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Accepted by Z. Nagy: 22 Jul. 2016; published: 29 Aug. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4158 (2): 203
–
212
http://www.mapress.com/j/zt/
Article
203
http://doi.org/10.11646/zootaxa.4158.2.3
http://zoobank.org/urn:lsid:zoobank.org:pub:4E5AB214-9CD0-4A60-8C16-E847433A9409
Recharacterization of Rhinophis dorsimaculatus Deraniyagala, 1941
(Serpentes: Uropeltidae), including description of new material
DAVID J. GOWER
1,3
& L. J. MENDIS WICKRAMASINGHE
2
1
Department of Life Sciences, The Natural History Museum, London SW7 5BD, UK
2
Herpetological Foundation of Sri Lanka, 31/5, Alwis Town, Hendala, Wattala, Sri Lanka
3
Corresponding author. E-mail: d.gower@nhm.ac.uk
Abstract
The Sri Lankan uropeltid (shieldtail) snake Rhinophis dorsimaculatus Deraniyagala, 1941 was described originally from
two specimens that were subsequently lost. The small amount of previously published data and lack of published colour
photographs made this one of South Asia’s most poorly known snake species, and this resulted in at least one instance of
taxonomic misidentification. An additional 10 specimens from a historical collection from the vicinity of the type locality
recently came to light. This material is reviewed and documented and the species recharacterized. An additional locality
for the species is reported. The newly reported material helps to corroborate the taxonomic validity and distinctiveness of
Rhinophis dorsimaculatus. The species is readily distinguished from congeners by having 227 or more ventral scales; a
large, dorsally carinate rostral shield; posterior margins of paired anals that are largely separated by the posteriormost ven-
tral scale; and a distinctive colour pattern with bilaterally asymmetrical dark blotches within a broad, pale middorsal stripe
and regularly punctate flanks.
Key words: Rhinophis porrectus, Rhinophis punctatus, shieldtail, Sri Lanka, systematics, taxonomy
Introduction
The uropeltid snake Rhinophis dorsimaculatus was described on the basis of two specimens from a single locality
in Sri Lanka, deposited in the Department of National Museums, Colombo, Sri Lanka (NMSL: Deraniyagala
1941). Those two specimens could not be located by us or several other people during multiple visits to the NMSL
collection since 2001 and are presumed lost or destroyed along with other older NMSL specimens (R. Pethiyagoda,
pers. comm. 2006). Gans (1966) reported that the two R. dorsimaculatus specimens were lost during evacuation,
citing “Director of National Museums, in litteris”. Deraniyagala (1955) presented a colour painting of the species,
confirming his previous description of a distinctive orange and black dorsum. However, the loss of the types, lack
of any additional material, and lack of colour photographs of living or preserved specimens has subsequently
caused at least one instance of taxonomic confusion in which a previously undescribed species (R. zigzag Gower &
Maduwage, 2011) was identified as R. dorsimaculatus (Somaweera 2006; Wickramasinghe et al. 2009). Uropeltid
taxonomy in general is in need of reassessment and revision (e.g., Gower et al. 2008).
Material from the late Carl Gans’ collection that has been deposited relatively recently in the California
Academy of Sciences, San Francisco (CAS) includes 10 specimens of R. dorsimaculatus. In addition, photographs
of live animals document a new locality for the species and further information on variation. The purpose of this
paper is to recharacterize the species and to clarify its taxonomic status.
Materials and methods
Relevant type and comparative material was examined in NMSL and the Natural History Museum, London, UK
(BMNH prefix). Total length was measured to the nearest 1 mm using a ruler or tape measure. Circumference was
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measured to the nearest 1 mm using a piece of thread and a ruler. All other measures were taken under stereo
dissecting microscopes with vernier calipers to 0.1 mm. Ventral scale counts include all midventral scales between
the mental and anal scales, following Gower & Ablett (2006). Dorsal scale-row reduction formulae are based on
Dowling (1951; see Appendix). Selected specimens were sexed by looking for oviducts and/or ova for females and
epididymis and/or testes for males, through an approximately midventral incision into the coelom. Uropeltid
taxonomy follows McDiarmid et al. (1999).
