Paroedura rennerae, Miralles & Bruy & Crottini & Rakotoarison & Ratsoavina & Scherz & Schmidt & Köhler & Glaw & Vences, 2021

Miralles, Aurelien, Bruy, Teddy, Crottini, Angelica, Rakotoarison, Andolalao, Ratsoavina, Fanomezana M., Scherz, Mark D., Schmidt, Robin, Koehler, Joern, Glaw, Frank & Vences, Miguel, 2021, Completing a taxonomic puzzle: integrative review of geckos of the Paroedura bastardi species complex (Squamata, Gekkonidae), Vertebrate Zoology 71, pp. 27-48 : 27

Paroedura rennerae sp. nov.

Figs 4 , 7 , 8 , 9

Remarks.

This species was previously named P. sp. aff. bastardi Ca01 “Marofandilia/Miandrivazo” by Cocca et al. (2018) and Paroedura bastardi by Aprea et al. (2013) and Köhler et al. (2019; partim).

Holotype.

ZSM 849/2010 (ZCMV 12740), adult female, from Kirindy reserve CNFEREF, Camp de base, 20.0674°S, 44.6569°E, ca. 55 m above sea level, Atsimo-Andrefana Region, western Madagascar, collected on 2 December 2010 by A. Miralles and A. Rakotoarison.

Paratypes

(n=2). ZSM 779/2009 (ZCMV 13023), subadult specimen of unknown sex, from Ambalavao, Anja reserve, 21.8522°S, 46.8443°E, 972 m a.s.l., Haute Matsiatra Region, Madagascar, collected on 08 December 2009 by A. Crottini, D.J. Harris, I.A. Irisarri, A. Lima, S. Rasamison, and E. Rajeriarison; and ZSM 850/2010 (ZCMV 12791), adult specimen from Anja reserve, 21.8519°S, 46.8440°E, Haute Matsiatra Region, Madagascar, collected on 08 December 2010 by A. Miralles and F. M. Ratsoavina.

Additional non-type material.

ZFMK 59808, juvenile (but not neonate) specimen from Isalo. Two specimens deposited at the University of Antananarivo (field numbers ZCMV 12789 and 12790, not presently examined), collected on 08 December 2010 by A. Miralles and F. M. Ratsoavina, both from Anja reserve, 21.8519°S, 46.8440°E, Haute Matsiatra Region, Madagascar.

Diagnosis.

Paroedura rennerae sp. nov. is characterized by the unique combination of the following characters: (1) presence of prominent dorsal tubercles arranged in regular longitudinal rows, (2) presence of three broad light crossbands on the dorsum in juveniles and subadults, (3) spines on the tail, (4) nostril separated from rostral scale by prenasal, and (5) a curly-bracket shaped marking in the occipital region.

Paroedura rennerae sp. nov. can be distinguished from most other currently recognized Paroedura species by the presence of only three broad light crossbands on the dorsum in juveniles and subadults (the first one between forelimbs, the second one at midbody, and the third one between hindlimbs) versus four light crossbands in all other species except those of the P. bastardi clade ( P. bastardi , P. guibeae , P. ibityensis , P. neglecta , and P. tanjaka , which all have three crossbands) and P. oviceps and P. vahiny (in which the juvenile coloration is still unknown). It can be distinguished from P. gracilis by larger dorsal scales, absence of a white tip to the original tail, absence of a raised vertebral ridge on the dorsum and shorter forelimbs, which do not extend forward beyond tip of snout; from P. masobe by much smaller eyes and absence of a dorsal row of paired spines on the tail; from P. fasciata , P. homalorhina , P. hordiesi , P. vahiny , and P. spelaea by presence of spines on the original tail (versus absence); from P. gracilis , P. homalorhina , P. kloki , P. maingoka , P. masobe , P. oviceps (from its type locality Nosy Be), P. picta , P. spelaea , most P. tanjaka , and P. vahiny by the presence of prominent dorsal tubercles arranged in regular longitudinal rows (versus rather irregular rows of dorsal tubercles).

