Research Article |
Corresponding author: Regiane Saturnino ( sf.regiane@gmail.com ) Academic editor: Ricardo Pinto-da-Rocha
© 2020 Paulo Pantoja, Marcos Drago-Bisneto, Regiane Saturnino.
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:
Pantoja P, Drago-Bisneto M, Saturnino R (2020) Updates on Berlandiella (Araneae: Philodromidae): a new species, description of the male of B. querencia and new diagnosis for the genus. Zoologia 37: 1-13. https://doi.org/10.3897/zoologia.37.e37384
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Berlandiella Mello-Leitão, 1929 is currently composed of six Neotropical species, of which Berlandiella querencia Lise & Silva, 2011 is known only from female specimens; the other species of the genus were described based on both males and females. In this paper, we describe and illustrate Berlandiella zabele sp. nov., based on a few individuals collected in Sete Cidades National Park, Piracuruca and Brasileira, state of Piauí, Brazil. We illustrate and describe the previously unknown male of B. querencia, based on a specimen collected from Reserva Mocambo, Belém, state of Pará, Brazil. The taxa described herein have scopula in the tarsi and metatarsi, and the males have a cymbial process, characters recorded for the first time for the genus. Additionally, we present an updated diagnosis for Berlandiella.
Neotropical, Northern, spiders, taxonomy
Philodromidae
Thorell, 1870 currently comprises 31 genera, of which 18 occur in the Neotropics (
According to
Based on the revision provided by
The specimens were analyzed with a Leica MZ16 stereomicroscope and the illustrations were made with the assistance of a camera lucida attached to a stereomicroscope. Descriptions follow
Berlandiella Mello-Leitão, 1929: 107, 122, 260, 264, 265 (type species Berlandiella insignis Mello-Leitão, 1929, by original designation). Lise & Silva, 2011: 350–373.
Berlandiella is similar to Paracleocnemis, Procleocnemis Mello-Leitão, 1929 and Metacleocnemis Mello-Leitão, 1929 due to general morphology, however, differs from Metacleocnemis for having little evident thoracic fovea; from Procleocnemis due to smaller and closer PME relative to the PLE, oval abdomen and legs III-IV of similar length to legs I-II; they differ from Paracleocnemis in having a longer than wide cephalothorax and posterior eye row more recurved than the anterior. Berlandiella is distinguished from the other neotropical genera of Philodromidae by the following set of characters: carapace covered by erect and robust bristles (Figs
Holotype. BRAZIL, 1 female from Piracuruca, Parque Nacional de Sete Cidades, 04°05’55.4”S, 41°42’33.2”W, 24 Jun. 2005, LS Carvalho leg. (MPEG.ARA 34482). Paratypes. BRAZIL, 1 male from Brasileira, Parque Nacional de Sete Cidades, 04°08’08.2”S, 41°43’08.4”W, 29 Jan. 2007, LS Carvalho et al. leg. (MPEG.ARA 34480); 1 male, same collection data as for preceding, 04°05’39.9”S, 41°43’53.3”W, 26 Jan. 2007, LS Carvalho et al. leg. (CHNUFPI 3398); 1 male, same collection data as for preceding, 24 Jan. 2007, LS Carvalho et al. leg. (MPEG.ARA 15585); 1 female, same collection data as for preceding, 22 Jan. 2007, LS Carvalho leg. (MPEG.ARA 15579).