Rhinophis dorsimaculatus Deraniyagala, 1941
Rhinophis dorsimaculatus Deraniyagala, 1941: Deraniyagala (1941: 800–802, fig. 1, plate 1); Smith (1943: 88, 526); Taylor
(1950: 533); Deraniyagala (1955: 12, 15, plate 36); Gans (1966: 13–14); de Silva (1980: 113, 131, 183, 190, 191, 195);
Mahendra (1984: 63, 65); de Sliva (1990: 45, 69); Cadle et al. (1990); de Silva (1996: 70); de Silva (1998: 111);
McDiarmid et al. (1999: 136); Abeysiriwardana et al. (2000: 195); Bambaradeniya & Samarasekara (2001: 36); de Silva
(2001: 63); Bossuyt et al. (2004: fig. 2C); Somaweera (2006: 227, 234, 235) (in part); Wickramasinghe et al. (2009: 1, 6,
11) (in part); Gower & Maduwage (2011: 55, 59, 60, 63, 65); Pyron et al. (2013: 977, fig. 1); Wallach et al. (2014: 638).
Holotype. The “type” designated by Deraniyagala (1941). NMSL 86, collected 1938 or 1941 from “the coastal
village of Marichchukate [= Marichchikattuwa; Marichchukkaddi], North Western Province, Ceylon [= Sri
Lanka]”, now lost (Gans, 1966: 14). Approximate coordinates from maps: 8º 35’ N, 79º 57’ E, < 40 m elevation.
“Paratype”. NMSL unnumbered, collected 1938 or 1941 from the type locality. In his description of the
species, Deraniyagala (1941) mentioned a second specimen. However, he did not explicitly designate it as a type or
report a specimen number. There is no evidence that any of the data or figures reported for R. dorsimaculatus
pertain to this second specimen, which we thus do not consider a paratype. The second specimen, which is also lost
(Gans 1966: 14), was referred to as a paratype by Gans (1966: 13).
Referred material. Ten specimens, all from the type locality, newly referred here. CAS 225802, 225803 and
225804 (collected 27 November, 1974), 225842 and 225843 (29 April, 1976), 226077 and 226078 (9 February,
1977), 226662 (29 May, 1974), and 244583 and 244584 (4 December, 1974). These specimens were in the personal
collection of, and under tight proprietary control by, Carl Gans until they were donated to the CAS shortly before
his death (C.J. Bell pers. comm. 2016). The CAS material is referable to Rhinophis dorsimaculatus on account of it
being topotypic, and matching the holotype in its long and dorsally carinate rostral, large posteriormost ventral
projecting between the posterior margins of the anal scales, high number of ventrals (227+), and distinctive colour
pattern with a highly regularly punctate flanks and belly and approximately midvertebral, somewhat irregular black
blotches lying in a broad, pale, middorsal stripe.
Revised diagnosis. In having 227 or more ventral scales, R. dorsimaculatus differs from all congeners except
R. punctatus Müller, 1832 and R. porrectus Wall, 1921. Rhinophis dorsimaculatus differs from R. porrectus in
having fewer than 260 (227–250 among the only known material) versus more than 260 ventral scales (283 in
holotype of R. porrectus). Rhinophis dorsimaculatus differs from R. punctactus in its colour pattern. Although both
species have a broad pale dorsal stripe, in the former this bears dark dorsal blotches and in the latter a continuous
narrow dark vertebral line. Additionally, the colour of R. punctatus resembles that of R. porrectus in that the pale
dorsal stripe is bordered by a broad dark, dorsolateral stripe (broader than the dark lines on the lower flanks and
belly), whereas in R. dorsimaculatus the pale stripe is bordered by a narrow dark line that is undifferentiated from
the similar lines on the lower flanks and belly.
Rhinophis dorsimaculatus also differs clearly from the non-Rhinophis Sri Lankan uropeltids that, although
currently classified in other genera (Pseudotyphlops, Uropeltis, e.g., McDiarmid et al., 1999), likely form part of
the Sri Lankan uropeltid radiation together with Sri Lankan Rhinophis (Cadle et al. 1990; Bossuyt et al. 2004;
Pyron et al. 2013). Thus, R. dorsimaculatus differs (beyond in its distinctive colour pattern) from Pseudotyohlops
philippinus Müller, 1832, Uropeltis phillipsi (Nicholls, 1929) and U. melanogaster (Gray, 1858) in having far more
than 170 ventrals and a strongly dorsally carinate rostral. The same differences apply to U. ruhunae Deraniyagala,
1954, known from a single specimen that otherwise bears features found only among some Indian uropeltids.
