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Accepted by T. Nguyen: 7 Feb. 2019; published: 26 Apr. 2019
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2019 Magnolia Press
Zootaxa 4590 (2): 201
–
230
https://www.mapress.com/j/zt/
Article
201
https://doi.org/10.11646/zootaxa.4590.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:D86EA283-398B-47D4-90F0-751949D97DF0
A new species of Indo-Papuan groundsnake, genus Stegonotus Duméril et al.,
1854 (Serpentes, Colubridae), from the Bird’s Head Peninsula of West Papua,
Indonesia, with comments on differentiating morphological characters
CHRISTINE M. KAISER
1,2,5
, MARK O’SHEA
3
& HINRICH KAISER
2,4
1
Department of Animal Evolution and Systematics and Zoological Collection Marburg, Faculty of Biology, Philipps-Universität Mar-
burg, Karl-von-Frisch-Straße 8, 35032 Marburg, Germany
2
Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA
3
Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton, West Midlands WV1 1LY, United
Kingdom; and
West Midland Safari Park, Bewdley, Worcestershire DY12 1LF, United Kingdom
4
Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA
5
Corresponding author. E-mail: c_kaiser@rocketmail.com
Abstract
We describe a new species of Indo-Papuan groundsnake (Stegonotus) from a single adult male specimen collected in 1953
near Kamro, a village in Maybrat Regency, West Papua, Indonesia. The specimen had been considered a member of S.
batjanensis, a well-defined species from the northern Maluku Islands over 500 km to the northwest with which it shares
the key characteristic of having the 3
rd
, 4
th
, and 5
th
supralabial scales touching the eyes. The new species can be differen-
tiated from S. batjanensis as well as all other species of Stegonotus by having its 5
th
supralabial scale projecting forward
from behind the eye to form a narrow contact zone with the eye. In addition, it is differentiated by the combination of the
following characteristics: seven supralabials, the 3
rd
–5
th
touching the eye; eight infralabials, the 1
st
–4
th
touching the anterior
genial; four scales separating the posterior genial and the first gastrostege; dorsal scales in 17 rows, diminishing to 15 pos-
teriorly; a low number of ventrals (181 in the holotype) combined with a high number of subcaudals (105 in the holotype),
the latter comprising 37% of the scales on the ventral surface, the highest proportion in the genus. The description of this
species is of interest beyond adding to the species diversity of Stegonotus: it allowed us to explore additional characteris-
tics to resolve taxonomic questions in a morphologically conservative genus, it illustrates the need for additional herpeto-
logical survey work on the Bird’s Head Peninsula, and its initial misidentification serves as a reminder of the continued
relevance and importance of natural history collections as repositories for specimens and data that influence our knowl-
edge today by reaching out from the past.
Key words: groundsnake, Stegonotus, new species, Bird’s Head Peninsula, West Papua, Indonesia, New Guinea
Introduction
The Bird’s Head region of West Papua Province, Indonesian New Guinea (Fig. 1), is an area that potentially has a
high diversity of amphibians and reptiles (e.g., Günther 1999, 2003a,b, 2015; Philipp 1999; Richards & Iskandar
2000, 2001; Richards et al. 2000, 2002; Böhme & Jacobs 2001; Murphy 2012; O’Shea & Kaiser 2016). At the
same time, it is also an area where research is limited by significant logistical challenges, especially by a lack of a
transportation infrastructure (World Bank 2009). The specific area where the type specimen of our new species was
found is part of the sparsely populated Kais River drainage system, comprising six lake and river systems draining
into the Seram Sea (Fig. 1A; Boeseman 1963). A focal point and a claim to fame for the area is Lake Ayamaru, the
home of Melanotaenia boesemani, a stunningly iridescent rainbow fish (Allen & Cross 1980) that has become so
popular among aquaculturists that it is on the brink of extinction (Allen 1996; Nugraha et al. 2015). This region is
the territory of the Ayamaru people, and its habitats are characterized by a thin surface layer of soil exhibiting a
poor nutrient content (Elmberg 1968).
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FIGURE 1. Map of the Banda Sea (dark blue area in the center of the map, showing significant deep-water sea floor
topography), adjacent bodies of water, and surrounding landmasses in the eastern part of Wallacea. The yellow buttons indicate
type localities for species of the genus Stegonotus, with yellow lines encircling the estimated distributions of these taxa.
Included are S. sutteri on Sumba (1); S. florensis on Flores (2); S. lividus on Semau (3); S. modestus on Ambon (4); S.
batjanensis on Bacan (5); S. iridis on Batanta (6); S. derooijae on Salawati (7); S. keyensis in the Kei Islands (8); S. aruensis in
the Aru Islands (9); S. cucullatus near Manokwari, West Papua (10); Lycodon magnus, a synonym of S. cucullatus, on Supiori
(11); and S. parvus on Yapen (12). The insert (A) provides a close-up of the area in the south-central Bird’s Head Peninsula,
West Papua Province, Indonesian New Guinea, near the type locality of S. ayamaru sp. nov. Buttons indicate the type locality
(red), and two locations of note (white), including the town of Teminabuan (1), Lake Ayamaru (2), and Kamro Village (3).
Stegonotus Duméril et al., 1854 is a genus of groundsnakes that currently comprises 22 species and displays a
considerable amount of regional endemism (Ruane et al. 2017; Kaiser et al. 2018a). The distribution of these snakes
reaches from the Sunda Shelf in the west (Borneo) to central Melanesia in the east (Bismarck and Louisiade
Archipelagos
1
), and from the eastern-central Philippines in the north (Leyte and Samar) to Queensland, Australia
2
in
the south (Kaiser et al. 2018a). Work on the taxonomy and biogeography of these snakes was, until recently, very
limited (e.g., McDowell 1972; Ruane et al. 2017; Kaiser et al. 2018a) and only their ecological aspects have received
attention in Australia (e.g., Madsen & Shine 1994; Brown et al. 2005; Dubey et al. 2008, 2009; Trembath et al. 2009).
The genus Stegonotus was originally named to accommodate a species from the Philippines that did not fit
1. There exists only a single specimen of Stegonotus from a locality farther east (AMNH R-42402), which was ostensibly
collected on Bougainville in the Solomon Islands during the Whitney South Sea Expedition of the American Museum of
Natural History in the late 1920s or early 1930s (Burt & Burt 1932). We consider this record questionable (see Kaiser et al.
2018a), given that no Stegonotus specimens have been collected there since that time, despite the activities of other
collectors. The primary focus of the Whitney Expedition was ornithological, but a significant number of reptiles was also
collected. Since the expedition did reprovision on Samarai Island in southern New Guinea (Mayr 1943) and visited a
variety of other islands to the west of New Guinea (Burt & Burt 1932), we consider it possible that some specimens,
including this Stegonotus, may have been collected in one location but cataloged as having been found in another.
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NEW STEGONOTUS FROM WEST PAPUA
with the definition of Lycodon, at the time considered a cohesive genus. Thus, Duméril et al. (1854:680) decided to
place their new species into a new genus, based primarily on specific features regarding the dentition and the
overall body shape, with the type species S. muelleri. After a comprehensive review of the genus (Kaiser et al.
2018a) we recognized that there were many specimens of Stegonotus in museum collections that could not be
aligned with members of existing species. The specimen we describe as a new species herein is one example of
those unaligned forms and with its description, we continue our revisionary work on the genus.
Materials and methods
Museum specimens were generally measured according to the methodology of Kaiser et al. (2018a) and
photographed using the protocols recommended by Kaiser et al. (2018b). We found the use of photographs of
particular benefit and viewing and comparing them over and over again has allowed us to identify some new
characters that we consider relevant and informative for species definitions in the genus Stegonotus. We also
determined that counting scales and making certain measurements from photographs (comparative measurements,
such as to determine length ratios, not for absolute measurements) yielded more consistent and more trustworthy
results than counting scales or making measurements directly on a specimen. The use of photographs also resulted
in significant timesavings when it came to assessing time required when visiting a museum collection (Kaiser et al.
2018b). We present an abbreviated listing of measurements and scale characters in Table 1. We define these
characteristics in a manner that may not always be consistent with historical usage, given that relatively loose
definitions for some characteristics may have caused some misunderstandings in the past. The reader is referred
particularly to the footnote in Table 1 for the definitions of scales on the belly of snakes.
Basic measurements. We obtained snout–vent length (SVL) and tail length (TL) using a non-elastic string and
a measuring tape, with an accuracy we determined to be reliable at 50 mm and 1 mm, respectively (Natusch &
Shine 2012; Kaiser et al. 2018a). We combined these two values to calculate total length (TTL).
Head scale and suture measurements, ratios, and angles. We took a series of measurements on the head
scales of specimens, including some scale dimensions, suture lengths, and angles (Fig. 2) to obtain relative length
comparisons. Straight-line measurements were taken using dorsal head photographs (Kaiser et al. 2018b) and a
ruler, supported by the imaging software AnalyzingDigitalImages (Pickle & Gullage 2015), to compare relative
dimensions of scales and sutures (e.g., as ratios). For the purposes of these measurements, we considered sutures as
straight lines, even though there is sometimes minor waviness in the line created by abutting scales.
Suture lengths were defined by the exact endpoints where scale edges met, irrespective of whether these were
straight or angled with respect to the head’s midline. For example, the prefrontal suture (PfS) would have its
anterior endpoint where the anteriormost edges of the prefrontals contact each other. Even though prefrontals and
internasals are paired structures, their sutures do not necessarily line up in the midline of the head. Thus, the
posterior endpoint of the internasal suture (InS) is not necessarily the same point as the anterior endpoint of the
prefrontal suture (Fig. 2A).
We defined external skull length (ESL) as the length from the anterior endpoint of the prefrontal suture to the
posterior endpoint of the parietal suture in order to permit comparison of other features on the head against a
measure of overall skull length. We realize that this measurement does not approximate the extent of the skull’s
underlying bony elements, but we were seeking a means to obtain relative lengths for the snout, the eye, and
selected head scales when compared to the entire head, and this measurement provides a suitable stand-in. For the
purposes of these measurements, the parietal suture (ParS) is defined by an anterior endpoint at the posterior vertex
of the frontal and a posterior endpoint where the edges of the parietals diverge posteriorly (Fig. 2A). We measured
2. Three Stegonotus specimens (AMS R325–26, NTM R2337) have localities in New South Wales, Australia. In the
OZCAM database, specimen NTM R2337 is listed as having been collected at a locality called “Patonga,” with the GPS
coordinates placing the locality a little north of Sydney in an unlikely habitat for a tropical snake. However, there is a
locality by the same name in Kakadu National Park, Northern Territory, Australia (ca. 12.9001°S, 132.5780°E), where
Stegonotus are known to occur (Rick Shine, in litt.) and which we believe to be the most likely correct locality. Along
Australia’s eastern coast, snakes may experience waif dispersal due to the transport of tropical fruits on trucks from areas
where Stegonotus is common, and with their ability to climb (Greer 1997), these snakes could be moved large distances in
such a manner. A translocation to a location in northernmost New South Wales, as indicated for the two AMS specimens,
would therefore be possible. Thus, just as with the locality in Bougainville, we urge caution when it comes to the accuracy
of unexpected localities (see O’Shea & Kaiser 2018), which appear to be rather far to the south for a type of tropical snake.
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frontal length from the posterior endpoint of the prefrontal suture to the posterior vertex of the frontal, and frontal
width at the widest point of the frontal, irrespective of where on the scale the widest point occurred (Fig. 2A). The
resulting measurements were used to calculate ratios, including InS/PfS (snout scale ratio, SSR) and PfS/ESL
(head scale ratio, HSR) to provide estimates of relative snout lengths and relative head lengths (Table 1). In order to
be able to measure the length of the mental groove consistently and to allow comparisons with species whose
posterior genials diverge medially, we measured the mental grove from the posterior end of the mental scale along
the suture between the two first infralabial scales and along the medial edge of the anterior genial.
During our examination of hundreds of specimens (see Appendix), we came to realize that the angles of some
scale edges and sutures were of potential diagnostic interest, as measured using a digital angle finder (GemRed
82305, 0.3° accuracy; Gemred Sensor Technology Co., Guilin, China) on dorsal head photographs, which leads us
to define the following geometric characteristics (Fig. 2B), with the angle opening towards the snake’s snout
(forward-facing) or neck (rear-facing). The posterior frontal angle (PF ğ) is the forward-facing angle produced by
the sutures forming the posterior vertex of the frontal. Qualitatively, we considered this angle with the character
states of < 90° and ≥ 90°, although perhaps with a larger series of specimens (unavailable for most species of
Stegonotus) quantitative measurements may be useful as well. The anterior parietal angle (AP ğ) is the rear-facing
angle (measured on the parietal) formed by the suture between parietal and frontal, and the suture between
supraocular and the parietal. Qualitatively, we considered whether the lateral ray of the angle extends away from
the apex of the angle in a lateral (difference in angle from the line passing through the two lateral apices of the
frontal ≤ 10°) or posterolateral (> 10°) direction. Again, this angle could be quantified if large numbers of
specimens allow an assessment of intraspecific and interspecific variation. We also considered the position of the
eyes relative to the head scales and identified their position by drawing a transverse line across the head that
intercepts the anteriormost points of the eyes, and where this line lies in relation to the frontal (Fig. 2B). This line,
which we call the anterior eye line (AE)
3
, has three character states. When the eyes are positioned a little farther
toward the back of the head, the AE may cross the anterior portion of the frontal, a character state we call “behind”
because the AE lies behind the anterior edge of the frontal. The AE may also cross the head at the same level
(“same level”) as the anterior edge of the frontal or even slightly anterior to the frontal (“in front”), which indicate
a more forward eye position.
Scale characteristics. For our description of pholidosis, we used the standard nomenclature, including rostral,
nasal, internasal, prefrontal, frontal, loreal, preocular, supraocular, postocular, parietal, temporal, supralabial,
mental, infralabial, and genial
4
scales. We counted dorsal scale rows at three places, one head length posterior to the
head, at midbody, and one head length anterior to the cloaca, to obtain a count involving three numbers (e.g., 17-
17-15) listing the numbers of dorsal scales at these positions from front to back. The projection of the rostral onto
the dorsal surface of the snout caught our attention as a novel feature to assist in species differentiation. We defined
six character states (Fig. 3, Table 2) that appear to provide useful, partially diagnostic information. These include
(1) not visible, when the scale is not visible when viewed from above and only its tip may show; (2) gull wing,
when the scale is barely visible and has slightly curved edges that form a point medially; (3) gull wing +, when the
scale’s edges form the same curves as in (2) but more of the scale is visible; (4) shallow V, when the suture between
the rostral and the internasals forms a wide open V (an obtuse angle, >> 90°) with nearly straight rays that are not
curved into a gull-wing shape; (5) deep V, when the point of the rostral extends well onto the dorsal surface of the
head, forming an angle close to 90°; and (6)
U
-shaped, when the rostral does not form a point but a rounded shape
on the dorsal surface.
