Research Article |
Corresponding author: Jaime Pizarro-Araya ( japizarro@userena.cl ) Academic editor: Danilo Harms
© 2024 Andrés A. Ojanguren-Affilastro, Fermín M. Alfaro, Martín J. Ramírez, Bernardino Camousseigt-Montolivo, Jaime Pizarro-Araya.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ojanguren-Affilastro AA, Alfaro FM, Ramírez MJ, Camousseigt-Montolivo B, Pizarro-Araya J (2024) A new species of genus Urophonius Pocock, 1893 (Scorpiones, Bothriuridae), from Andean Mauline Chilean forests, with a phylogenetic re-analysis of the genus. Zoosystematics and Evolution 100(2): 469-482. https://doi.org/10.3897/zse.100.119153
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Urophonius trewanke sp. nov. is described from the Mauline Andean woods of northern Chilean Patagonia. This species belongs to the granulatus species group, which includes the most basal species within the genus. This species is only active in summer as in all species of its group. We performed a phylogenetic analysis of the genus Urophonius based on morphological characters to establish the position and relationships of the new species in the genus.
Bothriuridae, Chile, Mauline woods, new species, phylogeny
The scorpion genus Urophonius Pocock, 1893 comprises small burrowing species from southern South America. In the last two decades, there has been a remarkable increase in the knowledge of this genus; in this period, the number of described species has almost doubled (
Urophonius is remarkable amongst all known scorpion genera from temperate and cold areas because of its adaptations to low temperatures (
The first morphological phylogeny of the genus (
On the other hand, some species of the genus still retain the summer activity period, which is common to most species of the family and the order (Ojanguren-Affilastro and Kovarik 2013). These summer Urophonius species are grouped in the granulatus species group and occur exclusively in cold areas of southern South America, both in steppes and temperate woods.
In a recent comprehensive study of arthropods conducted at the “Fundo La Escuadra” (Figs
In this contribution, we describe Urophonius trewanke sp. nov. (Fig.
Taxa. The matrix in the cladistics analysis comprises a total of 21 species, 17 species of Urophonius and four outgroups. We used the same species as
Specimens examined are deposited in the following collections: Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires, Argentina (
We used a matrix, based in 114 morphological characters. The complete list of characters and the matrix are available as supplementary material online, as Suppl. materials
Analyses were made with TNT 1.5 (
All new material reported here was collected by the authors; most specimens were collected at night by UV detection. Some specimens were also collected in daytime under logs or stones. Measurements, taken using an ocular micrometer, are recorded in mm. Descriptive terminology follows
The analysis under equal and implied weights resulted in highly concordant trees. All the analyses under implied weights with concavity constant below 20 produced the resolution of Fig.
In our phylogenetic tree (Fig.
We recovered two major clades confirming the results of
On the other hand, we recovered all species with winter activity as another monophyletic group (Fig.
Chile
, Maule Region (VII), Maule Valley, Fundo La Escuadra: Holotype ♂ (MNHN 8411), Bocatoma-Ojos de Agua (35°46'06.1"S, 70°47'44.4"W), 1009 m a.s.l.; 14–17/X/2022, Pizarro-Araya, Alfaro & Calderón coll. Paratipes: 2 ♂, same data as holotype (MACN); Laguna Invernada (35°43'16.1"S, 70°47'04.9"W), 1260 m a.s.l.; 14–17/X/2022, Pizarro-Araya, Alfaro & Calderón coll. 2 ♀, 10 ♂ (
The specific epithet “trewanke” is a noun in apposition meaning scorpion in Mapungudun, the language of the Mapuche people, the original inhabitants from most parts of southern and central Chile.
Urophonius trewanke sp. nov. is most closely related to U. tregualemuensis from south-central Chile (Fig.
measurements in mm of the holotype male (MNHN) and a female paratype (MACN) of Urophonius trewanke sp. nov.
