Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 October 2021 | 13(12): 19753–19761
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.7500.13.12.19753-19761
#7500 | Received 06 June 2021 | Final received
07 August 2021 | Finally accepted 26 September 2021
New record of Myrmarachne
melanocephala MacLeay, 1839 (Araneae:
Salticidae) from Jharkhand, India and biogeographical
implications of the co-occurrence of its ant model Tetraponera
rufonigra Jerdon, 1851
Rahul Kumar 1, Mirtunjay Sharma 2
& Ajay Kumar Sharma 3
1,3 University Department of Zoology,
Vinoba Bhave University, Hazaribagh, Jharkhand
825301, India.
1 Department of Zoology, Sheodeni Sao College, Kaler, Arwal, Bihar 824127, India.
2 Forest Department Office,
Hazaribagh, Jharkhand 825301, India.
1 rahuldayanand33@gmail.com
(corresponding author), 2 sharmamirtunjay@gmail.com, 3 ajaysharmavbu@gmail.com
Editor: John T.D. Caleb, ERI, Loyola
College, Chennai, India. Date of publication:
26 October 2021 (online & print)
Citation: Kumar, R., M. Sharma & A.K.
Sharma (2021). New record of Myrmarachne
melanocephala MacLeay, 1839 (Araneae:
Salticidae) from Jharkhand, India and biogeographical
implications of the co-occurrence of its ant model Tetraponera
rufonigra Jerdon, 1851.
Journal of Threatened Taxa 13(12): 19753–19761. https://doi.org/10.11609/jott.7500.13.12.19753-19761
Copyright: © Kumar et al. 2021. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: None.
Competing interests: The authors
declare no competing interests.
Author details: Rahul Kumar is working as an Assistant
Professor at Department of Zoology, Sheodeni Sao
College (Magadh University), Kaler, Arwal, Bihar. His doctoral research is going on from
University Department of Zoology, Vinoba Bhave
University, under Dr. Ajay Kumar Sharma. His doctoral work concerns with
mimicry among terrestrial arthropods and new approaches to explore integrative
taxonomy. He has research experience in molecular biology and functional
genomics from School of Life Sciences, JNU, New Delhi as UGC-JRF; in
nanotechnology and biotechnology from Department of Anatomy, AIIMS, New Delhi
as DST-Nanomission JRF; and in biosystematics and
integrative taxonomy from Division of Entomology, ICAR-IARI, New Delhi as
DST-SERB JRF. Mirtunjay Sharma is a forest department employee,
a Limca Book of World Record holder hobbyist
entomologist-arachnologist and a famous nature photographer from Hazaribagh,
Jharkhand. He is author of the book “Spiders of Jharlkhand”
published by Jharkhand Biodiversity Board. Dr.
Ajay Kumar Sharma is working as Associate Professor at University
Department of Zoology, Vinoba Bhave University,
Hazaribagh, Jharkhand and is also assigned with the positions of University
Dean of Science and course coordinator of the Department of Physiotherapy in
the same university. He has more than 40 years of research and teaching
experience. His thrust areas of research are molecular biology, entomology and
ecology.
Author contributions: Conceptualization of the research
and manuscript writing was done by RK. Site visits, data collections and
analyses were done by RK, MS and AKS. Manuscript was reviewed and finalized by
AKS before communication. Correspondence to the journal and subsequent revisions
were done by RK.
Acknowledgements: We acknowledge Dr. John Caleb from the Zoological Survey of India,
Kolkata, and Miss Stuti Rai and Mr. Anand Harshana from Division of Entomology, Indian Agricultural
Research Institute (IARI), Pusa, New Delhi, for
sharing their valuable taxonomic insights. We also acknowledge Dr. Shashank P.R., Division of Entomology, IARI, for
allowing us to use the microscopy facility of National Pusa
Collection, Division of Entomology, IARI, for our studies. Two anonymous reviewers
are thanked for their suggestions and critical comments which helped us to
improve our manuscript.
Abstract: We report the occurrence of the
ant-mimicking jumping spider Myrmarachne melanocephala MacLeay, 1839 for the first time from
Hazaribagh Wildlife Sanctuary, Jharkhand, India. Digital illustrations and
descriptions of the spider, the female’s exuviae, and
video records of a live male are also presented. The distribution pattern of M.
melanocephala has not been studied in detail
across India whereas its ant model, Tetraponera
rufonigra Jerdon, 1851
is known to have a wide distribution. Co-occurrence of the mimic and the model
implies a wider range of biogeographical distribution of these species in
India.
