1 Chapter 2

2 Class Collembola
3
4Nikolas G. Cipola
5Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil.
6
7Diego D. da Silva
8Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil.
9
10José G. Palacios-Vargas
11Universidad Naćional Autónoma de México, México
12
13Bruno C. Bellini
14Universidade Federal do Rio Grande do Norte, Natal, Brazil
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16
17Introduction
18Systematics of Collembola
19Order Poduromorpha
20Order Entomobryomorpha
21Order Neelipleona, Family Neelidae
22Order Symphypleona
23Limitations
24Terminology and Morphology
25Material Preparation and Preservation
26Key to Freshwater Collembola
27References
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51 INTRODUCTION
52
53 Springtails (Hexapoda: Collembola) are small terrestrial arthropods very similar to
54insects by tagmosis patterns and presence of one pair of antennae and three pairs of thoracic
55legs. They differ from all other hexapods by presence of three abdominal appendages: ventral
56tube (or collophore), tenaculum (or retinaculum) and furcula. These appendages are only seen
57in springtails and are considered as synapomorphies of the taxon (Hopkin, 1997). The
58taxonomic name “Collembola” was proposed by Lubbock in 1870 based on the presence of
59the collophore in species of the genus Podura Linnæus. In this sense, Lubbock observed that
60the appendage was responsible to attach specimens on a surface and used the latin/greek
61terminology to systematic redefine Podura: colla (Latin) or kolla (Greek) means glue, and
62embolon (Greek) means “that what has been thrown into something”, in this case, like a plug
63(Bellinger et al., 2016).
64 Historically springtails were considered firstly as insects and secondly as a non-insect
65taxon of entognathous hexapods. Entognathy is a condition of hexapods in which the
66mouthparts are almost completely hidden within head capsule and cannot be clearly seen,
67contrasting to the condition of true insects which bear large exposed mouthparts and for that
68are called “ectognathous”. Since proturans and diplurans also share entognathy, the three taxa
69were grouped in “Entognatha”, a taxon not supported by current phylogenetical surveys (e. g.
70Misof et al., 2014). In fact, nowadays, Hexapoda validity is questionable and it is supposed
71neither Collembola nor Protura are closely related to Diplura+Insecta. In this case, at least
72Collembola apparently evolved directly from marine crustacean ancestors, possibly non-
73related to other hexapods (Bellinger et al., 2016).
74 Regardless of the phylogenetic position of Collembola among other mandibulate
75arthropods, the group is a well-established monophyletic taxon based on several
76morphological features, especially abdominal appendages. Nowadays most classification
77systems still retains Collembola inside Hexapoda, as an independent class with four orders
78(Poduromorpha, Entomobryomorpha, Symphypleona and Neelipleona, Fig. 1) which hold
79together about 8,600 species in 700 genera and 29 families (Soto-Adames et al., 2008;
80Bellinger et al., 2016). However, this substantial species richness possibly represents a small
81slice of the real diversity of living forms, which may surpass 50,000 species (Hopkin, 1997;
82Deharveng et al., 2008; Cicconardi et al., 2013).
83 The potential high species richness of springtails is possibly related to small size of
84specimens, restricted dispersion capacity, widespread distribution in terrestrial and part of
85aquatic ecosystems and specialization of these populations to different microhabitats.
86Concerning aquatic habitats, springtails also present specializations to different (micro)
87habitats and can be ecologically categorized as primary (water-dependent) or secondary
88(epigean hydrophilous) aquatic associate species (Waltz & McCafferty, 1979; Deharveng et
89al., 2008). The primary aquatic associate springtails can be: cryophilous species, which live
90permanently on the top of snow in temperate mountains or glaciers; epineustic species, found
91effectively on the surface of freshwater, usually with adaptive morphological structures, such
92in some species of Sminthurides, Arlesminthurus and Pseudobourletiella; and intertidal (or
93littoral) species which live associate to beach soil, rocks and water puddles and are capable of
94being submerged, as species of Spinactaletes and Isotogastrura. The secondary aquatic
95associate springtails are usually hygrophilous epiedaphic species which accidentally fall on
96the surface of freshwater (Arlé, 1971; Waltz & McCafferty, 1979; Christiansen & Snider,
972008; Palacios-Vargas, 2013). However, the semiaquatic ecology is a derived condition that
98evolved several times independently in springtails (D’Haese, 2002).

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 99 Although several species of Collembola can be hygrophilous and may be found both
100in land or directly associate to water habitats, hardly these aquatic associations are reported on
101taxonomical or ecological papers. Until now 525 water-dependent species of springtails were
102reported worldwide, of which 465 are freshwater dependent. To Neotropical Region
103proximally 45 water-dependent species were recorded so far, of which 28 are freshwater
104dependent (Deharveng et al., 2008). ONLY FOR MEXICO THERE ARE 26 RECORDS,
105SO I THINK THERE IS A MISSINOFORMATION.
106
107Systematics of Collembola Orders
108
109 Phylogenetic relationships among Collembola orders are much discussed and
110numerous hypotheses have been proposed so far. Poduromorpha and Neelipleona are possibly
111monophyletic, as observed by different analyses (D'Haese 2002, 2003; Gao et al., 2008;
112Xiong et al., 2008; Schneider et al. 2011; Schneider & D'Haese, 2013). In the other hand
113Entomobryomorpha and Symphypleona appear as either monophyletic or polyphyletic taxa in
114different approaches (D'Haese 2002; Xiong et al., 2008; Schneider & D'Haese, 2013).
115 Due to the body similarities (including elongated trunk with clear tergal segmentation)
116Entomobryomorpha and Poduromorpha were grouped in past as Arthropleona. Since this
117group were not supported by phylogenetic analyses and elongated segmented trunk is
118considered as a plesiomorphic character among springtails, Arthropleona fell into disuse
119(Xiong et al., 2008; Schneider et al., 2011; Schneider & D'Haese, 2013; Bellinger et al.,
1202016). D'Haese (2003) presents Poduromorpha as the sister group of all other orders, and
121Entomobryomorpha as the sister group of Symphypleona plus Neelipleona. In the same sense,
122the relationships between springtails with globular body (Symphypleona and Neelipleona) are
123also uncertain. Some believe that body condition is a derived form of Collembola, in which
124Symphypleona would be sister group of Neelipleona (D'Haese, 2003; Xiong et al., 2008).
125Other analyses indicate that Symphypleona is the sister group of Poduromorpha plus
126Entomobryomorpha (Gao et al., 2008;), or all other orders (Schneider & D'Haese, 2013).
127
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129Order Poduromorpha (sensu D'Haese, 2002)
130
131 Poduromorpha (Fig. 1a) specimens are characterized by cuticle strongly granulated
132and body with distinct tergal segmentation and protergite present with setae or vesicles. The
133order has about 3342 species in 333 genera, in which from Neotropics were recorded at least
13470 genera (Mari-Mutt, 1990). These species belong to five superfamilies and 11 families:
135Neanuroidea (Brachystomellidae and Neanuridae), Poduroidea (Poduridae), Hypogastruroidea
136(Hypogastruridae and Paleotullbergiidae), Gulgastruroidea (Gulgastruridae) and
137Onychiuroidea (Isotogastruridae, Pachytullbergiidae, Odontellidae, Onychiuridae and
138Tullbergiidae) (Bellinger et al., 2016).
139 Among the above cited families, Neanuridae is the most diverse, with 167 genera and
140about 1400 species described worldwide. They are characterized by reduced (rarely totally
141absent) mandibles without molar plate, modified maxillae with or without denticles and large
142fringed lamellae and mucro never with three lamellae. Brachystomellidae is now considered
143as the sister group of Neanuridae, since phylogenetic studies indicate independent mandible
144loss is Brachystomellidae, which combined with maxillae morphology are the main
145characteristics used to differentiate both families (Massoud, 1967; D’Haese 2002; Najt et al.,
1462005; Bellinger et al., 2016). Genera as Aethiopella, Friesea, Furculanurida, Neotropiella and
147Pseudachorutes of Neanuridae, and Brachystomella, Brachystomellides and Rapoportella of

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148Brachystomellidae may have aquatic or semiaquatic species in Neotropical Region (Waltz &
149McCafferty, 1979; Christiansen & Snider, 2008).
150 Poduridae presents only two species described worldwide. Podura aquatica has
151Holartic distribution and is typically found on water. Podura species are characterized by
152dentes more than 3 times as long as manubrium, with distal rings of granules and modified
153mucro (Fig. ).
154 Hypogastruridae is the second largest family of Poduromorpha with about 700 species
155in 40 genera, of which Ceratophysella, Hypogastrura, Schoettella and Xenylla are found in
156aquatic environments in Neotropical Region. Hypogastruridae is defined on
157symplesiomorphies as presence of molar plate on mandibles and absence of tergal pseudocelli
158(D’Haese, 2002; Deharveng, 2004). Paleotullbergiidae is a monotypic family from Africa
159without any aquatic association (Bellinger et al., 2016).
160 Odontellidae, Isotogastruridae and Pachytullbergiidae are considered families
161nowadays within Onychiuroidea, as well as Onychiuridae and Tullbergidae (D’Haese 2002,
1622003). Onychiuridae is the most diverse family of Onychiuroidea with approximately 650
163species in 56 genera, followed by Tullbergiidae with 226 species and 32 genera. Onychiuridae
164is apparently polyphyletic, due to divergence in ancestry between the subfamilies:
165Onychiurinae and Tetrodontophorinae. Isotogastruridae and Pachytullbergiidae are small taxa
166that together comprise 13 species in three genera. Pachytullbergiidae species are distinguished
167by abdomen with lateral projections, and Sensiphorura (Pachytullbergiidae) apparently is a
168semiaquatic genera. Others Onychiuroidea genera that can be semiaquatic are:
169Lophognathella, Onychiurus, Superodontella, Thalassaphorura and Tullbergia (Waltz &
170McCafferty, 1979; Christiansen & Snider, 2008).

