Hidden in the Heart of Borneo-shedding light on Some Mysteries of an Enigmatic Lizard: First Records of Habitat Use, Behavior, and Food Items of Lanthanotus borneensis Steindachner, 1878 in Its Natural Habitat

Abstract

A short assessment of the behavior, habitat and habitat use of the sole member of its monotypic family, the Earless Monitor lizard Lanthanotus borneensis was undertaken for the first time. Field data of this important species were lacking for more than a century since its description in 1878. Nearly nothing is known about life history or even its natural habitat. The only previously known facts were taken from a few captive animals held in the early 60s and 70s of the last century. Metric data of size, weight, body temperature, and sex ratio of 19 individuals were examined. Natural food items were identified. The population structure of a subpopulation was estimated. A brief survey of the additional herpetofauna is given for the area.

Full text

HIDDEN IN THE HEART OF BORNEO-SHEDDING LIGHT ON SOME MYSTERIES
OF AN ENIGMATIC LIZARD: FIRST RECORDS OF HABITAT USE, BEHAVIOR,
AND FOOD ITEMS OF Lanthanotus borneensis STEINDACHNER, 1878
IN ITS NATURAL HABITAT
Christian Langner1
Submitted June 22, 2016
A short assessment of the behavior, habitat and habitat use of the sole member of its monotypic family, the Earless
Monitor lizard Lanthanotus borneensis was undertaken for the first time. Field data of this important species were
lacking for more than a century since its description in 1878. Nearly nothing is known about life history or even its
natural habitat. The only previously known facts were taken from a few captive animals held in the early 60s and
70s of the last century. Metric data of size, weight, body temperature, and sex ratio of 19 individuals were
examined. Natural food items were identified. The population structure of a subpopulation was estimated. A brief
survey of the additional herpetofauna is given for the area.
Keywords: Squamata: Lanthanotidae: Lanthanotus borneensis; habitat; measurements; behavior; nutrition;
possible threats.
INTRODUCTION
Although Lanthanotus borneensis was described al-
ready in 1878 by the well-known Austrian herpetologist
Franz Steindachner, this enigmatic species fell asleep for
roughly the next 100 years before a few tens of animals
reemerged from their century of “hibernation.” After
some severe flooding in Sarawak, Borneo, in the early
1960s, several earless monitors appeared and were sent
from Sarawak Museum to different institutions in Europe
and the US. Unfortunately these individuals were
brought to the museum by road workers and locals,
so again no details of their life history were available.
Some of these Lanthanotus were kept alive for several
years in captivity and the first important behavioral stud-
ies of this rare and unsearchable species could be made
and some mysteries could be lifted (Harrisson T., 1961,
1963, 1966; Harrisson B., 1961, Harrisson and Haile,
1961; Mertens, 1964, 1966, 1970; Proud, 1978). All fur-
ther assumptions about the natural history, behavior and
specialized morphological adaptations of this unique spe-
cies, remained highly speculative (Sprackland, 1971,
1976).
Apart from that, only phylogenetic research was un-
dertaken which illustrated what an outstanding position
in platynotan phylogeny Lanthanotus occupies. The
earless monitor represents the Lanthanotidae, a monoty-
pic family of its own (McDowell and Bogert, 1954; Mer-
tens, 1961; Rieppel, 1980). After this promising but short
period, the subsequent decades revealed no further
knowledge on the species and it remained untraceable for
the scientific world.
Only in the recent past have a few animals turned up
again. Now, for the first time in Kalimantan, the Indone-
sian part of Borneo, where the species’ occurrence was
expected but had not been previously verified. Yaap et al.
(2012) described an individual from the north of
Kalimantan Barat, and Vergner (2013) published the first
verification for the easternmost province Kalimantan
Timur, where it was found in the northeastern region,
close to the Malaysian border. Unfortunately detailed
research was not undertaken, as in both cases the discov-
eries were made by uninitiated researchers, not her-
petologists, but ethnologists and entomologists. Fortu-
nately, the significance of their discoveries was realized
and the articles were published in collaboration with a
Herpetologist.
During previous research surveys conducted during
1999 and 2006, primarily in the northeastern part of
Kalimantan Timur (Kayan Menterang National Park), we
received multiple plausible reports from local commu-
1026-2296/2017/2401-0001 © 2017 Folium Publishing Company
Russian Journal of Herpetology Vol. 24, No. 1, 2017, pp. 1 – 10
1Alstätte 23, 48727 Billerbeck, Germany; e-mail: langner.ch@gmx.de

