identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
8BEE4000238959A1861757426110D5E4.text	8BEE4000238959A1861757426110D5E4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Duttaphrynus brevirostris (Rao 1937)	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Duttaphrynus brevirostris (Rao, 1937)</p>
            <p>Figs 1, 2, 3, 4</p>
            <p>Original name and description.</p>
            <p> Bufo brevirostris Rao, 1937. Rao, C. R. N. 1937. On some new forms of  Batrachia from S. India. Proceedings of the Indian Academy of Sciences. Section B 6: 387-427. Type locality. "Kempholey, Hassan District, Mysore State," Karnataka, India. Current status of specific name. Valid name, as  Duttaphrynus brevirostris (Rao, 1937). </p>
            <p>Material studied.</p>
            <p> Topotype.   An adult male, BNHS 6126 (SVL 45 mm), from Kempholey Ghat region in Sakleshpur taluk,  Hassan district , Karnataka State, India  ,   collected by S.  D. Biju and Sonali Garg in June 2013. Other referred specimens. An adult male, SDBDU 2008.410 (SVL 48.6 mm), from Bhagamandala,  Kodagu district , Karnataka State  ;   an adult male, SDBDU 2015.3075 (SVL 46 mm), from Manipal,  Udupi district , Karnataka State  ;   and a subadult, SDBDU 4714 (SVL 25 mm), from Someshwara,  Udupi district , Karnataka State  . </p>
            <p>Rediscovery and validation of taxonomic status.</p>
            <p> This species was described based on a single specimen ("snout to vent, 27.00 mm") deposited in the Central College, Bangalore (CCB). This original name-bearing type specimen is considered lost (Dubois 1984; Biju 2001) and the species currently is known only from its original description. Rao (1937) enumerated several morphological character states to describe this taxon, but did not provide comparisons with other species. Our collection from a region of Kempholey Ghat in Sakleshpur taluk, that is part of the type locality (Rao 1937), is comparable with the original description with respect to several mentioned characters such as "canthus rostralis angular," "nostril nearer to the end of the snout than to the eye," "first finger equal to the second," "parotoids elongate, moderately prominent," and "upper surface of the skin covered with small uniformly distributed tubercles; with a small row of larger warts on the median line of the back." The primary inconsistencies between  Rao’s described specimen and our new collection involve snout-vent length, SVL 45 mm (vs. "27.00 mm") and weakly developed or inconspicuous cephalic ridges (vs. "crown without bony ridge"). The cephalic ridges in our new collection are relatively smooth, depressed, or less conspicuous (Figs 1A, C, 2A) when compared to other species of the  Duttaphrynus melanostictus group from Peninsular India. Hence, presence or absence of this character may be considered a matter of interpretation depending on degree of its prominence. Furthermore, the body size disparity between our collection and that of Rao (1937) also suggests that the type specimen he described could have been a subadult. We examined another subadult specimen from Someshwar (SDBDU 4714; SVL 25 mm), previously reported along with DNA sequence data (Van Bocxlaer et al. 2009), and found some comparable characters such as "a small row of larger warts on the median line of the back," "a network of dark lines," and "a dark temporal line extending to the sides," which can usually also be observed in subadults of  Duttaphrynus melanostictus group species (S.  D. B., personal observations). The Someshwar specimen is genetically identical to our Sakleshpur collection. Together, these two populations are also morphologically and genetically similar to our additional collections from other localities within the Malenadu (Malnad) and adjoining coastal regions of Karnataka (see 'Material  studied’ ). Altogether, we consider the available morphological and molecular evidence reliable for assigning all the mentioned populations to  D. brevirostris (Rao, 1937). </p>
            <p> Since the absence of a name-bearing type has contributed towards poor knowledge and uncertainty regarding the taxonomic identity of this taxon, as evident from the absence of new records, below we provide a detailed description of a newly-collected voucher specimen from the original type locality (Kempholey Ghat region in Sakleshpur taluk, Hassan district, Karnataka State, India: BNHS 6126), which is largely consistent with what is known of the former name-bearing type (Rao 1937). The topotype description provided below, augmented by a range of variation observed in vouchered specimens and genetic data from additional localities (Table 1; Suppl. materal 1: Tables S3, S4), validate the identity of  D. brevirostris and also serve as a redescription of this poorly known species for the benefit of future taxonomic work. </p>
            <p>Description of topotype, BNHS 6126</p>
            <p>(measurements in mm). A medium-sized, robust adult male (SVL 45.0); head of moderate size, wider (HW 16.9) than long (HL 14.0); snout subovoid in dorsal and ventral view, not projecting, its length (SL 6.1) longer than horizontal diameter of eye (EL 5.9); loreal region obtuse with sharp canthus rostralis; distance between posterior borders of the eyes (IBE 13.9) 2.2 times the distance between the anterior borders (IFE 6.3); interorbital space 1.2 times wider (IUE 5.1) than upper eyelid width (UEW 4.1); nostril oval without lateral flap of skin, closer to tip of snout (NS 1.7) than to eye (EN 3.2); tympanum distinct (TYD 2.6), vertically oval, 44.1% of eye diameter (EL 5.9), tympanum to eye distance (TYE 0.7); pineal ocellus absent; vomerine ridge and teeth absent; tongue small, oval, entire, median lingual projection absent; parotoid glands present, oval, flat, without spines and warts, longer (PL 6.2) than wide (PW 3.4), shorter than distance between them (PD 8.7); supraorbital and postorbital ridges weakly developed.</p>