Description of referred specimen CAS 225842. Generally good condition; split ventrolaterally and ventrally
at vent. Head small; snout pointed. Rostral large, protruding far anterior (1.8 mm) to mouth, pointed, trihedral
anteriorly, much longer than wide, dorsally strongly carinate and raised/arched (in lateral view); rostral widest at
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level of anterior superior corner of first supralabials. Rostral many times longer (in dorsal view) than short rostral-
frontal gap. Frontal subtriangular with convex anterior margin, distinctly longer than wide, little break in angle
between anterolateral (ocular) and posterolateral (parietal) margins. Frontal much shorter than, as wide as, rostral.
Paired nasals (and most of prefrontals) separated from each other by rostral. External naris small, subcircular,
slightly countersunk within small depression, located in anteroventral corner of nasal. Nasal contacts supralabials 1
and 2. Prefrontals briefly (for less than 25% of their length) in contact with each other along midline, briefly
separating frontal from rostral. Prefrontals taller than long, longer than frontal. Supralabials four, first smallest,
making the least contribution to margin of mouth; second larger but only slightly longer at mouth; fourth much the
largest. Ocular contacts supralabials 3 and 4. Eye small but distinct, diameter less than one third length of ocular,
located near anteroventral corner of ocular, bulging slightly from ocular surface, pupil subcircular. Paired parietals
longer than frontal, broadly rounded posteriorly (a little > 90°). Opposite parietals in brief midline contact, left
overlapping right. Parietals a little longer than wide, wider than frontal and rostral. Each parietal contacts four
scales other than head shields and supralabials. Three infralabials, first and third subequal in length, notably shorter
than second. First infralabials not in midline contact behind mental. First ventral longer than wide, second to fourth
approximately as long as wide, fifth and subsequent ventrals wider than long. First ventral not in contact with
mental.
Seven maxillary and seven or eight mandibular teeth on each side. Teeth simple, pointed, distinctly retrorse,
straight, evenly spaced. Anteriormost maxillary tooth aligned approximately with posterior end of lower margin of
second supralabial, posteriormost maxillary tooth close to posterior edge of lower margin of third supralabial;
mandibular row similar in length and alignment, with anteriormost member a little further forward than maxillary
row. Inside of mouth, including tongue, unpigmented.
FIGURE 1. Rhinophis dorsimaculatus. Topotype CAS 225842, whole specimen in two views. Scale bar in mm.
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FIGURE 2. Rhinophis dorsimaculatus. Topotype CAS 225842, head (upper) and tail (lower) ends in four views. Scales in mm.
Body subcylindrical to slightly dorsoventrally compressed. Head and body scales macroscopically smooth,
lacking keels. Body scales generally evenly sized on dorsum and along body. Midline ventral scales between
mental and anal of even size though anterior- and posteriormost ones gradually narrow. Posteriormost ventral much
larger, V-shaped, separating posterior margins of paired anals. Right anal possibly slightly overlaps left, but overlap
almost entirely absent. Ventrals 230, at midbody same width as exposed part of adjacent first dorsal scale row.
Posteriormost three or four ventrals slightly wider, 1.05 times as wide as adjacent first dorsal row; final ventral
much wider, V-shaped, 1.5 times as wide as first dorsal row. Dorsal scale rows 19 anteriorly, reducing to 17 along
most of body, until slightly anterior of vent. Scale row formula:
4+5 (11) 3+4 (224)
19 ------------------- 17 ------------------- 15
4+5 (11) 3+4 (225)
Dorsal scale rows approximately 13 at base of tail. Paired anal scales slightly larger than posteriormost dorsal
scales and subcaudals, slightly smaller than last ventral; only briefly in midline contact, substantially overlapped by
last ventral. Distal margin of each anal overlaps three other scales in addition to anteriormost subcaudals.
