcininae. This lineage exhibits considerable carapace diversity but all its members have the same kind of sternum, male abdomen and gonopod. These diagnostic characters consistently distinguish Eux 3 from the Euxanthinae s. str. (Eux 1) and are strongly supported by the molecular dataset. This study restricts and reinterprets the Euxanthinae with a fresh set of characters, while substantially redefining the Antrocarcininae to accommodate other taxa including two xanthines, Xanthias teres and Euryxanthops dorsiconvexus. One euxanthine, Glyptoxanthus (Eux 2), possesses a suite of remarkable adult characters that merit establishment of a separate subfamily for this genus (Mendoza and Guinot, in press). This change in status for the monotypic taxon is strongly corroborated by the DNA evidence presented here. The Etisinae also requires revision. While Etisus is polyphyletic, the true relationships of species excluded from the core group, such as E. odhneri (Eti 1), will need further clarification. The revision of this polyphletic genus is currently being undertaken by J.C.E. Mendoza and B. Thoma (pers. comm.). A few subfamilies are well supported including Zalasiinae, Polydectinae, Cymoinae, and, with the exclusion of Garthiella, Chlorodiellnae. The Zalasiinae is a good example of our multi-disciplinary approach to the present study. The carapace shape of its constituent genera are divergent whereas molecular analyses strongly support the morpho- logical rationale by Guinot (1976) for recognising them as a coherent group. The value of her characters, and their use in the present study, is underlined by the discovery that an actaeine genus, Pseudactaea surprisingly clustered in the Zalasiinae, but indeed has all the diagnostic sternal, abdominal and gonopodal characters of the subfamily. The fact that the carapace of Pseudactaea is nevertheless typically “actaeine” and its placement in the subfamily has never been questioned, only underscores the futility of relying on such characters for a phylogenetic classification. Some clades are wholly unexpected and emphasise the complexity of xanthid evolution. Clade Xan 10 contains three traditional xanthines, Paraxanthias, Lachnopodus and Metaxanthops as well as three euxanthine species of Medaeus. The larval data here are not helpful, and the adult morphological differences are relatively subtle (mainly in the thoracic sternum and abdomen). This lack of differentiation may be due to the basal position of the clade, with its constituent taxa exhibiting many plesiomorphic characters. The clade that comprises Garthiella also contains one species of Medaeus (Chl 2). While the available morphological features suggest that the two clades may be natural, evidence presented here is insufficient to justify recognising them as distinct subfamilies, at least for the time being. 5.1. Summary The present study highlights numerous problems associated with current xanthid systematics and nomenclature. Most traditional characters are considered convergent and not phylogenetically informative at subfamilial level, although they are diagnostic at species and generic rank. In comparison, the usually neglected ventral characters as originally proposed by Guinot (1967a,b,c, 1968a,b,c, 1969a,b,c,d, 1971, 1976, 1977a,b, 1978, 1979) including sternal and gonopodal morphology, appear to be more reliable as diagnostic characters and more phylogenetically informative at this level. Zoeal characters remain phylogenetically valuable, especially when used in conjunction with other evidence such as DNA sequences and adult morphology. The molecular xanthid tree presented here confirms that 7 out of 13 xanthid subfamilies as currently understood (see Ng et al., 2008) are para- or polyphyletic. The Xanthidae require a radical systematic revision, which may necessitate a number of nomenclatural changes, including the erection of new higher subfamily level taxa. Many existing groups will need substantial redefinition. Finally, the current systematic study was largely restricted to Indo-Pacific taxa with only a few Atlantic species included. Therefore a larger study including American Xanthidae, as well an even greater coverage of Indo-Pacific taxa is required, before the complex systematic relationships within the Xanthidae and Xanthoidea can be resolved. Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Acknowledgements This study was funded jointly by a European Distributed Institute of Taxonomy Integrating Research Grant, second call, to encourage increased collaboration between taxonomic researchers (awarded November 2009) and the Raffles Museum of Biodiversity Research, National University of Singapore. Joelle Lai would like to thank Jacqueline Mackenzie-Dodds, manager of the Wolfson Wellcome Biomedical Laboratories; Julia Llewellyn-Hughes, manager of the sequencing facility; and Peter Foster, manager of the Molecular Biology Computing Facility, all Department of Zoology, Natural History Museum, London, for all their support and help during her visit in January 2010. JC Mendoza wishes to thank the following for their kind assistance during museum visits and for facilitation of specimen loans: Régis Cleva (MNHN), Marivene Manuel-Santos (NMCR), Jody Martin, Regina Wetzer, Dean Pentcheff, Kathy Omura (LACM), Rafael Lemaitre, Karen Reed, Chad Walter (USNM), Gustav Paulay, Arthur Anker, John Slapcinsky, Sandy Beemis (UF), Charles Fransen, Jacques Smit (RMNH), Hironori Komatsu (NMST), Hiroshi Namikawa (KPMNH), Tohru Naruse (RUMF), Michael Türkay (SFM), Peter Davie (QM), Shane Ahyong (AMS) and Tan Siong Kiat (ZRC). Danièle Guinot and Paul Clark would