Urophonius trewanke sp. nov. | ||
Holotype ♂ | Paratype ♀ | |
Total length | 32.53 | 38.72 |
Carapace, length | 3.71 | 5.01 |
Carapace, anterior width | 2.58 | 3.39 |
Carapace, posterior width | 4.12 | 5.17 |
Mesosoma, total length | 8.88 | 9.69 |
Metasoma, total length | 19.94 | 21.02 |
Metasomal segment I, length | 1.94 | 2.58 |
Metasomal segment I, width | 2.42 | 3.23 |
Metasomal segment I, height | 2.02 | 2.58 |
Metasomal segment II, length | 2.34 | 2.82 |
Metasomal segment II, width | 2.18 | 2.91 |
Metasomal segment II, height | 1.85 | 2.50 |
Metasomal segment III, length | 2.58 | 3.47 |
Metasomal segment III, width | 2.15 | 2.74 |
Metasomal segment III, height | 1.85 | 2.42 |
Metasomal segment IV, length | 3.07 | 4.04 |
Metasomal segment IV, width | 2.02 | 2.58 |
Metasomal segment IV, height | 1.85 | 2.42 |
Metasomal segment V, length | 4.68 | 5.41 |
Metasomal segment V, width | 2.10 | 2.83 |
Metasomal segment V, height | 1.77 | 2.42 |
Telson, length | 5.33 | 5.70 |
Vesicle, width | 1.77 | 2.26 |
Vesicle, height | 1.53 | 1.82 |
Chela, length | 6.06 | 7.62 |
Chela, width | 1.85 | 1.94 |
Chela, height | 1.64 | 2.10 |
Both species can also be separated by the shape of pedipalp chela, which is stouter in U. trewanke sp. nov. (Fig.
They can also be separated by some details of the hemispermatophore; in U. tregualemuensis, the bifid lobe of the internal lobe is connected to the superior concavity of the basal portion by a thick carina (Fig.
There are also some differences in the development of the ventral carinae of metasomal segments I and II, which are clearly more developed in U. tregualemuensis (Fig.
Based on the holotype ♂ (MNHN) and the paratypes ♀ (
Total length : 30–41 mm in ♂ (n = 12; mean = 34.7); 34.5–41 mm in ♀ (n = 4; mean = 38.88).
Colour
: Base colour yellowish, with dark brown spots (Figs
Carapace
: lateral surfaces granular (♂♀), medially smooth (♀) or slightly granular (♂). Anterior margin straight. Anterior longitudinal sulcus shallow; interocular sulcus weakly developed; posterior longitudinal and lateral sulci well developed. Median ocular tubercle not very pronounced, median ocelli large, ca. one diameter apart; with one macroseta behind each eye and one microseta in front of each eye. Three pairs of small lateral ocelli on each side of carapace, posterior ocellus slightly smaller than the rest of the ocelli; anterior and median ocelli almost in the same horizontal axis, posterior ocellus situated clearly dorsal to others; lateral ocelli pattern type 3A (
Pedipalps
: Femur with DI, DE and VI carinae granular, extending the entire length of segment (Fig.
Left pedipalp. a–e, g–i. Urophonius trewanke sp. nov. a–e. Chela, ♂; a. Dorsal aspect; b. Prolateral aspect; c. Ventro-prolateral aspect; d. Ventral aspect; e. Retrolateral aspect; g. Chela ♀, prolateral aspect; h. Femur, ♂, dorsal aspect; i. Patella, ♂, retrolateral aspect; f. Urophonius tregualemuensis Cekalovic, 1981. Left pedipalp chela, ♂, prolateral aspect. Scale bars: 1 mm.
Pectines : Tooth count: 15–17 in ♂ (n = 12, median = 16) and 14–15 in ♀ (n = 4, mean = 15).