Subcaudals 7 (left), 8 (right), either all paired/divided with posteriormost on right much wider than others, or under
an alternative interpretation posteriormost subcaudals all single/undivided. Tail scales macroscopically smooth
though many with two or three inconspicuous, low keels on posterior portion of posteriormost subcaudals on lower
flanks. Caudal 'shield' bluntly conical, forming tip of tail, longer than wide in dorsal view, only slightly shorter than
shielded part of head, more visible from above than below, base (only a littler narrower than base of tail)
surrounded by last pair of subcaudals and 10 other scales. In posterior view shield regular broad oval, widest point
approximately mid height. Shield surface matt, covered with tiny tubercles in approximately radial distribution.
Narrow halo around base of shield glossy and without tubercles.
Left hemipenis everted, perhaps not fully. Moderately long (ca. 4.8 mm), slender, subcylindrical, slightly
tapering distally. Asulcate surface ornamented with short, slender, curved, proximally-directed, evenly spaced
spines extending to base of organ. Sulcate surface mostly lacking spines or other ornamentation, more
encroachment of spines towards sulcus margins distally than proximally. Sulcus spermaticus shallow,
inconspicuous, walls smooth.
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RHINOPHIS DORSIMACULA TUS
FIGURE 3. Map indicating position of type locality (Marichchikattuwa = Marichchukate) and newly reported locality
(Sannar) for Rhinophis dorsimaculatus in northwestern Sri Lanka.
FIGURE 4. Rhinophis dorsimaculatus. Two live, uncollected specimens from Sannar near Vidattalativu. Photographs by
L.J.M.W.
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RHINOPHIS DORSIMACULA TUS
In alcohol, background body colour pale tan with much darker (brown to blackish) markings. Broad, regular,
pale middorsal stripe/band across five scale rows extending from shortly behind head to tail shield. Pale middorsal
band marked with asymmetrical dark blotches, mostly discontinuous, more continuous anteriorly, darker
posteriorly, becoming blackish on tail. All other dorsal scale rows of body and all ventrals each with single dark dot
together forming series of regular, narrow, punctate lines along length of body. Dark spots on body scales confined
to scale bases, distal margins paler and translucent. Tail with more unmarked scales laterally and ventrolaterally
than on body. Anals coloured as posteriormost ventrals and dorsal scales of lower flanks. Tail shield matt, mostly
pale orange with broad, irregular, subterminal pale brown ring. Head brownish grey with small pale flecks, pale
borders to many shields. Rostral somewhat yellowish, paler anteriorly and ventrally. Posteromedial parts of
parietals pale. Edges of mouth (‘lips’) paler than dorsal and ventral surface of head.
Vari a t i o n. See Table 1 for meristic and morphometric details. CAS 225843 and 226077 (the latter with much
of inside of mouth blackish, this atypical condition possibly artefactual) in generally good condition; other CAS
specimens dehydrated or otherwise damaged. CAS 225804 incomplete, in several parts, and poorly preserved;
CAS 244583 eviscerated, largely skinned and missing anterior of head; CAS 244584 partly eviscerated and lacking
most of vertebral column, end of tail including shield missing. CAS 225842 and the other new specimens closely
match the lost holotype. The holotype was seemingly less attenuate than the best-preserved CAS specimens (Table
1). The number and disposition of head shields (including left over right overlap of parietals) is similar in all
specimens as far as can be ascertained. Mental notably more prominent in CAS 225843 and separated from
anteriormost chin shields (as also in CAS 226078) by midline contact between first pair of infralabials.
Ventrals 230–250; subcaudals 6 to 8 on each side. In some specimens (e.g., CAS 225843) it is difficult to
determine whether some of the posteriormost scales on the ventral surface of the tail are ‘true’ subcaudals or not.
Anals always paired, making little or no midline contact except slight right over left overlap in CAS 226662.
Dorsal scale rows always 19 anteriorly, 17 by level of 13th ventral and then without reductions until close to the
vent (see Appendix). Tail shield slightly variable in shape and size, for example it is more squarely-ended and
strongly overhangs the posteriormost subcaudal in CAS 225843 and, to a lesser extent, in 226077. The shield
tubercles are all small but vary from being low and worn (e.g., CAS 225842, 226662) to more prominent, pointed
and spine-like (e.g., CAS 225843).