Keywords: Distribution, eastern India, exuviae, Hazaribagh Wildlife Sanctuary, jumping spider,
mimicry, myrmecomorphy.
Abbreviations: ALE—anterior lateral eye |
AME—anterior median eye | md—mid-dorsal | pd—pro-dorsal
| PLE—posterior lateral eye | PME—posterior median eye | pv—proventral | rv—retroventral.
INTRODUCTION
Ant-mimicry (myrmecomorphy) is
not very uncommon among arthropods. Most myrmecomorphs display Batesian mimicry
where non-ant species mimic unpalatable and aggressive ants in order to avoid
predatory attacks which is the result of adaptive evolution (Cushing 1997).
Myrmecomorphy has achieved a distinct level of perfection among salticids
(jumping spiders). The genus Myrmarachne
MacLeay, 1839 is one of the largest salticid genera consisting of 186 accepted
species globally out of which 23 are found in India but only six of them are
well characterized and all of these species exhibit Batesian mimicry displaying
profound morphological as well as behavioural resemblances towards ants (Caleb
2016; Prószyński 2016; Wanless
1978; World Spider Catalog 2021; Yamasaki & Ahmad
2013; Yamasaki & Edwards 2013; Caleb & Benjamin 2017; Caleb &
Sankaran 2021).
The genus Myrmarachne
forms part of a phylogenetic group of the subfamily Salticinae
Blackwall, 1841 (Maddison 2015). Myrmarachne
melanocephala MacLeay, 1839 is the type species
of the genus Myrmarachne which was originally
described from Bengal (which corresponds to present day West Bengal in India
and Bangladesh combined) (Edwards & Benjamin 2009; World Spider Catalog 2021). M. melanocephala
mimics the worker of Tetraponera rufonigra Jerdon, 1851, an
arboreal ant with conspicuous eyes and long slender body (Pocock 1909). This
species was originally described from the Carnatic-Malabar region (which
corresponds to present day Indian states: Karnataka, Kerala, Tamil Nadu, and
Andhra Pradesh) (Ward 2001). The genus Tetraponera
Smith, 1852 belongs to subfamily Pseudomyrmecinae of Formicidae which is represented by 95 extant species
globally, of which 10 are found in India (Bharti & Akbar 2014; Bolton
2021).
This paper is concerned with the
discovery of M. melanocephala from Hazaribagh
Wildlife Sanctuary, Hazaribagh, Jharkhand with detailed morphological
descriptions and observation on its ant model T. rufonigra.
For the first time, in addition an undamaged whole body exuviae
of a freshly moulted female M. melanocephala
has been used here as material for
morphological descriptions of this spider. Both taxonomic as well as
behavioural (mimicry related) aspects have been noted. The biogeographical
implications of the co-occurrence of the mimic and the model together in the
same habitat has also been discussed in the Indian context.
MATERIALS
AND METHODS
Sampling and collection were
performed at Hazaribagh Wildlife Sanctuary, Hazaribagh, Jharkhand, India (Image
1). The spiders and ants were observed on and around Sal Shorea
robusta trees. One male and two female spider
specimens were spotted on a Sal tree trunk. The male specimen was captured
alive and one freshly moulted female specimen and its undamaged whole body exuviae were manually collected and preserved in 70%
ethanol for further investigations. The live male spider and ant specimens were
photographed and videoed using a cellphone camera
(Samsung M42). Measurements of the live male spider specimen were taken in millimeters (mm) using ocular micrometer
placed within the eyepiece of a stereoscopic microscope by keeping the material
on a cavity slide with a drop of water. A coverslip was temporarily placed over
the cavity of the cavity slide in order to keep the material static for quick
measurements. The water drop was added to create surface tension in order to
keep the cover slip in firm position without killing the spider. The specimen
was kept alive for behavioural studies. Note: one leg of the male spider got
detached while trying to measure it alive. The ventral and dorsal sides of the
palp and dentition of the live male spider were studied using hand-held
magnifying glasses and stereoscopic microscope. The ethanol preserved specimen
of the female spider was dissected for its epigyne,
palp, and head containing chelicerae and fangs using a fine surgical scalpel. Epigyne was cleared by boiling it in 10% potassium
hydroxide (KOH) solution for a minute. After boiling, the epigyne
was rinsed thoroughly in water to remove excess KOH and was temporarily mounted
in a drop of glycerol using coverslip on a glass slide for microscopic
observations. The ethanol preserved female spider (before dissection), its
undamaged whole body exuviae, and dissected epigyne, palp & head with chelicerae & fangs were
photographed using Leica DFC 425C digital camera mounted over Leica M205FA stereozoom automontage microscope
at National Pusa Collection, Division of Entomology,
Indian Agricultural Research Institute, Pusa, New
Delhi 110012. Measurements were taken in millimeters
(mm) using inbuilt settings of the automontage. The format of the description follows
Yamasaki (2010), Caleb (2016), and Ward (2001). The studied ethanol preserved
specimen of the female spider and its exuviae were
deposited in the museum collections of University Department of Zoology, Vinoba
Bhave University, Hazaribagh.
