Accepted Manuscript A new species of Ilyoplax (Decapoda, Brachyura, Dotillidae) from Gujarat, India N. Trivedi Jigneshkumar, M. Soni Gunjan, J. Trivedi Dhruva, D. Vachhrajani Kauresh, Professor of Zoology PII:
S2287-884X(15)00013-8
DOI:
10.1016/j.japb.2015.02.005
Reference:
JAPB 59
To appear in:
Journal of Asia-Pacific Biodiversity
Received Date: 8 January 2015 Revised Date:
16 February 2015
Accepted Date: 27 February 2015
Please cite this article as: Trivedi Jigneshkumar N, Soni Gunjan M, Trivedi Dhruva J, Vachhrajani Kauresh D, A new species of Ilyoplax (Decapoda, Brachyura, Dotillidae) from Gujarat, India, Journal of Asia-Pacific Biodiversity (2015), doi: 10.1016/j.japb.2015.02.005. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
ACCEPTED MANUSCRIPT
A new species of Ilyoplax (Decapoda, Brachyura, Dotillidae) from Gujarat, India
RI PT
Trivedi Jigneshkumar N., Soni Gunjan M., Trivedi Dhruva J. and Vachhrajani Kauresh D.* Marine Biodiversity and Ecology Lab., Department of Zoology, Faculty of Science, The M. S. University of Baroda, Vadodara, Gujarat, India.
*Corresponding author detail:
SC
Dr. Kauresh D. Vachhrajani Professor of Zoology,
Faculty of Science, The M. S. University of Baroda, Vadodara, Gujarat, India. E-mail Id -
[email protected]
TE D
Phone no: 91-9427839382.
M AN U
Department of Zoology,
Abstract: A new species, Ilyoplax sayajiraoi belonging to the family Dotillidae is described from Gujarat state, India. It differs from its closely related species, I. stevensi (Kemp, 1919) and I. frater (Kemp, 1919) by differences in shape and proportions of the carapace, shape and setae
EP
arrangement pattern on chela and morphologies of male and female abdomen. The details on the
AC C
morphological differences are presented in the study.
Key words: Ilyoplax; Brachyuran crab; Mudflat; Gulf of Khambhat; Gujarat
Introduction
Brachyuran crabs belonging to genus Ilyoplax Stimpson, 1858 are commonly found on
the intertidal mudflats distributed from temperate to tropical Indo- Western Pacific region. Although, currently the genus consists of 27 well established species (Kitaura and Wada 2006; Ng et al 2008; Davie and Naruse, 2010; Fatemi et al 2011), the genus has not been studied in detail for its taxonomy. Earlier, Serene and Lundoer (1974) studied the taxonomy of the genus
ACCEPTED MANUSCRIPT
based on male gonopod structure and distributed all the identified species into three different groups. Recently, Kitaura et al (1998) and Kitaura and Wada (2006) studied the taxonomy of the genus using molecular tools and identified three distinct lineages in the genus. The taxonomy and diversity of brachyuran crabs belonging to family Dotillidae Stimpson, 1858 found in coastal
RI PT
area of India have been studied in detail by few workers (Alcock 1900; Kemp 1919). So far total 11 species and 4 genera belonging to family Dotillidae have been reported from India (Alcock 1900; Kemp 1919). Out of these 11 species, only two species belonging to genus Ilyoplax i.e. Ilyoplax gangetica (Kemp 1919) and Ilyoplax stapletoni (de Man 1908) are reported from India
SC
(Alcock 1900; Kemp 1919). Species belonging to genus Ilyoplax are smaller in size but occur in large number on the sandy and muddy shores and they play significant role in the ecology of the
M AN U
habitat (Nishihira 1984; Snowden et al 1991).The species belonging to particular genus also exhibit typical behavior patterns including chela waving display which provided basis for many behavioral experiments related to the reproductive and courtship aspect of the species as well as it is one of the important characters for species identification (Kosuge et al 1994; Wada and Wang 1998; Kitaura and Wada 2006).
The new species reported here was collected as a part of the research project
TE D
“Documentation of crustacean biodiversity of Gujarat” (Sponsored by Gujarat State Biodiversity Board, Government of Gujarat, India) of 2014-2015 undertaken by us at Marine Biodiversity and Ecology Lab., Department of Zoology, Faculty of Science, The M. S. University of Baroda, Vadodara, Gujarat, India. The details of morphological characters of the new species are given in
EP
the paper.
