George D. F. Wilson

Arthropod Cladistics: Combined Analysis of Histone H3 and U2 snRNA Sequences and Morphology

Morphological, developmental, ultrastructural, and gene order characters are catalogued for the same set of arthropod terminals as we have scored in a recent study of histone H3 and U2 snRNA sequences (D. J. Colgan et al., 1998, Aust. J. Zool. 46, 419–437). We examine the implications of separate and simultaneous analyses of sequence and non‐sequence data for arthropod relationships. The most parsimonious trees based on 211 non‐sequence characters (273 apomorphic states) support traditional higher taxa as clades, including Mandibulata, Crustacea, Atelocerata, Myriapoda, and Hexapoda. Combined analysis of morphology with histone H3 and U2 sequences with equal character weights differs from the morphological results alone in supporting Progoneata + Hexapoda (= Labiophora) in favor of a monophyletic Myriapoda, resolves the entognathous hexapods as a grade, and supports pycnogonids as sister group to Euchelicerata (rather than as basal euarthropods). Monophyly of Chelicerata (including pycnogonids), Mandibulata, Crustacea, Progoneata, Chilopoda, and Hexapoda is maintained under a range of transition/transversion and third codon weights, whereas Atelocerata and Myriapoda/Labiophora do not withstand all sensitivity analyses.

Historical influences on deep-sea isopod diversity in the Atlantic Ocean

Most isopod crustaceans in the North Atlantic deep sea belong to the suborder Asellota. In contrast, South Atlantic isopod faunas have a significant component of flabelliferan isopods, a phylogenetic clade that contains suborders derived evolutionarily later than the Asellota. The flabelliferans decrease diversity from shallow water to deep water and on a south-to-north latitudinal gradient. Although many asellote families are endemic to the deep sea, none of the flabelliferan families appear to have evolved in the abyss. Recent colonisations of the deep sea, which may have been limited to the southern hemisphere by oceanographic conditions, have significant consequences for observed regional diversities of some taxa. Instability in oceanographic conditions owing to glaciation and benthic storms may have further limited benthic species richness of the North Atlantic deep-sea benthos.

Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater

The isopod crustaceans are diverse both morphologically and in described species numbers. Nearly 950 described species (∼9% of all isopods) live in continental waters, and possibly 1,400 species remain undescribed. The high frequency of cryptic species suggests that these figures are underestimates. Several major freshwater taxa have ancient biogeographic patterns dating from the division of the continents into Laurasia (Asellidae, Stenasellidae) and Gondwana (Phreatoicidea, Protojaniridae and Heterias). The suborder Asellota has the most described freshwater species, mostly in the families Asellidae and Stenasellidae. The suborder Phreatoicidea has the largest number of endemic genera. Other primary freshwater taxa have small numbers of described species, although more species are being discovered, especially in the southern hemisphere. The Oniscidea, although primarily terrestrial, has a small number of freshwater species. A diverse group of more derived isopods, the ‘Flabellifera’ sensu lato has regionally important species richness, such as in the Amazon River. These taxa are transitional between marine and freshwater realms and represent multiple colonisations of continental habitats. Most species of freshwater isopods species and many genera are narrow range endemics. This endemism ensures that human demand for fresh water will place these isopods at an increasing risk of extinction, as has already happened in a few documented cases.

The lacinia mobilis and Similar Structures - a Valuable Character in Arthropod Phylogenetics?

Abstract Homology between the lacinia mobilis of peracarid crustaceans (Malacostraca) and movable appendages of the mandibular edge in other crustaceans, hexapods, and myriapods has been advocated in classical as well as recent phylogenetic studies, and in some cases this feature has been attributed major significance in arthropod systematics. A comparative SEM survey of the lacinia mobilis in Peracarida and its alleged homologues (prostheca of Hexapoda, internal tooth of Diplopoda, ‘lacinia mobilis’ of Symphyla and Remipedia) rejects the primary homology of these varied structures. The lacinia mobilis of Peracarida can be characterized precisely by asymmetry on the left and right mandibles, as a strong tooth-like structure on the left mandible which is oriented at a right angle to the remaining edge and as a stalked, spine-like structure on the right mandible. A fundamental difference to other Malacostraca is that the peracarid lacinia mobilis characterizes the adult mandibles. Supposed homologues of the peracarid lacinia mobilis are instead a modified pectinate lamella (Diplopoda, Chilognatha) or a movable appendage that is associated with the pars molaris rather than the pars incisivus (Symphyla), or an inner separation of the incisor process (Remipedia). The detailed structure of the prostheca in Campodeoida and Ephemeroptera weakens interpretations of its origin as a separation of the incisor process.