Legs : Surfaces smooth in ♀, granular in ♂. Basitarsi each with two well developed, equal length, pedal spurs. Telotarsi elongated, shallow, each with well-developed ventromedian row of hyaline setae and paired rows of ventrosubmedian spiniform setae with the following counts on each telotarsus: I: 1/1, II: 2/2, III: 5–6, IV: 6–6/6–7. The only pair of spines of telotarsus I and the first pair of spines of telotarsus II are less sclerotised than the remaining spines, the rest are well sclerotised. Ungues slightly curved, equal in length.
Tergites : Surfaces, I–VI: anterior area smooth, posterior and lateral margins finely granular; more so in ♂; VII with sparse, coarse granules in posterolateral margins, with paired dorso-submedian carinae in posterior third and paired dorso-lateral carinae in posterior two-thirds of the segment.
Sternites
Surfaces, III–VI smooth, with small elliptical spiracles; VII, surface sparsely granular, more so in ♂; in ♀ with two VM and two VL barely visible carinae in posterior third of the segment, not conspicuous in ♂ (Fig.
a–c, f–h. Urophonius trewanke sp. nov., a, b. Sternite V and metasomal segments I and II, ventral aspect a. ♂ and b. ♀; c. metasomal segment V, ♂, ventral aspect; f. Metasomal segment V, ♀, ventral aspect; g. Telson, ♂, lateral aspect; h. Telson, ♀, lateral aspect; d, e. Urophonius tregualemuensis Cekalovic, 1981, sternite V and metasomal segments I and II, ventral aspect; d. ♂; e. ♀. Scale bars: 1 mm.
Metasoma
: Metasomal segment I, dorsal surface finely granular; DL carinae granular, extending the entire length of segment with anterior and posterior granules more developed than the rest; dorso-lateral margins granular, LSM carinae represented by some tiny granules in the posterior part of the segment, LM carina with an anterior blunt small keel and a granular part extending the posterior two-thirds of the segment; LIM carinae granular, restricted to the posterior half of the segment, with one macroseta; ventral surface smooth, VL carinae extending the entire length of the segment, granular in ♀, as an elevation of the tegument in ♂, VSM carinae well developed and granular in ♀, barely visible in ♂, with two pairs of VSM macrosetae and three pairs of VL macrosetae (Fig.
Telson
: Vesicle, shallow, more lobular in ♀; ventral surface with medium sized granules in ♀, less granular in ♂; dorsal surface smooth, with (♂) or without (♀) an elliptical median well-developed depression corresponding to the telson gland. Aculeus short, shallowly curved (Fig.
Hemispermatophore
: Basal portion well developed. Distal lamina well developed, ca. 30% shorter than basal portion; distal crest almost straight, orientated almost in same direction to the posterior margin of the DL; frontal crest (distal posterior flexure) present; internal lobe with two well-developed denticles, not connected with the distal lamina (Fig.
a, c, d, Urophonius trewanke sp. nov., a. Left hemispermatophore, external aspect; c. Left hemispermatophore, lobe region, internal aspect; d. Right hemispermatophore, lobe region, internal aspect; b, e, f, Urophonius tregualemuensis Cekalovic, 1981; b. Left hemispermatophore, external aspect; e. left hemispermatophore, lobe region, internal aspect; f. Right hemispermatophore, lobe region, internal aspect. Scale bars: 1 mm.
This species has only been collected in its type locality, in the small preserved area of “Fundo La Escuadra”, 35°46'06.1"S, 70°47'44.4"W (Fig.
The area where Urophonius trewanke sp. nov. has been collected is located within the landscape of the “Estepa de los Andes Maulinos” (Mauline Andean Steppe) Botanical Formation. This distinctive formation represents the southernmost extension of the high Andean steppes. From this point, towards the south, a change in ecological conditions is perceived, characterised by an increase in precipitation and snowfall. This transition marks a natural limit for the distribution of numerous southern and boreal species (
The ecological matrix of this region is made up of various types of vegetation, which has defined its heterogeneity. Dry forests intersect with shrub steppes, creating a plant mosaic. Notable vegetation components include species such as Chuquiraga oppositifolia D.Don, Gochnatia foliorosa D.Don and Proustia cuneifolia D.Don (Asteraceae), each of which plays a role in shaping the unique habitat of Urophonius trewanke sp. nov. (Fig.
