Urkuphryne merinoi Ortega, Cisneros-Heredia, Camper, Romero-Carvajal, Negrete & Ron, 2025, sp. nov.
Creators: name: Ortega, Jhael A.
name: Cisneros-Heredia, Diego F.
name: Camper, Jeffrey D.
name: Romero-Carvajal, Andrés
name: Negrete, Leonardo
name: Ron, Santiago R.
Publisher: name: Zenodo
Year: 2025
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Strabomantidae
Urkuphryne
Urkuphryne merinoi

description: Urkuphryne merinoi sp. nov.
LSID: urn:lsid:zoobank.org:act: FD715F02-0CB4-4A5F-93C2- BFD649C05240
Holotype (Figs 28, 29): QCAZ 66078 (field no.SC-PUCE 48592) adult female from Ecuador, Carchi Province, Canton Espejo, El Goaltal Parish, protective forest Cerro Golondrinas, trail from the Heliconia Cabin to the Cortadera (0.8204°N, 78.0944°W), 2600 m a.s.l. Collected by Diego Almeida, Kunam Nusirquia, Darwin Núñez, Fernando Ayala, David Mantilla, Santiago Recalde, Carlos Castro, and Polibio Malte on 12 October 2016.
Paratypes (N = 8; Fig. 30): All collected in Ecuador, Carchi Province. QCAZ 66080 juvenile, QCAZ 66081– 82 adult males, QCAZ 66079, QCAZ 66083, and QCAZ 66091 adult females, same locality as holotype; QCAZ 66085 adult male from protective forest Cerro Golondrinas, Waterfall trail to Las Juntas (0.8217°N, 78.0951°W), 2544 m a.s.l.; collected by Diego Almeida, Kunam Nusirquia, Darwin Núñez, Fernando Ayala, David Mantilla, Santiago Recalde, Carlos Castro, and Polibio Malte on 12 October 2016. QCAZ 41813 adult female from Ecuador, Carchi Province, La Comadre waterfall (0.8233°N, 78.0232°W), 2740 m a.s.l. Collected by Juan Fernando Dueñas and Ítalo Tapia on 18 September 2008.
Proposed standard English name: Merino leaflitter frog.
Proposed standard Spanish name: Cutín de hojarasca de Merino.
Definition (Figs 28–33; Tables 2 and 3): We assign the new species to the genus Urkuphryne based on its phylogenetic relationships. The new species is characterized by: (1) skin on dorsum smooth to finely shagreen, dorsolateral folds absent, dorsal surfaces of thighs smooth, finely shagreen or shagreen, skin on throat, chest and belly smooth, discoidal fold present, ventral surfaces of thighs finely shagreen, skin on flanks shagreen to weakly areolate; (2) tympanic membrane, tympanic annulus, supratympanic fold, and postrictal tubercle present; (3) snout rounded to broadly rounded in dorsal and lateral views; (4) upper eyelid without tubercles, cranial crests absent; (5) vomerine teeth present; (6) vocal slits present, nuptial pads absent; (7) fingers not expanded distally, finger tips rounded to subacuminate, lacking papillae, finger I shorter than finger II, supernumerary tubercles present; (8) fingers lacking lateral fringes, pads present, lacking circumferential grooves; (9) distal phalanges of all toes and fingers narrower distally, with the tip pointed; phalangeal formula of hand 2-2- 3-3; (10) ulnar tubercles absent; (11) knees and heels without tubercles, outer edge of tarsus without tubercles, inner edge of tarsus bearing an inconspicuous fold; (12) inner metatarsal tubercle elongate in ventral view and rounded in lateral view, bigger than rounded outer metatarsal tubercle; (13) toes not expanded distally, with rounded to subacuminate toe tips, lacking papillae, supernumerary tubercles absent, lateral fringes on toes absent, toe basal webbing present, all toes bearing well-defined pads lacking circumferential grooves, toe V shorter than toe III; (14) in life, dorsal surfaces dark brown, bearing darker circular sacral lateral patches delineated by an orange line; dark brown facial mask from the tip of snout to near the groins, bordered dorsally by an orange line; throat dark brown or cream in females and almost black in males; ventral surfaces light brown or brown, bearing white spots; and (15) SVL in adult males 17.22 mm (N = 3) and SVL in adult females 19.69 mm (N = 5) (Table 4).
