Shift in symbiotic lifestyle as the major process shaping the evolution of pea crabs (Decapoda: Brachyura: Pinnotheroidea).

The pea crabs, superfamily Pinnotheroidea, are exceptional among brachyuran crabs in their diverse symbiotic associations involving both inquilinism and protective symbiosis. While this group presents a rare opportunity for evolutionary comparative study of host switching and morphological evolution in marine macroinvertebrates, previous phylogenetic studies have been focused on systematics. Here, we reconstructed the most extensive phylogeny of Pinnotheroidea based on two mitochondrial and six nuclear markers, with the aim of elucidating the host switching pathways and the correlation between symbiotic lifestyles and selected morphological adaptations. Ancestral state reconstruction of host association revealed a monophyletic origin of symbiosis in the form of inquilinism. Subsequent shifts in microhabitat preference for burrows or worm tubes, and the move to protective symbiosis, primarily in the switch to mollusc endosymbiosis, contributed to radiation in Pinnotheridae. Further parallel colonisations of echinoderms and tunicates occurred but did not lead to extensive diversification, except in the Clypeasterophilus + Dissodactylus lineage, which experienced a unique switch to echinoderm ectosymbiosis. The evolution of the third maxillipeds, carapace shape and ambulatory pereiopods suggests a rather strong coupling with the symbiotic lifestyle (whether inquilinism or protective symbiosis). Phenotypic diversity of these characters was higher among species engaged in protective symbiosis, with convergence in form (or function) among those sharing the same host affiliation. Species having different host affiliations or symbiotic lifestyles might also exhibit convergence in the form of the three morphological traits, suggesting a common adaptive value of the specialisations. Pinnotherid crabs overall exhibited a lower trait diversity than the also symbiotic palaemonid shrimps with comparable species diversity. This may plausibly be attributed to differences in potential for morphological modification to serve additional functions among the traits analysed in the two groups, the less frequent host switching and the less diverse host affiliations, and thus a less complicated evolutionary history in pinnotherids.

Evolution of protective symbiosis in palaemonid shrimps (Decapoda: Caridea) with emphases on host spectrum and morphological adaptations.

Palaemonidae is the most speciose caridean shrimp family, with its huge biodiversity partially generated via symbiosis with various marine invertebrates. Previous studies have provided insights into the evolution of protective symbiosis in this family with evidence for frequent inter-phyla host switches, but the comprehensiveness of evolutionary pathways is hampered by the resolution of the previous phylogenetic trees as well as the taxon coverage. Furthermore, several critical issues related to the evolution of a symbiotic lifestyle, including the change in host spectrum and corresponding morphological adaptations, remain largely unresolved. We therefore performed a much extended phylogenetic comparative study on Palaemonidae, rooted in a comprehensive phylogeny reconstructed by a supermatrix-supertree approach based on a total of three mitochondrial and five nuclear markers. Ancestral state reconstruction of host associations revealed at least three independent evolutions into symbiosis, with potentially a drive to seek protection fuelling incipient symbiosis. Yet, most of the observed symbiotic species diversity was radiated from a single cnidarian associate. The evolution of mandibles and ambulatory dactyli suggests a general lack of correlation with host affiliation (except sponge endosymbionts), implying limited morphological adaptations following host switching, despite being putatively a major adaptive consequence of symbiosis. Our analyses of host spectrum, in terms of basic and taxonomic specificity, revealed no apparent phylogenetic signal but instead resolved a dynamic pattern attributable to frequent host switching. Uncoupling between host spectrum and the degree of morphological specialisation is the norm in palaemonids, suggesting that morphological characters are not fully in tune with host spectrum, in addition to host affiliation. This study demonstrates the complexity in the evolution of symbiosis, pointing to the presence of cryptic adaptations determining host spectrum and governing host switch diversification, and provides a clear direction for the evolutionary study of symbiosis in other marine symbiotic groups involving host switching.

An Updated Checklist of Sea Slugs (Gastropoda, Heterobranchia) from Hong Kong Supported by Citizen Science.

Research on heterobranch sea slug diversity in Hong Kong was flourishing in the 1980-90s, with the first checklist and guidebook published and many new species and records documented later. This golden era of sea slug research ended abruptly, and no review nor any extensive collection has been done in the past 20+ years. In view of the extensive revision in heterobranch systematics in recent years, the current checklist can be considered out of date in both nomenclature and composition. Therefore, we are dedicated to providing a synthesis of the current state of knowledge on Hong Kong sea slug diversity by reviewing previous records and engagement of citizen science. An integration of previously published and unpublished records revealed the presence of 186 species, of which 107 species were documented by our citizen science programme with a further report of 71 new records. This yields a total of 257 species from 126 genera, 53 families and ten higher taxa, of which seven species have not been reported elsewhere from Hong Kong. While the present study has markedly illuminated our understanding of sea slug diversity in Hong Kong, the true biodiversity may be underestimated given the high proportion of novel records, especially when many previous records were not herein observed. Hong Kong likely harbours a greater sea slug diversity which awaits discovery from more rigorous sampling and a more inclusive citizen science programme.

Distinct suites of pre- and post-adaptations indicate independent evolutionary pathways of snapping claws in the shrimp family Alpheidae (Decapoda: Caridea).

One of the most notable evolutionary innovations of marine invertebrates is the snapping claw of alpheid shrimps (Alpheidae), capable of generating a powerful water jet and a shock wave, used for defense, aggression, excavation, and communication. Evolutionary analysis of this character complex requires the study of a suite of complementary traits to discern pre-adaptations or post-adaptations of snapping behavior. A comprehensive phylogenetic analysis of the Alpheidae based on two mitochondrial and four nuclear markers, covering 107 species from 38 genera (77.6% generic coverage), is presented. Ancestral state reconstruction analyses revealed five independent origins of snapping, two of which relate to the morphologically similar but phylogenetically distant genera Alpheus and Synalpheus, highlighting significant convergence. The evolution of the five complementary traits (adhesive plaques, tooth-cavity system, dactylar joint type, chela size enlargement, and orbital hood) did not always show a significant correlation with the evolution of snapping overall, sometimes only in a few lineages, suggesting different evolutionary pathways were involved and demonstrating the versatility in the evolution of the snapping mechanisms.

Isolation and Characterization of Polymorphic Microsatellite Loci for Caridina cantonensis and Transferability Across Eight Confamilial Species (Atyidae, Decapoda).

Lai Him Chow, Ka Yan Ma, Jerome H. L. Hui, and Ka Hou Chu (2018) Caridina cantonensis is a common freshwater shrimp found in Guangdong Province, China. The species is landlocked, and this life history contributes substantially to its strong population differentiation at the very small geographical scale. Given its widespread distribution, it serves as an excellent model for examining population connectivity and phylogeographical history of freshwater invertebrates in South China. This study isolated ten microsatellite loci from genome sequences of C. cantonensis and four from the transcriptome assemblies of Neocaridina davidi. The microsatellites were then characterized in 24 C. cantonensis individuals from one population. The number of alleles ranged from seven to 18. Observed and expected heterozygosities ranged from 0.050 to 0.958 (averaged 0.649) and 0.754 to 0.919 (averaged 0.833), respectively. We also assessed the cross-species transferability of the markers developed across eight confamilial species. On average, nine markers can be amplified in each species, and five markers can be amplified across all eight species. The markers developed in this study would enable evaluation of genetic diversity and population structure of these species for conservation management planning.