Arthur Anker

Morphological phylogeny of alpheid shrimps : Parallel preadaptation and the origin of a key morphological innovation, the snapping claw

Abstract The Alpheidae—possibly the most diverse family of recent decapod crustaceans—offers attractive opportunities to study the evolution of many intriguing phenomena, including key morphological innovations like spectacular snapping claws, highly specialized body forms, facultative and obligate symbioses with many animal groups, and sophisticated behaviors like eusociality. However, studies of these remarkable adaptations remain hampered by insufficient phylogenetic information. We present the first phylogenetic hypothesis of relationships among 36 extant genera of alpheid shrimps, based on a cladistic analysis of 122 morphological characters from 56 species, and we use this hypothesis to explore evolutionary trends in morphology and species diversity. Our results strongly supported a monophyletic Alpheidae that included two hitherto difficult-to-place genera (Yagerocaris and Pterocaris). Of 35+ nodes among genera, all were supported by at least one morphological character (24 were supported by two or more) and 17 received greater than 50% jackknife support. Unfortunately, many basal nodes were only weakly supported. Six genera appeared nonmonophyletic, including the dominant genus Alpheus (paraphyletic due to inclusion of one clade with three minor genera). Evolutionary trends in alpheid claw form shed some revealing light on how key innovations evolve. First, several functionally significant features of the cheliped (claw bearing leg) evolved independently multiple times, including: asymmetry, folding, inverted orientation, sexual dimorphism, adhesive plaques that enhance claw cocking, and tooth-cavity systems on opposing claw fingers, a preadaptation for snapping. Many conspicuous features of alpheid claw form therefore appear prone to parallel evolution. Second, although tooth-cavity systems evolved multiple times, a functional snapping claw, which likely facilitated an explosive radiation of over 550 species, evolved only once (in Synalpheus + [Alpheus + satellite genera]). Third, adhesive plaques (claw cocking aids) also evolved multiple times, and within snapping alpheids are associated with the most diverse clade (Alpheus + derivative genera). This pattern of parallel preadaptation—multiple independent evolutionary origins of precursors (preadaptations) to what ultimately became a key innovation (adaptation)—suggests alpheid shrimp claws are predisposed to develop features like tooth-cavity and adhesive plaque systems for functional or developmental reasons. Such functional/developmental predisposition may facilitate the origin of key innovations. Finally, moderate orbital hoods—anterior projections of the carapace partly or completely covering the eyes—occur in many higher Alpheidae and likely evolved before snapping claws. They are unique among decapod crustaceans, and their elaboration in snapping alpheids suggests they may protect the eyes from the stress of explosive snaps. Thus one key innovation (orbital hoods) may have facilitated evolution of a second (snapping claws).

Acanthaster planci Outbreak: Decline in Coral Health, Coral Size Structure Modification and Consequences for Obligate Decapod Assemblages

Although benthic motile invertebrate communities encompass the vast majority of coral reef diversity, their response to habitat modification has been poorly studied. A variety of benthic species, particularly decapods, provide benefits to their coral host enabling them to cope with environmental stressors, and as a result benefit the overall diversity of coral-associated species. However, little is known about how invertebrate assemblages associated with corals will be affected by global perturbations, (either directly or indirectly via their coral host) or their consequences for ecosystem resilience. Analysis of a ten year dataset reveals that the greatest perturbation at Moorea over this time was an outbreak of the corallivorous sea star Acanthaster planci from 2006 to 2009 impacting habitat health, availability and size structure of Pocillopora spp. populations and highlights a positive relationship between coral head size and survival. We then present the results of a mensurative study in 2009 conducted at the end of the perturbation (A. planci outbreak) describing how coral-decapod communities change with percent coral mortality for a selected coral species, Pocillopora eydouxi. The loss of coral tissue as a consequence of A. planci consumption led to an increase in rarefied total species diversity, but caused drastic modifications in community composition driven by a shift from coral obligate to non-obligate decapod species. Our study highlights that larger corals left with live tissue in 2009, formed a restricted habitat where coral obligate decapods, including mutualists, could subsist. We conclude that the size structure of Pocillopora populations at the time of an A. planci outbreak may greatly condition the magnitude of coral mortality as well as the persistence of local populations of obligate decapods.

Ecological speciation in anemone-associated snapping shrimps (Alpheus armatus species complex).