3. In geometric notation, a line is a specifically defined feature indicated by a line placed above the letters indicating
endpoints. In our terminology, we use the correct notation in our figures, but the typesetting software produces irregular,
unsightly line gaps when placing a line above letters. As a consequence, we adapted the notation and use an underline to
indicate this feature.
4. The terminology for scales on the chin of snakes has not been stable. Whereas the terms genial scales or chin shields have
been commonly used to refer to the paired scales following the mental, Ruane et al. (2017) recently reintroduced the term
inframaxillary for these scales, a term most commonly used in the 19
th
Century. We here use the technical term genial in
favor of chin shields because we believe the latter to be too generic a term; any scale on the chin could be considered a
chin shield. We avoid the use of inframaxillary because this relates an external, clearly visible feature to an internal,
invisible one (the maxilla, a bone), which we feel is not an optimal use of terminology in a kinetic skull where the bony,
internal features may shift. We refer to all other scales on the chin of snakes as “chin scales,” to differentiate them from the
genials.
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FIGURE 2. Characters used to assess the head scale pattern for species in the genus Stegonotus. (A) Positions of head scales,
sutures, and measurements, including internasals (IN) and the internasal suture (InS), prefrontals (PF) and the prefrontal suture
(PfS), the frontal (F) with its length (FL) and width (FW), the supraoculars (SO), and the parietals (P) with the parietal suture
(ParS). (B) Positions of the posterior frontal angle (PF ğ), the anterior parietal angle (AP ğ) and its lateral ray (yellow line),
and the anterior eye line (AE; white line). The underlying specimen is the holotype of Stegonotus ayamaru sp. nov.
(RMNH.RENA 31199).
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FIGURE 3. Six conditions of the rostral scale when viewed from above in species of Stegonotus. (A) Not visible. The body of
the scale is not visible, and only its tip may show. Pictured is the holotype of S. aruensis (MSNG 30186). (B) Gull wing. The
rostral scale is barely visible, and its suture with the internasals forms slightly curved edges that come to a medial point.
Pictured is the holotype of S. sutteri (NMBA 14872). (C) Gull wing +. The rostral scale is more visible than in (B). Pictured is
the holotype of S. ayamaru sp. nov. (RMNH.RENA 31199). (D) Shallow
V
. The suture between rostral and internasals forms
an obtuse angle (>> 90°) with straight or nearly straight rays (i.e., not curved into a gull-wing shape). Pictured is the holotype of
S. florensis (ZMA 11080). (E) Deep
V
. The point of the rostral comes up far onto the dorsal surface of the head and lies between
the nostrils. It forms an angle close to 90°. Pictured is a specimen of S. cucullatus (RMNH.RENA 47736). (F) U-shaped. The
rostral does not have a point on the dorsal surface of the head but the edge of the suture is rounded. Pictured is the holotype of
S. poechi (NMW 23406).
Coloration. Colors were estimated from digital images of snakes displayed on the uncalibrated screen of a
MacBook Pro using the guide of Köhler (2012). We realize that displayed colors may not necessarily be true, but
colors in preservatives are estimates at best regardless of calibration.
Sex. To determine the sex of specimens we used the presence of the m. retractor penis magnus and/or the
presence of the vas deferens for males when presence of the m. retractor penis magnus could not be determined
(such as in specimens with very dry tails). In females, we did not consider the absence of the m. retractor penis
magnus alone a diagnostic criterion and required the presence of oviducts, follicles, or eggs for positive
identification. Sex determination required dissection, but in a large majority of specimens suitable slits or body
cavity openings already existed. In order to locate the m. retractor penis magnus in undissected specimens, we used
a pointed scalpel (No. 11 blade) to make a 2-cm incision in the ventral midline of the tail, beginning after the first
three subcaudal scale rows. Gentle expansion of the slit using a blunt probe and forceps would reveal paired
muscular structures in males, whereas no such structures are present in females. Females do possess paired scent
glands in this area, however, which should not be confused with the paired hemipenial muscles. When dissection
along the body of specimens became necessary, we made an incision in the area between 30 and 40 ventral scales
anterior to the cloaca, which would allow us to locate relevant anatomical features.
Miscellanea. Museum abbreviations are used according to the list of Sabaj (2016). In a genus whose taxonomy
is still becoming understood, our inter-species comparisons are based on our own data collection, with the
exception of information for S. admiraltiensis, for which we obtained data from the text, illustrations, and appendix
in Ruane et al. (2017).
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TABLE 1. List of measurements and scale characters used to differentiate and describe species of Stegonotus.
Character
Abbreviation
Description
Snout–vent length
SVL
from the tip of the snout to the posterior edge of the cloacal scale
Tail length
TL
from the posterior edge of the cloaca to the tip of the tail
Total length
TTL
SVL + TL
Internasal suture, length
InS
the straight-line length of the suture between the two internasal scales
Prefrontal suture, length
PfS
the straight-line length of the suture between the two prefrontal scales
Snout scale ratio
SSR
InS / PfS
Head scale ratio
HSR
PfS / ESL
Frontal scale, length
FL
the length of the frontal scale measured along the midline of the head
Frontal scale, width
FW
the widest width of the frontal scale
Loreal scale ratio
LSR
height of the loreal scale/length of the loreal scale
Preocular scales, number
PR
number of scales immediately anterior to the eye
Postocular scales, number
PO
number of scales immediately posterior to the eye
Parietal suture, length
ParS
the straight-line length of the suture between the two parietal scales
Frontal-parietal ratio
FPR
length of the frontal scale as a percentage of ParS
External skull length
ESL
straight distance measurement along the midline of the head, from the anteriormost point of the prefrontal suture to the
posteriormost point of the parietal suture
Dorsal scales, number
D
enumerated one head length behind the head, at midbody, and one head length anterior to the cloaca
Ventral scales, number
V
gastrostegesa, counted between the neck and the cloaca, including the cloacal scale
Subcaudal scales, number
SC
scales on the ventral side of the tail, counted on one side including the tail tip
Subcaudal ratio
SCR
SC / (V + SC)
Supralabial scales, number
SL
enlarged scales of the upper lip, counted to the angle of the jaw and excluding rostral
Supralabials touching the eye
SLE
the numbered supralabials in contact with the eye
Infralabial scales, number
IL
enlarged scales of the lower lip, counted opposite the supralabial scale row and excluding mental
Infralabials touching anterior genial
ILG
the numbered infralabial scales making contact with the anterior genial
Genial scales
G
an anterior and a posterior pair of enlarged scales on the chin
Ventral head length
VHL
measured from the posterior end of the mental scale to the posterior edge of the first gastrostege
Posterior frontal angle
PF∠
forward-facing angle formed by the sutures at the posterior apex of the frontal scale
Anterior parietal angle
AP
∠
rear-facing angle formed by the suture between parietal and frontal scales, and the suture between supraocular and
parietal
aIn common usage, the term “ventrals” is used to describe the series of broad, transverse scales along a snake’s belly, which is used as an important character in
snake taxonomy. Kaiser et al. (2018a) noted that a “ventral” position would in the strict sense describe any scales along the ventral surface of a snake (including
ventral head scales, neck scales, belly scales, and scales under the tail). There are technical terms in use to allow differentiation of these scales, including the
terms “genial” (for the paired chin scales), “gastrostege” (for the broad ventral scales), “cloacal plate” (for the scale or scales covering the cloacal opening), and
“urostege” (commonly known as subcaudal scales). We here follow Kaiser et al. (2018a) in limiting the jargon in our presentation, unless it is necessary to
specifically differentiate scales. For example, there is a variable number of ventral scales in the midline of the neck region between the genials and the first
gastrosteges, and there is no technical term for these scales.
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Species description
Stegonotus ayamaru sp. nov.
Ayamaru Groundsnake
(Figs. 4, 5, 6A, 6A’, 7A, 7A’, Table 2)
Holotype. RMNH.RENA 31199, an adult male from “Komara” [Kamro], Aitinyo District, Maybrat Regency, West
Papua Province, Indonesia (1.5103°S, 132.3763°E; elevation ca. 140 m; Fig. 1). The type locality is a village of the
Ayamaru people inhabiting the central Bird’s Head Peninsula, situated in lowland primary rainforest. The specimen
was collected in February 1953 by the Reverend Herbert Marcus (1883–1961) and his wife Mieneke Marcus-van
den Nieuwenhuizen
5
(1911–2009). In addition to the village name, the original specimen label also gives the local
name for the specific collection site as “apan bebach,” but this appears to be a local designation and we were
unable to determine its precise locality. In order to determine the detailed locality of the village listed as “Komara,”
we located the name on Map SA 53-1 (Army Map Service 1942), the most likely map of the area available to
missionaries in the 1950s. On this map, there is a village listed under the name Komara
6
, which we confirmed as
the correct locality based on the accounts of the missionaries (Marcus-van den Nieuwenhuizen 2009). The map in
Elmberg (1968: Fig. 2) reflects this arrangement and indicates that Komara was a village with school activity,
making it a very likely residence for missionaries. This location is now known as Kamro. The initial identification
of the specimen as S. batjanensis was done by Dr. Maria S. (“Riet”) Knaap-van Meeuwen (born 1936) in March
1962.
Diagnosis. Stegonotus ayamaru appears to be a relatively small-sized (SVL of only known specimen = 493
mm) member of the genus with a relatively long tail (SCR
♂
= 0.37). It can be distinguished from all other known
Stegonotus by the following combination of characters: (1) rostral extending onto the dorsal part of the head and
not intruding into the internasal area, resulting in a relatively long internasal suture (character state: gull wing +;
Fig. 3C); (2) area of prefrontals 1.5 times that of the internasals, internasal suture two-thirds the length of the
prefrontal suture (Fig. 7A); (3) frontal clearly pentagonal with well-formed corners and a slightly convex anterior
suture; (4) AE lies behind the anterior end of frontal (Fig. 7A’); (5) length of frontal equal to that of the parietal
suture; (6) PF ğ ≥ 90° (Fig. 7A’); (7) AP ğ = 135° with lateral ray directed laterally (Fig. 7A’); (8) three temporal
scales touching the parietals, the lengths of the two anterior temporals combined equaling the length of the
posterior temporal, temporal formula 2+2+3; (9) three neck scales contacting parietals (10) loreal two-thirds as
long as wide; (11) a single preocular with dimensions similar to those of the loreal, curving around the anterior
border of the eye (Fig. 5B’); (12) seven supralabials, three (SL3–5) touching the eye; SL5 projecting forward from
behind the eye to form a narrow contact zone with the eye (Figs. 5B, B’, 6A’); (13) eight infralabials, four (IL1–4)
touching the anterior genial, two-thirds of IL4 in contact with the anterior genial (Fig. 5D’); (14) four chin scales
separating the posterior genial and the anteriormost gastrostege; (15) 17-17-15 dorsals; 181 ventrals, 105 paired
subcaudals; (16) cloacal plate entire; (17) color in preservative (65 years post-collection): dorsum Maroon (29)
fading laterally to Hazel (26), head Maroon with areas on the frontal and parietals that are a lighter color (Hazel);
nasals Light Buff (2) with some darker areas (Hazel); venter Light Buff (Figs. 4, 5). On the ventral surface, the
anterior to medial portions of the subcaudals are Hazel, creating a dark tail (Fig. 4C).
Comparisons. In the comparisons that follow, characteristics of S. ayamaru are listed in parentheses. Species
names are followed by the number of specimens used to determine ranges of continuous characters. Since the only
known specimen of S. ayamaru is a male, we limit our comparisons of characteristics with potential sexual
dimorphism (see Kaiser et al. 2018a) to male specimens; gender is indicated throughout by subscripted male (♂)
and female (♀) symbols for ease of reference. A listing of those characters most relevant for interspecies
comparisons is provided in Table 2.
5. The Reverend Herbert F. Marcus was a Dutch Mennonite (Doopsgezind in Dutch) missionary in the Vogelkop Peninsula of
Dutch New Guinea in the 1950s. His wife Mieneke Marcus-van den Nieuwenhuizen served as a government physician in
support of the missionary work.
6. There is an obvious difficulty with the identification of towns whose names are so similar in writing and pronunciation, a
problem compounded by local customs. It is, for example, quite possible that entire villages move, either to farm new
areas or to escape bouts of illness or death (or due to spiritual concerns), with the new village at a different location given
the same name as the old one. In this case, we have relied specifically on the collectors’ account (Marcus-van den
Nieuwenhuizen 2009) as it matches the contemporary map. This situation is compounded by the presence in southern New
Guinea (Papua Province, Indonesia) of the Kamoro people (Muller 2004).
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FIGURE 4. Holotype of Stegonotus ayamaru sp. nov. (RMNH.RENA 31199) in (A) dorsal and (B) ventral views. (C)
Enlargement of the subcaudal region to show the extent of dark patterning.
Stegonotus admiraltiensis (n = 1) differs from S. ayamaru by scale counts of V
♂
= 208 (181), SC
♂
= 98 (105),
SCR
♂
= 0.32 (0.37), D = 17-19-15 (17-17-15), SL = 8 (7), SL
E
= 4+5 (3+4+5), IL = 9 or 10 (8), and IL
G
= 6 (4).
Additional differences include: (1) the rostral of S. admiraltiensis is clearly visible on the dorsal surface of the
head, where it forms a shallow V (Fig. 3D) when viewed from above (gull wing +); (2) the rostral intrudes upon the
internasal space, shortening the internasal suture (does not extend into internasal space, internasal suture not
impacted); (3) InS = б PfS (InS = ⅔ PfS); (4) the lateral ray of AP ğ is directed posterolaterally (laterally); (5) FL
is slightly shorter than ParS (FL = ParS); (6) two temporals contact the parietals (3); and (7) two neck scales
contact the parietals (3). The closest known locality for S. admiraltiensis is on Manus Island, Papua New Guinea
(PNG), ca. 1600 km by air from the type locality of S. ayamaru.
Stegonotus aruensis (n = 1) differs by scale counts of V
♂
= 190 (181) and SL
E
= 3+4 (3+4+5). Additional
differences include (1) the rostral of S. aruensis is not visible on the dorsal surface of the head (Fig. 3A) (gull wing
+); (2) InS = ¾ PfS (InS = ⅔ PfS); (3) the lateral ray of AP ğ is directed posterolaterally (laterally); and (4) FL = е
ParS (FL = ParS). The closest known locality for S. aruensis is in the Aru Islands, Maluku Province, Indonesia, ca.
500 km by air from the type locality of S. ayamaru.
