Research Article
Print
Research Article
The Hercules pseudoscorpions from Madagascar: A systematic study of Feaellidae (Pseudoscorpiones: Feaelloidea) highlights regional endemism and diversity in one of the “hottest” biodiversity hotspots
expand article infoMichelle Lorenz§, Stephanie F. Loria, Mark S. Harvey|, Danilo Harms
‡ Museum of Nature Hamburg – Zoology, Leibniz Institute for the Analysis of Biodiversity Change (LIB), Hamburg, Germany
§ University of Hamburg, Hamburg, Germany
| Western Australian Museum, Collections & Research, Welshpool, Australia
¶ University of Western Australia, Crawley, Australia
Open Access

Abstract

Madagascar is amongst the “hottest” biodiversity hotspots with extreme levels of diversity and endemism. Throughout the last decades, there has been substantial progress in documenting the Malagasy invertebrate fauna but no study has ever focused on pseudoscorpions (Arachnida: Pseudoscorpiones) in the arachnid fauna. Here we review the Malagasy fauna of Hercules pseudoscorpions (family Feaellidae), which are common in soil habitats of arid biomes across Madagascar. Using morphology and molecular data, we recover three reciprocally monophyletic clades that correspond to three new genera in well-defined biogeographical regions and identify twelve new species: Antsiarananaella gen. nov. for Antsirananaella lorenzorum sp. nov., Antsiarananaella leniae sp. nov., Antsiarananaella faulstichi sp. nov. and Antsiarananaella marlae sp. nov.; Mahajanganella gen. nov. for Mahajanganella fridakahloae sp. nov., Mahajanganella heraclis sp. nov. and Mahajanganella schwarzeneggeri sp. nov.; Toliaranella gen. nov. for Toliaranella fisheri sp. nov., Toliaranella griswoldi sp. nov., Toliaranella mahnerti sp. nov., Toliaranella meridionalis sp. nov. and Toliaranella pumila sp. nov. Local endemism in this fauna is high and most species have small distributions, ranging from 20 km to 350 km linearly. Genetic distances between populations are also high, suggesting restricted dispersal or selection against dispersal in this fauna. Species’ ranges seem to be delimited by geological barriers including volcanic fields (Ambre-Bobaomby in the north of Madagascar), mountain ranges (foothills of the Central Highland Plateau), and rivers (Manankolana, Mandrare, Manombo and Onilahy Rivers and their anabranches), but mainly by different biome habitats. Overall, Madagascar emerges as a global “hotspot” of feaellid radiation and these animals may be used in future studies to test biogeographical hypotheses across xeric biomes on this island.

Keywords

biodiversity, false scorpions, new species, systematics, taxonomy

1. Introduction

Madagascar is one of the world’s priority conservation hotspots and amongst the most biodiverse countries with extreme levels of endemism and high species diversity across rich vegetation zones, including tropical rainforest in the east, spiny desert in the south and dry deciduous rainforest in the west (Mittermeier et al. 2004). Sadly, more than 90% of original habitats are already lost and major habitat types are irreversibly damaged (Ganzhorn et al. 2008). Madagascar’s high levels of biodiversity and endemism are correlated with landscape heterogeneity, bioclimatic zonation and a long history of isolation that includes rifting from West Gondwana (Africa and South America) beginning as early as 166 Ma and ending by 116 Ma, and from India-Seychelles 85–90 Ma, during the break-up of Gondwana (Ali and Aitchison 2008; Vences et al. 2009). Ecological disparities between the arid west and humid east, montane and lowland rainforest refugia, and habitats separated by canyon and riverine barriers have driven adaptive trait diversification among the island’s fauna and flora, allowing species to occupy new niches as they emerged (Wood et al. 2015). In addition to ongoing deforestation and overharvesting, anthropogenic climate change is increasingly threatening the remaining natural habitats and accelerating biodiversity loss in this natural laboratory (Morelli et al. 2020).

In the past two decades, there has been major progress in documenting the Malagasy invertebrate fauna at the taxonomic level, including many arachnid lineages (e.g., Griswold et al. 2012; Saucedo et al. 2015; Wood and Scharff 2018). Several studies have addressed patterns and causes of invertebrate speciation (e.g., Miraldo et al. 2011; Wesener et al. 2011; Agnarsson et al. 2015), but no data are presently available for pseudoscorpions, a mesodiverse arachnid lineage, following the more diverse spiders and mites, which are found in all ground habitats on Madagascar (WPC 2022). No detailed taxonomic studies have ever been conducted on Malagasy pseudoscorpions and 15 species are presently recognized in eight families – a relatively small number compared to regions such as Kenya (133 species) or South Africa (ca. 140 species). In a brief contribution, Heurtault (1986) listed eight of the presently 25 recognized pseudoscorpion families on Madagascar and identified twelve putatively endemic genera in these families but also wrote that “Les Pseudoscorpions (…) ne présentent pas de cas “d’explosion” ou de “pulverisation” spécifique” (“The pseudoscorpions do not represent a case of explosive speciation”) in contrast to many other faunal groups. She was also amongst the first to note the presence of the pseudoscorpion family Feaellidae Ellingsen, 1906 on Madagascar and mapped a record from Toliara Province but did not describe any species, however.

Feaellid pseudoscorpions have a unique morphology that includes a dorsoventrally compressed body, a thick and granulate cuticle, the presence of 2–6 lobes on the anterior margin of the carapace, and raptorial pedipalps with heavily armed and robust pedipalpal femora, resulting in a somewhat hulky appearance, hence the name “Hercules pseudoscorpions” that is introduced here to symbolize this morphology for non-arachnologists. Together with their sister-family, the Pseudogarypidae Chamberlin, 1923, this family constitutes one of the two basal suborders of pseudoscorpions, Atoposphyronida Harvey 2019, which is characterized by the unique presence of carapaceal lobes, spiracles near sternites IV and V, a heavily sclerotized anal plate, and the absence of venom glands in the chelal fingers (Harvey 1992; Benavides et al. 2019). Atoposphyronida may have originated in the Permian or even Carboniferous (Benavides et al. 2019). Today, the Feaellidae are restricted to the Mediterranean and tropical biomes of former Gondwanan landmasses (Fig. 1), although compression and amber fossils from the Triassic and Paleogene of Europe highlight a once wide distribution (Henderickx and Boone 2014, Kolesnikov et al. 2022). The taxonomy of Feaellidae is still in flux and there is no recent generic revision but six genera and subgenera are presently recognized in two extant subfamilies. Cybellinae Judson, 2017 includes a single genus, Cybella Judson, 2017 with four species from subterranean habitats in the Southeast Asian countries of Cambodia, Malaysia and Vietnam (Judson 2017; Harvey 2018). The second subfamily, Feaellinae Ellingsen, 1906, includes the monotypic Iporangella Harvey et al., 2016 from the Brazilian Atlantic rainforests, and Feaella Ellingsen, 1906, which is divided into three subgenera based on the number of carapaceal lobes: Feaella (Feaella) Beier, 1955 with three species from tropical Africa (Guinea-Bisseau, Côte d’Ivoire, Congo and Kenya) having six carapaceal lobes; Feaella (Tetrafeaella) Murthy and Ananthkarishnan, 1977 with twelve Recent species from southern Africa, India, Sri Lanka, the Seychelles, Maldives, Western Australia and a Baltic amber fossil (Harvey et al. 2016a, 2016b; Novák et al. 2020; Harvey 2022); and Feaella (Difeaella) Beier, 1966 with a single species from Kruger National Park, South Africa (Beier 1966). Although all species in the subgenus Feaella (Tetrafeaella) have four carapaceal lobes, it is likely that this subgenus is polyphyletic as it includes species from diverse climates (Mediterranean, tropical) and habitats (e.g., rocky outcrops in Western Australia, savannah and coastal habitats in continental Africa; Beier 1955; Harvey et al. 2016a), and with diverse morphologies.

Figure 1. 

Global distribution of the pseudoscorpion family Feaellidae Ellingsen, 1906.

Before the onset of phylogenetics, most feaellid species were considered widespread, e.g., Feaella (Feaella) mirabilis Ellingsen, 1906 across western and central Africa (Heurtault-Rossi and Jézéquel 1965; Heurtault 1983) or Feaella (Tetrafeaella) indica (Chamberlin, 1931) across Bangladesh, India and Sri Lanka (Chamberlin 1931; Batuwita and Benjamin 2014). However, recent molecular studies have indicated extreme local endemism in a morphologically cryptic fauna with slow rates of evolution, implying that many species hypotheses will have to be revised (Harvey et al. 2016a; Novák et al. 2020). In contrast to the historical literature, Feaellidae may in fact turn out to be a highly suitable group for vicariance biogeography because they seem to disperse poorly and occur almost exclusively on continental landmasses despite a record from the Maldives that might indicate trans-oceanic dispersal (Novák et al. 2020). Certainly, the present distribution is indicative of diversification from continental drift and the Late Cretaceous (Cenomanian: 99 Ma), Burmese amber fossil, Protofeaella peetersae Henderickx and Boone, 2014 shares prime synapomorphies of feaellids, providing unequivocal evidence for an ancient radiation beyond age estimates provided by molecular clock analyses (Henderickx and Boone 2016).

The presence of feaellids in Madagascar went largely unnoticed even in the arachnological community apart from sporadic records (Vachon 1960; Heurtault 1986) but comprehensive biodiversity surveys conducted by Brian L. Fisher and his team have established records of Hercules pseudoscorpions from all of Madagascar’s drier vegetation zones but not from eastern forests. Unlike other regions of the world where feaellids are highly elusive and rarely collected, Hercules pseudoscorpions are amongst the more abundant pseudoscorpions in sclerophyll and savannah habitats and have been frequently collected in pitfall traps, often in large numbers. However, occurrences seem to be localized and strictly tied to specific microhabitats or localities. Following our detailed study, the Malagasy fauna also falls into distinct morphological clades that show strict regional zonation: a northern clade (in the former Antsiranana Province), a western clade (in the former Mahajanga and Toliara Provinces), and a southern clade (in the former Toliara Province) (Fig. 2). These three clades share the presence of four carapaceal lobes and hence resemble Feaella (Tetrafeaella) rather than Feaella (Feaella) or Feaella (Difeaella). However, considering that each of these three clades present unique diagnostic characters, are genetically well-differentiated and correspond to specific regions of Madagascar, they are each recognized here as new genera.

Figure 2. 

Distribution of examined Malagasy genera: Antsirananaella, gen. nov. (turquoise), Mahajanganella, gen. nov. (yellow) and Toliaranella, gen. nov. (pink).

This study aims to: (1) document the Malagasy fauna of Hercules pseudoscorpions both at the generic and species level across dry and subarid bioclimatic zones in Madagascar; (2) provide the first comprehensive monograph of a Malagasy pseudoscorpion lineage to date; (3) use taxonomic data to discuss endemism and distributional patterns with regards to hotspot biogeography of other invertebrates; and (4) discuss potential reasons for the unprecedented levels of feaellid diversity in Madagascar.

2. Methods

2.1. Morphology

All specimens were collected between 2002–2006 and are deposited in the California Academy of Sciences (CAS), San Francisco and the Museum of Nature Hamburg – Zoology (formerly Zoological Museum Hamburg (ZMH)). Specimens sequenced as outgroups are lodged in the Western Australian Museum (WAM), Perth. Most Malagasy specimens were collected by Dr. Brian Fisher during extensive surveys of Malagasy invertebrates and preserved in 75% ethanol. Specimens were sexed, identified and sorted using a Leica M125C stereomicroscope. Dissected parts were kept in microvials. Measurements (in mm) were taken with a Leica M205A stereomicroscope and Leica Application Suite X Version 3.0.1. Digital images were taken with a custom-made BK Plus Lab System by Dun, Inc. using a Canon EOS 7D Mark II camera, an attached 5×-magnification microscopic lens in a P-51 CamLift-System installation controlled by P-51 Camlift Controller ver. 2.8.0.0 and by Capture One ver. 9.3. Scanning electron images were taken from temporarily mounted specimens using a Hitachi TM4000Plus scanning electron micrograph. Since many specimens were coated with a thick layer of dirt or particles, ultrasound cleaning was used to improve imaging results, however, success was limited. Drawings were made by hand using original images that were checked against primary specimens after illustration. Maps were created using QGIS Version 3.0 (https://www.qgis.org) and coordinates taken from original labels. Images, trees and maps were edited with Adobe Photoshop Version CC 2017.

Abbreviations. OR (outer row); MR (middle row); IR (inner row); af (anterior furca); al (anteriolateral); am (anteriomedial); pf (posterior furca); pl (posteriolateral); pm (posteriomedial)

2.2 DNA Extraction, Sequencing and Taxon Sampling

DNA extraction and Sanger sequencing was trialed for representative specimens (one per locality) but because all specimens were collected almost two decades ago, amplification of mitochondrial or ribosomal markers failed. Still, we were able to amplify a fragment of the nuclear protein-coding Histone 3a locus (H3), using primers and protocols from a previous study (Harms et al. 2019). Amplification of this short gene fragment (324 bp in total) was successful for eighteen Malagasy samples, representing eighteen localities (Table 1). Two additional Feaella (Tetrafeaella) specimens were also sequenced, including Feaella (T.) cf. anderseni (Harvey, 1989) from the Bonaparte Archipelago in Western Australia and Feaella (T.) cf. mucronata (Tullgren, 1907) from Eastern Cape, South Africa. We also sequenced ten representative outgroup taxa including two species of Pseudogarypidae [Pseudogarypus bicornis (Banks, 1895) and Neopseudogarypus scutellatus Morris, 1948], the sister family of Feaellidae, and eight species of Heterosphyronida including taxa from both families, Chthoniidae Daday, 1889 and Pseudotyrannochthoniidae Beier, 1932, representing the second most plesiomorphic suborder within Pseudoscorpiones. A sequence of a scorpion, Urodacus planimanus Pocock, 1893, was downloaded from GenBank and included as the root for phylogenetic analyses (Table 1).

Table 1.

Details of the species sequenced for H3 and GenBank accession codes.

