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Table 2 in New species and new records of camallanid nematodes (Nematoda, Camallanidae) from marine fishes and sea snakes in New Caledonia
<p><b>Table 2.</b> Comparison of measurements of <i>Camallanus carangis</i> females from marine fishes in New Caledonia.</p><table><tbody><tr><th>Host</th><th><i>Atule</i></th><th><i>Carangoides</i></th><th><i>Selar</i></th><th><i>Parupeneus</i></th><th><i>Pristipomoides</i></th><th><i>Epinephelus</i></th></tr></tbody><tbody><tr><th></th><td><i>mate</i></td><td><i>dinema</i></td><td><i>crumenophthalmus</i></td><td><i>cyclostomus</i></td><td><i>filamentosus</i></td><td><i>retouti</i></td></tr><tr><th>No. of specimens 1 (grav.)</th><td>2 (subgr.)</td><td>3 (gr., subgr.)</td><td>2 (subgr.)</td><td>1 (subgr.)</td><td>1 (subgr.)</td></tr><tr><th>Body length (in mm)</th><td>14.93</td><td>4.75 –7.00</td><td>14.63–17.00</td><td>10.23–11.13</td><td>14.89</td><td>10.95</td></tr><tr><th>Body width</th><td>340</td><td>163–245</td><td>381–517</td><td>340–408</td><td>313</td><td>354</td></tr><tr><th>Buccal capsule – length</th><td>195</td><td>159–180</td><td>186–225</td><td>159</td><td>150</td><td>183</td></tr><tr><th>Buccal capsule – width</th><td>198</td><td>144–180</td><td>204–233</td><td>144</td><td>159</td><td>195</td></tr><tr><th>No. of ridges</th><td>32</td><td>37–40</td><td>35–50</td><td>?</td><td>32</td><td>44</td></tr><tr><th>Basal ring – length</th><td>30</td><td>27</td><td>24–36</td><td>21–24</td><td>24</td><td>30</td></tr><tr><th>Basal ring – width</th><td>111</td><td>96</td><td>108–117</td><td>96–99</td><td>90</td><td>108</td></tr><tr><th>Length of tridents</th><td>141</td><td>135</td><td>195–240</td><td>105–195</td><td>141</td><td>153</td></tr><tr><th>Oesoph. cup – length</th><td>27</td><td>24–30</td><td>30–39</td><td>33</td><td>15</td><td>36</td></tr><tr><th>Oesoph. cup – width</th><td>36</td><td>30–36</td><td>39–45</td><td>36</td><td>33</td><td>30</td></tr><tr><th>Musc. oesoph. – length</th><td>1550</td><td>721–1047</td><td>1333–1850</td><td>816–979</td><td>911</td><td>1469</td></tr><tr><th>Musc. oesoph. – width</th><td>136</td><td>90</td><td>123–177</td><td>122–136</td><td>163</td><td>122</td></tr><tr><th>Gland. oesoph. – length</th><td>1333</td><td>653–911</td><td>1156–1659</td><td>775–925</td><td>993</td><td>1156</td></tr><tr><th>Gland. oesoph. – width</th><td>136</td><td>96–99</td><td>114–190</td><td>122–163</td><td>163</td><td>136</td></tr><tr><th>Musc./gland. oesoph.</th><td>1:0.86</td><td>1:0.87–0.91</td><td>1:0.87–0.98</td><td>1:0.94–0.95</td><td>1:1.09</td><td>1:0.79</td></tr><tr><th>length ratio</th></tr><tr><th>% of buc. c. and oesoph.</th><td>21</td><td>22–40</td><td>18–25</td><td>17–19</td><td>14</td><td>26</td></tr><tr><th>of body</th></tr><tr><th>Excretory pore</th><td>1401</td><td>707–911</td><td>1591</td><td>?</td><td>?</td><td>1224</td></tr><tr><th>Vulva from ant. end (in</th><td>7.51</td><td>2.56–3.93</td><td>7.13–8.80</td><td>4.92</td><td>7.00</td><td>5.74</td></tr><tr><th>mm)</th></tr><tr><th>% of vulva of body</th><td>50</td><td>54–56</td><td>49–55</td><td>48</td><td>47</td><td>52</td></tr><tr><th>Tail</th><td>272</td><td>90–136</td><td>225</td><td>144–147</td><td>177</td><td>190</td></tr></tbody></table>
Table 1 in New species and new records of camallanid nematodes (Nematoda, Camallanidae) from marine fishes and sea snakes in New Caledonia
