Research Article |
Corresponding author: Charlene Janion-Scheepers ( cjanion@gmail.com ) Academic editor: Wanda M. Weiner
© 2017 Mikhail Potapov, Charlene Janion-Scheepers, Louis Deharveng.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Potapov M, Janion-Scheepers C, Deharveng L (2017) Taxonomy of the Cryptopygus complex. II. Affinity of austral Cryptopygus s.s. and Folsomia, with the description of two new Folsomia species (Collembola, Isotomidae). ZooKeys 658: 131-146. https://doi.org/10.3897/zookeys.658.11227
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Folsomia minorae sp. n. and F. australica sp. n. are described from New Zealand and Australia, respectively. Their possible affinity to two different groups of Cryptopygussensu stricto is discussed. Attention is paid to the variability of sensillary patterns of the genital segment in Cryptopygus: mainly, all s-chaetae are subequal, but in more advanced forms the dorsal triplet, lateral duplet or either of them become macrochaeta-like in length. Cryptopygus ulrikeae (= Folsomia ulrikeae Najt & Thibaud, 1987), comb. n. is given a new generic position.
Australia, New Zealand, taxonomy
The genus Cryptopygus Willem, 1902 sensu stricto has not received its modern generic diagnosis and most of its “austral” species need to be revised (
The notation system accp-as-al (
Abd. I–VI abdominal segments I–VI
accp accessory p-row s-chaeta
Ant. I–IV antennal segments I–IV
as anterosubmedial s-chaeta
a.s.l. above sea level
bms basal micro s-chaeta on antennal segments
e7 ‘guard’ of labial papilla E
Leg I, II, III first, second and third pairs of legs
M macrochaeta
ms micro s-chaeta(e) (=microsensillum(a) auct.)
s macro s-chaeta or s-chaetae (=macrosensillum(a) or sensillum(a) auct.)
s.s. sensu stricto
PAO postantennal organ
Th.II–III thoracic segments II and III
U3 inner edge of unguis
MNZTPT Museum of New Zealand Te Papa Tongarewa, Wellington
SAMA South Australian Museum, Adelaide
MSPU Moscow State Pedagogical University, Russia.
Holotype: adult ♀. New Zealand, southern South Island, Central Otago, Pisa Range, 44°52'03"S, 169°9'33"E, 1700 m a.s.l., in soil and debris under Dracophyllum muscoides cushion, 18.ii.2014, coll. M. Minor (on slide). Paratypes. 10 paratypes, subadult ♀♀ and ♂♂of similar size with holotype, 7 of which from the same locality (and in close proximity), and 3 from Central Otago, The Remarkables Mts, 45°3'42"S, 168°48'40"E, 1829 m a.s.l., herbaceous snowbank, in soil, 19.ii.2014, coll. M. Minor (all on slides). Holotype and 4 paratypes kept in MNZTPT; 5 paratypes in MSPU.
One ♀ identified in all details as F. minorae sp. n. by A. Fjellberg (not seen by us): New Zealand, South Island (northern part), Avalanche Peak trail above Arthur Pass, 42°56'26"S, 171°33'29"E, forest litter, 23.i.2004, coll. A. Fjellberg.
Folsomia species with 5+5 ocelli; slender subapical organite of Ant.IV; clavate tibiotarsal hairs; outer teeth on claws; stout dens with few chaetae and a large mucro; and characteristic ‘3+2’ sensillary pattern of s-chaetae on Abd.V.
Body size of the only adult female 1.75 mm. Dark blue, appendages paler. Body cylindrical (Fig.
The name is given after Maria Minor, who kindly provided some of the material on the new species.
To date eight species of Folsomia are known from New Zealand (
Differentiating characters of the new species are: five ocelli, unique subapical organite of Ant.IV, clavate tibiotarsal hairs, presence of outer teeth on claws, stout dens with few chaetae, and a very large mucro. Well differentiated ‘3+2’ sensillary pattern of s-chaetae on Abd.V is also characteristic (see below). Three long and slender dorsal s-chaetae of Abd.V are found in species of several groups of Folsomia of the Holarctic (i.e. F. penicula Bagnall, 1939, F. quadrioculata (Tullberg, 1871) and F. sensibilis Kseneman, 1936), which belong to either ‘3+2’ or ‘3+1+1’ patterns.
F. minorae sp. n. is known from three localities in South Island, New Zealand. It is probably a species restricted to mountainous areas.
Folsomia minorae sp. n. (7–12) and F. australica sp. n. (13) 7 apex of Ant.IV, lateral view 8 chaetotaxy of posterior part of Abd.IV, Abd.V and VI 9 furca, tenaculum, and furcal subcoxae, posterior view 10–11 apical part of Leg II (10) and III (11) 12–13 s-patterns chaetae of Abd.IV–V (lateral s of Abd.IV not shown). org—organite, sms—subapical ms, pin—pin-chaeta. Scal bar 0.1 mm in 8, 0.03 mm in 9, others, 0.01 mm.
