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New species of Furculanurida (Collembola, Neanuridae, Pseudachorutinae) from the Luquillo Mountains, Puerto Rico
expand article infoClaudia M. Ospina-Sánchez, José G. Palacios-Vargas§, Grizelle González
‡ USDA-FS, International Institute of Tropical Forestry, Río Piedras, United States of America
§ Universidad Nacional Autónoma de México, México, Mexico
Open Access

Abstract

A new species of Furculanurida is described and illustrated. Furculanurida bistribus sp. nov. differs from other species of the genus by the presence of three eyes, three setae on the dens, and the white and purple coloration pattern. A key for identification of the world species of the genus is included.

Keywords

Island, Luquillo Experimental Forest, subtropical forest, taxonomy

Introduction

In Puerto Rico, most studies of arthropod community dynamics have been done in the Luquillo Mountains. Designated as a US Experimental Forest in 1956, it became part of the International Network of Biosphere Reserves in 1976 (González and Barberena 2018; Quiñones et al. 2018; Richardson 1999). Four easily distinguishable forest types are dominated by an assortment of distinctive tree species. The Tabonuco forest (Dacryodes excelsa Vahl) occupies areas below 600 m, in the mid-elevation zone. Palo Colorado forest (Cyrilla racemiflora L.) occurs in areas above the cloud condensation level from 600 to 900 m a.s.l. The Elfin forest (dominant tree Tabebuia rigida Urban), with stunted vegetation and waterlogged anoxic soils, is located only on the highest peaks above 900 m. Palm forests (Prestoea montana (R. Grah.) Nichols) occur at all elevations, predominantly on windward slopes, in wet gullies, and in stream valleys (Gould et al. 2006). These forests represent subtropical wet and subtropical rain forest life zones in Puerto Rico (Ewel and Whitmore 1973).

In most studies of litter and soil fauna in the Luquillo Experimental Forests (hereafter LEF), Collembola are an important group because of their numerical dominance, combined with their key responses to changes in disturbance, altitude, and vegetation type (Schowalter and Ganio 1999; Schowalter et al. 2003; Richardson et al. 2005; Richardson et al. 2010; Schowalter et al. 2014). In Puerto Rico, Collembola are well known in comparison to other groups of soil arthropods. However, not all Collembola species from LEF have been identified.

In a recent survey between 2014 and 2015, we identified 16 families (sensu Deharveng 2004; Soto-Adames et al. 2008; Bellinger et al. 1996–2019), 37 genera, and probably 60 species, of which 15 are new. As a result of this survey, the inventory of Collembola in the LEF increased to 44 genera and 70 species. The purpose of this paper is to describe a new species of Furculanurida Massoud, 1967, a genus not previously reported from Puerto Rico.

The genus Furculanurida was created to relocate Micranurida africana Massoud, 1963 because of the development of its furcula (Massoud 1967). The main characters of this genus have been discussed previously (Palacios-Vargas and Gao 2009; Queiroz and Fernandes 2011; Zon et al. 2014, Neves et al., 2019), and some species are dubiously placed within Furculanurida (Table 1). To date, the genus has 16 nominal species distributed in the Neotropical, Ethiopian, and Nearctic regions, with seven, six, and one described species, respectively. Reported in the Neotropical Region are F. arawakensis Thibaud & Massoud, 1983 from the Lesser Antilles; F. guatemalensis Palacios-Vargas & Gao, 2009 and F. septemoculata Palacios-Vargas & Gao, 2009 from Guatemala; F. nessimiani Fernandes & Mendonça 2002, F. belemensis Arlé & Rufino, 1976, F. goeldiana Arlé & Rufino, 1976, and F. tropicalia Queiroz & Fernandes, 2011 from Brazil; and F. longisensillata Najt, Thibaud & Weiner, 1990 from French Guiana.

Furculanurida species with disputed generic placement and their taxonomic history.

