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Cladocera: Anomopoda: Daphniidae) from the Lower Cretaceous of Australia

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Cladocera: Anomopoda: Daphniidae) from the Lower Cretaceous of Australia Palaeontologia Electronica palaeo-electronica.org Ephippia belonging to Ceriodaphnia Dana, 1853 (Cladocera: Anomopoda: Daphniidae) from the Lower Cretaceous of Australia Thomas A. Hegna and Alexey A. Kotov ABSTRACT The first fossil ephippia (cladoceran exuvia containing resting eggs) belonging to the extant genus Ceriodaphnia (Anomopoda: Daphniidae) are reported from the Lower Cretaceous (Aptian) freshwater Koonwarra Fossil Bed (Strzelecki Group), South Gippsland, Victoria, Australia. They represent only the second record of (pre-Quater- nary) fossil cladoceran ephippia from Australia (Ceriodaphnia and Simocephalus, both being from Koonwarra). The occurrence of both of these genera is roughly coincident with the first occurrence of these genera elsewhere (i.e., Mongolia). This suggests that the early radiation of daphniid anomopods predates the breakup of Pangaea. In addi- tion, some putative cladoceran body fossils from the same locality are reviewed; though they are consistent with the size and shape of cladocerans, they possess no cladoceran-specific synapomorphies. They are thus regarded as indeterminate diplostracans. Thomas A. Hegna. Department of Geology, Western Illinois University, Macomb, IL 61455, USA. ta- [email protected] Alexey A. Kotov. A.N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071, Russia and Kazan Federal University, Kremlevskaya Str.18, Kazan 420000, Russia. alexey-a- [email protected] Keywords: Crustacea; Branchiopoda; Cladocera; Anomopoda; Daphniidae; Cretaceous. Submission: 28 March 2016 Acceptance: 22 September 2016 INTRODUCTION tions that the sparse known fossil record does not correlate with a meager past diversity. The rarity of Water fleas (Crustacea: Cladocera) are small, the cladoceran fossils is probably an artifact, a soft-bodied branchiopod crustaceans and are a result of insufficient efforts to find them in known diverse and ubiquitous component of inland and new palaeontological collections (Kotov, aquatic communities (Dumont and Negrea, 2002). 2007). Over the past two decades, our knowledge Despite this, their pre-Quaternary fossil record is of fossil cladocerans has increased dramatically very sparse — owing to the fact that they are uni- due to the close examination of ancient lacustrine versally small (majority are less than 1 mm, a few sediments by cladocerans experts. The result of are about 5–6 mm long) and are hard to identify as this has been a flurry of new species and the dis- compression fossils. There are intriguing indica- Hegna, Thomas A. and Kotov, Alexey A. 2016. Ephippia belonging to Ceriodaphnia Dana, 1853 (Cladocera: Anomopoda: Daphniidae) from the Lower Cretaceous of Australia. Palaeontologia Electronica 19.3.40A: 1-9 palaeo-electronica.org/content/2016/1623-cretaceous-ephippia Copyright: Paleontological Society November 2016 HEGNA & KOTOV: CRETACEOUS EPHIPPIA covery of two extinct, order-level clades of cladoc- et al., 2005; Szeroczyńska and Sarmaja-Korjonen, erans, the Cryptopoda Kotov, 2007 and the 2007). Thus, the fossils discussed herein are kept Proanomopoda Kotov, 2013 (Kotov, 2007, 2013). in open nomenclature at the genus level. The spec- Two of the four major orders of cladocerans, imen was found while one of the authors (TAH) Haplopoda Sars, 1865 and Onychopoda Sars, was reviewing the figured specimens of the anos- 1865, have no pre-Quaternary fossil record, and a tracan species from the Koonwarra Fossil Bed (Jell third order, Ctenopoda Sars, 1865 (aside from and Duncan, 1986, further details below under Quaternary records) is only known from four mid- Occurrence). Specimens were photographed with to late Mesozoic Mongolian and Asian Russia a Leica and Zeiss binocular microscope and digital lacustrine localities (Kotov, 2007; Kotov and Kor- camera. The contrast was enhanced using Adobe ovchinsky, 2006). The most diverse modern order Photoshop. Drawings were produced using Adobe of cladocerans, the Anomopoda Sars, 1865, has a Illustrator. The modern specimens of Ceriodaphnia meager representation in the fossil record with a sp. and Daphnia magna were lyophilized (freeze- handful of species known from body (excluding dried), coated with gold, and imaged with a JEOL- ephippia) fossils (Smirnov, 1992; Kotov, 2007, 840A scanning electron microscope. 