Stegonotus australis (n = 21) differs by scale counts of V
♂
= 191–220 (181), SC
♂
= 74–88 (105), SCR
♂
=
0.25–0.29 (0.37), SL
E
= 4+5 (3+4+5); SL = 8 or 9 (7); IL = 9 or 10 (8), and IL
G
= 5 (4). Additional differences
include (1) the rostral of S. australis is clearly visible on the dorsal surface of the head, where it forms a shallow V
when viewed from above (gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal
suture (does not extend into the internasal space, internasal suture not impacted); (3) InS = ⅓ PfS (InS = ⅔ PfS); (4)
the lateral ray of AP ğ is directed posterolaterally (laterally); (5) FL is slightly shorter than ParS (FL = ParS); and
(6) two temporals contact the parietals (3). The closest known locality for S. australis is in northern Queensland,
Australia, ca. 1500 km by air from the type locality of S. ayamaru.
Stegonotus batjanensis (n = 15) differs by scale counts of V
♂
= 221–236 (181), SC
♂
= 78–89 (105), SCR
♂
=
0.25–0.29 (0.37), SL = 8, or rarely 7 (7), IL = 9, rarely 8 or 10 (8), and IL
G
= 5 (4). In addition, (1) in S. batjanensis
SL3 projects posteriorly towards the eye to form a narrow contact zone (Fig. 6B, B’) (SL5 projects anteriorly
towards the eye; Fig. 6A, A’); (2) the rostral of S. batjanensis is clearly visible on the dorsal surface of the head,
where it forms a shallow V when viewed from above (gull wing +); (3) the rostral intrudes upon the internasal
space, shortening the internasal suture (does not extend into internasal space, internasal suture not impacted); (4)
InS = ¼ PfS (InS = ⅔ PfS); (5) AE lies in front of the anterior edge of the frontal (AE lies behind anterior edge of
the frontal); (6) FL is slightly shorter than ParS (FL = ParS); (7) one, rarely two, neck scales contact the parietals
(3); and (8) a distinctive color pattern is present (no distinctive color pattern). The closest known locality for S.
batjanensis is on Halmahera Island, North Maluku Province, Indonesia, ca. 500 km by air from the type locality of S.
ayamaru.
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FIGURE 5. Head of Stegonotus ayamaru sp. nov. (holotype, RMNH 31199) in both photographs (left column) and line
drawings (right column). Shown are dorsal (A, A’), right lateral (B, B’), left lateral (C, C’), and ventral (D, D’) views. Scale
abbreviations are as in Fig. 2, with the addition of rostral (R), nasals (N), loreals (L), preoculars (PR), postoculars (PO), anterior
temporals (AT), posterior temporals (PT), supralabials (SL), infralabials (IL), mental (M), anterior genials (AG), posterior
genials (PG), and ventrals (V).
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Stegonotus borneensis (n = 1) is a geographically distant species from Borneo, an island to the west of
Wallace’s Line, with scale counts differing in V
♂
= 196 (181), SC
♂
= 78 (105), SCR
♂
= 0.28 (0.37), SL
E
= 4+5
(3+4+5), SL = 9 (7), IL = 10 (8), and IL
G
= 5 (4). Additional differences include (1) InS = Э PfS (InS = ⅔ PfS); (2)
the lateral ray of AP ğ is directed posterolaterally (laterally); (3) FL = е ParS (FL = ParS); and (4) two temporals
contact the parietals (3). The closest known locality for S. borneensis is in northern Borneo, Sabah State, Malaysia,
ca. 1900 km by air from the type locality of S. ayamaru.
Stegonotus cucullatus (n = 3) differs by scale counts of V
♂
= 205–216 (181), SC
♂
= 84–95 (105), SCR
♂
=
0.29–0.31 (0.37), SL
E
= 4+5 (3+4+5), SL = 8 (7), IL = 9 or 10 (8), and IL
G
= 5 (4). Additional differences include
(1) the rostral of S. cucullatus is clearly visible on the dorsal surface of the head, where it forms a deep V (Fig. 3E)
when viewed from above (gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal
suture (does not extend into internasal space, internasal suture not impacted); (3) InS = ¼ PfS (InS = ⅔ PfS); (4)
AE lies at the same level as the anterior end of the frontal (AE lies behind the anterior edge of the frontal); (5) FL =
⅔ ParS (FL = ParS); and (6) two temporals are in contact with the parietals (3). The closest known locality for S.
cucullatus is in Manokwari, West Papua, Indonesia, ca. 200 km by air from the type locality of S. ayamaru.
Stegonotus derooijae (n = 1) differs by scale counts of SC
♂
= 94 (105), SL
E
= 3+4 (3+4+5), and IL
G
= 5 (4).
Additional differences include (1) AE lies at the same level as the anterior end of the frontal (AE lies behind the
anterior edge of the frontal); (2) PF ğ is < 90° (≥ 90); and (3) three to five neck scales contact the parietals (3). The
closest known locality for S. derooijae is on Salawati Island, Raja Ampat Archipelago, West Papua Province,
Indonesia, ca. 200 km by air from the type locality of S. ayamaru.
Stegonotus diehli (n = 1; juvenile, sex unknown) differs by scale counts of SC
?
= 82 (SC
♂
= 105), SCR
?
= 0.31
(SC
♂
= 0.37), D = 15-15-15 (17-17-15), and SL
E
3+4 (3+4+5). Additional differences include (1) the rostral of S.
diehli is barely visible on the dorsal surface of the head, where it forms a gull wing (Fig. 3B) when viewed from
above (gull wing +); (2) InS = ½ PfS (InS = ⅔ PfS); (3) PF ğ < 90° (≥ 90°); and (4) four neck scales contact the
parietals (3). The closest known locality for S. diehli is Bogadjim, Madang Province, PNG, ca. 1500 km by air from
the type locality of S. ayamaru.
Stegonotus dorsalis (n = 1) differs by scale counts of V
♂
= 208 (181), SC
♂
= 88 (105), SCR
♂
= 0.30 (0.37), D =
15-15-15 (17-17-15), SL
E
4+5 (3+4+5), SL = 8 (7), IL = 10 (8); and IL
G
= 5 (4). Additional differences include (1)
the rostral of S. dorsalis is clearly visible on the dorsal surface of the head, where it forms a deep V when viewed
from above (gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal suture (does
not extend into internasal space, internasal suture not impacted); (3) InS = ¼ PfS (InS = ⅔ PfS); (4) AE lies at the
same level as the anterior end of the frontal (AE lies behind the anterior edge of the frontal); (5) FL = е ParS (FL =
ParS); and (6) three neck scales contact the parietals, two of them enlarged (3, equally sized). The closest known
locality for S. dorsalis is in Madang Province, PNG, ca. 4.2280°S, 144.9333°E (see Kaiser et al. 2018a), ca. 1400
km by air from the type locality of S. ayamaru.
Stegonotus florensis (n = 1) differs by scale counts of V
♂
= 230 (181), SC
♂
= 82 unpaired (105, paired), SCR
♂
= 0.26 (0.37), D = 21-21-19 (17-17-15), SL = 8 or 9 (7), IL = 10 (8), and IL
G
= 5 (4). Additional differences include
(1) the rostral of S. florensis is clearly visible on the dorsal surface of the head, where it forms a shallow V when
viewed from above (gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal suture
(does not extend into internasal space, internasal suture not impacted); (3) InS = б PfS (InS = ⅔ PfS); (4) AE lies
at the same level as the anterior end of the frontal (AE lies behind anterior edge of the frontal); (5) the lateral ray of
AP ğ is directed posterolaterally (laterally); (6) FL is slightly shorter than ParS (FL = ParS); (7) PF ğ < 90° (≥
90°); and (8) a distinctive color pattern is present (no distinctive color pattern). The closest known locality for S.
florensis is on Flores Island, East Nusa Tenggara Province, Indonesia, ca. 1400 km by air from the type locality of
S. ayamaru.
Stegonotus guentheri (n = 29) differs by scale counts of V
♂
= 185–198 (181), SC
♂
= 64–80 (105), SCR
♂
=
0.25–0.29 (0.37), D = 15-15-13 (17-17-15), SL
E
4+5, or rarely 3+4 (3+4+5), SL = 8, or rarely 7 (7), IL = 9, or
rarely 8 or 10 (8), and IL
G
= 5 (4). Additional differences include (1) InS = б PfS (InS = ⅔ PfS); and (2) FL is
slightly shorter than ParS (FL = ParS). The closest known locality for S. guentheri is on Fergusson Island, Milne
Bay Province, PNG, ca. 2100 km by air from the type locality of S. ayamaru.
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FIGURE 6 (A–D). Right lateral views of four species of Stegonotus as photographs (upper images) and line drawings (lower
images), illustrating the position of supralabial scales (SL) and the eye. (A, A’) S. ayamaru sp. nov. (holotype, RMNH.RENA
31199), showing SL = 7 and a 3+4+5 pattern with SL5 projecting forward from a position behind the eye to form a narrow
contact zone with the eye. (B, B’) S. batjanensis (holotype, BMNH 1946.1.11.36), showing SL = 8 and a 3+4+5 pattern with
SL3 projection backwards from a position in front of the eye to form a narrow contact zone with the eye. (C, C’) S. florensis
(holotype, ZMA 11080), showing SL = 9 and a 3+4+5 pattern similar to S. batjanensis. In this specimen, SL5 displays a
developmental aberration on both sides of the head by being divided into two scales (as indicated by red stippling). (D, D’) S.
sutteri (holotype, NMBA 14872), showing SL = 9 and a 3+4+5 pattern similar to S. batjanensis and S. florensis.
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FIGURE 6 (E–G). Right lateral views of four species of Stegonotus as photographs (upper images) and line drawings (lower
images), illustrating the position of supralabial scales (SL) and the eye. (E, E’) S. cucullatus (RMNH.RENA 47736), showing
SL = 8 and a 4+5 pattern with the contact zone of SL5 only slightly greater than the one of SL4. (F, F’) S. keyensis (holotype,
MSNG 7521), showing SL = 9 and a 4+5 pattern with the contact zone of SL5 much greater than the one of SL4. (G, G’) S.
modestus (holotype, RMNH.RENA 324), showing SL = 7 and a 3+4 pattern with the contact zone of SL4 much greater than the
one of SL3. (H, H’) S. parvus (neotype, RMNH.RENA 46844), showing SL = 7 and a 3+4 pattern similar to that found in S.
modestus. In this specimen, the lower postocular scale appears to be fused to SL4.
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FIGURE 7 (A–D). Dorsal head views of four species of Stegonotus, shown to illustrate the position of measurements, sutures,
and angles as explained in Fig. 2. The placement of AE in A’–H’ documents the position of the eyes relative to the anterior
border of the frontal. The yellow line in A’–H’ shows the lateral ray of AP ğ. (A, A’) S. ayamaru sp. nov. (holotype,
RMNH.RENA 31199). (B, B’) S. batjanensis (holotype, BMNH 1946.1.11.36). (C, C’) S. florensis (holotype, ZMA 11080).
(D, D’) S. sutteri (holotype, NMBA 14872), showing a forward extension of the anterior border of the frontal.
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FIGURE 7 (E–G). Dorsal head views of four species of Stegonotus, shown to illustrate the position of measurements, sutures,
and angles as explained in Fig. 2. The placement of AE in A’–H’ documents the position of the eyes relative to the anterior
border of the frontal. The yellow line in A’–H’ shows the lateral ray of AP ğ. (E, E’) S. cucullatus (RMNH.RENA 47736). (F,
F’) S. keyensis (holotype, MSNG 7521). (G, G’) S. modestus (holotype, RMNH.RENA 324). (H, H’) S. parvus (neotype,
RMNH.RENA 46844).
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TABLE 2. Comparative data for species in the genus Stegonotus, including S. ayamaru sp. nov. Displayed characteristics include the number of specimens examined (n) and maximum lengths for
males and females. Asterisks (*) denote specimens with an incomplete tail. For missing data, we use the symbol when no specimens exist. Abbreviations include snout–vent length (SVL), tail length
(TL), total length (TTL), snout scale ratio (SSR), anterior eye line (AE), frontal-parietal ratio (FPR), number of supralabials contacting the eye (SLE), number of infralabials contacting the anterior genial
(ILG), and subcaudal ratio (SCR). Conditions that are relatively rare (occurring in < 20% of specimens) are identified by a superscripted dagger symbol (†). Where a lack of available specimens required
the use of unsexed material, this is indicated by the letter U. The character states of the rostral scale include: not visible when viewed from above, gull wing, gull wing +, V-shaped, or U-shaped (see
text). For definitions of the other listed characteristics, see Table 1.
Character
Sex
Taxon
ayamaru
sp. nov.