The female of Berlandiella zabele sp. nov. is similar to B. magna and B. querencia by the long and ellipsoid spermathecae (Figs
Female (holotype – MPEG.ARA 34482). Carapace slightly longer than wide, wider at the level of legs III, narrowed anteriorly, with many sockets of bristles; median region yellow with a brown central spot and wide dark brown bands on each side, formed by a tangle of dark spots. Clypeus vertical with many sockets of bristles, with a median brown spot and irregular spots on lateral. Chelicerae yellow with two teeth on the promargin of the fang furrow, with a large dark brown spot on the anterior face. Labium orange with rounded apex, reaching almost to the middle of the endites. Endites pale yellow, excavated diagonally. Sternum pale yellow. Legs covered by various types of setae and with many robust macrosetae on tibiae and metatarsi. Left and right tibiae I with 0–2–2–0 ventral macrosetae, left metatarsus I with 2–2–0 and right metatarsus I with 2–2–1 ventral macrosetae. Left and right tibia II with 0–2–2–0 ventral macrosetae and right metatarsus II with 2–2–1 ventral macrosetae. Tarsi and metatarsi with scopula. Long and thin trichobothria in tarsi and metatarsi, with distal and proximal plate limit well differentiated; the distal margin of the trichobothrial hood is well defined; hood entire, differentiated with transverse ridges; alveolus notched; distal plate smooth, slightly elevated. Femura yellow with brown spots, more abundant in prolateral sides of femora I-III, and a small white spot at the apex; patellae I-III yellow with brown spots; patella IV yellow; tibiae and metatarsi I-III yellow, shaded with brown, but tibia and metatarsus III darker; tibia IV yellow with a basal dark spot on prolateral side; tarsi I-IV yellow, with a dark ring at their bases. Tarsal claw pectinate and conspicuous with claw apex tooth curved and seven short and robust teeth. Palp yellow with brown spots. Abdomen yellow, longer than wide, densely covered by setae sockets, the dorsum covered by a wide brown spot with two longitudinal yellow bands around the heart mark; laterals yellow shaded with brown and the venter yellow stippled with brown. Triangular epigynal groove, shallow, demarcated by sclerotized edges, small and widely spaced copulatory openings, elliptical and elongated spermathecae, thickly covered with punctuations.
Total length 3.72. Carapace 1.85 long, 1.62 wide, 0.59 high. Chelicerae 0.57 long, 0.33 wide. Labium 0.30 long, 0.23 wide. Endites 0.43 long, 0.20 wide. Sternum 0.96 long, 0.89 wide. Clypeus 0.27 high. Abdomen 2.58 long, 1.66 wide, 1.22 high. Leg: I. femur 1.54; patella 0.72; tibia 1.27; metatarsus 1.04; tarsus 0.50; total length 5.07; II. 1.68; 0.72; 1.39; 1.13; 0.76; 5.68; III. 1.42; 0.63; 1.07; 0.97; 0.51; 4.60; IV. 1.17; 0.63; 1.12; 1.18; 0.61; 4.71. Leg formula II-I-IV-III. Eye diameters and eye interdistances. AME 0.08, ALE 0.09, PME 0.07, PLE 0.08, AME-AME 0.15, AME-ALE 0.08, PME-PME 0.22, PME-PLE 0.27. MOQ 0.25 long in frontal view, 0.22 in dorsal view, anterior width 0.30, posterior width 0.36.
(paratype – MPEG.ARA 34480). Shape and bristles of carapace (Figs
Total length 3.72. Carapace 1.49 long, 1.51 wide, 0.51 high. Chelicerae 0.44 long, 0.25 wide. Labium 0.17 long, 0.22 wide. Endites 0.37 long, 0.17 wide. Sternum 0.82 long, 0.75 wide. Clypeus 0.17 high. Abdomen 1.41 long, 1.00 wide, 0.66 high. Legs: I. femur 1.24; patella 0.59; tibia 1.03; metatarsus 0.89; tarsus 0.63; total length 4.38; II. 1.50; 0.60; 1.25; 1.00; 0.68; 5,03; III. 1.24; 0.50; 0.97; 0.76; 0.50; 3.97; IV. 1.35; 0.54; 1.08; 1.05; 0.51; 4.53. Leg formula: II-IV-I-III. Eye diameters and eye interdistances. AME 0.08, ALE 0.07, PME 0.06, PLE 0.10, AME-AME 0.12, AME-ALE 0.05, PME-PME 0.20, PME-PLE 0.18. MOQ 0.21 long in frontal view, 0.19 in dorsal view, anterior width 0.24, posterior 0.32.