Colour pattern generally constant and matching holotype (as figured by Deraniyagala 1941). Posteromedial
edges of parietals generally pale (as in holotype and CAS 225842), pigmented in CAS 225843. Tail shield always
somewhat orange, but brown marks more or less diffuse and variable in shape (e.g., U- rather than O-shaped in
CAS 226662). Pale dorsal band on body always present but variable. For example, it is three (rather than five)
scales wide on the anterior half of CAS 226662 and anterior two thirds to three quarters of CAS 225843, 226077,
and 244584. The narrow, punctate, darker lines on the other dorsal scale rows and ventrals are a constant feature.
Last ventral unpigmented in CAS 225843, 226077, 226662, and 244584.
Suggested ‘common’ names: Blotched Rhinophis, Marichchukate Rhinophis (English). The names ‘orange
shield tail’ (English) and ‘thambapanni walga ebaya’ (Sinhalese) were used by Wickramasinghe (2012: 112).
Distribution, natural history, and conservation: The lost types and only known existing museum specimens
of Rhinophis dorsimaculatus are all from a single locality, Marichchukate, on the northwestern coast of Sri Lanka
(Fig. 3). This is lowland and in the dry zone (sensu Legg & Jewell 1995), in contrast to most Sri Lankan uropeltids
that occur in the hills of the wet zone. We here report a second locality for R. dorsimaculatus. On 21 and 23 January
2014, eight live R. dorsimaculatus were seen at Sannar near Vidattalativu, Northern Province (8º 59’ 30.69” N, 80º
06’ 26.34” E, < 20 m elevation) while digging in loose soil and/or on the surface at night following monsoon
rainfall. This locality is also northwestern coast dry zone lowland, approximately 40 km north of the type locality
(Fig. 3). The habitat was typical dry-zone evergreen forest mixed with shady home garden vegetation. None of the
animals was collected but two were photographed (Fig. 4). Data were not recorded for the two photographed
animals but their colour pattern, head shields, and vertebral scale counts (235 and 238) closely match those of the
lost holotype and the CAS specimens of R. dorsimaculatus reported here. One of the photographed specimens (Fig.
4, left panel) has a more orange dorsal band (as reported for the type material by Deraniyagala 1941, 1955) and the
other (Fig. 4, right panel) a paler tan band. Both specimens differ slightly from the CAS specimens in that the dark
dorsal blotches are more continuous and less asymmetric, and the pale band has more scales with dark marks.
The type and the newly reported locality were, until recently, inaccessible during three decades of civil unrest
in Northern Province. After the war (itself cited as a cause of habitat degradation by Abeysiriwardana et al. 2000),
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the region has experienced rapid deforestation for human settlement. Much of the forest cover in the region of
Marichchukate has been cleared. In addition to expanding human settlements, forest has been cleared for
agriculture, which might also pose a conservation threat to R. dorsimaculatus (see also Abeysiriwardana et al.
2000). The second author and colleagues have also seen roadkill specimens of R. dorsimaculatus in the region.
Discussion
Although the holotype of Rhinophis dorsimaculatus is lost, there is no need for a neotype to be designated.
Deraniyagala’s (1941) description and illustrations are clear, there are reasonable locality data, topotypic material
matches Deraniyagala’s description, and the species is valid. In Deraniyagala’s (1955) colour plate of R.
dorsimaculatus (reproduced in Somaweera 2006: 235 plate B), the dorsal scale rows are too irregular, tail shield
too flat, and shield spines too large. In reality, the dorsal scales of R. dorsimaculatus are regular.
Hemipenis morphology of uropeltids has been little studied to date. The two species of Rhinophis for which
hemipenis descriptions are available (R. dorsimaculatus, R. lineatus) differ in the relative size and disposition of
spines. As is apparent from their figure, Gower & Maduwage (2011: fig. 5) were in error in describing the
hemipenis of R. lineatus as “deeply forked”; instead it is simple, as in R. dorsimaculatus.
Species of Rhinophis generally resemble other uropeltids in having a pair of large anal shields that are in broad,
overlapping contact and that are not substantially overlapped by the last ventral (e.g., Wickramasinghe et al. 2009;
Gower & Maduwage 2011). In this respect, R. dorsimaculatus is atypical—it has a pair of less enlarged anals that
are not, or only barely, in contact and which are substantially overlapped by the last ventral (the tip of which
overlaps the first subcaudals). We suspect that the anteriormost subcaudals were misidentified as the anals by
Deraniyagala (1941: fig. 1C), such that the type likely had seven rather than six subcaudals, the first of which are
paired. The condition of the anals in R. dorsimaculatus (relatively small and being overlapped substantially by the
posteriormost ventral) is similar to that in the BMNH material of R. porrectus and R. punctatus.