admiraltiensis
aruensis
australis
batjanensis
n
1♂
1♂
1♀ 1♂
10♀ 21♂
9♀ 15♂
Maximu m known length (mm)
SVL + TL = TTL
♀
527 + 208 = 735
815 + 232 = 1047
912 + 259 = 1171
♂
493 + 217 = 710
658 + 185 = 843
582 + 233 = 815
1330 + 275 = 1605
1490
+ 365
*
= 1845
Head characters
Rostral, condition
gull wing +
shallow V
not visible
shallow V
shallow V
SSR
2/3
2/5
3/4
1/3
1/4
Lateral ray of AP∠
, orientation
lateral
posterolateral
posterolateral
posterolateral
lateral
PF∠
98°
96°
91°
90°
94°
Position of
AE in relation to frontal
behind
behind
same level or behind
same level or behind
in front
FPR (%), mean
104
91
80
93
90
Temporals contacting parietal on one side
3
2
3
2
2 or 3
Neck scales contacting parietals
3
2
3
3
or 4
1 or 2†
Supralabials, number (SL
E)
7 (3+4+5)
8 (4+5)
7 or 8
(3+4)
8 or 9 (4+5)
8 (3+4+5)
Infralabials, number (IL
G)
8 (4)
9 or 10 (6)
8 or 9 (4)
9†
or 10 (5)
9† or 10 (5)
Body characters
Dorsal scale rows, numbers
17
-17-15
17-
19-15
17
-17-15
17
-17-15
17
-17-15
Gastrosteges, mean ± SD
(Range)
♀
191
200 ± 10.7 (178
–
215)
211 ± 4.5 (203
–216)
♂
181
208
190
211 ± 6.9 (191
–220)
229 ± 4.3 (221
–236)
Cloacal plate, condition
single
single
single
single
single
Subcaudals,
condition
paired
paired
paired
paired
paired
Subcaudals, mean ± SD
(Range)
♀
78
78 ± 5.7
(69–87)
84 ± 3.8 (80
–90)
♂
105
98
103
82 ± 4.5
(74–88)
85 ± 3.3 (78
–89)
SCR, mean ± SD
(Range)
♀
0.29
0.28 ± 0.01 (0.28
–
0.29)
0.29 ± 0.01 (0.27
–
0.30)
♂
0.37
0.32
0.35
0.28 ± 0.01 (0.25
–
0.29)
0.27 ± 0.01 (0.25
–
0.29)
……continued on the next page
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TABLE 2. (Continued)
Character
Sex
Taxon
borneensis
cucullatus
derooijae
diehli
dorsalis
florensis
n
1♀ 1♂
3♂
1♂ 3U
1U (juv)
1♂
1♀ 1♂
Maximum known
length (mm) SVL + TL = TTL
♀
715 + 221 = 936
196 + 56 =
252
447 + 109 =
556
♂
866 + 237 = 1103
769 + 254 = 1023
455 + 193 = 648
805 + 245 = 1050
850 + 195 = 1045
Head characters
Rostral, condition
gull wing +
deep V
gull wing +
gull
wing
deep V
shallow V
SSR
1/5
1/4
2/3
1/2
1/4
2/5
Lateral ray of AP∠
, orientation
posterolateral
lateral
lateral
lateral
lateral
posterolateral
PF∠
100°
104°
86°
82°
103°
66°
Position of
AE in relation to frontal
same level or behind
same level
same level
behind
same level
same level
FPR (%), mean
80
70
100
100
82
94
Temporals contacting parietal on one side
2
2
3
3
3
3
Neck scales contacting parietals
3
3
3
–5
4
3
2 or 5
Supralabials, number (SL
E)
9 (4+5)
8 (4+5)
7
(3+4)
7 (3+4)
8 (4+5)
9 (3+4+5)
Infralabials, number (IL
G)
10 (5)
10 (5)
8 (5)
8 (4)
10 (5)
10 (5)
Body characters
Dorsal scale rows, numbers
17
-17-15
17
-17-15
17
-17-15
15
-15-15
15
-15-15
21
-21-19
Gastrosteges, mean ± SD
(Range)
♀
193
179
217
♂
196
210 ± 5.6
(205
–216)
181
208
230
Cloacal plate, condition
single
single
single
single
single
single
Subcaudals, condition
paired
paired
paired
paired
paired
unpaired
Subcaudals, mean ± SD
(Range)
♀
78
82
65
♂
78
90 ±
5.6 (84–
95)
94
88
82
SCR, mean ± SD
(Range)
♀
0.29
0.31
0.23
♂
0.28
0.30 ± 0.01
(0.29–0.31)
0.34
0.30
0.26
……continued on the next page
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TABLE 2. (Continued)
Character
Sex
Taxon
guentheri
heterurus
iridis
keyensis
lividus
melanolabiatus
n
9♀ 29♂
3♀ 13♂
6♂
3♀ 4♂
2♀
5♀ 5♂
Maximu m known length (mm)
SVL + TL = TTL
♀
683 + 165 = 848
575 + 221 = 796
861 + 180 = 1041
489 + 118 = 607
652 + 222 = 874
♂
996 + 196*
= 1192
633 + 218 = 851
845 + 250 = 1095
1012 + 245*
= 1257
690 + 225*
= 915
Head characters
Rostral, condition
gull wing +
shallow V
deep V
gull wing
gull wing +
gull wing
SSR
2/5
2/5
–1/2
2/5
1/5
2/5
2/5
Lateral ray of AP∠
, orientation
lateral
posterolateral
lateral
lateral
lateral
posterolateral
PF∠
91°
86°
95°
101°
77°
93°
Position of
AE in relation to frontal
same level or behind
same level
same level
same level
same level
same level
FPR (%), mean
89
80
78
76
103
74
Temporals contacting parietal on one side
2
–4
2 or 3
2 or 3
2
2 or 3
2 or 3
Neck scales contacting parietals
3
2 or 3
3 or 4
3†
or 5
2 or 4
3 or 5
Supralabials, number (SL
E)
7†
or 8
(3+4
† or 4+5)
7 (3+4)
8 (4+5)
8†
or 9 (4+5)
7 (3+4)
7 (3+4)
Infralabials, number (IL
G)
9 or 10†
(5)
8 or 9†
(4)
10 (5)
9†
or 10 (4 or 5)
9 (4)
8 (5)
Body characters
Dorsal scale rows, numbers
15
-15-13
17
-17-15
17
-17-15 or 17-19
-15
17
-17-15
17
-17-15
17
-17-15
Gastrosteges, mean ± SD
(Range)
♀
187 ± 6.4 (174
–
198)
189 ± 5.1 (183
–
193)
193 ± 6.1 (188
–
200)
198
() (197–
199)
184 ± 4.3 (180
–
191)
♂
191 ± 3.5 (185
–
198)
188 ± 5.2 (179
–
198)
204 ± 4.4 (198–211)
204 ± 3.5 (200
–
207)
196 ± 4.7 (190
–
201)
Cloacal plate, condition
single
single
single
single
single
single
Subcaudals, condition
paired
unpaired
paired
paired
paired
paired
Subcaudals, mean ± SD
(Range)
♀
74 ± 4.9
(64–80)
89 ± 5.5
(83–93)
63 ± 3.5
(59–66)
68 (
) (67–68)
96 ± 4.9
(89–100)
♂
73 ± 4.3
(64–80)
82 ± 3.9
(76–88)
84 ± 3.7
(78–88)
74 ± 5.1
(70–80)
95 ± 3.3
(93–100)
SCR, mean ± SD
(Range)
♀
0.28 ± 0.02
(0.25
–
0.31)
0.32 ± 0.01
(0.31
–0.33)
0.25 ± 0.02
(0.23
–0.26)
0.25
0.34 ± 0.02
(0.32
–0.35)
♂
0.28 ± 0.01
(0.25
–0.29)
0.31 ± 0.01
(0.29
–0.32)
0.29 ± 0.01
(0.28
–0.30)
0.27 ± 0.01
(0.26
–0.28)
0.33 ± 0.01
(0.32
–0.34)
……continued on the next page
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NEW STEGONOTUS FROM WEST PAPUA
TABLE 2. (Continued)
Character
Sex
Taxon
modestus
muelleri
parvus
poechi
reticulatus
sutteri
n
4♀ 14♂
12♀ 18♂
1♀ 1♂
1♀
34♀ 82♂
1♀ 2♂
Maximu m known length (mm)
SVL + TL = TTL
♀
542 + 141 = 683
1242 + 323 = 1565
217 +
75 = 292
860 + 164*
= 1024
1077 + 254 = 1331
644 + 157 = 801
♂
915 + 277 = 1192
1545 + 450 = 1995
1413 + 325*
= 1738
567 + 144 = 711
Head characters
Rostral, condition
gull wing +
gull wing +
not visible
U
-shaped
gull wing +
gull
wing
SSR
2/5
2/5
2/5
1/2
2/5
1/2
Lateral ray of AP∠
, orientation
posterolateral
posterolateral
posterolateral
posterolateral
laterally
posterolateral
PF∠
96°
97°
81°
76°
98°
91°
Position of
AE in relation to frontal
same level
behind
behind
behind
same level
behind
FPR (%), mean
88
79
83
108
85
82
Temporals contacting parietal on one side
2 or 3
3
3
3 or 4
2
3
Neck scales contacting parietals
3
3
4
3
3 or 4
3
Supralabials, number (SL
E)
7 (3+4)
8 or 9†
(4+5)
7 (3+4)
9
(4+5+6)
8 or 9
(3+4
†
or 4+5)
9 (3+4+5)
Infralabials, number (IL
G)
8
or 9 (4)
9†
or 10 (4)
8 (4)
10 (4)
9 or 10 (4)
9 or 10 (4 or 5)
Body characters
Dorsal scale rows, numbers
17
-17-15
19
-17-15
17
-17-15
19
-19-17
17
-17-15
21
-21-19
Gastrosteges, mean ± SD
(Range)
♀
194 ± 3.5 (190
–198)
227 ± 4.7 (217
–234)
173
200
202 ± 8.2 (186
–215)
210
♂
202 ± 3.3 (195
–206)
235 ± 3.1 (226
–241)
177
208 ± 7.7 (188
–230)
220 (
) (210–
230)
Cloacal plate, condition
single
single
single
single
single
single
Subcaudals, condition
paired
paired
paired
paired
paired
unpaired
Subcaudals, mean ± SD
(Range)
♀
63 ± 4.4
(58–68)
97 ± 5.3
(86–103)
87
55*
82 ± 4.3
(73–88)
76
♂
84 ± 4.5
(78–94)
97 ± 7.0
(85–108)
100
83 ± 5.2
(71–92)
83
SCR, mean ± SD
(Range)
♀
0.24 ± 0.01
(0.23–
0.26)
0.29 ± 0.02
(0.26–
0.31)
0.33
0.29 ± 0.01
(0.27–
0.30)
0.27
♂
0.29 ± 0.01
(0.28–
0.31)
0.29 ± 0.02
(0.26–
0.31)
0.36
0.29 ± 0.01
(0.26–
0.33)
0.27
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Stegonotus heterurus (n = 13) differs by scale counts of V
♂
= 179–198 (181), SC
♂
= 76–88 unpaired (105
paired), SCR
♂
= 0.29–0.32 (0.37), and SL
E
= 3+4 (3+4+5). Additional differences include (1) the rostral of S.
heterurus is generally clearly visible on the dorsal surface of the head, where it forms a shallow V when viewed
from above (gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal suture (does
not extend into internasal space, internasal suture not impacted); (3) InS = б–½ PfS (InS = ⅔ PfS); (4) AE lies at
the same level as the anterior end of the frontal (AE lies behind anterior edge of the frontal); (5) the lateral ray of
AP ğis directed posterolaterally (laterally); (6) FL = е ParS (FL = ParS); and (7) PF ğ< 90° (≥ 90°). The closest
known locality for S. heterurus is Duke of York Island, East New Britain Province, PNG, ca. 2200 km by air from
the type locality of S. ayamaru.
Stegonotus iridis (n = 6) differs by scale counts of V
♂
= 198–211 (181), SC
♂
= 78–88 (105), SCR
♂
= 0.28–0.30
(0.37), SL
E
4+5 (3+4+5), SL = 8 (7), IL = 9 or 10 (8), and IL
G
= 5 (4). Additional differences include (1) the rostral
of S. iridis is clearly visible on the dorsal surface of the head, where it forms a deep V when viewed from above
(gull wing +); (2) the rostral intrudes upon the internasal space, shortening the internasal suture (does not extend
into internasal space, internasal suture not impacted); (3) InS = б PfS (InS = ⅔ PfS); (4) AE lies at the same level
as the anterior end of the frontal (AE lies behind anterior edge of the frontal); (5) FL = е ParS (FL = ParS); and (6)
a distinctive color pattern is present (no distinctive color pattern). The closest known locality for S. iridis is on
Salawati Island, Raja Ampat Archipelago, West Papua Province, Indonesia, ca. 200 km by air from the type
locality of S. ayamaru.
Stegonotus keyensis (n = 4) differs by scale counts of V
♂
= 200–207 (181), SC
♂
= 70–80 (105), SCR
♂
=
0.26–0.28 (0.37), SL
E
= 4+5 (3+4+5), SL = 8, or rarely 9 (7), IL = 8, 9, or rarely 10 (8), and IL
G
= 5 (4). Additional
differences include (1) the rostral of S. keyensis is barely visible on the dorsal surface of the head, where it forms a
gull wing when viewed from above (gull wing +); (2) InS = Э PfS (InS = ⅔ PfS); (3) AE lies at the same level as
the anterior end of the frontal (AE lies behind anterior edge of the frontal); (4) FL = ¾ ParS (FL = ParS); (5) two
temporals contact the parietals (3); and (6) five (rarely 3) neck scales contact the parietals (3). The closest known
locality for S. keyensis is in the Kei Islands, Maluku Province, Indonesia, ca. 400 km by air from the type locality
of S. ayamaru.
Stegonotus lividus (n = 2, female specimens) differs by scale counts of V
♀
= 197–199 (V
♂
= 181), SC
♀
= 67–68
(SC
♂
= 105), SCR
♀
= 0.25 (SCR
♂
= 0.37), and SL
E
= 3+4 (3+4+5). Additional differences include (1) InS = бPfS
(InS = ⅔ PfS); (2) AE lies at the same level as the anterior end of the frontal (AE lies behind anterior edge of the
frontal); (3) PFğ < 90° (≥ 90°); and (4) two or four neck scales contact the parietals (3). The closest known locality
for S. lividus is on Semau Island, East Nusa Tenggara Province, Indonesia, ca. 1350 km by air from the type
locality of S. ayamaru.
Stegonotus melanolabiatus (n = 5) differs by scale counts of V
♂
= 190–201 (181), SC
♂
= 93–100 (105), SCR
♂
= 0.32–0.34 (0.37), SL
E
= 3+4 (3+4+5), and IL
G
= 5 (4) Additional differences include (1) the rostral of S.
melanolabiatus is barely visible on the dorsal surface of the head, where it forms a gull wing when viewed from
above (gull wing +); (2) InS = б PfS (InS = ⅔ PfS); (3) AE lies at the same level as the anterior end of the frontal
(AE lies behind anterior edge of the frontal); (4) the lateral ray of AP ğ is directed posterolaterally (laterally); and
(5) FL = ¾ ParS (FL = ParS). The closest known locality for S. melanolabiatus is at Bobole, Hela Province, PNG
(Ruane et al. 2017), ca. 1300 km by air from the type locality of S. ayamaru.
Stegonotus modestus (n = 14) differs by scale counts of V
♂
= 195–206 (181), SC
♂
= 78–94 (105), SCR
♂
=
0.28–0.31 (0.37), and SL
E
= 3+4, rarely 4+5 (3+4+5). Additional differences include (1) InS = б PfS (InS = ⅔
PfS); (2) AE lies at the same level as the anterior end of the frontal (AE lies behind anterior edge of the frontal); (3)
the lateral ray of AP ğ is directed posterolaterally (laterally); (4) FL is slightly shorter than ParS (FL = ParS); and
(5) almost all known specimens of S. modestus have a distinctive, incomplete, light-colored band that marks the
end of the head (no neck band). The closest known locality for S. modestus is on Seram Island, Maluku Province,
Indonesia, ca. 250 km by air from the type locality of S. ayamaru.
Stegonotus muelleri (n = 18) is from the south-central Philippine Islands and differs by scale counts of V
♂
=
226–241 (181), D = 19-17-15 (17-17-15), SC
♂
= 85–108,
SCR
♂
= 0.26–0.31 (0.37), SL
E
= 4+5 (3+4+5), SL = 8, or
rarely 9 (7), and IL = 10, or rarely 9 (8). Additional differences include (1) InS = б PfS (InS = ⅔ PfS); (2) the
lateral ray of AP ğ is directed posterolaterally (laterally); and (3) FL = е ParS (FL = ParS). The closest known
locality for S. muelleri is on the Philippine island of Mindanao, ca. 1200 km by air from the type locality of S.
ayamaru.
