Cyrtodactylus mcdonaldi, Shea & Couper & Wilmer & Amey, 2011

Shea, Glenn, Couper, Patrick, Wilmer, Jessica Worthington & Amey, Andrew, 2011, Revision of the genus Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Australia 3146, Zootaxa 3146 (1), pp. 1-63 : 34-40

Cyrtodactylus mcdonaldi sp. nov.

Figs. 17–18

Holotype. QM J87075 , female, The Archways Cave section, Chillagoe–Mungana Caves National Park, 350 m above sea level (asl) (17º 05' 29" S 144º 23' 28" E) (P. Couper & K. McDonald, 28.ix.2008).

Paratypes. MCZ 152021, Laura River , Cook Hwy, 2 [28] mi. W, 32 mi. S Cooktown (15º 55' S 144º 50' E) ; QM J19327 , Mt Molloy (16º 41' S 145º 20' E) ; J30062 –63, 21.4 km E Chillagoe (17º 16' S 144º 37' E) ; J31272 , J48084 , Chillagoe (17º 09' S 144º 31' E) ; J45365 , Mt Mulligan (16º 53' S 144º 51' E) ; J60725 –26, Little Forks , Annan River (15º 49' S 145º 13' E) ; J61772 –73, nr Chillagoe township (17º 09' S 144º 31' E) ; J63482 , Windsor Tableland (16º 18' S 145º 05' E) ; J87047 –48, Ship Rock , Mt Poverty, Grey Range (15º 51' 47" S 145º 12' 19" E) ; J87070 , Mt Windsor National Park (16º 18' 06" S 145º 05' 25" E) ; J87071 , Donner Cave , Chillagoe–Mungana Caves National Park (17º 09' 57" S 144º 30' 57" E) ; J87072 –74, 87076–77, The Archways Cave section, Chillagoe–Mungana Caves National Park (17º 05' 29" S 144º 23' 28" E) ; J87078 , Granite Gorge , via Wakamin (17º 02' 50" S 145º 21' 15" E) ; J87083 , Royal Arch Tower section, Chillagoe–Mungana Caves National Park (17º 11' 06" S 144º 29' 57" E) ; J87084 , Royal Arch Tower section, Chillagoe–Mungana Caves National Park (17º 11' 00" S 144º 29' 51" E) ; J88027 –88028, Parrot Creek Falls , via Shiptons Flat (15º 48' 08" S 145º 15' 24" E) .

Diagnosis. A medium-sized Cyrtodactylus (SVL to 105 mm) with large, weakly to moderately projecting tubercles on antebrachium, strongly developed dorsal tubercles in 18–23 longitudinal rows at the midpoint of the trunk (axilla-groin interval), 28–41 ventral scale rows at the same level, a continuous series of 34–46 enlarged femoroprecloacal scales extending from one knee to the other, in males, pores are in three sections with 4–9 unpored scales between; mental with a posterior extension extending between postmentals; lips marbled or stippled with brown, dark dorsal bands on trunk usually three, with a narrow dark edge posteriorly but usually not anteriorly; pale interspaces between dark body bands usually unspotted, rarely with a few dark macules; basal tail bands only a little wider than pale interspaces.

Description. Size medium (males 69.5–95 mm, mean = 85.6 mm, sd = 8.19, n = 7; females 36–105 mm, mean = 90.1 mm, sd = 15.34, n = 19).