nity members that earless monitors do exist in this area.
The first indication of the species’ presence in this region
was published by Aulya (2006), and confirmed by
Vergner (2013). In April 2014, a new effort was made to
shed some light on the totally unknown life history of
Lanthanotus borneensis, with the intention to find them
for the first time in their natural habitat. The starting
point of this new survey took place in northeast Kaliman-
tan, which we were familiar with from previous expedi-
tions. Unfortunately, we had no success finding any spec-
imens of Lanthanotus again despite positive feedback
from the locals. So we decided to change locations and
head over to the west coast. Due to the narrow time frame
of our expedition, we had only a few days left and we
were not very confident that we would be successful in
this area either. However, the main intention was to eval-
uate and get an impression of the local conditions and
habitat. Three days before having to leave homeward, we
received another promising hint from some local villag-
ers. Still not too optimistic, we decided to make our last
effort and planned a night survey for the late evening,
since night surveys in Borneo are always a pleasure, es-
pecially in an area we have never been before. At a small,
not very promising-looking, downhill stream, surrounded
mainly by palm oil plantations, really and truly our life-
time dream came true. The first earless monitor appeared
in the cone of our headlights. Needless to say what an
awesome memorable moment it was to bring this long
missing phantom to the surface.
MATERIAL AND METHODS
The following material was used during fieldwork:
kern pocket balances CM 320-1N; pH-Fix test strip, mea-
suring range 4.5 – 10.0; steel ruler 500 mm Holex; Digi-
tal vernier caliper, Burgwächter Profiscale-Precise PS
7215; Tinytag plus, Gemini Data Logger; Garmin GPS
Map 62 S; Barigoprecision Hygrometer Supratherm;Dig-
2 Christian Langner
Fig. 1. Habitat of Lanthanotus borneensis: downhill section of the
habitat.
Fig. 2. Microhabitat of Lanthanotus borneensis in a more uphill sec-
tion of the habitat.

ital Laboratory Thermometer GTH 175/MO1000 Grei-
singer Electronic. Length specification was made in
0.1 mm steps; weighing was made in in 0.1 g steps; air
temperature and humidity was quantified 1 m above
ground level; water temperature was quantified at a depth
of approximately 20 cm in a shallow pool; body tempera-
ture was taken from the cloaca; temperature was mea-
sured in °C; one data logger was deposited in a tree hole
at 2 m above ground level. A second one was placed in
a crevice between boulders at ground level above the
maximum flood line; pH measurements were made at
several pools, as well as within flowing parts of the water
channel.
RESULTS AND DISCUSSION
Habitat. The small stream flowed through a larger
riverbed infrequently framed with sandstone boulders,
rocks and large pebbles, in a series of several downhill
steps (Fig. 1). The surfaces of rocks were often over-
grown with moss. The water-bearing part of the stream
consisted of some pools, ~2 m wide and up to 50 cm
deep, at maximum (Fig. 2). In the more uphill region the
remaining stream was predominantly comprised of a
small rivulet, only a few centimeters in width and depth.
The water was clear, not pigmented from tannins.
The streambed soil surface was sandy, interspersed with
pebbles of various sizes. In some pools, the substrate was
a very fine sediment material which muddied the water
quickly after movement. This fine material indicated that
the period of the slow flowing, low water level had lasted
for some time. Furthermore, the stream pool substrate
was covered by a dense layer of dead leaves. Underwater
vegetation was completely absent.
The whole (mostly dry) streambed was ~4 m wide.
The actual stream would certainly be much broader dur-
ing the peak of rainy season. The surveyed section of the
stream where Lanthanotus borneensis were found was
~400 m long.
The altitude of the locality rises from 178 m at the
beginning of the searched part to 197 m a.s.l. at the end of
the 400 m survey section.
The vegetation was dominated by secondary growth
with low trees, bushes, reeds, and bamboo, interspersed
with fragments of primary dipterocarp forest patches and
some remaining sole larger trees. Some trees were used
to harvest caoutchouc. Neighboring close-by was farm-
land; mostly palm oil plantations but also rice, manioc
and taro fields. In some distance, larger areas of pristine
primary rainforest still exist in the adjacent hills.
First Records of Lanthanotus borneensis Steindachner, 1878 in Its Natural Habitat 3
Fig. 3. Submerged male Lanthanotus borneensis in situ.

