Bronchocela jubata Duméril and Bibron, 1837

Amarasinghe, A. A. Thasun, Ineich, Ivan, Riyanto, Awal, Hallermann, Jakob, Andayani, Noviar, Abinawanto, A. & Supriatna, Jatna, 2022, Taxonomy and distribution of a common arboreal lizard, Bronchocela jubata Duméril & Bibron, 1837 (Reptilia: Agamidae), with designation of its lectotype from Java, Indonesia, Zootaxa 5150 (1), pp. 65-82 : 69-75

Bronchocela jubata Duméril and Bibron, 1837

( Figs. 4 , 5 ; Tables 2 , 3 )

Bronchocela jubata Duméril & Bibron, 1837

Bronchocele (sic) intermedia Berthold, 1842 (not Calotes intermedia Peters & Doria, 1878 )

Calotes intermedius — Berthold 1846

Calotes jubatus — Boulenger 1885, de Rooij 1915, Smith 1935

Bronchocela jubata — Moody 1980, Manthey & Schuster 1992, Hallermann 2005, Manthey 2008, Amarasinghe et al. 2022 Lectotype (designated herein). MNHN-RA 2542, adult male (SVL 130 mm), collected from Java, Indonesia by Jean-René Constant Quoy and Joseph-Paul Gaimard in 1828. Note: The original syntypes were composed of four specimens: (i) MNHN-RA 2541 and (ii) MNHN-RA 2541A from Java, Indonesia (donated from the Leyden Museum) labelled as Calotes gutturosus on the original tag of Leyden, currently lost; (iii) MNHN-RA 2542 from Java, Indonesia (collected by Quoy and Gaimard); and (iv) MNHN-RA 2543 from Pondichéry, India (collected by Leschenault). Only the latter two syntypes are present at MNHN, and here we designate the one from Java as the lectotype. For the justification for lectotype designation see discussion.

Diagnosis. A species of Bronchocela inhabiting the Greater Sunda Islands (Sumatra, Java, Borneo, and Bali), characterized as follows: morphologically most similar to its allopatric congener on Sumatra Island, B. hayeki (see Amarasinghe et al. 2022) and sympatric congener, B. cristatella , but differs by having a comparatively larger gular sac (smaller in B. hayeki and B. cristatella ), ventral scales arranged in 10–12 non-enlarged rows (vs. 8–10 enlarged rows in B. hayeki and 10–14 slightly-enlarged rows in B. cristatella ), lower number of mid body scale rows, 33–59 (vs. 64–75 in B. hayeki and 50–106 in B. cristatella ), 1 or 2 upper dorsal scale rows directed upward (vs. 5–7 rows in B. hayeki and 4–10 in B. cristatella ), well-developed nuchal crest (weakly-developed in B. cristatella ) with crescent-shaped scales longer than ED (vs. lanceolate and shorter than ED in B. cristatella ), enlarged scales on temporal region (vs. absent in B. hayeki ), 3 rd finger shorter than fourth (vs. longer in B. hayeki ), the orbital area and the tympanum mostly pale (vs. mostly black in B. hayeki ), and tail colouration banded (vs. uniform in B. hayeki ).

In addition, Bronchocela jubata is distinguished from other congeners by having the following combination of characters: adults reach a maximum SVL of 141.0 mm in males and 142.0 in females, 9–11 supralabials, 10–12 nuchal crest scales to the level of axilla, 56–73 ventrals, 30–37 lamellae on fourth toe, third finger shorter than the fourth; large lateral scales directed downward anteriorly and straight backwards posteriorly, 1 or 2 upper dorsal scale rows on the lateral body directed upward along the body, mid gular scales enlarged, abdominal scales acuminated and enlarged compared to pectoral, keeled temporal scales with some enlarged scales and 6 or 7 rows between orbit and tympanum, tympanum more than half the size of orbit.

Description of lectotype. MNHN-RA 2542, an adult male, SVL 130 mm. Head moderately large, elongate, HL 27.3% of SVL, narrow, subtriangular in dorsal and ventral aspects, HW 52.2% of HL; distinct from neck; snout elongate, snout length greater than eye diameter, ED 48.8% of ES; interorbital distance broad; eye large, pupil rounded; diameter of eyes slightly shorter than eye-tympanum distance, ED 98.7% of TYE; ear opening shallow, its greatest diameter dorsolaterally, tympanum smaller than orbit, nearly 60% of orbit diameter; tympanum surrounded by keeled scales; several temporal scales enlarged, keeled, juxtaposed, six scale rows between orbit and tympanum; forehead concave; scales on interorbital and supercilium area keeled; scales on snout keeled, larger in size than those of occipital region; a well-developed nuchal crest continuing dorsally as a dorso-nuchal crest; dorsal crest rudimentary, consisting of 12 scales up to the level of axilla, no crest on the tail; rostral scale width greater than its height, ventro-posteriorly in contact with first supralabial, contacting posteriorly five more or less equal-sized postrostral scales; around nostrils on each side two supranasals, three postnasals, a single prenasal, and two subnasals, which separate the nasal from the supralabials; nostrils round located middle of the undivided nasal plate; canthus rostralis and supraciliary edges sharp; five canthal scales between supranasal and anterior margin of orbit; no distinct parietal plate; mental subtriangular, flat posteriorly, shorter than wide, posterior-laterally in contact with two enlarged postmentals separated by a smaller scale; each postmental pair bordered posteriorly by four smooth scales, including the medial scale, but exclusive of infralabials; chin scales keeled; gular pouch present, midgular scales enlarged; throat scales and midgular scales keeled, mucronate, and imbricate; three scale rows separate orbit from supralabials; supralabials eleven (ninth in midorbit position); infralabials eleven, decreasing in size toward gape.