Species Collection Number Locality GenBank Accession Code
Heterosphyronida
Allochthonius coreanus Morikawa, 1970 WAM T120699 South Korea OP589972
Pseudotyrannochthonius giganteus Beier, 1971 WAM T120449 Australia: Western Australia OP589948
Ephippiochthonius boldorii (Beier, 1934) WAM T120677 Germany: Saxony OP589971
Ephippiochthonius tetrachelatus (Preyssler, 1790) WAM T120678 Germany: Saxony-Anhalt OP589973
Mundochthonius ussuricus Beier, 1979 WAM T120369 Russia: Primorsky Krai OP589945
Tyrannochthonius terribilis malaccensis Beier, 1952 WAM T114617 Malaysia: Selangor OP589946
Atoposphyronida
Neopseudogarypus scutellatus Morris, 1948 WAM T104213 Australia: Tasmania OP589951
Pseudogarypus bicornis (Banks, 1895) WAM T114846 U.S.A.: Oregon OP589950
Feaella (Tetrafeaella) cf. anderseni (Harvey, 1989) WAM T120617 Australia: Western Australia OP589952
Feaella (Tetrafeaella) cf. mucronata (Tullgren, 1907) WAM T120637 South Africa: Eastern Cape OP589949
Antsirananaella lorenzorum sp. nov. CAS 9071844 Madagascar: Antsiranana region OP589964
Antsirananaella faulstichi sp. nov. CAS 9071843 Madagascar: Antsiranana region, Reserve special d’Ankarana
Antsirananaella leniae sp. nov. CAS 9072445 Madagascar: Antsiranana region, Foret d’Antsahabe OP589968
CAS 9072435 Madagascar: Antsiranana region, Foret d’Binara OP589966
CAS 9072455 Madagascar: Antsiranana region, Foret d’Bekaraoka OP589967
Antsirananaella marlae sp. nov. CAS 9072444 Madagascar: Antsiranana region, Foret d’Ampondrabe OP589965
Mahajanganella heraclis sp. nov. CAS 9071835 Madagascar: Province Mahajanga, Parc National d’Ankarafantsika OP589970
Mahajanganella fridakahloae sp. nov. CAS 9071518 Madasgacar: Province Toliara, Foret de Kirindy OP589969
Mahajanganella schwarzeneggeri sp. nov. CAS 9071739 (not sequenced) Madagascar: Province Mahajanga, Parc National de Baie de Baly
Toliaranella pumila sp. nov. CAS 9071606 Madagascar: Province Toliara, Foret de Tsinjoriaky OP589953
CAS 9071743 Madagascar: Province Toliara, Foret de Beroboka
Toliaranella fisheri sp. nov. CAS 9071590 Madagascar: Province Toliara, Foret de Bealoka OP589962
CAS 9071578 Madagascar: Province Toliara, Foret de Mahavelo OP589963
Toliaranella griswoldi sp. nov. CAS 9071500 Madagascar: Province Toliara, Parc National d’Andohahela OP589960
CAS 9071509 Madagascar: Province Toliara, Foret d’Andohahela OP589959
Toliaranella mahnerti sp. nov. CAS 9071763 Madagascar: Province Toliara, Antafoky OP589955
CAS 9071770 Madagascar: Province Toliara, Sept Lacs region OP589954
CAS 9071522 Madagascar: Province Toliara, Mandarano OP589957
CAS 9071753 Madagascar: Province Toliara, Mandarano OP589958
CAS 9072461 Madagascar: Province Toliara, Mandarano OP589956
Toliaranella meridionalis sp. nov. CAS 9071748 Madagascar: Province Toliara, Reserve Speciale de Cap Sainte Marie OP589961

2.3. Phylogenetic Analyses

The H3 dataset comprising the eighteen Malagasy feaellids, two Feaella species and eleven outgroup taxa for a total of 31 terminals, was aligned using MAFFT version 7 online server (https://mafft.cbrc.jp/alignment/server), applying the G-INS-i method of alignment, which assumes global homology (Katoh et al. 2002, Katoh and Toh 2008, Katoh and Standley 2013). The total length of the aligned H3 locus was 340 bp. A -f -a maximum likelihood tree search was performed with 1000 bootstrap iterations, applying a MULTIGAMMA model, and using the RAxML (Stamatakis 2006, 2014) graphical user interface raxmlGUI 2.0 (Edler et al. 2020) in the CIPRES Science Gateway V. 3.3. (Miller et al. 2010). Mean uncorrected pairwise (p) genetic distances were calculated among all feaellid taxa using MEGA v. 11 (Tamura et al. 2021).

3. Results

3.1. Phylogenetic Analyses

Maximum likelihood analysis of the H3 dataset recovered Feaellidae as monophyletic with T. cf. mucronata and T. cf. anderseni forming a clade, sister to the Malagasy feaellids, which in turn, formed three geographically delimited clades (Fig. 3): a northern clade represented by five terminals from dry Diana and Sava Regions (formerly Antsiranana Province), a central clade represented by two terminals from the dry Boeny and Menabe Regions (formerly Mahajanga and upper Toliara Provinces), and a southern clade represented by eleven terminals from the sub-arid Androy, Anosy and Atsimo-Andrefana Regions (formerly Toliara Province). Bootstrap support values were moderate to high for all three clades: 52%, 85% and 95% for the central, southern and northern clades, respectively.

Figure 3. 

Phylogeny of Malagasy Feaellidae Ellingsen, 1906 obtained by maximum likelihood analysis of the H3 locus (340 aligned sites) for 31 terminals. Nodes above branches indicate bootstrap support values.

Uncorrected p-distances between Malagasy terminals ranged from 0% (MGF 034 and MGF 033) to 15.2% (between Sept Lacs and Forêt de Bekaraoka). Mean uncorrected p-distances between the Malagasy species and two Feaella species was 15.5% and among the Malagasy clades were as follows: central-southern clades (11.4%), northern-central clades (12.0%) and northern-southern clades (13.1%). Mean uncorrected p-distances within each clade were 5% for the northern and southern clades, and 6% for the central clade. Subdivision into three geographically delimited clades is also mirrored by morphological differences, primarily morphometrics and chaetotaxy of the pedipalps, which are commonly used in pseudoscorpion taxonomy for species delimitation. Based on results of the phylogenetic analyses, genetic distances and supporting morphological data, these three geographically-delimited Malagasy clades are recognized as genera below.

3.2 Systematics

Family Feaellidae Ellingsen, 1906

Subfamily Feaellinae Ellingsen, 1906

All genera and species described below can be placed in the subfamily Feaellinae based on the presence of pleural platelets, the absence of the cheliceral rallum and an unmodified coxa III. Cybellinae lack pleural platelets, coxa III is modified, and the rallum comprises two blades (Judson 2017; Harvey 2018).

Note that there are significant morphological differences between the species included in the three different subgenera of Feaella which are retained here for practical reasons but almost certainly polyphyletic and in need of revision. The status and composition of Feaella (Tetrafeaella) is problematic due to many poorly described species, including the type species Feaella (T.) indica Chamberlin, 1931, and the knowledge that the Australian fauna is misplaced in the genus (Harvey et al. 2016a).

The newly defined genera also differ from all previously described genera of Feaellinae and can be differentiated using characters in the key below:

Key to genera and subgenera of Feaellinae

1a Carapace with 2 or 4 anterior lobes 2
1b Carapace with 6 anterior lobes Feaella Beier, 1955
2a Carapace with 2 anterior lobes 3
2b Carapace with 4 anterior lobes 4
3a Specialized setae on the retrolateral face of the movable pedipalpal finger absent Iporangella Harvey et al., 2016
3b Specialized setae on the retrolateral face of the movable pedipalpal finger present Feaella (Difeaella) Beier, 1966
4a Coxa I with 3–4 coxal spines Feaella (Tetrafeaella) Beier, 1955 from Australia
4b Coxa I with 1–2 coxal spines 5
5a Coxa I with deep basal depression stretching widely into coxa II, framed by large irregularly sized spines; sternum heavily granulated (Fig. 25B, C) Feaella (Tetrafeaella) from Africa
5b Coxa I with shallow basal depression framed by irregularly sized spines from coxa II; sternum slightly or moderately granulated 6
6a. Movable chelal finger with 5 specialized setae between trichobothrium b and terminal teeth arranged in a transverse row Antsirananaella, gen. nov.
6b. Movable chelal finger with 5 specialized setae between trichobothrium b and terminal teeth arranged in a compact group 7
7a. Cuticle of coxa I with perforated pit organ in a basal depression Toliaranella, gen. nov.
7b. Cuticle of coxa I without perforated organ in a basal depression Mahajanganella, gen. nov.

Antsirananaella gen. nov.

Type species

Antsirananaella lorenzorum sp. nov.

Diagnosis

Antsirananaella gen. nov. differs from other Malagasy genera, Toliaranella gen. nov. and Mahajanganella gen. nov., by the following characters: the presence of five specialized setae on the retrolateral face of the movable chelal finger that are arranged in a row between trichobothrium b and the terminal chelal teeth (arranged in a group in Toliaranella gen. nov. and Mahajanganella gen. nov.); the presence of five terminal teeth (including one large tooth) on the fixed chelal finger and seven on the movable chelal finger (number of terminal teeth varying from 7–9 teeth in fixed and movable fingers in other Malagasy genera); anteromedial lobes of carapace closer to each other and slightly longer than anterolateral lobes (all four lobes equidistant in other Malagasy genera); and larger body size (1.80–2.17/2.17–2.47 ♂/♀ in Antsirananaella gen. nov. and ca. 1.56–1.89/1.96–2.33 ♂/♀ in all other genera). Antsirananaella gen. nov. differs from Cybella (Southeast Asia) by having platelets on the pleural membrane (absent in Cybella), from Iporangella (Brazil) by the presence of specialized setae on the movable chelal finger (absent in Iporangella), from Feaella (Difeaella) (South Africa) and Feaella (tropical Africa) by the presence of four anterior carapaceal lobes (two and six, respectively) , from Feaella (Tetrafeaella) in continental Africa by having a less pronounced depression on the base of coxa I and on top of coxa II (distinctly more pronounced in Feaella (T.) cf. mucronata in Figs 2527), and from the Australian species presently attributed to Feaella (Tetrafeaella) by having fewer coxal spines (one pair versus three or four in the Australian taxa).

Etymology

This genus is named after the former Antsiranana Province, the area where the specimens were found. The gender is feminine.

Description

The following description is based on examination of all species in Antsiranana. Typical feaellid habitus with a spherical abdomen, short and robust pedipalps, four prominent carapaceal lobes and dark reddish-brown body colour. — Carapace: (Figs 5A, 7A, 8A, 9A, 10A): With four distinctly pointed anterior lobes; anteriolateral lobes slightly broader and wider than anteriomedial ones; anteriomedial lobes distinctly longer and closer to each other than anteriolateral ones; two pairs of eyes with lenses, equal in size, second pair partly covered by cuticula; four prominent posterior lobes (pm, pl in Figs 7A, 8A, 9A, 10A) and two longitudinal furrows more distally (af, pf in Figs 7A, 8A, 9A, 10A); with two medio-lateral mounds; and two postero-lateral processes. — Pedipalp: With a distinct conical protuberance on trochanter (Figs 4A, B, 7D, 8D, 9D, 10D); femur broad and with one prolateral triangular process plus a retrolateral hump, patella cone-shaped. Chelal hand very small and with one large medial tooth at the base of each finger. Fixed finger with 9 trichobothria including dt, movable finger with 4 trichobothria (Figs 6A, B, C, 7C, 8C, 9C, 10C). Movable finger with 5 specialized setae on ventral face, arranged in a transverse row between b and terminal teeth (Fig. 6A). Chelal teeth large and retrorse, arranged in three rows on both chelal fingers. Terminal teeth situated in compact groups facing medial, four equally sized teeth and one larger tooth on fixed finger, seven equally sized teeth on movable finger (Figs 6A, B, C, 7C, 8C, 9C, 10C). With 2 sensory setae (dt in Figs 6B, 7C, 8C, 9C, 10C) on dorsal tip of fixed finger. — Chelicera: Palm with five long and several short setae; is and ls close; sbs proximal on fixed finger; movable finger with 1 subdistal seta (gs); es on palm close to base of movable finger; galea absent; spinneret conical; serrula exterior with 16–20 blades; no rallum could be found in any of the Malagasy species because of dirt; movable finger short and without teeth. — Coxal region (Fig. 5B, C): Pedipalpal coxae with one spike laterally near base (Fig. 5B); coxae I with a basal depression and one small coxal spine only (Fig. 5B); coxae II with irregular shaped spines various in number framing depression of coxa I (Fig. 5B); all coxae reaching towards midline and with coxa IV bigger in size than coxa I–III; cuticle within depression strongly granulate. — Legs (Figs 4B, 7B, 8B, 9B, 10B): Trochanter I and II rather circular, trochanter III and IV rather elliptical; femur I and II slightly longer than patella I and II; femur III and IV shorter than patella III and IV; all tarsi long and slender, without specialized tactical trichobothria; subterminal tarsi with two curved and smooth claws; all setae acuminate; arolium much shorter than claws and with fimbriate distal margin; claws divided. — Abdomen (Fig. 4A, B, C): Paired tergites and sternites medially divided; anal plate strongly sclerotized and with a circular raised rim in the center; pleural membrane with two rows of pleural platelets, 15 in the dorsal row, 14 in the ventral row. — Genital region (Figs 4B, 5B): Details like outer genital setae and inner structures not visible because of dirt on all specimens.

Figure 4. 

Antsirananaella, gen. nov., habitus. Specimen used for imaging: A. leniae sp. nov. holotype (CAS 9072445). A dorsal; B ventral; C lateral. Scale bar: 0.5 mm.

Figure 5. 

Antsirananaella gen. nov., scanning electron micrographs. Specimen used for imaging: A. leniae sp. nov. male paratype (ZMH-A0016685). A carapace, dorsal; B sternum, ventral; C coxal spines (highlighted in red). Scale bars: 0.2 mm (A, B); 0.1 mm (C).

Figure 6. 

Antsirananaella, gen. nov., scanning electron micrographs. Specimen used for imaging: A. faulstichi sp. nov. allotype (ZMH-A0016680). A right chela from ventral with specialized setae in a row in detail; B left chela with double trichobothrium dt in detail; C left chela from medial with terminal teeth in a compact group in detail. Scale bar: 0.2 mm.

Included species

Antsirananaella lorenzorum sp. nov.; Antsirananaella faulstichi sp. nov.; Antsirananaella leniae sp. nov.; and Antsirananaella marlae sp. nov..

Distribution

Antsirananaella gen. nov. is endemic to dry deciduous forests in Diana and Sava Regions (formerly Antsiranana Province) of northern Madagascar and separated by other Malagasy genera by more than 500 km of terrain. The climate is arid or “transition tropical” with temperatures ranging from 10.5–41.5 °C. Annual precipitation ranges from 400–1000 mm.

Antsirananaella lorenzorum sp. nov.

Type material examined

MADAGASCAR: Antsiranana Province [Diana Region]: Holotype ♂ (CAS 9071844 [BLF3182]), allotype ♀ (ZMH-A0016678), paratypes: 2 ♀ (CAS 9071844 [BLF3182]), 1 ♀ (ZMH-A0016679), 12°19′22″S 49°20′17″E, December 2003, B.L. Fisher.

Diagnosis

Differs from all congeners by body length (1.94/2.33 ♂/♀ versus 1.80/2.17 ♂/♀ in A. leniae sp. nov., and 2.14/2.47 ♂/♀ in A. faulstichi sp. nov.); smallest ratio of pedipalpal femora (1.70/1.74 ♂/♀ versus 1.80/1.84 ♂/♀ in A. marlae sp. nov., and 1.75/1.79 ♂/♀ in A. faulstichi sp. nov.); pedipalpal chaetotaxy with sb not on same sagittal level as est but slightly more distal (on same sagittal level in A. marlae sp. nov. and A. leniae sp. nov.), it between eb and et closer to eb than to et (equidistant in A. leniae sp. nov.), et situated midway between dt and eb on fixed finger (situated in distal third between dt and eb in A. marlae sp. nov. and A. faulstichi sp. nov.).

Etymology

This species is a patronym for the senior author’s family that include parents Sylvia and Günther Lorenz and brother Henrik.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 7A): 1.38 (♂), 1.35–1.47 (♀) times longer than broad. — Pedipalp (Fig. 7C): Trichobothrial pattern: esb and est in the proximal half of the retrolateral face; ib, isb and ist situated basally as a slightly curved line, isb and ib closer to each other than isb and ist; eb and it situated sub-distally and very close to each other, with it more distal and more medial than eb; et situated distally and approximately the same distance to dt than to eb; dt situated very distal in a plain pit; st situated sub-basally at the same sagittal level as esb; st and est also on the same level as ist; t distinctly closer to sb than to b; sb distinctly closer to t than to st. Chelal fixed finger with 10 teeth in the OR, 20 in the MR and 15 in the IR; movable finger with 11 teeth in the OR, 14–15 teeth in the MR, and 14 teeth in the IR. — Dimensions (mm): Holotype ♂: Body length 1.94; abdomen length 1.33; abdomen width 1.12 (without pleura), 1.24 (with pleura); carapace length 0.58; carapace width 0.42. Pedipalp: trochanter 0.22; femur length 0.54; femur width 0.32; patella 0.46; chela (without pedicel) 0.54; hand length (without pedicel) 0.12; width 0.14; movable finger length 0.42. Leg I: trochanter 0.11; femur 0.19; patella 0.17; tibia 0.17; tarsus 0.27. Leg IV: trochanter 0.17; femur 0.13; patella 0.28; tibia 0.29; tarsus 0.35. Allotype ♀: Same as male except body length 2.26; abdomen length 1.59, width 1.36 (without pleura), 1.49 (with pleura); carapace length 0.66, width 0.49. Pedipalp: trochanter 0.30, femur length 0.75, width 0.44, patella length 0.61, chela (without pedicel) 0.64, hand length (without pedicel) 0.14, width 0.18. Leg I: trochanter 0.14, femur 0.25, patella 0.22, tibia 0.20, tarsus 0.29. Leg IV: trochanter 0.23, femur 0.15, patella 0.36, tibia 0.42, tarsus 0.39.