The locality is in the central northern part of the Indo-
nesian province Kalimantan Barat, not far from the Ma-
laysian border of Sarawak. GPS coordinates were re-
corded but are not stated here for obvious reasons. Unfor-
tunately, in the recent past several earless monitors have
been offered multiple times in the pet trade. It is not yet
clear how vulnerable the species is regardless of the high
population density described below. Efforts are under-
taken to include the currently not-listed earless monitor
in the highest protection category “A” of the Convention
on International Trade in Endangered Species of Wild
Fauna and Flora (CITES), Nijman and Stoner, 2014,
CITES 2015). In a period of about one and a half hours
after the discovery of the first Lanthanotus 8 additional
specimens were located; all were active on the ground.
Earless monitors were observed on land (Fig. 4) as well
as submerged in water (Fig. 3) at a rate of 10:9; always
in close proximity to the water. In three nights, a total of
19 earless monitors where observed at this location and at
two additional neighboring streams. The weather, even in
the early dry season, was nonetheless rainy. A light driz-
zle started in the early afternoon and held on with short
breaks throughout the whole night. This pattern contin-
ued in the following days. The first earless monitor was
found at half past eight in the evening, shortly after we ar-
rived in the area. The air temperature was 28.4°C at this
time and dropped to 26.2°C by midnight. Humidity
changed only in relation to rain intensity and ranged from
89 to 96%. Water temperature was stable at 26.1°C. The
pH was at a neutral range of 7.0. Body temperature was
measured in three females and one male. The range in
body temperature was at a relatively consistent, from
26.1 to 26.6°C (see Table 1). As accompanying herpeto-
fauna we could verify the following species: Limnonec-
tes kuhlii,Ansonia spinulifer,Duttaphrynus melanostic-
tus,Leptobrachella cf. mjobergi,Leptobrachium abotti,
4 Christian Langner
Fig. 4. Female Lanthanotus borneensis on land in situ.

Cyrtodactylus consobrinus,Cnemaspis cf. kendallii,
Phoxophrys nigrilabris,Gonocephalus grandis,Taky-
dromus sexlineatus,Tropidophorus brookei,Varanus du-
merilii,Python breitensteini, and Xenopeltis unicolor.
The only tadpoles detected in the pools belonged to Lep-
tobrachium abotti in the developmental stages 34 – 46
(after Gosner, 1960). Freshly metamorphosed froglets of
this species, as well as adults, were also found in the leaf
litter around the pools.
Measurement and sex ratio. Due to the limited time
frame, a detailed study of the habitat was conducted only
at the stream habitat surveyed on the first night. Only
metric and weight data were recorded for the Lanthano-
tus discovered during the subsequent evening surveys of
the additional stream habitats. All but one metric data
were within the expectable parameters known from the
limited references of the species’ size, Mertens (1961),
Harrisson (1963). The largest individual was a male with
a total length of 441 mm, slightly larger than the previ-
ously recorded maximum size of 438 mm (Proud, 1978).
The determined female:male sex ratio was 2:1. One fe-
male appeared noticeably heavier and was likely gravid.
Another female was very skinny and looked weak and
unhealthy with a shrunken pelvis. All other individuals
appeared strong and healthy. The individual sex could be
determined with relative ease, even in the field, due to a
distinct sexual dimorphism. The males have distinctly
broader heads as well as a broadened tail base (Figs. 5
and 6). Furthermore, hemipenes could be partially
everted in all males, with gentle manipulation at the tail
base. No juveniles were found.
Behavior. Nearly all males exhibited some forms of
healed injuries. Toes and parts of the tail were missing,
and in two males a complete amputation of the shank of a
hind leg was observed (Fig. 7.). Most of the males had
healed white scares on the head, neck and/or dorsum.
Only one smaller male had no old trauma visible. Fe-
males had no signs of partially missing extremities, only
First Records of Lanthanotus borneensis Steindachner, 1878 in Its Natural Habitat 5
TABLE 1. Recorded Field Data of Lanthanotus borneensis
Weight,
g
Snout-vent
length,
mm
Tail
length,
mm
Head
width,
mm
Sex Remarks
86 180 175 34 xShank of the right back leg missing, tail partly missing
60 177 187 22 x
84 178 186 19 }
87 190 214 26 xScars on right front-leg and neck, 3 fingers missing on feet of same leg
55 159 188 17 }Body temperature 26.4°C
67 171 192 18 }
88 193 204 23.5 xOnly short stump of right back femoral was left, tail tip missing, body temperature 26.1°C
48 158 174 16.5 }
120 220 221 23 x2 fingers are missing on left front leg, one on right front leg
51 160 176 17 }Tail tip missing, body temperature 26.6°C
107 182 205 20 }Presumably gravid
65 168 189 18 }Body temperature 26.4°C
90 200 90 25 xMore than half of the tail missing, scars on head and back, on both hind feet 2 toes missing
68 170 190 18 }
67 170 188 17.5 }Tail tip missing
50 156 174 17 }
42 162 175 16.5 }Skinny
65 169 187 18 }
Fig. 5. Sexual dimorphism in Lanthanotus borneensis, distinct shape
of tail base: left, female; right, male.