Species Original genus Generic placement + Character ++
africana (Massoud, Z, 1963) Micranurida Furculanurida (type species) Massoud 1967 Furcula developed
arlei Thibaud & Massoud, 1980 Furculanurida Stachorutes Weiner and Najt (1998) Presence of a microsensillum on Ant IV, mandible with only 2 teeth and a reduced furcula with very small mucro
Furculanurida Thibaud and Palacios–Vargas (2000) Presence of Ant. IV with 8 thick sensilla on antennal segment IV and long sensorial setae on the body and a small tooth in the unguis
ashrafi (Yosii, 1966) Micranurida Stachorutes Deharveng and Lienhard (1983) Furcula reduced
Furculanurida Thibaud and Palacios–Vargas 2000 Presence of four teeth in the mandible
furculata (Salmon, 1956) Kenyura Furculanurida Massoud (1967) Furcula developed and post-antennal organ present
perplexa (Salmon, 1956) Hypanurida Furculanurida Massoud (1967) Reduced furcula
Hypanurida Thibaud and Palacios–Vargas (2000) Reduced furcula and 3–4 setae in dens

Initially, the genus Furculanurida was established using the combination of the following characters: postantennal organ (PAO) present, eyes absent, maxilla styliform, and furcula present (Massoud 1967). According to Queiroz and Fernandes (2011) species of the genus have Ant IV with a trilobed apical bulb, dorsolateral microsensillum present or absent, 4–7 sensilla, and long ordinary setae; PAO circular or elliptical with 4–22 vesicles; eyes from zero to eight per side; mandible with 2–10 teeth; maxilla styliform with two fused lamellae; tenent hair on tibiotarsi acuminated; ventral tube with 3 or 4 setae on each side; tenaculum with 2 or 3 teeth on each ramus; furcula complete, with well-developed dens and mucro; dens with 5 or 6 setae on each side; mucro separated from dens and with two tapering lamellae, sensilla on body always long. However, the more recently described Furculanurida species include specimens without the full development of the furcula (Zon et al. 2014). Zon et al. concluded that Furculanurida can only be separated from Pseudachorutes Tullberg, 1871 by a conditional combination of characters, i.e., “eyes less than 8+8, or, when 8+8, microchaeta ms absent on Ant. IV”. Similarly, differences with Stachorutes Dallai, 1973 would be “mucrodens complete, or, if mucro absent, ms absent on Ant. IV” (Zon et al. 2014: 496). Therefore, these genera need an extensive revision to clarify the morphological differences between them.

Materials and methods

Abbreviations

a.s.l above sea level

Abd abdominal segment

Ant antennal segment

Cx coxa

Fe femur

M long macroseta

ms microsensillum

mi microseta

PAO Postantennal organ

S sensillum

Sgd dorsal guard sensillum of Ant. III

Sgv ventral guard sensillum of Ant. III

ss sensorial seta

Th thoracic segment

Ti tibiotarsus

Tr trochanter

VT Ventral Tube

The material used to describe the species was collected during the Collembola microhabitats project at the Luquillo Mountains, as part of a survey conducted in three forest types. Collembola were extracted using Berlese-Tullgren funnels into 95% ethanol. They were cleared using Nesbitt solution and fixed on slides using Mac André II solution (Mari Mutt 1979). The slides were then dried in a slide warmer at 45–50 °C for seven days. Finally, each specimen was labeled with its collecting data. Specimens were examined with a Leica DM500 phase-contrast microscope. The drawings were made with the aid of a drawing tube. All the type material is deposited at International Institute for Tropical Forestry laboratory.

Taxonomy

Furculanurida bistribus sp. nov.

Figures 1, 2, 3–8, 9–13, Tables 1, 2

Type material

Holotype (female on slide) and 8 paratypes (2 males, 4 females, and 2 juveniles, each one on slides). Puerto Rico, Luquillo, Luquillo Mountains, Pico del Este, 18°16’17”N; 65°45’40”W; 987.6 m a.s.l.; ex mosses, Tabebuia rigida forest type, from leaf litter and epiphytes, 04 Nov 2014, leg. CM Ospina.

Other material

2 females on slides, Puerto Rico, Luquillo, Luquillo Mountains, Pico del Este, Tabebuia rigida forest type, epiphyte, 987.6 m a.s.l., 19 May 2015, leg. CM Ospina. 1 female on slide, Puerto Rico, Luquillo, Luquillo Mountains, Pico del Este, Tabebuia rigida forest type, epiphyte, 987.6 m a.s.l., 11 Feb 2015, leg. CM Ospina. 1 juvenile on slide Puerto Rico, Luquillo, Luquillo Mountains, Pico El Yunque, Tabebuia rigida forest type, leaf litter, 1044.8 m a.s.l., 4 Nov 2014, leg. CM Ospina. 1 female on slide Puerto Rico, Luquillo, Luquillo Mountains, Pico El Yunque, Tabebuia rigida forest type, leaf litter, 1044.8 m a.s.l., 19 May 2015, leg. CM Ospina.