Figure 7. 

Drawings of Antsirananaella lorenzorum sp. nov.. A carapace, dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.94 (♂), 2.26–2.45 (♀); abdomen length 1.33 (♂), 1.59–1.71 (♀); abdomen width (without pleura) 1.12 (♂), 1.27–1.40 (♀); abdomen width (with pleura) 1.24 (♂), 1.41–1.58 (♀), 1.19 (♂), 1.20–1.27 (♀) times longer than broad; carapace length 0.58 (♂), 0.66–0.75 (♀), width 0.42 (♂), 0.48–0.51 (♀), 1.38 (♂), 1.36–1.47 (♀) times longer than broad. Pedipalp: trochanter 0.22 (♂), 0.27–0.30 (♀); femur 0.54 (♂), 0.71–0.79 (♀) in length, 0.32 (♂), 0.40–0.47 (♀) in width, 1.70 (♂), 1.68–1.80 (♀) times longer than broad; patella 0.46 (♂), 0.58–0.63 (♀) long; chela (without pedicel) 0.54 (♂), 0.61–0.67 (♀); hand (without pedicel) 0.12 (♂), 0.13–0.14 (♀) in length, 0.14 (♂), 0.17–0.18 (♀) in width, 0.86 (♂), 0.76–0.82 (♀) times longer than broad; movable finger 0.42 (♂), 0.47–0.54 (♀), 2.77 (♂).

GenBank Code

OP589964. The species differs from all congeners by more than 6.9% in the H3 dataset.

Habitat

Dry deciduous forest between the coast and a rising limestone plateau.

Distribution

Only known from the type locality (BLF3182) in Diana Region (formerly eastern Antsiranana Province), northern Madagascar.

Antsirananaella faulstichi sp. nov.

Type material examined

MADAGASCAR: Antsiranana Prov. [Diana Region]: Holotype ♂ (CAS 9071843 [BLF3012]), allotype ♀ (ZMH-A0016680), paratypes: 7 ♂, 1 ♀ (CAS 9071843 [BLF3012]), 3 ♂ (ZMH-A0016681 – ZMH-A0016683), Reserve special d’Ankarana, 12°51′49″S 49°13′33″E, December 2003, B.L. Fisher.

Additional material examined

MADAGASCAR: Antsiranana Prov. [Diana Region]: 8 ♂, 4 ♀, 7 nymphs (CAS 9072436 [BLF11400]), Reserve Analamerana, 12°44′48″S 49°29′41″E, 5 December 2004, B.L. Fisher; 1 ♂ (CAS 9008191 [BLF2972]), Reserve Speciale de l’Ankarana, 12°54′32″S 49°06′35″E, 10–16 February 2001, Fisher, Griswold et al.; 1 ♀ (CAS 9071838 [BLF2858]), Reserve Speciale de l’Ankarana; 12°54′32″S 49°06′35″E, 10–16 February 2001, Fisher, Griswold et al.; Antsiranana Prov. [Sava Region]: 1 nymph (CAS 9072439 [BLF9656]), Foret de Binara, 13°15′48″S 49°36′12″E, 3 December 2003, B.L. Fisher.

Diagnosis

Smallest body sizes of all species of Antsirananaella gen. nov. (e.g., body length 1.80/2.17 ♂/♀ versus 1.94–2.17/ 2.21–2.47 ♂/♀; carapace length 0.55/0.65 ♂/♀ versus 0.58–0.69/ 0.70–0.77 ♂/♀). Unique cheotaxy: sb not on the same sagittal level as est but slightly more distal (on the same sagittal level in A. marlae sp. nov. and A. leniae sp. nov.); it between eb and et closer to eb than to et (equidistant in A. leniae sp. nov.); et in the distal third between dt and eb on the fixed finger (midway in between them in A. lorenzorum sp. nov. and A. leniae sp. nov.).

Etymology

This species is named after my former Latin teacher, Gerhard Faulstich, who might be one of the wisest men ML has ever known.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 8A): 1.36–1.55 (♂), 1.43–1.50 (♀) times longer than broad. — Pedipalps (Fig. 8C): Trichobothrial pattern: sb on the movable finger distinctly distal to est on the fixed finger; it between eb and et on the fixed finger closer to eb than to et; et situated in the distal third between dt and eb on the fixed finger. Chelal fixed finger with 12 teeth in the OR, 16 in the MR and 15 in the IR; movable finger with 13 teeth in the OR, 13 in the MR and 16 in the IR. — Dimensions (mm): Holotype ♂: Body length 1.98; abdomen length 1.35, width (without pleura) 1.16, 1.25 (with pleura); carapace length 0.62, width 0.42; Pedipalp: trochanter 0.25; femur length 0.57, width 0.35; patella 0.46; chela (without pedicel) 0.57; hand length 0.13, width 0.15; movable finger 0.44; Leg I: trochanter 0.11, femur 0.20, patella 0.16, tibia 0.18, tarsus 0.27; Leg IV: trochanter 0.18, femur 0.12, patella 0.33, tibia 0.29, tarsus 0.32. Allotype ♀: Body length 2.28; abdomen length 1.53, width (without pleura) 1.34, 1.45 (with pleura); carapace length 0.73, width 0.51; Pedipalp: trochanter 0.32; femur length 0.71, width 0.38; patella 0.56; chela 0.65; hand length 0.14, width 0.18; movable finger 0.50; Leg I: trochanter 0.13; femur 0.25; patella 0.22; tibia 0.20; tarsus 0.33; Leg IV: trochanter 0.23; femur 0.14; patella 0.36; tibia 0.36; tarsus 0.36.

Figure 8. 

Drawings of Antsirananaella faulstichi sp. nov.. A carapace, dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.95–2.02 (♂), 2.14–2.28 (♀); abdomen length 1.29–1.37 (♂), 1.46–1.53 (♀), width (without pleura) 1.12–1.23 (♂), 1.24–1.34 (♀), 1.20–1.30 (♂), 1.33–1.45 (♀) (with pleura); 1.09–1.22 (♂), 1.14–1.18 (♀) times longer than broad; carapace length 0.58–0.66 (♂), 0.69–0.73 (♀), width 0.42–0.45 (♂), 0.46–0.51 (♀), 1.36–1.51 (♂), 1.43–1.50 (♀) times longer than broad. Pedipalp: trochanter 0.24–0.31 (♂), 0.26–0.32 (♀); femur length 0.57–0.67 (♂), 0.65–0.71 (♀), width 0.31–0.36 (♂), 0.36–0.38 (♀); 1.62–2.00 (♂), 1.81–1.87 (♀) times longer than broad; tibia 0.46–0.51 (♂), 0.52–0.56 (♀); chela (without pedicel) 0.55–0.58 (♂), 0.62–0.65 (♀); movable finger 0.44–0.47 (♂), 0.49–0.50 (♀); hand length (without pedicel) 0.10–0.13 (♂), 0.10–0.14 (♀), width 0.12–0.16 (♂), 0.16–0.18 (♀); 0.63–1.00 (♂), 0.63–0.78 (♀) times longer than broad.

GenBank Code

Genetic data is unavailable for this species.

Habitat

Tropical dry forest at 290 m altitude.

Distribution

Known from the type locality (BLF3012) and four additional locations (BLF11400, BLF2858, BLF2972 and BLF9656) in the Diana and Sava Regions (formerly Antsiranana Province).

Antsirananaella leniae sp. nov.

Type material examined

MADAGASCAR: Antsiranana Province [Sava Region]: Holotype ♂ (CAS 9072445 [BLF10116]), allotype ♀ (ZMH-A0016684), paratypes: 7 ♂, 3 ♀ (CAS 9072445 [BLF10116]), 5 ♂, 3 ♀ (ZMH-A0016685 – ZMH-A0016692), Forêt d’Antsahabe, 13°12′42″S 49°33′24″E, 12 December 2003, B.L. Fisher.

Additional material examined

MADAGASCAR: Antsiranana Province [Sava Region]: 10 ♂, 8 ♀, 15 nymphs (CAS 9072435 [BLF9556]), Foret de Binara, 13°15′18″S 49°37′00″E, 1 December 2003, B.L. Fisher; 1 ♂, 6 ♀, 8 nymphs (CAS 9072455 [BLF9872]), Foret de Bekaraoka, 13°10′00″S 49°42′36″E, 7 December 2003, B.L. Fisher.

Diagnosis

Very similar to A. lorenzorum sp. nov. but has the largest female body size of Antsirananaella gen. nov. (e.g., body length 2.47 (♀) versus 2.17 (♀) in A. faulstichi sp. nov., 2.21 (♀) in A. marlae sp. nov., 2.33 (♀) in A. lorenzorum sp. nov.; carapace length 0.77 (♀) versus 0.65 (♀) in A. faulstichi sp. nov., 0.70 (♀) in A. lorenzorum sp. nov., 0.71 (♀) in A. marlae sp. nov.); most slender chelal hand of females in Antsirananaella gen. nov. (0.85 (♀) × longer than wide versus 0.71–0.81 in other species of Antsirananaella gen. nov.). Difference in trichobothrial pattern: sb on the same sagittal level as est (more distal in A. lorenzorum sp. nov. and A. faulstichi sp. nov.); it between eb and et equidistant to both of them (closer to eb than to et in all other species of Antsirananaella gen. nov.); et midway between dt and eb on fixed finger (in the distal third between them in A. marlae sp. nov. and A. faulstichi sp. nov.).

Etymology

This species is a patronym for DH’s beloved daughter.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 9A): 1.36–1.51 (♂), 1.33–1.45 (♀) times longer than broad. — Pedipalps (Fig. 9C): Trichobothrial pattern: sb on the movable finger and est on the fixed finger on the same sagittal level; it between eb and et on the fixed finger with the same distance to eb and et. Chelal fixed finger with 12 teeth in the OR, 19 in the MR and 11 in the IR; movable finger with 11 teeth in the OR, 17 in the MR and 13–17 in the IR. — Dimensions (mm): Holotype ♂: Body length 2.21; abdomen length 1.54, width (with pleura) 1.34, 1.22 (without pleura); carapace length 0.68, width 0.45; Pedipalp: trochanter 0.27; femur length 0.63, width 0.34; patella 0.50; chela (without pedicel) 0.58; hand length (without pedicel) 0.11, width 0.15; movable finger 0.54; Leg I: trochanter 0.10; femur 0.27; patella 0.19; tibia 0.20; tarsus 0.29; Leg IV: trochanter 0.18; femur 0.13; patella 0.34; tibia 0.35; tarsus 0.38. Allotype ♀: Body length 2.46; abdomen length 1.71, width (without pleura) 1.51, (with pleura) 1.66; carapace length 0.72, width 0.54; Pedipalp: trochanter 0.36; femur length 0.79, width 0.44; patella 0.62; chela (without pedicel) 0.70; hand length 0.15, width 0.19; movable finger 0.54; Leg I: trochanter 0.14, femur 0.29, patella 0.17, tibia 0.20, tarsus 0.28; Leg IV: trochanter 0.23, femur 0.15, patella 0.41, tibia 0.48, tarsus 0.43.

Figure 9. 

Drawings of Antsirananaella leniae sp. nov.. A carapace dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 2.01–2.24 (♂), 2.18–2.70 (♀); abdomen length 1.36–1.54 (♂), 1.46–1.89 (♀), width 1.17–1.39 (♂), 1.26–1.59 (♀) (without pleura), 1.32–1.42 (♂), 1.39–1.78 (♀) (with pleura), 1.00–1.26 (♂), 1.11–1.23 (♀) times longer than broad. Pedipalp: trochanter 0.21–0.28 (♂), 0.27–0.36 (♀); femur length 0.59–0.65 (♂), 0.71–0.87 (♀), width 0.33–0.35 (♂), 0.40–0.46 (♀); 1.71–1.91 (♂), 1.68–1.89 (♀) times longer than broad; tibia 0.48–0.52 (♂), 0.57–0.66 (♀); chela 0.55–0.62 (♂), 0.65–0.72 (♀); movable finger 0.44–0.55 (♂), 0.50–0.56 (♀); hand length (without pedicel) 0.11–0.15 (♂), 0.14–0.17 (♀), width 0.15–0.17 (♂), 0.18–0.19 (♀); 0.69–0.94 (♂), 0.74–0.89 (♀) times longer than broad.

GenBank Code

OP589966, OP589967, OP589968. The species differs from all congeners by more than 4.8% in the H3 dataset.

Habitat

Tropical dry forest at 550 m altitude. All specimens were found in sifted litter (leaf mold, rotten wood).

Distribution

. Presently known from three localities: type locality (BLF10116) and two additional localities (BLF9556 and BLF9872) in the Sava Region (formerly Antsiranana Province).

Antsirananaella marlae sp. nov.

Type material mxamined

MADAGASCAR: Antsiranana Province [Sava Region]: Holotype ♂ (CAS 9072444 [BLF9974]), allotype ♀ (ZMH-A0016693), paratypes: 3 ♀, 2 ♂ (CAS 9072444 [BLF9974]), 2 ♀, 2 ♂ (ZMH-A0016694 – ZMH-A0016697), Forêt d’Ampondrabe, 26.3km 10° NNE Daraina, 175 m, 12°58′12″S 49°42′00″E, 10 December 2003, B.L. Fisher.

Diagnosis

Differs from congeners by large body size (e.g., male body length 2.17 versus 1.80–2.14, male carapace length 0.69 versus 0.55–0.66); a slender carapace (1.47 times longer than broad versus 1.36–1.40); medio-lateral mounds more pronounced than in A. lorenzorum sp. nov.; a slender pedipalpal femur (1.80 (♂), 1.84 (♀) times longer than broad versus 1.70–1.79 (♂), 1.74–1.80 (♀)); pedipalpal trichobothrial pattern: sb on movable finger on same sagittal level as est on fixed finger (slightly more distal in A. lorenzorum sp. nov. and in A. faulstichi sp. nov.), et situated in the distal third between dt and eb on fixed finger (situated midway between dt and eb on fixed finger in A. lorenzorum sp. nov. and in A. leniae sp. nov.); it between eb and et situated closer to eb than to et (equidistant in A. leniae sp. nov.).