some tails appeared possibly incomplete. No evidence of
tail regeneration could be determined. The lower number
of males observed, together with the high rate of injuries
sustained, led to the conclusion of a highly territorial
aggressive behavior between male earless monitors. The
sharp teeth are able to inflict serve injuries and a whole
extremity could be easily ripped off completely. These
sharp-edged dentition structures could be interpreted as
an adaptation to their slimy and hard-shelled food re-
sources (e.g., catfish, crustaceans, worms). The behav-
ioral pattern of handled Lanthanotus was mostly docile
and passive; some showed a distinct thanatosis behavior.
However, they were able to bite suddenly with unex-
pected high speed, and to inflict deep, heavy bleeding
lacerations with their razor blade-like, sharp teeth.
A deep cut, resulting from a bite to the author’s left
pointer finger healed without complications after a few
days. No signs of intoxication from saliva or bacteria
were noticed. Males were much more aggressive than fe-
males when handled. When detected, the earless moni-
tors remained motionless “frozen” on the ground in the
head torch beam with one exception: in one case an indi-
vidual sprinted with unexpected speed into a crevice be-
tween two large boulders, out of reach for further inspec-
tion. In opposition to the observation of captive animals
made by Harrisson and Haile (1961) and Mertens (1966)
which were described as lethargic and not very agile,
they are quite capable of surprising speeds at short dis-
tances. At individuals we encountered submerged the
lower transparent eyelids were predominantly closed
(Figs. 8 and 9). A further noticeable observation was a
rapid movement of the throat; an oscillation similar to
that visible sometimes in the throat pouches of frogs.
Tounge flicking behavior was notisted infrequently just
in a few cases in handled individuals (Fig. 11). Another
interesting and unexpected detection was the ability of
vocalization! A hushed gentle squeak was recognized on
two separate occasions while handling females. In con-
trast to the observation made by Yaap et al. (2012), no
Lanthanotus were observed in the open during the day.
We found one female earless monitor in its hiding place
under a large rock fragment, ~70 × 40 × 40 cm, at 09.30
the next morning (Fig. 10). The temperature within this
retreat was 26.8°C. The substrate was moist and the cav-
ity which remained after turning the rock filled slowly
with water. Some cockroaches of the semi aquatic spe-
cies Rhicnoda natatrix were found at the same hiding
place between dead leaves. At night this cockroach spe-
cies was seen frequently submerged in the pools of the
stream, a potential food item for Lanthanotus. Data log-
gers were deposited for future visits to determine the cli-
mate conditions in the seasonal change of a longer pe-
riod. Telemetry research is planned in the future.
Nutrition. Data on the natural food items of earless
monitors are very poor, if not completely unknown.
Solely, Pregill et al. (1986) cite unpublished data of
6 Christian Langner
Fig. 6. Sexual dimorphism in Lanthanotus borneensis, distinct wide of
the head: left, male; right, female.
Fig. 7. Amputated right hind shank of a male Lanthanotus borneensis.