Body slender; lateral body scales large, equal, strongly keeled and imbricate; scales on lateral body slightly smaller than on the venter at same level, directed backward and downward anteriorly and directed straight backward posteriorly; lateral body scales on the posterior body slightly larger than the anterior body scales; 1 or 2 upper dorsal scale rows directed backward and upward along the body; 37 scales around the midbody; pectoral scales and abdominal scales keeled, acuminated, imbricate and keels forming regular and parallel continuous ventral ridges; abdominal scales larger than pectoral scales; 10–12 rows enlarged ventrally, without clear margin with the lateral scales; 64 ventrals.

Forelimbs moderately short; no oblique fold (pit) present on shoulders, but shoulder scales keeled and smaller; dorsal scales on fore- and hind limbs keeled, enlarged, imbricate and mucronate; ventral scales on upper arm and lower arm keeled, imbricate, and mucronate; hind limbs relatively longer than forelimbs; scales on ventral surface of thigh keeled, enlarged, imbricate and mucronate; tibia comparatively longer than femur; keels on tibia forming a series of continuous parallel ridges; digits elongate, slender; relative length of digits (fingers) 4> 3> 2> 5> 1; (toes) 4> 3> 5> 2> 1; all bearing slightly recurved claws, claws are sharp and elongate; subdigital lamellae entire, bicarinate, and regular, 30 (left) subdigital lamellae on toe IV.

Tail elongated and complete, 447 mm. Ventral scales on tail base keeled and imbricate, smaller in size than on dorsal tail; dorsal scales on tail enlarged, imbricate, keeled, mucronate, and keels forming continuous parallel ridges; tail with subcaudals on median row not enlarged, subequal, imbricate, keeled, and mucronate.

Colouration.–– In preservative, colours faded and dorsum pale cream due to colour having been bleached.

In life, based on field observations, dorsum bright luminous green to dark olive green; few sky blue markings on the lateral body; lateral head lemon green; nuchal crest scales blonde yellow, green or cream, dorsal head greyish or brownish green; ventral head bluish green or pale green; orbit, labial band including the tympanum same as body colours or sometimes pale or whitish; dorsal crest scales luminous green or same colour as body; knee, elbow, wrist, and heel darker; dorsal fingers and toes, posterior 2/3 rd of the tail greenish brown; ventral body, limbs, anterior tail, and mid gular lighter luminous green; ventral digits light brown. In some populations, especially in males, prominent white patch below the tympanum (sometimes this patch may extend to the lips as a labial band), lateral neck (below the nuchal crest), and the lateral sky blue markings may be visible as whitish stripes across lateral body (similar to Calotes calotes in Sri Lanka).

Dentition. Based on our examination of three B. jubata specimens (MZB 2963, 3791, and 3898) from Java, all have two premaxillary teeth, 13 maxillary teeth, and 15 dentary teeth.

Hemipenes. Based on MZB 6639 ( Fig. 6 ), the hemipenis of B. jubata is well developed, single, width of the organ greater than its length; hemipenial lobes slightly divided for approximately 10% of its length; apex divided into four segments, two short dorsal segments and two long ventral segments by lateral and medial sulcus; calyculate ornamentation present on each lobe; thick-walled smooth calyces forming deep oval pits; apical calyces smaller than ventral and dorsal calyces; sulcus spermaticus converged and narrowly open at apex, proximal half deep, distal half shallow; sulcal lips smooth; a fleshy cardioid structure at the base of the ventral sulcus.