Diagnosis (Tables 2 and 3): Urkuphryne merinoi resembles species of the genus Phyllonastes but can be distinguished from them by the presence of vomerine teeth (absence of vomerine teeth is a synapomorphy of Phyllonastes) and by the absence of circumferential grooves on toes (present in all Phyllonastes species). Urkuphryne merinoi can be differentiated easily from Barycholos pulcher (Boulenger, 1898) by the presence of vomerine teeth in two small rows, oblique, broadly separated, and posteromedial to choanae (vomerine teeth in two long transverse rows posterior to the level of choana, narrowly separated in Barycholos pulcher).
Description of the holotype (Figs 28 and 29): Adult female (QCAZ 66078). Measurements (in millimetres): SVL, 19.53; tibia length, 7.79; foot length, 8.66; head length, 5.74; head width, 7.05; eye diameter, 2.14; tympanum diameter, 1.18; interorbital distance, 1.93; upper eyelid width, 1.31; internarial distance, 1.79; eye–nostril distance, 1.51.
Head wider than long, head wider than body; canthus rostralis slightly convex in lateral view; loreal region slightly concave in dorsal view; cranial crests absent; upper eyelids bearing no tubercles. Tympanic annulus weakly defined, differentiated only at its lower half; tympanic membrane present; small supratympanic fold present; postrictal tubercle present. Snout rounded in dorsal and lateral views, without rostral papilla. Vomerine teeth present, small, oblique, broadly separated, posteromedial to choanae; each vomer bearing small teeth; vocal slits and nuptial pads absent, 30 teeth visible in the upper maxilla.
Skin on dorsum smooth, skin on flanks shagreen; dorsolateral folds present; skin on throat, chest, and belly smooth; ventral surfaces of thighs finely shagreen; discoidal fold absent; skin in ventral cloacal region areolate. Ulnar tubercles absent; palmar tubercles prominent, outer palmar tubercle circular on right hand and slightly elongate on left hand, thenar tubercle large and elongate; proximal subarticular tubercles well defined, round in ventral and lateral view; distal subarticular tubercles not visible; minute and round supernumerary tubercles all over palmar surface; lateral dermal fringes absent; discs narrow, rounded, and bearing well-defined rounded pads without circumferential grooves; fingers not expanded distally, tip of fingers rounded without papillae; phalangeal formula of 2-2-3-3; relative length of fingers is I <II <IV <III.
Hindlimbs robust; dorsal surface of hindlimbs smooth; posterior and ventral surfaces of thighs shagreen; heel and knee without tubercles; outer tarsal folds absent; inner tarsal fold present, inconspicuous and thin; outer tarsal tubercles absent; inner tarsal tubercle absent; inner metatarsal tubercle small (1.08 mm), prominent, elongate in ventral view and rounded in lateral view, 1.86 × longer than rounded; outer metatarsal tubercle small (0.75 mm), well defined, rounded, and prominent; plantar surface without supernumerary tubercles; basal subarticular tubercles well defined, round in dorsal view; distal subarticular tubercles not visible; toes without lateral dermal fringes; basal webbing between toes present; discs narrow, rounded and bearing well-defined rounded pads without circumferential grooves; toes not expanded distally, tip of toes rounded without papillae; relative length of toes is I <II <V <III <IV; toe III longer than toe V (toe III surpasses the distal border of the second subarticular tubercle of toe IV; toe V reaches the proximal border of the second tubercle of toe IV).
Colour of holotype in life (based on digital photographs) (Fig. 28): The dorsal surface is dark brown, with a few scattered white and low tubercles, and covered by small orange dots that group in two conspicuous lateral lines (one on each side) from the arm insertion to the tip of the snout through the upper eyelids. Posteriorly, the dorsum bears two dark brown circular sacral lateral patches, with a few orange flecks and delineated by an orange line formed by the aggregation of the same orange dots, more densely grouped medially at the level of the sacrum. The dorsal surface of limbs is dark brown, covered with white and orange flecks; it bears a few irregular dark brown blotches delineated by aligned orange dots. Dorsal feet surface dark brown, covered by orange and yellowish brown flecks and with irregular dark brown blotches with yellowish brown borders formed by the alignment of the flecks. Laterally, a dark brown facial mask extends as a dark band from the tip of snout to near the groins. The facial mask bears irregular white marks and orange and yellowish brown flecks more conspicuous rostrally. The flecks form ill-defined bars in the upper lip, with white marks. Along its path, the mask is bordered dorsally by orange spots, longitudinally aligned. Groins are dark brown, uniformly covered by small orange dots and with some scattered white marks. Throat uniformly dark brown, darker than belly and limb ventral surfaces, with white scattered marks. Belly and limb ventral surfaces with dark brown flecks densely aggregated in many irregular patches alternated with yellowish brown irregular blotches, with fewer dark brown flecks. Posterior surfaces of thighs dark brown with orange flecks; flecks form an orange stripe in the upper cloacal region that extends over the thighs.