Head relatively long (HL/SVL 27.9–34.1%, mean = 29.9%; sd = 1.58, n = 25) and wide ( HW / HL 63.6 – 78.8 %, mean = 70.5%; sd = 3.62, n = 25), moderately depressed ( HD / HL 35.8 – 43.4 %, mean = 40.7%, sd = 1.95, n = 25), distinct from neck. Loreal region moderately inflated, canthus rostralis poorly defined. Interorbital region and top of snout concave, deepest and widest just anterior to level of rostral canthus of eye. Snout moderately long (SL/HL 28.4–41.0%, mean = 37.2%, sd = 2.19, n = 25; only one individual less than 35.1%); EN/HL 25.2–30.0%, mean = 27.6%, sd = 0.98, n = 25), much longer than eye diameter (SL/EYE 118.3–193.0%, mean = 167.8%, sd = 15.99, n = 25, only one less than 143.6%), and a little longer than eye-ear interval ( EE / HL 27.6 – 33.1 %, mean = 29.9%, sd = 1.42, n = 25). Eye large ( EYE / HL 19.2 –27.0%, mean = 22.3%, sd = 2.03, n = 25), pupil vertical with crenated margin, forming about 3–4 low lobes along each edge of pupil. Supraciliaries in a double row, large, frill-like, well-differentiated from adjacent more medial granules of the brow ridge, and largest anteriorly. Ear opening small ( EAR / HL 6.7 – 12.3 %, mean = 8.8%, sd = 1.21, n = 25), usually a little taller than long and slightly angled posterodorsally, but sometimes rounder. Rostral wider than high, height at centre less than that more laterally, dorsal part usually divided by a relatively straight median groove (groove with small side branch in QM J87078 , 'T'-shaped in QM J87072 , terminates as a small isolated scale within the rostral shield in QM J60726 and replaced by a small scale deeply penetrating the upper medial edge of the rostral in QM J87047 ) that extends approximately ½ the midline height of the scale, and fails to reach the oral margin. Usually two enlarged supranasals separated by usually a single, less enlarged internasal (supranasals in contact in QM J87076 and QM J87084 , and two internasals present in QM J87048 ). External nares circular, bordered by first supralabial, rostral, supranasal, nasal (extending into posterior part of nostril) and 1–3 smaller granular scales between nasal and first supralabial. Nares moderately separated ( IN / HL 10.9 – 15.6 %, mean = 12.5%, sd = 0.91, n = 25). Supralabials anteriorly large, distinct from adjacent loreal granules, 8–10 (mode = 8 (48.0%), mean = 8.7, sd = 0.75, n = 25) to level of mid-orbit, then inflecting dorsally and posteriorly, and becoming smaller, to gradually blend along rictal margin with adjacent small granules; supralabials separated from orbital margin by at least two rows of small granular scales at narrowest point. Mental wider than deep, with a strong median extension, a little to moderately narrower (50% of specimens) or equal to rostral, and bordered posteriorly by a single elongate pair of large postmentals ( Fig. 10B ). Infralabials anteriorly much larger than adjacent gular scales, becoming smaller posteriorly, 8–12 (mode = 10 (44.0%), mean = 9.8, sd = 0.82, n = 25). First infralabial with ⅔–¾ of ventral border contacting postmental, ¼ –⅓ by anteriormost enlarged subinfralabial (fully contacting postmental in QM J87072 and QM J87084 ). Subinfralabial scales anteriorly large, flattened, and polygonal, becoming smaller, more rounded and granular posteriorly and medially (towards gular area) .

Body moderately robust (AGL/SVL 33.5–47.1%, mean = 42.2%, sd = 2.82, n = 25), with low, but distinct, ventrolateral skin folds approximately marking the transition between the enlarged flattened ventral scalation and the smaller, more rounded, granular lateral scalation. Scales on dorsum of head, body and limbs small, juxtaposed, rounded granules, with interspersed much larger tubercles. Granular scales finest over parietal region of head, becoming coarser over body, then more polygonal and flatter on tail. On head dorsum, tubercles small and only slightly projecting, anteriorly commencing in the posterior interocular area, becoming larger, more projecting and with a more conical, slightly posteriorly-tilted apex over nape ( Fig. 11B ). Tubercles on body dorsum larger again ( Fig. 12B ), but with a more longitudinally ovoid base, often with a weak median keel, and relatively low on anterior body, but becoming much more projecting posteriorly on body, over sacrum and onto tail base, where they are markedly conical. Tubercles persist along tail, one to two whorls per segment, becoming lower and less differentiated until eventually losing their distinction by about the fifth dark band. Large tubercles on body dorsum separated by 2-4 smaller granular scales, those on head and nape more widely separated by several scales. Tubercles on body arranged in about 18–23 (mean = 20.5, sd = 1.36, n = 25) roughly longitudinal rows. Dorsum of brachium relatively homogenous, sub-imbricate to imbricate; antebrachium with more imbricate, larger scales distally and over manus, and with dispersed, larger tubercles ( Fig. 13B ). Dorsum of thigh and crus with small, juxtaposed granules and densely packed, large tubercles, only dorsum of pes with imbricate scales ( Fig. 13G ).

Laterally, tubercles commence over temporal region and in postinfralabial area, where they are noticeably larger ( Fig. 11G ) than those of the head dorsum, then along nape and body, where they are smaller and noticeably less protuberant than those dorsally, and along tail, commencing on tail base as prominent, protuberant, conical scales, then rapidly losing differentiation by second dark tail band ( Fig. 14B ).