<p><b>Table 1.</b> Comparison of measurements of <i>Camallanus carangis</i> males from marine fishes in New Caledonia.</p><table><tbody><tr><th>Host</th><th><i>Atule mate</i></th><th><i>Carangoides fulvoguttatus</i></th><th><i>Cephalopholis sonnerati</i></th><th><i>Epinephelus retouti</i></th></tr></tbody><tbody><tr><th>No. of specimens</th><td>2</td><td>1</td><td>3</td><td>1</td></tr><tr><th>Body length (in mm)</th><td>12.17–13.67</td><td>6.90</td><td>9.93–11.32</td><td>frag. 5.40</td></tr><tr><th>Body width</th><td>299–354</td><td>218</td><td>258–326</td><td>258</td></tr><tr><th>Buccal capsule – length</th><td>165–189</td><td>162</td><td>183–186</td><td>159</td></tr><tr><th>Buccal capsule – width</th><td>159–189</td><td>156</td><td>156–183</td><td>165</td></tr><tr><th>No. of ridges</th><td>33–35</td><td>38</td><td>34–35</td><td>40</td></tr><tr><th>Basal ring – length</th><td>21–24</td><td>27</td><td>27–33</td><td>27</td></tr><tr><th>Basal ring – width</th><td>90–105</td><td>90</td><td>96–102</td><td>93</td></tr><tr><th>Length of tridents</th><td>150</td><td>150</td><td>171–219</td><td>153</td></tr><tr><th>Oesoph. cup – length</th><td>21–30</td><td>30</td><td>30–36</td><td>36</td></tr><tr><th>Oesoph. cup – width</th><td>24–33</td><td>39</td><td>36–39</td><td>30</td></tr><tr><th>Musc. oesoph. – length</th><td>1183–1414</td><td>1020</td><td>1333–1170</td><td>1102</td></tr><tr><th>Musc. oesoph. – width</th><td>122–136</td><td>81</td><td>122–136</td><td>84</td></tr><tr><th>Gland. oesoph. – length</th><td>1156–1333</td><td>857</td><td>1034–1238</td><td>1673</td></tr><tr><th>Gland. oesoph. – width</th><td>136</td><td>96</td><td>122–136</td><td>90</td></tr><tr><th>Musc./gland. oesoph. length ratio</th><td>1:0.94–0.98</td><td>1:084</td><td>1:0.88–0.93</td><td>1:0.79</td></tr><tr><th>% of buc. c. and oesoph. of body</th><td>21</td><td>30</td><td>23–24</td><td>–</td></tr><tr><th>Excretory pore</th><td>1047–1387</td><td>?</td><td>1115–1251</td><td>?</td></tr><tr><th>Right spicule</th><td>306–309</td><td>300</td><td>294–312</td><td>–</td></tr><tr><th>Tail</th><td>75–81</td><td>102</td><td>84–117</td><td>–</td></tr></tbody></table>
Sexual dimorphism in size and shape of the head in the sea snake Emydocephalus annulatus
<p>In snakes, divergence in head size between the sexes has been interpreted as an adaptation to intersexual niche divergence. By overcoming gape-limitation, a larger head enables snakes of one sex to ingest larger prey items. Under this hypothesis, we do not expect a species that consumes only tiny prey items to exhibit sex differences in relative head size, or to show empirical links between relative head size and fitness-relevant traits such as growth and fecundity. Our field studies on the sea snake <i>Emydocephalus annulatus</i> falsify these predictions. Although these snakes feed exclusively on fish eggs, the heads of female snakes are longer and wider than those of males at the same body length. Individuals with wider heads grew more rapidly, reproduced more often, and produced larger litters. Thus, head shape can affect fitness and can diverge between the sexes even without gape-limitation. Head size and shape may facilitate other aspects of feeding (such as the ability to scrape eggs off coral) and locomotion (hydrodynamics); and a smaller head may advantage the sex that is more mobile, and that obtains its prey in narrow crevices rather than in more exposed situations (i.e., males).</p>
Data for: Sex-based divergence in tidal, lunar and seasonal cycles of activity in the olive sea snake, Aipysurus laevis (Elapidae, Hydrophiinae)