Colour of holotype in preservative (Fig. 28): Dorsal surfaces dark brown, with darker sacral lateral circular patches surrounded by a poorly defined cream edge. Facial mask dark brown (darker than dorsal surface) to black extending from the tip of snout to near the groins, delimited dorsally by a thin cream line. Throat dark brown, belly and ventral surfaces of limbs with a mosaic of cream and dark brown blotches bearing dark brown flecks. Posterior surfaces of thighs dark brown, with a darker triangular blotch in the cloacal region limited on its upper side by a thin cream line. Variation (Fig. 30): Vocal slits are present in adult males (e.g. QCAZ 66085); dorsum skin might be finely shagreened (e.g. QCAZ 66079 and QCAZ 66083) and on flanks (e.g. QCAZ 41813); the latter can also be weakly areolate (QCAZ 66083). Snout might be broadly rounded (QCAZ 66083). Supernumerary palmar tubercles might be inconspicuous (e.g. QCAZ 66091); tip of fingers might be subacuminate (e.g. QCAZ 66081). Dorsal surfaces of limbs might be weakly shagreened (e.g. QCAZ 66085 and QCAZ 66091) or shagreen (e.g. QCAZ 41813); toe III might be subacuminate (QCAZ 66082). Live individuals might have dark brown, almost black (e.g. 66082), or cream (e.g. QCAZ 66091) throat and chest; the round sacral lateral patch might expand anteriorly to the iliac region (e.g. QCAZ 66085) or be inconspicuous (QCAZ 66083); belly and dorsal surfaces might be light brown (e.g. QCAZ 66085); the entire coloration can be lighter (e.g. QCAZ 66083 and QCAZ 66091). Morphometric variation is detailed in Table 4.
Osteology
The osteological description is based on micro-CT images of the holotype (adult female QCAZ 66078) and paratypes (adult females QCAZ 41813 and QCAZ 66091).
Skull (Fig. 31): The skull is slightly wider than long; the widest section is at the quadratojugal–maxillary joint. The longest part of the skull, measured from the anterior face of the premaxilla to posterior border of the exoccipital, is 92.5% of its widest section. The rostrum is short, with a distance from the anterior edge of the frontoparietals to the anterior face of the premaxilla of ~32.2% of the skull length (measured from the anterior face of the premaxilla to posterior face of the exoccipital). At the level of the anterior edge of orbits, skull is ~62.2% of the maximum skull width and reaches ~88.5% of the maximum skull width at the level of midorbit. The posterior edge of orbits reaches 99.2% of the maximum skull width and is located posterior to it.
The braincase contains well-ossified elements. The frontoparietals are well-developed bones, markedly longer than wide and slightly narrower anteriorly than posteriorly; frontoparietals are ossified and sutured along their entire length, but they leave a thin, short, unossified fontanel at their anterior end. At their posterior end, the frontoparietals are completely suturedwiththeotoccipitals, togetherformingprominentridgeson each side and a completely closed braincase. The ridges formed have very small unossified patches. The otoccipitals are formed by well-fused prootic and exoccipital, with small unossified patches in the crests they formed when fused. Ridges formed between frontoparietals + otoccipitals and prootic + exoccipitals form a continuous V-shaped ridge, with the anterior ramus (fro ntoparietals + otoccipitals) more prolonged (posterior ramus is 61.96% of the anterior ramus). The apex between rami is medial and angled at slightly>90°. At their anterior and lateral end, the frontoparietals fuse with the sphenethmoids. The latter have complete ossification and are fused ventrally with the anterior portion of the cultriform process of the parasphenoid. Ventrally and posteriorly, the sphenethmoids surpass the midpoint of the orbits.