TAXONOMIC
NOTES ON THE MYRMECOMORPH
Myrmarachne melanocephala
MacLeay, 1839
(Images 2A–D, 3 & 4A–E; supplementary
video 1)
Myrmarachne melanocephala MacLeay, 1839: 11, pl. 1, fig.
4; Galiano, 1969: 146; Edwards & Benjamin, 2009: 5, figs. 1A–H, 2A–D, 3A–D,
4A–E, 5A–D; Yamasaki & Edwards, 2013: 15, figs. 46–58; Yamasaki &
Ahmad, 2013: 541, figs. 32A–G, 33A–H, 34A–C; Benjamin, 2015: 17, figs. 17A–D,
18A–D, 19A–D; Caleb, 2016: 409, figs 20–30.
Further references may be read in
World Spider Catalog (2021).
Materials examined (n=3). 1 preserved freshly moulted
female specimen, 1 preserved complete whole-body exuviae
of female, and 1 live male. Hazaribagh Wildlife Sanctuary (24° 8′ 14.7372″ N,
85° 23′ 1.3956″ E ), Hazaribagh, Jharkhand, India, 20.iv.2021, R. Kumar &
M. Sharma.
Diagnosis (following Yamasaki & Edwards
2013). Pedicel in both sexes as long as ALE-PLE. Males are further
distinguished from other congeners by the shape and dentition of chelicerae.
Females can be distinguished by abdominal markings and structure of epigyne (Yamasaki & Edwards 2013). For a complete
diagnosis and description see Edwards & Benjamin (2009) and Benjamin (2015).
Female. Body legth
7.8; carapace length 2.25, width 1.5; abdomen length 3.4, width 1.66. Width of
eye row I 1.11; II 1.1; III 1.2. Eye sizes and interdistances:
AME 0.3, ALE 0.2, PME 0.05, PLE 0.2; ALE-PLE 0.8; ALE-PME 0.39. Leg spination: tibia I pv 4, rv 5; metatarsus I pv 2, rv 2; tibia II pv 3, rv 3; metatarsus II pv 2, rv 2. Pedicel 0.6 long. Cephalic region rugulose
and dark brown to black, covered with white hairs; thoracic region reddish
brown, sparsely covered with white hairs. Cephalic region slightly higher than
thoracic region. Sternum brown. Chelicerae dark brown, geniculate with seven
teeth on promargin and 8 teeth on retromargin
(Image 2D). Legs I and II light yellow, coxae I brown, tarsi I light brown
which gets darker near the tip, legs III, IV brown, trochanter and patella of
leg IV yellowish, patella of leg III lighter in colour. Leg IV longest. Palp
paddle shaped and fringed with preening setae (Image 2C). Abdomen elongate, and
slightly constricted in the anterior third, covered with white hairs, almost
oval (Image 2A). Epigyne with laterally oriented
large oval copulatory atria; copulatory ducts twist to form butterfly shaped
structure just before reaching ovoid spermatheca; lateral pockets present
between artia just at the bases of copulatory ducts
(Image 2B).
Exuviae of female: The undamaged whole body exuviae revealed morphological features of the female
spider (Image 3). Exuviae contains imprints of the
outer surface of spider’s body. Abdominal markings (dark and lightly pigmented
areas) are sharply revealed in the exuviae which is
otherwise not clearly visible on the spider body. The inflated abdominal region
displays the site of exit of the spider after moulting from its own exuviae. Locations of four pairs of limbs, limb markings, geniculate
chelicerae, fangs, and paddle shaped pedipals are
clearly visible.
Male: Body legth
7.5; carapace length 3, width 1.2; abdomen length 2.9, width 1.5; chelicerae 2.