171
172Order Entomobryomorpha (sensu Soto-Adames et al., 2008)
173
174 Entomobryomorpha (Fig. 1b) is the most diverse order of Collembola, with more than
1754,000 described species within 200 genera, which belong to 8 families and 4 superfamilies:
176Coenaletoidea, Entomobryoidea, Isotomoidea, Tomoceroidea (Soto-Adames et al., 2008).
177 Entomobryoidea (Entomobryidae, Microfalculidae and Paronellidae) is a
178monophyletic group, characterized by body setae mostly multilaterally ciliate, macrosetae
179sometimes cylindrical, usually truncate or broadened at the tip, postocular bothriotrichum
180present, and second and third abdominal segments with 2 and 3 bothriotricha respectively
181(Zhang et al., 2015). Microfalculidae is a monotypic family restricted from Magadascar.
182Entomobryidae is the most diverse family of Collembola, with 1,839 species in 64 genera,
183and as Paronellidae, it is widely distributed in the world and found in almost all terrestrial and
184freshwater environments. After Zhang et al. (2015), both families are paraphyletic, but they
185are distinguished from each other by crenulate dentes and claw-like mucro (falcate or
186bidentate) in Entomobryidae, while the Paronellidae present smooth dentes and square to
187elongate mucro with up to 10 teeth (Soto-Adames et al., 2008).
188 Isotomoidea (Actaletidae and Isotomidae), as well as Coenaletoidea (Coenaletidae),
189are characterized by body without scales and setae usually smooth or unilaterally ciliate,
190dorsal chaetotaxy more homogenous when compared to Entomobryoidea and, if present,
191macrosetae always acuminate. The phylogenetic relationships of these groups are still unclear,
192but Coenaletidae differs from Isotomoidea by presenting third abdominal segment fused with
193the fourth. Actaletidae and Coenaletidae are families with few species recorded from Mexico,
194Central America and the Caribbean Islands, which are present in marine water the first, and
195associated with hermit crabs of the genus Coenobita (Crustacea: Decapoda) the second.

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196Within Isotomoidea, Isotomidae differs from Actaletidae by fourth to sixth abdominal
197segments distinct; if fused, then shorter than the length of mesothorax to third abdominal
198segment and tenent hairs variously shaped but never leaf-like. Isotomidae is the second most
199diverse family of the order, with 112 genera and about 1,400 species, which are usually
200related to the soil environment (Bellinger, 1985; Soto-Adames et al., 2008; Bellinger et al.,
2012016). However, at least 30 genera of Isotomidae have been recorded in freshwater in North
202America until now (Waltz & McCafferty, 1979; Christiansen & Snider, 2008).
203 Tomoceroidea (Oncopoduridae and Tomoceridae) is possibly a monophyletic group,
204but with unclear relationships among other groups of Entomobryomorpha. They are similar to
205Entomobryidae by body setae mostly ciliate, but differ by lacking bothriotricha on head and
206reduced second and third abdominal segments. Tomoceridae species are distinguished by dens
207distally with spines, while in Oncopoduridae the dens lacks spines; or if present, spines are
208only seen on its basal half (Soto-Adames et al., 2008; Bellinger et al., 2016). Genera found in
209freshwater of these families are Tomocerus from North America and Oncopodura from
210Brazilian Amazon (Arlé, 1960; Waltz & McCafferty, 1979).
211
212Order Symphypleona (sensu Bretfeld, 1994, 1999)
213
214 Symphypleona (Fig. 1c) springtails present antennae longer than head and globular
215body, formed by fusion of most thoracic and abdominal segments. They are widely distributed
216and quite diverse with about 1,300 species in 115 genera, 10 families and 5 superfamilies:
217Dicyrtomoidea (Dicyrtomidae), Katiannoidea (Arrhopalitidae, Collophoridae, Katiannidae
218and Spinothecidae), Sminthuridoidea (Mackenziellidae and Sminthurididae), Sturmioidea
219(Sturmiidae) and Sminthuroidea (Bourletiellidae and Sminthuridae) (Bellinger et al., 2016).
220 Sminthuridoidea differs from other superfamilies by prehensile modifications in
221second and third antennal segments of males. Sminthurididae differ from Mackenziellidae by
222presence of mandibles and hypognathous head. In Neotropical Region, species of
223Sminthurididae can be found associated to water puddles such as Denisiella, Sminthurides
224and Sphaeridia (Waltz & McCafferty, 1979; Bretfeld & Gauer, 1994; Fjellberg, 2007;
225Christiansen & Snider, 2008).
226 The remaining superfamilies of Symphypleona form a monophyletic clade, the
227Appendiciphora, characterized by the presence of a pair of modified setae (subanal appendix)
228on inferior lobes of the small abdomen of females. This subanal appendix may be directed to
229the anus (up) in Dicyrtomoidea and Sminthuroidea, posteriolly in Sturmioidea, or to the
230genital opening (down) in Katiannoidea (Bretfeld, 1999).
231 Sminthuridae is the most diverse family of Symphypleona, representing 58% of the
232genera and 38% of the species within the group. Sminthuridae differs from Bourletiellidae by
233fifth abdominal segment with a pair of bothriotricha on each side and pretarsus with a pair of
234setae. Dicyrtomidae resembles Sminthuridae for subanal appendix directed upward, but it is
235easily distinguished by fourth antennal segment very small compared to the third.
236 In Katiannoidea, Katiannidae is the most diverse family with about 210 species in 19
237genera, of which at least six of them are recorded from Neotropical Region. Katiannidae
238differs from other families of the group by presence of 6-8 eyes and tibiotarsus with spatulate
239setae. Collophoridae and Arrhopalitidae are characterized by reduction of eyes (0-4). They
240differ mainly by the presence of a capitate bothriotricha on each side of the small abdomen
241small in Collophoridae (Bretfeld, 1999; Bellinger et al., 2016).
242 The Appendiciphora genera considered aquatic or semiaquatic recorded in freshwater
243in North America are: Arrhopalites (Arrhopalitidae), Collophora (Collophoridae),
244Sminthurinus (Katiannidae), Bourletiella, Deuterosminthurus, Heterosminthurus,

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245Pseudobourletiella (Bourletiellidae), Neosminthurus, Sminthurus and Sphyrotheca
246(Sminthuridae) (Waltz & McCafferty, 1979; Christiansen & Snider, 2008).
247
248Order Neelipleona, Family Neelidae
249
250 Neelidae (Fig. 1d) is the only family in Neelipleona, and similarly to Symphypleona,
251specimens are characterized by globular body pattern. They differ from Symphypleona due to
252very minute body size, antennae shorter than head, eyes always absent, large abdomen with
253sensory field and small abdomen ventrally hidden (Schneider & D'Haese, 2013).
254 Currently Neelidae comprises 44 species in 5 genera, of which Megalothorax,
255Neelides, Neelus and Spinaethorax were recorded in Neotropical Region. The species are
256usually associated with euedaphic environments and caves, but species such as Megalothorax
257aquaticus Stach are found in freshwater in Europe, as well as Neelus spp. in North America
258(Christiansen & Snider, 2008; Bellinger et al., 2016).
259
260 LIMITATIONS
261
262 Since springtails are small but remarkable complex organisms, identification of
263specimens can be difficult and often may lead to dubious or erroneous taxonomic identities.
264Some other factors related to limitations to correctly identifying springtails are listed below.
265 Limited descriptions: Collembola species are mostly compared and described
266nowadays by, among other features, chaetotaxy (distribution, morphology, size and presence
267or absence of some head, trunk or appendicular setae). Due to complexity of such evaluation,
268past descriptions not always were based in such characterization (or presented summarized
269versions of it). The lack of details in descriptions sometimes can hide several species under
270one solely species name or difficult comparisons among species. In Brazil, a clear example is
271seen to Entomobrya species, which in most cases were described based in color patterns, a
272character currently seen as highly plastic and of second importance when compared to
273chaetotaxy (Heckman, 2001; Katz et al., 2015). A similar problem is seen to Seira prodiga
274Arlé, a name which possibly holds a complex of quite different species concerning other
275morphological characters than color pattern and body shape. So, very detailed descriptions
276can potentially prevent future synonyms and even help to identify cryptic species (Arlé, 1959;
277Katz et al., 2015; Tully & Potapov, 2015; Soto-Adames, 2016).
278 Descriptions of immature forms or unclear data about intraspecific morphological
279variation: immature springtails are different from adults in size, coloration, morphometry,
280absence of genital pore and chaetotaxy. Due this, immature forms and adults analyzed by the
281scrutiny of currently taxonomy can look like different species in some cases. In the same
282sense springtails species usually present some range of polymorphic characters, as expected to
283most animal populations. For example, it was recorded to different Entomobryoidea species
284variations in color pattern, some dorsal body setae, labial chaetotaxy and number of teeth on
285unguis within specimens of the same population (Cipola et al., 2016; Soto-Adames, 2016).
286Some specimens can even present punctual asymmetries of chaetotaxic elements when left
287and right sides are compared. Because of this, descriptions of springtails should be based in a
288large number of type specimens, which should be carefully revised to detect possible
289polymorphic characters.
290 Terminology confusion: a minor problem concerning taxonomy in Collembola is the
291use of different terminologies to same structures. Appendages like ventral tube, tenaculum
292and furca can also be called collophore, retinaculum and furcula respectively. A synonym to
293“setae” is “chaetae” and “bothriotrichum” can also be “trichobothrium”. The labial triangle
294region is also called ventral basomedian field. To body setae this is more problematic.