            <p>Forelimbs short; forearm length (FAL 10.8) shorter than hand length (HAL 11.3); fingers rather thin, FLI nearly equal to FLII, FLIII longest (6.3); relative length of fingers: I=II&lt;IV&lt;III; tips of fingers rounded; subarticular tubercles prominent, single on fingers I, II, IV, double in finger III, oval, all present; prepollex oval, distinct; single rounded prominent palmar tubercle; numerous supernumerary tubercles irregularly set on palm.</p>
            <p>Hind limbs relatively long and thin, thigh length (TL 17.8) shorter than shank length (SHL 18.8) and foot length (FOL 18.5); relative length of toes: I&lt;II&lt;V&lt;III&lt;IV; tips of all toes rounded, without discs; webbing between toes present, small: I1+-2II1+-3III2-3⅔IV3⅔-2V; well-developed dermal fringes present on all toes; subarticular tubercles rather distinct, oval, all present; inner metatarsal tubercle present, prominent, its length (IMT 1.6) nearly half the length of outer metatarsal tubercle (OMT 3.1); numerous supernumerary tubercles irregularly set on foot.</p>
            <p>Skin. Dorsal and lateral surfaces of head and snout, and skin between eyes relatively smooth; anterior and posterior parts of back with flat and smooth glandular projections; flanks glandular without horny spinules or warts; dorsal surfaces of thigh, shank, and tarsus with smooth glandular warts. Ventral surfaces of throat, chest, belly, and thighs glandular.</p>
            <p>Secondary sexual character. Male: light brown granular projections on lateral surfaces of fingers I, II, and III.</p>
            <p>Colour in preservation. Dorsum and limbs slate grey to buff coloured; lateral surfaces of head, flank, and groin slightly lighter than dorsum; ventral surfaces (including limbs) off-white; throat with a faint light bluish-grey calling patch (Fig. 1). Colour in life: dorsum uniformly golden yellow with a brown tinge; limbs darker than dorsum; ventral surfaces white with a prominent bluish-yellow calling patch on throat.</p>
            <p>Variation.</p>
            <p>Adult size range: SVL 45-49 mm. Morphometric data from three adult males, including the described topotype, is given in Table 1. Dorsal colour varies from dark brown to golden yellow with a brown or reddish tinge; prominence of cephalic ridge varies from being inconspicuous to rather prominent; parotoid glands more prominent in life and relatively flattened in preservation; dorsal skin texture varies from having smooth glandular projections to glandular warts.</p>
            <p>Comparisons.</p>
            <p> Duttaphrynus brevirostris differs from other congeners that have relatively prominent cephalic ridges (  D. chandai ,  D. himalayanus ,  D. kiphirensis ,  D. mamitensis ,  D. manipurensis ,  D. melanostictus ,  D. microtympanum ,  D. mizoramensis ,  D. nagalandensis ,  D. parietalis ,  D. scaber ,  D. silentvalleyensis ,  D. stuarti ,  D. wokhaensis ,  D. crocus ,  D. kotagamai ,  D. noellerti , and  D. totol ) by its relatively smooth and inconspicuous cephalic ridges (vs. prominent and often with carotenoid margins or spinules), and smooth glandular dorsal skin (vs. presence of prominent glandular warts with horny spinules). Specifically, it also differs from the Indian species by the following characters: from  D. chandai , by its shorter male snout-vent length, SVL 45-49 mm (vs. longer, SVL 67-89 mm), absence of canthal, parietal, and cranial ridges (vs. present), and distinct tympanum (vs. inconspicuous externally); from  D. himalayanus ,  D. kiphirensis ,  D. mamitensis ,  D. manipurensis ,  D. melanostictus ,  D. microtympanum ,  D. mizoramensis ,  D. nagalandensis ,  D. parietalis ,  D. scaber ,  D. silentvalleyensis , and  D. wokhaensis , by absence of canthal, preorbital, and supratympanic ridges (vs. present), relatively flat parotoid glands (vs. prominently raised), and ventral surfaces of hand, fingers, foot, and toes with smooth tubercles (vs. raised and spinular tubercles); and from  D. beddomii ,  D. hololius ,  D. peninsularis , and  D. stomaticus by the presence of supraorbital and postorbital ridge (vs. absent).  Duttaphrynus brevirostris specifically also differs from  D. beddomii by its finger and toe tips lacking expanded discs (vs. with weakly-expanded discs), relatively reduced foot webbing, I1+-2II1+-3III2-3⅔IV3⅔-2V (vs. extensive, I1-1II1-1III1-2IV2-1V), and absence of prominently glandular warts or horny spinules on dorsum (vs. present); from  D. hololius , by its robust body (vs. dorso-ventrally flattened body), absence of mid-dorsal line (vs. present), sharp canthus rostralis (vs. rounded), snout rounded in lateral view (vs. acute), and more extensive foot webbing, I1+-2II1+-3III2-3⅔IV3⅔-2V (vs. rudimentary); from  D. stomaticus , by its shorter male snout-vent length, SVL 45-49 mm (vs. longer, SVL 54-69 mm), snout subovoid in dorsal view (vs. rounded), canthus rostralis sharp (vs. rounded), and relatively reduced foot webbing, I1+-2II1+-3III2-3⅔IV3⅔-2V (vs. more extensive: I1-1II1-2-III1-3IV3-1V); and from  D. peninsularis , by its canthus rostralis sharp (vs. rounded), snout length longer than eye diameter, SL/EL ratio 1.2-1.3 mm (vs. nearly equal), and relatively reduced foot webbing, I1+-2II1+-3III2-3⅔IV3⅔-2V (vs. more extensive: I1+-2II1+-3-III  1½-3IV3-1½ V). </p>
            <p>Phylogenetic relationships and genetic distances.</p>
            <p> Duttaphrynus brevirostris is a member of the  Duttaphrynus melanostictus group (Fig. 3), within which it is more closely related to  D. melanostictus ,  D. cf. microtympanum (  D. “sp” , Van Bocxlaer et al. 2009), and  D. parietalis (Fig. 3). All populations of  D. brevirostris exhibit intraspecific distances of 0-0.2% in 16S. The sequence divergence for  D. brevirostris from other members of the  Duttaphrynus melanostictus group was as follows: 2.1-3.3% from  D. melanostictus , 2.2-2.6% from  D. cf. microtympanum , 2.8-3.2% from  D. parietalis , 3.0-4.3% from  Duttaphrynus sp. 1, and 2.4-5.6% from  Duttaphrynus sp. 2 (Suppl. materal 1: Table S4). </p>