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Stegonotus parvus (n = 1; numeric data are from the original species description
7
, specimen destroyed) differs
by scale counts of V
♂
= 177 (181), SC
♂
= 100 (105), and SL
E
= 3+4
(3+4+5). Additional differences include (1) the
rostral of S. parvus is not visible on the dorsal surface of the head (gull wing +); (2) InS = б PfS (InS = ⅔ PfS); (3)
the lateral ray of AP ğ is directed posterolaterally (laterally); (4) FL = е ParS (FL = ParS); (5) PF ğ < 90° (≥ 90°);
and (6) four neck scales contact the parietals (3). The closest known locality for S. parvus is on Yapen Island,
Papua Province, Indonesia, ca. 400 km by air from the type locality of S. ayamaru.
Stegonotus poechi (n = 1, a female specimen) differs by scale counts of V
♀
= 200 (181), D = 19-19-17 (17-17-
15), SL
E
= 4+5+6 (3+4+5), SL = 9 (7), and IL = 10 (8). Additional differences include (1) the rostral of S. poechi is
clearly visible on the dorsal surface of the head, where it forms a U (Fig. 3F) when viewed from above (gull wing
+); (2) the rostral intrudes upon the internasal space, somewhat shortening the internasal suture (does not extend
into internasal space, internasal suture not impacted); (3) InS = ½ PfS (InS = ⅔ PfS); (4) the lateral ray of AP ğ
directed posterolaterally (laterally); and (5) PF ğ < 90° (≥ 90). The closest known locality for S. poechi is in Madang
Province, PNG, ca. 4.2280°S, 144.9333°E (see Kaiser et al. 2018a), ca. 1400 km by air from the type locality of S.
ayamaru.
Stegonotus reticulatus (n = 82) differs in counts of V
♂
= 188–230 (181), SC
♂
= 71–92 (105), SCR
♂
= 0.26–0.33
(0.37), SL
E
= 4+5, rarely 3+4 (3+4+5), SL = 8 or 9, or rarely 7 (7), and IL = 9 or 10 (8). Additional differences
include (1) InS = б PfS (InS = ⅔ PfS); (2) AE lies at the same level as the anterior end of the frontal (AE lies
behind anterior edge of the frontal); (3) FL is slightly shorter than ParS (FL = ParS); (4) two temporals contact the
parietals (3); and (5) there is a very prominent color pattern in S. reticulatus, where each dorsal scale has dark
posterior edging to produce a reticulated pattern (no distinctive color pattern). The closest known locality for S.
reticulatus is along the Ok Ma Road in Western Province, PNG, ca. 1100 km by air from the type locality of S.
ayamaru. We have also examined a specimen (USNM 119549) from the Padaido Islands, Papua Province,
Indonesia (ca. 500 km by air from the type locality of S. ayamaru), whose color pattern approximates that of S.
reticulatus. With all other specimens of S. reticulatus occurring in localities south of New Guinea’s mountainous
spine or along its southern versant, we set this specimen aside for further examination as S. cf. reticulatus.
Stegonotus sutteri (n = 2) differs in counts of V
♂
= 210–230 (181), D = 21-21-19 (17-17-15), SC
♂
= 83
unpaired (105, paired), SCR
♂
= 0.27 (0.37), SL = 9 (7), and IL = 9 or 10 (8). Additional differences include (1) the
rostral of S. sutteri is barely visible on the dorsal surface of the head, where it forms a gull wing when viewed from
above (gull wing +); (2) InS = ½ PfS (InS = ⅔ PfS); (3) the lateral ray of AP ğ is directed posterolaterally (laterally);
and (4) FL = е ParS (FL = ParS). The closest known locality for S. sutteri is on Sumba Island, ca. 1600 km by air
from the type locality of S. ayamaru.
Description of the holotype – metrics (in mm) and pholidosis. An adult male; length 493 SVL + 217 TL =
710 TTL, TL 30.5% of TTL; head distinct from body; snout rounded in dorsal view, angled in lateral view, mouth
subterminal. Rostral barely extending onto the dorsal part of the head and not intruding into the internasal area
(character state: gull wing +; Figs. 3C, 5A, B). Internasals nearly as long as wide, with InS = ⅔ PfS. Prefrontals
slightly wider than long and about 1.5 times the area of the internasals; PfS = Э ESL. Frontal pentagonal with well-
formed corners and a slightly convex anterior suture; AE lies behind anterior edge of the frontal; FL = ParS (Fig.
5A); PFğ = 98°. Supraoculars relatively large and triangular, projecting beyond anterior and posterior extent of the
eye. Parietals angular, about twice as long as wide; APğ = 135°, with lateral ray directed laterally. Three temporals
touching the parietals, the lengths of the two anterior temporals combined equal the length of the posterior
temporal; temporal formula 2+2+3; three neck scales in contact with the parietals.
Nasal slightly larger than loreal, half as tall as wide, its widest point on the posterior border; naris one-third the
length and half the height of the nasal; loreal two-thirds as long as wide; a single preocular of similar dimensions as
the loreal, curving around the anterior border of the eye; two postoculars, the lower scale the size of the upper.
Seven supralabials; SL3–5 touching the eye, SL5 projecting forward from behind the eye to form a narrow contact
zone with the eye (Fig. 5B, B’); relative areas of supralabials 5 > 6 > 4 > 3 = 7 > 2 = 1, SL4 about half the size of
7. Based on the high value of SCR (0.36, the second-highest in the genus), Kaiser et al. (2018a) determined that the syntype
used for the numeric description by Meyer (1874) was a male. The neotype of S. parvus (RMNH 46844) proposed by
Kaiser et al. (2018a) is female, so it does not allow for direct numeric comparisons to the male holotype of S. ayamaru. We
used the neotype to determine characters other than those that might be sexually dimorphic.
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SL5 (Fig. 6A’). Eye length 75% of eye–naris distance; pupil round; internarial distance 62% of interorbital
distance. Eight infralabials; IL1–4 contacting anterior genials, with two-thirds of the medial border of IL4 in
contact with the anterior genial; four scales separate the posterior genial and the anteriormost gastrostege; length of
mental groove one-third of ventral head length (Fig. 5D’). Dorsals in 17-17-15 rows, 181 ventrals, cloacal plate
entire, 105 paired subcaudals, SCR
♂
= 0.37.
Description of the holotype – coloration in preservative (65 years after collection). Dorsum Maroon (29),
possibly with a more deeply colored middorsal area, color fading laterally to Hazel (26). Head Maroon with Hazel
areas on the frontal and parietals and extending over most of the rostral and all preoculars, postoculars, and
temporals (Fig. 5A). Much of the supralabials colored Hazel (Fig. 5B, C), with posterior edges on SL2, 3, and 5 of
a lighter color, Light Buff (2). Nasals Light Buff with their anterior third as well as a small area on the posterior
border of the nostril colored Hazel. Infralabials Hazel with posterior edges of IL2–6 and all of IL7 Light Buff (Fig.
5D). Venter colored Light Buff, well demarcated in color from the lateral areas. The ventral area of the tail is dark
because the anteromedial parts of the subcaudals are darkly colored in Hazel (Fig. 4C).
Natural history. Members of the genus Stegonotus are widely distributed geographically in the Indo-
Australian archipelago and occupy habitats ranging from swamps and lowland tropical forests (e.g., S. modestus) to
montane dipterocarp forests (e.g., S. borneensis in Sabah). They are able to adapt to disturbed areas, such as
gardens and coconut plantations (e.g., Karin et al. 2018: S. keyensis), and they may display arboreality in epiphytes
or even in honeycombed ant plants (Myrmecodia) (O’Shea 1996; Greer 1997). These snakes are known to consume
a variety of small vertebrates, particularly small lizards and reptile eggs (Brown et al. 2002; Trembath et al. 2009).
Given that there is only a single specimen, there is no specific information about this species. It was collected in
moist lowland rainforest.
Etymology. The species name ayamaru is a noun in apposition. It references the Ayamaru people of Maybrat
Regency, West Papua, Indonesian New Guinea and their homonymic language. We select this name not only to
indicate the type locality but also to highlight the Ayamaru people’s struggle to protect their forests and waterways
from exploitation. The Ayamaru Lakes are a case in point. One of the several endemic fishes in the lake
(Melanotaenia boesemani) has been over-collected and is currently listed as Endangered on the IUCN Red List
(Allen 1996). West Papua was recently declared a conservation province (Anonymous 2018), and it is hoped that
this will have a long-term, positive effect on regional development.
Discussion
The taxonomic history of the genus Stegonotus since its original description in 1854 has been rather convoluted,
and only recently have contemporary methods led to a better understanding of both diversity and species
boundaries in the genus (Ruane et al. 2017; Kaiser et al. 2018a). It is therefore no surprise that many species
assignments over the last 150 years were vague or incorrect: there was simply insufficient information on which to
base stable taxonomic decisions. As a case in point, even in his valiant attempt to assess Stegonotus populations in
Papua New Guinea, McDowell (1972) ended up with something he called “Stegonotus spec. cf. parvus,” for whose
taxonomic status the author was unable to corral a defining suite of data.
History of the Ayamaru specimen. Stegonotus ayamaru was initially identified as S. batjanensis in 1962 by
Riet Knaap-van Meeuwen, a postdoctoral researcher at the RMNH and a student of the noted Dutch herpetologist
Leo Daniël Brongersma (1907–1994). As curator and later museum director Brongersma and his students were
likely continuously in the process of finding names for those portions of the museum’s herpetological collection
that were continually expanding due to the Dutch presence in its former or contemporary colonies (the status of
Dutch New Guinea was only resolved, quite unsatisfactorily from the Dutch government’s point of view, in 1963;
Platje 2001; Webster 2013), and to which he himself added periodically (Boschma 1972). With Dr. Knaap-van
Meeuven, he published the description of a new elapid snake (Brongersma & Knaap-van Meeuwen 1964), now
known as Cryptophis boschmai. The misidentification as S. batjanensis, a long-established species, was easy to
make on account of the diagnostic pattern of supralabials 3+4+5 touching the eye and, perhaps, unfamiliarity with
the color pattern of S. batjanensis. Whereas simple scale counts (and consideration of the rather disparate
geographic origins) would have attested to a noteworthy find, the specimen appears to have been considered as
“one more sample” of S. batjanensis.
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Ecology. The type locality for S. ayamaru (1.5103° S, 132.3763° E) is present-day Kamro Village, one of 17
villages in the Aitinyo District of Maybrat Regency. This area is in lowland primary rainforest with a relatively low
human population density (21 per km
2
) and an average of only 300 inhabitants per village (calculated from
information presented by Brinkhoff 2018), with relatively high rainfall. There is to date very little in the way of
human development in the area and, absent the arrival of palm oil or extractive development, there is the potential
for ecotourism due to the presence of a high diversity of bird species, including at least 15 species of birds-of-
paradise (Paradisaeidae) (Frith et al. 2018).
How to define species boundaries in Stegonotus? There exists a general conundrum with determining
species boundaries in snakes using external morphological data. Whereas the set of traditional characteristics
related to scale counts and some measurements is readily and reliably obtained, these characteristics at the same
time pose a limitation when they are insufficient to define and identify species. This is the case in the genus
Stegonotus, which has so far defied attempts to produce a reliable character set diagnostic for species-level
taxonomy (e.g., McDowell 1972; Ruane et al. 2017). When comparing Stegonotus specimens from far-distant or
isolated localities, we realized that there could be considerable, confusing similarity in individuals, whose overall
gestalt led us to hypothesize that they should be different, but whose assembled character set did not support our
hypothesis. Thus emerged the need for a more detailed analysis. Given that there are relatively few external
characteristics that can be enumerated along the body of snakes (including length measurements as well as the
numbers of dorsals, ventrals, and subcaudals), we realized that the greatest potential data source was the head
scalation.
While looking at the overall shape of head scales in hundreds of specimens of Stegonotus, it became clear to us
that, sometimes, significant differences exist. However, we found it difficult to quantify these characters in a
simple way, while simultaneously allowing them to remain useful to field biologists. For example, there are several
easily recognizable angles formed by the shapes of scales, and we believe that two of these (PF ğ, AP ğ), as well
as the direction of the lateral ray of AP ğ, are taxonomically informative. PF ğ is an indicator of the overall shape
of the frontal. When this angle is ≥ 90° this translates to the frontal having a typical pentagonal shape with well-
defined angles. When the angle is < 90° then the frontal may still appear to be pentagonal, but its posterior portion
is thinner and more elongate overall, producing larger angles where the lateral edges of the frontal contact the
posteromedial border of the supraocular and the anteromedial border of the parietal (Fig. 2B). The AP ğ helps
quantify the shape of the area formed by the two sutures created by the posterior edges of the frontal, the anterior
edges of the parietal, and the posterior edges of the supraoculars. In conjunction with the size of the AP ğ, the lateral
ray of the AP ğ is directed either laterally or posterolaterally.
Another characteristic we found to be diagnostic is the influence of the rostral on the scale arrangement of the
dorsal surface in the snout region. The rostral scale is the anteriormost scale of a snake’s body, and as it emerges
from the front of the head onto the dorsum several potentially related traits can be identified: (1) The degree of the
rostral’s emergence onto the dorsal surface can be divided into several character states (Fig. 3, Table 2), as defined
in the Methods section above. (2) The specific extent of the rostral’s position on the dorsum can influence the
length of the internasal suture and, therefore, the SSR (ratio of internasal and prefrontal suture lengths). The highest
value of SSR among the species of Stegonotus we studied is ¾ (in S. aruensis), and we generally observed higher
SSR values in specimens whose rostral is less visible (character state: gull wing; e.g., S. diehli, S. sutteri). As the
rostral emerges more and more onto the dorsum (character states: gull wing +, shallow V, deep V, U-shaped,
respectively) then the SSR decreases from ½ to a minimum of Э. There are a few instances (S. florensis, all but one
specimen of S. keyensis) when the anterior edges of the prefrontals form an obtuse angle (>> 90°), which also
reduces the length of the internasal suture. It should be noted that the SSR is influenced not only by the emerging
rostral or a forward-pushing prefrontal edge, the length of the internasals along the midline (i.e., having “short”
internasals) also changes the SSR value. Such synergistic effects quantitatively impact the SSR values in S. diehli,
S. keyensis, and S. sutteri, whose “gull wing” rostral characteristic alone would lead to a prediction of greater SSR
values than actually observed. (3) Lastly, an issue related to the emergence of the rostral onto the dorsum is the
position of its point in relation to the nostrils. In some species, the rostral point lies anterior to the nostrils (e.g., Fig.
3A–D), but in others (Fig. 3E) the point lies at the same level of the nostrils.