Rhinophis dorsimaculatus was described as less attenuate than R. punctatus and R. porrectus (Deraniyagala
1941) but our observations suggest the three species are similar in this respect. Total length divided by midbody
width is 46–56 in the four measured specimnes of R. dorsimaculatus housed at CAS (43 reported for the type
material: Deraniyagala 1941), 67 in the holotype of R. porrectus (BMNH 1920.8.25.1), and 48–65 in four R.
punctatus specimens (BMNH 71.11.13.1-2; 74.4.29.220-221). Reported differences among R. dorsimaculatus, R.
porrectus and R. punctatus in the ratio of the lengths frontal:rostral and frontal:parietal (Deraniyagala 1941) do not
hold up based on the material examined here for the three species. A photograph of R. punctatus in life
(Somaweera 2006: 247 plate A) shows a whitish/pinkish pale dorsal stripe and thin, dark continuous, punctate
vertebral line. However, from this photograph we count approximately 270 vertebral scales, and thus we identify it
instead as more probably R. porrectus. An “undescribed species” of Rhinophis (Das & de Silva 2005: 67) that has
the colour pattern of a R. porrectus or R. punctatus is here identified tentatively as R. punctatus on the basis of the
reported ventral count of 251.
Smith (1943: 526) suggested that R. dorsimaculatus was “closely related to R. punctatus” (under Smith’s
taxonomy, R. punctatus is a senior subjective synonym of R. porrectus). We agree with Smith’s interpretation,
based on the shared derived conditions of a high number of ventrals, greatly attenuate body, dorsally strongly
carinate rostral (also in some other Rhinophis), small frontal, large posteriormost ventral projecting between
posterior margins of anals, and pale, broad, dorsal midline band. Biochemical data did not resolve the relationships
of R. dorsimaculatus (Cadle et al. 1990) and the only DNA sequence-based phylogenies published to date (Bossuyt
et al. 2004; Pyron et al. 2013) did not sample R. punctatus or R. porrectus.
Acknowledgements
We thank Jens Vindum for his patient loaning of CAS specimens to London. For practical help with various aspects
of this study, we are grateful to Mohomed M. Bahir, Lalith Kariyawasam, Kalana Maduwage, Kelum
Manamendra-Arachchi, Sudath Nanayakkara, Rohan Pethiyagoda, Dinarzarde Raheem, and Ruchira Somaweera.
DJG thanks the Zoology Department of the Natural History Museum, London (NHM) for funding a visit to Sri
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Lanka that benefited this project. Harry Taylor (NHM) produced Figures 1 and 2, and Prasanna Samarawickrama
produced Figure 3. LJMW thanks Dilmah Conservation for funding project activities, and especially Asanka
Abayakoon, Department of Wildlife Conservation, Sri Lanka for permission granted, and Jaliya Dehideniya and
Ruwan Jayalath Dandeniya for their hospitality at Vidattalativu, Commando Training School. Christopher Bell and
Jennifer Olori provided constructive criticisms of an earlier draft.
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APPENDIX
Scale row formulae (based on Dowling, 1951) for five of the 10 CAS specimens of Rhinophis dorsimaculatus. CAS 225804 is
too incomplete to be included, and data were not recorded for other specimens. Possible scale row reduction in final 20-30
ventrals was not assessed for CAS 225802 or 225803. Upper and lower values correspond to right and left side, respectively.
4+5 (11) 3+4 (229)
CAS 225843: 19 ------------------- 17 ------------------- 15
4+5 (13) 3+4 (231)
4+5 (13) 3+4 (231)
CAS 226662: 19 ------------------- 17 ------------------- 15
4+5 (12) 3+4 (241)
4+5 (11) 3+4 (224)
CAS 225842: 19 ------------------- 17 ------------------- 15
4+5 (11) 3+4 (225)
3+4 (12)
CAS 225802: 19 ------------------- 17
3+4 (11)
5+6 (9)
CAS 225803: 19 ------------------- 17
5+6 (9)