Within the P. bastardi clade, the species can easily be distinguished from P. tanjaka and P. neglecta by the absence of contact between the nostril and the rostral scale (versus presence). It can be distinguished from P. ibityensis by larger maximum SVL (> 70 mm versus 61 mm). In comparison with P. bastardi sensu novo and P. guibeae , the new species can be distinguished by the presence of a very sharp and contrasting dark transverse pattern, evoking the shape of a thin curly-bracket ({), in the occipital region and delimiting the skull from the neck. Moreover, Paroedura rennerae sp. nov. is unambiguously larger in size than P. guibeae (adult SVL> 70 mm versus <60 mm in P. guibeae ), and its dorsal tubercles are more prominent. It also lacks striped fingers (versus striped in P. guibeae ), and the light patch on its head lacks concave anterior edge and central vacuity in juveniles (versus both present in P. bastardi ).

Description of the holotype.

Adult female in very good condition, with the exception of the regenerated tail tip, which is amputated (ca. 10 mm missing). Head distinctly wider than neck, as wide as the body. Canthal ridges relatively well developed with a marked median depression. Ear opening is a vertical slit. Tail regenerated, nearly round (slightly flattened dorso-ventrally) in cross section in its proximal part; ventral pygal section of tail with a pair of poorly developed postcloacal sacs. Digits distinctly expanded at tips. Rostral scale rectangular, more than two times wider than tall and barely wider than mental. Nostrils separated from the rostral by prenasals. The two enlarged prenasals in contact with rostral and first supralabials, both separated by a single small granular scale. 12/12 (left/right) smooth supralabials, followed by two carinated tubercules above the mouth commissure. Eyes desiccated. Scales covering canthal ridges, loreal, temporal and periphery of the parietal region distinctly enlarged, spiny and tuberculate. Scales covering the dorsolateral side of neck and body heterogeneous, with enlarged, spiny, carinate and tuberculate scales regularly separated from each other by one (most often transversally) to three (most often longitudinally) rows of small, flat and juxtaposed scales or, along the vertebral line, by a single distinct row of smaller spiny tubercles. Seventeen longitudinal rows of tuberculate scales at midbody. Dorsal scales of forelimbs and hindlimbs mostly tuberculate and keeled, with a tetrahedral outline. Ventral scales of forelimbs distinctly smaller than surrounding ventral scales of the body. Three transverse rows at the base of the tail with six very spiny pygal scales per row. Ventrally, six rows of pygal scales squared and flat. Tail segments with irregular transverse row of spiny tubercles. Mental triangular, bordered posteriorly by a pair of elongated, irregular hexagonal postmentals. Each postmental in contact with six scales: other postmental, mental, first infralabial, one enlarged lateral gular, one smaller posterolateral gular, and one larger central gular. First three infralabials slightly larger (taller) than others. Gulars small, slightly granular. Ventrals of chest and abdomen flat and roundish. Proximal subdigitals in rows of mostly two. One pair of squarish terminal lamellae. Claws curving downwards between terminal pads of digits.

Measurements of the holotype (in mm): SVL = 73.6; TaL = 34.2 (tail regenerated and incomplete, distal tip of ca. 10 mm missing); HL = 21.0; HW = 16.9; HH = 10.1; AGL = 32.4; distE = 2.7, ED = 5.3, EO = 2.7; HAL = 8.6; TIBL = 13.0; FoL = 11.5.

After nine years in alcohol (Fig. 9 ), head dorsally ochre colored with a pair of dark temporal bands, running from the eyes to contact each other in the nuchal region. The contrast between the ochre dorsal side of the head and the darker temporal/nuchal bands is amplified by a dark blackish curly-bracket shaped transverse stripe in the occipital region (Fig. 8 ). Area along the upper lip alternating taupe-gray and cream. Body dorsally brown with three distinct lighter ochre (strongly contrasting thanks to very dark anterior and posterior borders) crossbands fading at the flanks: one transverse light crossband below forelimb insertion (width along the vertebral axis 6.1 mm), one distinctly broader light bow-tie-shaped crossband at midbody (width along the vertebral axis 9.6 mm), and one slightly less distinct band between the hindlimbs (6.3 mm). Dorsal surfaces of forelimbs and hindlimbs slightly marbled with brown and ochre (hindlimbs not darker than forelimbs). Flank coloration lighter than dorsum, fading gradually towards the ventral surface. Ventral coloration (throat, chest, abdomen, ventral parts of forelimbs and hindlimbs) cream (very slightly pigmented on the throat and chest).