Among the more traditional characteristics used when comparing snake species are the numbers of dorsal and
ventral scales (including gastrosteges and subcaudals) as well as supra- and infralabial counts. We find those useful
in combination, as they can provide a general sense of differentiation. For example, some species have high ventral
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and low subcaudal counts and others low ventral and high subcaudal counts, there is a definitive division by dorsals
into species with 15, 17, 19, or 21 middorsal scale rows, with the number of scales reducing posteriorly in some
species (e.g., 15-15-15 in S. diehli, but 15-15-13 in S. guentheri), and there are patterns among the supra- and
infralabial counts. It is unusual to have a species where specimens have more than one middorsal scale row
number, and this may be due to injury (where sometimes even numbers become possible) or the exact place along
the body where the count is made. While it is not too uncommon to see two or even three different supra- or
infralabial numbers within a species (and sometimes even on different sides of the same specimen), we have never
seen an intraspecific variation such that either two or three supralabials may touch the eye – it is invariably a case
that two supralabials touch the eye (3+4 or 4+5), except in those species where three touch the eye (3+4+5 or
4+5+6). It therefore appears as if some characteristics provide a stronger signal (e.g., three supralabials touching
the eye, number of middorsal scales) than others when it comes to delineating species.
Morphology, biogeography, and evolution. With the inclusion of some of the new characters we identified
and defined above, a more complete suite of morphological characters is now available to assist when identifying
species of Stegonotus. Considering some of the specifics of the geography and morphology of S. ayamaru,
identification is now readily possible by looking at certain diagnostic characters. For example, S. batjanensis has a
much higher ventral scale count with a much lower subcaudal count, resulting in a relatively short tail (low SCR
♂
).
Along with higher counts of SL, IL, and IL
G
, the condition of the rostral, a short internasal suture, and the color
pattern in life, distinguishing S. ayamaru from S. batjanensis now appears to be straightforward – but it has taken
some intensive and detailed study to reach this point, using data and/or specimens not available to Brongersma,
Knaap-van Meeuwen, or McDowell.
The same experience applies to other, geographically close species. For example, Stegonotus iridis has a much
higher ventral count and a much lower subcaudal count, resulting in a relatively short tail (low SCR
♂
). Along with
higher SL and IL
G
counts, SL
E
= 4+5, the condition of the rostral, and the striking color pattern in life S. iridis
appears much different from S. ayamaru. Even though S. derooijae is morphologically most similar to S. ayamaru,
it differs in three readily identified characteristics, including SL
E
= 3+4, a higher IL
G
count, and a PF∠ of < 90°.
Furthermore, among the nearly 1200 specimens we list as Stegonotus spp. in the Appendix, there are 32 specimens
from roughly the same area of Maybrat Regency (RMNH.RENA 46851, 46884–85, 46888–92, 46953, 47533,
47535–45, 47550–53, 47738, 47744–48, 47750) as where S. ayamaru occurs. Geographic proximity (the
occurrence of “topotypes”) could lead to the assumption that these all might be conspecific, with a single species of
Stegonotus found in this region. However, based on even a cursory examination of characters, we can easily
dismiss this notion: among the 32 specimens none have the characteristic SL
E
= 3+4+5 pattern of S. ayamaru, but
some have SL
E
= 3+4 and some SL
E
= 4+5, with concomitant SL and IL counts. Additional differences to S.
ayamaru include the morphology of other head scales as well as the counts of ventral and subcaudal scales. As a
consequence, we are confident that none of these specimens belongs to S. ayamaru, and we hypothesize that they
represent two unnamed species. This should come as no surprise given the presence of up to three species of
Stegonotus even on islands (e.g., those of the Raja Ampat Archipelago; Ruane et al. 2017). It appears that species
of Stegonotus with only limited gross morphological differences are able to divide resources in a habitat to support
the coexistence of multiple species in a limited area.
There is, at this point, no clear emerging pattern to explain either the biogeography or the evolutionary history
of species in the genus Stegonotus. While we have come to the realization that careful specimen work does allow us
to differentiate between different forms of Stegonotus and test taxonomic hypotheses, there is still much α-
taxonomy to be done before it is possible to assess broader patterns. However, the characters we have assembled at
least allow progress when it comes to a definition of species boundaries in Stegonotus and we intend to use them
for additional species assignments. This is critical in a genus for which hundreds of museum specimens are
unassigned to species or assigned incorrectly and no (or few) tissue samples would allow the use of molecular data
for making the correct determinations. We have also begun an analysis using geometric morphometrics (see
Zelditch & Swiderski 2018; Zelditch et al. 2012, 2013), but while this is a useful approach to solidify taxonomic
decisions, we strive to ensure that our species are also identifiable during fieldwork. As a consequence, we will
continue our examination of α-taxonomy using morphological data from existing museum specimens knowing that
we can now produce a more complete picture of the diversity in the genus Stegonotus.
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Acknowledgment
For specimen loans, for access to specimens under their care, and for additional details on the collection, we thank
the curators and collection managers at the AMS (Cecilie Bateson, Sandy Ingleby, Jodi Rowley, Ross Sadlier),
AMNH (David Dickey, David Kizirian, Christopher Raxworthy), ANWC (Margaret Cawsey, Leo Joseph, Alex
Drew, Robert Palmer, Eric Rittmeyer), BMNH (Patrick Campbell, David Gower, Colin McCarthy), BPBM (Allen
Allison, Nicholas Griffiths, Molly Hagemann, Kathleen Imada), CAS (Erica Ely, Lauren Scheinberg, Jens Vindum,
Laura Wilkinson), FMNH (Alan Resetar), IRSNB (Sebastien Bruaux, Olivier Pauwels), KU (Rafe Brown, Jack
Lapin, Luke Welton), MCZ (Joseph Martinez, José Rosado), MNHN (Ivan Ineich, Annemarie Ohler, Nicolas
Vidal), MNHP (Jiří Moravec), MSNG (Giuliano Doria, Maria Bruna Invernici, Massimo Petri), MWNH (Fritz
Geller-Grimm), MTKD (Markus Auer, Raffael Ernst), MZB (Irvan Sidik), NMBA (Denis Vallan, Urs Wüest),
NMV (Bentley Bird, Katie Date), NMW (Georg Gaßner, Heinz Grillitsch, Silke Schweiger), NRM (Bodil Kajrup,
Sven Kullander), PNGM (Bulisa Iova), QM (Andrew Amey, Geoff Thompson), RMNH/ZMA (Esther Dondorp),
SAMA (Carolyn Kovach, Steve Richards), SMF (Gunther Köhler, Linda Mogk), UCM (Emily Braker), UPNG
(Paulus Keip), UPS (Hans Mejlon), USNM (Kevin de Queiroz, Steve Gotte, Jeremy Jacobs, Roy McDiarmid, Ken
Tighe, Robert Wilson, George Zug), WAM (Paul Doughty, Brad Maryan), ZMB (Mark-Oliver Rödel, Frank
Tillack), and ZSM (Michael Franzen, Frank Glaw). Thanks to Jack Lapin, who sexed, examined, and photographed
the KU holdings of S. muelleri. We are very grateful to Sven Mecke (HLMD), Max Kieckbusch, and Lukas
Hartmann (both Philipps-Universität, Marburg) for expertly helping with measuring and counting specimens in
several museums. We extend our gratitude to Nicolas Vidal (MNHN) for sexing the holotypes of S. cucullatus and
S. muelleri, and to Patrick Campbell (BMNH) for sexing those of S. guentheri, S. heterurus, and S. reticulatus. We
especially thank Rick Shine, for his quick responses to emailed queries and for helping us resolve the status of
incongruous New South Wales Stegonotus localities, Glenn Shea, for various significant bits of advice regarding
Australian reptile taxonomy, and Sam Sweet for his constructive review of the manuscript. CMK thanks George
Zug (USNM) for explaining in word and deed how to properly sex snakes, and Lothar Beck (Philipps-Universität,
Marburg) for supporting her dissertation work and for providing some financial assistance for visits to Germany.
We also thank Isabel Kaiser for her guidance in adapting Köhler’s (2012) color guide for use with preserved
specimens. This paper is part of the first author’s dissertation at the Philipps-Universität, Marburg, Germany.
References
Allen, G. (1996) Melanotaenia boesemani. The IUCN Red List of Threatened Species, 1996, e.T13058A3408005. Available
from: http://www.iucnredlist.org/details/13058/0 (accessed 3 Jul. 2018)
Allen, G.R. & Cross, N.J. (1980) Description of five new rainbowfishes (Melanotaeniidae) from New Guinea. Records of the
Western Australian Museum, 8, 377–396.
Anonymous (2018) West Papua as Conservation Province. Conservation International Indonesia, Jakarta. Available from:
https://www.conservation.org/publications/Documents/CI_SLP-West-Papua-Conservation-Province.pdf (accessed 20 Oct.
2018)
Army Map Service (1942) Cape de Goede Hoop. New Guinea. Map SA 53-1. United States Army, Washington, D.C. [map]
Böhme, W. & Jacobs, H.J. (2001) Varanus macraei sp. n., eine neue Waranart der V. prasinus-Gruppe aus West Irian, Indonesia.
Herpetofauna, 23 (133), 5–10.
Boeseman, M. (1963) Notes on the fishes of western New Guinea. I. Zoologische Mededelingen, 38 (14), 221–242.
Boschma, H. (1972) Notes on the scientific career of Professor Dr. L.D. Brongersma, compiled at the occasion of his sixty-fifth
birthday. Zoologische Mededelingen, 47, VIII–XXII.
Brinkhoff, T. (2018) City Population. Aitinyo District. Accessible from: http://www.citypopulation.info/php/indonesia-
papuabarat-admin.php?adm2id=9110020 (accessed 20 Oct. 2018)
Brongersma, L.D. & Knaap-van Meeuwen, M.S. (1964) On a new species of Denisonia (Reptilia, Serpentes) from New
Guinea. Zoologische Mededelingen, 39, 550–554.
Brown, G.P., Shine, R. & Madsen, T. (2002) Responses of three sympatric snake species to tropical seasonality in northern
Australia. Journal of Tropical Ecology, 18, 549–568.
https://doi.org/10.1017/S0266467402002365
Brown, G.P., Shine, R. & Madsen, T. (2005) Spatial ecology of slatey-grey snakes (Stegonotus cucullatus, Colubridae) on a
KAISER ET AL.
226
·
Zootaxa 4590 (2) © 2019 Magnolia Press
tropical Australian floodplain. Journal of Tropical Ecology, 21, 605–612.
https://doi.org/10.1017/S0266467405002671
Burt, C.E. & Burt, M.D. (1932) Herpetological results of the Whitney South Sea Expedition. VI. Pacific island amphibians and
reptiles in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural
History, 63, 461–597.
Dubey, S., Brown, G.P., Madsen, T. & Shine, R. (2008) Male?biased dispersal in a tropical Australian snake
(Stegonotus cucullatus, Colubridae). Molecular Ecology, 17, 3506–3514.
https://doi.org/10.1111/j.1365-294X.2008.03859.x
Dubey, S., Brown, G.P., Madsen, T. & Shine, R. (2009) Sexual selection favours large body size in males of a tropical snake
(Stegonotus cucullatus, Colubridae). Animal Behaviour, 77, 177–182.
https://doi.org/10.1016/j.anbehav.2008.09.037
Duméril, A.M.C., Bibron, G. & Duméril, A.H.A. (1854) Erpétologie Générale ou Histoire Naturelle Complète des Reptiles.
Vol. 7. Partie 1. Roret, Paris, xvi + 780 pp.
Elmberg, J.-E. (1968) Aspects of Tradition and Change among the Mejprat of Irian Barat. Etnografiska Museet, Stockholm,
322 pp.
Frith, C., Frith, D. & Bonan, A. (2018) Birds-of-paradise (Paradisaeidae). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie,
D.A. & de Juana, E. (Eds.), Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. Available from: https://
www.hbw.com/node/52367 (accessed 31 Oct. 2018)
Greer, A.E. (1997) The Biology and Evolution of Australian Snakes. Surrey Beatty, Sydney, New South Wales, xii + 358 pp.
Günther, R. (1999) Morphological and bioacoustic characteristics of frogs of the genus Platymantis (Amphibia, Ranidae) in
Irian Jaya, with descriptions of two new species. Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische
Reihe, 75, 317–335.
https://doi.org/10.1002/mmnz.19990750210
Günther, R. (2003a) Three new species of the genus Oreophryne from western Papua, Indonesia (Amphibia, Anura,
Microhylidae). Spixiana, 26, 175–191.
Günther, R. (2003b) First record of the microhylid frog genus Cophixalus from western Papua, Indonesia, with descriptions of
two new species (Anura: Microhylidae). Herpetozoa, 16, 3–21.
Günther, R. (2015) Two new Oreophryne species from the Fakfak Mountains, West Papua Province of Indonesia (Anura,
Microhylidae). Vertebrate Zoology, 65, 357–370.
Kaiser, C.M., Kaiser, H. & O’Shea, M. (2018a) The taxonomic history of Indo-Papuan groundsnakes, genus Stegonotus
Duméril et al., 1854 (Colubridae), with some taxonomic revisions and the designation of a neotype for S. parvus (Meyer,
1874). Zootaxa, 4512 (1), 1–73.
http://dx.doi.org/10.11646/zootaxa.4512.1
Kaiser, C.M., Kaiser, H., Rickerl, K. & O’Shea, M. (2018b) A portable, low-cost approach for photographing fluid-preserved
snake specimens: recommendations with comments on optimizing specimen photography in natural history collections.
Herpetological Review, 49 (4), 666–667.
Karin, B.R., Stubbs, A.L., Arifin, U., Bloch, L.M., Ramadhan, G., Iskandar, D.T., Arida, E., Reilly, S.B., Kusnadi, A. &
McGuire, J.A. (2018) The herpetofauna of the Kei Islands (Maluku, Indonesia): comprehensive report on new and
historical collections, biogeographic patterns, conservation concerns, and an annotated checklist of species from Kei Kecil,
Kei Besar, Tam, and Kur. Raffles Bulletin of Zoology, 66, 704–738.
Köhler, G. (2012) Color Catalogue for Field Biologists. Herpeton, Offenbach, 49 pp.
Madsen, T. & Shine, R. (1994) Toxicity of a tropical Australian frog, Litoria dahlii, to sympatric snakes. Wildlife Research, 21,
21–25.
https://doi.org/10.1071/WR9940021
Marcus-van den Nieuwenhuizen, M. (2009) Autobiography of Rev. Herbert Marcus, Doopsgezind (Dutch Mennonite)
Missionary Based in the Bird's Head of Netherlands New Guinea (West Papua) in the 1950s. Available from: http://
papuaweb.org/dlib/bio/marcus/autobio.pdf (accessed 1 Oct. 2018)
Mayr, E. (1943) A journey to the Solomon Islands. Natural History, 52, 30–37 + 48.
Meyer, A.B. (1874) [Eine Mittheilung von Hrn. Dr. Adolf Meyer] über die von ihm auf Neu-Guinea und den Inseln Jobi,
Mysore und Mafoor im Jahre 1873 gesammelten Amphibien. Monatsberichte der Königlich Preussischen Akademie der
Wissenschaften zu Berlin, 1874, 128–140.
McDowell, S.B. (1972) The species of Stegonotus (Serpentes, Colubridae) in Papua New Guinea. Zoologische Mededelingen,
47 (2), 6–26.