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295Different nomenclature systems refer to the same setae with different names (or labels), which
296may difficult identification or comparison of different species. The language often present
297important differences naming the same structure, for instance tenent hair (English), ergot
298(French) spürhaar (German) and sometimes even in the same language it can vary as in
299Spanish pelo tibiotarsal and espolón mazudo for the same structure.
300 Loss of type material and missing collection data: studying springtails demands
301mounting specimens under glass slides, which include the use of different techniques and
302chemicals to clear the specimens and addition of a fluid medium to conserve them after
303preparation (see material preparation and preservation). The misuse of corrosive chemicals
304during clarification, improper choice of conservation medium or storage of slides under
305inadequate conditions can lead to material degradation. Another problem is the absence of
306important collecting data to the material, especially regarding the type locality. An example of
307this is Dicranocentrus silvestrii Absolon, which its type locality was presented by the author
308as “South America” and for this the species has already been suggested as incertae sedis
309(Mari Mutt, 1979). Mastigoceras camponoti Handschin, a monotypic endemic taxon from
310Brazil, was presented by the author as collected from “South of Minas Gerais state”, which is
311one of the largest federal units of Brazil (Heckman, 2001). Another example is provided by
312Börner (1906) who described three species from “São Franscisco, Brazil”, which could be
313several locations in different regions of the country.
314 Regional reality: concerning Neotropical Region, two main conditions also obscure
315the real identities of springtails. Firstly, lack of samplings or taxonomical data from different
316ecosystems or even large phytogeographic domains in Central and South America. Most of
317the species recorded from the New World were from United States, Mexico, Puerto Rico and
318Brazil, with few or no species to other countries (Mari-Mutt & Bellinger, 1990). At the same
319time, large countries with several records like Brazil are still under sampled to many
320ecosystems and/or federative units (Zeppelini et al., 2016). Historically this may be related to
321a restricted number of taxonomists working with the Neotropical fauna, a number which is
322now rising at least in Brazil and Mexico. Secondly Neotropical Region potentially holds a
323large biodiversity of land arthropods, including Collembola. So, it is expected that potential
324new species can be commonly found in unexplored habitats. Such lack of knowledge
325combined to a potential undescribed fauna limits the overall comprehension of distribution
326and composition of Collembola fauna in Neotropical Region. Regarding the aquatic fauna,
327only 45 species in 15 genera were recorded so far, which we believe does not reflect the real
328species richness of aquatic springtails to the region since these numbers were gathered from
329isolated data, not from directed collections (Deharveng et al., 2008). (CHECK MY
330COMMENTS ON PAGE 3)
331 All these presented factors also lead to the lack of identification keys and specific
332diagnoses for freshwater springtails from neotropics. The only specific key to freshwater
333springtails of the New World was proposed by Waltz & McCafferty (1979) including 11
334families and 25 genera from North America taxa and therefore does not reflect the diversity of
335the Neotropical springtails.
336
337 TERMINOLOGY AND MORPHOLOGY
338
339 The general morphology of Collembola presented below is based on the following
340references: Hopkin (1997), Christiansen & Bellinger (1998), Fjellberg (1999, 2007),
341Heckman (2001), Deharveng (2004), Zeppelini & Bellini (2004), Bellinger et al. (2016).
342Specific references for each order follows: Massoud (1967), Jordana et al. (1997), Thibaud et
343al. (2004) and Najt et al. (2005) for Poduromorpha; Mendonça & Arlé (1987), Potapov
344(2001), Liu et al. (2008), Soto-Adames et al. (2008, 2014), Bellini & Zeppelini (2011) and

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345Soto-Adames (2016) for Entomobryomorpha; Richards (1968), Arlé (1971), Betsch (1980),
346and Bretfeld (1999) for Neelipleona and Symphypleona.
347 Springtails, as well as other hexapods, have the body divided in three tagmata: head,
348thorax and abdomen. The body is entirely covered by a thin granulous cuticle, which is
349variable hydrophobic. The granules of epicuticle are typically hexagonal patches formed by
350the connection of primary triangular microgranules. This pattern of cuticular organization is
351quite evident in scanning electron microscope photographs of Poduromorpha species. Other
352taxa may present reduction or fusion of such granules giving the epicuticle another
353microprofile. To some species of Symphypleona, the cuticular granules can also be square
354(formed by four linked granules), not hexagonal. The cuticle has numerous different setae
355(usually smooth or ciliated), spines, sensilla and/or scales, which combined are mostly
356diagnostic to different taxonomical groups. An example of this is the presence of scales only
357in some Entomobryomorpha.
358 Head is dorsally rounded, elliptical or subtriangular and bears a pair of antennae,
359eyepatches and postantennal organs (which can be totally absent) and mouthparts. The mouth
360projection position makes springtails prognathous (most taxa of Poduromorpha and
361Entomobryomorpha) or hypognathous (Neelipleona and most taxa of Symphypleona).
362 Antennae are inserted anterodorsally on head and present intrinsic muscles that allows
363independent movement of all 4 antennal articles. Antennae size is variable within different
364taxa and springtails can present antennae smaller than head (e.g. Neelidae) to antennae much
365longer than the whole body (e.g. Campylothorax, Salina and Temeritas). The first two
366antennal articles are usually simple and undivided, but in some cases the first (e.g.
367Mastigoceras) and the second (e.g. Dicranocentrus) segments may be subdivided, causing an
368impression that antennae have five or six segments. The third antennal segment is usually
369undivided and with an apical sense organ with peculiar setae or sensilla associate. In males of
370Sminthurididae (e.g. Sminthurides) and in Coenaletidae (Coenaletes) there are projections and
371spines in the second and third antennal segments which, along with the morphology of the
372segments themselves, provide a prehensile capacity to the antennae to hold females during the
373courtship. The fourth antennal segment can be undivided (more common), annulated (e.g.
374Mastigoceras) or pseudosegmented (e.g. Sminthurides, Temeritas). This segment can hold an
375apical bulb and numerous sensory modifications (sensilla) of taxonomic significance (e.g.
376Neanuridae, Brachystomellidae). To most Collembola groups this segment is the largest
377antennomere, but there are exceptions like in Dicyrtomidae and some Tomoceridae, in which
378it is smaller than the third segment.
379 The eyepatches (if present) are dorsal and bear a small set of eyes (ommatidia) with up
380to 8 lenses on each side. Eyepatches are usually pigmented (black, brown or dark blue).
381Reduction in number of ommatidia is common and basically all families present forms with
382less than 8+8 eyes. For example, the Collophoridae has 4+4 eyes, Arrhopalitidae 2+2 or less,
383and Neelidae and Cyphoderinae (Paronellidae) are completely blind.
384 Between the eyes and antennae in some families a pair of postantenal organs can be
385present (e.g. Brachystomellidae, Neanuridae, Isotomidae and rarely in Entomobryidae),
386whose structure is variable and of taxonomic importance. They can be totally absent (e.g.
387Symphypleona), simple and oval (e.g. part of Isotomidae) to quite complex, formed by
388several circular vesicles (moruliform, e.g. part of Anurida). The postantennal organ is a
389hydrophilic structure and it is possibly a chemo, thermo or hygroreceptor.
390 The dorsal region of mouth is covered by: a variably developed clypeus usually
391carrying some setae and which ends in a prelabrum region (also often with four setae); and an
392anterior labrum usually with a set of 14 setae.
393 Ventrally the head presents the labial region with associate papillae and setae, a pair of
394maxillary palps (more laterally), and a post labial region divided in left and right sides by the

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395cephalic groove. The labial papillae consist of a conjunct of anterior projections ventral to the
396mouth opening surrounded by some accessory setae. Most springtails present five main
397papillae but reduction or absence of them was recorded in some Brachystomellidae,
398Onychiuridae and Tullbergiidae. The set of papillae are often seen near a set of four to seven
399proximal setae. The maxillary palps (or outer lobes) are basically conical projections seen
400laterally of labial papillae. Each palp usually has an apical and basal seta with rare exceptions
401of one or two additional setae in Isotomidae (e.g. Desoria). The sublobal plate may have up to
402four setae-like projections which can be totally absent. Finally, labial and post-labial regions
403present several typical setae of taxonomic importance.
404 Inside the cephalic capsule are the mouthparts, mostly formed by a pair of strong
405chewing mandibles and a pair of lateral maxillae. The mandibles consist apically by an inner
406row of incisors teeth (4 to 6) followed by a basal molar plate with numerous conical denticles.
407Mandibles are completely absent in most of Brachystomellidae and at least lacking molar
408plates in Neanuridae. The maxillae are usually formed by a variable number of apical teeth
409plus some fringed associated lamellae. Maxillae teeth and lamellae could also be reduced or
410modified, as seen in styliform maxillae of Pseudachoritinae (Neanuridae). The mandibles and
411maxillae can be elongated in Rhynchocyrtus, Stenognathriopes and Stenognathellus.
412 Trunk is formed by a thorax with ambulacral legs and a legless abdomen. It can be
413cylindrical or elongated and present clearly tergal segments (Entomobryomorpha and
414Poduromorpha), or such segments can be mostly fused to each other generating a rounded or
415globose body (Symphypleona and Neelipleona).
416 Thorax has three segments, prothorax, mesothorax and metathorax, each with a pair of
417legs. Prothorax is the shortest segment and can present or lack its tergal cover (pronotum).
418The presence or absence of pronotum in clearly segmented springtails defines two groups:
419Poduromorpha (with pronotum) and Entomobryomorpha (lacking the pronotum). Meso and
420metathorax are simple and similar to each other, although in a few cases can be strongly
421projected in its midline (e.g. Lepidocyrtoides, Lepidocyrtinus and Campylothorax).
422 Each leg is formed by coxa, trochanter, femur, tibiotarsus, pretarsus and one claw
423(with or without an empodial appendix - unguiculus). Coxae are usually subdivided into
424epicoxa (or subcoxa 1), subcoxa (or subcoxa 2) and coxa and present only few setae. In some
425Symphypleona such as Sminthuridae, the third coxae present a robust spine-like seta.
426Trochanters are small segments of legs like the coxae and may contain specialized features as
427well. The third pair of trochanters of Entomobryoidea presents a very distinct set of small
428smooth spine-like setae named trochanteral organ. Symphypleona can also present
429trochanteral organs which may not be homologous to the Entomobryoidea, with a single spine
430as seen in some Katiannidae and Sminthuridae.
431 Femur (plural, femora) is an undivided segment covered by different types of setae
432and/or scales. This segment usually has minor taxonomic value but a remarkably exception is
433Tyranosseira, in which males have broadened femora with distinct spines on the first pair of
434legs.
435 Tibiotarsus (plural, tibiotarsi) is supposedly formed by the fusion of tibia and part of
436tarsomeres seen in other hexapods. This segment usually is long and covered by numerous
437setae, which are almost all ciliate in Entomobryoidea and smooth in Poduromorpha,
438Isotomidae, Symphypleona and Neelipleona. Tibiotarsi can present very different setae, scales
439and/or spines in distinct taxa. The anterodistal region of tibiotarsican present one (e.g.
440Entomobryidae) or more (e.g. Katiannidae and Bourletiellidae) modified setae called tenent
441hairs, which can be pointed, apically capitate or even clavate. In Symphypleona the tibiotarsi
442may also present distinct organs such as oval organs, which aid to diagnose some genera and
443species.