            <p>Distribution and natural history.</p>
            <p> Duttaphrynus brevirostris is endemic to the Western Ghats, where it currently is known only from the State of Karnataka. Here, we report this species from Hassan district (Sakleshpur taluk, encompassing the type locality Kempholey Ghat), Kodagu district (Bhagamandala), and Udupi district (Someshwara and Manipal). Furthermore, we confirm the following available DNA sequences for this species: Someshwara (FJ882786, Van Bocxlaer et al. 2009), specimen examined herein; Bajipe (AB530640) and Shirva (AB530642), specimen vouchers unavailable and reportedly released (Hasan et al. 2014); and another sample EU071759 from an unknown locality in India (Shouche and Ghate, unpublished GenBank data). Based on available evidence,  D. brevirostris is confirmed to occur in Malnad or Malenadu regions as well as coastal regions (districts of Mangalore and Udupi) of Karnataka State and, therefore, has a wider distribution than previously surmised (Fig. 4). </p>
            <p>Most individuals were located during night searches (between 17:00-21:00 hours) in secondary forests or open urban areas. Calling males, usually with yellow dorsal colouration, were observed in June, away from the bodies of water. Specimens found closer to water were generally greyish-brown. A cursory tadpole description was provided along with the original description (Rao 1937).</p>
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	https://treatment.plazi.org/id/8BEE4000238959A1861757426110D5E4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Bisht, Karan;Garg, Sonali;Sarmah, A. N. D. Akalabya;Sengupta, Saibal;Biju, S. D.	Bisht, Karan, Garg, Sonali, Sarmah, A. N. D. Akalabya, Sengupta, Saibal, Biju, S. D. (2021): Lost, forgotten, and overlooked: systematic reassessment of two lesser-known toad species (Anura, Bufonidae) from Peninsular India and another wide-ranging northern species. Zoosystematics and Evolution 97 (2): 451-470, DOI: http://dx.doi.org/10.3897/zse.97.61770, URL: http://dx.doi.org/10.3897/zse.97.61770
03CC36A23EDB5D288DE7DFAE477B4697.text	03CC36A23EDB5D288DE7DFAE477B4697.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Duttaphrynus peninsularis (Rao 1920) Bisht & Garg & Sarmah & Sengupta & Biju 2021	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Duttaphrynus peninsularis (Rao, 1920) comb. nov.</p>
            <p>Figs 1, 2, 3, 4, 5</p>
            <p>Original name and description.</p>
            <p> Bufo stomaticus peninsularis Rao, 1920. Rao, C. R. N. 1920. Some South Indian batrachians. "Journal of the Bombay Natural History Society" 27: 119-127. Holotype. ZSIC 19176, SVL 45.1 mm (designated by Chanda et al. 2001  “2000” ), from "Mavkote and Watekolle, Coorg," Karnataka State, India. Current status of specific name. Valid name, as  Duttaphrynus peninsularis (Rao, 1920), comb. nov. </p>
            <p>Material studied.</p>
            <p> Topotype. An adult male, SDBDU 6370 (SVL 50.8 mm), collected by S.  D. Biju, from Wattakolli, Karnataka State. Other referred specimens. Four adult males, SDBDU 4018 (SVL 51.8 mm), SDBDU 4019 (SVL 45.5 mm), SDBDU 4020 (SVL 49.5 mm), and SDBDU 4021 (SVL 46.5 mm), from Coimbatore, Tamil Nadu State. </p>
            <p>Reassessment and validation of taxonomic status.</p>
            <p> Rao (1920) described a new variety of  Bufo stomaticus from "Mavkote and Watekolle, Coorg" as "  Bufo stomaticus peninsularis var. nov." The original description mentioned two specimens ("Type and syntype in the Indian Museum") and subsequently Chanda et al. (2001  “2000” ) proposed ZSIC 19176 to be the holotype. Currently a single specimen is available in the ZSIC (Kolkata) collection (S.  D. B., personal observation). It is noteworthy that, prior to describing this taxon, Rao (1920) took an opinion from Boulenger (then Curator, British Museum Natural History, London), who was not in favour of separating this collection from  D. stomaticus . However, Rao being unconvinced mentioned "no doubt about their being racially distinct" in the original description and went on to formally describe  Bufo stomaticus peninsularis as a new variety of  D. stomaticus . This nomen was considered to be a synonym of  Bufo stomaticus (=  Duttaphrynus stomaticus ) by Daniel (1963), without any justification or comparison, other than considering the characters mentioned by Rao (1920) as variation, based on examination of  D. stomaticus specimens from Bombay. This action was followed by Dubois (1974) and Dutta (1997). In later years, regional anuran lists reported  Duttaphrynus stomaticus from Peninsular India based on earlier reports and photographs, without citing any voucher specimens (Hegde 2012; Ramachandra et al. 2012; Seshadri et al. 2012). Srinivasulu et al. (2013) identified the  “captioned-photographs” of Seshadri et al. (2013) and Hegde (2012) as belonging to  D. scaber , a species that is widely distributed in Peninsular India (Dutta 1997; Chanda 2002; Daniels 2005; Dinesh et al. 2009; Padhye et al. 2013). Srinivasulu et al.'s (2013) notes concerning the misidentifications of  D. scaber as  D. stomaticus (and not  D. peninsularis ) was by implication considered as a synonymisation action of  Bufo stomaticus peninsularis with  D. scaber by Frost (2021). </p>
            <p> In order to verify the above, we compared the type specimen and the original description of  Bufo stomaticus peninsularis Rao, 1920. Although the holotype (ZSIC 19176) was found to be in a severely damaged and dehydrated condition (Fig. 1), the head portion was relatively better preserved. Diagnostic morphological characters, such as absence of prominent cephalic ridges, weakly developed parotoid glands, distinct tympanum (about 63% of the eye), and the relatively smooth skin texture of the head and dorsum, match with the original description of  Bufo stomaticus peninsularis Rao, 1920. Additionally, Rao (1920) clearly stated six differences between his new variety and the typical form of  Bufo stomaticus from "Indian Museum nos., 16067, 16068, 17254 and 17274" (see the detailed comparison section), which we further re-examined to confirm distinctness of the two taxa. </p>