Muller, K. (2004) The Biodiversity of New Guinea. Available from: http://papuaweb.org/dlib/up/muller-ngb/index.html
(accessed 1 Oct. 2018)
Murphy, J.C. (2012) Synonymised and forgotten, the Bird’s Head stout-tailed snakes, Calamophis Meyer (Squamata:
Serpentes: Homalopsidae). Raffles Bulletin of Zoology, 60 (2), 515–523.
Natusch, D.J.D. & Shine, R. (2012) Measuring body lengths of preserved snakes. Herpetological Review, 43, 34–35.
Nugraha, M.F.I, Pouyaud, L., Carman, O., Widyastuti, U., Zairin Junior, M., Kadarusman & Avarre, J.C. (2015) Genetic
diversity of Boeseman’s Rainbowfish (Melanotaenia boesemani) reared in Indonesian farms compared to endangered
natural populations. Tropical Conservation Science, 8 (3), 796–812.
Zootaxa 4590 (2) © 2019 Magnolia Press
·
227
NEW STEGONOTUS FROM WEST PAPUA
https://doi.org/10.1177/194008291500800315
O’Shea, M. (1996) A Guide to the Snakes of Papua New Guinea. Independent Publishing, Port Moresby, xii + 239 pp.
O’Shea, M. & Kaiser, H. (2016) The first female specimen of the poorly known Arfak Stout-tailed Snake, Calamophis
sharonbrooksae Murphy, 2012 (Serpentes: Colubroidea: Homalopsidae), from the Vogelkop Peninsula of Indonesian West
New Guinea, with comments on the taxonomic history of primitive homalopsids. Amphibian & Reptile Conservation, 10
(2), 1–10.
O’Shea, M. & Kaiser, H. (2018) Erroneous environs or aberrant activities? Reconciling unexpected collection localities for
three New Guinea Worm-eating Snakes (Toxicocalamus, Serpentes, Elapidae) using historical accounts. Herpetological
Review, 49, 189–207.
Philipp, K.M. (1999) Niche partitioning of Varanus doreanus, V. indicus and V. jobiensis in Irian Jaya: preliminary results.
Mertensiella, 11, 307–316.
Pickle, J. & Gullage, D. (2015) Manual for AnalyzingDigitalImages. Available from: https://www.umassk12.net/adi (accessed
27 Apr. 2018)
Platje, W. (2001) Dutch Sigint and the conflict with Indonesia 1950–62. Intelligence and National Security, 16, 285–312.
https://doi.org/10.1080/714002840
Richards, S. & Iskandar, D. (2000) A new minute Oreophryne (Anura: Microhylidae) from the mountains of Irian Jaya,
Indonesia. Raffles Bulletin of Zoology, 48, 257–262.
Richards, S. & Iskandar, D. (2001) A new species of tree frog (Anura, Hylidae, Litoria) from the mountains of Irian Jaya,
Indonesia. Alytes, 18, 141–152.
Richards, S., Iskandar, D.T. & Allison, A. (2000) Amphibians and reptiles of the Wapoga River area, Irian Jaya, Indonesia. In:
Mack, A.L. & Alonso, L.E. (Eds.), A Biological Assessment of the Waponga River Area of Northwestern Irian Jaya,
Indonesia. RAP Bulletin of Biological Assessment, 14, 54–57 + 113–120.
Richards, S., Iskandar, D.T., Tjaturadi, B. & Krishar, A. (2002) Amphibians and reptiles of the Yongsu area, Papua, Indonesia.
In: Richards, S.J. & Survadi, S. (Eds.), A Biodiversity Assessment of Yongsu—Cyclops Mountains and Southern
Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment, 25, 73–75 + 162–163.
Ruane, S., Richards, S.J., McVay, J.D., Tjaturadi, B., Krey, K. & Austin, C.A. (2017) Cryptic and non-cryptic diversity in New
Guinea ground snakes of the genus Stegonotus Duméril, Bibron and Duméril, 1854: a description of four new species
(Squamata: Colubridae). Journal of Natural History, 2017, 1–28.
Sabaj, M.H. (2016) Standard Symbolic Codes for Institutional Resource Collections in Herpetology and Ichthyology: an Online
Reference. Version 6.5 (16 August 2016). American Society of Ichthyologists and Herpetologists, Washington, D.C.
Electronically accessible. Available from: http://www.asih.org.
Trembath, D.F., Fearn, S. & Undheim, E.A.B. (2009) Natural history of the slatey grey snake (Stegonotus cucullatus)
(Serpentes: Colubridae) from tropical north Queensland, Australia. Australian Journal of Zoology, 57, 119–124.
https://doi.org/10.1071/ZO08091
Webster, D. (2013) Self-determination abandoned: the road to the New York Agreement on West New Guinea (Papua),
1960–62. Indonesia, 95, 9–24.
https://doi.org/10.1353/ind.2013.0006
World Bank (2009) Infrastructure Strategies for Papua and West Papua. World Bank Group, Washington, D.C., 35 pp.
Available from: http://siteresources.worldbank.org/INTINDONESIA/Resources/Publication/280016-1235115695188/
5847179-1263873728984/infrastructure.en.pdf (accessed 3 Jul. 2018)
Zelditch, M.L., Swiderski, D.L. & Sheets, H.D. (2012) Geometric Morphometrics for Biologists: a Primer. Academic Press,
London, 488 pp.
Zelditch, M.L., Swiderski, D.L. & Sheets, H.D. (2013) A Practical Companion to Geometric Morphometrics for Biologists:
Running Analyses in Freely-Available Software. Available from: http://booksite.elsevier.com/9780123869036/content/
Workbook.pdf (accessed 22 Oct. 2018)
Zelditch, M.L. & Swiderski, D.L. (2018) A Practical Companion to Geometric Morphometrics for Biologists: Running
Analyses in Freely-Available Software. 2
nd
Edition. Available from: https://www.researchgate.net/publication/
328215969_A_Practical_Companion_to_Geometric_Morphometrics_for_Biologists_Running_analyses_in_freely-
available_software_Second_Edition (accessed 22 Oct. 2018)
APPENDIX
Specimens Examined. With exact locality data (if known) for the specimens listed below available from online resources or
from collection staff, we limit the listing we present below to countries, provinces, and major political or geographic divisions.
Specimens are listed alphabetically by species, with those not yet assigned to species placed into the category Stegonotus spp.
Locality names include country names in capital letters, lesser administrative units (e.g., provinces, states, territories) in small
caps, and more specific localities (e.g., islands, regions) in regular font. Where locality data are missing, the phrase “no further
locality” is written in the style of the appropriate rank at which the data are missing. Thus, if the country is known and no
further locality, then the specimens are listed under a section with the font showing N
O
F
URTHER
L
OCALITY
. We did not
examine specimens of Stegonotus admiraltiensis under that name, although we did examine several specimens from the
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Admiralty Archipelago, which are listed in the section that includes Stegonotus sp. with locality Manus Province.
Stegonotus aruensis (n = 2).—INDONESIA, M
ALUKU
P
ROVINCE
, Aru Islands: MSNG 30186, SMF 18055.
Stegonotus australis (n = 37).—AUSTRALIA, Q
UEENSLAND
: AMNH R-69262, 69264, 82227, AMS 2160, 31894 (holotype of
Lycodon darnleyensis), 31895 (holotype of Herbertophis plumbeus), 43911–12, 44232, 46004, 55594, 55683, 57126–27,
58999, 62429, 64224, 74310, 74678, 82585, 82586, 84299, 86779, 94364–65, 99712, 104829, BMNH 1946.1.14.93
(holoty pe), FMNH 97832–35, MCZ 128118, 128185–86, 137639–40, MSNG 7513, SMF 65223.
Stegonotus ayamaru (n = 1).—INDONESIA, West Papua Province, Kamoro: RMNH.RENA 31199.
Stegonotus batjanensis (n = 29).—INDONESIA, M
ALUKU
P
ROVINCE
, Ambon: ZMA 17660A–B. N
ORTH
M
ALUKU
P
ROVINCE
,
Bacan: BMNH 1946.1.11.36 (holotype). Halmahera: AMS 140778, MCZ 33510, MSNG 54816, 30187-1–3, NMW 19538,
SMF 18066–67, 18069–70, USNM 216000–01, 237129, ZMB 12052, 16128. Morotai: RMNH.RENA 327A–B, USNM
159972, ZMA 17658. Ternate: MSNG 34687, 7512-1–4.
Stegonotus borneensis (n = 5).—MALAYSIA, S
ABAH
: FMNH 251054, USNM 130244, ZMB 47866. S
ARAWAK
: FMNH
164746, 188500.
Stegonotus cucullatus (n = 3).—INDONESIA, W
EST
P
APUA
P
ROVINCE
, Manokwari: MNHN 3412 (holotype), RMNH.RENA
47736–37.
Stegonotus derooijae (n = 5).—INDONESIA, W
EST
P
APUA
P
ROVINCE
, Batanta: SAMA R70467 (paratype). Salawati: MZB
Ophi.3288 (holotype), Ophi.3293 (paratype). Waigeo: MZB Ophi.3292, SAMA [SJR 7651].
Stegonotus diehli (n = 1).—PAPUA NEW GUINEA, M
ADANG
P
ROVINCE
: MWNH 1244 (holotype).
Stegonotus dorsalis (n = 1).—PAPUA NEW GUINEA, M
ADANG
P
ROVINCE
: NMW 14861 (holotype).
Stegonotus florensis (n = 2).—INDONESIA, E
AST
N
USA
T
ENGGARA
P
ROVINCE
, Flores: WAM 104606, ZMA 11080
(holotype).
Stegonotus guentheri (n = 38).—PAPUA NEW GUINEA, M
ILNE
B
AY
P
ROVINCE
, Fergusson: AMS 137977, BMNH
1946.1.11.37 (paralectotype), 1946.1.11.38 (lectotype), 1946.1.11.39 (paralectotype), BPBM 16529–31, 17292. Kuia:
AMS 86835. Normanby: AMS 129734, 129743, BPBM 16532–33, 17289–91, 20821, 39689–96. No further locality:
AMNH R-42374–75, 42377–81, 42389, 73964, 76702, 76704, 76712, 76715.
Stegonotus heterurus (n = 19).—PAPUA NEW GUINEA, E
AST
N
EW
B
RITAIN
P
ROVINCE
, Duke of York: BMNH 1946.1.14.95
(lectotype), 1946.1.15.10 (paralectotype). No further locality: AMNH R-107181, BMNH 1946.1.14.91 (paralectotype),
MCZ 140716, SAMA R60253, ZMB 15191. New Britain Island, no further locality: AMS 58749–50, NMW 22627.1–4,
SMF 18064. N
EW
I
RELAND
P
ROVINCE
: AMS 28683, BPBM 12017–18, 12045, USNM 340182, ZMB 24023. N
O
F
URTHER
L
OCALITY
: SAMA R64702, R64824, R64825.
Stegonotus iridis (n = 5).—INDONESIA, W
EST
P
APUA
P
ROVINCE
, Batanta: MZB Ophi.3303 (paratype), Ophi.3306 (holotype).
Salawati: MZB Ophi.3301, Ophi.3302 (paratype). Waigeo: MZB Ophi.3305 (paratype).
Stegonotus keyensis (n = 7).—INDONESIA, M
ALUKU
P
ROVINCE
, Kei Islands: AMS 64782, 147649, MSNG 7521 (holotype),
NMW 22593, SMF 18056–57, ZMA 17667.
Stegonotus lividus (n = 2).—INDONESIA, E
AST
N
USA
T
ENGGARA
P
ROVINCE
, Semau: RMNH.RENA 325A (lectotype), 325B
(paralectotype).
Stegonotus melanolabiatus (n = 10).—PAPUA NEW GUINEA, H
ELA
P
ROVINCE
: AMS 115360, 122359. S
IMBU
P
ROVINCE
:
AMS 115320 (paratype), 115321, 115342, 115343 (holotype), 115361 (paratype). S
OUTHERN
H
IGHLANDS
P
ROVINCE
:
AMS 122360, 122906 (paratype), 122907.
Stegonotus modestus (n = 18).—INDONESIA, M
ALUKU
P
ROVINCE
, Ambon: BMNH 1946.1.13.74 (holotype of Ablabes
greineri), FMNH 134324, 166902, 169758, 169761–62, RMNH.RENA 324 (holotype), SMF 18054. Buru: RMNH.RENA
5136, 43599, ZMA 17666A–C, ZMB 9368. Seram: BMNH 1998.328–29, 1946.1.11.40 (holotype of Coluber holochrous),
RMNH.RENA 4066 (holotype of Coronella rosenbergii).
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229
NEW STEGONOTUS FROM WEST PAPUA
Stegonotus muelleri (n = 34).—PHILIPPINES, D
INAGAT
I
SLANDS
P
ROVINCE
, Dinagat: KU 310178. Leyte: KU 311286,
328843, 344097. Mindanao: KU 320001–04, 326661–64, 334773–75, SMF 75167. Samar: CAS 13233, KU 310853–54,
310869, 328844, 338848, 338850–53, 338855, 344534–38, ZMB 4294 (holotype of Spilotes samarensis). N
O
FURTHER
LOCALITY
: MNHN 848 (holotype of Stegonotus muelleri), ZSM 97/1978.
Stegonotus parvus (n = 2).—INDONESIA, P
APUA
P
ROVINCE
, Yapen: MTKD 876A (holotype, destroyed), RMNH.RENA
46844 (neotype).
Stegonotus poechi (n = 1).—PAPUA NEW GUINEA, M
ADANG
P
ROVINCE
: NMW 23406 (holotype).
Stegonotus reticulatus (n = 145).—PAPUA NEW GUINEA, C
ENTRAL
P
ROVINCE
: AMNH R-59065, 82524, 103673, 115061,
117758, AMS 14541, BMNH 1935.5.10.160, BPBM 18937–38, 19480, 19483–84, 37707–08, MCZ 59055, 145948,
146061, MSNG 52310, NMW 22592.1–4, PNGM 22139, 22810, 24496, 24502–05, 24511, UPNG 1700, 7102. S
IMBU
P
ROVINCE
: AMNH R-72783–86, 98865, 98867–69, 98871, AMS 115351, 115359, 115362–64, BPBM 5026, MCZ
121566. E
AST
S
EPIK
P
ROVINCE
: AMNH R-75358. E
ASTERN
H
IGHLANDS
P
ROVINCE
: AMNH R-85717–19, 134134. G
ULF
P
ROVINCE
: AMS 13302, BMNH 1936.7.7.31, MCZ 59052, UPNG 6076, ZSM 107–17/2000, 119–21/2000. M
ADANG
P
ROVINCE
: PNGM 24495, UPNG 971. M
ILNE
B
AY
P
ROVINCE
: AMNH R-73957, 73967, 76718, AMS 2431, 2615,
124933, 124976, 129733, 129744, BMNH 1946.1.14.87 (lectotype), 1946.1.14.88 (paralectotype), BPBM 15158, 15159,
16528, 17278–80, 39687, MCZ 149741, MSNG 38732, 52312, 52309-1–3, NMW 22591.2, 22591.4, ZMB 14194.