Ventrally, gular scales small, rounded and juxtaposed, becoming larger, flat and more imbricate over body venter, from clavicular region. Ventral scales at midbody, between ventrolateral skin folds 28–41 (mean = 34.4, sd = 3.74, n = 26). Ventral scales on brachium and antebrachium like gular scales. On ventral surface of thighs, but not on crus or in precloacal region, an abrupt junction between enlarged imbricate scales and much smaller scales posteriorly, enlarged scales 37–45 between distal extent on each thigh (mean = 41.7, sd = 2.37, n = 26). Ventral scales of tail base like those of body, most of tail venter with a single median series of very broad scales about four times the width of adjacent ventrolateral scales.

Precloacal and femoral pores present in males, separated by unpored scales into three patches. Distal femoral pores 8–11 (mean = 8.9, sd = 0.67, n = 14), separated by 4–9 unpored scales (mean = 6.7, sd = 1.32, n = 14) from the 13–16 (mean = 13.9, sd = 1.46, n = 7) proximal femoral + precloacal pores. Unilaterally on three individuals, a single pored scale located within the unpored series, towards its distal end.

Pores best developed in precloacal region where they are deep and transversely oriented, becoming much shallower, smaller and rounder distally under thigh. No pubic groove. About three large blunt-tipped postcloacal spurs on ventrolateral surface of tail base, more projecting in adult males than females or juveniles.

Forelimbs and hindlimbs well-developed (FLL/SVL 14.0–16.9%, mean = 15.3%, sd = 0.88, n = 25; HLL/SVL 16.5–19.2%, mean = 17.9%, sd = 0.72, n = 25). Digits well-developed, reflected dorsally at proximal interphalangeal joint, and all bearing robust, strongly curved claws sheathed at the base by two scales. Subdigital lamellae expanded basally, beginning on pes over distal part of metatarsals and ending at point of reflection of toes, lamellae distal to this point not expanded. Lamellae under first toe 5–9 expanded (mean = 7.1, sd = 0.95, mode = 7 (48.0%)) + 8–11 narrow (mean = 9.0, sd = 0.73, mode 9 (60.0%)), total 13–18 (mean = 16.1, sd = 1.17, mode = 16 (48.0%), n = 25). Lamellae under second toe 7–10 expanded (mean = 8.5, sd = 0.82, mode = 9 (44.0%)) + 8–11 narrow (mean = 9.8, sd = 0.78, mode = 10 (48.0%)), total 16–20 (mean = 18.2, sd = 0.97, mode = 18 (40.0%), n = 25). Lamellae under third toe 7–12 expanded (mean = 9.5, sd = 1.12, mode = 9 (36.0%)) + 9–12 narrow (mean = 10.8, sd = 0.90, mode = 11 (44.0%)), total 17–23 (mean = 20.3, sd = 1.38, mode = 21 (48.0%), n = 25). Lamellae under fourth toe 8–13 expanded (mean = 11.1, sd = 1.50, mode = 12 (48.0%)) + 10–13 narrow (mean = 11.0, sd = 0.87, mode = 11 (52.0%)), total 19–25 (mean = 22.1, sd = 1.41, mode = 23 (36.0%), n = 25). Lamellae under fifth toe 7–10 expanded (mean = 8.6, sd = 0.76, mode = 8 (44.0%)) + 10–13 narrow (mean = 11.4, sd = 0.96, mode = 12 (40.0%)), total 18–22 (mean = 20.0, sd = 1.06, mode = 20 (40.0%), n = 25). Relative lengths of digits on manus I<II<V<III<IV; on pes I<II<III=V<IV. Very slight traces of webbing between bases of fingers; weak webbing between bases of toes 2–3, 3–4 and 4–5.

Tail a little longer than body (TL/SVL 117.0–128.7%, mean = 123.6%, sd = 4.40, n = 5), narrow at base (TW/ SVL 5.8–11.1%, mean = 7.8%, sd = 1.24, n = 25) and tapering evenly to a conical tip. Tail segments externally identifiable by straight scale junctions, segments about 7–9 scales long when counted to include tubercles. Cloacal sacs present in both sexes, larger in males, external orifices just posterior to vent, laterally.