2009a, 2009b, 2013). The status of the purported Paleozoic fossil 'anomopod' Ebullitiocaris Ander- SYSTEMATIC PALAEONTOLOGY son, Crighton, and Hass, 2004 reported by Wom- Phylum ARTHROPODA Latreille, 1829 ack et al. (2012) is highly dubious as the position of Subphylum CRUSTACEA Brünnich, 1772 the “second antennae” relative to the carapace Class BRANCHIOPODA Latreille, 1817 conflicts with the body plan of all other known cla- Subclass DIPLOSTRACA Gerstaecker, 1866 docerans — living or extinct. ?Diplostraca indet. Anomopod egg cases or ephippia (dormant Figure 1.3-1.4 embryos), provide another glimpse into their fossil record—one with a better preservation potential Material. NMVP 109450 and NMVP 109462 from than fossils of the body. Ephippia enclose resting the Museum of Victoria, Australia. eggs (typically the product of sexual reproduction) Description. Only unclear impressions are found. encased in a protective outer covering formed from Some traces of the head (Figure 1.3; hd), valves part of the dorsal chitinous carapace of the mother (vv), postabdomen (pd), antennae II (aII), and tho- (Dumont and Negrea, 2002). Anomopods are the racic limbs (tl) are observable, but no detail could only group of cladocerans to produce such cases be unambiguously identified. In some impressions, (Kotov, 2009a, 2013). Some major groups of living there are obvious mandibles (Figure 1.4; md) of the anomopods (i.e., from the families Daphniidae branchiopod-type, asymmetrical, with bent distal Straus, 1820 and Moinidae Goulden, 1968) pro- end supplied with mastiscatory surface (Figure duce characteristic ephippia that are diagnostic 2.2). No further details are recognizable. In addi- down to the genus level (Berner, 1985; Scourfield, tion, the specimens seem to be strongly deformed, 1902; Kokkinn and Williams, 1987; Lu, 2001; and may represent either molted exoskeletons or Vanderkerhove et al., 2004; Mergeay et al., 2005). carcasses that had undergone significant decay of The aim of this study is to update the taxo- soft tissue prior to preservation. nomic status of some cladoceran remains from the Discussion. Jell and Duncan (1986) reported on freshwater lacustrine sediments of the Lower Cre- several fossil specimens identified as daphniid taceous Koonwarra Fossil Bed, Australia. body fossils. However, they list no daphniid-spe- cific synapomorphies to support their identifica- MATERIAL AND METHODS tions. Furthermore, many of their interpretive drawings suggest a chydorid-like body shape Although ephippia have a higher preserva- (round with a pointed rostrum) rather than a daph- tional potential than cladoceran body fossils, they niid-like shape (more elliptical, rounded head are not used to describe modern species — in fact, shield, often with a posterior apical spine). Size they are undescribed for a number of extant spe- and shape are really the only features that can be cies (Kotov, 2013). However, representative ephip- unambiguously described from these specimens— pia are known from many cladoceran genera, and and they are nearly impossible to translate into some of those genera are characterized by distinc- unambiguous synapomorphies. The specimens tive ephippial morphotypes that often allow for described above could also belong to Ceriodaph- identification at the genus level (Kokkin and Wil- nia (family Daphniidae), but no real evidence for liams, 1987, Vanderkerhove et al., 2004; Mergeay this could be found; the impressions are too 2 PALAEO-ELECTRONICA.ORG FIGURE 1. Fossil diplostracan remains from Koonwarra. 1, An image of a fossil ephippium belonging to Ceriodaphnia sp. from the Koonwarra Fossil Beds (photographed with obliquely directed light), NMV P332637, 2, Interpretive draw- ing of Figure 1.1, scale bar for 1-2 equals 0.1 mm. 3, Diplostracan indet, NMV P109450 (figure 70G and 71C in Jell and Duncan, 1986), See Figure 1.4 for scale bar. 4, Diplostracan indet, NMV P109462 (figure 70H and 71E in Jell and Duncan, 1986). Scale bar in 1.4 applies to 1.3 as well. Scale bar for 1.3-4 equals 1 mm. Abbreviations: aII - antenna II; hd - head; tl – thoracic limbs; vv - valves. deformed and badly preserved. The specimens Ceriodaphnia sp. could easily belong to other anomopod genera, Figure 1.1-1.2, Appendix 1 ctenopod cladocerans, or possibly even juvenile Material. NMV P332637, Museum of Victoria. The clam shrimp (Spinicaudata, Laevicaudata, or rock which contains this specimen also contains Cyclestherida). fossil anostracans (see Jell and Duncan, 1986, fig- Occurrence. Lower Cretaceous Koonwarra Fossil ure 73A). Additional specimens are figured in the Bed, Strzelecki Group, south Gippsland, Victoria, Appendix 1. Australia. The age of this bed is constrained to a Description. Ephippium shaped as half an oval maximum age of between 115±6 Ma and 118±5 Ma bisected along the short axis, straight dorsal mar- (Aptian) by fission track radiometric ages derived gin, length = 0.6 mm, height/length = 0.8, posterior from volcanogenic apatites isolated from above margin is symmetrically curved with a possible faint and below the fossil-bearing strata (Lindsay, 1982, rim preserved around the ventral margin. No orna- see Drinnan and Chambers, 1986). This is consis- mentation is preserved, as in some extant
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