<p>Marine environments show strong cycles at daily (tidal), monthly (lunar) and seasonal timeframes, and the behavioural responses of marine organisms to such cycles may depend upon ecological and behavioural traits that differ between the sexes. Underwater observations of free-ranging olive sea snakes (Aipysurus laevis, Hydrophiinae, Elapidae) at a site on the southern Great Barrier Reef revealed sex-based divergences in the effects of abiotic cycles on snake activity. Female snakes were active primarily on high and rising tides that allowed access to shallow-water sites for foraging. In contrast, male snakes were active primarily on low and falling tides, especially near the time of the full moon (when tidal range is highest), conditions that may restrict a female snake's ability to evade a courting male. Males were common on the coral-reef site during winter (the mating season), but were rarely seen during summer, whereas females remained on the reef year-round. This highly sexually dimorphic species shows strong temporal separations between the sexes in patterns of activity.</p>
Sexual dimorphism in aipysurine sea snakes
<p class="MsoNormal">The transition from terrestrial to aquatic life by hydrophiine elapid snakes modified targets of natural selection and likely affected sexual selection also. Thus, the shift to marine life also might have affected sexual dimorphism. Our measurements of 419 preserved specimens of six species of aipysurine snakes (genera <em>Emydocephalus</em> and <em>Aipysurus</em>) revealed sexual dimorphism in mean adult snout-vent length (= SVL), body width relative to SVL, lengths and widths of heads and tails relative to SVL, and eye diameter relative to head length. Females averaged larger than males in all taxa, and generally were wider-bodied with shorter and wider tails and smaller eyes. For other traits, sexual dimorphism varied among species: for example, relative head length ranged from male-biased to female-biased, and head shape (width relative to length) was highly dimorphic only in <em>A. laevis</em>. The transition to marine life appears to have eliminated male-male combat (reducing selection for large males) and favoured visual rather than pheromone-based mate-searching (favouring larger eyes in males). Variation in head-size dimorphism may reflect intersexual niche partitioning, with different taxa following different trajectories. Repeated evolutionary transitions from terrestrial to aquatic life in snakes provide a powerful opportunity to explore selective forces on sexually dimorphic traits.</p>
Sexual dimorphism in size and shape of the head in the sea snake Emydocephalus annulatus
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Swim with the tide: tactics to maximise prey detection by a specialist predator, the greater sea snake (Hydrophis major)
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Data for: Sex-based divergence in tidal, lunar and seasonal cycles of activity in the olive sea snake, Aipysurus laevis (Elapidae, Hydrophiinae)
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Sexual dimorphism in aipysurine sea snakes
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Colour polymorphism in the sea snake Emydocephalus annulatus
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FIGURE 2 in Description of a new snake eel Ophichthus olivaceus (Teleostei: Anguilliformes Ophichthidae) from the Red Sea
FIGURE 2. Ophichthus olivaceus sp. nov., SMF 35878 [KAU17-4], holotype. A: underside of the head; B: head and anterior trunk close-up; C: cephalic sensory pores system; IO-infraorbital pores, LL-lateral line pores, M-lower jaw pores, PO-preopercular pores, SO-supraorbital pores, ST-supratemporal pores. Photos by S.V. Bogorodsky (A & B), drawing by J.E. McCosker (C).