Colour in preservative. Dorsal pale ground colour fawn. Head dorsum ( Fig. 11B ) without brown mottling and bordered posterolaterally by a fine pale edge. Pale nape zone bordered posteriorly by a U-shaped mid to dark brown chevron on nape, widest vertebrally, and extending anteriorly over temporal region to eye, then visible as a narrowing, increasingly diffuse streak over the lores to the nostril. Second broad dark transverse dorsal band over shoulders, lateral margins extending more narrowly anteroventrally in front of forelimbs. Three dark bands over trunk, extending lateroventrally with even width, but dissipating over flanks. A dark band over hips in 54% of specimens (n = 26), a pale band in 46%. Tail with dark bands over most of length which become paler and less defined near the tip. When they can be counted to the distal end of tail, dark tail bands 10–14 (mean = 11.7, sd = 1.30, n = 12). On nape and body, dark bands usually somewhat wider than pale interspaces, and with abrupt straight edges; pigmentation darkest along the posterior margin but sometimes a darker edging is also evident along the anterior edge. Bands on tail ( Fig. 14B ) of similar width to body bands, and usually significantly wider than pale interspaces, but darker and more solidly dark than those of body. Pale interspaces on body generally clean.

Upper and lower lips ( Fig. 11G ) cream, finely stippled with brown or with brown mottling. Dorsum of forelimbs and hindlimbs usually with little indication of pattern, although obscure bars are sometimes present on the thighs.

Entire ventral surface usually immaculate cream, including the underside of the tail. Some specimens with diffuse brown mottling which is strongest in the gular region.

Description of holotype. The holotype of C. mcdonaldi is a mature-sized female, with the following character states of those variable for the taxon: SVL 100 mm, AGL 45 mm, TL 125 mm, TW 8.0 mm, HL 28.0 mm, HW 21.1 mm, HD 11.7 mm, IN 3.4 mm, SL 10.6 mm, EN 7.9 mm, EYE 5.7 mm, EE 9.0 mm, EAR 2.4 mm, FLL 15.3 mm, HLL 18.4 mm, lamellae below digits I–V 8+9, 10+9, 10+11, 12+11, 9+10 respectively, supralabials 10, infralabials 10, rows of dorsal tubercles 20, transventral rows 34, femoroprecloacal scales 43, dark band across hips present and dark tail bands 12.

Etymology. Named after Keith R. McDonald (b: 1950), Principal Senior Technical Officer, Threatened Species Unit, Queensland Department of Environment and Resource Management, and an Honorary Research Associate of the Queensland Museum, resident of Atherton, and discoverer of numerous north Queensland reptile and amphibian species. It was Keith who first alerted PC that Australian ' C. louisiadensis ' had an interesting, fragmented distribution and warranted further investigation. Keith was instrumental in collecting many of the initial tissue samples on which this revision is based.

Distribution. From Parrot Creek Falls, near Shiptons Flat, in the north, south to Chillagoe and 21.4 km E Chillagoe ( Fig. 15 ).

Conservation status. Using the IUCN criteria, this species most closely fits the Least Concern category (LC), in that it has relatively large distribution with little evidence of continuing declines or fluctuations and with extensive parts of its distribution in protected areas.

Comparison with other species ( Table 6). Cyrtodactylus mcdonaldi is geographically proximate to C. tuberculatus with the two closest localities for each species (Mt Leswell C. tuberculatus and Parrot Creek Falls C. mcdonaldi ) being separated by only 3 km. Cyrtodactylus mcdonaldi is also a sister taxon of C. tuberculatus with an average sequence divergence of 11.07% ( Table 2). It may be differentiated morphologically from C. tuberculatus by males having the distal femoral pores separated from the proximal femoroprecloacal pores by 4–9 unpored scales. It is further differentiated from C. tuberculatus by its smaller size (SVL males 69.5–100 vs 96–116.5 mm; females 88.5–105 vs 46–119 mm), less tuberculate skin (most obvious in the postlabial area, which is smooth vs usually tuberculate in C. tuberculatus , and on the antebrachium, which has protruding tubercles only on the largest individuals, but flat tubercles often present vs strongly projecting tubercles in all adult C. tuberculatus ). The colour pattern is generally 'cleaner' than in C. tuberculatus , with the crown of the head only with grey smudges (vs usually strongly mottled), the labial areas less strongly mottled, the postlabial area usually immaculately pale (vs mottled), the pale interspaces between dark body bands lacking dark macules (vs usually with some dark spots), and the antebrachium nearly plain (vs usually more mottled). The dark body bands usually have only a prominently dark posterior margin (both prominent dark anterior and posterior edges in C. tuberculatus ). A dark band over the hips is often lacking in C. mcdonaldi , but consistently present in C. tuberculatus .