Etymology

The specific epithet is a patronym in honour of Marla Elisa Nibasumba, the senior author’s godchild.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 10A): 1.43–1.49 (♂), 1.30–1.46 (♀) times longer than broad. — Pedipalps (Fig. 10C): Trichobothrial pattern: sb on the movable finger and est on the fixed finger on the same sagittal level. et situated in the distal third between dt and eb on the fixed finger. Chelal fixed finger with 11 teeth in the OR, 21 teeth in the MR, and 12 teeth in the IR; movable finger with 11 teeth in the OR, 13 teeth in the MR, and 13 teeth in the IR. — Dimensions (mm): Holotype ♂: Body length 1.81; abdomen length 1.25; abdomen width 0.99 (without pleura), 1.11 (with pleura); carapace length 0.55; carapace width 0.37. Pedipalp: trochanter 0.24; femur length 0.54; femur width 0.29; patella 0.43; chela (without pedicel) 0.49; hand length (without pedicel) 0.11; width 0.13; movable finger length 0.40. Leg I: trochanter 0.10; femur 0.13; patella 0.16; tibia 0.14; tarsus 0.25. Leg IV: trochanter 0.16; femur 0.11; patella 0.29; tibia 0.28; tarsus 0.31. Allotype ♀: Body length 2.30; abdomen length 1.60; abdomen width 1.21 (without pleura), 1.44 (with pleura); carapace length 0.67; carapace width 0.46. Pedipalp: trochanter 0.31; femur length 0.68; femur width 0.39; patella length 0.54; chela (without pedicel) 0.60; hand length (without pedicel) 0.14; hand width 0.18; movable finger length 0.48. Leg I: trochanter 0.11; femur 0.24; patella 0.22; tibia 0.15; tarsus 0.25. Leg IV: trochanter 0.21; femur 0.14; patella 0.35; tibia 0.41; tarsus 0.39.

Figure 10. 

Drawings of Antsirananaella marlae sp. nov.: A carapace dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.70–1.91 (♂), 2.05–2.19 (♀); abdomen length 1.14–1.32 (♂), 1.37–1.46 (♀), width 0.98–1.07 (♂), 1.18–1.25 (♀) (without pleura), 1.02–1.18 (♂), 1.29–1.38 (♀) (with pleura), 1.17–1.28 (♂), 1.14–1.24 (♀) times longer than broad; carapace length 0.52–0.58 (♂), 0.63–0.67 (♀), width 0.36–0.39 (♂), 0.46–0.48 (♀), 1.43–1.49 (♂), 1.30–1.36 (♀) times longer than broad; Pedipalp: trochanter 0.22–0.25 (♂), 0.27–0.30 (♀); femur length 0.52–0.56 (♂), 0.70–0.71 (♀), width 0.29–0.32 (♂), 0.38–0.40 (♀), 1.73–1.86 (♂), 1.78–1.87 (♀) times longer than broad; tibia 0.39–0.46 (♂), 0.54–0.55 (♀); chela 0.45–0.51 (♂), 0.56–0.60 (♀); movable finger 0.37–0.41 (♂), 0.46–0.49 (♀); hand length (without pedicel) 0.10–0.12 (♂), 0.12–0.14 (♀), width 0.12–0.14 (♂), 0.15–0.17 (♀), 0.79–0.92 (♂), 0.75–0.88 (♀) times longer than broad.

GenBank Code

OP589965. The species differs from all congeners by more than 4.8% in the H3 dataset.

Habitat

Isolated tropical dry forest at a moderate elevation (100–500 m) located northwest of Ambarilao. The forest leads up to the mountain ranges of the Central Highlands. The specimens were found in sifted litter (leaf mold, rotten wood).

Distribution

Only known from the type locality (BLF9974) in Sava Region (formerly eastern Antsiranana Province) of northern Madagascar.

Mahajanganella gen. nov.

Type species

Mahajanganella heraclis sp. nov.

Diagnosis

Mahajanganella gen. nov. is morphologically unique by having the following characters, which differ from other feaelloid members in Madagascar: 5 specialized setae on the retrolateral face of the movable chelal finger arranged in a group between trichobothrium b and terminal teeth (arranged in a row in Antsirananaella gen. nov.); smaller overall size than Antsirananaella gen. nov.; stronger granulate cuticle than Antsirananaella gen. nov.; all four anterior lobes of carapace with the same distance to each other (smaller distance between anteriomedial lobes than to anteriolateral ones in Antsirananaella gen. nov.). Like Antsirananaella gen. nov. and Toliaranella gen. nov., it differs from Cybella by having platelets on the pleural membrane (absent in Cybella), from Iporangella by the presence of specialized setae on the movable chelal finger (absent in Iporangella), from Feaella (Difeaella) and Feaella (Feaella) by the presence of four anterior carapaceal lobes (two and six, respectively) , from Feaella (Tetrafeaella) in continental Africa by having a less pronounced depression on the base of coxa I and on top of coxa II (distinctly more pronounced in Feaella (T.) cf. mucronata), and from the Australian species presently attributed to Feaella (Tetrafeaella) by having fewer coxal spines (one pair versus three or four in the Australian taxa).

Etymology

The genus is named after the former Mahajanga Province, where many of these feaellids occur. The gender is feminine.

Description

The following description is based on holotype and allotype of M. heraclis sp. nov.Carapace (Figs 12A, 14A, 15A, 16A): Moderate granulate and all four anterior lobes with same distance to each other. — Pedipalp (Figs 11A, B, 13A, B, C, 14C, D, 15C, D, 16C, D): Trichobothrial pattern: esb between st and ist on the sagittal level with ist slightly more distal and st slightly more proximal than esb; t distinctly closer to sb than to b; sb distinctly closer to t than to st; it on the fixed finger between eb and et distinctly closer to eb than to et. Movable finger with five specialized setae on the ventral face, arranged in a group and slightly more distally situated than b (Fig. 13A). Terminal teeth varying in number, both in fixed and the movable fingers each with 7–8 terminal teeth (one large tooth included on fixed finger). — Chelicera: Most specimens were too dirty to get secure information about characters. Coxal region (Fig. 12B, C), legs (Figs 11B, 14B, 15B, 16B), abdomen (Fig. 11A, B, C) and genital region (Fig. 12B) with the same characters as Antsirananaella gen. nov.

Figure 11. 

Mahajanganella gen. nov., habitus. Specimen used for imaging: M. schwarzeneggeri sp. nov. allotype (ZMH-A0016716). A dorsal; B ventral; C lateral. Scale bar: 0.2 mm.

Figure 12. 

Mahajanganella gen. nov., scanning electron micrographs. Specimens used for imaging: A M. fridakahloae sp. nov. female paratype (ZMH-A0016709); B, C M. schwarzeneggeri sp. nov. male paratype (ZMH-A0016717). A carapace, dorsal; B sternum ventral; C coxal spines (highlighted in red). Scale bars: 0.2 mm (A, B); 0.1 mm (C).

Figure 13. 

Mahajanganella gen. nov., scanning electron micrographs. Specimens used for imaging: A M. heraclis sp. nov. male paratype (ZMH-A0016701); B M. fridakahloae sp. nov. male paratype (ZMH-A0016715); C M. schwarzeneggeri sp. nov. male paratype (ZMH-A0016717). A right chela from ventral with specialized setae in a group in detail (highlighted in red); B left chela with double trichobothrium dt in detail (highlighted in red); C left chela from medial with terminal teeth in a compact group in detail. Scale bar: 0.2 mm.

Included species

Mahajanganella fridakahloae sp. nov.; Mahajanganella heraclis sp. nov.; and Mahajanganella schwarzeneggeri sp. nov..

Distribution

Specimens of Mahajanganella gen. nov. have been found in dry localities (dry tropical forests) in the Boeny and Menabe Regions (formerly Mahajanga and Toliara Provinces) along the western coast of Madagascar.

Mahajanganella heraclis sp. nov.

Type material examined

MADAGASCAR: Province de Mahajanga [Boeny Region]: Holotype ♂ (CAS 9071835 [BLF3599]), allotype ♀ (ZMH-A0016698), paratypes: 6 ♂, 6 ♀, (CAS 9071835 [BLF3599, BLF3597]), 4 ♂, 4 ♀ (ZMH-A0016699 – ZMH-A0016706), Parc National d’Ankarafantsika, Foret de Tsimaloto, 16°13′41″S 46°8′37″E, 2–8 April 2001, B.L. Fisher, Griswold et al.

Additional material examined

MADAGASCAR: Mahajanga Province [Boeny Region]: 23 ♂, 18 ♀, 29 nymphs (CAS 9008212 [BLF3571]), Parc National d’Ankarafantsika, 16°17′56″S 46°48′47″E, 26 March–1 April 2001, Fisher, Griswold et al.; 4 ♂, 2 ♀, 11 nymphs (CAS 9072477 [BLF6506]), Parc National de Namoroka, 16°22′36″S 45°19′36″E, 8–12 November 2002, Fisher, Griswold et al.; 2 ♂, 2 nymphs (CAS 9071651 [BLF6446]), Parc National de Namoroka, 16°28′00″S 45°21′00″E, 4–8 November 2002, Fisher, Griswold et al.

Diagnosis

Largest female body length of Mahajanganella gen. nov. (2.15 versus 2.03–2.08); robustest pedipalps of this new genus measured in ratios (e.g., femur 1.64/1.62 ♂/♀ times longer than broad versus 1.69–1.75/1.69–1.74 ♂/♀); e.g., femur 1.14 ♂ and ♀ times longer than tibia versus 1.27/1.21–1.27/1.27 ♂/♀; e.g., femur 1.02/1.06 ♂/♀ longer than chela versus 1.12/1.19–1.14/1.19 ♂/♀); anteriomedial lobes distinctly pointed and moderate carapace cuticle above second eyes (in other species of Mahajanganella gen. nov. distinctly more granulate carapace especially above second eyes).

Etymology

This species is named after the divine hero Herakles (lat. Hercules) who, in Greek mythology, is known for his power and courage. He is often portrayed with a mace, which resembles the strong pedipalps of this species. It is to be treated as a noun in apposition.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 14A): 1.28–1.49 (♂), 1.21–1.36 (♀) times longer than broad. — Pedipalp (Fig. 14C, D): Femora very robust 1.50–1.68 (♂), 1.56–1.69 (♀) times longer than broad. See trichobothrial pattern in genus description. Chelal fixed finger with 10 teeth in the OR, 15 in the MR and 11 in the IR; movable finger with 9 teeth in the OR, 12 in the MR and 8 in the IR. Terminal teeth with 7 equally sized and 1 larger tooth on the fixed finger, and 8 equally sized teeth on the movable finger. — Dimensions (mm): Holotype ♂: Body length 1.74; abdomen length 1.17, width (without pleura) 0.95, (with pleura) 1.08; carapace length 0.50, width 0.38. Pedipalp: trochanter 0.25; femur length 0.47, width 0.28; patella 0.39; chela (without pedicel) 0.43; hand length 0.10, width 0.13; movable finger 0.35. Leg I: trochanter 0.08; femur 0.17; patella 0.17; tibia 0.12; tarsus 0.22. Leg IV: trochanter 0.14; femur 0.12; patella 0.23; tibia 0.21; tarsus 0.22. Allotype ♀: Body length 2.19; abdomen length 1.57, width (without pleura) 1.25, (with pleura) 1.41; carapace length 0.55, width 0.45. Pedipalp: trochanter 0.25; femur length 0.52, width 0.32; patella 0.47; chela (without pedicel) 0.48; hand length 0.10, width 0.15; movable finger 0.42. Leg I: trochanter 0.09; femur 0.22; patella 0.16; tibia 0.17; tarsus 0.30. Leg IV: trochanter 0.16; femur 0.14; patella 0.34; tibia 0.33; tarsus 0.30.

Figure 14. 

Drawings of Mahajanganella heraclis sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.67–1.84 (♂), 2.06–2.22 (♀); abdomen length 1.13–1.26 (♂), 1.42–1.59 (♀); width (with pleura) 1.04–1.13 (♂), 1.32–1.46 (♀); width (without pleura) 0.92–1.05 (♂), 1.14–1.30 (♀); 1.13–1.28 (♂), 1.17–1.32 (♀) times longer than broad; carapace length 0.50–0.58 (♂), 0.55–0.61 (♀); width 0.37–0.40 (♂), 0.43–0.48 (♀); 1.28–1.49 (♂), 1.21–1.36 (♀) times longer than broad. Pedipalp: trochanter 0.17–0.25 (♂), 0.22–0.26 (♀); femur length 0.41–0.47 (♂), 0.50–0.57 (♀); width 0.26–0.28 (♂), 0.30–0.36 (♀); 1.50–1.37 (♂), 1.56–1.68 (♀) times longer than broad; patella 0.38–0.41 (♂), 0.39–0.50 (♀); chela (without pedicel) 0.42–0.45 (♂), 0.46–0.53 (♀); hand length 0.08–0.10 (♂), 0.09–0.11 (♀); width 0.12–0.13 (♂), 0.14–0.16 (♀); 0.62–0.83 (♂), 0.56–0.73 (♀) times longer than broad; movable finger 0.31–0.35 (♂), 0.37–0.42 (♀); femur 1.05–1.21 (♂), 1.08–1.16 (♀) times longer than patella; femur 0.95–1.09 (♂), 1.02–1.12 (♀) times longer than chela.

GenBank Code

OP589970. The species differs from all congeners by more than 6.5% in the H3 dataset.

Habitat

The specimens were found in sifted litter of tropical dry forest at an elevation of 135 m.

Distribution

Presently known only from the type locality (BLF3599) and from three additional locations (BLF3571, BLF6506 and BLF6446) in the Boeny Region (formerly Mahajanga Province).

Mahajanganella fridakahloae sp. nov.

Type material examined

MADAGASCAR: Province de Toliara [Menabe Region]: Holotype ♂ (CAS 9071518 [BLF4605]), allotype ♀ (ZMH-A0016707), paratypes: 9 ♂, 6 ♀ (CAS 9071518 [BLF4605]), 4 ♂, 4 ♀ (ZMH-A0016708 – ZMH-A0016715), Foret de Kirindy, 20°02′42″S 44°39′44″E, 28 November–2 December 2001, B.L. Fisher, Griswold et al.

Additional material examined

MADAGASCAR: Province de Toliara [Menabe Region]: 2 ♂, 1 ♀, 4 nymphs (CAS 9071535 [BLF4432]), Parc National de Tsingi de Bemaraha, 19°42′34″S 44°43′5″E, 16–20 November 2001, Fisher, Griswold et al.; 17 ♂, 7 ♀, 16 nymphs (CAS 9071781 [BLF4726]), Toliara Province [Menabe Region]: Parc National de Kirindy Mite, 20°47′43″S 44°08′49″E, 6–10 December 2001, Fisher, Griswold et al.

Diagnosis

Very similar to the type species but remarkably more granulated cuticle on the carapace, especially above the posterior pair of eyes; anterior lobes on the carapace are not pointed but rounded at the tip.

Etymology

This species is named after the Mexican artist Frida Kahlo de Rivera († 1954) whose unmistakable character were her striking eyebrows, which she included in many of her self-portraits. The species is reminiscent of her because of the strongly granulated cuticle above the second pair of eyes, which resemble ‘eyebrows’.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 15A): Cuticle strongly covered with granulate especially above the second eye pair which makes it look like an additional lateral pair of mounds behind the anterior margin; anterior lobes on carapace not pointed but rounded at the tips; 1.26–1.42 (♂), 1.18–1.34 (♀) times longer than broad. — Pedipalps (Fig. 15C, D): See trichobothrial pattern in genus description. Chelal fixed finger with 9–12 teeth in the OR, 14–15 in the MR and 8–12 in the IR; movable finger with 6–9 teeth in the OR, 9–11 in the MR and 9–10 in the IR; fixed finger with 6 equally sized and one larger terminal tooth, movable finger with 7 equally sized terminal teeth. — Dimensions (mm): Holotype ♂: Body length 1.71; abdomen length 1.15, width (with pleura) 1.01, width (without pleura) 0.90; carapace length 0.49, width 0.39. Pedipalp: trochanter 0.24; femur length 0.50, width 0.27; patella 0.34; chela (without pedicel) 0.41; hand length 0.08, width 0.12; movable finger 0.34. Leg I: trochanter 0.09; femur 0.14; patella 0.13; tibia 0.12; tarsus 0.21. Leg IV: trochanter 0.14; femur 0.10; patella 0.24; tibia 0.25; tarsus 0.23. Allotype ♀: Body length 2.22; abdomen length 1.58, width (with pleura) 1.35, width (without pleura) 1.14; carapace length 0.60, width 0.45. Pedipalp: trochanter 0.26; femur length 0.54, width 0.35; patella 0.44; chela (without pedicel) 0.49; hand length 0.10, width 0.14; movable finger 0.37. Leg I: trochanter 0.10; femur 0.17; patella 0.19; tibia 0.14; tarsus 0.22. Leg IV: trochanter 0.18; femur 0.13; patella 0.29; tibia 0.30; tarsus 0.31.

