taxonID	type	description	language	source
610487BBFF9BFFE20326F9B6C9124A40.taxon	description	Figure 4 A – F Pattern description. Yellow-orange fluorescence. Entirety of shell covered by banded axial waves. Waves are irregular in shape, sometimes connecting to the adjacent wave and other times remaining separate. The waves on each whorl do not align with adjacent whorls, i. e., they are not continuous. Frequently the posterior of the whorl will show a dashed checkerboard-like pattern (arrowheads in Figure 4 A, D), rather than the continuation of the thicker main waves (Figure 4 E, F). There is also occasionally partial pigmentation in the spiral striae, separate from the main patterning. The inner lip fluoresces solidly.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF9BFFE20326F9B6C9124A40.taxon	discussion	Comment. The extinct M. murrayanus is morphologically very similar to extant Maoricolpus roseus. The original description of M. murrayanus by Tate (1893 a) differentiated it from M. roseus by only “ the spiral ornament of the posterior whorls develops into ribs on the anterior whorls ”. Garrard (1972) proposed there are strong grounds for considering M. murrayanus and M. roseus as synonymous due to the high morphological similarities. Whilst the protoconch was not preserved in the specimens in this study, the protoconch of both species has previously been described as multispiral, with three to four microscopic flatly globose whorls, apparently sinusigeroid (Garrard 1972). The main argument presented for not synonymising the species is the differing age and geographical ranges (Beu 2009). M. roseus is one of two extant species in the genus Maoricolpus, together with Maoricolpus finlayi. Both are natively endemic to New Zealand, but M. roseus is much more common and has a wider distribution (Powell 1940; Donald and Spencer, 2014). M. roseus is now a dominant species in New Zealand waters, often the most dominant benthic species present (McKnight, 1969). This is like what is observed in M. murrayanus in the Murbko Marl — it is the most common species present (Figure 3 D for abundance). M. roseus has been anthropogenically introduced to the east-coast of Australia, arriving in Tasmania in the 1920 s and now found as far north as Botany Bay (Gunasakera et al., 2005). It is now considered a pest in Australian waters (Bax et al., 2003). M. roseus was previously separated into two subspecies, M. roseus roseus and M. roseus manukauensis, based on slight morphological variances and differences in distribution. Donald and Spencer (2014) used molecular data to conclude these subspecies were not valid, and comprised a single, morphologically variable species. M. roseus has also been merged with a fossil species, the now invalid Maoricolpus proroseus, described from Tertiary rocks of the Gisborne District in the northeast of New Zealand (MolluscaBase eds., 2023). Thus, there is a precedent for a fossil species to be combined with M. roseus. The colour patterning revealed here strengthens the argument that M. murrayanus and M. roseus are a single species with slight morphological variations through time which are insufficiently different to justify separation. The patterns on each species show banded axial waves. Both species show a variability in the presence of a checkerboard-like pattern towards the outer end of the whorl, as described above. This seems to be more commonly present on M. murrayanus than M. roseus but is present on specimens such as the ones in Figure 4 H, I. Essentially all 387 specimens of Murraycolpus murrayanus studied fluoresced very strongly. Unlike most of the other taxa, no bleach treatment was applied to the photographed M. murrayanus specimens as trials found no benefit due to the already highly fluorescent condition. In situ observation of M. murrayanus underneath UV light also displayed clear fluorescence when viewed in weak shade (see Supplementary Figure 1). Several specimens were found to partially retain the original pigmentation. The most intact of these specimens (Figure 4 F) shows clearly defined pigmentation on the body whorl and first two spire whorls. The pigment is only present on the photographed side, while the side of the shell with the aperture lacks pigment, likely due to sun exposure. The other shells show a much fainter and smaller area of partial pigmentation. The fluorescence is not as strong in the partially pigmented specimens as it is in the non-pigmented specimens. The shells of modern relative M. roseus do not fluoresce under UV light.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE20188FA6DCC574931.taxon	description	Figure 5 F Pattern description. The tip of the apex strongly fluoresces yellow-orange, gradually fading in fluorescence away from the tip. The aperture fluoresces solidly. The upper half of the fasciolar band fluoresces strongly. The body whorl does not fluoresce but appears a darker, grey shade under UV light, though this colour difference did not translate well in photography.