Materials and methods
AC C
The specimens of the species were collected from the coastal areas of Kamboi (220 12′ 59″ N; 72036′ 59″ E) located on the extreme interior of Gulf of Khambhat closer to lower estuarine region of Mahi Sagar River on the western side of Bharuch District of Gujarat state, India. The coastal areas of the sampling site supports mudflat habitat. The type material used for the present study is deposited in the Zoology Museum, Department of Zoology, Faculty of Science, The Maharaja Sayajirao University of Baroda, Vadodara, Gujarat, India. The general terminologies follow Davie and Naruse, 2010. The measurements are provided for the carapace width (CW)
ACCEPTED MANUSCRIPT
and carapace length (CL). Male G1 refers the male first gonopod. All the measurements were taken using digital vernier calipers (0.01mm accuracy).
RI PT
Systematic account Family Dotillidae Stimpson, 1858 Subfamily Dotillinae Stimpson, 1858
SC
Genus Ilyoplax Stimpson, 1858
Ilyoplax sayajiraoi sp. nov.
Type. Holotype:
M AN U
(Figure 1-3)
, ZL-AR-CR-76 (CW- 9.4 mm, CL- 6.3 mm), Kamboi (220 12′ 59″ N; 720
36′ 59″ E), Gujarat, India, mudflat habitat, 11th April, 2014 (Jignesh Trivedi and Gunjan Soni). Paratype: 5 , (ZL-AR-CR-76 [1-5]) (CW- 8.7 mm, CL-5.9 mm; CW- 9.1 mm, CL- 6.1 mm;
TE D
CW- 6.5 mm, CL- 4.7 mm; CW- 6.7 mm, CL- 4.9 mm; CW- 8.2 mm, CL- 5.2 mm) and two (ZL-AR-CR-76 [6-7]) (CW- 12.3 mm, CL-9.0 mm; CW- 12.1 mm, CL-8.8 mm), Kamboi, 14th May, 2014 (leg. Gunjan Soni and Dhruva Trivedi); 2 , (ZL-AR-CR-76 [8-9]) (CW- 6.4 mm,
EP
CL-4.2 mm; CW- 5.7 mm, CL-3.6 mm), 3 , (ZL-AR-CR-76 [10-12]) (CW- 12.1 mm, CL- 8.7 mm; CW- 12.2 mm, CL-8.9 mm; CW- 12.5 mm, CL- 8.5 mm), Kamboi, 15th August, 2014 (leg.
AC C
Kauresh Vachhrajani, Jignesh Trivedi and Gunjan Soni); 3 , (ZL-AR-CR-76 [13-15]) (CW- 8.4 mm, CL- 5.9 mm; CW- 7.7 mm, CL- 3.6 mm; CW- 7.3 mm, CL- 5.7 mm) and 3 , (ZL-AR-CR76 [16-18]) (CW- 11.0 mm, CL- 8.2 mm; CW- 12.3 mm, CL- 8.6 mm; CW- 10.7 mm, CL- 8.0 mm), Kamboi, 21st December, 2014 (leg. Kauresh Vachhrajani, Jignesh Trivedi, Dhruva Trivedi and Gunjan Soni).
ACCEPTED MANUSCRIPT
Comparative material. Ilyoplax frater (Kemp, 1919). 1♂, ZL-AR-CR-79, (CW- 7.4 mm, CL5.2 mm), Beyt Dwarka (220 46′ 72″ N; 690 11′ 54″ E), Gujarat, India, mudflat habitat, 13th November, 2014. (Jignesh Trivedi and Gunjan Soni); 1♂, (ZL-AR-CR-79 [1]), (CW- 6.3 mm,
RI PT
CL- 4.1mm), Beyt Dwarka (220 46′ 72″ N; 690 11′ 54″ E), Gujarat, India, mudflat habitat, 17th November, 2014. (leg. Jignesh Trivedi, Gunjan Soni and Richa Pareek).
Description. Carapace transversely oblong, anterior breadth one and half time the length, the upper surface of carapace slightly convex on both sides and moderately sculptured (Figure 3a).