Figure 15. 

Drawings of Mahajanganella fridakahloae sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.71–1.86 (♂), 1.92–2.26 (♀); abdomen length 1.15–1.27 (♂), 1.50–1.67 (♀); width (with pleura) 1.01–1.08 (♂), 1.20–1.41 (♀); width (without pleura) 0.90–0.99 (♂), 1.03–1.17 (♀); 1.22–1.35 (♂), 1.23–1.43 (♀) times longer than broad; carapace length 0.50–0.55 (♂), 0.52–0.61 (♀); width 0.38–0.41 (♂), 0.42–0.47 (♀); 1.26–1.42 (♂), 1.18–1.34 (♀) times longer than broad. Pedipalp: trochanter 0.19–0.24 (♂), 0.20–0.27 (♀); femur length 0.44–0.51 (♂), 0.53–0.60 (♀); width 0.26–0.30 (♂), 0.32–0.36 (♀); 1.57–1.85 (♂), 1.54–1.75 (♀) times longer than broad; patella 0.34–0.40 (♂), 0.44–0.47 (♀); chela (without pedicel) 0.39–0.45 (♂), 0.44–0.49 (♀); hand length 0.07–0.11 (♂), 0.90–0.12 (♀); width 0.12–0.13 (♂), 0.13–0.16 (♀); 0.54–0.92 (♂), 0.56–0.80 (♀) times longer than broad; movable finger 0.32–0.34 (♂), 0.37–0.40 (♀); femur 1.16–1.47 (♂), 1.20–1.36 (♀) times longer than patella; femur 1.04–1.22 (♂), 1.13–1.34 (♀) times longer than chela.

Habitat

Specimens were found in tropical dry forest in sifted litter (leaf mold, rotten wood) at an elevation of 100–150 m.

GenBank Code

OP589969. The species differs from all congeners by more than 6.5% in the H3 dataset.

Distribution

Known from the type locality (BLF4605) and two additional localities (BLF4432 and BLF4726) in the Menabe Region (formerly Toliara Province) of western Madagascar.

Mahajanganella schwarzeneggeri sp. nov.

Type material examined

MADAGASCAR: Mahajanga Province [Boeny Region]: Holotype ♂ (CAS 9071739 [BLF6812]), allotype ♀ (ZMH-A0016716), paratypes: 1 ♂ (ZMH-A0016717), 1 ♀ (CAS 9071739 [BLF6812]), Parc National de Baie de Baly; 16°00′36″S 45°15′54″E, 26–30 November 2002, B.L. Fisher, Griswold et al.

Diagnosis

Very similar to Mahajanganella heraclis sp. nov. but smaller female body size (body length 2.03 versus 2.15 in M. heraclis sp. nov.), and most slender carapace (1.47 (♂)/1.41 (♀) × longer than broad versus 1.35 (♂)/1.29 (♀) in M. heraclis); most slender femora of Mahajanganella gen. nov. (1.57/1.74 ♂/♀ × longer than broad versus 1.64/1.62 ♂/♀ in M. heraclis sp. nov.).

Etymology

This species is a patronym honoring Arnold Schwarzenegger, a famous former bodybuilder from Austria, known as an actor in the movie “Terminator”, former governor of California and now supporting conservation programs.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 16A): 1.47 (♂), 1.38–1.43 (♀) times longer than broad. — Pedipalps (Fig. 16C, D): see trichobothrial pattern in genus description. Chelal fixed finger with 8 teeth in the OR, 13–17 in the MR and 9–12 in the IR; movable finger with 8–9 teeth in the OR, 11–13 in the MR and 9–12 in the IR; terminal teeth build a group of 6 equally sized and 1 larger tooth on the fixed finger and 8 equally sized teeth on the movable finger. — Dimensions (mm): Holotype ♂: Body length 1.93; abdomen length 1.34; width (with pleura) 1.17; (without pleura) 1.07; carapace length 0.56; width 0.38. Pedipalp: trochanter 0.23; femur length 0.56; width 0.32; patella 0.44; chela (without pedicel) 0.47; hand length 0.09; width 0.13; movable finger 0.36. Leg I: trochanter 0.09; femur 0.21; patella 0.14; tibia 0.15; tarsus 0.26. Leg IV: trochanter 0.15; femur 0.13; patella 0.29; tibia 0.24; tarsus 0.31. Allotype ♀: Body length 2.04; abdomen length 1.44; width (with pleura) 1.30; (without pleura) 1.18; carapace length 0.60; width 0.42. Pedipalp: trochanter 0.27; femur length 0.56; width 0.34; patella 0.48; chela (without pedicel) 0.51; hand length 0.10; width 0.15; movable finger 0.39. Leg I: trochanter 0.09; femur 0.21; patella 0.20; tibia 0.13; tarsus 0.28. Leg IV: trochanter 0.16; femur 0.13; patella 0.34; tibia 0.31; tarsus 0.33.

Figure 16. 

Drawings of Mahajanganella schwarzeneggeri sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.85–1.93 (♂), 2.02–2.04 (♀); abdomen length 1.25–1.34 (♂), 1.43–1.44 (♀); width (with pleura) 1.14–1.17 (♂), 1.29–1.30 (♀); width (without pleura) 1.03–1.07 (♂), 1.17–1.18 (♀); 1.21–1.25 (♂), 1.22 (♀) times longer than broad; carapace length 0.56 (♂), 0.58–0.60 (♀); width 0.38 (♂), 0.42 (♀); 1.47 (♂), 1.38–1.43 (♂) times longer than broad. Pedipalp: trochanter 0.23 (♂), 0.25–0.27 (♀); femur length 0.52–0.56 (♂), 0.56–0.58 (♀); width 0.29–0.32 (♂), 0.33–0.34 (♀); 1.75–1.79 (♂), 1.65–1.76 (♀) times longer than broad; patella 0.42–0.44 (♂), 0.48 (♀); chela (without pedicel) 0.44–0.47 (♂), 0.51 (♀); hand length 0.09–0.10 (♂), 0.09–0.10 (♀); width 0.13–0.14 (♂), 0.15 (♀); 0.67–0.69 (♂), 0.67–0.71 (♀) times longer than broad; movable finger 0.36 (♂), 0.39–0.40 (♀); femur 1.24–1.27 (♂), 1.17–1.21 (♀) times longer than patella; femur 1.18–1.19 (♂), 1.10–1.14 (♀) times longer than chela.

GenBank Code

Genetic data is unavailable for this species.

Habitat

Specimens were found in sifted litter in tropical dry forest at an elevation of 10 m very close to the coastline.

Distribution

Known only from the type locality (BLF6812) in the Boeny Region (formerly Mahajanga Province) of western Madagascar.

Toliaranella gen. nov.

Type species

Toliaranella pumila sp. nov.

Diagnosis

Toliaranella gen. nov. differs from all other Malagasy feaellids by the presence of a perforated organ above the coxal spines and smallest body size of any Malagasy groups. Like Antsirananaella gen. nov. and Mahajanganella gen. nov., it differs from Cybella by having platelets on the pleural membrane (absent in Cybella), from Iporangella by the presence of specialized setae on the movable chelal finger (absent in Iporangella), from Feaella (Difeaella) and Feaella (Feaella) by the presence of four anterior carapaceal lobes (two and six, respectively) , from Feaella (Tetrafeaella) in continental Africa by having a less pronounced depression on the base of coxa I and on top of coxa II (distinctly more pronounced in Feaella (T.) cf. mucronata) and from the Australian species presently attributed to Feaella (Tetrafeaella) by having fewer coxal spines (one pair versus three or four in the Australian taxa).

Etymology

This genus is named after the former Toliara Province, where all specimens were collected. The gender is feminine.

Description

The following description is based on holotype and allotype. — Carapace (Figs 18A, 20A, 21A, 22A, 23A, 24A): Strongly granulate, all four anterior lobes with same distance to each other and rounded at tips. — Pedipalps (Figs 17A, B, 19A–C, 20C, D, 21C, D, 22C, D, 23C, D, 24C, D): Terminal teeth with 7–8 teeth on both the fixed and the movable fingers (including 1 large tooth on fixed finger). — Coxal region (Figs 17B, 18B, C): Pedipalpal coxae each with one spike laterally near base; coxae I with basal depression, each side with one small coxal spine; slightly more cranial of coxal spine one novel organ on each side: rose–shaped, not granulate but perforated organ (Fig. 18B, C) filling the base of the depression; coxae II with irregular shaped spines framing depression of coxae I.

Figure 17. 

Toliaranella gen. nov.. Specimen used for imaging: T. griswoldi sp. nov. holotype (CAS 9071500). A dorsal overview; B ventral overview; C lateral overview. Scale bar: 0.2 mm.

Figure 18. 

Toliaranella gen. nov., scanning electron micrographs. Specimens used for imaging: A T. griswoldi sp. nov. male paratype (ZMH-A0016740); B, C T. meridionalis sp. nov. male paratype (ZMH-A0016753). A carapace, dorsal view; B sternum ventral overview; C coxal spines and perforated organ in detail (highlighted in red). Scale bars: 0.2 mm (A, B); 0.1 mm (C).

Included species

Toliaranella fisheri sp. nov.; Toliaranella griswoldi sp. nov.; Toliaranella mahnerti sp. nov.; Toliaranella meridionalis sp. nov.; and Toliaranella pumila sp. nov.

Distribution

Members of this genus are localized to the southern part of Madagascar in the Androy, Anosy and Atsimo-Andrefana Regions (formerly Toliara Province) where the main landscape is covered by arid spiny bush vegetation.

Toliaranella pumila sp. nov.

Type material examined

MADAGASCAR: Holotype ♂ (CAS 9071606 [BLF5966]), allotype ♀ (ZMH-A0016718), paratypes: 25 ♂, 4 ♀ (CAS 9071606 [BLF5966]), 8 ♂, 3 ♀ (ZMH-A0016719 – ZMH-A0016729), Toliara Province, Forêt de Tsinjoriaky, 22°48′08″S 43°25′14″E, 6 March 2002, B.L. Fisher.

Additional material examined

MADAGASCAR: Toliara Province [Atsimo-Andrefana Region]: 25 ♂, 10 ♀, 39 nymphs (CAS 9071598 [BLF 6070]), 6 ♂, 1 ♀, 5 nymphs (CAS 9071743 [BLF 6068]), Foret de Beroboka, 22°13′59″S 43°21′59″E, 12–16 March 2002, Fisher, Griswold et al.

Diagnosis

Smallest body sizes of Toliaranella species and of all Malagasy feaellids (e.g., body length 1.56/1.96 ♂/♀ versus 1.60–1.85/2.04–2.33 ♂/♀; Pedipalps: femur length 0.44/0.53 ♂/♀ versus 0.45–0.55/0.56–0.67 ♂/♀); most robust male pedipalpal femur of Toliaranella (femur 1.63 × longer than broad versus 1.65–1.72); cheotaxy: est on fixed finger on same sagittal level as sb on movable finger (est more distal than sb in T. griswoldi sp. nov. and T. fisheri sp. nov.).

Etymology

This species is named after the latin ‘pumilus, -a, -um’ which means “dwarf”. The specific epithet references the small size of all specimens.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 20A): 1.29–1.44 (♂), 1.24–1.47 (♀) times longer than broad. — Pedipalp (Fig. 20C, D): Femora very robust (1.55–1.69 (♂), 1.43–1.80 (♀) times longer than broad; trichobothrial pattern: same as in Mahajanganella gen. nov., except: est on fixed finger on same sagittal level as sb on movable finger. Chelal fixed finger with 9–10 teeth in the OR, 12–15 in the MR and 8–12 in the IR; movable finger with 6–10 teeth in the OR, 9–11 in the MR and 9–10 in the IR; fixed finger with 6 equally sized and one larger terminal tooth, movable finger with 7 equally sized terminal teeth. — Chelicera, legs, abdomen and genital region: With same characters as all members before. — Coxal region: See genus description. — Dimensions (mm): Holotype ♂: Body length 1.47; abdomen length 1.02, width (with pleura) 0.93, width (without pleura) 0.86; carapace length 0.45, width 0.32. Pedipalp: trochanter 0.21; femur length 0.45, width 0.27; patella 0.35; chela (without pedicel) 0.38; hand length 0.08, width 0.11; movable finger 0.30. Leg I: trochanter 0.08; femur 0.12; patella 0.15; tibia 0.11; tarsus 0.22. Leg IV: trochanter: 0.13; femur 0.11; patella 0.23; tibia 0.21; tarsus 0.26. Allotype ♀: Body length 2.08; abdomen length 1.51, width (with pleura) 1.39, width (without pleura) 1.15; carapace length 0.55, width 0.40. Pedipalp: trochanter 0.26; femur length 0.58, width 0.36; patella 0.42; chela (without pedicel) 0.43; hand length 0.11, width 0.13; movable finger 0.35. Leg I: trochanter 0.09; femur 0.18; patella 0.19; tibia 0.15; tarsus 0.26. Leg IV: trochanter 0.15; femur 0.13; patella 0.27; tibia 0.28; tarsus 0.33.

Figure 19. 

Toliaranella, gen. nov., scanning electron micrographs. Specimen used for imaging: T. pumila sp. nov. male paratype (ZMH-A0016722). A right chela from ventral with specialized setae in a group in detail (highlighted in red); B left chela with double trichobothrium dt in detail (highlighted in red); C left chela from medial with terminal teeth in a compact group in detail. Scale bar: 0.2 mm.

Figure 20. 

Drawings of Toliaranella pumila sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.47–1.66 (♂), 1.83–2.11 (♀); abdomen length 1.02–1.16 (♂), 1.33–1.51 (♀); width (with pleura) 0.93–0.98 (♂), 1.13–1.39 (♀); width (without pleura) 0.83–0.88 (♂), 0.96–1.15 (♀); 1.19–1.32 (♂), 1.25–1.47 (♀) times longer than broad; carapace length 0.43–0.47 (♂), 0.48–0.55 (♀); width 0.32–0.35 (♂), 0.36–0.41 (♀); 1.29–1.41 (♂), 1.24–1.47 (♀) times longer than broad. Pedipalp: trochanter 0.18–0.22 (♂), 0.22–0.27 (♀); femur length 0.42–0.46 (♂), 0.48–0.58 (♀); width 0.25–0.30 (♂), 0.30–0.36 (♀); 1.50–1.84 (♂), 1.43–1.80 (♀) times longer than broad; patella 0.33–0.35 (♂), 0.40–0.43 (♀); chela (without pedicel) 0.37–0.39 (♂), 0.42–0.46 (♀); hand length 0.07–0.10 (♂), 0.09–0.12 (♀); width 0.11–0.12 (♂), 0.13–0.14 (♀); movable finger 0.30–0.31 (♂), 0.35–0.37 (♀).

GenBank Code

OP589953. The species differs from all congeners by more than 5.3% in the H3 dataset.

Habitat

Specimens were found in sifted litter of tropical dry forest (leaf mold, rotten wood) at an elevation of 375 m.

Distribution

Presently known from the type locality (BLF9556) and one additional locality (BLF6070) in the Atsimo-Andrefana Region (formerly Toliara Province).

Toliaranella fisheri sp. nov.