            <p> We examined specimens from two populations of  Duttaphrynus "  Duttaphrynus stomaticus ," sampled from different localities (including Wattakolli) in Peninsular India, which were found to be comparable to the original description and type specimen of  Bufo stomaticus peninsularis Rao, 1920 with respect to snout-vent length, absence of cephalic ridges, weakly developed parotoid glands, and relatively smooth skin. Based on re-examination of the holotype and assessment of newly-collected material, and molecular data, we conclude that  Bufo stomaticus peninsularis Rao, 1920 and  Bufo stomaticus Lütken , 1864 represent two distinct species, both individually diagnosable from other Indian congeners and each other. Hence, we formally resurrect  Bufo stomaticus peninsularis Rao, 1920, as a distinct species:  Duttaphrynus peninsularis (Rao, 1920), comb. nov. Furthermore, since the holotype is poorly preserved, we also provide a detailed redescription of this species, based on new topotypic material from Wattakolli, which matches the original description and the type. </p>
            <p>Description of topotype, SDBDU 6370</p>
            <p>(measurements in mm). A medium-sized, robust adult male (SVL 50.9); head of moderate size, wider (HW 18.0) than long (HL 14.0); snout truncate in dorsal and ventral view, rounded in lateral view, projecting beyond the mouth, its length (SL 5.8) nearly equal to horizontal diameter of eye (EL 5.7); loreal region acute with rounded canthus rostralis; distance between posterior borders of the eyes (IBE 13.9) 1.6 times the distance between the anterior borders (IFE 8.2); interorbital space about 1.4 times wider (IUE 6.2) than upper eyelid width (UEW 4.5); nostril oval without lateral flap of skin, closer to tip of snout (NS 1.7) than eye (EN 3.2); tympanum distinct (TYD 3.1), vertically oval, about 56.4% of eye diameter (EL 5.5), tympanum to eye distance (TYE 1.0); pineal ocellus absent; vomerine ridge and teeth absent; tongue small, oval, entire, median lingual projection absent; parotoid glands present, oval, flat, without spines and warts, slightly longer (PL 10.4) than wide (PW 5.5), distance between them (PD 6.2) more than the width.</p>
            <p>Forelimbs short; forearm length (FAL 11.5) longer than hand length (HAL 10.9); fingers rather thin, FLI longer than FLII, FLIII longest (5.6); relative length of fingers: II&lt;IV&lt;I&lt;III; tips of fingers rounded; subarticular tubercles prominent, single, all present; prepollex oval, distinct; single rounded prominent palmar tubercle; numerous supernumerary tubercles irregularly set on palm.</p>
            <p> Hind limbs relatively long and thin, thigh length (TL 19.7) longer than shank (SHL 17.8) and foot (FOL 18.4) length; relative length of toes: I&lt;II&lt;V&lt;III&lt;IV; tips of all toes rounded, without discs; webbing between toes present, small: I1+-2II1+-3-III  1½-3IV3-1½ V; dermal fringes present on all toes; subarticular tubercles rather weakly developed, oval; inner metatarsal tubercle present, prominent, its length (IMT 1.6) shorter than outer metatarsal tubercle (OMT 1.8); numerous weakly developed supernumerary tubercles set on foot. </p>
            <p>Skin. Dorsal and lateral surfaces of head and snout, and skin between eyes relatively smooth to sparsely granular; anterior and posterior parts of back with flat and smooth glandular projections; flanks glandular without horny spinules or warts; dorsal surfaces of thigh, shank, and tarsus with smooth glandular warts. Ventral surfaces of throat, chest, belly, and thighs glandular.</p>
            <p>Male secondary sexual character. Light brown granular projections on the lateral surfaces of fingers I, II, and III.</p>
            <p>Colour in preservation. Dorsum and limbs greyish-brown without any prominent markings; lateral surfaces of head, flank, and groin slightly lighter than dorsum; ventral surfaces (including limbs) greyish-white, throat with a faint light blue calling patch (Fig. 1). Colour in life: dorsum yellowish-brown with reddish patches; limbs yellowish brown; ventral surfaces white with a prominent bluish-yellow calling patch on throat (Fig. 2).</p>
            <p>Variation.</p>
            <p>Adult size range: male SVL 45-52 mm. Morphometric data from five adult males, including the described topotype, is given in Table 1. The dorsal colour is highly variable in life: SDBDU 4018: light brown with light grey patches, SDBDU 4019: light brown with reddish blotches, and SDBDU 4020: uniformly olive green.</p>
            <p>Comparisons.</p>
            <p> Duttaphrynus peninsularis differs from the Indian congeners:  D. chandai ,  D. himalayanus ,  D. kiphirensis ,  D. mamitensis ,  D. manipurensis ,  D. melanostictus ,  D. microtympanum ,  D. mizoramensis ,  D. nagalandensis ,  D. parietalis ,  D. silentvalleyensis ,  D. scaber ,  D. stuarti , and  D. wokhaensis , and species from other regions:  D. crocus (Myanmar),  D. kotagamai and  D. noellerti (Sri Lanka), and  D. totol (Indonesia), by the absence of conspicuous cephalic ridges (vs. present), absence of prominent or raised parotoid glands (vs. present), and dorsal skin without distinct glandular warts or horny spinules (vs. present in all species). Due to the lack of conspicuous cephalic ridges  D. peninsularis could be confused with four Indian species  D. beddomii ,  D. brevirostris ,  D. hololius , and  D. stomaticus . However, it differs from  D. beddomii in having a relatively larger tympanum (vs. smaller), finger and toe tips without discs (vs. with weakly developed discs), relatively reduced foot webbing, I1+-2II1+-3-III  1½-3IV3-1½ V (vs. extensive, I1-1II1-1III1-2IV2-1V), and absence of prominent glandular warts or horny spinules on dorsum (vs. present).  