Greene — in a dissected specimen from the Museum of
Comparative Zoology at Harvard University. Fragments
of earthworm setae were found in the specimens’ intesti-
nal tract. In another case, an unpublished observation by
Green and Schwenk was cited, stating that Lanthanotus
feed on earthworms and crustaceans (Losos and Greene,
1988). No further comments or details were given and it
remains unclear in both cases, if the statements refer to
captive individuals. Furthermore, conclusion of the food
items could be only made from captive animals, where
earthworms, strips of fish (Pleuronectes platessa, exclu-
sively), egg yolk of Chelonia mydas as well as chicken
and pig liver stripes were accepted as food (Harrisson,
1961; Mertens, 1966; Proud, 1978). Additionally impor-
tant results of the natural composition of natural food
items could be presented hereby first time from live ani-
mals. Two females regurgitated slightly digested fresh-
water shrimps of the species Macrobrachium cf. pilima-
nus. Numerous individuals of this crustacean species
were observed in the leaf litter layer of the pools.
In another case a male Lanthanotus regurgitated two
small pieces of highly digested fishbone vertebra which
were identified, via DNA analysis, as belonging to the
catfish genus Clarias, most likely imbedded in species
complex of C. tejsmanni. The only fish species we could
verify in several pools was an unidentified snakehead
species of the genus, Channa. Close to the daytime re-
treat of the female Lanthanotus borneensis we detected a
partly crushed and eaten shell fragment of a land crab,
presumably of the genus Barytelphusa. It is rather un-
likely that this large crustacean would be a part of the
earless monitor’s diet. The crab was more likely con-
sumed by one of the Duméril’s monitors (Varanus dume-
rilii) we encountered in the same habitat. These monitors
are known to feed largely on landcrabs (Krebs, 1979).
Possible threats. The habitat is influenced by vari-
ous human activities in multiple manners. Local villagers
use the stream for washing, fishing and as a source for
drinking water. Earless monitors were reportedly seen by
locals very infrequently, just every few years. Some in-
digenous people from the area stated, that they had never
seen them. Some were not aware of their existence at all.
All human activities in the habitat are restricted to day-
time hours which limit interactions. The earless monitor
is not especially feared, and is not hunted or killed but it
is known that they are able to inflict painful bites. The
First Records of Lanthanotus borneensis Steindachner, 1878 in Its Natural Habitat 7
Fig. 8. Head portrait of a male Lanthanotus, note the partly closed semitransparent lower eye lid.

fact that we found earless monitors in three neighboring
habitats in an unexpected high population density in ad-
dition to the circumstance that local people responded
mostly positive of the species existence in the various
areas surveyed in northwestern and northeastern Kali-
mantan, together with the repeated note that they were
found even in rice fields and drainage channels, I come to
the cautious optimistic conclusion that the species is
much more common and widespread than previously
supposed. Aside from the abundant synanthropic house
gecko species of the genus Hemidactylus of Southeast
Asia, I have rarely seen such a dense population in a liz-
ard species. The sympatric, water skink (Tropidophorus
brookei), occurring in the same microhabitat, for exam-
ple lives in small family groups at a distance of
400 – 600 m away from the next group. A fortiori it is un-
intelligible that nonetheless earless monitors stayed hid-
den for so long considering the extensive herpetological
work in recent years. The apparently stable population of
Lanthanotus borneensis in a heavy anthropogenically in-
fluenced environment draws the conclusion that the spe-
cies copes much better under these conditions than it
8 Christian Langner
Fig. 9. Head portrait of a female Lanthanotus, note the completely closed semitransparent eyelid that was predominantly closed in submerged
individuals.
Fig. 10. Frontal head portrait of female Lanthanotus borneensis.

First Records of Lanthanotus borneensis Steindachner, 1878 in Its Natural Habitat 9
Fig. 12. Wider vicinity of the locality of Lanthanotus borneensis in north western Kalimantan Barat, Indonesia. Recently established palm
oil plantation visible in background.
Fig. 11. Tongue-flicking behavior of a Lanthanotus borneensis female.

would be suspected, taking the rare discoveries in the
past as a basis. The natural habitat (rock edged streams)
will remain mostly untouched, even if the surrounding
habitat will undergo considerable changing and habitat
degradation. The most devastating problem nowadays on
Borneo is the rapidly growing palm oil production
(Fig. 12) with its disastrous impacts of one of the most di-
vers and important ecosystems on earth.
Acknowledgments. I am very grateful to various people
for their kind support in the field. For the possibility to partici-
pate research projects in Kalimantan in the past, I would like to
thank Dr. Peter Sound, Boppard, and Prof. Dr. Michael Veith,
University of Trier, Germany. Further I would like to thank the
Eva-Mayr-Stihl-Stiftung for their kind financial support of
these projects. I have to thank Dr. Kristina von Rintelen, Leib-
niz-Institut für Evolutions- und Biodiversitätsforschung, Ber-
lin, Germany, and Werner Klotz, Rum, Austria, for their kind
and immediate help in determine the freshwater shrimp species.
For the DNA sequencing and determination of Clarias I would
like to thank Claudia Etzbauer, Simon Walter, and Mathias Gei-
ger, Museum Alexander Koenig, Bonn, Germany, for their kind
support. Finally I have to thank Prof. Dr. Wolfgang Böhme,
Museum Alexander Koenig, Bonn, Germany, and Rob Nixon
Tucson, Arizona, USA, for their valuable comments on an ear-
lier draft of the Manuscript.
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