to thank all the curators and Greasi Simon of the Raffles Museum for hosting their visit in March 2010. Paul Clark gratefully acknowledges the following grants and support while collecting ovigerous xanthids and panopeids: a Smithsonian Short term Visitor grant to the Smithsonian Marine Station at Link Port, Fort Pierce Florida (via Ray Manning); the Albion Fisheries Research Centre, Ministry of Fisheries and Marine Resources, Petite Rivière, Albion, Mauritius, and the Smithsonian Institution for logistical support and providing SCUBA cylinders; support from European research project, INCO-DC no. IC18-CT96-0127, “Macrobenthos of Eastern African mangroves: life cycles and reproductive biology of exploited species” to work on Inhaca Island, Mozambique, and the Estação de Biologia Marítima da Inhaca and for funds to attend the 7th Colloquium Crustacea Decapoda Mediterranea in Lisbon (via José Paula); a fellowship from the Conservation Fund of the National University of Singapore and Nanyang Technological University, Singapore, and the Research Fund from the Raffles Museum, National University of Singapore (via Peter Ng); two Visiting Scientist grants from the Muséum national d’Histoire naturelle, Paris (via Alain Crosnier); British Council, Jeddah, for providing finance for travel, the Dean of the Faculty of Marine Sciences, King Abdulaziz University for funding a stay in Jeddah; British Airways for logistical support; the Zoology Research Fund, Zoology Department and an Enhancement Grant, both from The Natural History Museum, London. We are indebted to John Bishop, Marine Biological Association, England and Dave McGrath, Galway Mayo Institute of Technology, Ireland, for collecting specimens of Xantho hydrophilus and X. pilipes, respectively and making them available for this study. 439 Appendix A. Adult material of Xanthoidea examined XANTHIDAE MacLeay, 1838. Actaeinae Alcock, 1898: Actaea calculosa (H. Milne Edwards, 1834), 1 male, coll. May 1934, reef, Lindeman Is., Queensland, Australia, ZRC 1965.761–6; Actaea polyacantha (Heller, 1861), 2 males, coll. 21 May 1985, Chatan Village, Sunabe, Okinawa, Japan, ZRC 1999.0261; Actaea semblatae Guinot, 1976, 1 male, coll. S.H. Fan, Aug. 1996, Sa Yu Harbour, Fujian Province, China, ZRC 2002.0577; Actaeodes hirsutissimus (Rüppell, 1830), 2 males, coll. L.W.H. Tan, 1986, Sentosa Reef, Singapore, ZRC 1987.551–552; 1 ovigerous female, coll.T. Tan, May 2000, Pulau Bintan, Indonesia, ZRC 2008.1143; Actaeodes mutatus Guinot, 1976, 1 male, 1 female, coll. B. Goh, 16 Oct. 1986, on Pavona coral, Raffles Lighthouse, Singapore, ZRC 1993.0287–0288; Actaeodes tomentosus (H. Milne Edwards, 1834), 1 male, coll. PANGLAO 2004, Jun. 2004, stn M1, Panglao Is., Philippines, ZRC; 3 males, 1 female, coll. C.D. Schubart, 25 Jun. 2000, Pulau Sapi, Sabah, Malaysia, ZRC 2000.1673; Epiactaea nodulosa (White, 1848), 7 males, 2 females, coll. D.G.B. Chia and T.L. Koh, Jan. 1992, Sentosa Is., Singapore, ZRC 1995.0333; Gaillardiellus orientalis (Odhner, 1925), 1 male, coll. C.M. Yang et al., 22 Jul. 1997, Pulau Seringat, Singapore, ZRC 2000.1196; Gaillardiellus rueppelli (Krauss, 1843), 1 male, coll. PANGLAO 2004, Jun. 2004, Bohol Is., Philippines, ZRC 2010.0162; Lobiactaea lobipes (Odhner, 1925), 1 male, coll. P.K.L. Ng et al., Jul. 2003, tangle net, Balicasag Is., Philippines, ZRC; Novactaea bella Guinot, 1976, 2 males, coll. P.K.L. Ng et al., May 1993, Lagoi, Pulau Bintan, Indonesia, ZRC 1998.0981; Odhneria echinus (Alcock, 1898), 1 male, coll. SANTO 2006, Sep. 2006, NW coast of Malo Is., Vanuatu, ZRC 2009.1177; Paractaea rufopunctata (H. Milne Edwards, 1834), 1 male, coll. PANGLAO 2004, Feb. 2004, stn PN1, tangle net, Balicasag Is., Philippines, ZRC; 1 male, coll. L. Kirkendale, 30 Apr. 1999, rubble, 10 m, Pago Bay, Guam, ZRC 2000.0718; Psaumis cavipes (Dana, 1852), 1 male, 1 female, coll. PANGLAO 2004, 20 Jun. 2004, Psa-cav-1, stn B18, 3–5 m, Sungcolan Bay, Panglao Is., Philippines, ZRC 2010.0157; Pseudactaea corallina (Alcock, 1898), 1 male, coll. PANGLAO 2004, 4 Jul. 2004, stn T38, beam trawl, 80–140 m, Balicasag Is., Philippines, ZRC; 1 ovigerous female, coll. H.H. Tan and P.F. Clark, 27 Jul. 2003, near Balicasag Island Dive Resort, Balicasag Is., Philippines, ZRC 2003.0276; Pseudoliomera variolosa (Borradaile, 1902), 1 male, 1 female, coll. R. de Felice, 26 Jan. 2000, Moku Manu Is., north of Mokapu Peninsula, Oahu, Is., Hawaii, ZRC 2000.0508; Serenius kuekenthali (De Man, 1902), 1 male, 1 female, coll. PANGLAO 2004, 2 Jun. 2004, Surkue-2, stn B5, 4 m, Biking, Panglao Is., Philippines, ZRC 2010.0159; Serenius pilosus (A. Milne-Edwards, 1867), 1 female, coll. PANGLAO 2004, 10 Jun. 2004, Ser-pil-2, stn B10, 3–14 m, Momo Beach, Panglao Is., Philippines, ZRC 2010.0158. Author's personal copy 440 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Antrocarcininae Ng and Chia, 1994: Antrocarcinus petrosus Ng and Chia, 1994, paratype female, coll. B. Richer de Forges, 29 Jan. 1993, stn DW170, 241–244 m, New Caledonia, ZRC 1995.212; Cyrtocarcinus truncatus (Rathbun, 1906), 1 ex., Ouvéa, east of Mouly, New Caledonia, RUMFZC-01302 [tissue only]; Glyptocarcinus lophopus Takeda, 1973, 1 ex., east of Muko-jima, Japan, NSMT-SY-09-04 [tissue only]; Glyptocarcinus politus Ng and Chia, 1994, paratype female, coll. B. Richer de Forges, 28 Jan. 1993, stn DW156, 262–275 m, New Caledonia, ZRC 1995.213; 1 ex., Kaikata Seamount, Japan, NSMT KT-09-2 KK1-2(1) [tissue only]. Chlorodiellinae Ng and Holthuis, 2007: Chlorodiella barbata (Borradaile, 1900), 1 male, coll. A. Anker and R. Lasley, 17 Feb. 2009, reef flat, 1–2 m, Lizard Is., Queensland, Australia, UF 