Brookesia nofy Rakotoarison, Hasiniaina, Glaw & Vences, 2024, sp. nov.
Creators: name: Rakotoarison, Andolalao
name: Hasiniaina, Alida Frankline
name: Glaw, Frank
name: Vences, Miguel
Publisher: name: Zenodo
Year: 2024
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Squamata
Chamaeleonidae
Brookesia
Brookesia nofy

description: Brookesia nofy sp. nov.   
Figs. 3, 5  
 Holotype. ZSM 30 /2024 (field number ZCMV 13721), adult male with fully everted hemipenes, found by locals and collected by Miguel Vences, Sandratra Rakotomanga, Solohery Rasamison, and Pedro Galan, at Ankanin’ny Nofy forest close to Palmarium Beach Hotel, eastern Madagascar, geographical coordinates ca. -18.6261, 49.2037, near sea level on 22 November 2022.   
 Paratype. UADBA-R-70860 (field number ZCMV 13722), probably a subadult male, collected by Andolalao Rakotoarison, Alida F. Hasiniaina and Paul at the same locality as the holotype on 15 September 2023.   
 Diagnosis. A miniaturized chameleon assigned to the subgenus Evoluticauda in the genus Brookesia on the basis of its very small body size (SVL of adult male <21 mm), short tail, presence of rows of dorsolateral tubercles along vertebral column, presence of pelvic spine, and molecular phylogenetic relationships. Brookesia nofy sp. nov. is distinguished by the following unique suite of morphological characters: (1) male SVL 20.5 mm; (2) male TL mm 32.8 mm; (3) TaL/SVL ratio 0.52–0.60; (4) absence of lateral or dorsal spines on the tail; (5) absence of dorsal pelvic shield in sacral area; (6) presence of distinct pelvic spine; (7) 5–6 dorsolateral pointed tubercles; (8) hemipenis of balloon-like shape and without obvious ornamentation.   
 Within the genus Brookesia, B. nofy sp. nov. can easily be distinguished from adults of all species that are not members of the B. minima species group (i.e., the subgenus Evoluticauda) by its diminutive size (SVL <21 mm vs. ≥ 34 mm).   
 Within the B. minima species group, we here focus on comparisons with B. ramanantsoai which is undoubtedly most similar and phylogenetically sister to the new species, and only give a subset of most relevant diagnostic characters to the other species. The new species can be distinguished from many species by the smaller total length (TL) of males: the known adult male (33.0 mm) is substantially smaller than all known male specimens of B. desperata (39.7–42.9 mm), B. exarmata (39.8–40.1 mm), and probably also B. karchei (only female size reliably known: 51.0 mm) and B. dentata (no precise measurements known). It is substantially larger than the extremely miniaturized species B. micra (male TL 22.5–23.6 mm) and B. nana (male TL 21.6 mm). The presence of distinct pelvic spines differentiates the new species from B. minima (absent or indistinct pelvic spines). The short relative tail length (tail length/SVL) constitutes a difference to B. minima (0.65–0.73), B. tedi (0.74–0.92), B. tristis (0.71–0.72), and B. tuberculata (0.68–0.88). Finally the globular (balloon-like) hemipenis without obvious ornaments strongly differs from the genital organs of B. tuberculata (elongated, tubular), B. micra (elongated with apical papillae), B. confidens (short and cone-like), B. desperata, B. peyrierasi and B. tristis (with small apical spines or apical lobes).   
 The new species is morphologically most similar to its sister species, B. ramanantsoai. It differs by smaller total length (male TL 32.8 mm vs. 34.5–44.8 mm), which is primarily caused by relative tail length being consistently smaller in all available males, females and juveniles (TAL/SVL 0.52–0.60 vs. 0.61–0.83). It also appears to differ by a minimally smaller snout–vent length (male SVL 20.5 vs. 20.6–25.0) and perhaps by relatively larger heads (HW/SVL 0.19–0.20 vs. 0.14–0.17; HH/SVL 0.20–0.21 vs. 0.15–0.18; based on our own measurements only). Also distinguished by a cephalic ridges pattern with U-shaped ridge with two inner ridge (vs. U-shaped ridge absent and three inner ridges; Fig. 5).   
 Description of the holotype. Adult male in excellent state of preservation, right forelimb removed as tissue sample (Figs. 3, 5). See Table 1 for measurements and ratios.   
Lateral crest on head poorly recognizable; orbital crest well developed, denticulated, consisting of conical tubercles with one prominent supraocular cone directed anteriorly; transversal row of enlarged tubercles at the posterior edge of head not recognizable; moderately distinct border between head and body; a pair of curved crests that start above the eyes and fade in the neck followed by a U-shaped ridge posteriorly (Fig. 5); depression between the eyes lacking any further crests; one pointed tubercle on each side of head; few scattered, slightly enlarged tubercles on lateral surfaces of head; supranasal cone distinct, not projecting beyond tip of snout; chin and throat without enlarged tubercles. Dorsal surface of body without vertebral ridge or keel; 6/5 (left/right) dorsolateral pointed tubercles that form an incomplete longitudinal line, ending behind midbody; anteriormost pointed dorsolateral tubercle being largest; pointed dorsolateral tubercles along vertebral column almost equally spaced; dorsal surface of tail lacking distinctly enlarged tubercles; no dorsal pelvic shield in sacral area, but distinct small pelvic spine directed posterolaterally; lateral surface of body with few indistinct, irregularly spaced enlarged tubercles; venter without enlarged tubercles; several distinctly enlarged tubercles on limbs; no pointed tubercles around cloaca; longitudinal rows of slightly enlarged tubercles lateral on anterior tail; no dorsal, lateral, or ventral spines on tail; no enlarged tubercles on ventral surfaces of tail.  
