Huicundomantis, Paez, Nadia B. & Ron, Santiago R., 2019

Paez, Nadia B. & Ron, Santiago R., 2019, Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species, ZooKeys 868, pp. 1-112 : 1

Huicundomantis subgenus nov.

Type species.

Pristimantis phoxocephalus (Lynch, 1979).

Definition.

This clade is strongly supported by genetic evidence ( Fig. 3 ). Morphological synapomorphies are unknown. Members of this clade are characterized by: (i) dorsolateral folds absent (except for P. atratus ); (ii) cranial crests absent (except for P. atratus , P. percultus , and P. spinosus ); (iii) tympanic membrane and tympanic annulus prominent (both absent in P. philipi ); (iv) dentigerous processes of vomer present; (v) small to prominent tubercles on heel; (vi) fingers with lateral fringes; (vii) basal webbing between toes (except for P. balionotus , P. hampatusami , and P. philipi which lack webbing); (viii) Toe V longer or much longer than Toe III ( Figs 7 - 9 ); (ix) in life, groins and concealed surfaces of thighs with distinctive coloration patterns including flash colors and light or bright colored flecks or spots on a darker background; colors, shapes and sizes of these ornaments are variable among species; (x) SVL females 24.9-46.8 mm; SVL males 16.1-34.5 mm. Species of this clade may bear a fleshy keel or a papilla at the tip of the snout, and lateral, middorsal, or postocular folds.

Content.

This clade comprises 23 described species (11 of them described below): P. atillo sp. nov., P. atratus , P. balionotus , P. chomskyi sp. nov., P. cryptomelas , P. gagliardoi , P. gloria sp. nov., P. hampatusami , P. jimenezi sp. nov., P. lutzae sp. nov., P. multicolor sp. nov., P. muscosus , P. nangaritza sp. nov., P. percultus , P. philipi , P. phoxocephalus , P. spinosus , P. teslai sp. nov., P. tinguichaca , P. torresi sp. nov., P. totoroi sp. nov., P. versicolor , P. verrucolatus sp. nov. This clade encompasses the P. phoxocephalus and P. cryptomelas species groups.

Distribution.

Huicundomantis occurs in Eastern and Western Andean slopes and Inter-Andean valleys of southern and central Ecuador, and Eastern Andean slopes of northern Peru. They inhabit the following Natural Regions: Deciduous Costa Forest, Western Foothill Forest, Western Montane Forest, Paramo, Inter-Andean Shrub, Eastern Montane Forest, and Eastern Foothill Forest, between elevations of 230 and 4200 m a.s.l.

Etymology.

We name this clade Huicundomantis because these frogs are frequently found inside bromeliad plants. Huicundo is a word in Quechua, an indigenous South American language, locally used to referring to bromeliads.

Pristimantis phoxocephalus species group new taxon

Definition. The P. phoxocephalus species group is strongly supported in our phylogeny. Members of this group share the following morphological traits: (i) dorsolateral folds absent (except for P. atratus ); (ii) snout with a fleshy keel or papilla at the tip; (iii) cranial crests absent (except for P. atratus and P. percultus ); (iv) tympanic membrane and tympanic annulus prominent; (v) dentigerous processes of vomer present; (vi) males with vocal slits (except for P. versicolor ); (vii) fingers and toes with lateral fringes; (viii) basal webbing between toes (except for P. balionotus and P. hampatusami ); (ix) Toe V longer or much longer than Toe III ( Fig. 7 ); (x) in life, groins and concealed surfaces of thighs with distinctive coloration, including flash colors and light or bright colored flecks or spots on a darker background; colors, shapes, and sizes of ornaments variable among species; (xi) SVL females 24.9-46.8 mm; SVL males 16.8-34.5 mm.

Content. Currently, the P. phoxocephalus group comprises 17 described species (10 of them are described below): P. atillo sp. nov., P. atratus , P. balionotus , P. chomskyi sp. nov., P. gloria sp. nov., P. hampatusami , P. jimenezi sp. nov., P. lutzae sp. nov., P. multicolor sp. nov., P. percultus , P. phoxocephalus , P. teslai sp. nov., P. tinguichaca , P. torresi sp. nov., P. totoroi sp. nov., P. versicolor , P. verrucolatus sp. nov.

Distribution. Eastern and Western Andean slopes and Inter-Andean valleys of central and southern Ecuador, in ten provinces: Azuay, Bolívar, Cañar, Chimborazo, Cotopaxi, El Oro, Loja, Morona Santiago, Tungurahua, and Zamora Chinchipe. They inhabit Deciduous Costa Forest, Western Foothill Forest, Western Montane Forest, Paramo, Inter-Andean Shrub, and Eastern Montane Forest Natural Regions, between 230 and 4100 m a.s.l.

Remarks. The P. phoxocephalus species group is sister to the P. cryptomelas species group.

Figures

Figure 3. Phylogenetic relationships of Pristimantis, sugenus Huicundomantis. Maximum likelihood tree for genes 16 S, ND 1 and RAG 1. Bootstrap values (%) followed by Bayesian posterior probabilities are shown under the corresponding branches. Asterisks indicate support values of 100 (bootstrap) or 1 (posterior probabilities); missing values indicate values below 50 (bootstrap) or 0.5 (posterior probability). The collection number, identification, province and locality of the samples are shown next to each terminal; all samples are from Ecuador. Outgroup is not shown. Abbreviations: CCS = confirmed candidate species, NP = national park, PF = protected forest, UCS = unconfirmed candidate species.

Figure 7. Palmar and plantar surfaces of Pristimantis atillo sp. nov., Pristimantis chomskyi sp. nov., Pristimantis gloria sp. nov., and Pristimantis jimenezi sp. nov. Photographs of left hand and foot of the holotypes A Pristimantis atillo sp. nov.: QCAZ 42500, male B Pristimantis chomskyi sp. nov.: QCAZ 47515, male C Pristimantis gloria sp. nov.: QCAZ 57201, female D Pristimantis jimenezi sp. nov.: QCAZ 45170, male. All specimens are shown at the same scale.

Figure 9. Palmar and plantar surfaces of Pristimantis teslai sp. nov., Pristimantis torresi sp. nov., Pristimantis totoroi sp. nov., and Pristimantis verrucolatus sp. nov. Photographs of hand and foot of the holotypes of the following species: A Pristimantis teslai sp. nov.: QCAZ 46213, male B Pristimantis torresi sp. nov.: QCAZ 47342, female C Pristimantis totoroi sp. nov.: QCAZ 25105, male D Pristimantis verrucolatus sp. nov.: QCAZ 46982, male. All specimens are shown at the same scale.