Is the HEBBLE isopod fauna hydrodynamically modified? A second test

Abstract Several times per year, the High Energy Benthic Boundary Layer Experiment (HEBBLE) site (4820 m depth, 40°27′N 62°20′W) experiences benthic storms during which near-bottom flows can erode millimeters of sediment. Thistle and Wilson (Deep-Sea Research, 34 1987, 73–87) predicted that isopods that inhabited the surface of the sediment would be relatively rare at the HEBBLE site compared to those at quiescent deep-sea sites. They tested this prediction by comparing the composition of the HEBBLE isopod fauna to that of a quiescent site and found a significant difference in the predicted direction. Although this result was encouraging, the strength of their inference was limited because only one site from each type of environment had been compared. We performed a second test of Thistle and Wilsons hypothesis by comparing the composition of the isopod fauna from two additional physically quiescent locations (4500 m depth, 14°40′N 125°26′W, and 4800 m depth, 12°57′N 128°19.5′W) to that of the HEBBLE site. Those isopods that are thought to be exposed to the erosion caused by storms occurred in a significantly greater proportion of the samples at the quiescent sites than at the HEBBLE site, a result consistent with Thistle and Wilsons hypothesis.

Evidence for Permo- Triassic colonization of the deep sea by isopods

The deep sea is one of the largest ecosystems on Earth and is home to a highly diverse fauna, with polychaetes, molluscs and peracarid crustaceans as dominant groups. A number of studies have proposed that this fauna did not survive the anoxic events that occurred during the Mesozoic Era. Accordingly, the modern fauna is thought to be relatively young, perhaps having colonized the deep sea after the Eocene/Oligocene boundary. To test this hypothesis, we performed phylogenetic analyses of nuclear ribosomal 18S and 28S and mitochondrial cytochrome oxidase I and 16S sequences from isopod crustaceans. Using a molecular clock calibrated with multiple isopod fossils, we estimated the timing of deep-sea colonization events by isopods. Our results show that some groups have an ancient origin in the deep sea, with the earliest estimated dates spanning 232–314 Myr ago. Therefore, anoxic events at the Permian–Triassic boundary and during the Mesozoic did not cause the extinction of all the deep-sea fauna; some species may have gone extinct while others survived and proliferated. The monophyly of the ‘munnopsid radiation’ within the isopods suggests that the ancestors of this group evolved in the deep sea and did not move to shallow-water refugia during anoxic events.

Gondwanan groundwater: subterranean connections of Australian phreatoicidean isopods (Crustacea) to India and New Zealand

Phreatoicidea Stebbing, 1893 live in freshwaters of Gondwana: Australia, South Africa, India and New Zealand. Many of these isopods have a subterranean lifestyle. Parsimony analysis of morphological data of generic exemplars and a Triassic fossil was used to explore the timing of this habitat adaption. The monophyly of the Hypsimetopidae Nicholls, 1943, including blind taxa Hyperoedesipus Nicholls & Milner, 1923 (Western Australia), Nichollsia Chopra and Tiwari, 1950 (Ganges Plain, India) and Phreatoicoides Sayce, 1900 (Tasmania and Victoria) was strongly supported. Crenisopus Wilson and Keable, 1999 (Kimberleys, Western Australia) and the PonderellidaeWilson & Keable, 2004 (Queensland mound springs) may be sister to hypsimetopids. Blind Phreatoicidae found only in south-eastern Australia and in New Zealand were also monophyletic. The hypogean habitat, blindness, fossil and plate tectonic evidence were mapped on the cladogram to estimate timing of this adaptation. A subterranean adaptation before 130 million years ago was supported for hypsimetopids. Phreatoicus Chilton, 1891 and Neophreatoicus Nicholls, 1944 (hypogean in New Zealand) were in a monophyletic clade with epigean Phreatoicidae, Crenoicus Nicholls, 1944 (south-eastern Australia) and Notamphisopus Nicholls, 1943 (New Zealand). Blindness in epigean taxa is consistent with recolonisation of surface waters from underground refuges. Because Crenoicus is sister-group to the New Zealand clade, and because overseas dispersal between Australia and New Zealand is unlikely, the minimum age for these blind phreatoicids is ~80 million years. This evidence is consistent with a subterranean freshwater fauna surviving the presumed Oligocene inundation of New Zealand.