The cultriform process of the parasphenoid is complete and well ossified, with a biconvex shape, with anterior and posterior ends pointed and thinner than its medial segment; at its base (at the anterior border of the alar processes of the parasphenoid) is ~8.21% of the maximum skull width and reaches its maximum thickness at its midpoint (24.09% of the maximum skull width). It is important to mention that the alar processes do not emerge from the posterior end of the cultriform process, but a few millimetres anteriorly, hence the posterior tip of the cultriform process protrudes posteriorly. Fenestra vestibule (parasphenoid alar processes + otoccipitals) is poorly ossified, and their shape cannot be detailed.
The neopalatines are thin and separated from one another. They articulate with the ventral face of the sphenethmoid by their upper tip and with the internal face of the maxilla in its anterior third, by their lower end forming the anteroventral corner of the orbit. The septomaxilla is small and horseshoe shaped, with a medial and slightly posterior opening; both ends are pointed, and the centre is widened and flattened laterally; both ends are free and do not articulate with any bone. The prevomers are large, located medial, superior, and posterior to the septomaxilla; they are broadly separated from one another medially; the separation is greater rostrally. The dentigerous processes are conspicuous, more visible ventrally, in the left prevomer. The columella (or stapes) is large and well ossified. The nasals are thin and posterolaterally expanded. They are ossified only in their posterior portion, hence they are completely distant from the alar processes of the premaxilla. Posteroventrally they are fused with the sphenethmoids and posteriorly with the anterior border of the frontoparietals; their posterolateral expansion is close but not fused to the neopalatine + maxillary joint.
The maxillary arch bears many small and moderately sized teeth on the maxillae and on the transversal section of the premaxillae. The premaxillae are partly separated medially, and their rostral alary processes rise divergent from the midline. Alary processes of the premaxillae are prominent and widened in the opposite direction to their base, rising perpendicular to the cross-section of the premaxillae. They are separated ventrally from each other. The short premaxilla and maxilla are in lateral contact by juxtaposed articulation. The maxilla becomes narrow posteriorly, reaching an acuminate caudal end, which is in contact, on its inner side, with the quadratojugals. The triradiate pterygoid bears a long, curved rostral ramus oriented anterolaterally that fuses with the internal face of the maxilla near its midpoint; this ramus is the longest of the three, it is flattened laterally, and its anterior end is thinner than the posterior one. The caudal ramus is shorter than the rostral one and as wide as the latter in its initial portion, and likewise, it sharpens towards the tip (posterior end), which merges with the posterior tip of the angulosplenial; this ramus is also flattened laterally. The medial ramus has the same length as the caudal, is widened in its terminal portion and flattened anteroposteriorly; on its posterior face it articulates with the anterolateral surface of the otoccipital.
The quadratojugal is well ossified; it articulates by its dorsoposterior face with the ventral ramus of the T-shaped squamosal. The squamosal has three rami; the posteroventral ramus widens towards its free (posterior) end as it moves away from its base; this ramus is flattened anterodorsally, it is slightly shorter than the posterodorsal ramus and longer than the zygomatic or anterior ramus. The posterodorsal ramus is flattened dorsoventrally, and its root is very close to the columella; the small zygomatic ramus is flattened laterally. Both branches (posterodorsal and anterior) become pointed towards their free ends and do not articulate with any bony structure.
The mandible is slim and completely edentate. The mentomeckelians are small, medially separated from each other, in the shape of a cylinder widened towards their ends; they are in contact posteriorly with the dentaries, which are short, acuminate towards the posterior end, and flattened anterodorsally. From the middle of its posterior border and with its posterior end, the dentary bones articulate with the anterior face of the angulosplenial.The angulosplenial is long, arcuate, and expanded at its posterior end; it articulates broadly by its anterolateral face with the posteromedial face of the slim and relatively small dentaries. The only ossified portions of the hyoid apparatus are the two posteromedial processes, which are moderately expanded anteriorly and posteriorly. Both posteromedial processes present a steep anteroventral inclination and are moderately separated from one another at the anterior ends; the separation between them increases posteriorly.
Postcranium (Fig. 32): There are eight non-imbricate presacral vertebrae. The first presacral vertebra (cervical vertebra) is wider than posterior vertebrae and has no diapophyses. The cervical vertebra has a type I cotylar arrangement. The cervical cotyles receive the occipital condyles of the cranium, which are widely separated from each other, leaving a foramen magnum of 21.69% of the maximum skull width.