17105; Chlorodiella corallicola Miyake and Takeda, 1968, 1 male, coll. PANGLAO 2004, 25 Jun. 2004, stn B25, 16 m, Pamilacan Is., Philippines, ZRC 2010.0296; Chlorodiella cytherea (Dana, 1852), 1 male, 2 females, coll. H.H. Tan and P.F. Clark, 25 Jul. 2003, reef flat, Alona Beach, Panglao Is., Philippines, ZRC 2008.0644; Chlorodiella laevissima (Dana, 1852), 1 male, coll. S. Smith and K. Mills, 20 Feb. 2009, rubble, 25–30 m, Yonge Reef, Lizard Is., Queensland, Australia, UF 17262; Chlorodiella nigra (Forskål, 1775), 1 male, coll. V. Bonito et al., 24 Jan. 2005, under rocks, 1–3 m, Bar Al-Hikman Peninsula, Oman, UF 17948; Chlorodiella xishaensis Chen and Lan, 1978, 1 male, 1 female, coll. PANGLAO 2004, 23 Jun. 2004, stn P4, tangle net, Balicasag Is., Philippines, ZRC 2010.0289; Cyclodius ungulatus (H. Milne Edwards, 1834), 15 males, 18 females, coll., C.A. Gibson-Hill, 1941, Cocos Keeling Is., ZRC 1965.11.11.16–25; 3 males, coll. P.K.L. Ng et al., Jul. 2003, tangle net, Balicasag Is., Philippines, ZRC 2010.0138; Garthiella aberrans (Rathbun, 1906), 1 female, coll. G. Paulay, east of Opunohu Pass, Moorea, Society Is., Tuamotu Archipelago, UF 15624; Garthiella sp. nov., 1 female, coll. PANGLAO 2004, 2 Jul. 2004, Gar-sik-3, stn B37, 19–20 m, Balicasag Is., Philippines, ZRC 2010.0142; Pilodius areolatus (H. Milne Edwards, 1834), 5 males, 1 female, coll. P.K.L. Ng and S.H. Tan, 22 Jan. 2000, on sea wall, surf zone, Ala Moana, Waikiki, Oahu Is., Hawaii, ZRC 2000.0438; 2 males, 1 female, coll. G. Paulay, Apr. 2000, SW Orote Peninsula, Guam, ZRC 2000.0720; Pilodius paumotensis Rathbun, 1907, 1 male, coll. 12 Mar. 2002, Pulau Jemaja, Anambas Is., Indonesia, ZRC 2003.0559; Pilodius pilumnoides (White, 1848), 1 male, coll. M. Ward, Lindeman Is., Queensland, Australia, ZRC 1965.11.11.150; Pilodius pugil Dana, 1852, 1 male, coll. M. Puglisi, 8 Jun. 1998, coral rubble, 9 m, Cocos lagoon, Guam, ZRC 2000.0722; Sulcodius deflexus (Dana, 1852), 3 males, 1 female, coll. B. Richer de Forges, Ile Ouen–Bay du Prony, New Caledonia, ZRC 1998.793; Vellodius etisoides (Takeda and Miyake, 1968), 1 male, 1 female, coll. S.L. Yang, 25 Mar. 1982, Paracel Is. (Xisha Is.), South China Sea, ZRC 1998.50. Cymoinae Alcock, 1898: Cymo melanodactylus Dana, 1852, 2 males, 1 female, coll. S.H. Tan et al., 23 Jul. 1997, Pulau Seringat, Singapore, ZRC 1997.0347; 1 male, coll. S.H. Tan et al., 27 Jun. 1998, Pulau Seringat, Singapore, ZRC 1999.0339; Cymo quadrilobatus Miers, 1884, 1 male, 1 ovigerous female, coll. B. Henke, 3 Aug. 2001, Hospital Point, 9 m, Guam, ZRC 2001.0755. Etisinae Ortmann, 1893: Etisus anaglyptus (H. Milne Edwards, 1834), 1 male, coll. P.K.L. Ng, 27 Oct. 1992, Pulau Semakau, Singapore, ZRC 1992.10564; 1 female, coll. P.K.L. Ng et al., 24 Jun. 1999, Paya Beach, Pulau Tioman, Malaysia, ZRC 1999.0931; Etisus dentatus (Herbst, 1785), 1 male, coll. H.C. Liu, 31 Jul. 2001, Dadi Beach, Guam, ZRC 2001.0753; Etisus electra (Herbst, 1801), 1 male, 2 females, coll. R. de Felice, 19 Jan. 2000, Heeia Kea, Kaneohe Bay, Oahu Is., Hawaii, ZRC 2000.0503; Etisus frontalis (Dana, 1852), 1 male, coll. L. Kirkendale, May 1999, under rubble, 5 m, Double Reef, Guam, ZRC 2000.0694; Etisus odhneri Takeda, 1971, 1 female, coll. PANGLAO 2004, 11 Jun. 2004, Eti-odh-2, stn B11, 2–4 m, Pamilacan Is., Philippines, ZRC 2010.0140; Etisus splendidus Rathbun, 1906, 1 male, coll. local villagers, 23 Sep. 2003, Bunaken Is., Sulawesi, Indonesia, ZRC 2008.0526; Etisus utilis Jacquinot, in Jacquinot and Lucas, 1853, 1 female, Ao Nam Bay, Phuket Is., Thailand, ZRC 2002.0586; Etisus aff. villosus Clark and Galil, 1995, 1 male, coll. J.C.E. Mendoza and J. Arbasto, 21 Dec. 2007, tangle net, 120–160 m, Panglao Is., Philippines, ZRC 2008.1480. Euxanthinae Alcock, 1898: Alainodaeus akiaki Davie, 1993, holotype male, coll. SMCB/J. Poupin, 10 Mar. 1989, trap, 230–240 m, Rurutu, Austral Is., French Polynesia, MNHN B22243; Alainodaeus filipinus Mendoza and Ng, 2008, 1 female, coll. J.C.E. Mendoza and J. Arbasto, Dec. 2008, NW Panglao Is., Philippines, ZRC 2009.0142; Cranaothus deforgesi Ng, 1993, 1 male, 1 female, coll. PANGLAO 2004, 31 May 2004, stn B3, base of reef slope, 8 m, Arco Point, Panglao Is., Philippines, ZRC 2008.1363; Crosnierius carinatus Serène and Vadon, 1981, 1 male, coll. PANGLAO 2005, 24 May 2005, stn CP2349, 219–240 m, off Pamilacan Is., Philippines, NMCR 27332; 1 female, coll. AURORA 2007, 1 Jun. 2007, stn CP2737, 269–272 m, east coast of Luzon Is., Philippines, ZRC 2008.1370; Danielea noelensis (Ward, 1942), 1 male, Laamu Atoll, Maldives, ZRC 2007.0757; Epistocavea mururoa Davie, 1993, 1 male, coll. PANGLAO 2004, 21 Jun. 2004, stn L36, 85–90 m, Looc, Panglao Is., Philippines, ZRC 2008.1371; Euxanthus exsculptus (Herbst, 1790), 2 males, 1 female, coll. Honors Field Class NUS, Sep. 2002, Paya Beach, Pulau Tioman, Malaysia, ZRC 2002.0535; Euxanthus herdmani Laurie, 1906, 1 male, coll. PANGLAO 2004, 28 Jun. 2004, stn 32, 2–3 m, Looc, Panglao Is., Philippines, NMCR 27334; Euxanthus huonii (Hombron and Jacquinot, 1846), 1 male, coll. P.K.L. Ng, Dec. 1984, Pulau Hantu, Singapore, ZRC 1987.471; 1 male, coll. PANGLAO 2004, 9 Jun. 2004, stn M17, Pontod, Panglao Is., Philippines, ZRC 2008.1376; Euxanthus ruali Guinot, 1971, 1 male, coll. SANTO 2006, Sep. 2006, Aimbue Bay, Aore Is., Espiritu Santo, Vanuatu, ZRC 2009.1178; Glyptoxanthus erosus (Stimpson, 1859), 1 male, Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 coll. M. Gray, 7 Apr. 1977, Sapelo Is., Georgia, USA, ZRC 1998.9; Hepatoporus orientalis (Sakai, 1935), 1 male, coll. LUMIWAN 2008, Apr. 2008, off western Batangas, Luzon Is., Philippines, ZRC 