FIGURE 4 in Description of a new snake eel Ophichthus olivaceus (Teleostei: Anguilliformes Ophichthidae) from the Red Sea
FIGURE 4. Maximum Likelihood phylogenetic tree based on partial sequences (652 bp) of the mitochondrial COI gene includ- ing available sequences of Ophichthinae and COI barcoding sequences from the two type specimens of Ophichthus olivaceus sp. nov. and several sequences of the subfamily Myrophinae as the outgroup. Branches with 100 percent support from bootstrapped analyses (1.000 replicates) are depicted as broad lines, other bootstrap values higher than 50 percent are shown. Scale bar represents the average number of nucleotide nucleotide substitutions per site.
FIGURE 5 in A new species of turtle-headed sea Snake (Emydocephalus: Elapidae) endemic to Western Australia
FIGURE 5. Holotype of Emydocephalus orarius sp. nov. WAM R165708 dorsal (A); Ventral (B); cephalic scalation (C); and paratype WAM R73661 (D)
FIGURE 7 in A new species of turtle-headed sea Snake (Emydocephalus: Elapidae) endemic to Western Australia
FIGURE 7. Emydocephalus species in life: Emydocephalus orarius sp. nov from Pilbara (A) and Shark Bay (B) (DPIRD), Emydocephalus annulatus from the Timor Sea (C) (Jenna Crowe-Riddell) and Emydocephalus ijimae from Okinawa (D) (Vladimir Dinets).
Peaceful coexistence between people and deadly wildlife: why are recreational users of the ocean so rarely bitten by sea snakes?
<p>1) Research on interactions between humans and deadly snakes has focused on situations that result in high rates of snakebite; but we can also learn from cases where snakes and people coexist peacefully. For example, coastal bays near Noumea, in the Pacific archipelago of New Caledonia, are used by thousands of tourists and snakes, but bites are rare.</p> <p>2) Our long-term studies clarify reasons for this coexistence. Although 97% of snakes encountered in standardized snorkel surveys were a harmless species (<em>Emydocephalus annulatus</em>), we recorded dangerously venomous taxa often enough (one snake per eight hours snorkelling) that we would expect many risky human- snake interactions in these crowded bays. However, the risk is reduced by low overlap between humans and snakes in the timing of activity, both seasonally and on the diel cycle. Mate-searching male snakes, the group most likely to approach divers, enter the bays only in cooler months of the year when few beach users are present. Also, snakes tend to be active by night whereas people are not.</p> <p>3) Risk is further reduced by spatial divergence: bare-footed beach users stay in sandy areas rather than the adjacent coral-reef areas that are preferred by snakes. The response of snakes to disturbance is important also: most sea snakes are reluctant to bite even when harassed. Water currents frequently push sea snakes against hard objects, perhaps explaining why the snakes do not interpret brief contact with a human as an attack. The ability of snakes to flee is increased by uniformly high body temperature, and a complex three-dimensional aquatic environment.</p> <p>4) Thus, the danger of snakebite for recreational users of these popular beaches is reduced by aspects of human and snake behaviour that (i) decrease encounter rates, and (ii) render snakes unlikely to bite even if contacted. The risk to snakes is also reduced, because snakes are more difficult to detect and kill underwater than on land. As a result, thousands of snakes and people coexist harmoniously within these small bays.</p>
Data from: Rates of population differentiation and speciation are decoupled in sea snakes
Comparative phylogeography can inform many macroevolutionary questions, such as whether species diversification is limited by rates of geographic population differentiation. We examined the link between population genetic structure and species diversification in the fully aquatic sea snakes (Hydrophiinae) by comparing mitochondrial phylogeography in 16 species from two closely related clades that show contrasting diversification dynamics across northern Australia. Contrary to expectations from theory and several empirical studies, our results show that, at the geographic scale studied here, rates of population differentiation and speciation are not positively linked in sea snakes. The eight species sampled from the rapidly speciating Hydrophis clade have weak population differentiation that lacks geographic structure. In contrast, all eight sampled Aipysurus-Emydocephalus species show clear geographical patterns and many deep intraspecific splits, but have three-fold slower speciation rates. Alternative factors, such as ecological specialisation, species duration, and geographic range size, may underlie rapid speciation in sea snakes.