Duttaphrynus peninsularis differs from  D. hololius by its robust body (vs. dorso-ventrally flattened), absence of mid-dorsal line (vs. present), snout rounded in lateral view (vs. acute), tympanum smaller than eye diameter (vs. nearly equal), and more extensive webbing between toes, I1+-2II1+-3-III  1½-3IV3-1½ V (vs. rudimentary).  Duttaphrynus peninsularis differs from  D. stomaticus by its relatively shorter snout-vent length, male SVL 45-52 mm (vs. longer, male SVL 54-69 mm), its snout truncate in dorsal and ventral view (vs. rounded), snout longer than eye diameter (vs. nearly equal), dorsal skin granulation relatively smooth (vs. with prominent glandular warts), and relatively reduced foot webbing, I1+-2II1+-3-III  1½-3IV3-1½ V (vs. more, I1-1II1-2-III1-3IV3-1V). For comparisons to  D. brevirostris , see the respective comparison section. </p>
            <p> We quantitatively assessed the degree of morphometric differentiation of  Duttaphrynus peninsularis from the other two Indian members of the  Duttaphrynus stomaticus group (  D. hololius and  D. stomaticus ). An ordination of the first two principal components resulted in formation of three distinct clusters, what we consider to be three species (Fig. 5). The first two principal components (PC) accounted for 50.73% of the total variance, of which PC1 was able to explain 32.08%, and PC2 explained 18.65% of the variation in the dataset. Variables with the highest factor loadings for PC1 were HW, TYD, EL, IUE, and IN, while PC2 was highly loaded for UEW. The third and fourth principal components (PC3 and PC4) accounted for 9.37% and 9.07% of the total variance, respectively, taking the cumulative variance for the first four components to 69.17% (Suppl. materal 1: Table S5). The Box and whiskers plots of the five most significant characters recovered from PCA showed diagnostic differences between the three species (Fig. 5). Of the three species,  D. hololius was more distinct for all the studied characters, whereas  D. peninsularis and  D. stomaticus could be clearly delineated based on SVL, EL/SVL, TYD/SVL, and IN/SVL. </p>
            <p>Phylogenetic relationships and genetic distances.</p>
            <p> Duttaphrynus peninsularis is a member of the  Duttaphrynus stomaticus group (Fig. 3), within which it is more closely related to  D. stomaticus and  D. '  Duttaphrynus olivaceus ' than to  D. dhufarensis and  D. hololius . The studied populations of  D. peninsularis exhibit intraspecific distances of 0-0.4% in 16S. The sequence divergence of  D. peninsularis from other members of the  Duttaphrynus stomaticus group was as follows: 2.3-3.8% from  D. dhufarensis , 5.2-5.4% from  D. hololius , 1.3-2.6% from  D. stomaticus , and 1.0-1.5% from  D. '  Duttaphrynus olivaceus ' (Suppl. materal 1: Table S4). </p>
            <p>Distribution and natural history.</p>
            <p> Duttaphrynus peninsularis is currently known only from the Peninsular Indian States of Karnataka, Tamil Nadu, and Maharashtra. Genetically confirmed records are from Karnataka: Kodagu district (Wattakolli); Tamil Nadu: Coimbatore district (Coimbatore); and Maharashtra: Solapur district (Barshi and Solapur). We have also observed this species at Namakkal district (Kolli Malai) of Tamil Nadu. DNA sequences of this species were previously reported as  D. stomaticus (FJ882787, Van Bocxlaer et al. 2009). Another genetically identical sample from an unknown locality in India is currently available (EU071742, Shouche and Ghate, unpublished GenBank data). Given that this species currently has a disjunct distribution based on available genetically confirmed records, it is likely to be more widely distributed in the intervening regions of Peninsular India (Kerala, Tamil Nadu, and Karnataka, up to southern Maharashtra). Furthermore, its most closely related congener  D. stomaticus is frequently and widely reported in Peninsular India, which could be misidentifications of  D. peninsularis ; hence the identity of all '  D. stomaticus ' records from this region require further verification. Based on the present study, the geographical boundary between  D. peninsularis (southern species) and  D. stomaticus (northern species) could lie in the northern Western Ghats regions of Maharashtra state, where we have observed and genetically confirmed the presence of both these species (see Distribution and Natural History section of  D. stomaticus ). Further extensive sampling will be necessary to understand the patterns of population structure and delineate the ranges of these two species, using integrative approaches focusing on quantified ranges of phenotypic variation, traditional morphology, bioacoustics, ecological information, and phylogeny. </p>
            <p> Most individuals reported here were located during night searches (between 17:00-21:00 hours) largely in vegetated urban areas. The species were also found in secondary forest patches adjacent to human settlements. Ganesh et al. (2020) reported this species as  D. stomaticus from Tuticorin, Tamil Nadu. </p>
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	https://treatment.plazi.org/id/03CC36A23EDB5D288DE7DFAE477B4697	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Bisht, Karan;Garg, Sonali;Sarmah, A. N. D. Akalabya;Sengupta, Saibal;Biju, S. D.	Bisht, Karan, Garg, Sonali, Sarmah, A. N. D. Akalabya, Sengupta, Saibal, Biju, S. D. (2021): Lost, forgotten, and overlooked: systematic reassessment of two lesser-known toad species (Anura, Bufonidae) from Peninsular India and another wide-ranging northern species. Zoosystematics and Evolution 97 (2): 451-470, DOI: http://dx.doi.org/10.3897/zse.97.61770, URL: http://dx.doi.org/10.3897/zse.97.61770