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE20183FB72CDFC4BA7.taxon	description	Figure 5 D Pattern description. Rounded patches located on the sculptured nodules of the shell fluoresce a pale orange. This continues throughout the whorls of the shell but is less pronounced closer to the apex.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE201A9FD0DCD264DB1.taxon	description	Figure 6 C Pattern description. Fluorescent spiral band positioned right after suture to middle whorl height. The spiral band consists of very closely related axial lines of variable thickness and brightness of the fluorescing areas. The axial lines are consistently thick near the edges of the spiral band, regardless of their thickness at the middle of the band.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE20301FBF7C9384B4B.taxon	description	Figure 5 A Pattern description. Yellow-orange fluorescence. Thin, wavy axial lines. Lines vary on specimens from being relatively straight to more curved. Lines do not frequently overlap with other lines; they are independent.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE20339FA41CBC04EC0.taxon	description	Figure 5 B Pattern description. Yellow or orange fluorescence. Most notable area of fluorescence is the aperture, which fluoresces solidly. The fluorescence is strongest on the lips but is present across the entire aperture and interior siphonal canal surface. This is also apparent at the varices where the old outer lips were located during shell growth. There is also faint fluorescence on the body of the shell, but this is not clear enough to discern any pattern.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE201B2FE96CC7C4C47.taxon	description	Figure 6 F Pattern description. Solid fluorescence on aperture, inner lip, and a band along the external side of the outer lip.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE201B2FE96CC7C4C47.taxon	discussion	Comment. In the photographed specimen the fluorescence is much more orange and intense on the outer lip than it is on the spire and inner lip. The fluorescence is also restricted to the section which is smooth and shiny.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF95FFE2018AFB98CDED4ADC.taxon	description	Figure 5 C Pattern description. Thin, yellow, fluorescent line running spirally through the centre of the whorls, concurrent with the apex of the sculptured peaks.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE001C9FCC1CBEE4AB5.taxon	description	Figure 5 G Pattern description. Irregular, wavy pattern aligned to axis. Waves are not consistent in size or shape. Waves often overlap and connect with other waves. Pattern continuous throughout entire shell (excluding protoconch) but is greater defined on the body whorl compared to the spire, where the whorls are small and comparatively more sculptured.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE601AAF996CAA44FE0.taxon	description	Figure 6 E Pattern description. Yellow-orange fluorescence. The ridged surface of the whorls shows fluorescence of lined chevrons, the apex of which point dextrally. Occasional fainter fluorescence in the depressed areas of the whorls showing vertical wavy lines continuing from the chevrons.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE601AAF996CAA44FE0.taxon	discussion	Comment. The patterns revealed here support the recent reassignment of L. murrayana from Lucerapex to Lophiotoma, where it had previously been placed. The move to Lucerapex was made by Powell (1963) who felt the original description of the species better matched Lucerapex, but did not study any specimens directly. Darragh (2024) returned L. murrayana back to Lophiotoma. Modern Lophiotoma frequently display patterns like what is observed here, whereas modern Lucerapex are described to typically be devoid of pattern and colour, with most species being pure white (Powell, 1966). Thus, we support the assignment to Lophiotoma proposed by Darragh (2024).	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE0030AFF41C9784AAD.taxon	description	Figure 6 A Pattern description. “ False ” triangle pattern (defined from the lack of pigment in a pigmented background) present on body whorl and first two whorls of spire. Pattern appearance the same as Nannamoria sp., aside from two medium-weight lines running spirally through the body whorl. These lines are interrupted by triangles.