Diagnosis

Eyes 3+3 eyes. Post antennal organ in rosette with 5 or 6 vesicles. Ant IV with six sensilla. Seta a0 on head absent. Mandible with four teeth. Dens with three setae. Unguis without internal tooth.

Description

Average body length: adults 1009 µm (n = 5); juveniles 847 µm (n = 2). Specimens in ethanol with antenna and abdomen evenly grey, ocular patch dark; head, legs III, and furcula light grey; thorax, legs I and II white to light purple (Fig. 1). Granulation coarse. Body setae comprising short, smooth and thin setae, and long and smooth sensorial setae.

Figure 1. 

Furculanurida bistribus sp. nov. Specimen in ethanol 96%.

Head: antenna shorter (0.6) than head diagonal. Ant III and IV fused dorsally, ventral separation clearly marked. Ant IV dorsally with trilobed apical vesicle, six subcylindrical thin sensilla and 14 long setae; subapical organite present; dorsoexternal microsensillum absent (Fig. 3); no sensorial field ventrally on Ant IV (Fig. 4); Ant III sense organ with two small internal slightly bent sensilla, two subcylindrical guard sensilla, Sgv larger than Sgd, ventral microsensillum present (Figs 3, 5); Sgd apically displaced, towards Ant IV, aligned to S2 and S3; Ant II with 10 setae; Ant I with seven setae (Fig. 3). Eyes 3+3 on a pigmented eyepatch; PAO with five or six vesicles disposed as a rosette (Fig. 2). Head dorsal chaetotaxy as in Figure 2; seta a0 absent; d row with 4 setae, sd row with three setae; setae Oc 1–3 present; c setae absent; p1–3 setae present. Buccal cone elongate, labium with complete chaetotaxy, A to G setae, C and D apically displaced (Fig. 6). Pre-labral/Labral chaetotaxy 4/2322 (Fig. 7). Mandible with four teeth, two apical short and subequal, one medial and one basal large and subequal; maxilla styliform with one lamella (Fig. 8).

Figure 2. 

Furculanurida bistribus sp. nov. Dorsal chaetotaxy of body (female).

Figures 3–8. 

Furculanurida bistribus sp. nov. 3 Ant I–IV, dorsal view 4 Ant IV and III, ventral view 5 Sensorial Organ in Ant. III 6 Labium 7 Labrum 8 Maxilla and mandible.

Dorsal chaetotaxy: ordinary setae smooth, distributed as in Figure 2. Th I with 2+2 setae; Th II and III with one dorsolateral seta posteriorly displaced. Sensory setae (s) clearly differentiated, in position p3 and p6 in Th II and III and in position p3 in Abd I–V; S-chaetotaxy formula = 022/11111.

Legs chaetotaxy: subcoxae 1, two; subcoxae 2, one; Cx, three; Tr, four; Fe, 10, and Ti 19 setae (Fig. 9). Seta M present between B4 and B5 without displacement. Tenent hair acuminate. Claws without teeth, unguis larger than Ti (ratio tibiotarsus: unguis = 1:1.2:); unguiculus absent.

Ventral chaetotaxy as in Figure 10. VT with 3+3 setae; tenaculum with 3+3 teeth and without setae; furcula well developed, manubrium with 14 setae, dens with three setae, mucro straight with a broad hook-like apex (Fig. 11). Ratio mucro: dens = 1: 1.3. Female genital plate with 2+2 pregenital setae, four circumgenital setae, and 1+1 eugenital setae (Fig. 12). Male genital plate with 2+2 pregenital setae, ten circumgenital setae, and 4+4 eugenital setae (Fig. 13).

Figures 9–13. 

Furculanurida bistribus sp. nov. 9 Tibiotarsus II 10 Ventral quetotaxy 11 Manubrium and furcula 12 Female genital plate 13 Male genital plate.

Etymology

Bistribus, Latin for two times three, in reference to the presence of 3+3 eyes and 3+3 setae on dens, diagnostic characters of the species.

Distribution

This species is only the Furculanurida known from the Luquillo Mountains in the Tabebuia rigida forest type, on Pico del Este 18°16’17”N; 65°45’40”W; 987.6 m a.s.l. and Pico El Yunque 18°18’37”N; 65°47’26”W; 1044.8 m a.s.l.