2008.1379; Hypocolpus abbotti (Rathbun, 1894), 1 male, coll. P.K.L. Ng et al., 2 Mar. 2004, tangle net, 50–200 m, Balicasag Is., Philippines, NMCR 27335; Hypocolpus diverticulatus (Strahl, 1861), 1 male, coll. A. Anker et al., 2009, NW Nosy Komba, Madagascar, UF 14076; Hypocolpus pararugosus Crosnier, 1997, 1 male, coll. P.K.L. Ng et al., Jan. 2004, tangle net, 50–200 m, Balicasag Is., Philippines, ZRC 2008.1389; Hypocolpus perfectus GuinotDumortier, 1960, 1 female, coll. A. Crosnier, May 1960, Fort Dauphine, Madagascar, MNHN-B6516; Ladomedaeus fungillus Manuel-Santos and Ng, 2007, paratypes, 12 males, 8 females, coll. PANGLAO 2005, 22 May 2005, stn CP 2332, 396–419 m, Maribohic Bay, off Panglao Is., Philippines, ZRC 2010.0130; Ladomedaeus serratus (Sakai, 1965), 1 male, coll. T.Y. Chan et al., 2008, stn K13, near shallow hydrothermal vent, off Tashi Port, Taiwan, ZRC 2009.0342; Medaeops granulosus (Haswell, 1882), 2 males, 3 females, coll. M. Ward, May 1934, Lindeman Is., Queensland, Australia, ZRC 1965.11.10.46-50; 1 female, coll. S.H. Tan, 22 Aug. 1997, Pulau Seringat, Singapore, ZRC 1999.0856; Medaeops neglectus (Balss, 1922), 1 male, coll. A. Anker et al., 20 May 2008, lagoonal fringing reef slope, 1–2 m, Trois Freres islet, near Nosy Be, Madagascar, UF 14599; Medaeus danielita Mendoza and Ng, 2010, 1 male, coll. S. McKeon et al., 12 Oct. 2008, off Sheraton Hotel beach, Moorea, Society Is., French Polynesia, UF 15469; Medaeus elegans A. Milne-Edwards, 1867, 1 male, coll. H.T. Conley, 12 Mar. 2002, Glass Breakwater, 3–6 m, Apra Harbour, Guam, UF 2053; Medaeus ornatus Dana, 1852, 3 males, 2 females, coll. Albatross, stn 3872, Auau Channel, between Maui Is. and Lanai Is., Hawaii, USNM 29523; 1 male, coll. K. Kirkendale, 8 Apr. 1998, Glass Breakwater, 9 m, Apra Harbour, Guam, ZRC 2009.1180; Medaeus sp. nov., holotype male, coll. SANTO 2006, 7 Oct. 2006, stn FR 62, scuba, 2–3 m, Port Benier, Aore Is., Espiritu Santo, Vanuatu, ZRC 2010.0163; Miersiella cavifrons Takeda, 1989, 2 males, 2 females, coll. PANGLAO 2004, 16 Jun. 2004, stn T11, 78–96 m, Maribohoc Bay, off Bohol Is., Philippines, ZRC 2008.1396; Miersiella haswelli (Miers, 1886), 1 male, 2 females, coll. PANGLAO 2005, 23 May 2005, stn CP2343, 273–356 m, off Pamilacan Is., Philippines, ZRC 2008.1400; Monodaeus couchii (Couch, 1851), 3 males, coll. Thalassa Expedition, 1973, stn Z427, 330 m, Golfe de Gascogne, France, MNHN B30718; Monodaeus cristulatus Guinot and Macpherson, 1988, holotype male, coll. E. Macpherson/Benguela VI Expedition, dredge, 232 m, off Namibia, MNHN B17498; Monodaeus rectifrons (Crosnier, 1967), holotype male, coll. J. Marteau, 1 Aug. 1973, trawl, 80 m, off Pointe-Noire, Congo, MNHN B9580; Monodaeus aff. tuberculidens (Rathbun, 1911), 2 males, coll. P. Guéze, 1 Mar. 1973, trap, 420 m, Baie St. Paul, Reunion Is., MNHN B16164; New genus 2 sp. nov., paratypes, 2 males, coll. AURORA 2007, 28 May, 2007, stn CP2712, 144 m, east of Luzon Is., Philippines, ZRC 441 2010.0164; New genus 3 sp. nov. 1, paratype female, coll. PANGLAO 2004, 2 Jul. 2004, stn. B38, 17–18 m, Balicasag Is., Philippines, ZRC 2010.0152; New genus 3 sp. nov. 2, holotype male, coll. P.K.L. Ng et al., Nov. 2003, tangle net, Balicasag Is., Philippines, ZRC 2010.0153; Olenothus uogi Ng, 2002, holotype male, coll. L. Kirkendale, 17 Nov. 1998, Tepungan Channel, 2 m, Guam, UF 2096; paratype female, coll. 2000–2001, Guam, ZRC 2002.176; Paramedaeus globosus Serène and Vadon, 1981, 7 males, 5 females, coll. PANGLAO 2004, 1 Jun. 2004, stn T4, beam trawl, 84 m, Bolod, Panglao Is., Philippines, NMCR 27352; Paramedaeus aff. simplex A. Milne-Edwards, 1873, 1 male, 1 ovigerous female, coll. PANGLAO 2004, 20 Jun. 2004, stn B18, 3–5 m, Sungcolan Bay, Panglao Is., Philippines, ZRC 2008.1408; Pseudomedaeus distinctus (Rathbun, 1898), 1 male, coll. G. Paulay, 28 Nov. 2004, Capetown dredge, 39 m, Gulf of Mexico, off St. Petersburg, Florida, USA, UF 6659; Rizalthus anconis Mendoza and Ng, 2008, holotype male, coll. PANGLAO 2004, 8 Jun. 2004, stn R30, reef slope, 15–37 m, Napaling, Panglao Is., Philippines, NMCR 27507; paratype ovigerous female, coll. PANGLAO 2004, 2 Jul. 2004, stn B39, reef wall with caves, 17–25 m, Pontod Lagoon, Panglao Is., Philippines, ZRC 2008.0215; Visayax osteodictyon Mendoza and Ng, 2008, holotype male, coll. PANGLAO 2004, 21 Jun. 2004, stn S22, 15–22 m, Pamilacan Is., Philippines, NMCR 27508; 2 females, coll. PANGLAO 2004, 1 Jun. 2004, stn B4, reef slope with overhangs, 24 m, BBC Point, Panglao Is., Philippines, ZRC 2008.0753. Kraussiinae Ng, 1993: Kraussia rugulosa (Krauss, 1843), 1 male, Littlke Liuchiu, Taiwan, ZRC 2009.0335; Palapedia integra (De Haan, 1835), 1 male, coll. 2 Jan. 1980, Ping-tan Is., Fujian Province, China, ZRC; Palapedia roycei (Serène, 1972), 1 male, coll. P.K.L. Ng et al., 2 Mar. 2004, Pal-roy-1, tangle net, Balicasag Is., Philippines, ZRC; Palapedia sp. A, 1 male, coll. SANTO 2006, Sep. 2006, Tutuba Is., Vanuatu, ZRC 2009.1175; Palapedia sp. B, 1 male, coll. SANTO 2006, Sep. 2006, Segond Channel, Vanuatu, ZRC 2009.1176. Liomerinae Sakai, 1976: Liomera bella (Dana, 1852), 2 males, 5 females, coll. P.K.L. Ng, Apr. 1992, coral reef near Bise Village, Motobu-cho, Okinawa, Japan, ZRC 1993.646–652; Liomera cinctimana (White, 1847), 1 male, coll. J. Starmer and L. Kirkendale, 17 Jan. 1997, fore reef, 3–20 m, Gun Beach, Guam, ZRC 2000.0705; 1 male, Apra Harbour, Guam, ZRC 2000.0730; Liomera tristis (Dana, 1852), 1 male, coll. C.A. Gibson-Hill, 1941, Cocos Keeling Is., ZRC 1965.11.8.99–108; 2 males, 2 females, coll. P.K.L. Ng, Apr. 1992, Bise, Motobu-cho, Okinawa, Japan, ZRC 1993.653–656; 2 males, 2 