Presacral vertebrae II–VIII bear well-developed diapophyses. The transverse processes of the presacrals increase in length as they move away from the skull, with the transverse processes of presacral II being the shortest. The transverse processes of presacral II are horizontal and have a dorsoventral flattening and slightly widened lateral free ends. The transverse processes of presacral III are slightly rotated forwards and present an anteroposterior flattening; in addition, their lateral free ends are evidently wider. From presacral IV to VII, the transverse processes are rotated 90° and are also flattened anteroposteriorly; these processes are not completely ossified toward their ends. In width, the processes of presacral III are followed by those of presacral II, the second widest slightly expanded distally, and those of presacral IV, the third widest, with their edges parallel along its entire length. Transverse processes of presacrals V–VIII are the smallest and similar in size (length and width) between them. This vertebrae of this species are characterized by having a holochordal centrum.
The sacrum bears moderately expanded diapophyses. Transverse processes are oriented laterally and slightly dorsally, with an angle of dorsal opening of ~140°, and articulate distally with the ilia, a few millimetres anterior to its front end. The sacrum is articulated caudally with the coccyx or urostyle by a bicondylar articulation. The long and thin urostyle is slightly longer than the presacral portion of the vertebral column (the presacral portion of the vertebral column is 91.85% of the urostyle) and bears a well-developed longitudinal ridge, which is largest anteriorly (at the point of its articulation with the sacrum) and gradually decreases in height posteriorly. The urostyle does not have complete transverse processes nor remnants and is not fully ossified at its posterior end. The sacrum–ilia articulation is not visible in the micro-CT scan. The ilia articulate posteromedially with each other and posteriorly with the ischia and present a long shaft.
In the pectoral girdle, the clavicles are long and slim, oriented anteromedially, curved with an anterior opening concavity, with the medial tips articulated between them. The coracoids are stout, with the anterior edge curved, with a pronounced anterior opening concavity, and with a straight posterior edge; their medial tips are separate from each other, and their sternal and glenoidal ends are widened, the sternal ones more than the glenoidal ones. The scapula is long, with the anterior edge slightly oriented anteromedially. Epicoracoids are visible. The sternum has no well-ossified elements. The omosternum is absent.
Manus and pes (Fig. 33): All phalanges are well ossified, with a phalangeal formula for the fingers and toes of 2-2-3-3 and 2-2-3-4-3, respectively. The increasing order of finger length is I <II <IV <III, and that of toes is I <II <III <V <IV. Distal phalanges of all toes and fingers are narrower distally, with the tip pointed. Bones of the carpus and metacarpus are completely ossified.
Distribution, natural history, and conservation status (Fig. 3): Urkuphryne merinoi is known from the surroundings of Protective Forest Cerro Golondrinas from 2500 to 2800 m a.s.l. in Western Montane Forest. Surrounding the protective forest, in both highlands and lowlands, there are deforested areas, some in regeneration, near human settlements and artificial open areas for agriculture and cattle raising. Individuals were collected in secondary forest, in the afternoon (between 12:30 and 13:30 h) on mossy rock walls with high humidity or buried in leaf litter, the adult male QCAZ 66091 was found vocalizing at 12:30 h buried in soil; and at night (19:00–22:30 h) on very damp rock walls ~ 150 cm from the ground, on leaf litter and mossy soils near creeks, or buried ~ 3 cm deep in soil. All females found were next to clutches with 8– 12 eggs. One female (QCAZ 41813) was collected between 9:00 and 10:30 h on the roadside in a mossy habitat with herbaceous plants, buried under moss next to a clutch of white eggs.
Because of the lack of information on population size and geographical range, we assign U. merinoi to the Data Deficient IUCN Red List Category (based on IUCN Standards and Petitions Committee 2023).
Etymology: The specific name merinoi is a noun in the genitive case and is a patronym for Andrés Merino-Viteri, an Ecuadorian herpetologist, professor at Pontificia Universidad Católica del Ecuador (PUCE). During his career, Andrés Merino has contributed significantly to the study of chytridiomycosis and possible consequences of climate change in anurans. At PUCE, he is director of The Threatened Amphibian Conservation Initiative of Ecuador ‘Balsa de los Sapos’, one of Ecuador’s largest and long-lasting ex situ amphibian conservation projects in the world.