304D1635ED2A5B81B320F58739B79EE7.text	304D1635ED2A5B81B320F58739B79EE7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Duttaphrynus stomaticus (Luetken 1864)	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> 
Duttaphrynus stomaticus (
Luetken
, 1864)
</p>
            <p>Figs 1, 2, 3, 4, 5</p>
            <p>Original name and description.</p>
            <p> Bufo stomaticus Lütken , 1864.  Lütken , C. F. 1864  “1863.” Nogle ny Krybyr og Padder. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i  Kjøbenhavn , Serie 2, 4: 292-311. Syntypes. Three adult females, ZMUC 131137 [ex 196], ZMUC 131365 [ex 198], and one unnumbered, from  “Assam;” two adult males, ZMUC 131136 [ex 195] and one unnumbered, from  “Assam;” and three subadults, ZMUC 131366 [ex 199] from  “Hoogly,” ZMUC 131363 [ex 193] from  “Calcutta,” and ZMUC 131364 [ex 194] from  “Calcutta.” Type locality.  “Assam,” India, based on two specimens used in the original description (  Lütken , 1864). Current status of specific name. Valid name, as  Duttaphrynus stomaticus (  Lütken , 1864). </p>
            <p>Material studied.</p>
            <p>  Syntypes:  Three adult females, ZMUC 131137 [ex 196] (SVL 60.9 mm), ZMUC 131365 [ex 198] (SVL 55.2 mm), and one unnumbered (SVL 61.4 mm), from  “Assam;” two adult males, ZMUC 131136 [ex 195] (SVL 55 mm) and one unnumbered (SVL 59.2 mm), from  “Assam;” and three subadults, ZMUC 131366 [ex 199] (SVL 26.4 mm) from  “Hoogly,” ZMUC 131363 [ex 193] (SVL 33.4 mm) from  “Calcutta” (Kolkata), and ZMUC 131364 [ex 194] (SVL 30.0 mm), from  “Culcutta” (Kolkata).  Other referred specimens: three adult males, SDBDU 2018.4109 (SVL 57.6 mm), SDBDU 2018.4110 (SVL 69.2 mm), and SDBDU 2018.4111 (SVL 55.1 mm), from  Sonitpur district ,  Assam State ; two adult males, SDBDU 2018.3717 (SVL 56.2 mm) and SDBDU 2018.3750 (SVL 54.2 mm), from  Dehradun ,  Uttarakhand State ; an adult female, SDBDU 2012.2172 (SVL 67.5 mm), from  Delhi ; an adult female, SDBDU 2012.2269 (SVL 68.7 mm), from  
Kaitha in 
Banka district ,  Bihar State ; an adult male, SDBDU 2012.2170 (SVL 51.0 mm), from Jaipur, Rajasthan State  . </p>
            <p> Taxonomic history of  Bufo stomaticus Lütken , 1864. </p>
            <p> In the original description,  Lütken (1864) mentioned that the Zoological museum, Copenhagen received six specimens of a toad from "Hr. Grosserer Westerman" (= Mr. Wholesales man Westermann) from  “ostindiske” (= East India). Subsequent researchers stated the type locality of this species to be 'East  India’ where it was later restricted to Assam (Boulenger 1891). Dutta (1997) stated that the type specimens are untraceable. We (SDB and SG) studied the types that are available at ZMUC, Copenhagen, and found a total of eight specimens (see 'Other material  studied’ ). According to the museum catalogue and bottle labels, all the adult animals are from  “Assam,” one juvenile from  “Hoogly,” and two juveniles from  “Culcutta” (Kolkata). All the specimens belong to the same species and the morphological characters were in agreement with the brief original description. Boulenger (1891) had mentioned after examining the syntypes that the exact locality from where these were procured is unknown and believed they originated from Assam or "they are perhaps from Bengal." However, while describing  Bufo stomaticus Lütken (1864) provided four measurements from two specimens, without mentioning the voucher numbers-"en Han" (one male) and "en Hun" (one female) "Fra Snudespidsen til Gattet" (= from snout to cloaca) 54 mm and 61 mm, respectively. Among the eight located syntypes, two similar-sized specimens were found bearing small tags on the hind limbs stating  ‘type’ . </p>
            <p> Based on the available information, it is apparent that only two specimens, ZMUC 131137 [ex 196] and ZMUC 131136 [ex 195], were used for  Lütken’s (1864) description of  Bufo stomaticus ; hence only these can be considered as potential syntypes. However, since the type series contains both adult and subadult specimens originating from different localities, it has led to confusion regarding the type locality and type status (Boulenger 1891). In order to clarify the taxonomic status of  B. stomaticus , we provide a detailed redescription for one potential syntype, ZMUC 131137 [ex 196], an adult female, SVL 60.9 mm, from  “Assam.” The below redescription, along with live photographs, interspecific comparisons, and enumeration of diagnostic characters, may be useful for differentiating this taxon from other known  Duttaphrynus species. We also provide additional information on new topotypic material, including live photographs, genetic data, inferred phylogenetic relationships, and extended geographical records, based on morphologically-characterised and genetically-confirmed records-all of which shows that  D. stomaticus (as understood here) is consistent with what is known of the name-bearing types. </p>
            <p>Description of syntype, ZMUC 131137 [ex 196]</p>
            <p>(measurements in mm). A medium-sized, robust adult female (SVL 60.9). Head of moderate size, wider (HW 22.7) than long (HL 17.8); snout rounded in lateral, dorsal, and ventral view, projecting beyond the mouth, its length (SL 6.8) longer to horizontal diameter of eye (EL 6.0); loreal region acute with rounded canthus rostralis; distance between posterior borders of the eyes (IBE 16.2) 1.8 times the distance between the anterior borders (IFE 9.2); interorbital space concave, 1.3 times wider (IUE 6.6) than upper eyelid width (UEW 5.0); nostril oval without lateral flap of skin, closer to tip of snout (NS 1.8) than to eye (EN 3.5); tympanum distinct (TYD 3.6), rounded, 58.1% of eye diameter (EL 6.2), tympanum to eye distance (TYE 1.6); pineal ocellus absent; vomerine ridge and teeth absent; tongue small, oval, entire, median lingual projection absent; parotoid glands present, oval, elongate, without spines and warts, longer (PL 13.9) than wide (PW 6.5) and distance between them (PD 10.0) wider than their width; cephalic ridges absent.</p>