Divergent natural selection driven by competition for limited resources can promote speciation, even in the presence of gene flow. Reproductive isolation is more likely to result from divergent selection when the partitioned resource is closely linked to mating. Obligate symbiosis and host fidelity (mating on or near the host) can provide this link, creating ideal conditions for speciation in the absence of physical barriers to dispersal. Symbiotic organisms often experience competition for hosts, and host fidelity ensures that divergent selection for a specific host or host habitat can lead to speciation and strengthen pre‐existing reproductive barriers. Here, we present evidence that diversification of a sympatric species complex occurred despite the potential for gene flow and that partitioning of host resources (both by species and by host habitat) has contributed to this diversification. Four species of snapping shrimps (Alpheus armatus, A. immaculatus, A. polystictus and A. roquensis) are distributed mainly sympatrically in the Caribbean, while the fifth species (A. rudolphi) is restricted to Brazil. All five species are obligate commensals of sea anemones with a high degree of fidelity and ecological specificity for host species and habitat. We analysed sequence data from 10 nuclear genes and the mitochondrial COI gene in 11–16 individuals from each of the Caribbean taxa and from the only available specimen of the Brazilian taxon. Phylogenetic analyses support morphology‐based species assignments and a well‐supported Caribbean clade. The Brazilian A. rudolphi is recovered as an outgroup to the Caribbean taxa. Isolation–migration coalescent analysis provides evidence for historical gene flow among sympatric sister species. Our data suggest that both selection for a novel host and selection for host microhabitat may have promoted diversification of this complex despite gene flow.

Atyidae and Palaemonidae (Crustacea: Decapoda: Caridea) of Bocas del Toro, Panama

The present contribution is a preliminary report on the freshwater caridean fauna of Bocas del Toro province, northeastern Panama, based on field collections carried out during a Shrimp Taxonomy Workshop at the STRI station in Bocas del Toro in August 2008. A total of eight species from two families, Atyidae and Palaemonidae, were collected at 17 different collection sites in the rivers, streams and ponds on several islands of the Bocas del Toro archipelago and the adjacent mainland. The species reported herein are Atya scabra (Leach, 1815), Jonga serrei (Bouvier, 1909), Micratya poeyi (Guerin-Meneville, 1855), Potimirim glabra (Kingsley, 1878), P. potimirim (Muller, 1881) (Atyidae), Palaemon pandaliformis (Stimpson, 1871), Macrobrachium acanthurus (Wiegmann, 1836) and M. crenulatum Holthuis, 1950 (Palaemonidae). The record of J. serrei is the first for Panama, and M. poeyi and P. glabra the first for Bocas del Toro province.

Descriptions of two new species of alpheid shrimps from Japan and Australia, with notes on taxonomy of Automate De Man, Coronalpheus Wicksten and Bermudacaris Anker and Iliffe (Crustacea: Decapoda: Caridea)

Two new species of the caridean family Alpheidae are described from distant Indo-Pacific localities: Automate hayashii sp. nov. from Hakodate Bay, southern Hokkaido, Japan, and Bermudacaris australiensis sp. nov. from the North-West Shelf off Western Australia. Automate hayashii appears closest to the poorly known A. salomoni Coutière, 1908. Bermudacaris australiensis sp. nov., the second species of the genus, represents the first discovery of the genus in the Indo-Pacific. The type species of Bermudacaris Anker and Iliffe, 2000, B. harti Anker and Iliffe, 2000, was described from anchialine caves of Bermuda, while the unique specimen of Bermudacaris australiensis sp. nov. was collected from an apparently typical marine environment. Relationships among the species of Automate De Man, 1888, Bermudacaris and Coronalpheus Wicksten, 1999 are discussed. Characters separating these three closely related genera, including the development of the rostrum, the shape of the eye-stalks, the absence of the appendix masculina, and the features of the first pereopods, are reassessed. Three informal species groups are recognized in Automate, showing certain heterogeneity of this genus.

Description of Alpheus cedrici sp. n., a strikingly coloured snapping shrimp (Crustacea, Decapoda, Alpheidae) from Ascension Island, central Atlantic Ocean

Abstract Alpheus cedrici sp. n. is described based on two specimens collected under rocks while scuba diving off the coast of Ascension Island, central Atlantic Ocean. The new species belongs to the Alpheus macrocheles (Hailstone, 1835) species complex and appears to be most closely related to the eastern–central Atlantic Alpheus macrocheles, the western Atlantic Alpheus amblyonyx Chace, 1972, and the eastern Pacific Alpheus bellimanus Lockington, 1877 and Alpheus rectus Kim & Abele, 1988. However, it differs from all these species by a combination of morphological characters and by a diagnostic and striking colour pattern.

Decapod crustaceans from the state of Ceará, northeastern Brazil: an updated checklist of marine and estuarine species, with 23 new records