Type material examined

MADAGASCAR: Toliara Province [Anosy Region]: Holotype ♂ (CAS 9071590 [BLF5316]), allotype ♀ (ZMH-A0016730), paratypes: 6 ♂, 7 ♀, (CAS 9071590 [BLF5316]), 4 ♂, 4 ♀ (ZMH-A0016731 – ZMH-A0016738), Reserve Prive Berenty, Foret de Bealoka, 24°57′25″S 46°16′17″E, 3–8 February 2002, B.L. Fisher, Griswold et al.

Additional material examined

MADAGASCAR: Toliara Province [Anosy Region]: 1 ♂, 1 nymph (CAS 9071750 [BLF5314]), Reserve Prive Berenty, Foret de Bealoka, 24°57′25″S 46°16′17″E, 3–8 February 2002, B.L. Fisher, Griswold et al. 76; ♂, 55 ♀, >100 nymphs (CAS 9071589 [BLF5426]), Reserve Prive Berenty, Foret de Malaza, Mandrare River, 25°00′28″S 46°18′22″E, 6 February 2002, Fisher, Griswold et al.; 1 ♂, 2 ♀, 2 nymphs (CAS 9071578 [BLF5278]), Foret de Mahavelo, Isantoria River, 24°45′13″S 46°09′5″E, 31 January 2002, Fisher, Griswold et al.

Diagnosis

Most similar to T. griswoldi sp. nov., but females with most slender femora of this new genus (1.71 × longer than broad versus 1.62–1.69) and different body size.

Etymology

This species is a patronym honoring Brian Fisher who collected many Malagasy feaellids.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 21A): 1.25–1.39 (♂), 1.27–1.39 (♀) times longer than broad. — Pedipalps (Fig. 21C, D): See trichobothrial pattern in description of T. griswoldi sp. nov. Chelal fixed finger with 10 teeth in the OR, 13–16 in the MR and 10–13 teeth in the IR; movable finger with 8–10 teeth in the OR, 9–12 in the MR and 8–11 teeth in the IR. Fixed finger with 6–7 equally sized and one large terminal tooth, movable finger with 7 terminal teeth equal in size. — Dimensions (mm): Holotype ♂: Body length 1.56; abdomen length 1.10, width (with pleura) 1.00, width (without pleura) 0.85; carapace length 0.44, width 0.34. Pedipalp: trochanter 0.24; femur length 0.43, width 0.27; patella 0.36; chela (without pedicel) 0.39; hand length 0.09, width 0.12; movable finger 0.32. Leg I: trochanter 0.07; femur 0.16; patella 0.14; tibia; 0.13; tarsus 0.22. Leg IV: trochanter 0.15; femur 0.10; patella 0.25; tibia 0.26; tarsus 0.26. Allotype ♀: Body length 2.15; abdomen length 1.49, width (with pleura) 1.35, width (without pleura) 1.13; carapace length 0.57, width 0.44. Pedipalp: trochanter 0.24; femur length 0.60, width 0.33; patella 0.45; chela (without pedicel) 0.50; hand length 0.11, width 0.15; movable finger 0.41. Leg I: trochanter 0.10; femur 0.20; patella 0.16; tibia 0.15; tarsus 0.28. Leg IV: trochanter 0.19; femur 0.13; patella 0.30; tibia 0.31; tarsus 0.35.

Figure 21. 

Drawings of Toliaranella fisheri sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.54–1.72 (♂), 1.83–2.15 (♀); abdomen length 1.07–1.16 (♂), 1.30–1.54 (♀); width (with pleura) 0.97–1.05 (♂), 1.16–1.38 (♀); width (without pleura) 0.83–0.92 (♂), 1.00–1.17 (♀); 1.23–1.29 (♂), 1.27–1.40 (♀) times longer than broad; carapace length 0.44–0.50 (♂), 0.51–0.58 (♀); width 0.34–0.38 (♂), 0.39–0.44 (♀); 1.25–1.39 (♂), 1.27–1.39 (♀) times longer than broad. Pedipalp: trochanter 0.23–0.27 (♂), 0.22–0.27 (♀); femur length 0.43–0.48 (♂), 0.51–0.60 (♀); width 0.26–0.29 (♂), 0.29–0.35 (♀); 1.59–1.78 (♂), 1.56–1.84 (♀) times longer than broad; patella 0.36–0.39 (♂), 0.41–0.47 (♀); chela (without pedicel) 0.39–0.43 (♂), 0.44–0.50 (♀); hand length 0.08–0.11 (♂), 0.10–0.12 (♀); width 0.11–0.13 (♂), 0.13–0.15 (♀); 0.67–0.92 (♂), 0.67–0.86 (♀) times longer than broad; movable finger 0.31–0.35 (♂), 0.36–0.41 (♀).

GenBank Code

OP589962, OP589963. The species differs from all congeners by more than 3.7% in the H3 dataset.

Habitat

Specimens were found in sifted litter (leaf mold and rotten wood) in gallery forest and spiny forest thicket at an elevation of 35–115 m.

Distribution

Known from three localities along the Mandrare River in the Anosy Region (formerly Toliara Province) in southern Madagascar.

Toliaranella griswoldi sp. nov.

Type material examined

MADAGASCAR: Toliara Province [Anosy Region]: Holotype ♂ (CAS 9071500 [BLF4810]), allotype ♀ (ZMH-A0016739), paratypes: 11 ♂, 1 ♀ (CAS 9071500 [BLF4810]), 4 ♂, 2 ♀ (ZMH-A0016740 – ZMH-A0016745), Parc National d’Andohahela, Forêt de Manantalinjo, 24°49′1″S 46°36′36″E, 12–16 January 2002, B.L. Fisher, Griswold et al.

Additional material examined

MADAGASCAR: Toliara Province [Anosy Region]: 4 ♂, 4 ♀, 9 nymphs (CAS 9071509 [BLF4984]), Parc National d’Andohahela, Forêt d’Ambohibory, 24°55′48″S 46°38′44″E, 16–20 January 2002, Fisher, Griswold et al.; 1 ♂, 1 ♀ (CAS 9071516 [BLF4815]), Parc National d’Andohahela, Foret de Manantalinjo, 24°49′1″S 46°36′36″E, 12–16 January 2002, Fisher, Griswold et al.

Diagnosis

Differs from T. pumila sp. nov. in different cheotaxy: est on fixed finger situated between sb and t of the movable finger on sagittal level (est on same sagittal level with sb in T. pumila sp. nov.); most compact carapace of the southern feaellid members (1.27/1.23 ♂/♀ times longer than broad versus 1.31–1.37/1.31–1.39 ♂/♀).

Etymology

This species is a patronym honoring Charles Griswold who collected many Malagasy feaellids.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 22A): Very compact, heavily granulate, 1.20–1.31 (♂), 1.18–1.31 (♀) times longer than broad. — Pedipalps (Fig. 22C, D): Similar trichobothrial pattern as T. pumila sp. nov. but the following difference: est on fixed finger situated between sb and t of the movable finger on sagittal level. Chelal fixed finger with 9–10 chelal teeth in the OR, 12–14 in the MR and 9–11 in the IR; movable finger with 6–8 in the OR, 7–11 in the MR and 7–8 in the IR. Fixed finger with 6–7 terminal teeth equal in size and one large one, movable finger with 7–8 equally sized terminal teeth. — Dimensions (mm): Holotype ♂: Body length 1.62; abdomen length 1.13, width (with pleura) 1.05, width (without pleura) 0.91; carapace length 0.46, width 0.35. Pedipalp: trochanter 0.22; femur length 0.49, width 0.27; patella 0.35; chela (without pedicel) 0.35; hand length 0.10, width 0.11; movable finger 0.26. Leg I: trochanter 0.07; femur 0.14; patella 0.13; tibia 0.11; tarsus 0.21. Leg IV: trochanter 0.13; femur 0.10; patella 0.21; tibia 0.20; tarsus 0.26. Allotype ♀: Body length 2.04; abdomen length 1.47, width (with pleura) 1.33, width (without pleura) 1.11; carapace length 0.53, width 0.45. Pedipalp: trochanter 0.25; femur length 0.59, width 0.35; patella 0.46; chela (without pedicel) 0.44; hand length 0.10, width 1.14; movable finger 0.35. Leg I: trochanter 0.11; femur 0.16; patella 0.14; tibia 0.16; tarsus 0.25. Leg IV: trochanter 0.18; femur 0.14; patella 0.27; tibia 0.31; tarsus 0.30.

Figure 22. 

Drawings of Toliaranella griswoldi sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variations

Body length 1.54–1.67 (♂), 1.96–2.24 (♀); abdomen length 1.08–1.18 (♂), 1.41–1.63 (♀); width (with pleura) 0.98–1.09 (♂), 1.27–1.41 (♀); width (without pleura) 0.84–0.95 (♂), 1.10–1.19 (♀); 1.20–1.29 (♂), 1.27–1.37 (♀) times longer than broad; carapace length 0.42–0.48 (♂), 0.53–0.57 (♀); width 0.35–0.38 (♂), 0.41–0.45 (♀); 1.20–1.31 (♂), 1.18–1.29 (♀) times longer than broad. Pedipalp: trochanter 0.19–0.22 (♂), 0.24–0.26 (♀); femur length 0.40–0.49 (♂), 0.54–0.59 (♀); width 0.25–0.28 (♂), 0.33–0.37 (♀); 1.60–1.81 (♂), 1.53–1.69 (♀) times longer than broad; patella 0.33–0.36 (♂), 0.45–0.48 (♀); chela (without pedicel) 0.35–0.38 (♂), 0.43–0.47 (♀); hand length 0.08–0.11 (♂), 0.10–0.11 (♀); width 0.11–0.13 (♂), 0.13–0.14 (♀); 0.73–0.91 (♂), 0.71–0.79 (♀) times longer than broad; movable finger 0.26–0.30 (♂), 0.35–0.37 (♀).

GenBank Code

OP589959, OP589960. The species differs from all congeners by more than 3.7% in the H3 dataset.

Habitat

Specimens were found in sifted litter of spiny forest thicket and tropical dry forest at an elevation of 150–300 m.

Distribution

Currently known from the type locality (BLF4810) and two additional localities of (BLF4984 and BLF4815) in the Anosy Region (former Toliara Province) in southern Madagascar.

Toliaranella mahnerti sp. nov.

Type material examined

MADAGASCAR: Toliara Province [Atsimo-Andrefana Region]: Holotype ♂ (CAS 9071763 [MGF001]), allotype ♀ (ZMH-A0016746), paratypes: 4 ♂, 2 ♀ (CAS 9071763 [MGF001]), 3 ♂, 1 ♀ (ZMH-A0016747 – ZMH-A0016750), Antafoky, 23°28′45″S 44°3′58″E, 9 February 2002, Frontier Project.

Additional material examined

MADAGASCAR: Toliara Province [Atsimo-Andrefana Region]: 3 nymphs (CAS 9071531 [MGF004]), Antafoky, 23°29′16″S 44°04′39″E, 9 February 2002, Frontier Project; 3 ♂, 1 ♀, 6 nymphs (CAS 9071523 [MGF020]), Sept Lacs, 23°31′42″S 44°09′20″E, 8 March 2002, Frontier Project; 5 ♂, 3 ♀, 6 nymphs (CAS 9071770 [MGF028]), Sept Lacs, 23°31′29″S 44°09′33″E, 10 March 2002; Frontier Project; 3 ♂, 3 ♀, 10 nymphs (CAS 9071522 [MGF030]), Manderano, 23°31′38″S 44°05′15″E, 10–14 March 2002, Frontier Wilderness Project; 6 ♂, 6 ♀, 8 nymphs (CAS 9071753 [MGF033]), Manderano, 23°31′27″S 44°05′34″E, 29 May–1 June 2002, Frontier Wilderness Project; 3 ♂, 4 ♀, 5 nymphs (CAS 9072461 [MGF034]), Manderano, 23°31′24″S 44°05′40″E, 8–12 June 2002, Frontier Wilderness Project; 1 ♂, 2 nymphs (CAS 9071731 [MGF037]), Sept Lacs, 23°31′15″S 44°8′35″E, 20–24 August 2002, Frontier Wilderness Project; 3 nymphs (CAS 9071780 [MGF040]), 1 nymph (CAS 9072465 [MGF043]), Fiherenana, 23°10′37″S 43°57′39″E, 21–24 October 2002, Frontier Wilderness Project; 2 ♂, 2 ♀, 2 nymphs (CAS 9071745 [MGF050]), Fiherenana, 23°14′07″S 43°52′15″E, 1–4 December 2002, Frontier Wilderness Project; 2 ♂, 1 ♀, 3 nymphs (CAS 9071774 [MGF056]), Ranobe, 23°02′03″S 43°36′43″E, 5–9 February 2003, Frontier Wilderness Project; 2 ♂, 1 nymph (CAS 9071517 [MGF061]), Lake Ranobe, 23°02′23″S 43°36′39″E, 17–21 February 2003, Frontier Wilderness Project; 3 ♂, 4 ♀, 1 nymph (CAS 9072418 [MGF064 (14120(1))]), Lake Ranobe, 23°02′21″S 43°36′42″E, 25–28 April 2003, Frontier Wilderness Project; 1 nymph (CAS 9071722 [MGF070]), Lake Ranobe, 23°02′44″S 43°36′58″E, 17–21 May 2003, Frontier Wilderness Project; 1 ♀, 1 nymph (CAS 9071644 [MGF074]), Fiherenana, 23°13.351′S 43°52.853′E, 5–7 August 2003, Frontier Wilderness Project; 1 nymph (CAS 9071649 [MGF077]), Fiherenana, 23°10.619′S 43°57.685′E, 18–19 August 2003, Frontier Wilderness Project; 1 ♂, 1 ♀ (CAS 9072464 [MGF092]), Ranobe, 23°02.667′S 43°36.917′E, 26–29 January 2004, Frontier Wilderness Project; 21 ♂, 8 ♀, 47 nymphs (CAS 9071597 [BLF5850]), Foret de Mite, 23°31′27″S 44°07′17″E, 27 February–3 March 2002, Fisher, Griswold et al.

Diagnosis

Very similar to T. pumila sp. nov. but larger (e.g., body length 1.76/2.23 ♂/♀ versus 1.56/1.96 ♂/♀ in T. pumila sp. nov.; femur length 0.49/0.61 ♂/♀ versus 0.44/0.53 ♂/♀ in T. pumila sp. nov.); more slender femora (1.66/1.69 ♂/♀ times longer than broad versus 1.63/1.63 ♂/♀ in T. pumila sp. nov.).

Etymology

This species is a patronym honoring Volker Mahnert who worked on African pseudoscorpions.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 23A): 1.33–1.42 (♂), 1.32–1.37 (♀) times longer than broad. — Pedipalps (Fig. 23C, D): See trichobothrial pattern in description of T. pumila sp. nov.; Chelal fixed finger with 10–11 chelal teeth in the OR, 12–16 in the MR and 8–14 in the IR; movable finger with 6–8 in the OR, 9–11 in the MR and 8–10 teeth in the IR; 7 equally sized and one larger terminal tooth on fixed finger and 8 equally sized terminal teeth on movable finger. — Dimensions (mm): Holotype ♂: Body length 1.77; abdomen length 1.21, width (with pleura) 1.13, width (without pleura) 0.99; carapace length 0.52, width 0.39. Pedipalp: trochanter 0.26; femur length 0.49, width 0.30; patella 0.41; chela (without pedicel) 0.44; hand length 0.10, width 0.13; movable finger 0.36. Leg I: trochanter 0.08; femur 0.17; patella 0.15; tibia 0.15; tarsus 0.24. Leg IV: trochanter 0.15; femur 0.12; patella 0.26; tibia 0.23; tarsus 0.30. Allotype ♀: Body length 2.34; abdomen length 1.70, width (with pleura) 1.13, width (without pleura) 1.22; carapace length 0.59, width 0.43. Pedipalp: trochanter 0.26; femur length 0.61, width 0.37; patella 0.47; chela (without pedicel) 0.48; hand length 0.12, width 0.15; movable finger 0.39. Leg I: trochanter 0.10; femur 0.17; patella 0.20, tibia 0.18; tarsus 0.27. Leg IV: trochanter 0.18; femur 0.14; patella 0.31; tibia 0.32; tarsus 0.34.