females, coll. PANGLAO 2004, 1 Jun. 2004, Lio-tri-1, stn M7, 0–3 m, Momo Beach, Panglao Is., Philippines, ZRC 2010.0144; 1 male, 2 females, coll. P.K.L. Ng, Sep. 2002, Paya Beach, Pulau Tioman, Malaysia, ZRC 2003.0217; Liomera venosa (H. Milne Edwards, 1834), 1 female, coll. P.K.L. Ng et al., Jul. 1995, Tanjung Tandang, north coast of Pulau Bintan, Indonesia, ZRC 1999.0383; Lipkemera corallina (Takeda and Marumura, 1997), 1 male, coll. J. Arbasto, Nov. 2003–Apr. 2004, tangle net, 100–300 m, Author's personal copy 442 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 Maribojoc Bay, Bohol Is., Philippines, ZRC 2009.1172; Lipkemera holthuisi Mendoza, 2010, paratypes, 2 males, 1 female, coll. J. Arbasto, Jul. 2004-May 2005, tangle net, 50–200 m, north of Panglao Is., Philippines, ZRC 2009.1171; Neoliomera cerasinus Ng, 2002, holotype male, coll. L. Kirkendale, 1 Dec. 1999, Christmas Is., Indian Ocean, WAM; 1 male, Kume-jima, Ryukyu Is., Japan, ZRC 2001.2265; Neoliomera demani Forest and Guinot, 1961, 1 male, coll. S.H. Tan, 18 Mar. 2009, aquarium shop in Singapore, reportedly from Hawaii, ZRC 2009.1174; Neoliomera insularis (Adams and White, 1849), 1 male, coll. P.K.L. Ng et al., Dec. 2000, tangle net, Balicasag Is., Philippines, ZRC 2001.0666; Neoliomera striata Buitendijk, 1941, 1 female, coll. H.H. Tan, 27 Jul. 2003, near Balicasag Island Dive Resort, Balicasag Is., Philippines, ZRC 2008.0641. Polydectinae Dana, 1851: Lybia edmondsoni Takeda and Miyake, 1970, 3 females, coll. P.K.L. Ng and S.H. Tan, 22 Jan. 2000, Maipalaoa Beach, near Maili Point, Oahu Is., Hawaii, ZRC 2000.0435; Lybia hatgumoana Sakai, 1961, 1 male, coll. PANGLAO 2004, 14 Jun. 2004, Lyb-hat-3, stn T9, beam trawl, 97–120 m, off San Isidro, Panglao Is., Philippines, ZRC 2010.0147; Lybia leptochelis (Zehntner, 1894), 1 ovigerous female, coll. PANGLAO 2004, 11 Jun. 2004, Lyb-lep-3, stn B11, coral rubble, 2–4 m, Pamilacan Is., Philippines, ZRC 2010.0146; Lybia tesselata (Latreille, in Milbert, 1812), 1 female, coll. G. Paulay, Apr. 2000, fore-reef, SW of Orote Peninsula, Guam, ZRC 2000.0710; Polydectus cupulifer (Latreille, in Milbert, 1812), 1 female, coll. P.K.L. Ng and S.H. Tan, 22 Jan. 2000, Magic Is., Waikiki, Oahu Is., Hawaii, ZRC 2000.0441. Speocarcininae Števčić, 2005: Speocarcinus lobatus Guinot, 1969, 1 ex., Northern Gulf of Mexico, ULLZ 7820 [GenBank, Thoma et al., 2009]; Speocarcinus monotuberculatus Felder and Rabalais, 1986, South-western Gulf of Mexico, ULLZ 7562 [GenBank, Thoma et al., 2009]. Xanthinae MacLeay, 1838: Cycloxanthops truncatus (De Haan, 1837), 1 female, coll. P.K.L. Ng et al., Nov. 2003, Cyctru-1, tangle net, Balicasag Is., Philippines, ZRC 2010.0139; Demania cultripes (Alcock, 1898), 1 male, lobster pot, 30 m, Peace Island, Taiwan, ZRC 1998.530; Demania intermedia Guinot, 1969, 1 female, coll. J. Arbasto, Jan.–Dec. 2007, tangle net, 120–160 m, NW Panglao Is., Philippines, ZRC 2009.0187; Demania scaberrima (Walker, 1887), 1 male, coll. P.K.L. Ng et al., May 2000, Pichai fishport, Phuket Is., Thailand, ZRC 2000.0833; Demania splendida Laurie, 1906, 1 male, coll. P.K.L. Ng et al., Dec. 2000, tangle net, Balicasag Is., Philippines, ZRC 2001.0650; Eucratodes agassizii A. Milne-Edwards, 1880, 1 ex., ULLZ 8400 [GenBank, Thoma, et al. 2009] Euryxanthops dorsiconvexus Garth and Kim, 1983, 1 male, 2 females, coll. PANGLAO 2005, 26 May 2005, stn CP2358, 569–597 m, Bohol/Sulu seas sill, Philippines, ZRC 2010.0136; Euryxanthops orientalis (Sakai, 1939), 1 male, 1 female, coll. AURORA 2007, 29 May 2007, Eur-ori1, stn CP2716, 356 m, east of Luzon Is., Philippines, ZRC 2010.0141; Garthiope spinipes (A. Milne Edwards, 1880), 1 ex., Southern Gulf of Mexico, ULLZ 8131 [GenBank, Thoma et al., 2009]; Lachnopodus bidentatus (A. Milne-Edwards, 1867), 1 male, 1 female, coll. D. Vandenspiegel, 8 Mar. 1992, Laing Is., Papua New Guinea, ZRC 1995.0411; Lachnopodus subacutus (Stimpson, 1858), 1 ovigerous female, coll. P.F. Clark and H.H. Tan, Jul. 2003, reef flat, Balicasag Is., Philippines, ZRC 2003.0282; Leptodius exaratus (H. Milne Edwards, 1834), 1 female, coll. 18 Mar. 2002, Pulau Salor, Natuna Is., Indonesia, ZRC 2003.0549; Leptodius nigromaculatus Serène, 1962, 7 males, 1 female, coll. C.D. Schubart, Mar. 2000, Labrador Beach, Singapore, ZRC 2000.2010; 1 male, Raffles Lighthouse, Singapore, ZRC 2008.0652; Leptodius nudipes (Dana, 1852), 1 male, coll. C.D. Schubart, 25 Jun. 2000, Pulau Sapi, Sabah, Malaysia, ZRC 2000.1675; Liagore pulchella Ng and Naruse, 2007, 2 males, 2 females (paratypes), coll. SANTO 2006, 17 sep. 2006, stn EP15, tangle net, 103–105 m, off Espiritu Santo Is., Vanuatu, ZRC 2007.0642; Liagore erythematica Guinot, 1971, 3 males 4 females, coll. J.C.Y. Lai et al., 26 Nov. 2003, Ishigawa Port, Okinawa, Japan, ZRC 2005.0013; Liagore rubromaculata (De Haan, 1835), 1 male, coll. PANGLAO 2004, 21 Jun. 2004, Lia-rub-1, stn T23, 35–45 m, Cortes, Bohol Is., Philippines, ZRC 2010.0143; Macromedaeus crassimanus (A. Milne-Edwards, 1867), 1 female, coll. H.H. Tan and P.F. Clark, 27 Jul. 2003, reef, Balicasag Is., Philippines, ZRC 2003.0369; Metaxanthops acutus Serène, 1984, 1 ovigerous female, coll. PANGLAO 2004, 7 Jun. 2004, Met-acu-1, stn D6, dredge, 3 m, Bolod, Panglao Is., Philippines, ZRC 2010.0148; Micropanope sculptipes Stimpson, 1871, 1 ex., Southeastern Gulf of Mexico, ULLZ 6603 [GenBank, Thoma et al., 2009]; Nanocassiope alcocki (Rathbun, 1902), 1 male, coll. LUMIWAN 2008, 1 Apr. 2008, Nan-gra-1, stn CC2925, west of Batangas, Luzon Is., Philippines, ZRC 2010.0149; 1 male, coll. P.K.L. Ng et al., 25 Oct. 2003, Nan-nsp-1, tangle net, Balicasag Is., Philippines, ZRC 