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444 Pretarsus is a small membranous structure in which are attached an unguiculus and
445one unguis. They all together form the empodial complex of springtails. Pretarsus usually
446present two very small smooth setae, one on anterior and another on posterior face, but the
447latter is absent in Bourletiellidae and both are apparently absent in Campylothorax. Unguis
448(plural, ungues) is the single tibiotarsal claw of springtails. It presents an external face formed
449by a convex transverse plate tapering distally, which engage on its inner side (concave face) a
450central lamella which may or not contain basal paired teeth and/or unpaired distal teeth. In
451certain isotomids and sminthurids the external face of unguis may have one membranous and
452somehow inflated membrane called tunica, as well as sometimes dorsal and lateral teeth on a
453pair of lamellae, the pseudonychia. Unguiculus (plural, unguiculi) is the empodial appendix of
454the pretarsus and may look like a secondary smaller claw. It presents an outer plate and an
455inner lamella. The external side is apically acuminate and can hold an apical or a subapical
456filament on some Poduromorpha and Symphypleona. Unguiculus inner lamella can be
457acuminate, truncate or even concave. In several families of Poduromorpha, as Neanuridae,
458unguiculi can be are missing.
459 Abdomen in springtails has six segments and in Arthropleona (Poduromorpha +
460Entomobryomorpha) usually they are individualized, but in some Isotomidae genera two or
461three terminal segments can be completely fused to each other (e.g. Folsomina, Folsomia,
462Cryptopygus, Isotomiella, Hemisotoma). In Poduromorpha, Isotomidae and some
463Entomobryidae (e.g. Dicranocentrus, Mastigoceras) abdominal segments III and IV have
464similar length in the midline, while in most Entomobryidae and all Paronellidae the fourth
465abdominal segment is at least twice the length of the third segment. To Symphypleona large
466and small abdomen may include different thoracic and abdominal segments. In most families
467the large abdomen is formed by the last two thoracic segments and the first four abdominal
468segments, and the small abdomen presents the last two abdominal segments (as in
469Arrhopalitidae, Bourletiellidae, Katiannidae and part of Sminthuridae). In the other hand the
470large abdomen in some Sminthuridae includes also the first segment of the thorax. In
471Dicyrtomidae the large abdomen holds the fifth abdominal segment. Most remarkably is
472Collophoridae and Neosminthurus (Sminthuridae) in which the large abdomen includes
473almost all trunk segments, excluding only the sixth abdominal segment.
474 In the ventral abdomen is the collophore, tenaculum, furcula, genital plate and anus
475respectively in the first, third, fourth, fifth and sixth abdominal segment.
476The collophore (or ventral tube) is inserted in a basal plate, followed by a cylindrical part
477(except in Sturmiidae in which is biparted), where ends the ventral groove. The anterior,
478latero-distal and posterior faces of collophore can bear setae of taxonomical importance.
479Distally the collophore presents two eversible vesicles (or sacs), which can be short
480(Poduromorpha and Entomobryomorpha), or long (Symphypleona). Tenaculum (or
481retinaculum) is formed by a basal corpus, with or without setae, and a pair of lateral rami with
482(or without) a basal appendage and 2-4 distal teeth. In Symphypleona the corpus can be a
483posterior lobe (e.g. Sminthurididae) and an apical lobe with 1-4 setae. The tenaculum has the
484function of retaining the furcula (or furca) attached to the body before a leap. When the
485furcula is absent or vestigial, the tenaculum is often absent (e.g. Friesea, Americanura,
486Stachia and Stachiomella).
487 The furcula is the jumping appendage of springtails and can be long and well-
488developed (e.g. Entomobryoidea) to short, reduced or even absent in Onychiuridae,
489Tullbergiidae and part of Brachystomellidae, Hypogastruridae and Neanuridae. The furcula
490consists in: a proximal manubrium, a pair of dentes (singular dens) and a pair of distal
491mucros. Manubrium is a single distally bifid segment usually with setae associated. Some
492taxa can also present manubrial scales (e.g. part of Entomobryoidea), spines (Isotoma), or can
493even be devoid of any attachments (Archisotoma, Ballistura, Folsomides). Dentes are paired

19 10
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494cylindrical segments, generally longer than the manubrium and covered by different types of
495setae, spines and/or scales. Rarely the dens can present modifications such as appendages in
496anterior (e.g. neotropical Lepidocyrtus) and distal region (e.g. Salina). In Entomobryidae,
497some genera of Isotomidae and rarely in Arlesminthurus (Bourletiellidae) the dorsal surface of
498dens can be typically crenulated. Another modification is the annulated form of dens in
499Poduridae. Moreover, some Entomobryoidea, Isotomidae and Tomoceridae may have dorsal
500spines on dorsal dens or large scales-like setae. At the tip of each dens of furcula there is one
501mucro, with different shapes among families and genera. To most Poduromorpha mucro can
502be spoon-like, relatively large when compared to the dens length, with smooth edges and with
503or without a small lateral tooth. In Isotomidae mucro can bear 1 to 4 teeth and usually are
504very short. To Entomobryidae mucro is also short, with 1 or 2 teeth, with or without a single
505mucronal seta. In Paronellidae mucro is quite variable, and can be long with up to 10 teeth (as
506in Cyphoderinae), or short with up to 6 teeth (in Paronellinae) and in both cases lack any
507mucronal seta. In Symphypleona the mucro is large and flattened dorsoventrally forming
508concave spatula with smooth edges (Bourletiellidae), or a spoon-like structure with at least
509one serrated edge (most Sminthuridae). The mucro can or cannot present a small discrete
510basolateral seta.
511 The genital plate is ventral in fifth abdominal segment where the genital opening
512develops in adults. Females present a transverse opening with at least some anterior setae and
513males present a circular projection where a small longitudinal opening appears, usually
514surrounded by setae. In Entomobryomorpha male genital plate can be oligochaetotic (with
515few setae), multisetaceous (with many setae), circinate (almost smooth without setae) and
516papillate (with papillae).
517 Finally, the sixth abdominal segment is divided into two inferior and one superior
518lobes that together surround the triradiate anus in ventro-terminal region. The three anal lobes
519(or valves) bear setae of taxonomic value at least to Poduromorpha and Symphypleona. In this
520last taxon some setae are also sexually dimorphic, as the subanal appendix seen only in
521females of all families, except Sminthurididae. The dorsal anal lobe may present a pair (or
522more) of spines in some Hypogastruridae, Isotomidae, Neanuridae, Odontellidae,
523Onychiuridae and Tullbergiidae.
524
525
526Collembola terms are defined below.
527
528Bothriotrichum (or trichobothrium) – Differentiate long thin setae, smooth or ciliate. In plural:
529bothriotricha.
530Antennal bulb – Apical lobe or vesicle of fourth antennal segment.
531Antennal sense organ – Two latero-distal rods of third antennal segment with one guard
532sensillum on each side.
533Collophore (or ventral tube) – Ventral appendix on the first abdominal segment, formed by
534basal cylindrical tube with two distal eversible vesicles.
535Circumanal seta – Definition?
536Dens (plural, dentes) – Paired segment of furcula appendage, located between manubrium and
537mucro.
538Dental spine – Short smooth or ciliated spine-like setae arranged in rows or grouped on the
539dens.
540Empodial complex – Distal region of tibiotarsus formed by pretarsus, unguis and unguiculi.
541Furcula (or furca) – Jumping organ inserted ventrally in the fourth abdominal segment,
542composed by manubrium, a pair of dentes and a pair of distal mucros.

21 11
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543Large abdomen – Globular region of trunk composed by fusion of thoracic and abdominal
544segments of Symphypleona and Neelipleona.
545Macroseta (or macrochaeta) – Robust and long setae present on head, trunk and appendages.
546Microseta (or microchaeta) – Small setae present on head, trunk and appendages.
547Mandibles – Main chewing mouthparts, usually with apical teeth and basal molar plate.
548Manubrium – Unpaired basal appendage of furcula.
549Maxillae – Accessory chewing mouthparts, usually consisting of a capitulum (apical region
550usually with teeth and fringed lamellae) connected with 3 articulating arms: stipes, cardo and
551fulcrum.
552Mesothorax – Second segment of the thorax.
553Metathorax – Third segment of the thorax.
554Mucro – Furcula distal segment, with or without teeth and setae; sometimes claw-like.
555Neosminthuroid setae – Differentiate ciliate setae on posteroinferior region of large abdomen
556in some Symphypleona.
557Oval organ – Small, oval or elliptical integumentary depression with rims present in different
558segments of legs in Symphypleona.
559Pretarsus – Small membrane in the distal tibiotarsus connected to unguis and (if present)
560unguiculus.
561Postantennal organ (PAO) – Dorsal head paired sensory organ, seen (if present) between the
562anterior eyes and the antennal base.
563Prelabral setae – Transverse row of four setae inserted on anterior region of the head, between
564the labrum and the frontoclypeal field.
565Prothorax – First segment of the thorax. Can present a pronotum (or protergite) or not.
566Sensillum (plural, sensilla) – Small setiform projection generally with cylindrical apex and
567smooth surface
568Seta (or chaeta) – Structure slender, hair-like, usually a sensory extension of the cuticle,
569connected to the body wall by a socket.
570Sensory field – Definition?
571Small abdomen – Formed by terminal abdominal segments in Symphypleona and
572Neelipleona.
573Subanal appendage – Pair of modified setae present on inner side anal inferior lobes in part of
574Symphypleona females.
575Tenaculum (or retinaculum) – Ventral appendage of the third abdominal segment that holds
576the furcula at rest.
577Tenent hair – Dorsal or ventral differentiate setae of tibiotarsi apex. It can be pointed,
578knobbed or clavate.
579Tibiotarsus (plural, tibiotarsi) – Last segment of the legs, formed possibly by fusion of tibia
580and the first tarsomeres seen in other hexapods.
581Trochanteral organ – Conjunct of spines-like smooth setae seen on trochanters of third legs.
582(GOOD FOR ENTOMOBRYIDAE BUT NOT SYMPHYLEONA)
583Unguis (plural, ungues) – Leg distal claw inserted in the pretarsus.
584Unguiculus (plural, unguiculi) – A pretarsus appendix, similar to a smaller claw.
585
586
587 MATERIAL PREPARATION AND PRESERVATION
588
589 After collecting it is very important to properly preserve the specimens. It is preferable
590to keep the springtails in 70% ethanol or higher concentrations. To freshwater springtails we
591recommend the use of 95% ethanol. This will prevent loss of quality of the material and leave
592it in good conditions for preparation and identification.