The present study is the first major assessment of the marine decapod fauna of Ceará, northeastern Brazil, since contributions of J. Fausto-Filho in the 1960s-1970s. A fully updated checklist of all decapod crustaceans occurring in marine and estuarine habitats of Ceará is provided, based on literature records, specimens held in two carcinological collections of the Universidade Federal do Ceará (UFC), and material collected mainly by the authors between 2011 and 2014. A total of 337 decapod species are listed, distributed among the following taxa: Achelata (8 species), Anomura (42 species), Astacidea (1 species), Axiidea (11 species), Brachyura (162 species), Caridea (83 species), Dendrobranchiata (20 species), Gebiidea (9 species), and Stenopodidea (1 species). Among them, 23 species represent new records for Ceará, with 14 species, viz. Alpheus peasei (Armstrong, 1940), A. thomasi Hendrix & Gore, 1973, Ambidexter symmetricus Manning & Chace, 1971, Axianassa australis Rodrigues & Shimizu, 1992, Biffarius biformis (Biffar, 1971), B. fragilis (Biffar, 1970), Leptalpheus axianassae Dworschak & Coelho, 1999, L. forceps Williams, 1965, Lysmata bahia Rhyne & Lin, 2006, L. intermedia (Kingsley, 1878), Merhippolyte americana Holthuis, 1961, Neocallichirus maryae Karasawa, 2004, Ogyrides hayi Williams, 1981, and Typton carneus Holthuis, 1951, now having Ceará as the northern-most limit in their distribution range along the Brazilian coastline. One shrimp species, Lysmata lipkei Okuno & Fiedler, 2010, which was also found in the state of Rio Grande do Norte, possibly represents an invasive taxon in Brazil and the western Atlantic, originating from the Indo-West Pacific. Alpheus buckupi Almeida, Terossi, Araújo-Silva & Mantelatto, 2013, previously recorded from Ceará based on a colour photograph, is confirmed from this state, with specimens from several new localities. A few doubtful records from Ceará are briefly discussed. Colour photographs are provided for most of the taxa newly recorded from Ceará; some species are illustrated in colour for the first time.

Re-examination of the eastern Pacific and Atlantic material of Alpheus malleator Dana, 1852, with the description of Alpheus wonkimi sp. nov. (Crustacea, Decapoda, Alpheidae)

The bumpy-clawed snapping shrimp, Alpheus malleator Dana, 1852 (Alpheidae), is revised based on the recently collected and older museum material from the eastern Pacific (Panama, Ecuador), Caribbean (Panama, Puerto Rico, Trinidad & Tobago), Brazil (São Paulo), and West Africa (Cape Verde, Senegal, Guinea, Equatorial Guinea, Congo). The eastern Pacific material is assigned to A. wonkimi sp. nov., based on one morphological difference in the colour and thickness of the uropodal spiniform seta, as well as previously published molecular data. The Caribbean, Brazilian and West African material is considered to represent a single, widespread, morphologically variable, amphi-Atlantic taxon, A. malleator. Alpheus pugilator A. Milne-Edwards, 1878 is retained as ajunior synonym of A. malleator, whereas A. tuberculosus Osorio, 1892, A. malleator var. edentatus Zimmer, 1913 and A. belli Coutière, 1898, the latter two based on juvenile specimens, are tentatively placed in the synonymy of A. malleator. Illustrations, including colour photographs, are provided for A. wonkimi sp. nov. and A. malleator and their morphological variability is discussed and illustrated.

Confirmation of the presence of Janicea antiguensis (Chace, 1972) (Decapoda: Barbouriidae) in northeastern and eastern Brazil

The barbouriid shrimp Janicea antiguesis (Chace, 1972) is reported from Porto de Galinhas and Tamandare in Pernambuco and Guarapari in Espirito Santo, Brazil. These records confirm the presence of J. antiguensis in northeastern and eastern Brazil, considerably extending its Brazilian range from Fernando de Noronha to southern Espirito Santo and also representing the first record of this species from mainland coastal reefs. In Brazil, J. antiguensis occurs in or near marine reef caves, typically at depths between 5 and 15 m, and can be most easily observed while scuba diving at night. Colour photographs of J. antiguensis from various Atlantic localities are provided and its colour pattern is compared to that of the closely related Indo-West Pacific barbouriid shrimp, Parhippolyte misticia (Clark, 1989). Some in situ observations are provided for the Porto de Galinhas population of J. antiguensis. Multiple observations of pairs with both individuals brooding embryos at different developmental stages suggest protandric simultaneous hermaphroditism in J. antiguensis.

A distinctive new species of the alpheid genus Salmoneus Holthuis, 1955 (Crustacea: Decapoda: Caridea) from the northwestern Pacific

A new species of the alpheid shrimp genus Salmoneus Holthuis, 1955, S. pinguis sp. nov., is described based on the holotype from Okinawa, Ryukyu Archipelago, Japan. An additional, much smaller and morphologically slightly different specimen from Panglao, Bohol Sea, the Philippines, is tentatively assigned to S. pinguis. sp. nov. The new species is one of the most unusual members of Salmoneus presenting a combination of several unique characters, including the unusually stout, plump body; the carapace strongly domed dorsally, with a broadly rounded rostrum, and without orbital teeth; the second abdominal pleuron conspicuously enlarged; the fourth pleuron greatly overlapping the fifth pleuron; the antennular and antennal flagella furnished with long setae; and the major cheliped comparatively small, and with the distally blunt, slightly bulbous fingertips.