SC
Anterolateral border slightly and evenly convex, the breadth in the middle decidedly larger than the breadth between orbital angles. An inconspicuous median furrow starts from the frontal
M AN U
region and ends in the middle of the gastric region, one large grove about one third of the breadth of the carapace defines the posterior limit of the gastric region. Shallow depression present parallel to the orbital region on both the sides, the depression excavates near the median furrow. On the branchial regions small tubercles bearing setae arranged in four rows, the anterior row small and indistinct made up of one or two tubercles, second row distinct with 2-4 tubercles, third row made up of 6-8 tubercles; most posterior row longest and bearing 6-10 tubercles; the
TE D
last row much transverse than longitudinal and if the row continued forward the line so form would pass through the proximal part of orbits. Sharp, perfectly transverse and straight ridge present above the posterior carapace margin. Front obliquely deflexed, apex broadly rounded; the lateral border slightly constricted near the base; front breadth one sixth or seventh of the anterior
EP
breadth of the carapace.
Orbits slightly oblique, not transverse. The upper orbital border beaded with distal portion
AC C
beaded with large tubercles, it excavates near the base of the front and distinctly concave in the outer half. The lower orbital border finely crenulated. On the floor of the orbits, a ridge runs parallel to the lower border of the orbits throughout the length, outer orbital angle consists of an acute tooth directed outwards; a tuft of setae present near the base of the tooth. Small notch present behind the outer orbital angle which runs downs with series of minute tubercles. The lateral margins defined by sharp crest bearing long setae. Epistome short with narrow triangular median tooth. The buccal cavity completely closed by third maxillipeds, ischium of third maxillipeds sub quadrate in shape with anterior border lined by setae (Figure 3d); merus shorter than ischium and
ACCEPTED MANUSCRIPT
broader than long, ^ shaped furrow present interiorly, the outer grove of ^ shape turns upwards and forms a median furrow which runs towards the apical portion of ^ shape. The surface of the merus smooth. The exopod concealed under the maxillipeds and furnished with long slender flagellum.
RI PT
Chelipeds of male strong, slender, well developed and longer than female chelipeds, chelipeds length less than twice the length of the carapace. Merus trigonal with microscopically beaded edges, merus bears tympanum on the outer side and other large tympanum on the inner side, the outer side tympanum not always present. The carpus short smooth without tooth on the inner
SC
aspect; the inner margin of upper surface crested beneath it tuft of short setae present (Figure 3f). Fingers similar in length with the upper border of palm (Figure 3b), in the inner phase of palm,
M AN U
longitudinal row of setae present beneath the upper border, Four finely beaded carinae run backward from the tip of the fixed fingers on to the palm; the median two carinae run parallel and disappears before reaching the middle of the lower surface of palm; the inner most carinae curves upwards across the inner surface; the outer most runs along the lower part of the outer border till the proximal end of the palm (Figure 3c). The gap between the carinae without any tubercles; the rest of the surface including most of the outer part of palm smooth. The dactylus
TE D
slightly longer than the fixed finger, the distal half of lower border of dactylus and upper border of fixed finger bear crest of minute teeth.
Chelipeds of females weak and more slender than those of males, fingers twice the length of the upper border of palm. Carpus small, the upper border ridged and bears long setae; the palm
EP
crested above with long setae; the upper and lower margin of dactylus bear long setae; the upper margin of fixed finger also bear long setae; no denticulation on the dactylus and palm. Only two
AC C
carinae present runs backward from the tip of the fingers on to the palm, the carinae enclosing the lower flattened surface and bears long setae; the fingers gap wide near the base. Third walking leg longest, two and half times the length of the carapace; well defined tympana present on the upper and lower surfaces of merus of first two pairs and lower part of last two pairs; In the first two pairs, on dorsal surface a crenulated ridge runs parallel to the upper border of the proximal half of the merus; the edges of the merus are microscopically beaded. The carpus and propodus of second and third pair bear two carinae on the superior face; dactylus flattened and shorter than propodus. In large male a dense patch of tomentum present on the second walking leg extending from middle of the carpus to the distal third of the propodus
ACCEPTED MANUSCRIPT
(Figure 3e), in the young individuals no traces of tomentum observed. The basal segments of walking legs bears plumose setae. All the segment of male abdomen distinct (Figure 3g); second segment of abdomen narrower than first; the third and fourth segments rounded at the sides; the third segment little broader than
RI PT
fourth; the fourth segment little longer than third; the breath of fifth segment less than the half breadth of the fourth segment, fifth segment slightly constricted near its proximal half. The sixth segment less than twice as broad as long, the seventh segment triangular in shape, broader than long and more widely rounded distally. The female abdomen (Figure 3h) much broader than
SC
male but comparatively narrow at the base, the seventh segment triangular in shape.