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE0030AFF41C9784AAD.taxon	discussion	Comment. Nannamoria cadella Hawke, 2021 in its original description was said to be comparable to Nannamoria trionyma Darragh, 1988 but with a smaller size, fewer shoulder nodules, which are also less developed, and with occurrence only in the Murbko Marl. No colour pattern for N. trionyma is known. Darragh (2024) synonymised the two species. We have continued to use N. cadella here for the following reasons: N. trionyma has not previously been recorded from the Murbko Marl, and each of the specimens studied follow the features given for N. cadella by Hawke (2021), with smaller shell sizes and fewer shoulder nodules (between 5 and 7). As the colour pattern for N. trionyma is currently unknown and we do not have access to previously identified N. trionyma fossils, it is our hope that the colour pattern revealed here can be used in the future for comparison with analysis of Victorian material of N. trionyma to help address this issue, and either justify of refute the validity of N. cadella.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE00195FB5CCBCB4BA0.taxon	description	Figure 6 D Pattern description. Bright yellow fluorescence. The edge of the body whorl has a thick, solid band of fluorescence. The inner lip also fluoresces strongly. Scattered fluorescence along the whorl sutures.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF97FFE001A5FDB6CC6F4D2B.taxon	description	Figure 5 E Pattern description. Orange fluorescence. Pattern of short, thick dashed lines. Start and end points of lines typically align with those above and below. Pattern covers entirety of shell but is greater defined on the body whorl than the more densely sculptured spire.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF91FFE60311FB81C95A4B51.taxon	description	Figure 7 A, B Pattern description. Solid, bright orange fluorescence on the entire external shell surface. The internal surface does not fluoresce aside from the muscle scars and pallial line, which appear yellow. Comment. Fluorescence intensity varies among specimens, with some shells showing only the faintest external fluorescence even after bleach treatment.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF91FFE60336FA18CA23487C.taxon	description	Figure 7 D Pattern description. The external surface shows weak, yellow fluorescence as concentric banding. The internal surface is non-fluorescent, excluding the muscle scars and pallial line.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF91FFE60150FEF6CD40492B.taxon	description	Figure 7 C Pattern description. Yellow fluorescence. 1 – 2 thin transverse bands, located near the centre and just above the anterior aperture of the shell. Energy-dispersive X-ray Spectroscopy The EDX results on Maoricolpus murrayanus (Table 1) include data from both fluorescing and non-fluorescing areas of the shell. Whilst in the EDX machine it was not possible to see the fluorescing area due to a lack of UV light source, and SEM images being rendered without colour. To minimise this issue, a fluorescent section marked prior to analysis was examined as closely as possible. Thus, the elemental spectrum results are a combination of both the fluorescing area and non-pigmented area and are not able to be separated outside of visual interpretations made from the elemental distribution maps (Figure 8). EDX of the fossil shell M. murrayanus identified O, C, and Ca, which constitute the calcium carbonate of the shell, with Si (apparent concentration 4.80, 2.54 Wt % – i. e., weight percentage –, 1.6 At %) and Al (apparent concentration 1.70, 100 Wt %, 0.6 At %) as the only other elements of notable amounts. Na, Mg, S, Cl, K, Fe, and W are all also present in miniscule amounts (see Table 1). Visual examination of the distribution map (Figure 8) shows a weak correlation between the density of Si, and to a lesser extent Al, and the area of remaining pigmentation / fluorescence. There also appears to be less carbon at this location than there is in the surrounding area. Other recorded elements show no apparent difference in density between the fluorescent and non-fluorescent areas. In comparison, the EDX of the modern shell, M. roseus, indicated that there were no levels of anything over 1 At % (atomic percentage) except for the expected O (51.4 At %), C (36.9 At %), and Ca (11.2 At %). Na, S, Si, Cl, Fe, Mg, and Sr were present but in miniscule amounts. There was no apparent visual correlation in the elemental maps between the pigmented area and any of the identified elements (Supplementary Figure 2).	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
610487BBFF91FFE60332F912CB1A4EE0.taxon	description	Figure 7 E Pattern description. External surface fluoresces yellow-orange in an irregular, mottled pattern. Fluorescent bands generally restricted to ribs – pattern does not usually end in the middle of a rib, rather it spans the width of the rib. Internal surface is non-fluorescent, excluding the muscle scars and pallial line.	en	Fergusen, Mahala A., Reed, Elizabeth H., García-Bellido, Diego C. (2025): Investigating colour in marine Miocene molluscs: UV fluorescence patterns and pigment EDX spectroscopy in shells from the Murbko Marl, Murray Basin (South Australia). Palaeontologia Electronica (a 19) 28 (1): 1-25, DOI: 10.26879/1394, URL: https://doi.org/10.26879/1394