Habitat, natural history, and distribution. B. jubata is usually found in areas of open canopy in primary forests (mostly forest edge) or undisturbed secondary forests, forests ecotones, other vegetation (e.g. coffee and pepper plantations) and in well-maintained home gardens. This could be an artefact since the species will be more visible in such areas and probably more difficult to observe in true dense forest where it also occurs. They usually forage on tree trunks and branches at 1.5 to 4 meters above the ground during the daytime, especially when basking, mostly around 0900 hr. At night, the adults prefer the highest branches of trees to sleep, mostly in open canopy areas, while juveniles prefer tiny branches of shrubs. Puruhita (2014) observed insects of the orders Hymenoptera, Lepidoptera, Odonata, Hemiptera, Orthoptera, and Coleoptera in the gut contents of B. jubata populations in Central Java which highlights its insectivorous feeding habit.

This species is mostly distributed in the western parts of Java Island and southern parts of Sumatra Island (most frequently in Lampung Province). It is comparatively rare on the eastern part of Java and Bali Island. It seems extremely rare on Borneo Island and the only occurrence available to us was a couple of museum specimens collected from eastern Kalimantan. As we have noticed it seems that the abundance of B. jubata is correlated with the abundance of its sympatric congener B. cristatella . B. jubata is usually distributed in higher abundance when sympatric in areas where B. cristatella shows low abundance, probably in order to reduce the interspecific competition.

In addition, the species is sympatric with several other arboreal agamids such as Gonocephalus chamaeleontinus , and Pseudocalotes tympanistriga . We always found several B. jubata close together at elevations below 1,300 m above sea level. Most of the individuals were observed at 300–800 m elevations. The reports from the Malay Peninsula, northern parts of Sumatra, the Philippines, and Wallacea are doubtful and we remove them from its distribution range—see discussion.

Conservation status. Distribution of this species is scattered ( Fig. 1 ) due to forest fragmentation. Forest habitat fragments are further threatened by encroaching agricultural and industrial lands on Java Island. The application of the IUCN Red List criteria ( IUCN Standards & Petitions Subcommittee 2019) with our updated distribution data (excluding the single specimen reported from Borneo, ZMH R06163) shows that B. jubata is restricted to an area of occupancy (AOO) of 2752 km 2 and recorded from nearly 200 localities within 169,604 km 2 extent of occurrence (EOO).Although the species shows scattered distribution in severely fragmented forests patches in highly populated islands, given its wider area of occupancy, B. jubata should be considered as a Least Concern (LC) species.

Figures

FIGURE 1. Current distribution map showing the collection/observation localities of Bronchocela jubata (red) and its sympatric congener, B. cristatella sensu lato (blue) on Java and Bali Islands; sympatric localities filled partially; for the distribution of B. jubata on Sumatra Island, see Amarasinghe et al. 2022.

FIGURE 4. Bronchocela jubata lectotype (MNHN-RA 2542, adult male) from Java: (A) lateral view of the full body, (B) dorsal head, (C) lateral head, (D) ventral head.

FIGURE 5. Bronchocela jubata (A) in life (adult male, not collected) from Cibinong near Bogor, West Java.

FIGURE 6. Sulcate view of the left hemipenis of Bronchocela jubata (MZB 6639) from Sukamantri, Bogor, West Java.

Tables

TABLE 2. Some morphometric ratios and meristic characters of Bronchocela jubata and its sympatric congener, B. cristatella in Java, Indonesia (for accession data see Appendix).

Character		Bronchocela in Java ( Indonesia)
		B. jubata	B. cristatella
	Male (n =54)	Female (n =44)	Male (n =10)	Female (n =8)
AG/SVL%	49.1–52.8	47.9–49.8	48.0–49.6	50.7–52.3
HL/SVL%	28.7–28.9	26.5–28.5	28.9–30.9	27.3–28.4
HW/HL%	52.2–55.1	53.5–54.1	53.4–54.2	51.7–52.5
EN/HL%	21.3–23.2	21.0–23.1	20.2–20.3	20.4–22.2
ES/HL%	37.0–38.3	37.7–40.3	38.0–39.4	43.3–43.8
ED/HL%	28.6–31.6	28.4–31.6	31.8–32.2	37.3–38.7
TYE/HL%	19.3–22.7	17.5–20.7	19.0–19.9	19.1–19.4
LAL/SVL%	20.3–20.9	19.4–19.8	21.8–22.7	19.7–19.9
PLM/SVL%	21.8–22.4	20.2–20.9	20.2–20.3	19.9–20.4
TBL/SVL%	28.9–29.3	26.8–28.2	30.7–31.9	28.9–30.0
FOL/SVL%	36.2–41.3	34.0–38.9	38.9–41.7	37.3–37.4
TAL/SVL	2.4–3.4 (n =36) 3.3–3.4 (n =34)		3.1–3.6 (n =9)	3.4–3.5 (n =6)
supralabials		9–11		9, 10
infralabials		9–11		9, 10
scales from eye to tympanum		6, 7		8–10
nuchal crest scales		10–14		10–15
subdigital lamellae on toe IV		30–37		31–34
midbody scale rows	35–55	33–49	64–80	61–72
ventrals	56–73	56–70	63–86	60–75