Figure 23. 

Drawings of Toliaranella mahnerti sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Body length 1.64–1.88 (♂), 2.04–2.54 (♀); abdomen length 1.14–1.25 (♂), 1.44–1.86 (♀); width (with pleura) 1.03–1.15 (♂), 1.29–1.51 (♀); width (without pleura) 0.87–0.99 (♂), 1.14–1.31 (♀); 1.21–1.33 (♂), 1.26–1.42 (♀) times longer than broad; carapace length 0.50–0.53 (♂), 0.56–0.60 (♀); width 0.35–0.39 (♂), 0.42–0.44 (♀); 1.33–1.43 (♂), 1.32–1.40 (♀) times longer than broad. Pedipalp: trochanter 0.21–0.26 (♂), 0.24–0.29 (♀); femur length 0.46–0.52 (♂), 0.56–0.65 (♀); width 0.28–0.32 (♂), 0.34–0.37 (♀); 1.53–1.79 (♂), 1.61–1.78 (♀) times longer than broad; patella 0.37–0.41 (♂), 0.47–0.50 (♀); chela (without pedicel) 0.41–0.45 (♂), 0.48–0.52 (♀); hand length 0.07–0.11 (♂), 0.11–0.14 (♀); width 0.11–0.13 (♂), 0.14–0.15 (♀); 0.58–0.92 (♂), 0.79–0.93 (♀) times longer than broad; movable finger 0.33–0.36 (♂), 0.39–0.41 (♀).

Habitat

Specimens were found in leaf litter in spiny forest thicket, degraded and undegraded gallery forest, and in disturbed and undisturbed riparian forest; elevation 20–160 m. All close to a river stream.

GenBank Code

OP589954, OP589955, OP589956, OP589957, OP589958. The species differs from all congeners by more than 5.2% in the H3 dataset.

Distribution

Presently known from 17 localities in the Atsimo-Andrefana Region (formerly Toliara Province): type locality (MGF001) in southwestern Madagascar close to the Onilahy River delta and additional localities in the radius of 9 km upstream along the right side of the Onilahy River and around the Fiherenana River (MGF004, MGF020, MGF028, MGF030, MGF033, MGF034, MGF037, MGF043, MGF040, MGF050, MGF056, MGF061, MGF064, MGF070, MGF074, MGF077, MGF092).

Toliaranella meridionalis sp. nov.

Type material examined

MADAGASCAR: Toliara Province [Androy Region]: Holotype ♂ (CAS 9071748 [BLF5570]), allotype ♀ (ZMH-A0016751), 4 ♂ paratypes (CAS 9071748 [BLF5570 and BLF5650]), 2 ♂ (ZMH-A0016752, ZMH-A0016753), Reserve Speciale de Cap Sainte Marie, 25°35′40″S 45°8′49″E, 13–19 February 2002, B.L. Fisher, Griswold et al.

Additional material examined

MADAGASCAR: Toliara Province [Androy Region]: 8 ♂, 14 ♀, 5 nymphs (CAS 9071596 [BLF5500]), Reserve Speciale de Cap Sainte Marie, 25°34′54″S 45°10′6″E, 11–15 February 2002, Fisher, Griswold et al.

Diagnosis

Largest body size of the southern Malagasy feaellids (e.g., body length 1.85/2.33 ♂/♀ versus 1.56–1.80/1.96–2.23 ♂/♀; carapace length 0.53/0.64 ♂/♀ versus 0.45–0.51/0.52–0.58 ♂/♀; femur length 0.55/0.67 ♂/♀ versus 0.44–0.53/0.53–0.61 ♂/♀)

Etymology

This species is named after the type locality, which is the southernmost point of Madagascar.

Description

The following description is based on holotype and allotype. — Carapace (Fig. 24A): 1.25–1.38 (♂), 2.64 (♀) × longer than broad. — Pedipalps (Fig. 24C, D): See trichobothrial pattern in description of T. pumila sp. nov. Chelal fixed finger with 9–11 teeth in the OR, 13–14 in the MR and 10 in the IR; movable finger with 8–9 teeth in the OR, 11–14 in the MR and 9 teeth in the IR; terminal teeth: 7 equally sized and one large one on the fixed finger, 7 teeth on the movable finger equal in size. — Dimensions (mm): Holotype ♂: Body length 1.82; abdomen length 1.29, width (with pleura) 1.18, width (without pleura) 1.04; carapace length 0.50, width 0.40. Pedipalp: trochanter 0.24; femur length 0.59, width 0.34; patella 0.44; chela (without pedicel) 0.43; hand length 0.10, width 0.13; movable finger 0.34. Leg I: trochanter 0.10; femur 0.13; patella 0.17; tibia 0.17; tarsus 0.22. Leg IV: trochanter 0.16; femur 0.12; patella 0.26; tibia 0.24; tarsus 0.30. Allotype ♀: Body length 2.33; abdomen length 1.69, width (with pleura) 1.50, width (without pleura) 1.30; carapace length 0.64, width 0.46. Pedipalp: trochanter 0.27; femur length 0.67, width 0.40; patella 0.52; chela (without pedicel) 0.50; hand length 0.11, width 0.16; movable finger 0.41. Leg I: trochanter 0.11; femur 0.20; patella 0.21; tibia 0.15; tarsus 0.29. Leg IV: trochanter 0.18; femur 0.16; patella 0.35; tibia 0.34; tarsus 0.34.

Figure 24. 

Drawings of Toliaranella meridionalis sp. nov.: A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

Variation

Allotype ♀: Body length 1.75–1.90 (♂), 2.33 (♀); abdomen length 1.17–1.36 (♂), 1.69 (♀); width (with pleura) 1.11–1.24 (♂), 1.50 (♀); width (without pleura) 1.01–1.09 (♂), 1.30 (♀); 1.16–1.26 (♂), 1.30 (♀) × longer than broad; carapace in length 0.50–0.55 (♂), 0.64 (♀); width 0.40–0.42 (♂), 0.46 (♀); 1.25–1.38 (♂), 1.39 (♀) × longer than broad. Pedipalp: trochanter 0.21–0.29 (♂), 0.27 (♀); femur length 0.52–0.59 (♂), 0.67 (♀); width 0.32–0.35 (♂), 0.40 (♀); 1.59–1.74 (♂), 1.68 (♀) × longer than broad; patella 0.43–0.47 (♂), 0.52 (♀); chela (without pedicel) 0.42–0.45 (♂), 0.50 (♀); hand length 0.90–0.12 (♂), 0.11 (♀); width 0.13–0.14 (♂), 0.16 (♀); 0.64–0.86 (♂), 0.69 (♀) × longer than broad; movable finger 0.32–0.36 (♂), 0.41 (♀).

GenBank Code

OP589961. The species differs from all congeners by more than 3.3% in the H3 dataset.

Habitat

Specimens found in sifted litter from spiny forest thicket very close to the coastline at an elevation of 160 m.

Distribution

Known from the two southernmost localities (type locality and BLF5500) among all Malagasy feaellids in the Androy Region (formerly Toliara Province).

4. Discussion

4.1. Biogeography

The only available studies specifically addressing Malagasy pseudoscorpions are short notes (Vachon 1960, Heurtault 1986) pointing out that all families present, including Feaellidae, do not seem to have undergone excessive in-situ speciation and that many genera are shared with continental Africa, southern India, Australia or Southeast Asia (Heurtault 1986). The present contribution is the first detailed study on a pseudoscorpion lineage in Madagascar and we point out that, after detailed examination, it is evident that at least the Feaellidae of Madagascar are a highly endemic fauna at both the generic and species level. We argue that this is most likely true for other pseudoscorpion families occurring on Madagascar but none of these have been revised taxonomically. Although there appear to be no families restricted to the island, endemism might in fact be substantial at the genus and species level and initial assessment of low endemism may result from the general problems of pseudoscorpion taxonomy, such as cryptic morphologies that change little over time, perhaps because of niche conservatism and retainment of ancestral ecological characters in soil-dwelling fauna (Wiens and Graham 2005). Undoubtedly detailed morphological studies in conjunction with genetic data will be necessary to fully unravel the evolutionary history of the Malagasy pseudoscorpion fauna.

Feaellids are an ancient lineage with a fossil origin that dates back to the Late Triassic and Late Cretaceous (Henderickx and Boone 2016; Harms and Dunlop 2017; Kolesnikov et al. 2022) and molecular clock estimates suggest the lineage originated in the Paleozoic (Benavides et al. 2019). No molecular phylogeny currently exists for the family Feaellidae but it can be hypothesized that the origins of Malagasy Hercules pseudoscorpions is Gondwanan, similar to other ancient invertebrate lineages in soil habitats such as pill millipedes (Wesener et al. 2010). Under a continental vicariance scenario, we predict this fauna to be closely allied with that of the Seychelles and southern India, followed by southern Africa, and then tropical Africa (Yoder and Nowak 2006) – a scenario that could be tested in a molecular phylogenetic framework because feaellids occur in all three of these regions and at least those from India, Maldives and the Seychelles are morphologically quite similar (e.g., Mahnert 1978; Novák et al. 2020) but differ from those in Brazil and Australia (e.g., Harvey et al. 2016a, b) and also Southeast Asia (Judson 2017). We hypothesize further that a dated molecular phylogeny should reflect continental vicariance to some degree, although we cannot rule out the possibility that some of the new genera described herein are older. Not only are feaellids an ancient Pangaean lineage, but the molecular data presented here and elsewhere (Harvey et al. 2016a) clearly refute a historical concept based on morphology that considers many feaellid species to be widespread. In line with recent molecular studies on other soil-dwelling pseudoscorpions (e.g., Cosgrove et al. 2016; Harms 2018; Harms et al. 2019), the case is rather the opposite and feaellids appear to show extreme microendemism, habitat restriction and pronounced population structuring despite the absence of morphological disparity, perhaps comparable to some mygalomorph spiders that typically thrive in localized populations with limited dispersal between them (Greenberg et al. 2021) or mite harvestmen occurring also in soil habitats (De Bivord and Giribet 2010; Jay et al. 2016). Vicariance at various spatial scales (continental, regional and local) and different time periods seems to be a major factor that needs to be tested, although at least some feaellids occur in coastal habitats (Beier 1955) and might disperse, as recently suggested for a species found in the Maldives (Novák et al. 2020). Overall, our data are congruent with a hypothesis of pronounced endemism at all spatial scales, vicariance as a common case for diversification, and small species ranges in specific habitats in the presence of relative morphological stasis that may be caused by niche conservatism. Feaellids therefore emerge as a new model group to test hypotheses pertaining to vicariance biogeography.

4.2. In-situ speciation and causes

Vences et al. (2009) reviewed possible diversification mechanisms for Madagascar and distinguished between ecographical constraints (meaning adaptive radiation in response to climatic shifts), speciation in rainforest refugia, montane refugia, riverine barriers, and river catchments as testable scenarios. Many studies on invertebrates have emphasized the importance of mountain or rainforest refugia (Wesener et al. 2011; Wood et al. 2015) and pointed out that in-situ speciation in such refugia, together with time and adaptive niche radiation, has been fundamentally important in diversification. Many taxonomic revisions have also focused on taxa that are exclusively or predominantly found in the high-altitude forests of the east (Griswold 1997; Griswold et al. 2012; Wood and Scharff 2018) and documented substantial microendemism. For more arid habitats, a study by Wesener (2009) found that endemism in pill millipedes from dry forests was not as pronounced as that of species occurring in mesic habitats. Comprehensive taxonomic revisions on goblin spiders (Family Oonopidae Simon, 1890) have also documented major endemism at the generic level for the Malagasy fauna (Álvarez-Padilla et al. 2012; Saucedo et al. 2015), but wide species distributions in arid habitats that roughly correspond with the generic distributions in our dataset. Perhaps the highest similarity is with the distribution of the goblin spider genus Opopaea Simon, 1891, where species endemism is high even in arid habitats (Andriamalala and Hormiga 2013). So, what is driving the high diversity of feaellids in Madagascar?

Adaptive radiation in response to climatic zonation is a likely hypothesis for the three feaellid genera described in our paper. Antsiarananaella gen. nov. is clearly restricted to the dry deciduous savannah forests of the north, a biome that is well known as a center of endemism in both vertebrates and invertebrates (e.g., Brown et al. 2016). The Irodo and Loky Rivers seem to be major barriers here that define species ranges to some extent and divide the ranges of A. faulstichi sp. nov. and A. leniae sp. nov. and A. marlae sp. nov. although forest cover and topography also seem to be important. The distribution of Mahajanganella gen. nov. encompasses the dry deciduous forests of the western Malagasy biome and species seem a little more widespread and river drainages do not seem as important here, although our species hypotheses are primarily based on morphology and we cannot rule out the possibility that more widespread species such as M. heraclis sp. nov. might show significant genetic structuring or cryptic speciation. Toliaranella gen. nov. is restricted to the subarid biome of the south, which is primarily covered by arid spiny bush vegetation. Our maps and genetic data (whenever available) clearly show that river drainages can define species ranges, such as with T. mahnerti sp. nov. that is found in the Fiherenana and Onilahy drainages but nowhere else. The Mandrare drainage also has a distinct species (T. fisheri sp. nov.) although some species such as T. meridionalis sp. nov., are found in coastal drainages of the Cape St. Marie Special Reserve in the south. It appears that ecographical constraints are effective at the generic level in Malagasy Hercules pseudoscorpions, whereas river catchments and sclerophyll forest refugia offer a testable framework to derive species limits in this fauna. It might also be pointed out that feaellids are absent from two of the major Malagasy biomes which is the central highlands and the evergreen forests to the east. These are also the regions that receive the highest annual rainfall with >1500 ml per year, whereas areas occupied by feaellids altogether receive less than 1000 ml. It appears that Malagasy feaellids are absent from rainforest habitats, unlike the fauna of central and western Africa or species recorded from Brazil that have been collected from rainforest habitats (e.g., Harvey et al. 2016b) or Southeast Asia where the only records are from caves (Judson 2017; Harvey 2018). Ecologically this fauna seems to be more closely aligned with that of eastern Africa where feaellids have been collected from dry leaf litter in savannah or near-coastal habitats (Beier 1966; Mahnert 1982; Henderickx 2009). Interestingly, some other pseudoscorpion lineages in Madagascar show exactly the opposite pattern, such as the family Pseudotyrannochthoniidae Beier, 1932 that seems to be restricted to the central highlands and the evergreen forests according to survey records. Revising these taxonomically could complement our understanding of the Malagasy pseudoscorpion fauna and help generate a more comprehensive picture of pseudoscorpion diversification patterns on the island.