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593 For observation of the basic morphology and identification, most taxa need to be
594cleared and mounted under glass slides for microscopy (with magnification of 400x or more).
595It is desirable a phase-contrast equipment attached to a drawing tube.
596 Specimens need to be cleared especially when they have strong pigmentation. The
597main chemical reagents used to clarify are: 1) basic solution of potassium hydroxide 5-10%
598(KOH), which converts the pigmentation of Collembola for a red color, and must be used for
599a short time (5 to 10 min) which serves to dissolve the cuticular oil; 2) lactic acid (C3H6O3) or
600lactofenol (C3H6O3 + C6H6O) which serves to macerate the internal specimen tissues, and in
601which the specimen can be left for a long period of time (weeks), or be heated for some
602seconds; 3) Nesbitt’s solution (C2H3Cl3O2 + H2O + HCl) is also useful for rapid clarification
603when the medium is heated on a hot plate (55°, 10-20 min.), and also suitable for specimens
604that are preserved in ethanol for long time. These methods can be combined, modified and
605used for different periods depending on the specimen and the results to be obtained. For dark
606specimens it is appropriate to wash the specimens firstly in potassium hydroxide solution,
607remove the excess of KOH and then wash them in lactic acid or lactophenol, and drop the
608excess of chemical before mounting. Jordana et al. (1997) suggest the modification of
609Nesbitt’s fluid, adding chloral-lactophenol to the solution (C2H3Cl3O2 + C3H6O3 + C6H6O).
610 After clarification, a temporary mounting slide can be used when the specimen need to
611be analyzed only for a short time. A temporary slide can be made on cavity glass slides with
612lactic acid, lactophenol or glycerine. The advantage of this method is the specimen may be
613recovered by diluting medium, and stored back in alcohol. It is also an appropriate method
614when there are several specimens of the same taxon and there is no problem in damage and/or
615lose some individuals. In some cases, when the specimen is very large, it is appropriate to use
616an excavated slide to dissect it but not to mount.
617 The semi-permanent mounting is the most common method to preserve specimens for
618identification. The mounted springtails can be preserved in glass slide during long periods
619(over 50 years) without the need of remounting. The disadvantage of this method is that each
620specimen will remain in a fixed position and possibly some structures may be obscured. The
621most commonly medium used for semi-permanent slides is Hoyer’s solution (H2O distilled +
622fragments of arabic gum + glycerin + chloral hydrate). As a semi-permanent medium, Hoyer’s
623solution is soluble in water, so it is possible to dissolve the medium and carefully remount
624specimens, if necessary.
625 In institutional collections, as well as other small arthropods, springtails can be kept in
626liquid vials, preserved in 92% ethanol or higher concentrations, or in semi-permanent slides.
627In liquid, they are usually kept in small vial with ethanol (4 cm length x 0.5 in diameter)
628which can be closed with cotton and placed vertically in a large pot also containing ethanol.
629Permanent mounting can be stored in glass slides boxes in dry and fungi proof environments.
630 More information on preservation, clarification and mounting techniques are presented
631in Jordana et al. (1997), Christiansen & Bellinger (1998), Palacios-Vargas & Mejía-Recamier
632(2007).
633
634
635 KEY TO FRESHWATER COLLEMBOLA
636
637Collembola: Orders
638
6391 Elongated body, with distinct tergal segmentation (Fig. 1a, b) …………………………… 2
6401’ Globular body, formed by thoracic fusion and abdominal segments (Fig. 1c, d) ………… 3
641
6422(1) Prothorax distinct, with setae or papillae (Fig. 1a) …………………… Poduromorpha

25 13
26
6432’ Prothorax reduced, without setae or papillae (Fig. 1b) ……………. Entomobryomorpha
644
6453(1’) Eyes usually present; antennae longer than head; small abdomen distinct; specimens of
646coloration and sizes variable (Fig. 1c) …………………………………….… Symphypleona
6473’ Eyes absent; antennae shorter than head; small abdomen indistinct, hardly seen; very
648minute specimens; totally depigmented (Fig. 1d) ……………………………… Neelipleona
649
650
651Collembola: Poduromorpha Families
652
6531 Mandibles absent or rudimentary, if present lacking molar plates ………………………… 2
6541’ Mandibles present and developed, with molar plates (Fig. 2a) …………………………… 4
655
6562(1) Maxilla cardo present (Fig. 2b) ………………………………………………………… 3
6572’ Maxilla cardo absent (Fig. 2c) ……………………………………………… Odontellidae
658
6593(2) Maxilla capitulum with apical, well developed teeth (Fig. 2b); mandibles usually absent
660……………………………………………………………………………… Brachystomellidae
6613’ Maxilla capitulum otherwise, maxilar teeth usually reduced (Fig. 2d-g); mandibles usually
662present …………………………………………………………………………… Neanuridae
663
6644(1’) Dens, if present, short and straight, without distal rings (Fig. 3a) …………………… 5
6654’ Dens long and curved, distally crenulate (Fig. 3b) [MEX] …………………………………
666…………………………………… Poduridae, one species: Podura aquatica Linnaeus, 1758
667
6685(4) Head and trunk dorsally with pseudocelli (Fig. 3c) …………………………………… 6
6695’ Head and trunk dorsally without pseudocelli …………………………… Hypogastruridae
670
6716(5) Third antennal segment sense organ with 2 guard parallel sensilla; sense rods and guard
672sensilla hidden by well developed conical tegument papillae (Fig. 3d) …………………
673Onychiuridae
6746’ Third antennal segment sense organ with 1-3 guard sensilla; when more than one, they are
675curved to each other; sense rods partially hidden by weakly developed tegument papillae (Fig.
6763e) ……………………………………………………………………………… Tullbergiidae
677
678
679Collembola: Poduromorpha: Brachystomellidae Genera
680
6811 Maxilla globular with rostral apical teeth (Fig. 2b)………………………………… 2
6821’ Maxilla elongated with lateral teeth (Fig. 4a) [NEO] …… Rapoportella Ellis & Bellinger
683
6842 Fourth antennal segment ventral side with several modified short setae (Fig. 4b) [NEO] …
685………………………………………………………………………… Brachystomellides Arlé
6862’ Fourth antennal segment ventral side without modified short setae [NEO] …………………
687………………………………………………………………………… Brachystomella Ågren
688
689
690Collembola: Poduromorpha: Hypogastruridae Genera
691
6921 Sixth abdominal segment with 2 or 0 anal spines (Fig. 5a) ……………………………… 2

27 14
28
6931’ Sixth abdominal segment with 3 anal spines (Fig. 5b) [SA] ……… Triacanthella Schäffer
694
6952(1) Postantennal organ absent ……………………………………………………………… 3
6962’ Postantennal organ present ……………………………………………………………… 5
697
6983(2) Specimens pigmented, usually dark blue; 5+5 (rarely 4+4) eyes; furcula and tenaculum
699usually present ……………………………………………………………………………… 4
7003’ Specimens white, lacking pigments; eyes, furcula and tenaculum absent [NEO] …………
701………………………………………………………………………… Acherontiella Absolon
702
7034(3) Third antennal segment sense organ with thick far apart sense rods (Fig. 5c) [NEO] …….
704…………………………………………………………………………… Paraxenylla Murphy
7054’ Third antennal segment sense organ with small near to each other sense rods (Fig. 5d)
706[NEO] ……………………………………………………………………...… Xenylla Tullberg
707
7085(2’) Eyes and furcula absent ………………………………………………………………… 6
7095’ Eyes and furcula present ……………………...…………………………………………… 7
710
7116(5) Unguiculus present [NEO] ……………………………………………… Willemia Börner
7126’ Unguiculus absent [NA] …………………………………………………… Tafallia Bonet
713
7147(5’) Unguiculus present, with basal lamella enlarged (Fig. 5e) …………………………… 8
7157’ Unguiculus absent, or if present, setiform [NEO] …………………… Schoettella Schäffer
716
7178(7) Mandibles normal, with apical teeth and strong molar plate (Fig. 2a) ………………… 9
7188’ Mandibles reduced, without apical teeth and with molar plate rudimentary (Fig. 5f) [NA]
719… ……………………………………………………………………… Microgastrura Stach
720
7219(8) Antennal segments IV and III ventral junction with eversible sac, mucro spatulate (Fig.
7225g) [NEO] ……… …………………………………………………………………………
723Ceratophysella Börner
7249’ Antennal segments IV and III ventral junction without eversible sac, mucro triangular
725[NEO] ……………… …………………………………………………………………………
726Hypogastrura Bourlet
727
728
729Collembola: Poduromorpha: Neanuridae Genera
730
7311 Sixth abdominal segment dorsally bilobed (Fig. 6a); postantennal organ absent ………… 2
7321’ Sixth abdominal segment dorsally simple, rounded (Fig. 6b); postantennal organ present or
733absent ………………………………………………………………………………………… 5
734
7352(1) Eyes 2+2 to 0+0; specimens not blue, yellowish, redish or lacking pigments ………… 3
7362’ Eyes 3+3; specimens usually blue [NEO] ………………………… Neanura MacGillivray
737
7383(2) Fourth antennal segment sensilla subequal in thickness ……………………………… 4
7393’ Fourth antennal segment with one sensillum clearly thicker than others (Fig. 6c) [NA] …
740……………………………………………………………………… Sensillanura Deharveng
741
7424(3) Head dorsal pair of setae d1 absent (Fig. 6d) [NEO] ……………… Ectonura Cassagnau