Both hemipenes fully everted, 4 mm long, bulbous (Fig. 5). A pair of knob-like structures emerge from the apical surface, each of which being the apical end of a fleshy lobe. The truncus is smooth and lacks any trace of calyces.  
In life (when apparently largely unstressed, Fig. 3A), overall dorsal ground coloration was light brown, with a row of black tubercles dorsolaterally, starting behind the insertion of forelimbs; additional, partly indistinct spots scattered throughout the body and head, without forming any conspicuous pattern; limbs with slightly darker pattern, distal parts of fingers and toes black. Ground color of tail almost dirty white and distinctly lighter than body. Dorsal surface of head with some reddish-brown pattern, anterior head between the eyes with an orange-brown band, bordered by a contrasting darker brown line; a light, dirty white area below this band and the tip of snout, with scattered very fine red dots; a series of radiating brown spots and streaks is present around the eyes.  
After one year in ethanol, the body coloration is generally faded with less evident pattern. The ground coloration is pale gray on head and body, but still lighter on the tail. Head and body with diffuse pattern of different shades of brown, gray, and white. Dorsolateral tubercles and pelvic spines black. The dark radial streaks are still recognizable and the distal parts of finger and toes are still rather dark.  
 Variation. The subadult male paratype is in good state of preservation. Snout–vent length 15.5 mm, see Table 1 for additional measurements and ratios. In life (when apparently undisturbed), dorsal color beige to gray, with a row of dark gray tubercles along the dorsolateral border, starting behind the insertion of forelimbs and ending at the insertion of hindlimbs; additional smaller spots scattered throughout the body and head, but generally lacking any conspicuous pattern; limbs with darker pattern, especially on elbow; distal parts of fingers and toes black. Dorsal surface of head with some reddish-brown pattern, anterior head between the eyes with an orange-brown band, bordered by a contrasting darker line and small scattered black spots; a series of radiating blackish or reddish-brown streaks is present around the eyes; iris light (golden). When apparently stressed, limbs, ventrolateral border of body, and eyes can get almost black (Fig. 3D). The tail base is distinctly narrower than in the male holotype.   
 Etymology. The species name is a noun in apposition to the genus name, derived from the Malagasy word “nofy ”, which means “dream”. This choice is reflective of its association with the type locality Ankanin’ny Nofy. Additionally, the name refers to the currently ongoing regeneration of the forest at this locality (see discussion) which can be seen as a dream come true: a chance for sustainable conservation of these microendemic Brookesia.   
 Natural history. Specimens were found in the Ankanin’ny Nofy forest next to Palmarium Hotel which has been regenerating for about 20 years due to its use as touristic site by the hotel. One pair of B. nofy (not collected), with a slightly lighter colored male riding on the back of the female, was found head upwards on a thin vertical branch about 2 cm above the leaf litter (Fig. 3E) at Ankanin’ny Nofy on 14 September 2023. According to information from local people in September 2023, the species is apparently rather abundant in the rainy season. The nocturnal walk on 14 September 2023 began at 18:15 and lasted for 1 hour. In 28 minutes of searching along the forest trail, 8 specimens were discovered. A second pair of mating individuals was also observed on a small branch, approximately 2 cm above the ground, while the other four individuals were found at heights ranging from 4 to 8 cm. On 15 September 2023, no mating specimens were encountered, and only two individuals were observed during the same time frame as the previous night. We hypothesize that the decrease in activity on 15 September may be attributed to the absence of rain on 14 September. Heavy rain occurred on 13 September 2023, the first significant rainfall of the month. According to the guide, it was also the first time they observed Brookesia nofy sp. nov. mating during night tours with tourists. Thus, the heavy rain may have triggered reproductive activity.   
 Other herpetofauna found at Ankanin’ny Nofy and the neighboring Vohibola (updated after Gehring et al. 2010, 2011 with our new, unpublished observations in 2023) includes the frogs Anodonthyla cf. boulengeri, Blommersia dejongi, Boophis opisthodon, Boophis tephraeomystax, Guibemantis methueni, Heterixalus madagascariensis, Plethodontohyla notosticta; and the squamates Acrantophis madagascariensis, Calumma vohibola, Ebenavia sp., Furcifer pardalis, Furcifer willsii, Geckolepis sp., Hemidactylus mercatorius, Ithycyphus goudoti, I. cf. perineti, Lygodactylus fritzi, Madagascarophis colubrinus, Madascincus cf. melanopleura, Oplurus cuvieri (probably introduced), Phelsuma guttata, P. kely, P. laticauda (probably introduced), P. lineata, P. parva, P. ravenala, Trachylepis gravenhorstii, Uroplatus sameiti, and Zonosaurus cf. haraldmeieri (probably introduced).
description: Published as part of Rakotoarison, Andolalao, Hasiniaina, Alida Frankline, Glaw, Frank & Vences, Miguel, 2024, A new miniaturized species of leaf chameleon, genus Brookesia, from a littoral forest fragment in eastern Madagascar, pp. 533-547 in Zootaxa 5506 (4) on pages 541-544, DOI: 10.11646/zootaxa.5506.4.3, http://zenodo.org/record/13747416