Branchiopod crustaceans are represented by fairy, tadpole, and clam shrimps (Anostraca, Notostraca, Laevicaudata, Spinicaudata), which typically inhabit temporary freshwater bodies, and water fleas (Cladoceromorpha), which live in all kinds of freshwater and occasionally marine environments [1, 2]. The earliest branchiopods occur in the Cambrian, where they are represented by complete body fossils from Sweden such as Rehbachiella kinnekullensis [3] and isolated mandibles preserved as small carbonaceous fossils [4-6] from Canada. The earliest known continental branchiopods are associated with hot spring environments [7] represented by the Early Devonian Rhynie Chert of Scotland (410 million years ago) and include possible stem-group or crown-group Anostraca, Notostraca, and clam shrimps or Cladoceromorpha [8-10], which differ morphologically from their modern counterparts [1, 2, 11]. Here we report the discovery of an ephemeral pool branchiopod community from the 365-million-year-old Strud locality of Belgium. It is characterized by new anostracans and spinicaudatans, closely resembling extant species, and the earliest notostracan, Strudops goldenbergi [12]. These branchiopods released resting eggs into the sediment in a manner similar to their modern representatives [1, 2]. We infer that this reproductive strategy was critical to overcoming environmental constraints such as seasonal desiccation imposed by living on land. The pioneer colonization of ephemeral freshwater pools by branchiopods in the Devonian was followed by remarkable ecological and morphological stasis that persists to the present day.

After terrestrialization, the diversification of arthropods and vertebrates is thought to have occurred in two distinct phases, the first between the Silurian and the Frasnian stages (Late Devonian period) (425-385 million years (Myr) ago), and the second characterized by the emergence of numerous new major taxa, during the Late Carboniferous period (after 345 Myr ago). These two diversification periods bracket the depauperate vertebrate Romer's gap (360-345 Myr ago) and arthropod gap (385-325 Myr ago), which could be due to preservational artefact. Although a recent molecular dating has given an age of 390 Myr for the Holometabola, the record of hexapods during the Early-Middle Devonian (411.5-391 Myr ago, Pragian to Givetian stages) is exceptionally sparse and based on fragmentary remains, which hinders the timing of this diversification. Indeed, although Devonian Archaeognatha are problematic, the Pragian of Scotland has given some Collembola and the incomplete insect Rhyniognatha, with its diagnostic dicondylic, metapterygotan mandibles. The oldest, definitively winged insects are from the Serpukhovian stage (latest Early Carboniferous period). Here we report the first complete Late Devonian insect, which was probably a terrestrial species. Its 'orthopteroid' mandibles are of an omnivorous type, clearly not modified for a solely carnivorous diet. This discovery narrows the 45-Myr gap in the fossil record of Hexapoda, and demonstrates further a first Devonian phase of diversification for the Hexapoda, as in vertebrates, and suggests that the Pterygota diversified before and during Romer's gap.

A new notostracan crustacean, Strudops goldenbergi gen. et sp. nov., is described from the well-preserved terrestrial arthropod fauna of the Upper Devonian of Strud, Belgium. The fossil notostracan bears a close resemblance to modern notostracans in possessing a large, simple head shield covering almost half of the whole body, a set of phyllopodous thoracic appendages and a legless posterior abdomen with a telson bearing a caudal furca. The differentiation and relative size of mouthparts and limbs suggest that these specimens are all adults. The notostracans described herein are the earliest clear members of the total group Notostraca.

The ancestral states of bilaterian development, and which living groups have conserved them the most, has been a controversial topic in biology for well over a hundred years. In recent years, the idea that gastrulation primitively proceeded via the formation of a slit-like blastopore that then evolved into either protostomy or deuterostomy has gained renewed attention and some molecular developmental support. One of the key pieces of evidence for this 'amphistomy' theory comes from the onychophorans, which form a clear ventral groove during gastrulation. The interpretation of this structure has, however, proved problematic. Based on expression patterns of forkhead ( fkh), caudal (cad), brachyury (bra) and wingless (wg/Wnt1), we show that this groove does not correspond to the blastopore, even though both the mouth and anus later develop from it. Rather, the posterior pit appears to be the blastopore; the posterior of the groove later fuses with it to form the definitive anus. Onychophoran development therefore represents a case of 'concealed' deuterostomy. The new data from the onychophorans thus remove one of the key pieces of evidence for the amphistomy theory. Rather, in line with other recent results, it suggests that ancestral bilaterian development was deuterostomic.

The Non-Mineralized arthropod described herein is derived from the Sirius Passet fossil conservation deposit of North Greenland (82°47.6,N, 42°13.7ʹW), the oldest locality with exceptional preservation of soft tissues known from the Cambrian of Laurentia (Cambrian Series 2, Stage 3; Nevadella Zone). As such, it is broadly contemporaneous with the Chengjiang fauna of China (Hou et al., 2004) and some 10 million years older than the Burgess Shale fauna of British Columbia. The Sirius Passet fauna was first documented by Conway Morris et al. (1987) and its geological setting is discussed by Babcock and Peel (2007). In addition to the nevadiid trilobite Buenellus higginsi Blaker, 1988, the fauna is dominated by non-mineralized arthropods (Budd, 1993, 1995, 1997, 1999; Williams et al., 1996; Taylor, 2002). Other finds include sponges (Rigby, 1986), a lobopod (Budd and Peel, 1998), the earliest annelids (Conway Morris and Peel, 2008) and articulated halkieriids (Conway Morris and Peel, 1990, 1995), but most of the assemblage awaits description.