            <p>Forelimbs short; forearm length (FAL 11.5) shorter than hand length (HAL 13.7); fingers rather thin, FLI longer to FLII, FLIII longest (7.1 mm); relative length of fingers: I&lt;II&lt;IV&lt;III; tips of fingers rounded; subarticular tubercles prominent, single, all present; prepollex oval, distinct; single rounded prominent palmar tubercle; numerous supernumerary tubercles irregularly set on palm.</p>
            <p>Hind limbs relatively long and thin, thigh length (TL 21.3) shorter than shank (SHL 21.8) and foot (FOL 22.6) length; relative length of toes: I&lt;II&lt;V&lt;III&lt;IV; tips of all toes rounded without discs; webbing between toes present, small: I1-1II1-2-III1-3IV3-1V; dermal fringes present on all toes; subarticular tubercles rather well-developed, oval; inner metatarsal tubercle present, prominent, its length (IMT 3.1) shorter than outer metatarsal tubercle (OMT 3.7); numerous weakly developed supernumerary tubercles set on foot.</p>
            <p>Skin. Dorsal surfaces of head sparsely granular; lateral surfaces of head shagreened with scattered tubercles; upper eyelids with glandular warts possessing horny spinules; anterior and posterior parts of back with glandular warts possessing horny spinules, larger warts towards posterior back; flanks glandular without warts or horny spinules; dorsal surfaces of thigh, shank, and tarsus glandular. Ventral surfaces of throat, chest, belly, and thighs with fine glandular projections without horny spinules or warts.</p>
            <p>Secondary sexual characters. Female (ZMUC 131137): ova white, pigmented on pole (diameter 0.8-1.0 mm, N = 20); Male (SDBDU 2018.4111): light brown granular projections on the lateral surfaces of fingers I, II, and III. Colour in preservation: dorsal surfaces of head and body uniformly fawn, some spines brown; dorsal surface of fore-and hind limbs light fawn; ventral surfaces of head, body, and limbs light grey (Fig. 1). Colour in life (based on other material studied): dorsum yellowish-brown, straw, light brown, or olive green, with or without grey or brown patches; and a pair of faint discontinuous dorsolateral lines; ventral surfaces greyish-white (Fig. 2).</p>
            <p>Variation.</p>
            <p>Adult size range: male SVL 54-69 mm, female SVL 60-72 mm. Morphometric data from five adult males, including the described syntype, is given in Table 1. Dorsal colouration varies from light grey or brown to olive green; the amount and degree of prominence of granulation on dorsal skin variable.</p>
            <p>Comparisons.</p>
            <p> Duttaphrynus stomaticus differs from the Indian species:  D. chandai ,  D. himalayanus ,  D. kiphirensis ,  D. mamitensis ,  D. manipurensis ,  D. melanostictus ,  D. microtympanum ,  D. mizoramensis ,  D. nagalandensis ,  D. parietalis ,  D. silentvalleyensis ,  D. scaber ,  D. stuarti , and  D. wokhaensis , and other species found outside:  D. crocus (Myanmar),  D. kotagamai and  D. noellerti (Sri Lanka), and  D. totol (Indonesia), by the absence of cephalic ridges, absence of prominent or raised parotoid glands, and absence of distinct glandular warts or horny spinules (vs. present in all species). Due to the absence of cephalic ridges  D. stomaticus could be confused with three Indian species  D. beddomii ,  D. hololius , and  D. peninsularis . However,  D. stomaticus differs from  D. beddomii in having a tympanum larger than eye diameter (vs. smaller), finger and toe tips lacking expanded discs (vs. with weakly-expanded discs), relatively reduced foot webbing, I1-1II1-2-III1-3IV3-1V (vs. more extensive, I1-1II1-1III1-2IV2-1V), and less prominent glandular warts or horny spinules on dorsum (vs. more prominent); from  D. hololius , in having a stout body (vs. flattened or dorso-ventrally compressed), absence of a prominent or broad mid-dorsal line (vs. present), snout rounded in lateral view (vs. acute), dorsum with relatively more prominent smooth or spinular warts (vs. less prominent and scattered smooth tubercles), and moderate foot webbing, I1-1II1-2-III1-3IV3-1V (vs. rudimentary). For comparisons to  D. brevirostris and  D. peninsularis , see the respective comparison sections of those species. </p>
            <p>Phylogenetic relationships and genetic distances.</p>
            <p> Duttaphrynus stomaticus is a member of the  Duttaphrynus stomaticus group (Fig. 3), within which it is more closely related to  D. '  Duttaphrynus olivaceus ' and  D. peninsularis than to  D. dhufarensis and  D. hololius . The studied populations of  D. stomaticus exhibit intraspecific distances of 0-0.4% in 16S. The sequence divergence of  D. stomaticus from other members of the  D. stomaticus group is as follows: 0.2-0.6% from  D. '  Duttaphrynus olivaceus ', 1.3-2.6% from  D. peninsularis , 1.5-3.0% from  D. dhufarensis , and 3.4-5.6% from  D. hololius (Suppl. materal 1: Table S4). </p>
            <p> Relationships within  Duttaphrynus stomaticus group. </p>
            <p> The close phylogenetic relationship of  Duttaphrynus stomaticus with  D. dhufarensis ,  D. hololius ,  D. olivaceus , and  D. peninsularis is well-supported (Van Bocxlaer et al. 2009; Portik and Papenfuss 2015; present study). Martin (1972) also discussed the absence of conspicuous cephalic ridges as a potential morphological synapomorphy for these species. Within this group, subsequently referred to as the  Duttaphrynus stomaticus group (Inger 1972; Dubois and Ohler 1999; Silva and Mendelson 1999; Van Bocxlaer et al. 2009), the taxonomic identity of  D. olivaceus has been questionable due to the lack of sufficient morphological distinctness (Dubois 1984; Balletto et al. 1985; Minton 1966) as well as shallow genetic divergence (Portik and Papenfuss 2015; present study). Eiselt and Schmidtler (1973) regarded  D. olivaceus as the subspecies of  D. stomaticus . However, subsequent workers treated  D. olivaceus as a distinct species closely related to  D. stomaticus with relatively weak and variable morphological diagnostic characters, such as differences in the size of parotoid glands, number of subarticular tubercles on finger III, and weakly or well-developed tibial gland and tarsal folds (Schmidtler and Schmidtler 1969; Khan 1987; Auffenberg and Rehman 1997). The available genetic data for  D. stomaticus and  D. olivaceus , along with new samples reported in this study for various  D. stomaticus populations from India (including topotypic sequences) show a shallow divergence of 0.2-0.6% between the two species (Fig. 3). </p>