2010.0150; Neoxanthias michelae Serène and Vadon, 1981, 1 male, coll. S. Chaitiamvong et al., Dec. 1998, Pichai fish port, Phuket Is., Thailand, ZRC 1999.0516; Neoxanthops lineatus (A. MilneEdwards, 1867), 1 male, coll. P. Ng, 31 Dec. 1986, Sentosa Is., Singapore, ZRC 1987.0531; Neoxanthops quadrilobatus (Sakai, 1939), 1 male, stn 6199, 33 m, Beibu Wan, Taiwan, ZRC 1999.0022; New genus near Garthiope sp. nov., Northern Gulf of Mexico, ULLZ 8183 [tissue only]; Paraxanthias elegans (Stimpson, 1858), 1 female, coll. PANGLAO 2004, 3 Jun. 2004, Parx-ele-2, stn S2, 4–5 m, Baclayon, Bohol Is., Philippines, ZRC 2010.0155; Paraxanthias notatus (Dana, 1852), 7 males, 7 females, coll. 21 May 1985, Chutan, Sunabe, Okinawa, Japan, ZRC 1999.0260; Paraxanthias pachydactylus (A. Milne-Edwards, 1867), 1 male, 1 female, coll. PANGLAO 2004, 3 Jun. 2004, Parx-pac-1, stn S2, 4–5 m, Baclayon, Bohol Is., Philippines, ZRC 2010.0154; Paraxanthodes obtusidens (Sakai, 1965), 1 male, coll. AURORA 2007, 2 Jun. 2007, stn CP2747, 118–124 m, east of Luzon Is., Philippines, ZRC 2008.1410; Xanthias canaliculatus Rathbun, 1906, South Africa: Sodwana Bay, ULLZ 4381 [tissue only]; Xanthias lamarcki (H. Milne Edwards, 1834), 1 male, coll. C.C. Lim, 18 May 1988, Kaoshiung, Taiwan, Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 ZRC 1995.625; 1 ovigerous female, coll. PANGLAO 2004, 28 Jun. 2004, Xan-lam-1, stn B34, Sungcolan Inlet, Panglao Is., Philippines, ZRC 2010.0160; Xanthias latifrons (De Man, 1887), 1 male, coll. L. Kirkendale, 17 Nov. 1998, Tepungan Channel, 2 m, Guam, ZRC 2000.0728; Xanthias punctatus (H. Milne Edwards, 1834), 1 male, coll. P.K.L. Ng, 18 Dec. 2000, reef, Balicasag Is., Philippines, ZRC 2001.0309; Xanthias teres Davie, 1997, 1 male, coll. P.K.L. Ng, et al., 28 Nov. 2001, tangle net, 50–500 m, Balicasag Is., Philippines, ZRC 2001.0689; 1 male, coll. J. Arbasto, 2004–2005, tangle net, Maribojoc Bay, Panglao Is., Philippines, ZRC 2010.0161; Xantho hydrophilus (Herbst, 1790), 1 male, coll. J. Bishop, 15 Oct. 2009, intertidal, Plymouth, Cornwall, England, UK, ZRC 2011.0492. Xantho pilipes A. Milne-Edwards, 1867, 1 ex., Flaggy Shore, Ireland, NHM 2010.477 [tissue only]; Xantho poressa (Olivi, 1792), 4 males, coll. Z. Stevcic, 1 Sep. 1971, rocky shore, intertidal, Rovinj, Yugoslavia, ZRC 1988.694-697. Zalasiinae Serène, 1968: Banareia nobilii (Odhner, 1925), 1 male, coll. P.K.L. Ng et al., Jun. 2002, Ban-nob-1, tangle net, 200–300 m, Balicasag Is., Philippines, ZRC; 1 male, coll. PANGLAO 2004, 30 May 2004, Ban-nob-3, stn T1, 83–102 m, Bolod, Panglao Is., Philippines, ZRC 2010. 0131; Calvactaea tumida Ward, 1933, 1 male, coll. Colombo Museum, date unknown, Colombo, Sri Lanka, ZRC 1969.12.31.5; 1 female, coll. PANGLAO 2004, 30 May 2004, Cal-tum-2, stn T1, 83–102 m, Bolod, Panglao Is., Philippines, ZRC 2010.0137; Zalasius dromiaeformis (De Haan, 1839), 1 male, coll. P.K.L. Ng, Dec. 2000, tangle net, 200–300 m, Balicasag Is., Philippines, ZRC 2001.0383; 1 male, Taiwan, ZRC 2008.0680; Zalasius sakaii Balss, 1938, 1 male, coll. J.F. Huang, 7 Jun. 1992, Mitou, Kaoshiung County, Taiwan, ZRC 1997.0399. Zosiminae Alcock, 1898: Atergatis floridus (Linnaeus, 1767), 1 female, coll. S.H. Tan et al., 23 Jul. 1997, Pulau Seringat, Singapore, ZRC 2009.0373; Atergatis integerrimus (Lamarck, 1818), 1 male, coll. P.K.L. Ng, 27 Oct. 2003, Beting Bronok reef, Singapore, ZRC 2007.0252; Atergatis interruptus Takeda and Marumura, 1997, 2 males, coll. J.C.E. Mendoza and J. Arbasto, 21 Dec. 2007, tangle net, 120–160 m, NW Panglao Is., Philippines, ZRC 2008.1471; Atergatis obtusus A. Milne-Edwards, 1865, 2 males, coll. P.K.L. Ng, Dec. 1998, Phuket Is., Thailand, ZRC 1999.0521; 1 male, 1 female, coll. PANGLAO 2004, 28 Jun. 2004, stn R66, 1–3 m, Sungcolan, Panglao Is., Philippines, ZRC 2010.0132; Atergatopsis aff. amoyensis De Man, 1879, 1 female, coll. PANGLAO 2004, 30 May 2004, stn P1, tangle net, 90–200 m, Marinohoc Bay, Bohol Is., Philippines, ZRC 2010.0133; Atergatopsis germaini A. Milne-Edwards, 1865, 1 male, coll. H. P. Ho, 29 Oct. 1993, lobster pot, 40 m, Ho-Ping Is., Keelung, Taiwan, ZRC 2009.0298; Atergatopsis granulata A. Milne-Edwards, 1865, 1 female, coll. PANGLAO 2004, 5 Jun. 2004, stn B7, 4–30 m, Catarman, Panglao Is., Philippines, ZRC 2010.0134; Atergatopsis aff. obesa (A. Milne-Edwards, 1865), 1 female, coll. PANGLAO 2004, 4 Jul. 2004, stn B41, 17–19 m, Balicasag Is., 443 Philippines, ZRC 2010.0135; Lophozozymus anaglyptus (Heller, 1861), 1 female, coll. PANGLAO 2004, 11 Jun. 2004, Lop-ana-1, stn B11, coral rubble, 2–4 m, Pamilacan Is., Philippines, ZRC 2010.0145; Lophozozymus edwardsi (Odhner, 1925), 1 female, coll. H.H. Tan et al., 16 Mar. 2002, Pulau Laut, Natuna Is., Indonesia, ZRC 2003.0553; Lophozozymus pictor (Fabricius, 1798), 1 male, coll. 07 May 2004, Raffles Lighthouse, Singapore, ZRC 2009.0592; New genus 1 sp. nov. (close to Paratergatis), paratype male, coll. PANGLAO 2004,27 Jun. 2004, Neo-qua-5, stn P1, tangle net, Pamilacan Is., Philippines, ZRC 2010.0151; Paratergatis longimanus Sakai, 1965, 1 female, coll. M.S. Jeng, 26 Nov. 1997, Tai-chi Port, I-lan County, Taiwan, ZRC 1998.0047; 1 male, coll. S.H. Fan, Aug. 1996, Sa Yu Harbour, Fujian Province, China, ZRC 2002.0578; Platypodia eydouxi (A. Milne-Edwards, 1865), 1 male, 2 females, coll. P.K.L. Ng and S.H. Tan, 22 Jan. 2000, Malpalaoa Beach, Oahu Is., Hawaii, ZRC 2000.0440; Platypodia pseudogranulosa Serène, 1984, 1 male, coll. H.H. Tan, 25 Apr. 2008, Cyrene Reef, Singapore, ZRC 2008.0492; Platypodia tomentosa (De Man, 1902), 1 male, coll. PANGLAO 2004, 4 Jul. 2004, Pla-alc-2, stn T38, beam trawl, 80–140 m, Balicasag Is., Philippines, ZRC 2010. 