THE TRIASSIC ISOPOD PROTAMPHISOPUS WIANAMATTENSIS (CHILTON) AND COMPARISON WITH EXTANT TAXA (CRUSTACEA, PHREATOICIDEA)

Abstract Protamphisopus wianamattensis (Chilton, 1918) from the Middle Triassic (Anisian) Ashfield Shale in the Sydney Basin, Australia, is the earliest-known freshwater representative of the basal isopod suborder Phreatoicidea. In contrast, the late Paleozoic Palaeophreatoicidae species, which are morphologically distinct from extant families, are found in marine or estuarine facies. Comparison of Protamphisopus wianamattensis with living Phreatoicidea permits the external morphology of the fossils to be reconstructed and the species to be coded for cladistic analysis using a revised and expanded character set developed for living phreatoicideans. In resulting parsimonious trees as well as immediately suboptimal trees, Protamphisopus is nested within clades related to the family Amphisopodidae. Although not included in the analysis, the Late Permian Protamphisopus reichelti Malzahn (in Glaessner and Malzahn, 1962) appears to be a member of the Palaeophreatoicidae, rather than among the crown group of the Phreatoicidea. Therefore, a minimum age of Middle Triassic can be assigned to the basal branches within the phreatoicid crown group. The minimum age for the colonisation of fresh water by the suborder is also established although, given the advanced position of Protamphisopus wianamattensis in the cladograms, the habitat shift may have occurred earlier. The biogeographic distribution of extant Phreatoicidea on fragments of Gondwana is consistent with early Mesozoic origins for the major clades of this isopod suborder.

Monsoon-influenced speciation patterns in a species flock of Eophreatoicus Nicholls (Isopoda; Crustacea)

A species flock of the freshwater isopod genus Eophreatoicus Nicholls lives in seeps, springs and perched aquifers at the base of the Arnhem Plateau and associated sandstone outliers in Australias Northern Territory. These species have been found to have surprisingly high levels of genetic divergence and narrow range endemism, despite potential opportunities for dispersion during the summer monsoon season when streams flow continuously and have connectivity. Species of Eophreatoicus were identified morphologically as distinct taxa, sometimes with two or three species occurring at the same site. DNA sequence data from the mitochondrial 16S rRNA and cytochrome c oxidase subunit I genes corroborate our morphological concepts to a high level of resolution, with the exception of two distinct species that are identical genetically. The value of mtDNA data for identification of these species, therefore, is limited. These isopods disperse downstream from their home springs to a limited extent during the wet season, but the genetic data show that migration to non-natal springs, and reproduction there, may be rare. We argue that the multiplication of the narrow-range endemic species is the result of their homing behaviour combined with monsoonal alternation between aridity and flooding over recent and geological time scales since the Miocene period.

Occurrence of the Isopod Archaeoniscus coreaensis New Species from the Lower Cretaceous Jinju Formation, Korea

Abstract The fossil isopod crustacean genus Archaeoniscus has been known to occur in England, France and Germany during the Upper Jurassic, and in Mexico and Egypt during the Lower Cretaceous. The morphology of this genus is unique in having dorsoventrally compressed body, the cephalon set deeply into the first pereionite, pleon as wide as pereion, and a broad semicircular pleotelson. These features have resulted in placing the classification of the genus in the monotypic family Archaeoniscidae. However, due to the lack of detailed morphological data, suprafamilial classification of this genus has remained unclear, as well as its ecology and lifestyle. Here we report Archaeoniscus coreaensis n. sp. from the Jinju Formation, Gyeongsang Basin, Korea. The occurrence of Archaeoniscus in the East Asia implies that the genus may have had a worldwide distribution. The Gyeongsang Basin was a Cretaceous backarc basin, which consists of exclusively non-marine sedimentary sequences. The occurrence of this genus, therefore, indicates that Archaeoniscus successfully adapted to a freshwater ecosystem as well. Detailed anatomy including antennulae, antennae, pereiopods, and uropods was observed from well-preserved multiple specimens, which allows better understanding of the morphology of Archaeoniscus. The axial structure in the posterior part of the body, which was previously interpreted as a unique brood pouch characterizing the family, turned out to be a remnant of the hindgut. Females of all isopods and most of the members of the superorder Peracarida have a thoracic ventral brood pouch, modified from the thoracic coxal endites. Based on the morphology of the largely unmodified ambulatory pereiopods of A. coreaensis, the possibility of Archaeoniscus being ectoparasitic is discounted. Instead, the flattened body and the form of limbs of A. coreaensis would have been suitable for a benthic lifestyle.