Pedicel 0.5 long. Legs I and II light brown but not yellow as in female, coxae
and trochanters of legs I, II and III white, and tarsi I brown which gets
darker near the tip, legs III and IV brown in colour, trochanter and patellae
of leg IV white. Cephalic region rugulose and black;
thoracic region reddish brown. Cephalic region slightly higher than thoracic
region. Chelicerae black throughout except the region from where fangs arise
which is reddish brown. Chelicerae porrect with 10 teeth each on prolateral and
retrolateral margin (Images 4B, E; supplementary
video 1). Sternum
light brown. Abdomen elongate-oval, constricted in the anterior third. Palp
with oval cymbium, round tegulum
with distal-retrolateral C-shaped sperm duct, embolus
with two coils, spiralled helix like retrolateral
tibial apophysis with prominent flange (Images 4C, D).
Remarks: Exuviae has been used as a material for
species description by some workers, and some have used it even as a holotype (Kranzfelder et al. 2017; Lin et al. 2017). We have used it
here as additional material for
morphological studies. We suggest that exuviae can
also provide such useful information while describing a species which may not
be clearly recognizable in the type or non-type material.
Biology: The spider specimens were spotted
on a tree trunk at the Sal forest. Refer ecological notes on mimic-model
coexistence in a subsequent section for other important details.
Distribution: Pakistan to Indonesia (Edwards
& Benjamin 2009; World Spider Catalog 2021), the
records from India include states of Tamil Nadu, West Bengal, Assam,
Uttarakhand (Caleb 2016) and Jharkhand (present study) (Images 1 & 6).
TAXONOMIC NOTES ON THE ANT MODEL
Tetraponera rufonigra
Jerdon, 1851
(Image 5A–D; supplementary
video 2)
Tetraponera rufonigra
Jerdon, 1851: 111;
Smith, 1877: 68; Bingham, 1903: 108; Wheeler, 1922: 1015; Ward, 1990: 489;
Ward, 2001: 649.
Further references may be read in
Bolton (2021).
Materials examined (n= 6): 1 live solitary dealate gyne and 5 foraging live workers. Hazaribagh Wildlife
Sanctuary (24°8′14.7372″ N, 85°23′1.3956″ E), Hazaribagh, Jharkhand, India, 20.iv.2021,
R. Kumar & M. Sharma.
Diagnosis: (following Bolton 2021). Larger
than other Tetraponera species. Bicoloured
body. Head and gaster darker than mesosoma. In case
of the examined specimen, head and gaster are black
in contrast to the orange-brown mesosoma. The species is so distinctive that
its identity has never been a matter of confusion or doubt. For a complete
diagnosis and description see Ward (2001).
Dealate gyne: Large body (larger than other
species of Tetraponera) with broad head, with
small but conspicous crytalline
glass like compound eyes and three ocelli. Head densely punctate but without
clearly visible puncture interspaces. Clypeus with long and narrow median lobe.
Bicoloued body, head and gaster
dark in colour (dark brown to black) which contrast with the orange-brown
mesosoma. Antennae, mandibles, tarsi and protibia
light in colouration (light brown to yellowish-brown), matching with the colour
of mesosoma to some extant (Image 5B). The studied specimen also shows the
presence of a conspicuously wide semicircular band of
highly pigmented (dark brown to black) wing rudiment which contrasts with the
light orange background of the mesosoma which is characteristic feature of the
dealate gyne (Image 5D).
Worker: They display similar
morphological features and colouration pattern like the dealate gyne except the wing rudiment which is absent in workers.
Pronotum with well developed lateral margin and punctate
humeral corners (Image 5C). Workers are smaller than the dealate gyne. Abdomen is more tapering in the workers compared to
the more inflated abdomen of the dealate gyne (Image
5A).
Remarks: This ant is known to exhibit
regional morphological variations. For example, the ants studied from some
places in Sri Lanka are uniformly dark (mesosoma approaching colour of head and
gaster) in comparison to the typical bicoloured ants
like those found in India (head and gaster darker
than mesosoma). As already reported by various workers, common morphological
variations include variation in integument sculpture, colour of mesosoma, and pilosity (Ward 2001).
Biology: The ants were spotted on and
around trees of the Sal forest. Refer ecological notes on mimic-model coexistence
in a subsequent section for other important details.