The male G1 slender, tapering distally (Figure 3i), distal part of apical lobe deflexed outward
M AN U
(Figure 1j), long setae present on the outer surface and few long setae present along the inner part near the distal region.
Coloration. The carapace of the species is olive green in fresh specimen while the carapace color in preservation is grayish. The abdomen of male and female is off white or cream in color. The palm of male cheliped is pinkish purple in color up to proximal half while the distal half is
TE D
white in color. The color gets darker in the inner phase. The fingers are white in color. Distribution. The species is currently known from its type locality Kamboi (220 12′ 59″ N; 720 36′ 59″ E) which is coastal village located on the extreme interior of Gulf of Khambhat closer to
EP
lower estuarine mudflat region of Mahi Sagar River on the western side of Bharuch district of
AC C
Gujarat, State, India.
Habitat. The species occupies intertidal mudflat habitat (Figure 2b) where its distribution is restricted to upper intertidal area. The species is burrowing in nature and its density ranging from 15-25 burrows/m2. The burrow diameters of the species ranges from 5-11 mm while the burrow depth ranges from 15- 40 cm.
Etymology. The specific epithet sayajiraoi is named in honor of founder of Baroda Collage Sir Sayajirao Gaikwad III, Maharaja of the erstwhile state of Baroda, Gujarat.
ACCEPTED MANUSCRIPT
Discussion Serene and Lundoer (1974) have provided the identification keys for the species belonging to
RI PT
genus Ilyoplax based on the morphology of male G1. According to key provided by them, the present species falls in the group III which also includes species like I. stevensi and I. frater. The present species is very close to I. stevensi and I. frater which were described by Kemp, 1919 based on the material collected from the coastal area of Karachi, Pakistan. Though, I. stevensi
SC
and I. frater are very close to the present species, there are several significant morphological differences which separate the present species from other two closely related species. These
M AN U
significant morphological differences are listed below in the table (Table 1).
In addition to the above mentioned differences, the female abdomen of both the species also showed major differences. The different segments of female abdomen in the present species are
Acknowledgments
TE D
broader then the female abdomen of I. stevensi.
The authors are thankful to Gujarat State Biodiversity Board, Government of Gujarat for the financial support under the project “Documentation of Crustacean (Phylum Arthropoda)
EP
Biodiversity of Gujarat”. Trivedi Jigneshkumar is thankful to UGC, India for financial support under UGC-RFSMS scheme. The authors are also thankful to Dr. Reza Naderloo, Assistant
AC C
Professor, School of Biology and Center for Excellence in Phylogeny of Living Organisms, Collage of Science, University of Tehran, Iran for valuable comments on the identification of the species. The authors are also thankful to Ms. Barkha Purohit and Ms. Richa Pareek for technical support.
ACCEPTED MANUSCRIPT
References
Alcock A. 1900. Materials for the Carcinological Fauna of India (Catometopa or Grapsoidea).
RI PT
Journal of Asiatic Society of Bengal 69:279–456.
Davie PJF, Naruse T. 2010. A new species of Ilyoplax (Decapoda, Brachyura, Dotillidae) from Panglao, the Philippines. In Castro P, Davie PJF, Ng PKL et al. Studies on Brachyura: a Homage
SC
to Danièle Guinot. Crustaceana Monographs 11. Leiden: Brill. pp. 75-82.
De Man JG. 1908. The fauna of brackish ponds at Port Canning, Lower Bengal. Part 10.
M AN U
Decapod Crustacea, with an account of a small collection from brackish water near Calcutta and the Dacca district, Easter Bengal. Record of the Indian Museum 2: 211-231.