29 15
30
7434’ Head dorsal pair of setae d1 present (Fig. 6e) [NEO] ……………… Paleonura Cassagnau
744
7455(1’) Maxilla capitulum elongated, with or without teeth (Figs. 2e-g) ……………………… 6
7465’ Maxilla capitulum short and triangular, with two small lamellae toothed (Fig. 2d) [NEO]
747………………………………………………………...…………… Friesea von Dalla Torre
748
7496(5) Maxilla without fringed or toothed lamellae (Figs 2f-g) ……………………………… 7
7506’Maxilla with fringed and/or toothed lamellae (Fig 2e) [NEO] ……… Anurida Laboulbène
751
7527(6) Eyes 8+8 ………………………………………………………………………………… 8
7537’Eyes 7+7 or less ………………………………………………………………………… 10
754
7558(7) Postantennal organ present; furcula short, not reaching the ventral tube when near the
756trunk ………………………………………………………………………………………… 9
7578’ Postantennal organ absent; furcula long, trespassing the ventral tube when near the trunk
758[NEO] …………………………………………………………………… Pseudanurida Schött
759
7609(8) Postantennal organ moruliform (Fig. 6f) [NEO] ……………… Aethiopella Handschin
7619’ Postantennal organ with one ring of vesicles (Fig. 6g) [NEO] … Pseudachorutes Tullberg
762
76310(7’) Postantennal organ present ………………………………………………………… 11
76410’ Postantennal organ absent ……………………………………………………………… 13
765
76611(10) Postantennal organ moruliform (Fig. 6f) [NEO], eyes 5 + 5 … Neotropiella Handschin
76711’ Postantennal and eyes organ otherwise …………………………………………… 12
768
76912(11) Furcula present [NEO] ……………………………………… Furculanurida Massoud
77012’ Furcula absent or reduced to small ventral projections (Fig. 6h) [NEO] …………………
771…………………………………………………………………………… Micranurida Börner
772
77313(10’) Furcula absent or reduced to small ventral projections …………………………… 14
77413’ Furcula present and developed [NEO] ………………………………… Arlesia Handschin
775
77614(13) Fourth antennal segment with one sensillum clearly thicker than others (Fig. 6c);
777furcula reduced to small ventral projections (Fig. 6h) [NEO] ……………… Hylaeanura Arlé
77814’ Fourth antennal segment sensilla subequal in thickness; furcula completely absent [NEO]
779……………………………………………………………………………… Paranura Axelson
780
781
782Collembola: Poduromorpha: Odontellidae Genera
783
7841 Fourth antennal segment with apical bulb (Fig. 7a) [NEO] …………… Odontella Schäffer
7851’ Fourth antennal segment without apical bulb [NEO] ……………… Superodontella Stach
786
787
788Collembola: Poduromorpha: Onychiuridae Genera
789
7901 Furcula reduced to a finely granulated area with 2+2 microsetae in two rows (Fig. 7b); sixth
791abdominal segment with or without anal spines …………………………………………… 2

31 16
32
7921’ Furcula reduced to a finely granulated area with 4 microsetae in one transverse row (Fig.
7937c); sixth abdominal segment with anal spines [NEO] ………………… Onychiurus Gervais
794
7952 Postantennal organ with compound vesicles (Fig. 7d); distal tibiotarsus with 7 setae in a
796whorl; sixth abdominal segment devoid of anal spines [NEO] ……… Agraphorura Pomorski
7972’ Postantennal organ with simple vesicles (Fig. 7e); distal tibiotarsus with 8 or more setae in
798a whorl; sixth abdominal segment with or without anal spines [NEO] ………………………
799……………………………………………………………………… Thalassaphorura Bagnall
800
801
802Collembola: Poduromorpha: Tullbergiidae Genera
803
8041 Third antennal segment sense organ with two enlarged guard sensilla (Fig. 3e); first thoracic
805segment without pseudocelli [NEO] …………………………………… Mesaphorura Börner
8061’ Third antennal segment sense organ with three enlarged guard sensilla; first thoracic
807segment with or without one pair of pseudocelli [NEO] ………………… Tullbergia Lubbock
808
809
810
811
812Collembola: Entomobryomorpha: Families
813
8141 Antennae with 4 segments; fourth abdominal segment more than 1.5 the length of the third
815abdominal segment in the midline (Figs. 1b, 8a–c), when less, then first and/or second
816antennal segments subdivided at the base (Figs. 8d–e); scales present or not (Figs. 8a, 9a–b);
817body setae mostly ciliate, multilaterally ciliate macrosetae usually present, sometimes
818cylindrical and with truncate or broadened tips (Fig. 25) …………………………………… 2
8191’ Antennae with 4 segments undivided; fourth abdominal segment less than 1.5 the length of
820the third abdominal segment in the midline (Figs. 8f–h); scales absent; body setae usually
821smooth or unilaterally ciliate, multilaterally ciliate macrosetae can be present, but always
822acuminate (Figs. 9c–d) …………………………………………………………… Isotomidae
823
8242(1) Dens dorsally crenulate, mucro short with one or two teeth always designed dorsally
825(Fig. 10a, 11a) ………………………………………………………………… Entomobryidae
8262’ Dens straight, not crenulate; mucro short with at least two teeth designed apically (Figs.
82710b, 11b–c), when inserted dorsally then mucro with 1/3 of the length of dens or more (Fig.
82810c) …………………………………………………………………………… Paronellidae
829
830
831Collembola: Entomobryomorpha: Isotomidae genera
832
8331 Furcula present, reduced or long (Figs. 8f–h; 12) ………………………………………… 2
8341’ Furcula completely absent [NEO] ……………….……………… Pseudanurophorus Stach
835
8362(1’) Manubrium ventrally without distal setae (Fig. 12) …………………………………… 3
8372’Manubrium ventrally with distal setae (Figs. 13a–b) ……………………………………… 7
838
8393(2) Abdominal segments without bothriotricha; mucro spatulate and with 2 teeth, without
840setae (Figs. 15b–c) …………………………………………………………………………… 4

33 17
34
8413’ Fifth abdominal segment with bothriotricha; mucro with 3 teeth and with one basal seta
842(Figs. 15a and 16) [NA] …………………………………………… Archisotoma Linnaniemi
843
8444(3) Eyes present; abdominal segments individualized (Figs. 8F and 18) ………………… 5
8454’ Eyes absent; fifth and sixth abdominal segments fused (Figs. 8H and 17) [NEO] …………
846………………………………………………………………………… Isotomodes Linnaniemi
847
8485(4) Dens tuberculate posteriorly; mucro with lamellae and distinctly separated from dens
849(Fig. 14a; 15b) ……………………………………………………………………………… 6
8505’ Dens smooth; mucro (if present) without lamellae and weakly separated from dens (Fig.
85112) [NEO] ………………………………………………………………… Folsomides Stach
852
8536(5) Dens ventrally with setae (Fig. 14a) [NEO] …………………………… Ballistura Börner
8546’ Dens without ventral setae [NA] …………………… Bonetrura Christiansen & Bellinger
855
8567(2’) Fourth to sixth abdominal segment individualized (Fig. 8F) ………………………… 8
8577’ Fourth to sixth or fifth and sixth abdominal segment fused (Fig. 17) …………………… 15
858
8598(7) Postantennal organ present (Figs. 18–19) ……………………………………………… 9
8608’ Postantennal organ absent [NEO] ……………………………………… Axelsonia Börner
861
8629’(8) Manubrium ventrally with 11+11 or more setae (Fig. 13a) ………………………… 10
8638’Manubrium ventrally with 6+6 or less setae (Fig. 13b) [NEO] ………… Proisotoma Börner
864
86510(9) Third and/or fourth abdominal segments with bothriotricha (Fig. 16) ……………… 11
86610’ Abdominal segments without bothriotricha …………………………………………… 12
867
86811(10) Mucro with 4 teeth, mucronal spine present or absent (Fig. 14a) [NEO] ………………
869…………………………………………………………………………….. Isotomurus Börner
87011’ Mucro with 3 teeth, mucronal spine absent (Fig. 15c) [NEO] ……………………………
871…………………………………………….……… Psammisotoma Greenslade & Deharveng
872
87312(10’) Manubrium ventrodistal without modified setae (Figs. 13a–b) …………………… 13
87412’ Manubrium ventrodistal with thickened or spine-like setae (Fig. 13c) [NEO] ……………
875………………………………………………………………………………… Isotoma Bourlet
876
87713(12) Dens distally narrowed and crenulated (Fig. 14a) ……………………………….… 14
87813’ Dens distally tuberculated (Fig. 14b) [NA] ………………………… Granisotoma Stach
879
88014(13) Second tibiotarsus anteriorly with two modified setae, usually apically feathered (Fig.
88120a), [NA] …………………………………………………………….… Halisotoma Bagnall
88214’ Second tibiotarsus without modified setae [NEO] …………… Desoria Agassiz & Nicolet
883
88415(7’) Postantennal organ absent …………………………………………………………… 16
88515’ Postantennal organ present (Figs. 18–19) ……………………………………………… 17
886
88716(15) Fourth antennal segment with two apical oval sensilla; tergal segments only with
888smooth setae (Fig. 21a) [NEO] ……………………………………………… Folsomina Denis
88916’ Fourth antennal segment with six apical oval sensilla; tergal segments with some serrated
890setae (Figs. 21b and 22a) [NEO] ………………………………………… Isotomiella Bagnall