In addition, the landscape presents different herbaceous steppes, where species such as Acaena alpina Poepp ex. Walp. (Rosaceae) and Festuca acanthophylla Desv. (Poaceae) contribute to the overall floral composition. In the midst of this botanical diversity, the presence of the Cordillera cypress (Austrocedrus chilensis (D.Don) Pic-Serm. & Bizzarri.) stands out, which adds to the ecological tapestry with its characteristic shape and contributes to the general microhabitat where Urophonius trewanke sp. nov. occurs. This environment, with its varied types of vegetation and the inclusion of notable species, such as the Cypress of the mountain range, highlights the ecological importance of the “Estepa de los Andes Maulinos” Botanical Formation as a unique and valuable habitat for the diversity of arachnids, providing crucial information on the ecology and habitat preferences of Urophonius trewanke sp. nov.
The Maule Valley and its associated area along the Maule River in the Pehuenche Andean Pass, seems to constitute an area of endemism for the epigean fauna, separated from surrounding valleys by transverse mountain chains. This area is already known to harbour an endemic and highly restricted species of Anuran, Alsodes pehuenche Cei, 1976 (
In nearby localities outside the Maule Valley, but with similar habitats, U. trewanke sp. nov. is replaced by U. tregualemuensis, another species of the granulatus group, which occupies similar environments and niches as U. trewanke sp. nov., but has a wider distribution in south central Chile (Fig.
Urophonius trewanke sp. nov. has been collected in sympatry with an undescribed species of Brachistosternus, which also seems to be endemic of the Pehuenche Pass (Ojanguren-Affilastro et al. in prep.).
The description of U. trewanke sp. nov., a rare species belonging to the granulatus group from southern Chile, supports our previous hypothesis that the few species of Urophonius with a summer activity period, are restricted to central and southern Chile and to the colder areas of southern South America, whereas the species with a winter activity period are more widely distributed (
Urophonius trewanke sp. nov. has been collected in “Fundo La Escuadra”, meaning La Escuadra farm or ranch, a small preserved area currently under management by ENEL (“Empresa Nacional de Energía Eléctrica” or National Electric Energy Company). This area has been part of the “Cipreses” operational system of the Chilean Electric National System since 1955 and has, therefore, had highly restricted access for the last seventy years. This inaccessibility has resulted in an unintended, but yet remarkable, degree of preservation. This area is placed in the partially isolated Maule Valley which has proved to harbour an exceptional number of endemics, but lacks any kind of formal protection. We hope that the description of the actual diversity of this area could shed light on the need to preserve this important biological resource and lead to future preservation of the endemic species of the Maule Valley.
Our special thanks to the staff of the Los Cipreses hydroelectric power plant, especially to Christian Cartes for his help in logistics (La Escuadra, ENEL). Project funded by Environment & Permitting - HSEQ, Enel Green Power & Thermal Generation (ENEL). J.P-A thanks the Academic Excellence Scholarship (B134) from the Academic Vice-Rector’s Office, Research and Postgraduate Studies of the Universidad Santo Tomás, Santiago, Chile and ANID doctoral fellowship 2024. F.M.A. thanks the ANID doctoral fellowship 2023-21230592. We are indebted to the four reviewers of the manuscript Stepahnie Loria, Oscar Francke, Edmundo Gonzalez-Santillán and an anonymous reviewer, as well as to the editor Danilo Harms, for their helpful comments on the first version of the manuscript.
Studied material Urophonius phylogeny
Data type: docx
Characters Urophonius phylogeny
Data type: doc
Matrix Urophonius phylogeny
Data type: txt