BRAZIL. Piauí: Piracuruca, Parque Nacional de Sete Cidades, 04°05’39.9”S, 41°43’53.3”W, 02 Feb. 2007, LS Carvalho et al. leg. (1 female, MPEG.ARA 15576); same collection data as for preceding, 24 May. 2007, LS Carvalho et al. leg. (1 female, MPEG.ARA 15577); same collection data as for preceding, 22 Jan. 2007, LS Carvalho leg. (2 males, MPEG.ARA 15578); same collection data as for preceding, 02 Dec. 2006, NF Lo-Man-Hung leg. (1 female, MPEG.ARA 15580); same collection data as for preceding, 24 Jan. 2007, LS Carvalho leg. (2 males, MPEG.ARA 15581); same collection data as for preceding, 26 Jan. 2007, LS Carvalho et al. leg. (1 male, MPEG.ARA 15582); same collection data as for preceding, 02 Feb. 2007, LS Carvalho et al. leg. (1 male, MPEG.ARA 15583); same collection data as for preceding, 04°5’57.5”S, 41°43’0.7”W, 22 Jan. 2007, FM Oliveira-Neto leg. (1 male, MPEG.ARA 34478); Brasileira, 04°08’08.2”S, 41°43’08.4”W, 29 Jan. 2007, LS Carvalho et al. leg. (1 male, MPEG.ARA 34479); same collection data as for preceding, 28 Jan. 2007, LS Carvalho leg. (1 male, MPEG.ARA 34481).
Known only from the type locality (Fig.
The specific epithet refers to a mythological character known by indigenous people of Piauí. According to indigenous mythology, Zabelê, a member of the Amanajós tribe, fell in love with Metara, a member of an enemy tribe. When the romance of Zabelê and Metara was discovered, tribal warfare resulted in the death of both lovers. The god Tupã pitied the two lovers and so transformed them into birds that fly together for eternity.
The general color, legs especially, vary in the abundance of brown spots and some specimens may be darker or lighter than the paratype. The pattern of ventral macrosetae of the legs also varies between individuals.
Genitalia of Berlandiella zabele sp. nov.: (6) epigynum ventral; (7) epigynum dorsal; (8) palp, ventral; (9) palp, retrolateral; (10) palp, dorsal. CD: copulatory ducts, CO: copulatory opening, Co: conductor, E: embolus, EG: epigynal groove, FD: fertilization duct, GH: glandular head, CP: cymbial process, RTA: retrolateral tibial apophysis, S: spermatheca, VTA: ventral tibial apophysis. Scale bars: 0.3 mm.
Palp of Berlandiella zabele sp. nov.: (11) ventral; (12) detail of apical portion of tegulum; (13) dorsal (arrow indicates scopula in apex of cymbium); (14) scopula in cymbium (indicated by arrow); (15) detail of CP (indicated by arrow) and RTA; (16) detail showing excavation in the RTA. CP: cymbial process, RTA: retrolateral tibial apophysis. Scale bars: 11 = 200 µm, 12, 15, 16 = 50 µm, 13 = 150 µm, 14 = 20 µm.
Carapace of Berlandiella zabele sp. nov. male: (17) dorsal; (18) anterodorsal; (19) detail of tegument with many sockets of setae (indicated by arrows); (20) strong setae in the cephalic region; (21) hairs; (22) scales in the lateral of carapace. Scale bars: 7, 20 = 30 µm, 18 = 300 µm, 19, 22 = 50 µm, 20 = 30 µm, 21 = 10µm.
Berlandiella querencia Lise & Silva, 2011: 368, figs 1, 100–111 (holotype from Querência, Mato Grosso, Brazil, deposited in MPEG 17571; examined).