Ecology

Furculanurida bistribus sp. nov. was extracted from leaf litter and mosses in both dry and rainy seasons during November 2014, and May and August 2015.

Identification key to the species of Furculanurida Massoud, 1967

1 Eyes absent 2
Eyes present 4
2 Mucro developed, PAO with 8–10 vesicles 3
Mucro absent, PAO with 13–16 vesicles F. emucronata Zon, Tano & Deharveng, 2014
3 Internal tooth on unguis absent F. africana Massoud, 1963
Internal tooth on unguis present F. boiuna Neves, Mendonça & Queiroz, 2019
4 Eyes 8+8 F. tropicalia Queiroz & Fernandes, 2011
Eyes 7+7 or less 5
5 Tenaculum with 2+2 teeth 6
Tenaculum with 3+3 teeth 7
6 PAO with 11 vesicles F. duodecimoculata Thibaud & Massoud, 1980
PAO with 6 vesicles F. nessimiani Fernandes & Mendonça, 2002
7 Setae on dens 3, internal tooth onunguis absent F. bistribus sp. nov.
Setae on dens 5 or 6, internal tooth on unguis present 8
8 Setae on dens 5 F. langdoni Bernard, 2007
Setae on dens 6 9
9 Mandible with 10 teeth F. longisensillata Najt, Thibaud & Weiner, 1990
Mandible with 7 teeth or less 10
10 Ant IV with 7 sensilla, PAO with 4 vesicles, eyes 2+2 F. furculata Salmon, 1956
Ant IV with 6 sensilla, PAO with more than 4 vesicles 11
11 Eyes 4+4 or less, mandible with 7 teeth F. arawakensis Thibaud & Massoud, 1983
Eyes 5+5 or 7+7, mandible with less than 7 teeth 12
12 Eyes 7+7 13
Eyes 5+5 14
13 PAO with 7-10 vesicles, mandible with 4 teeth F. goeldiana Arlé & Rufino, 1976
PAO with 15 vesicles, mandible with 2 teeth F. septemoculata Palacios-Vargas & Gao, 2009
14 PAO with 8 or 9 vesicles in a circular form F. belemensis Arlé & Rufino, 1976
PAO with 9 or more vesicles in an elliptical form 15
15 Setae on ventral tube 3+3, PAO with 9 or 10 vesicles F. grandcolasorum Weiner & Najt, 1998
Setae on ventral tube 4+4, PAO with 15 vesicles F. guatemalensis Palacios-Vargas & Gao, 2009

Discussion

Furculanurida bistribus sp. nov. is placed in Furculanurida because many of its characters are similar with those of the other species of that genus, and it matches the current genus diagnosis: apical bulb trilobed, long setae present on Ant IV, maxilla styliform, furcula fully developed, and ordinary setae on the body short but sensory setae long (Queiroz and Fernandes 2011). Although the presence of one tooth on the unguis is observed in most Furculanurida, there are two exceptions where the unguis is toothless: F. africana and the new species described here. In any case, the inner tooth on the claw is usually considered as a specific, not a generic character (Zon et al. 2014). The antennal chaetotaxy is also useful to characterize the genus; the new species has antennal characters of Furculanurida: apical vesicle trilobed, microsensillum absent, and presence of six S-sensilla and long setae on Ant IV (Queiroz and Fernandes 2011). The low number of dental setae is a character shared with Hypanurida perplexa Salmon, 1956, but the position of this species is controversial because of the reduction of the furcula (Queiroz and Fernandes 2011).

Although morphological characters, including furcal reduction, appear similar between some Furculanurida (including F. bistribus sp. nov.) and Stachorutes (Zon et al. 2014), the geographic separation of the two genera is remarkable. The genus Furculanurida was established for three sub-Saharan African neanurids: Micranurida africana, Kenyura furculata Salmon,1956, and Hypanurida perplexa (see Massoud 1967). Subsequently, other species were described or included in Furculanurida from the Lesser Antilles, Guatemala, Brazil, French Guiana, Tanzania, Morocco, Nepal, and Ivory Coast (Queiroz and Fernandes 2011; Zon et al. 2014). The genus, thus, conforms to a general Gondwanan distribution, with a few exceptions, like Furculanurida langdoni which is found in North America. In contrast, Stachorutes exhibits mostly an Holarctic distribution, with species known from China, France, Poland, Russia, Slovakia, Spain, and North America, except for a single species from Africa (Tanzania) (Simon-Benito et al. 2005; Fanciulli et al. 2017).