Distribution: Pakistan to Indonesia including
India (Ward 2001). Widely distributed in India; the records from India include
Andaman & Nicobar Islands, Arunachal Pradesh, Assam, Bihar, Delhi, Goa,
Gujarat, Haryana, Himachal Pradesh, Jammu & Kashmir, Ladakh,
Jharkhand, Karnataka, Kerala, Maharashtra, Manipur, Meghalaya, Mizoram,
Nagaland, Orissa, Punjab, Rajasthan, Sikkim, Tamil Nadu, Tripura, Uttar
Pradesh, Uttarakhand, and West Bengal (Bharti et al. 2016) (Image 6).
DISCUSSION
Ecological notes on mimic-model
co-existence
As an ideal myrmecomorph, M. melanocephala shares many morphological features with
its ant model T. rufonigra. The most striking
feature is its ant like bicolored body and size. Both share similar body colour
pattern, body size and appearance. The female spider mimics the ant more
efficiently than the male spider due to the absence of long chelicerae found in
the male spider. The long pedicel of the spider mimics the whole petiolar and
post-petiolar structure of ants due to its comparable length. The inflated
elongate ovate abdomen of the spider and ant also appears similar in
appearance. The limbs of the spider also display a lighter colouration pattern
like that of the ant (Images 2A, 4A, B, 5A, B). Along with morphological
resemblances, Myrmarachne melanocephala MacLeay, 1839 also exhibits some
behavioural similarities to its ant model. In this regard, the first two pairs
of legs are lighter in colour (yellowish in female) than the third and fourth
pairs, and bear no stripes or markings in the spider (Image 2A). The spider
waves its first pair of limbs in the air to mimic the long antennae (which is
also yellowish) of the ant model occasionally. The spider also displays a
zig-zag movement like the ant along with occasional salutatory movement typical
of the jumping spiders (Supplementary video 1 & Supplementary
video 2). Therefore, M. melanocephala perfectly imitates the gait and gestures
of its ant model T. rufonigra.
Biogeographical implications of
mimic-model co-discovery
Most myrmecomorphic spiders have
been found to occupy spaces in close
vicinity to their ant models (Pekar & Jarab 2011a). Such associations provide a space devoid of
potential enemies as ants are mostly avoided by frequent predators who prey
upon birds, wasps and spiders (Edmunds 1974). Association of myrmecomorphs with
a particular ant species could be the result of either preference to ant
dominated habitat or ant’s prey preference (Pekár
2014). A large number of myrmecomorphic species go with the former strategy
where they prefer to occupy ant dominated habitats and they capture prey other
than ants. M. melanocephala fall under the
same category (Edmunds 1978; Oliviera 1988; Pekár 2014; Pekár & Jarab 2011b). Apparent from their global distribution
pattern, both M. melanocephala and T. rufonigra are adaptable to multiple climatic regions.
The association between M. melanocephala and
T. rufonigra was first observed by Pocock (1909).
Global natural distribution of the ant model T. rufonigra
completely overlaps with the reported distribution of its mimic M. melanocephala which means they follow a parallel
biogeographical distribution pattern (Ward 2001; Yamasaki 2010). But if we look
at India (which has the largest land area among the countries falling under the
habitat range of the spider and the ant), we find that the myrmecomorph is
confined to only five states (including the present study) as per available
reports, whereas the ant model is widely distributed across Indian Territory
(24 states including Jammu & Kashmir along with Ladakh
and three other union territories) (Bharti & Akbar 2014; Caleb 2016) (Image
6). Therefore, in India T. rufonigra shows
a continuous distribution pattern whereas M. melanocephala
shows a highly discontinuous distribution pattern. This big difference in the distribution
pattern may be attributed to the gap in studies pertaining to this spider in
India. This is also true for many other spider species in India. The spider
fauna of India has never been studied in entirety as noted by Keswani et al. (2012). This gap in study in turn may be
attributed to a very small number of arachnologists in India. It can also be
observed that all the states from which M. melanocephala
has been reported also have reports of T. rufonigra.
In the present study also, the spider mimic has been reported from the state
where the ant model is already known to exist. Local extinction of M. melanocephala in other states where it coexisted with T.
rufonigra in past or range expansion of T. rufonigra outside the states from where M. melanocephala have been reported may be speculated as a
reason of such a sharp difference in their distribution pattern but it cannot
be inferred with certainty because of the huge study gap and also due to high
adaptive capabilities of these animals to multiple climatic regions, there is
no apparent plausible reason to presume such extreme events in their natural history.
Therefore, we suggest that M. melanocephala
could also be present in other states from which the ant model has already been
reported. M. melanocephala could have a wider
distribution across different states of India vis-à-vis the distribution of the
ant model T. rufonigra.
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