N
ATIONAL
C
APITAL
D
ISTRICT
: BMNH 1987.915, BPBM 39686, PNGM 23032. N
ORTHERN
P
ROVINCE
: AMNH R-142876,
BPBM 39688, 39697, MCZ 141692. S
OUTHERN
H
IGHLANDS
P
ROVINCE
: AMS 122354–58, 122903, BPBM 28280, 34888,
ZSM 118/2000. W
ESTERN
P
ROVINCE
: AMNH R-57504, 111797–98, 111800, MSNG 54479-1, 54479-3, UPNG 1605,
UPNG 8593. W
ESTERN
H
IGHLANDS
P
ROVINCE
: AMNH R-82525, PNGM 22434. Southeast New Guinea, no further
locality: SMF 18115. “Southern New Guinea,” no further locality: MSNG 38733, 52311. “New Guinea,” no further
locality: AMS 39, A5770–73, “modestus” without specimen number, BMNH 97.12.10.120, PNGM 2213, 24748, 24810,
24821, UPNG 1208, 2568, 8848.
Stegonotus cf. reticulatus (n = 1).—INDONESIA, P
APUA
P
ROVINCE
, Padaido: USNM 119549.
Stegonotus sutteri (n = 3).—INDONESIA, E
AST
N
USA
T
ENGGARA
P
ROVINCE
, Sumba: NMBA 14872 (holotype), WAM
101597, 101680.
Stegonotus spp. (n = 1181).—AUSTRALIA, N
EW
S
OUTH
W
ALES
: NTM 2337. N
ORTHERN
T
ERRITORY
: AMS 14042, 31666,
31800, 37498, 39494, 40666, 51958, 73051, 73065, 177924–26, ANWC R00677, BMNH 1926.2.25.94–95, NMV D
8177, D 72572, D 8463–64, D 8466, DTD 1165–67, NTM R.40, R.164, R.227, R.253, R.838, R.1228, R.1939, R.3002,
R.3455, R.4015–16, R.4734, R.6189, R.6350, R.8251, R.8374–75, R.9843, R.9876, R.10471, R.13213, R.13315, R.13379,
R.13450, R.16553, R.17298, R.20750, R.21221, R.21663, R.23970, R.26275, R.26423, R.26454, R.26494, R.27045,
R.34200–04, WAM R30951, R40297, R40303. Q
UEENSLAND
: AMNH R-69262, 69264, 82227, ANWC R2204, R2415,
R2617, CAS 103015–17, FMNH 97832–35, MCZ 128118, 128185–86, 137639–40, AMS 2160, 41544, 43911–12, 44232,
46004, 55594, 55683, 57126–27, 58999, 62429, 64224, 74310, 74678, 82585–86, 84299–300, 86779, 94364–65, 99712,
104829, “Stego” Thursday Is. [unnumbered], MSNG 7513, NMV D 7575, D 8527, D 8670, D 8771, D 66684, DTD 1164,
SAMA R3559, SMF 65223, WAM R10699. Fitzroy Island: SAMA R22391. T
ORRES
S
TRAIT
I
SLANDS
: BMNH
78.10.16.26–28, 83.4.14.25–26, 85.6.30.65–66. N
O
F
URTHER
L
OCALITY
: ANWC R5026, SAMA R3726, R22596–97,
R49875, R54097. INDONESIA, P
APUA
P
ROVINCE
, Biak: MSNG 7522-2, RMNH.RENA 4706, 46879, 46883, 46922–23,
46941–42. Boven Digoel: ZMA 17668, 17671. Jayapura: BMNH 1938.6.9.41, 1938.6.9.44–45, CAS 89687,
RMNH.RENA 4707, 43595, 46886, 47601–03, ZMA 14091, 14095, 17665. Merauke: RMNH.RENA 36115, 46895.
Mimika River: BMNH 1913.10.31.198–99. Numfoor: MSNG 7514-1–2, 7522-3, NMW 22594, RMNH.RENA 43645–46,
47588. Octakwa River: BMNH 1913.11.1.105. Stekwa River: BMNH 1913.11.1.106–07. Timika: AMS 148822. Yapen:
RMNH.RENA 46843–47, 46850, 47751. No further locality: BMNH 78.2.11.13, BPBM 3179, 3182, 3933–34, CAS
14978, FMNH 43010–11, 43022, MCZ 49475, 49483–85, RMNH.RENA 5823, USNM 119507–08, 124635, 140256,
ZMA 13915, 17663A–B, 17669–70, ZMB 8794, ZSM 57/2015. W
EST
P
APUA
P
ROVINCE
, Ayamaru: RMNH.RENA 46851,
46953, 47539, 47550–53, 47738, 47744–48. Fak-Fak Peninsula: BMNH 1908.6.30.12, MCZ 7312–13, 178511–12, NMW
22584.1–3, RMNH.RENA 46848–49. Gag: WAM R131172. Manokwari: RMNH.RENA 46852–55, 46920–21, 46924–25,
46954, 47546–49, 47554–58, 47592, 47734. Maybrat: RMNH.RENA 46884–85, 46888–92, 47533, 47535–37, 47540–44.
Raja Ampat: BMNH 70.8.3.147. Tehak: RMNH.RENA 47538, 47545, 47750. “Irian Jaya,” no further locality: AMNH R-
62037–38, ZMB 23854. “Vogelkop,” no further locality: IRSNB 12652-1–2, MSNG 7522-1, 30184, 30188, NMW
22589.1–2, 22595, RMNH.RENA 14026, 43605, 46881–82, 46887, 46893–94, 46899–900, 46917–18, 46949.
“Wandamen Bay”: RMNH.RENA 14029. “Wasior”: RMNH.RENA 46880. PAPUA NEW GUINEA, A
UTONOMOUS
R
EGION
OF
B
OUGAINVILLE
: AMNH R-42402. C
ENTRAL
P
ROVINCE
: AMNH R-59087, 82522–23, 115062, 117757, BMNH
1908.10.14.9, 96.10.31.21–22, 97.12.10.119, BPBM 18939, 19479, 19481–82, 22557, 27333, 44325–26, CAS 118878,
118886, 118940, MCZ 59053, 104074–75, 111792–93, 116764, 116766, 116768–69, 142589, 142647, 145949, 146130,
KAISER ET AL.
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·
Zootaxa 4590 (2) © 2019 Magnolia Press
149743, 152427, MSNG 52313-1–2, PNGM 22335, 24494, 24497, 24506–10, UPNG 548, 1337, 1864, 2185, 5975,
6205–06, 6306, USNM 213453–59, 213537, ZSM 122/2000. E
AST
N
EW
B
RITAIN
P
ROVINCE
: AMS 7036, BMNH
77.2.24.20, BPBM 5128, NMW 22590.1–4, SAMA R8700–01, ZMB 14582. E
AST
S
EPIK
P
ROVINCE
: AMNH R-75000,
75023–26, 75249, 75359, AMS 28694–95, 95602–05, 129221–22, 129228, 129305, ANWC R1010, R1016, R1068,
R1106, BPBM 3774, 22194, 34779, 34784, 35952–53, 35957, CAS 116017, MCZ 153121, NMV D 7579, D 7694, PNGM
23230, 24688, ZSM 123/2000. E
ASTERN
H
IGHLANDS
P
ROVINCE
: AMNH R-85720, 85724, 95643, 98492, CAS 118877,
118887, MCZ 84146, 121569. G
ULF
P
ROVINCE
: AMS 14478, BMNH 1936.7.7.30, CAS 118879, MCZ 59051, 59056–57,
UPNG 7508, 8546, USNM 562930. M
ADANG
P
ROVINCE
: AMNH R-107182–93, AMS 15053, 19008–09, 25235,
25380–86, 25607, 25749, 28698–99, 28701–05, 28725, 28766, 31221, 31473–81, 68937, 120863–68, 121256, 121302,
121434–36, 121471, 124069, 124096, 124110, 124507, 124624, 124684–85, BMNH 1976.266, 1977.2231,
1922.11.24.36–38, 1926.5.31.15–17, 85.1.8.4–5, BPBM 5715, 21677–80, 31241, 31260, 33635, 34776–78, 35960, CAS
23724, 192893, 192943, IRSNB 13553, 13569, 13896–97, 16723–24, 16731–33, 16801–05, 16815–18, 16820, MCZ
49491, 132795, MSNG 30185, NMW 22585.1–5, 22586.1–2, 22586.4, 22587.1–2, 22588.1–2, 22596, PNGM 24513,
RMNH.RENA 46902–15, 47740–43, UPNG 961, 2402–03, USNM 567183, 567188, ZSM 124/2000. M
ANUS
P
ROVINCE
,
Manus: AMNH R-107189, AMS 28684–87, 130344, BPBM 2172, IRSNB 16679, 16806–07, MCZ 137655, 141003,
PNGM 22977, UPNG 8491–92, USNM 121260–61. Los Negros: AMS 129031–41, USNM 121736, 160009, 567184–87.
Pak: ZMB 28985. Pisik Lou: AMS 19305. Rambutyo: ZMB 27546. M
ILNE
B
AY
P
ROVINCE
: AMNH R-42379, 73951,
73956, 73964, 73971, 76614, 76622–24, 76677–78, 76702, 76704, 76712, 76715, AMS 6516, 69262, 86835–36, 86841,
124834, 124862, 124881–82, 124895, 124898–99, 125012, 129712, 129734, 129743, 137977, BPBM 16527, 16529–33,
17289–92, 17424, 17880, 17882–84, 20821, 39689–96, 39698–701, 42178–79, FK 10149, FK 10170, FK 10212, FK
10244–45, FK 10271, FK 10295, FK 10315, FK 10450, FK 10545, MSNG 57359, NMW 22591.1, 22591.3, 22597,
PNGM 22304, 22521, 23061, 24498–501, UPNG 5400, 5411. Trobriand Islands: BMNH 95.10.17.47. M
OROBE
P
ROVINCE
: AMNH R-66750, 66752, 85713–16, 85721–22, 95154–57, 95571–72, 95574–77, 95619–23, 101086,
101093–94, 101096–98, 103666–67, 103672, 104082–83, 107180, 142878, AMS 12973–74, 15966, BMNH 1965.595,
1922.11.24.40–41, BPBM 2777, 3716, 3919, 3924, 4160, 5134, 5439, 5441, 5450, 5453–54, 5505, 6234–37, 6243, 6277,
6516, 18940, 21674–76, 23703–04, 23736, 23737–38, 24145, 24147–48, 25444, 25590, 30804–05, 31729–32, 36147,
36166, 38841, CAS 113636, 113638, 113641, 193020, 198314, MCZ 44175, 121570–71, 145947, 149737–40, 149742,
NMV D 10475, PNGM 22782 "A", 22782 "B", UPNG 1450, 2728, 5736, USNM 119510–11, 119746, 159898–99, ZMB
21193, 21195–96, 22283. N
ATIONAL
C
APITAL
D
ISTRICT
: MCZ 59054, 137505, PNGM 24687, USNM 213444,
213449–50. N
EW
I
RELAND
P
ROVINCE
: SMF 18058–63, ZMB 24069, 26475. New Hanover: ZMB 8928. N
ORTHERN
P
ROVINCE
: AMNH R-111801–02, AMS 9572, 9847, 14859, 47461–62, BMNH 1922.11.24.39, 1935.5.10.159, BPBM
3787, 5012, 5495, 36142, 36144–45, 36189, 37588–89, 39810–12, 43030–31, 43033, MCZ 142371–72, PNGM 22976,
22303, 23245, UPNG 3900–02. S
ANDAUN
P
ROVINCE
: AMNH R-100036–45, AMS 127708, BPBM 3468, 23449–52,
NMV D 7573. S
IMBU
P
ROVINCE
: AMNH R-98866, 98870, 98872, AMS 115319–26, 115342–43, 115360–61, BPBM
5025, CAS 99918–20, 103376–77, 109105, 113634–35, 113639–40, 113642, 114110–14, 118881, 118888–89, 119176–77,
119179–80, 119182MCZ 83166–67, 83170, 85048, 116760–61, 116765, 121567, 121572, 121575–77, 121580–81, UPNG
8342–43, USNM 562929. S
OUTHERN
H
IGHLANDS
P
ROVINCE
: AMS 122359–60, 122904–07, BPBM 34386–88,
34880–81, MCZ 121568, NMV D 49960. W
EST
N
EW
B
RITAIN
P
ROVINCE
: CAS 113637, PNGM 22974. W
ESTERN
P
ROVINCE
: AMNH R-57530, 106279, 111794–96, AMS 121054, CAS 121222, CAS 121242, 127371, 127388,
132200–02, 132208, 132227, 133795, UPNG 3672, 5668, 7526, 7527, 8556, BMNH 85.6.30.32–33, 1987.916, MCZ
116767, 118676, 121573–74, 129136, 130033, 130316, 130333–36, 137551–52, 137641, 139515, 140752, 141002,
141067, 141320–22, 142345, 152426, MSNG 54479-2, 54480, 54487-1–2, PNGM 22975, 24650, UPNG 8617, USNM
213451–52. Bobo: MCZ 137488, 137546. Daru: MCZ 134881, 137489, 137545, 137617, 137954, 140808, 141360,
PNGM 22070, 22073, 24512. W
ESTERN
H
IGHLANDS
P
ROVINCE
: AMNH R-59891, 103668–71, AMS 14774–76, 14778,
14791–92, 16576, 16578, 16619, 16623, CAS 132931, PNGM 22268. “New Britain,” no further locality: FMNH 13880,
13885. N
O
F
URTHER
L
OCALITY
: ZMB 22179, 22211, 22235, 24063. “New Guinea,” no further locality: AMNH R-86558,
AMS 1015, 6968, 12774, 25505–07, 40348, 57291, 121225, 124482, 131142, BMNH 78.10.16.17, 80.9.23.5, 96.7.8.5–6,
1938.6.9.42–43, BPBM 38314, AA23803, FMNH 13872, 14059, IRSNB 530, 15814, 16812–13, MNHN 1320, NMP
32825, PNGM 24493, 24728–29, 24778, 24829–31, RMNH.RENA 5135, 5137–38, 5824–26, 5880, 5892, 6361, 9838,
14023, 14025, 14031–32, 47735, 47739, 47749, 47752, 47951, SMF 18053, 18064, UPNG 963–67, 7525, 7644, 7951,
8660, 8720, 8778–79, 8856, USNM 213537, ZMA 13436, 14089, 17661, 17664, ZMB 14800–01, 17663–64, 18260,
23987, 24030, 24177A–C, 64546–58, 78751–69, ZSM 68/2015. TIMOR-LESTE: USNM 579383–84, 580549–50.