Fatemi SMR, Vosoughi GH, Valinasab T, Savari A, Ghotbeddin N. 2011. First report of dotillid crabs (Decapoda, Brachyura) from the Northern Gulf of Oman, Iran. Crustaceana 84 (14): 1745-
TE D
1753.
Kemp S. 1919. Notes on Crustacea Decapoda in the Indian Museum. 12. Scopimera. Records of Indian Museum 16(5):305–348.
EP
Kitaura J, Wada K, Nishida M. 1998. Molecular phylogeny and evolution of uniquemud-using territorial behavior in ocypodid crabs (Crustacea: Brachyura: Ocypodidae). Molecular Biology
AC C
and Evolution 15 (6): 626-637.
Kitaura J, Wada K. 2006. New species of Ilyoplax (Brachyura: Ocypodidae: Dotillinae) from the Philippines and Indonesia: behavioral, molecular, and morphological evidence. Raffles Bulletin of Zoology 54 (2): 373-379.
Kosuge T, Poovachiranon S, Murai M. 1994. Male courtship cycles in three species of tropical Ilyoplax crabs (Decapoda, Brachyura, Ocypodidae). Hydrobiologia 285: 93-100.
ACCEPTED MANUSCRIPT
Ng PKL, Guinot D, Davie PJF. 2008. Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology (Supplement) 17: 1-286.
(Tesch), newly recorded from Okinawa, Japan. Galaxea 3: 97-101.
RI PT
Nishihira M. 1984. Ecological notes on the rock dwelling ocypodid crab, Ilyoplax integra
Serène R, Lundoer S. 1974. Observations on the male pleopod of the species of Ilyoplax Stimpson with a key to the identification of the species. Phuket Marine Biological Center
SC
Research Bulletin, 3: 1-10.
M AN U
Snowden RJ, Clayton DA, Al-Taher EY. 1991. Population biology of Ilyoplax stevensi (Brachyura: Ocypodidae) on a Kuwait mudflat. Marine Ecology Progress Series71: 219-225.
Stimpson W. 1858. Prodromus description isanimaliume vertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federatamissa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit W. Stimpson. Pars. V. Crustacea
TE D
Ocypodoidea. Proceedings of the Academy of Natural Sciences of Philadelphia,10: 93-110
Wada K, Wang CH 1998. Territorial and sexual behaviours in Ilyoplax formosensis and I.
AC C
Tables
EP
tansuiensis (Crustacea, Brachyura, Ocypodidae). Journal of Taiwan Museum 51:119-125.
Table 1. The morphological differences between I. sayajiraoi sp. nov., I. stevensi (Kemp, 1919) and I. frater (Kemp, 1919).
Figures
Figure 1. Ilyoplax sayajiraoi sp. nov. Holotype (ZL-AR-CR-76, male (CW- 9.4 mm, CL- 6.3 mm). (Line bar indicates 1 cm)
ACCEPTED MANUSCRIPT
Figure 2. Ilyoplax sayajiraoi sp. nov. (a) In life (b) Mudflat habitat of Kamboi
Figure 3. Ilyoplax sayajiraoi sp. nov. (a) Carapace dorsal view, (b) Male chela outer view (c) Male chela inner view (d) Third maxillipeds (e) Second walking leg outer view (f) cheliped
AC C
EP
TE D
M AN U
SC
RI PT
carpus inner view (g) Male abdomen (h) Female abdomen (i) Male G1 (j) Male G1 apical lobe.
ACCEPTED MANUSCRIPT
Ilyoplax stevensi (Kemp, 1919)
Ilyoplax frater (Kemp, 1919)
Characters
(Kemp, 1919)
(Kemp, 1919; Fatemi et al., 2011)
Lateral borders are slightly and
Lateral borders are straight and only
Lateral borders are slightly and
evenly convex thus the breadth of
convex posteriorly thus the breadth of evenly convex thus the breadth of the
the middle surface slightly more
the middle surface slightly less than
middle surface slightly more than the
than the anterior breadth.
the anterior breadth.
anterior breadth.
Four rows of tubercles present on
Three rows of tubercles present on
Three rows of tubercles present on
the branchial region.
the branchial region.
the branchial region.
Only first row is indistinct and made
The first two rows are indistinct and
The first two rows are indistinct and
up of two tubercles.
made up of one or two tubercles.
made up of one or two tubercles.