The male of B. querencia differs from the other species of the genus by having a sinuous CP, directed to the apex of the RTA (Figs
Male (MPEG.ARA 34477). Carapace slightly longer than wide, wider on the level of legs III, narrowed in front, with many sockets of bristles (Figs
Total length 2.84. Carapace 1.46 long, 1.03 wide, 0.39 high. Chelicerae 0.43 long, 0.25 wide. Labium 0.16 long, 0.23 wide. Endites 0.34 long, 0.17 wide. Sternum 0.81 long, 0.73 wide. Clypeus 0.16 high. Abdomen 1.38 long, 1.00 wide, 0.57 high. Legs: I. femur 1.06; patella 0.51; tibia 0.96; metatarsus 0.72; tarsus 0.52; total 3.77; II. 1.25; 0.53; 1.08; 0.84; 0.57; 4.26; III. 1.23; 0.44; 0.83; 0.71; 0.50; 3.71; IV. 1.22; 0.46; 1.00; 0.92; 0.47; 4.07. Leg formula II-I-IV-III. Eye diameters and eye interdistances: AME 0.08, ALE 0.7, PME 0.06, PLE 0.10, AME-AME 0.8, AME-ALE 0.05, PME-PME 0.15, PME-PLE 0.14, MOQ 0.19 long in frontal view, 0.19 in dorsal view, anterior width 0.25, posterior width 0.28.
See diagnosis and description in
BRAZIL. Pará: Belém, Reserva Mocambo, 01°26’26.7”S, 48°24’40.5”W, 10 Jul. 2003, JAP Barreiros and DR Santos de Sousa leg. (1 male, MPEG.ARA 34477); same collection data as for preceding, Museu Paraense Emílio Goeldi – Campus de Pesquisa, 01°27’04.44”S, 48°26’39.32”W, 2011, (1 female, MPEG.ARA 33972); same collection data as for preceding, Oct. 2011 (1 female, MPEG.ARA 33973); same collection data as for preceding, 12 May 2018, P Pantoja leg. (1 male, MPEG.ARA 35228); Mato Grosso: Querência, Fazenda Tanguro, 12º55’53.0”S, 52º26’49.1”W, 13 Jun. 2006, DF Candiani and NF Lo-Man-Hung leg. (1 female, MPEG.ARA 16716).
Known from Brazil (Mato Grosso and Pará) (Fig.
Leg I of Berlandiella zabele sp. nov. male: (23) tarsus and apex of metatarsus with scopula (indicated by arrows); (24) different types of seta in the metatarsus (smaller rectangle show tenent barbs of scopula); (25) claw and claw tufts; (26) detail of tenet barbs of claw tufts. Scale bars: 23 = 150 µm, 24, 25 = 20 µm, 26 = 5 µm.
Leg I of Berlandiella querencia: (34, 36, 38) female; (35, 37, 39) male; (34-35) metatarsus and apex of tarsus (arrows to scopular setae); (36) detail of setae in tarsus; (37) scopula, smaller rectangle show tenent barbs of scopular seta; (38-39) claw and claw tufts. Scale bars: 34, 35 = 150 µm, 36, 39 = 50 µm, 37, 38 = 20 µm.
Setae: (40) setae on the side tibia of B. querencia male; (41) pair of macrosetae in the venter of tibia I of B. zabele male; (42) trichobothrium on the dorsum of tarsus I of B. zabele male; (43) strong bristles on the side of the metatarsus of B. querencia female.Scale bars: 40, 43 = 50 µm, 41 = 100 µm, 42 = 20 µm.
We present an updated diagnosis of Berlandiella based on the revision carried out by
Scopula seem to be present only in sexually mature individuals, since we analyzed a juvenile of B. querencia without this structure. The scopular setae appeared in the anterior legs after ecdysis, when the specimen reached sexual maturity, becoming an adult male (MPEG.ARA 35228). Although we have not examined all species of the genus, it is clear that scopula should not be used to diagnose Berlandiella because the presence/absence of this structure varies among specimens of the same species and with degree of maturity.
We also removed from the diagnosis any mention about the height of the carapace, as well as the presence of dense claw tufts. In fact, some females of Berlandiella may have a relatively high carapace, and even males have convex carapace, although it is lower than females. As the understanding about the height of the carapace is subjective, we decided to remove this character from the diagnosis. The use of claw tufts in the genus diagnosis is not informative, since these structures are present in many others genera of Philodromidae.