According to Queiroz and Zeppelini (2017), there are key diagnostic characters that define two groups of Pseudachorutinae in the Neotropics, which are chaetotaxy of antennae, head, thorax, and tibiotarsi. Furculanurida bistribus sp. nov. exhibits many similarities with Arlesia group of genera. In the antennal chaetotaxy, the only difference was the absence of S10 sensu Queiroz and Zeppelini (2017). Regarding head and thorax chaetotaxy, the main characters are similar, i.e. the absence of setae c2 and c3 and the presence of p1, p2 and p3 on head, the presence of only 2+2 setae on Th I, and one posterior setae displaced laterally on Th II and III. In consequence, this description reinforces the need for a revision of Furculanurida, as not only F. bistribus sp. nov. but possibly other species of Furculanurida might present characters that fit the pattern displayed by the Arlesia-group of genera sensu Queiroz and Zeppelini (2017).

Despite the differences of the new species with the most recent genus diagnosis of Furculanurida (Queiroz and Fernandes 2011), we place F. bistribus sp. nov. in this genus because of the fully developed furcula in this species (Fig. 11) and its distribution, despite the number of setae on the dens. Inclusion of the new species in this genus thereby enlarges the generic diagnosis to include species with 3–6 setae on the dens. The characters that place the new species close to Stachorutes are of uncertain generic value and need more studies (Bernard 2007; Zon et al. 2014, Neves et al. 2019).

Furculanurida bistribus sp. nov. has this unique combination of characters: six sensilla on Ant IV, 3+3 eyes, three setae in the dens, and the absence of an internal tooth on the unguis, combined with its color pattern. Members of Furculanurida have between zero and eight eyes per side; F. bistribus sp. nov. has 3+3 eyes, though some specimens of F. arawakensis may have this number (usually 4+4 eyes). All the described species have a fully developed furcula, but more dental chaetae than the new species. Leaving aside the unique characters of F. bistribus sp. nov., it is more similar to F. arawakensis from which it differs by the presence of four teeth on mandibles (versus seven), less dental chaetae (3 versus 6), and the absence of tooth on unguis. The differences between all the species of the genus are summarized in Table 2.

Main characters of the all known species of Furculanurida, including species moved to other genera.

Species + Locality Eyes PAO vesicles Ant IV sensilla Ant IV ms Md. teeth Inner unguis tooth Dens setae Mucro VT setae Tenaculum teeth
africana Ivory Coast 0 8–10 6 - 9 absent 6 developed - -
arawakensis Lesser Antilles 4(2–4) 5–9 6 absent 7 present 6 developed 3
arlei (Stachorutes) Morocco 5 8 2 small
ashrafi (Micranurida) Nepal 9 3
belemensis Brazil 5 8–9 6 4–6 present 6
boiuna Brazil 0 8–9 6 absent 7 present 5–6 developed 3 3
duodecimoculata Morocco 11 4 3
emucronata Ivory Coast 0 13–16 absent present absent 3 3
furculata Rwanda 2 4 7 7 present 6 developed
goeldiana Brazil 7 7–10 7 4
grandcolasorum Tanzania 5 9–10 absent 3–6 3
guatemalensis Guatemala 5 15 6 present 4 present 6 developed 4 3
langdoni USA 5 14–22 19 present 6 present 5 developed 4 3
longisensillata French Guiana 6 6–7 6 absent 10–11 present 6 developed 3
nessimiani Brazil 6 absent 4 3
perplexa (Hypanurida) Rwanda 4 20–22 5 present 3–4 spiniform
septemoculata Guatemala 7 15 6 present 2 present 6 developed 4 3
tropicalia Brazil 8 8–10 6 absent 4 present 6 developed 3 3
bistribus sp. nov. Puerto Rico 3 6 6 absent 4 absent 3 developed 3 3

Acknowledgements

This research was supported by Grant DEB 1239764 and 1546686 from the US National Science Foundation to the Institute for Tropical Ecosystem Studies, University of Puerto Rico, and to the International Institute of Tropical Forestry (IITF) USDA Forest Service, as part of the Luquillo Long-Term Ecological Research Program. The US Forest Service (Department of Agriculture) Research and Development Unit and the University of Puerto Rico provided additional support. We also thank to María M. Rivera (IITF) who helped with fieldwork and Edward Quigley assisted with the digitization of the figures.

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