The direction of last row is much
The direction of last row is as much
The direction of last row is much
transverse than longitudinal.
as longitudinal as transverse.
transverse than longitudinal.
If the last row continued the line so
If the last row continued the line so
If the last row continued the line so
form would cross the proximal part
form would cross the proximal part
form would cross the outer end of
of orbits on the opposite side.
of front.
orbits on the opposite side.
A small notch present behind the
A small notch present behind the
A small notch is not present behind
tooth shaped outer orbital angle.
tooth shaped outer orbital angle.
the tooth shaped outer orbital angle.
Front
SC
M AN U
TE D
AC C
EP
Carapace
RI PT
Morphological Ilyoplax sayajiraoi sp. nov.
The breadth of the front is one sixth The breadth of the front is one The breadth of the front is one sixth or seventh of the anterior breadth of eleventh or twelfth of the anterior or seventh of the anterior breadth of the carapace. The lateral borders are breadth of the carapace The lateral the carapace. The lateral borders are slightly constricted near the base.
borders are slightly constricted near not constricted near the base. the base.
ACCEPTED MANUSCRIPT
The outer half of the orbits distinctly The outer half of the orbits perfectly The concave.
outer
half
of
the
orbits
conspicuously concave.
RI PT
Chelipeds of males are strong, Chelipeds of males are weak and Chelipeds of males are strongly slender, well developed and much only slightly stouter than females.
developed, much deeper and longer
stouter than female.
than females.
The fingers are more or less equal to The fingers are 1.5 times longer than The fingers are slightly longer than the length of the upper border of the length of the upper border of the length of the upper border of palm.
Palm is slender in shape.
Palm is slender in shape.
M AN U
palm.
palm. Palm is swollen in shape.
The upper margin of inner surface of The upper margin of inner surface of The upper margin of inner surface of carpus is crested and beneath it there carpus is crested and beneath it there carpus is crested and beneath it there is a tuft of short setae.
is a thick tuft of very long setae.
is a thick tuft of very long setae.
TE D
The space between four carinae The space between four carinae The space between four carinae running backwards from the tip of running backwards from the tip of the running backwards from the tip of the fix finger (inner face) on to the fix finger (inner face) on to the palm the fix finger (inner face) on to the
of tubercles.
EP
palm is not filled with any type/size is filled with extremely minute palm is filled with extremely minute tubercles.
AC C
Chelipeds
straight.
SC
Orbits
tubercles.
The dactylus is slightly longer than The dactylus is equal in length with The dactylus is equal in length with the fixed finger.
fixed finger.
fixed finger.
Dentition is present in the distal half Dentition is present in the proximal Dentition present throughout the of the dactylus.
half of the dactylus.
length of the dactylus with large tooth present in the centre of the
ACCEPTED MANUSCRIPT
dactylus. Small dentition is present on the
distal half of the fixed finger.
distal half of the fixed finger.
RI PT
Small dentition is present on the Fixed finger is without dentition.
Long setae not present on the lower Long setae present on the lower Long setae not present on the lower border of the fixed fingers.
border of the fixed fingers.
The edges of merus are beaded.
The edges of merus are spinulose.
The edges of merus are beaded.
SC
Walking legs
border of the fixed fingers.
Large tomentum present on the Large tomentum present on the Large tomentum not present on the
M AN U
carpus and propodus of the second carpus and propodus of the second carpus and propodus of the second walking leg.
walking leg.
Third segment is broader than Third and fourth segments are same Third and fourth segments are same fourth.
in breadth.
in breadth.
Fifth segment is less than the half Fifth segment is more than the half Fifth segment is more than the half breadth of the fourth segment.
TE D
breadth of the fourth segment.
breadth of the fourth segment.
The sixth segment is less than twice The sixth segment is twice as broad The sixth segment is twice as broad as long.
EP
as broad as long.
AC C
Male abdomen
walking leg.
as long.
AC C
EP
TE D
M AN U
SC
RI PT
ACCEPTED MANUSCRIPT
AC C
EP
TE D
M AN U
SC
RI PT
ACCEPTED MANUSCRIPT
AC C
EP
TE D
M AN U
SC
RI PT
ACCEPTED MANUSCRIPT