Since we documented the presence of the scopula in Berlandiella specimens, the absence of this character used to separated Berlandiella from Cleocnemis become invalid and was removed from the diagnosis. As consequence we could not establish a clear limit between Berlandiella and Cleocnemis. The original description of the type species, C. heteropoda Simon, 1886, as well as the redescription by
The limits between Berlandiella and Cleocnemis should be established based on the detailed analysis of the Cleocnemis type species, as well as a phylogenetic analysis to understand the relationship between the two genera. Both are beyond of the scope of this paper, although we have the concern of not describing a new taxa which can become a synonym of a species already described and allocated in Cleocnemis. Then, to make sure that B. zabele has not been already described under Cleocnemis we checked the type localities, original descriptions and illustrations for all 14 known species of this genus as follow summarized: C. bryantae (Gertsch, 1933), C. lanceolata Mello-Leitão, 1929, C. nigra Mello-Leitão, 1943, C. paraguensis (Gertsch, 1933), C. spinosa Mello-Leitão 1947, C. taquarae (Keyserling, 1891) and C. xenotypa Mello-Leitão, 1929 differ from B. zabele based on the genitalia. Cleocnemis moschata Mello-Leitão, 1943 and C. rosea Mello-Leitão, 1944 has several ventral macrosetae in the legs I-II and B. zabele has only 2–2 ventral macrosetae in this legs. Cleocnemis mutilata (Mello-Leitão, 1917), C. punctalata (Taczanowski, 1872), C. rudolphi Mello-Leitão, 1943 and C. serrana Mello-Leitão, 1929 has several somatic characters different from B. zabele, especially legs with unequal length. Cleocnemis nigra and C. rudolphi were registered from the state of Paraíba, relatively close to the type locality of B. zabele, however, differs from this species by the previously mentioned characters. The description of C. heteropoda is similar to B. zabele based on somatic characters, however, C. heteropoda occurs in Rio de Janeiro, relatively far from the type locality of B. zabele. Additionally, the heteropoda epithet refers to unequal legs, as indicated by
About the new characters described for Berlandiella here, males of B. zabele sp. nov. and B. querencia present a cymbial process, which differs from all other males of the genus. A similar structure occurs in several species of Philodromus Walckenaer, 1826 and has been called the paracymbial lamella (
Structures that emerge from the margin of the cymbium have been defined several times in the literature as paracymbium (see
Finally, despite the difficulty in distinguishing Cleocnemis from Berlandiella, this latter is a revised genus and presents uniformity in the diagnostic characters present in the type-species and in the others (see diagnosis above). For this reason, and due to the similarity between B. zabele and other Berlandiella species, we decided to allocate the species in this genus.
We would like to thank the Museu Paraense Emílio Goeldi (MPEG) by providing the laboratory structure to carry out this work, especially Alexandre Bonaldo, responsible for the Laboratório de Aracnologia of the MPEG. We also thank Hilton Tulio Costi and Laura Miglio for providing the use of the SEM lab of MPEG and for all support given to the authors. We are very grateful to the reviewers and the editor for their careful reading of the text, for all valuable comments and suggestions, which were very helpful in improving the paper: Cristina Anne Rheims, Martín Ramírez, Ricardo Pinto-da-Rocha and anonymous. Finally, we greatly thank André Prado for our discussions about Cleocnemis taxonomy. The authors were supported by the following grants: PP – Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, 130702/2019-6) and Programa de Pós-graduação em Zoologia – UFPA/MPEG; MD – Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, 88882.442371/2019-01) and Programa de Pós-graduação em Biodiversidade e Evolução – MPEG; RS – PandD ANEEL/CELPA # PD-0371-0017/2013. The laboratory facilities at Museu Paraense Emílio Goeldi were provided by Project #3362 CELPA/FADESP Monitoramento (REF.: 061/2013), also coded at Agência Nacional de Energia Elétrica (ANEEL) as PD-0371-0017/2013, entitled “Monitoramento dos possíveis impactos da linha de transmissão do Marajó sobre a fauna.” The last version of this paper was written when RS was supported by the Project #1199/FSADU (Fundação Sousândrade de Apoio ao Desenvolvimento da UFMA), entitled “Desenvolvimento experimental de metodologia para a detecção e redução de fatores de vulnerabilidade da vegetação que causam interrupções no fornecimento de energia”, funded by Companhia Energética do Maranhão-CEMAR.