Embryic development in U. merinoi and P. personinus
The U. merinoi clutch (Fig. 34A) was collected in the daytime, among moss with a lot of humidity on a rock wall. It was next to an adult female, presumably its mother. We were unable to observe behaviours related to mating or fertilization. It had 12 eggs, with relatively transparent jelly coats. Out of 12 eggs, 9 had developing embryos and 3 were unfertilized. Unfertilized eggs had a diameter of 4.03 ± 0.02 mm (SD). The embryos were at two stages: TS5 and TS11. Five embryos were at stage TS5 and had a length (from snout to tail tip) of 4.63 ± 0.01 mm. All these embryos failed to develop. The remaining four embryos were at stage TS11 and had a total length of 10.40 ± 0.33 mm. Development of these four embryos was followed until TS13+, when the last embryo died.
The P. personinus couple was found and collected in amplexus and transported to the laboratory. The clutch was laid during transport; thus, we were not able to observe mating or fertilization behaviours. The nest was deposited in a terrarium in the laboratory with leaf litter (Fig. 34B). It had two embryos and three unfertilized eggs. Unfertilized eggs had a diameter of 2.15 ± 0.01 mm. The remaining two embryos were at TS5 and had a total length of 2.97 ± 0.05 mm. Only one of these embryos hatched as a froglet.
For both species, embryos at TS5 have the shape and structures of most direct-developing embryos, with distinctive limb and tail buds and with a head with visible eyes. In both species, branchial arches and external gills were not apparent, and posterior limb buds were larger than the anterior ones. TS5 embryos in both species apparently lack oocyte-derived pigment, because both have clear unpigmented yolk and white embryonic integument. In P. personinus, melanocytes (neural crest-derived melanophores) are spread on the integument from the most dorsal part of the body but have not yet covered the yolk or the limb buds. Also, no retinal pigmentation was observed at this stage. Stages TS6–TS9 were recorded only in P. personinus. In TS6, pigmentation has advanced to the head and limb buds. At this stage, endolymphatic calcium deposits are rounded structures that protrude prominently from the posterior dorsal region of the head. During TS7 the eye develops retinal pigmentation distinct from the unpigmented lens vesicle (pupil), and melanocytes cover the whole dorsal body. By TS9, melanocytes and the body wall have covered one-third of the yolk surface and the endolymphatic calcium deposits, which now have a triangular shape. Compared with anterior limbs, at TS7, posterior limbs show more growth and differentiation, with distinctive paddles and knee joint constrictions. Forelimbs seem to develop joint constrictions later (TS9). At TS10, we detected hindlimb movement, and autopod paddles showed finger differentiation (not shown). Between TS6 and TS7, translucent membranous fins with vascularization were first apparent in the tail.
Urkuphryne merinoi TS 11 embryos are paler and seem to have less melanocytes that are spread across the whole body except for the ventral midline. In this species, endolymphatic calcium deposits were still visible and covered the dorsal surface of the hindbrain bilaterally, with projections towards the eyes.
In both species, the egg tooth is visible as an unpigmented monocuspid structure. Between TS12 and TS13, the egg tooth becomes keratinized and darkly pigmented. This was more evident in U. merinoi.
The tail develops to its maximum length by TS 11 in P. personinus and by TS 13 in U. merinoi. The tail of U. merinoi seems to be thinner and larger than that of P. personinus; however, both have symmetrical fins over the dorsoventral axis. The progress between TS13 and TS14 implies the reduction of yolk mass and the reabsorption of the tail, which remained as a translucent vestige in the P. personinus froglet. The progress from TS11 to TS 13 in U. merinoi lasted 10 days. The progress from TS5 to hatching in P. personinus lasted 32 days.
description: Published as part of Ortega, Jhael A., Cisneros-Heredia, Diego F., Camper, Jeffrey D., Romero-Carvajal, Andrés, Negrete, Leonardo & Ron, Santiago R., 2025, Systematics of minute strabomantid frogs allocated to the genus Noblella (Amphibia: Anura) with description of a new genus, seven new species, and insights into historical biogeography, pp. 1-60 in Zoological Journal of the Linnean Society 203 (1) on pages 44-53, DOI: 10.1093/zoolinnean/zlae162, http://zenodo.org/record/14832689