35 18
36
891
89217(15’) Fifth and sixth abdominal segments fused (Fig. 17) ……………………………… 18
89317’ Fourth to sixth abdominal segment fused [NEO] ………………………. Folsomia Willem
894
89518(17) Postantennal organ elliptical (Fig. 18); fifth and sixth abdominal segments without
896modified setae ……………………………………………………………………………… 19
89718’ Postantennal organ subdivided in the middle; fifth and/or sixth abdominal segments with
898foil setae (Figs 19 and 22b) [NEO] ……………………………………… Hemisotoma Bagnal
899
90019(18) Eyes absent; tenent-hairs pointed (Fig. 20a) [NA] ………………… Dagamaea Yosii
90119’ Eyes present; tenent-hairs clavate or pointed (Figs. 18 and 20a–b) [NEO] ………………
902…………………………………………………………………………… Cryptopygus Willem
903
904
905Collembola: Entomobryomorpha: Entomobryidae Genera
906
9071 Antennae with 5 or 6 segments (first and second segments subdivided); fourth abdominal
908segment less than 1.5 the length of the third abdominal segment in the midline (Figs. 8d–e)
909………………………………………………………………………………………………… 2
9101’ Antennae with 4 segments; fourth abdominal segment more than 2.0 the length of the third
911abdominal segment in the midline (Figs. 8a–c) ……………………………………………… 5
912
9132(1) Body with scales (Figs. 8d, 9b, 23a–e) ………………………………………………… 3
9142’ Body without scales (Fig. 9a) …………………………………………………………… 4
915
9163(2) Body dorsally with scales apically pointed, absent on antennae and furcula; antennae
917longer than body and with 5 segments (first segment subdivided); dens without spines (Figs.
9188e and 23a) [SA, Br] ……………………………………………… Mastigoceras Handschin
9193’ Body dorsally with scales apically rounded or truncate, present on antennae and furcula;
920antennae normal to long, with 6 segments (first and second segment subdivided); dorsal dens
921generally with basal spines (Fig. 8d; 23b–c, 24a) [NEO] ……………. Dicranocentrus Schött
922
9234(2’) Postantenal organ present; mucro with 1 tooth and without basal seta (Figs. 25 and 26a)
924[SA, Br] ……………………………………………………….……………… Notobrya Arlé
9254’ Postantenal organ absent; mucro with 2 teeth and with basal seta (Fig. 26c) [NA, Mex] …
926……………………….………………………………………………… Orchesella Templeton
927
9285(1’) Body without scales (Fig. 9a) ………………………….……………………………… 6
9295’ Body with scales (Figs. 8a–c, 9b, 23a–e) ………………………….……………………… 7
930
9316(5) Mucro falcate (Fig. 26b) [SA]… ……………………………………… Drepanura Schött
9326’ Mucro bidentate (Fig. 11a) [NEO] ………………………………… Entomobrya Rondani
933
9347(5’) Mucro falcate and without basal seta (Fig. 26a) ……………………………………… 8
9357’ Mucro bidentate and with basal seta (Fig. 26c) …………………………………… 9
936
9378(7) Antennae longer than or as long as the body length, mesothorax strongly projecting over
938the head; ungues with large outer tooth, dorsal dens with long blunt setae (Figs. 8b, 27a, 28b)
939[SA] …………………………………………………………………… Lepidocyrtinus Börner

37 19
38
9408’ Antennae smaller than the body length; mesothorax round and not projecting over the head,
941ungues with normal outer tooth, dorsal dens without blunt setae (Figs. 1b, 27b, 28b) [NEO]
942………………………………………………………………………………… Seira Lubbock
943
9449(7’) Dental spines absent (Fig. 28b) ……………………………………………………… 12
9459’ Dental spines present (Figs. 24b–c) ………………………………………….………… 10
946
94710(9’) Body with scales apically truncate and rounded (Figs. 23b–e), rarely pointed (Fig,
94823a); ventral dens with scales (Fig. 29a) …………………………………………………… 11
94910’ Body with scales apically pointed; ventral dens without scales (Figs. 23a and 29b) [SA,
950Br, Amazon] ………………………………………………… Amazhomidia Cipola & Bellini
951
95211(10) Body with scales heavily ciliate and apically pointed (Fig. 23e); mesothorax to first
953abdominal segment with macrosetae (Fig. 30a); ungues with large outer tooth (Fig. 27a);
954dental spines in rows along of the dens (Fig. 24b) [SA] …………… Acanthocyrtus Handschin
95511’ Body with scales gently ciliate and apically truncate (Fig. 23d), mesothorax to first
956abdominal segment only with microsetae (Fig. 30b); ungues with normal outer tooth (Fig.
95727b); dental spines agglomerate on dens base (Fig. 24c) [SA] …………………………………
958………………………………………………………………… Lepidocyrtus (in part) Bourlet
959
96012(9) Mesothorax usually projecting over the head (Figs. 8a and 8c); ventral dens with scales
961(Fig. 29a) …………………………………………………………………………………… 13
96212’ Mesothorax not projected over the head (Fig. 1b); ventral dens without scales (Figs. 29b)
963[NA and West Coast of SA] ………………………………………… Willowsia Shoebotham
964
96513(12) Body with scales gently striate and apically truncate (Fig. 23d); mesothorax to first
966abdominal segment without macrosetae (Fig. 30b); ungues with normal outer tooth (Fig. 27b);
967manubrial plate without macrosetae [NEO] ……………………… Lepidocyrtus (s.l.) Bourlet
96813’ Body with scales heavily striate and apically pointed (Fig. 23e); mesothorax to first
969abdominal segment with macrosetae (Fig. 30a); ungues with large outer tooth (Fig. 27a);
970manubrial plate with at least two macrosetae per side (Fig. 31) [SA] …………………………
971………………………………………………………………………… Lepidocyrtoides Schött
972
973
974Collembola: Entomobryomorpha: Paronellidae: Genera
975
9761 Body pale, without pigments and eyes (Fig. 32a and 33a); dens dorsally with fringed scale-
977like setae; mucro long (Fig. 10c) …………………………………………………………… 2
9781’ Body with or without pigments (Figs. 32b–d); eyes 0+0 to 8+8 (Figs. 33a–b); dens dorsally
979without fringed scale-like setae; mucro short (Figs. 10b, 11b–c) …………………………… 3
980
9812(1) Dens dorsally with ciliated setae between large scales (Fig. 10c) [NEO] ………………
982…………………………………………………………………………… Cyphoderus Nicolet
9832’ Dens dorsally without setae between large scales [SA, Rapa Nui and Bariloche] …………
984…………………………………………………………… Serroderus Delamare Deboutteville
985
9863(1’) Body with scales (Figs. 32d and 34a); dens dorsally with spines (Fig. 10b); mucro base
987without appendage (Fig. 11b) ……………………………………………………………… 4
9883’ Body unscaled (Fig. 34b); dens without spines; mucro base with enlarged (vesicular?)
989spatula-like appendage (Fig. 11c) [NEO] ……………………………… Salina MacGillivray*

39 20
40
 990
9914(3) Antennae longer than the body; metathorax enlarged and conspicuously humped,
992resulting in a bent body between metathorax and first abdominal segment (Figs. 32c and 35);
993third tibiotarsus only with ciliated setae (Fig. 36a) [NEO]…………… Campylothorax Schött
9944’ Antennae of the same length or shorter than the body (longer only in cave forms);
995metathorax normal (Fig. 32d); third tibiotarsus with one internal smooth setae (Fig. 36b)
996[NEO] …………………………………………………………………… Trogolaphysa Mills
997
998
999
1000Collembola: Symphypleona: Families
1001
10021 Female with subanal appendage (Figs. 37a–c); males antennal segments without any
1003peculiar modification ………………………………………………………………………… 2
10041’ Female without subanal appendage; males second and third antennal segments modified in
1005prehensile organ (Figs. 46a–b) ……………………………………………… Sminthurididae
1006
10072(1) Female subanal appendage directed toward the anus or posteriorly (Figs. 37a–b) …… 3
10082’ Female subanal appendage directed toward the genital opening (Fig. 37c) ……………… 6
1009
10103(2) Fourth antennal segment longer than third (Fig. 1c) …………………………………… 4
10113’ Fourth antennal segment much shorter than third (Fig. 38) ………………… Dicyrtomidae
1012
10134(3) Third antennal segment without papilla; female subanal appendage directed toward the
1014anus (Fig. 37a) ……………………………...……………………………………...………… 5
10154’ Third antennal segment with a low papilla at dorsal and medial position (Fig. 39); female
1016subanal appendage directed posteriorly (Fig. 37b) [NEO] ……………………………………
1017…………………………………………………… Sturmiidae, one genus: Sturmius Bretfeld
1018
10195(4) Third trochanter often with a posterior spine (Fig. 40); tibiotarsi often without capitate
1020tenent hairs; pretarsi with anterior and posterior setae ……….……………… Sminthuridae
10215’ Third trochanter without posterior spine; tibiotarsi always with capitate tenent hairs (Figs.
102258a–c); pretarsi only with anterior seta, sometime without anyone …….… Bourletiellidae
1023
10246(2’) Neck without peculiar organs (Fig. 1c) ………………………………………………….7
10256’ Neck with one pair of lateral soft organs (Fig. 41) [SA, Patagonia] ………………………
1026……………… Spinothecidae, one genus: Spinotheca Delamare Deboutteville sensu Bretfeld
1027Actually, there are two genera Spinotheca Delamare Deboutteville and Troglospinotheca
1028Palacios-Vargas
10297 Eyes at most with 4 lenses per side; tibiotarsi without capitate tenent hairs (Fig. 49) … 7
10307’ Eyes with 6–8 lenses per side; tibiotarsi with capitate tenent hairs (Fig. 42) ………………
1031………………………………………………………………………………….… Katiannidae
1032
10338(7) Fifth abdominal segment included in large abdomen; bothriotrichum D short and
1034capitated (Fig. 43) ………………………… Collophoridae, one genus: Collophora Richards
10358’ Fifth abdominal segment partially included in large abdomen; bothriotrichum D long and
1036with normal form [NEO] ………………… Arrhopalitidae, one genus: Arrhopalites Börner
1037
1038
1039Collembola: Symphypleona: Sminthurididae Genera