            <p> Recently, Safaei-Mahroo and Ghaffari (2020) discussed the taxonomic status of  D. olivaceus (Frost 2021). This study also proposed a new genus name  Firouzophrynus Safaei-Mahroo &amp; Ghaffari, 2020 to accommodate a single species  Duttaphrynus olivaceus (Blanford 1874), which rendered the genus  Duttaphrynus paraphyletic (Frost 2021). Subsequently, based on phylogenetic evidence from selected taxa, Dubois et al. (2021) redelimited  Firouzophrynus as a genus, while also stating the possibility of considering it as a subgenus, to include members of the  Duttaphrynus stomaticus group as defined by Inger (1972) and Dubois and Ohler (1999). However, as noted by Frost (2021), there continues to be lack of clarity regarding the morphological and phylogenetic affinities of some other members of the group, which may have implications on the monophyly of  Firouzophrynus . The composition of  Duttaphrynus stomaticus species group and its phylogenetic position have been discussed by numerous studies (Inger 1972; Martin 1972; Maxson 1981; Van Bocxlaer et al. 2009; Portik and Papenfuss 2015). However, only five species (  D. stomaticus ,  D. dhufarensis ,  D. hololius ,  D. olivaceus , and  D. peninsularis ) currently are included in this group based on morphological (Inger 1972; Martin 1972; Dubois and Ohler 1999; present study) and phylogenetic analyses (Frost et al. 2006; Van Bocxlaer et al. 2009; Portik and Papenfuss 2015; this study). At least two other species from Indonesia,  D. valhallae and  D. sumatranus , that are known to lack cephalic ridges, a characteristic of the group (Inger 1972; Dubois and Ohler 1999), require further studies to establish their systematic relationships. Although we do not doubt that  Firouzophrynus could be recognised as a genus or subgenus, we currently consider the taxonomic status of this taxon uncertain, pending additional studies which may provide clarity, because of its cursory description and lack of a clear definition. Because it is beyond the scope of the present work to address this question, we have provisionally referred our focal taxa to the genus  Duttaphrynus , sensu lato, and make use of previously defined species-groups, which could easily be adopted to an alternate classification, as more evidence concerning the recognition of  Firouzophrynus becomes available. </p>
            <p>Distribution and natural history.</p>
            <p> Duttaphrynus stomaticus is one of the most widely-distributed species of the genus, occurring between elevations of sea-level to 2500 m asl in India (through Indo-Gangetic Plains, upper and lower Indus Valleys) and the neighbouring Bangladesh, Nepal, Pakistan (Balochistan), Afghanistan, and Iran (Suppl. materal 1: Table S1). This species is known to occur in varying climatic conditions and habitats, ranging from dry scrub forests, arid and semi-arid regions, hot and humid mixed forests, plains, and grasslands to drier and colder regions, montane woodlands and forests (Choudhury et al. 2001; Mehta 2005; Deuti et al. 2014; Safaei-Mahroo et al. 2015). Genetically confirmed records of this species exist from India, Afghanistan, and Pakistan (Suppl. materal 1: Table S3). In the present study, we specifically confirm the presence of  D. stomaticus in the Indian States of Assam, Bihar, Delhi, Punjab, Rajasthan, and Uttarakhand (Suppl. materal 1: Table S3) and also clarify the identity of some previously published DNA sequences from Peninsular India (Van Bocxlaer et al. 2009; Shouche and Ghate 2007, unpublished GenBank data) as belonging to  D. peninsularis . Hence, records of  D. stomaticus from Peninsular India (south of Maharashtra and possibly Odisha) are currently presumed to be doubtful and will require verification of all known populations (see  D. peninsularis for discussion). The reports of  D. stomaticus from Karnataka and Tamil Nadu States (Hegde 2012; Ramachandra et al. 2012; Seshadri et al. 2012; Ganesh et al. 2020) likely refer to  D. peninsularis . A report of  D. olivaceus from Gurgaon, India (Ray and Deuti 2008) is also questionable (Heydari and Rastegar-Pouyani 2010) and considered to represent  D. stomaticus based on our fresh collections from Delhi and surrounding North Indian regions. </p>
            <p> Duttaphrynus stomaticus is predominantly a nocturnal species. In this study, we found individuals of this species in urban, rural, and secondary forested areas during the breeding season (usually between May-August). Calling and breeding activities were observed in agricultural fields and temporary puddles in urban and rural landscapes, whereas inside secondary forests breeding was observed in shallow parts of flowing streams. </p>
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	https://treatment.plazi.org/id/304D1635ED2A5B81B320F58739B79EE7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Bisht, Karan;Garg, Sonali;Sarmah, A. N. D. Akalabya;Sengupta, Saibal;Biju, S. D.	Bisht, Karan, Garg, Sonali, Sarmah, A. N. D. Akalabya, Sengupta, Saibal, Biju, S. D. (2021): Lost, forgotten, and overlooked: systematic reassessment of two lesser-known toad species (Anura, Bufonidae) from Peninsular India and another wide-ranging northern species. Zoosystematics and Evolution 97 (2): 451-470, DOI: http://dx.doi.org/10.3897/zse.97.61770, URL: http://dx.doi.org/10.3897/zse.97.61770