4.3. Conservation

Madagascar is amongst the “hottest” biodiversity hotspots and the conservation of the remaining natural or semi-natural habitats is crucial to ensure lineage survival under intense anthropogenic pressure. It is therefore relevant to note that none of the species described in this study come from strictly protected areas or reserves such as Tsingy de Bemaha Reserve, Betampona Reserve, Tsaratanana Reserve, or Zahamena Reserve, but that some species occur in areas that have gained some conservation status after the specimens were collected: Ankarafantsika National Park (since 2002) and Tsingi de Namoroka National Park (since 2002) for M. heraclis sp. nov. (collected in 2001), and Montagne des Français Protected Area (since 2013) for A. lorenzorum sp. nov. Feaellids habitats are also conserved in the Baie de Baly National Park (M. schwarzeneggeri sp. nov.), Kirindy Mitea National Park (M. fridakahloae sp. nov.), the Analamerana Reserve (A. faulstichi sp. nov.), Andranomena Reserve (M. fridakahloae sp. nov.), Cap Sainte Marie Reserve (T. meridionalis sp. nov.), and Ankarana Special Reserve (since 1956 for A. faulstichi sp. nov.). We emphasize here that many other species with narrow distributional ranges occur in habitats that do not have any conservation status at present, such as T. fisheri sp. nov. Conservation incentives should target these species and their habitats whenever possible.

4.4. Systematics & Taxonomy

In this paper, we describe three new genera of Hercules pseudoscorpions, each restricted to a specific biome and climate of Madagascar, and each of these containing multiple species with short ranges. Aside from biogeographical and conservation aspects, our study also highlights the need to revise generic concepts in the family Feaellidae, specifically the genus Feaella in the Afrotropics that is presently divided into the subgenera: Feaella (Feaella), Feaella (Tetrafeaella) and Feaella (Difeaella) based on the number of anterior lobes of the carapace ranging from two to six (Beier 1966; Mahnert 1978). The most common combination is four lobes as in Feaella (Tetrafeaella) and this condition is shared by all Malagasy species although these species are dissimilar from the type species, Feaella (Tetrafeaella) cf. mucronata (Figs 25, 26, 27) from Amanzimtoti in South Africa, both morphologically and genetically. Species from Australia, the Seychelles, the Maldives and the Indian subcontinent are also presently classified into the subgenus Tetrafeaella, which makes little sense biogeographically and it is evident that the distinguishing character (number of lobes) is of limited phylogenetic relevance.

Figure 25. 

Feaella (Tetrafeaella) cf. mucronata (Tullgren, 1907), scanning electron micrographs. Specimen used for imaging: F. cf. mucronata female paratype (ZMH-A0010793). A carapace, dorsal view; B sternum ventral overview; C coxal spines in detail (highlighted in red). Scale bars: 0.2 mm (A, B); 0.1 mm (C).

Figure 26. 

Feaella (Tetrafeaella) cf. mucronata (Tullgren, 1907), scanning electron micrographs. Specimen used for imaging: F. cf. mucronata female paratype (ZMH-A0010793). A right chela from ventral with specialized setae in a group in detail (highlighted in red); B left chela with double trichobothrium dt in detail (highlighted in red); C left chela from medial with terminal teeth in a compact group in detail. Scale bar: 0.2 mm.

Figure 27. 

Drawings of Feaella (Tetrafeaella) cf. mucronata (Tullgren, 1907): A carapace from dorsal; B leg IV; C left chela from dorsal with trichobothrial pattern; D right pedipalp from dorsal. Scale bars: 0.2 mm (AC); 0.4 mm (D).

It is not the aim of this study to test the generic limits within Feaellidae. However, a phylogenetic analysis with a detailed morphological investigation, including scanning electron microscopy and perhaps even micro-computed tomography, should be undertaken to resolve the phenetic classification. Our study may provide an avenue insofar as we define new genera and have identified morphological characters that had been overlooked or not given widespread attention, such as the sensory apparatus and setae, chelal tooth morphology, and cuticle sculpturing, which is an obvious feature in all feaellids that generally have a thick and armed body cuticle. Focusing on minute details unfortunately seems to be important in a morphologically highly conserved pseudoscorpion lineage where species are similar across continents despite significant genetic divergences (e.g., Harvey et al. 2016a). Analyzing the causes for such a conserved morphology and the resulting morphological crypsis of lineages, both at the species and generic level, might be an interesting task for the future, not only for Feaellidae, but also for some other pseudoscorpion lineages because strong selective pressures might exist limiting morphological change over millions of years and across continents. Alternatively, this pattern may also represent an exceptional case of ecological niche conservation reflected by morphological stasis. Since feaellids are an ancient invertebrate lineage with possible origins in the Carboniferous (Benavides et al. 2019), the low morphological diversity might be a result of the latter, but virtually nothing is known about feaellid biology and ecology and that discussion is left for the future.

5. Acknowledgements

We thank Brian Fisher and Charles Griswold (California Academy of Sciences) for their outstanding contribution to biodiversity research in Madagascar, to Lauren Esposito and Darrell Ubick for facilitating the loan of specimens from the California Academy of Sciences, to Christoph Abel (Universität Hamburg) for assistance with molecular laboratory work, and to Nadine Dupérré (LIB Hamburg) for help with imaging work and sample curation.

6. References

  • Agnarsson I, Jenik BB, Veve GM, Hanitriniaina S, Agostini D, Goh SP, Pruitt J, Kuntner M (2015) Systematics of the Madagascar Anelosimus spiders: remarkable local richness and endemism, and dual colonization from the Americas. ZooKeys 509: 13–52. https://doi.org/10.3897/zookeys.509.8897
  • Ali JR, Aitchison JC (2008) Gondwana to Asia: Plate tectonics, paleogeography and the biological connectivity of the India sub- continent from the Middle Jurassic through latest Eocene (166–35 Ma). Earth Science Reviews 88: 145–166. https://doi.org/10.1016/j.earscirev.2008.01.007
  • Álvarez-Padilla F, Ubick D, Griswold CE (2012) Noideattella and Tolegnaro, two new genera of goblin spiders from Madagascar, with comments on the gamasomorphoid and silhouettelloid oonopids (Araneae, Oonopidae). American Museum Novitates 3745: 1–76. https://doi.org/10.1206/3745.2
  • Andriamalala D, Hormiga G (2013) Systematics of the goblin spider genus Opopaea (Araneae, Oonopidae) in Madagascar. Bulletin of the American Museum of Natural History 380: 1–156. https://doi.org/10.1206/828.1
  • Beier M (1955) Pseudoscorpionidea, gesammelt während der schwedischen Expeditionen nach Ostafrika 1937–38 und 1948. Arkiv för Zoologi 7: 527–558.
  • Beier M (1966) Ergänzungen zur Pseudoscorpioniden-Fauna des südlichen Afrika. Annals of the Natal Museum 18: 455–470.
  • Benavides LR, Cogrove JG, Harvey MS, Giribet G (2019) Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones Tree of Life. Molecular Phylogenetics and Evolution 139: 106509. https://doi.org/10.1016/j.ympev.2019.05.023
  • Chamberlin JC (1931) The arachnid order Chelonethida. Stanford University Publications, Biological Sciences 7: 1–284.
  • Cosgrove JG, Agnarsson I, Harvey MS, Binford GJ (2016) Pseudoscorpion diversity and distribution in the West Indies: sequence data confirm single island endemism for some clades, but not others. Journal of Arachnology 44: 257–271. http://www.jstor.org/stable/24892348
  • De Bivort BL, Giribet G (2010) A systematic revision of South African Pettalidae (Arachnida: Opiliones: Cyphophthalmi) based on a combined analysis of discrete and continuous morphological characters with the description of seven new species. Invertebrate Systematics 24: 371–406. https://doi.org/10.1071/IS1001
  • Edler D, Klein J, Antonelli A, Silvestro D (2020) raxmlGUI 2.0: a graphical interface and toolkit for phylogenetic analyses using RAxML. Methods in Ecology and Evolution 12: 373–377. https://doi.org/10.1111/2041-210X.13512
  • Greenberg MR, Huey JA, Framenau VW, Harms D (2021) Three new species of mouse spider (Araneae: Actinopodidae: Missulena Walckenaer, 1805) from Western Australia, including an assessment of intraspecific variability in a widespread species from the arid biome. Arthropod Systematics and Phylogeny 79: 509–533. https://doi.org/10.3897/asp.79.e62332
  • Griswold CE, Wood HM, Carmichael AD (2012) The lace web spiders (Araneae, Phyxelididae) of Madagascar: phylogeny, biogeography and taxonomy. Zoological Journal of the Linnean Society 164: 728–810. https://doi.org/10.1111/j.1096-3642.2011.00779.x
  • Harms D (2018) The origins of diversity in ancient landscapes: deep phylogeographic structuring in a pseudoscorpion (Pseudotyrannochthoniidae: Pseudotyrannochthonius) reflects Plio-Pleistocene climate fluctuations. Zoologischer Anzeiger 273: 112–123. https://doi.org/10.1016/j.jcz.2018.01.001
  • Harms D, Roberts JD, Harvey MS (2019) Climate variability impacts on diversification processes in a biodiversity hotspot: a phylogeography of ancient pseudoscorpions in southwestern Australia. Zoological Journal of the Linnean Society 186: 934–949. https://doi.org/10.1093/zoolinnean/zlz010
  • Harvey MS (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6: 1373–1435. https://doi.org/10.1071/IT9921373
  • Harvey MS (2022) World Pseudoscorpiones Catalog (2022). Natural History Museum Bern, online at http://wac.nmbe.ch (accessed on 27.06.2022).
  • Harvey MS, Abrams KM, Beavis AS, Hillyer MJ, Huey JA (2016a) Pseudoscorpions of the family Feaellidae (Pseudoscorpiones : Feaelloidea) from the Pilbara region of Western Australia show extreme short-range endemism. Invertebrate Systematics 30: 491–508. https://doi.org/10.1071/IS16013
  • Harvey MS, Andrade Rd, Pinto-da-Rocha R (2016b) The first New World species of the pseudoscorpion family Feaellidae (Pseudoscorpiones: Feaelloidea) from the Brazilian Atlantic Forest. Journal of Arachnology 44: 227–234. https://doi.org/10.1636/JoA-S-16-007
  • Henderickx (2009) A new Feaella species (Pseudoscorpiones: Feaellidae) from Kenya. Phegea 37: 41–47.
  • Henderickx H, Boone M (2014) The first fossil Feaella Ellingsen, 1906, representing an unexpected pseudoscorpion family in Baltic amber (Pseudoscorpiones, Feaellidae). Entomo-Info 25: 5–11.
  • Henderickx H, Boone M (2016) The basal pseudoscorpion family Feaellidae Ellingsen, 1906 walks the earth for 98.000.000 years: a new fossil genus has been found in Cretaceous Burmese amber. Entomo-Info 27: 7–13.
  • Heurtault J (1983) Pseudoscorpions de Côte d’Ivoire. Revue Arachnologique 5: 1–27.
  • Heurtault J (1986) Les Pseudoscorpions de Madagascar: réflexions sur la répartition géographique. In: Eberhard WG, Lubin YD, Robinson BC (Eds) Proceedings of the Ninth International Congress of Arachnology, Panama 1983. Smithsonian Institution Press, Washington D.C., U.S.A., 127–129.
  • Heurtault-Rossi J, Jézéquel JF (1965) Observations sur Feaella mirabilis Ell. (Arachnide, Pseudoscorpion). Les chélicères et les pattes-mâchoires des nymphes et des adultes. Description de l’appareil reproducteur. Bulletin du Muséum national d’Histoire naturelle, Paris (2) 37: 450–461.
  • Jay KR, Popkin-Hall ZR, Coblens MJ, Oberski JT, Sharma PP, Boyer SL (2016) New species of Austropurcellia, cryptic short-range endemic mite harvestmen (Arachnida, Opiliones, Cyphophthalmi) from Australia’s Wet Tropics biodiversity hotspot. ZooKeys 586: 37–93. https://doi.org/10.3897/zookeys.586.6774
  • Katoh K, Misawa K, Kuma K, Miyata T (2002) MAFFT: A novel method for rapid multiple alignment based on fast Fourier transform. Nucleic Acids Research 30: 3059–3066. https://doi.org/10.1093/nar/gkf436
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
  • Katoh K, Toh H (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics 9: 286–298. https://doi.org/10.1093/bib/bbn013
  • Kolesnikov VB, Turbanov IS, Eskov KY, Propistsova EA, Bashkuev AS (2022) First non-amber Mesozoic pseudoscorpion from Upper Triassic deposits of eastern Europe, with a description of two new fossil subfamilies (Arachnida, Pseudoscorpiones, Feaellidae). Papers in Palaeontology 8(5): e1466. https://doi.org/10.1002/spp2.1466
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees, pp. 1–8. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA.
  • Mittermeier RA, Robles-Gil P, Hoffmann M, Pilgrim JD, Brooks TB, Mittermeier CG, Lamoreux JL, Fonseca GAB (2004) Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecoregions. CEMEX, Mexico City: 1–391.
  • Morelli TL, Smith AB, Mancini AN, Balko EA, Borgerson C, Dolch R, Farris Z, Federman S, Golden CD, Holmes SM, Irwin M, Jacobs RL, Johnson S, King T, Lehman SM, Louis Jr EE, Murphy A, Randriahaingo NT, Randrianarimanana HLC, Ratsimbazafy J, Razafindratsima OH, Baden AL (2020) The fate of Madagascar’s rainforest habitat. Nature Climate Change 10: 89–96. https://doi.org/10.1038/s41558-019-0647-x
  • Novák J, Lorenz M, Harms D (2020) Feaella (Tetrafeaella) obscura sp. nov. – a new pseudoscorpion species from the Maldives (Arachnida: Pseudoscorpiones), and an updated identification key to the subgenus Feaella (Tetrafeaella). Zoosystematics and Evolution 96: 769–779. https://doi.org/10.3897/zse.96.56885
  • Saucedo AD, Ubick D, Griswold CE (2015) The goblin spiders of the new genus Volborattella (Araneae: Oonopidae) from Madagascar. American Museum Novitates 3822: 1–72. https://doi.org/10.1206/3822.1
  • Vachon M (1960) Sur la présence a Madagascar d’un représentant de la famille des Feaellidae [sic] Ellingsen (Pseudoscorpions). Bulletin du Muséum National d’Histoire Naturelle, Paris 32(2): 165–166.
  • Wesener T (2009) Unexplored richness: discovery of 31 new species of Giant Pill-Millipedes endemic to Madagascar, with a special emphasis on microendemism (Diplopoda, Sphaerotheriida). Zootaxa 2097: 1–134. https://doi.org/10.11646/zootaxa.2097.1
  • Wesener T, Raupach MJ, Sierwald P (2010) The origins of giant pill-millipedes from Madagascar (Diplopoda: Sphaeroteriida: Arthrosphaeridae). Molecular Phylogenetics and Evolution 57: 1184–1193. https://doi.org/10.1016/j.ympev.2010.08.023
  • Wesener T, Raupach MJ, Decker P (2011) Mountain refugia play a role in soil arthropod speciation on Madagascar: A case study of the endemic giant fire-millipede genus Aphistogoniulus. PLoS ONE 6: e28035. https://doi.org/10.1371/journal.pone.0028035
  • Wiens JJ, Graham CH (2005) Niche Conservatism: Integrating evolution, ecology, and conservation biology. Annual Review of Ecology, Evolution, and Systematics 36: 519–539. https://www.jstor.org/stable/30033815
  • Wood HM, Gillespie RG, Griswold CE, Wainwright PC (2015) Why is Madagascar special? The extraordinarily slow evolution of pelican spiders (Araneae, Archaeidae). Evolution 69: 62–481. https://doi.org/10.1111/evo.12578
  • Wood HM, Scharff N (2018) A review of the Madagascan pelican spiders of the genera Eriauchenius O. Pickard-Cambridge, 1881 and Madagascarchaea gen. n. (Araneae, Archaeidae). ZooKeys 727: 1–96. https://doi.org/10.3897/zookeys.727.20222
  • Yoder AD, Nowak MD (2006) Has vicariance or dispersal been the predominant biogeographic force in Madagascar? Only time will tell. Annual Review of Ecology, Evolution, and Systematics 37: 405–431. https://www.jstor.org/stable/30033838