41 21
42
1040
10411 Third tibiotarsus without organs …………………………………………………………… 2
10421’ Third tibiotarsus with a distal pair of organs (Fig. 44) [NEO] …… Sminthurides Börner †
1043
10442 Mucro with basal seta (Fig. 45); male with second and third antennal segments highly
1045modified (Fig. 46a); first tibiotarsus with four proximal organs (Fig. 47) [NEO] ……………
1046…………………………………………………………………… Denisiella Folsom & Mills
10472’ Mucro without seta; male with second and third antennal segments subtly modified (Fig.
104846b); first tibiotarsus without organs [NEO] …………………… Sphaeridia Linnaniemi
1049
1050Collembola: Symphypleona: Dicyrtomidae Genera
1051
10521 Adults without bothriotrichum D; head clypeal region with four or more median setae (Fig.
105348a) ……………………………………………………………………………………… 2
10541’ Adults with bothriotrichum D; head clypeal region with two median setae (Fig. 48b) [NEO]
1055……………………………………………………………………… Ptenothrix Börner
1056
10572(1) Unguis with tunica well developed (Fig. 49) …………………………………………… 3
10582’ Unguis with tunica poorly developed [NA] …………………………… Dicyrtoma Bourlet
1059
10603(2) Head and anterior region of large abdomen with spininiform setae (Figs. 48b),
1061neosminthuroid setae absent [NEO] ………………………………… Dicyrtomina Börner
10623’ Head and anterior region of large abdomen with normal setae (Figs. 48a), neosminthuroid
1063setae present (Fig. 50a) [NEO] ………………………… Calvatomina Yosii
1064
1065
1066Collembola: Symphypleona: Sminthuridae Genera
1067
10681 Base of furcula with neosminthuroid setae (Fig. 50b–c) ………………………………… 2
10691’ Base of furcula without neosminthuroid setae …………………………………………… 5
1070
10712(1) Base of furcula with 1 neosminthuroid setae per side (Fig. 50b) ……………………… 3
10722’(1’) Base of furcula with 3 neosminthuroid setae per side (Fig. 50c) [NEO] ………………
1073…………………………………………………………………… Songhaica Betsch & Dallai
1074
10753(2) Fourth antennal segment well sub-segmented (Fig. 1c); large abdomen in adults with
1076bothriotricha B and D and without mesothoracic vesicles …………………………………… 4
10773’ Fourth antennal segment undivided or weakly sub-segmented; large abdomen in adults
1078without bothriotricha B and D, and with 1 mesothoracic vesicle per side (Fig. 51) [NA] ……
1079…………………………………………………………………………… Neosminthurus Mills
1080
10814(3) First and second trochanters without spine [NEO] ………………… Sphyrotheca Börner
10824’ First and third trochanters, (and sometime even the second) with spine (Fig. 40) [NEO]
1083………… ……………………………………………………………… Szeptyckithca Betsch
1084& Weiner
1085
10865(1’) Small abdomen with bothriotrichum D smooth and acuminate ……………………… 6
10875’ Small abdomen with bothriotrichum D clavated and ciliated (Fig. 52) [SA, Patagonia]
1088……… ……………………………… Austrosminthurus Delamare Deboutteville & Massoud
1089

43 22
44
10906(5) Third trochanter with spine (Fig. 40); head without post antennal seta …….……………7
10916’ Third trochanter without spine; head with post antennal seta present (Fig. 53) [NA] ………
1092…………………………………………………………………………… Sminthurus Latreille

10937(6) Unguis with normal shape, without any peculiarity (Figs. 58a–c) ……………………… 8
10947’ Unguis with an anterior external cavity (Fig. 54) [SA, Br] …………………………………
1095…………………………………………… Varelasminthurus Silva, Palacios-Vargas & Bellini
1096
10978(7) Fourth antennal segment very long, with 20-40 sub-segments; large abdomen without
1098macrosetae [NEO] ………………………………………………………… Temeritas Richards
10998’ Fourth antennal segment not very long, with less than 20; large abdomen with dorsal
1100macrosetae [NEO] ………………………………………… Pararrhopalites Bonet & Tellez
1101
1102
1103Collembola: Symphypleona: Bourletiellidae Genera
1104
11051 Third tibiotarsus with 3 capitate tenent hairs (Fig. 58a) ………………………………… 2
11061’ Third tibiotarsus with 2 capitate tenent hairs (Fig. 58b) ………………………………… 3
1107
11082 Unguiculus short, less than half the length of unguis; dens with long posterior internal and
1109external setae, longer than the space between them; males head frons with a median group of
1110modified setae [NEO] …………………………………………… Pseudobourletiella Stach
11112’ Unguiculus long, reaching or surpassing the apex of unguis; dens with normal setae; males
1112interocular region with a dorsal large salience (horn) with 2 or 3 long setae [NA, Virgin
1113Islands] ………………………………………………… Bovicornia Delamare Deboutteville
1114
11153 Large abdomen with dorsal spines (Fig. 55a) ……………………………………………….4
11163’Large abdomen without spines (Fig. 55b) ………………………………………………… 5
1117
11184(3) Head triangular (Fig. 56a); tibiotarsi inner surface with thick and serrated setae (Fig. 57);
1119unguiculus longer than unguis; large abdomen without neosminthuroid setae [NEO]
1120………………………………………………………… Stenognathriopes Betsch & Lasebikan
11214’ Head roundish (Fig. 55a); tibiotarsi inner surface with acuminated setae; unguiculus shorter
1122than unguis (Fig. 58a); large abdomen with neosminthuroid setae near the base furcula (Fig.
112355a) [NEO] …………………………………………………………… Adisianus Bretfeld
1124
11255(3) Tibiotarsi inner surface with acuminated setae (Fig. 58b); males with modified setae
1126between antennae and/or in small abdomen (Fig. 56b and 60) ……………………………… 6
11275’ Tibiotarsi inner surface with obliquely truncated or distally flattened setae (Fig. 58c); males
1128without any special sexual dimorphism [NEO] ……… Prorastriopes Delamare Deboutteville
1129
11306(5) Dens often crenulated with long internal and external setae (Fig. 59); males with space
1131between antennae with two pairs of spines (Fig. 56b) [NEO] …… Arlesminthurus Bretfeld*
11326’ Dens never crenulated with normal size setae; males with posterior region of large
1133abdomen and small abdomen with thick and modified setae (Fig. 60) [NEO] …………………
1134……………………………………………………………………………… Bourletiella Banks
1135
1136
1137Collembola: Symphypleona: Katiannidae Genera
1138

45 23
46
11391 Head roundish and mouthparts normal (Fig. 61a) ………………………………………… 2
11401’ Head and mouthparts elongated (Figs. 61b and 62) [NEO]………… Stenognathellus Stach
1141
11422(1) Fourth antennal segment subdivided, base of furcula without neosminthuroid seta …… 3
11432’ Fourth antennal segment not subdivided, base of furcula with neosminthuroid seta (Fig. 63)
1144………………………………………………………………………………………………… 4
1145
11463(2) Head with frontal thick spines [NEO] ………….…………. Katianna Maynard & Downs
11473’ Head only with normal setae [NEO] ………………………...……… Polikatianna Salmon
1148
11494(2’) Small abdomen with median circumanal seta bifurcated (Fig. 37c) [NEO] ……………
1150…………………………………………………………………………… Sminthurinus Börner
11514’ Small abdomen with median circumanal seta normal [BRA, Amz] ………………………
1152………………………………………………………………………...…. Arborianna Bretfeld
1153
1154
1155Collembola: Neelipleona: Neelidae Genera
1156
11571 Head and large abdomen with sensory fields (Figs. 64a–b); labrum without fringes or ciliate
1158edges ………………………………………………………………………………………… 2
11591’ Head and large abdomen without sensory fields; labrum with fringes and ciliate anterior
1160edges (Fig. 65) [NEO] ………………………………………………………… Neelides Caroli
1161
11622(1) Third and fourth antennal segments fused (Fig. 66); abdomen sensory field with 5
1163marginal setae (Fig. 64b) [NEO] ……………………………………… Megalothorax Willem
1164
11652’ Third and fourth antennal segments separated; abdomen sensory field with 2 marginal setae
1166(Fig. 64a) [NEO] ………………………………………………………… Neelus Folsom
1167
1168YOU FORFOT TO INCLUDE Spinothorax Papác et Palacios-Vargas, 2016
1169
1170REFERENCES
1171
1172Arbea, J.I. 2005. Agraphorura calvoi n. sp. from Venezuelan caves (Collembola:
1173Onychiuridae). International Journal of Speleology 34(1-2): 19-24.
1174
1175Arlé, R. 1959. Collembola Arthropleona do Brasil oriental e central. Separata dos arquivos do
1176Museu Nacional 49: 155-211.
1177
1178Arlé, R. 1960. Notas sôbre a família Oncopoduridae, com descrição de duas espécies novas
1179do Brasil (Collembola). Arquivos do Museu Nacional Rio de Janeiro 50: 9-23.
1180
1181Arlé, R. 1971. Collemboles d´Amazonie, III. Quelques Symphypléones du Bas-Amazone et
1182complément à la description de Deuterosminthurus aueti Arlé, 1961. Boletim do Museu
1183Paraense Emilio Goeldi 75: 1-11.
1184
1185Bellinger, P.F. 1985. A new family of Collembola (Arthropoda, Tracheata). Caribbean Journal
1186of Science 21(3-4): 117-123.
1187

47 24
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1188Bellinger, P.F., K.A. Christiansen, & F. Janssens. 2016. Checklist of the Collembola of the
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