0156; Pulcratis reticulatus Ng and Huang, 1997, 1 male, Ping-tung County, Taiwan, ZRC 1997.0402; Zosimus aeneus (Linnaeus, 1758), 1 male, coll. C.C. Lim, 19 May 1988, Kaoshiung, Taiwan, ZRC 1995.612; 1 male, coll. H.C. Liu, 19 May 1998, Heng Chun Peninsula, Ping-tung County, Taiwan, ZRC 1998.388; Zosimus hawaiiensis (Rathbun, 1906), 1 male, coll. J. Park, Jan. 2000, north of Oahu Is., ZRC 2000.0515; PANOPEIDAE Ortmann, 1893. Panopeinae Ortmann, 1893: Acantholobulus pacificus (Edmondson, 1931), 1 male, 1 female, Pearl Harbour, Oahu Is., Hawaii, ZRC 2000.0554; Glyptoplax smithii A. Milne Edwards, 1880, 1 ex., Southwestern Gulf of Mexico, USA, ULLZ 6793 [GenBank, Thoma et al., 2009]; Hexapanopeus paulensis Rathbun, 1930, 1 ex., Panama City, Northern Gulf of Mexico, Panama, ULLZ 8645 [GenBank, Thoma et al., 2009]; Panopeus herbstii H. Milne Edwards, 1834, 2 males, 2 females, coll. C.D. Schubart and J.A. Cuesta, 23 Aug. 2000, salt marsh, Masonboro Sound, Wilmington, North Carolina, USA, ZRC 2001.1015; Panopeus occidentalis Saussure, 1857, 1 male, ex. USU-1371, Brazil, ZRC 2008.0160; 1 ex., Panama City, Northern Gulf of Mexico, Panama, ULLZ 8643 [GenBank, Thoma et al., 2009]. PSEUDORHOMBILIDAE Alcock, 1900. Trapezioplax tridentata (A. Milne-Edwards, 1880), 1 ex., northern Gulf of Mexico, ULLZ 8054 [GenBank, Thoma et al., 2009]. Appendix B. List of 48 first stage zoeas of Xanthoidea examined and described XANTHIDAE Macleay, 1838. Actaeinae Alcock, 1898: Actaea areolata (Dana, 1852): coll. P. Clark and P. Ng, 27 July 2003, Balicasag, Philippines, Author's personal copy 444 J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 hatched 9 August 2003, NHM; Actaeodes hirsutissimus (Rüppell, 1830): coll. Ali Al-Aidaroos, 23 July 1990, Obhor Creek, 21◦ 40′ N, 39◦ 12′ E, Red Sea, ca. 20 km north of Jeddah, Saudi Arabia, hatched 26 July 1990, NHM 1994.3230; Actaeodes mutatus Guinot, 1976: coll. P. Clark, 30 July 2003, Buyong, Maribago, Moctan Island, Cebu Island, Philippines, hatched 6 August 2003, ZRC 2003.0278; Actaeodes tomentosus (H. Milne Edwards, 1834): coll. A. Al-Aidaroos, 23 July 1990, Obhor Creek, 21◦ 40′ N, 39◦ 12′ E, Red Sea, ca. 20 km north of Jeddah, Saudi Arabia, hatched 4 August 1990, NHM 1994.3229; Epiactaea nodulosa (White, 1848): coll. D. Lane, Southern Islands, Singapore, ZRC 1995.331; Gaillardiellus orientalis (Odhner, 1925): coll. P. Ng, from littoral coral reef, Sentosa Island, Singapore, October 1992, hatched 27 October 1992, NHM 1993.3408–3417; Novactaea bella Guinot, 1976: coll. P. Clark and P. Ng, 27 July 2003, Balicasag, Philippines; Psaumis cavipes (Dana, 1852): coll. P. Clark and P. Ng, 29 July 2003, west of Alona Beach, Panglao, Philippines, hatched 6 August 2003, ZRC 2003.0287; Pseudactea corallina (Alcock, 1898): coll. P. Clark and P. Ng, 27 July 2003, Balicasag, Philippines, hatched 28 July 2003, ZRC 2003.0276; Pseudoliomera speciosa (Dana, 1852): coll. by SCUBA from Albion Rocks, Baie de la Petite Rivière, off Avenue Victory, Petite Rivière, Albion, Mauritius, 3–7 m, 15 May 1995, hatched 18–19 May 1995, NHM 1997.1. Chlorodiellinae Ng and Holthuis, 2007: Chlorodiella nigra (Forskål, 1775): coll. P. Clark by SCUBA diving from Albion Rocks, Baie de la Petite Rivière, off Victory Road, Petite Rivière, Albion, Mauritius, ca. 20◦ 12.5′ S, 57◦ 23.5′ E, 3–7 m, 6 May 1995, hatched 19 May 1995, NHM 1998.602; Cyclodius obscurus (Jacquinot in Hombron and Jacquinot, 1846): coll. S. Lundoer, Phuket region, Thailand, 12 March 73, hatched 18 March 73, NHM 1998.601; Pilodius areolatus (H. Milne Edwards, 1834): coll. P. Clark, SCUBA diving from Albion Rocks, Baie de la Petite Rivière, off Victory Road, Petite Rivière, Albion, Mauritius, ca. 20◦ 12.5′ S, 57◦ 23.5′ E, 3–7 m, 6 May 1995, hatched 8 May 1995, NHM 1998.603; Pilodius paumotensis Rathbun, 1907: coll. P. Clark and J. Paula, polychaete worm reef, Cabo Inhaca, North East Inhaca Island, Mozambique, 13 November 1997, hatched 15 November 1997, NHM 2003.194; Pilodius pugil Dana, 1852: coll. P. Clark by SCUBA from Albion Rocks, Baie de la Petite Rivière, off Victory Road, Petite Rivière, Albion, Mauritius, ca. 20◦ 12.5′ S, 57◦ 23.5′ E, 3–7 m, 6 May 1995, hatched 20 May 1995, NHM 1998.604. Cymoinae Alcock, 1898: Cymo lanatopodus Galil and Vannini, 1990: coll. P. Clark and B. Galil by SCUBA, 4 m, 9 May 1995, Trou aux Biches, Mauritius, hatched 24 May 1995, NHM; Cymo melanodactylus Dana, 1853: coll. P. Clark and B. Galil by SCUBA, 4 m, 9 May 1995, Trou aux Biches, Mauritius, hatched 24 May 1995, NHM. Etisinae Ortmann, 1893: Etisus anaglyptus H. Milne Edwards, 1834: coll. P. Ng, Sentosa reefs, Singapore, ZRC 1985.1855; Etisus frontalis (Dana, 1852): coll. P. Ng, July 2001, Guam, NHM; Etisus utilis H. Jacquinot in H. Jacquinot and Lucas, 1854: coll. P. Clark by SCUBA, 7 December 1998, Chenal des 5 miles, au sud du récif Niagi, 22◦ 23′ S, 166◦ 45.37′ E, Nouvelle-Calédonie, hatched 17 December 1998, NHM. Euxanthinae Alcock, 1898: Medaeops granulosus (Haswell, 1882): coll. P. Ng, 22 May 1982, Singapore, ZRC 1985.1861; Monodaeus couchi (Couch, 1851): coll. P. Clark and J. Ellis, July 1979, trawled off Spanish Head, Isle of Man, NHM 1982.541–42. Kraussiinae Ng, 1993: Palapedia integra (De Haan, 1835): coll. H. S. Ko by SCUBA, 20 m, 2 July 2002, Jeju Island, 33◦ 25′ N, 126◦ 20′ E, South Korea, hatched 7 July 2002, Silla University Zoological Collection Cr 103240, South Korea; Palapedia valentini Ng, 1993: coll. C.G.S. Tan, 27 October 1992, under littoral coral rock, Palau Semakau, Singapore, ZRC 1993.4066. Liomerinae Sakai, 1976: Liomera bella (Dana, 1852): coll. P. Ng, January 2003, intertidal