Coloration in life (Fig. 9 ). The coloration of the preserved specimen is very similar to that of the living individual, although it is slightly duller (the contrasts are a little less strong and the colors a little less warm).

Variation.

Both paratypes, from Anja, present a lighter and more contrasted color pattern, with sharper dark lines (anterior and posterior margin of the light dorsal cross bands and dark curly-brackets delimiting the occipital region) (Figs 7 - 8 ). The tail of ZSM 779/2009, which is not regenerated, has seven pairs of regularly alternating light brown and cream crossbands delimited by dark (brown) transverse stripes, whereas the tail of ZSM 850/2010 (regenerated) is cream with five thin transverse zig-zagging dark brown stripes (Fig. 7 ). The specimen ZFMK 59808, juvenile (but not neonate) with its original tail, is relatively similar in coloration to the paratype ZSM 779/2009, although slightly paler. In contrast with adults with a regenerated tail, younger specimens (ZSM 779/2009, subadult and ZFMK 59808, juvenile) present very regular rows of spiny tubercules all along the tail (around 20 rows). See also Table 1 for the variation in measurements.

Etymology.

This new species, elegant and prickly, is dedicated to Susanne Renner, eminent botanist and evolutionary biologist, and Professor Emeritus of the University of Munich, in recognition of her substantial contributions to taxonomy and her invaluable collaboration in the framework of the “Taxon-omics” priority program of the German Research Foundation, DFG.

Habitat, habits, and distribution.

Paroedura rennerae is reliably known from five localities, some of them relatively distant from each other, suggesting this species is widely distributed in the central/southern region of Madagascar. In the dry forest of Kirindy CNFEREF, specimens have been observed on vertical surfaces (tree trunks, wooden walls of the CNFEREF camp huts), around 1 to 2 m above the ground. Like other members of the P. bastardi species complex, it is quick to bite when handled. In Anja, several specimens have been collected on granitic boulders. ZSM 779/2009 was found in a large cavity below two large granitic boulders, in a quite humid environment. In this cavity, ZSM 779/2009 and other individuals were found on the walls. In Isalo, specimens belonging to this species were found at two sites (Zahavola and Namazaha Valley). These individuals were found within the canyons of the sandstone Massif in shaded areas and in close proximity to a small cave or a small waterfall, again in quite humid microhabitats. Two additional 16S sequences confirm the presence of this species also in Marofandilia and Miandrivazo (GU129005 and GU128989, Aprea et al. 2013). All specimens have been observed at night or near dusk.

Figures

Figure 4. Multilocus phylogenetic trees (full dataset concatenated, nDNA and mtDNA concatenated trees), with haplotype networks reconstructed for each of the nuclear markers (after phasing). Photo credits: AM (P. rennerae sp. nov., P. picta, both from Kirindy), MV (P. ibityensis from Itremo), FG and JK (P. neglecta and P. tanjaka, both from Bemaraha), FG and MV (P. bastardi from Berenty, and P. guibeae from Tranoroa).

Figure 7. An overview of morphological diversity among the P. bastardi complex plus P. ibityensis. All specimens (adults, subadults and juveniles) are represented at the same scale (scale bar = 1 cm). Genotyped specimens are marked by the letter (G). Photo credit: FMNH 73049 © Field Museum of Natural History. Created by Field Museum of Natural History, Amphibian and Reptile Collection and licensed under CC-BY-SA 4.0. All other pictures taken by AM.

Figure 8. Details of the dorsal side of the head of the P. bastardi complex. Genotyped specimens are marked by the letter (G). Horizontal curly bracket ({) highlights the dark, contrasted and curved nuchal pattern evoking this symbol characterizing the specimens of Paroedura rennerae sp. nov. All pictures taken by AM.

Figure 9. Holotype of Paroedura rennerae sp. nov. (ZSM 849 / 2010, from Kirindy CNFEREF). The four top photographs are of the living specimen, whereas the bottom picture shows the preserved specimen after nine years in 70 % ethanol (pictures by AM).