Cyrtodactylus mcdonaldi differs from C. klugei and C. robustus in having many fewer femoroprecloacal pores (29–35 vs 66 or more; Kraus 2008) in males. The difference in number of pores is mirrored by the number of enlarged scales bearing them, which are countable in both males and females (37–45 vs 66 or more), and by the number of transventral scales (28–41 vs 41–54). It differs from both in having the femoroprecloacal pores broken into three series. Cyrtodactylus mcdonaldi is much more strongly tuberculate than C. klugei . It is further distinguished from C. klugei in having three dark bands across the trunk (vs usually two). It is further differentiated from C. robustus , with which it shares prominent tubercles, in its lesser size (SVL to 105 mm vs 161 mm; Kraus 2008), presence of a dark band across the hips (vs usually lacking), and marbled lips (vs pale lips). It also lacks the bright orange cloaca of C. robustus .

For comparisons with C. hoskini , C. adorus and C. pronarus , see the descriptions of those species.

Natural history. At Chillagoe, this species is found around limestone karsts, associated with poorly developed vine thickets, in open savannah woodland ( Fig. 16B ). A specimen from the Mareeba area was collected from an isolated granite outcrop, in granite hill country adjacent to a drainage line, in open forest with a grassy understory. Specimens from Mt Windsor National Park came from a road cutting in granite hills, surrounded by eucalypt-dominated woodland. The Parrot Creek (Shiptons Flat) site is comprised of exposed granite along a drainage line in wet sclerophyll forest. Genetic samples from Mt Poverty were collected from animals living on granitic outcrops in tall eucalypt forest (Keith McDonald, pers. comm.). Cyrtodactylus (not sampled but presumably C. mcdonaldi ) have been found in the Bloomfield area on trees in open woodland, where few rock outcrops occur (Lewis Roberts, pers. obs.). They are also known to colonise houses in this area and at Shiptons Flat (Lewis Roberts, pers. obs.).

An individual, presumably of this species, from Gap Creek via Ayton, near the Bloomfield River, was observed to catch and eat an adult gecko, Gehyra dubia (reported as Gehyra australis ) on the wall of a house ( Naylor 2000). Stomach contents of five individuals ( J19327 , J30062 –63, J45365 , J48084 ) were reported by Covacevich et al. (1996; as Cyrtodactylus louisiadensis ). Seven food items were reported: the spider Yiinthi chillagoe (Heteropodidae) , the scorpion Liocheles sp. (Ischnuridae) , two roaches ( Laxta sp. , Blaberidae ; Methana sp. , Blattidae ), two beetles ( Tenebrionidae and Curculionidae ), and a beetle larva, together with an unidentified nematode (possibly parasitic).

Figures

FIGURE 10. Chin shields of Australian Cyrtodactylus species. A. C. tuberculatus (QM J87009), B. C. mcdonaldi (QM J87075), C. C. hoskini (QM J86950), D. C. adorus (QM J86979), E. C. pronarus (QM J86900), F. C. pronarus (QM J86909).

FIGURE 11. Dorsal (A-E) and lateral (F-J) aspects of head of Australian Cyrtodactylus species. A,F. C. tuberculatus (QM J87009), B,G. C. mcdonaldi (QM J30062), C,H. C. hoskini (QM J86950), D, I. C. adorus (QM J86979), E,J. C. pronarus (QM J86900).

FIGURE 12. Dorsal aspect of trunk of Australian Cyrtodactylus species. A. C. tuberculatus (QM J87189), B. C. mcdonaldi (QM J87075), C. C. hoskini (QM J86950), D. C. adorus (QM J86979), E. C. pronarus (QM J86909).

FIGURE 13. Tubercles on front (A-E) and hind (F-J) limbs of Australian Cyrtodactylus species. A,F. C. tuberculatus (QM J87189), B,G. C. mcdonaldi (QM J87075), C,H. C. hoskini (QM J86950), D,I. C. adorus (QM J86979), E,J. C. pronarus (QM J86900).

FIGURE 14. Tubercles and colour pattern on tail base dorsum of Australian Cyrtodactylus species. A. C. tuberculatus (QM J87189), B. C. mcdonaldi (QM J30062), C. C. hoskini (QM J86950), D. C. adorus (QM J86979), E. C. pronarus (QM J86900).

FIGURE 15. Distribution of Cyrtodactylus tuberculatus (inverted triangles) and C. mcdonaldi (upright triangles). Open symbol indicates photographic record only.

FIGURE 16. Habitats of A. Cyrtodactylus tuberculatus at Black Mountain, Qld (Photo: A. Amey) and B. C. mcdonaldi at Chillagoe, Qld (Photo: P. Couper).

FIGURE 17. Holotype of Cyrtodactylus mcdonaldi (QM J87075).

FIGURE 18. Cyrtodactylus mcdonaldi from Chillagoe, Qld, in life (Photo: S. Wilson).