area, Maipalaoa Beach, near Maili Point, Maili, Waianae Coast, Hawaii, USA, NHM; Liomera cinctimana (White, 1847): coll. P. Ng, July 2001, Guam, NHM; Liomera laevis (A. Milne-Edwards, 1873): coll. P. Clark and P. Ng 5 May 2000, beach reef, Ao Tang Khaen, Phuket, Thailand, hatched 17 May 2000, NHM. Polydectinae Dana, 1851: Lybia plumosa Barnard, 1947: coll. P. Clark and J. Paula, Barreira Vermelha, Inhaca Island, Mozambique, 1 December 1997, hatched 8 December 1997, NHM 2003:195. Xanthinae MacLeay, 1838: Cataleptodius floridanus (Gibbes, 1850): coll. P. Clark, det. R. Manning, from south side of Fort Pierce Inlet, Florida, USA, 3 May 1997, hatched 13 May 1997, NHM 2001.118; coll. R. Ingle, Ferry Reach mud flats, opposite Biological Station, Bermuda, 2 October 1983, hatched 10 October 1983, NHM 1985:467; Garthiope barbadensis (Rathbun, 1921): coll. J.K. Reed by SCUBA, lockout diving, 19 May 1977, R/V Sea Diver, 80 m, Jeff’s Reef, 27◦ 32.8′ N, 79◦ 58.8′ W, 27 km north east of Fort Peirce, St. Lucie County, Florida, USA, hatched 1 June 1977, Harbour Branch Museum ID 89:4614; Lachnopodus subacutus (Stimpson, 1858): coll. P. Clark and P. Ng, 27 July 2003, Balicasag, Philippines, hatched 8 July 2003, ZRC 2003.0282; Leptodius exaratus (H. Milne Edwards, 1834): coll. by P. Clark and J. Paula, off Inhaca Marine Biological Station, Inhaca Island, Mozambique, 11 November 1997, hatched 12 November 1997, NHM 2000.1908; Leptodius sanquineus (H. Milne Edwards, 1834): Aldabra Island, hatched March 1968, NHM; Macromedaeus crassimanus (A. Milne-Edwards, 1867): coll. P. Clark, 27 July 2003, Balicasag, Philippines, hatched 9 August 2003, NHM; Microcassiope minor (Dana, 1852): coll. by hand on lower shore from Ponta Delgada, São Miguel, Açores, 5 August 2002, NHM 2002.2006; Nanocassiope granulipes (Sakai, 1939): coll. H. S. Ko by SCUBA diving, Cheju Island, off southern part of Korea, 13 July 1999, hatched 17 July 1999, Silla University Zoological Collection, South Korea; Nanocassiope melanodactyla (A. Milne-Edwards, 1867): coll. by SCUBA, 22 July 1989, 10 m, Caldeirinha, Guia, Ilha do Faial, Açores, hatched 30 July 1989, NMH 2002.4; Xantho Author's personal copy J.C.Y. Lai et al. / Zoologischer Anzeiger 250 (2011) 407–448 hydrophilus (Herbst, 1790): coll. R. Ingle and J. Paula, Mira Estuary, Portugal, 8 June 1988, NHM 2000.1907; Xantho pilipes A. Milne-Edwards, 1867: coll. J. Paula by SCUBA, July 1989, 10 m, crater of Guia Mount, Faial Island, Azores, reg. no. IPIMAR AdS10/1991; Xantho poressa (Olivi, 1792): coll. A. Rodríquez, El Chato Beach, Cádiz, Gulf of Cádiz, Spain, 14 June 1995, hatched 16 June 1995, reg. no. MNCN 20.04/3627. Zalasiinae Serène, 1968: Banareia subglobosa (Stimpson, 1858): coll. D.G.B. Chia, T.M. Sin, R. Teo and A. Wang, 12 September 1994, Johore Shoal, 01◦ 19′ 02′′ N, 104◦ 03′ 13′′ E, Singapore, hatched 19 September 1994, ZRC 1995.353. Zosiminae Alcock, 1898: Atergatis floridus (Linnaeus, 1767): coll. P. Ng, April 1982, Siloso Beach, Sentosa Island, Singapore, hatched 13 April 1982, ZRC 1984.608-4007; Atergatis subdentatus (De Haan, 1835): coll. P.-H. Ho, 5 May 1993, Heping Island, Keelung, Taiwan, hatched 17 May 1993, ZRC; Atergatopsis germaini A. Milne-Edwards, 1865: coll. P.-H. Ho, 15 April 1993, Heping Island, Keelung, Taiwan, hatched 17 May 1993, ZRC; Lophozozymus pictor (Fabricius, 1798): coll. P. Ng, Siloso Beach, Sentosa Island, Singapore, hatched 29 January 1992, ZRC 1997.771; Platypodia eydouxi (A. Milne-Edwards, 1865): coll. R. De Felice, 4 January 2002, hatched 7 January 2000 P. K. L. Ng, Heeia Kea, Kaneohe Bay, Oahu, Hawaii, NHM 2004.243; Platypodiella spectabilis (Herbst, 1794): coll. Zimmerman and J. Martin, 14 July 2000, hatched 14 July 2000, North Beach, Guana Island, British Virgin Islands, Los Angeles County Museum CR 2000006; Zozymodes xanthoides (Krauss, 1843): coll. P. Clark and J. Paula, polychaete worm reef, Cabo Inhaca, North East Inhaca Island, Mozambique, 13 November 1997, hatched 25 November 1997, NHM 2003:196. PANOPEIDAE Ortmann, 1893. Panopeinae Ortmann, 1893: Hexapanopeus paulensis Rathbun, 1930: coll. by otter trawl, 6–8 m on sandy bottom, near Ubatuba, 23◦ 26′ S, 45◦ 05′ W, São Paulo, Brazil, July 1989, NHM; Panopeus americanus De Saussure, 1857: coll. P. Clark, 3 May 1997, south side, Fort Pierce Inlet, Fort Pierce, St. Lucie County, Florida, USA, hatched 9 May 1997, NHM; Panopeus harti Smith, 1869: coll. P. Clark by dredging, 30 m, R/V Sunburst, 30 April 1997, off Bethel Shoals, 27◦ 40′ N, 80◦ 06′ W, Florida, USA, hatched 3 May 1997, NHM; Panopeus occidentalis De Saussure, 1857: coll. P. Clark, 3 May 1997, south side, Fort Pierce Inlet, Fort Pierce, St. Lucie County, Florida, USA, hatched 6 May 1997, NHM;Panopeus simpsoni Rathbun, 1930: coll. P. Clark, 4 May 1997, Jack Island, Indian River Lagoon, St. Lucie County, Florida, USA, hatched 16 May 1997, NHM. References Balss, H., 1957. VIII. Systematik. Decapoda. In: Bronns, H.G. (Ed.), Klassen und Ordnungen des Tierreichs. Leipzig, Akademische Verlagsgesellschaft. Bd5, I Abt., 7. Buch, 12 Lief., 1505–1672, abb. 1131–1199. 445 Bourdillon-Casanova, L., 1960. Le méroplancton du Golfe de Marseille: les larves de Crustacés Décapodes. Recl Trav. Stn mar. Endoume 30, 1–286. Clark, P.F., 2001. Interpreting patterns in chaetotaxy and segmentation associated with abbreviated brachyuran zoeal development. Invert. Rep. Dev. [2000] 38 (3), 171–181 (figs. 1–13). Clark, P.F., 2005. The evolutionary significance of heterochrony in the abbreviated zoeal development of pilumnine crabs (Crustacea: Brachyura: Xanthoidea). Zool. J. Linn. 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