Data from: Temporal and spatial activity-associated energy partitioning in free-swimming sea snakes
1. Partitioning energy between critical basal functions and activity-associated behaviours is a primary determinant of animal survival. Consequently, habitat selection is likely to be driven by the efficiency with which resources can be acquired from a heterogeneous energy landscape. 2. Determining how energy partitioning is achieved across temporal and spatial scales is particularly challenging in aquatic animals due to the logistical limitations in estimating field metabolic rates (FMR) while simultaneously examining habitat choice. 3. Here, accelerometry telemetry and bimodal respirometry were used to correlate vectorial dynamic body acceleration (<i>VeDBA</i>) with oxygen consumption rates (<i>V̇o<sub>2</sub></i>) of sea snakes (<i>Hydrophis curtus</i> and <i>H. elegans</i>) across an ecologically-relevant temperature range. Subsequently, <i>VeDBA</i> of free-roaming snakes was used to estimate activity-associated FMR within a near-shore environment over diel, seasonal and spatial scales. 4. Diel changes in activity explained short-term patterns in FMR, whereas seasonal changes in water temperature drove long-term patterns. Spatial analyses demonstrated that activity-associated FMR was elevated in productive seagrass and mudflat habitats, indicative of a concentration of foraging efforts. 5. Our findings illustrate for the first time how sea snakes partition activity-associated FMR across time and space, providing an approach by which we can monitor the impacts of, and vulnerabilities to, natural and anthropogenic disturbances like warming and trawl fisheries.
Data from: Recent rapid speciation and ecomorph divergence in Indo-Australian sea snakes
The viviparous sea snakes (Hydrophiinae) are a young radiation of at least 62 species that display spectacular morphological diversity and high levels of local sympatry. To shed light on the mechanisms underlying sea snake diversification, we investigated recent speciation and eco-morphological differentiation in a clade of four nominal species with overlapping ranges in Southeast Asia and Australia. Analyses of morphology and stomach contents identified the presence of two distinct ecomorphs: a 'macrocephalic' ecomorph that reaches >2 m in length, has a large head and feeds on crevice-dwelling eels and gobies; and a 'microcephalic' ecomorph that rarely exceeds 1 m in length, has a small head and narrow fore-body and hunts snake eels in burrows. Mitochondrial sequences show a lack of reciprocal monophyly between ecomorphs and among putative species. However, individual assignment based on newly developed microsatellites separated co-distributed specimens into four significantly differentiated clusters corresponding to morphological species designations, indicating limited recent gene flow and progress towards speciation. A coalescent species tree (based on mitochondrial and nuclear sequences) and isolation-migration model (mitochondrial and microsatellite markers) suggest between one and three transitions between ecomorphs within the last approximately 1.2 million to approximately 840 000 years. In particular, the macrocephalic 'eastern' population of Hydrophis cyanocinctus and microcephalic H. melanocephalus appear to have diverged very recently and rapidly, resulting in major phenotypic differences and restriction of gene flow in sympatry. These results highlight the viviparous sea snakes as a promising system for speciation studies in the marine environment.
FIGURE 1 in Hydrophis donaldi (Elapidae, Hydrophiinae), a highly distinctive new species of sea snake from northern Australia
FIGURE 1. Hydrophis donaldi sp. nov. (A) Dorsal aspect of holotype QM J90700 (before preservation), (B) Ventral aspect of holotype QM J90700 (before preservation), (C) strongly spinous dorsal scales, (D) micro-CT scan of the lateral view of the head of SAMA R65216 (Scale = 10 mm). Note that the maxillary bone does not extend forward far beyond the palatine and that the fang is followed by a diastema. both character states were used by Smith (1926) to diagnose Hydrophis.
FIGURE 2 in Hydrophis donaldi (Elapidae, Hydrophiinae), a highly distinctive new species of sea snake from northern Australia
FIGURE 2. Hydrophis donaldi sp. nov., holotype QM J90700 (A) lateral aspect of head, (B) Dorsal aspect of head. Scale =10 mm.
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