Holotype: adult ♀. Australia, Chiltern National Park, Victoria, heathy dry forest, mostly native, 36°7'53"S, 146°36'20"E, 12.iv.2015, M. Lythe leg. Four paratypes, one adult ♀, one adult ♂and two sub-adult ♀♀ of the same size as adults. Holotype and two paratypes in SAMA, two paratypes in MSPU.
10 specimens in ethanol (SAMA). Australia, Victoria, Mt Pilot National Park, 36°18'45"S, 146°33'16"E. 20.vii.2015, M. Lythe leg.
Folsomia species with 1+1 ocelli; chaetotaxy of dens 12/6; tridentate mucro; 2 lateral s-chaetae on Abd.V clearly longer than 3 dorsal ones; 2+2 chaetae on anterior side of manubrium.
Body size from 0.60 (adult male) to 0.75 mm (one of sub-adult females). White, with one black ocellus on each side of the head (Fig.
The name is given after the geographical distribution of the new species.
Folsomia australica sp. n. resembles the only other native Australian species of the genus, i.e. F. loftyensis (Womersley, 1934) (after the redescription of
Morphological features of the furca of F. australica and F. loftyensis, especially the tridentate mucro, are shared with several species of Cryptopygus: C. tricuspis Enderlein, 1909 (sub-Antarctic), C. insignis Massoud and Rapoport, 1968 (South America), C. patagonicus Izarra, 1972 (South America), and three unnamed species from South Africa (Cryptopygus sp. 5, C. sp. 6, and C. sp. 7, see below). These species probably represent another group of Cryptopyguss.s., dissimilar to C. antarcticus, which could be ancestral to the “Australian” species of Folsomia.
Folsomia australica sp. n. is known from two localities in south-eastern part of Australia (Victoria and New South Wales).
Folsomia australica sp. n. 14–15 chaetotaxy of posterior part of abdomen, lateral (14) and dorsal (15) views 16–17 furca, lateral (16) and anterior (17) view 18 manubrium, posterior view 19 apex of Ant.IV, dorsal view, left antenna 20 distal part of Leg III 21 ocellus and PAO. org—organite, sms—subapical ms. Scale bar 0.05 mm in 14, 15, others, 0.01 mm.
The fifth abdominal segment of all species of the genus Cryptopygus, as in Folsomia, has five s-chaetae on each side: accp1, accp2, accp3, accp4 and as. The s-pattern of C. antarcticus and several other species is probably the most primitive as it consists of regularly scattered sub-equal short and thin s-chaetae (Fig.
1. Weakly differentiated “3+2” pattern (Figs
Representatives:
– C. antarcticus antarcticus Willem, 1902. Material: several locations in Antarctic Peninsula (King George Isl., Deception Isl., Devil Isl.) leg. D. Russell.
– C. antarcticus maximus Deharveng, 1981. S-chaetae as in nominate subspecies. Material: Kerguelen Isl. leg. J. Travé.
– C. antarcticus reagens (Enderlein, 1909). S-chaetae of dorsal triplet almost twice longer than s-chaetae of duplet. Material: Crozet Isl. (sub-Antarctic), leg. J. Travé.
– C. antarcticus travei Deharveng, 1981. S-chaetae of dorsal triplet almost as long as those of duplet. Material: Marion Isl. (sub-Antarctic), leg. J. Travé.
– C. araucanus Massoud & Rapoport, 1968. S-chaetae of dorsal triplet slightly longer than s-chaetae of duplet. Material: syntypes, several locations in Argentina (Futalaufquen, Lago Curruhé, Lago Menendez). Collections of the Museum national d’Histoire naturelle (Paris, France).
– C. badasa Greenslade, 1995. All s-chaetae very short, “triplet” s-chaetae slightly longer. Material: Antarctic Peninsula (Devils Isl.), leg. D. Russell; South Georgia (sub-Antarctic), leg. V. Bulavintsev.
– C. binoculatus Deharveng, 1981. S-chaetae subequal. Material: holotype, Crozet Isl. (sub-Antarctic).
– C. insignis Massoud & Rapoport, 1968. S-chaetae subequal. Material: syntypes, Lago Menendez (Argentina). Collections of the Museum national d’Histoire naturelle (Paris, France).
– C. hirsutus (Denis, 1931). S-chaetae subequal. Material: possible syntypes, Costa Rica. Collections of the Museum national d’Histoire naturelle (Paris, France).
– C. lawrencei Deharveng, 1981. “Triplet” s-chaetae slightly longer. Material: Kerguelen Isl. (sub-Antarctic), leg. J. Travé.
– C. pilosus (Womersley, 1934). S-chaetae as in C. antarcticus. Material: South Australia, Lofty Ranges, leg. P. Greenslade.
– C. tricuspis Enderlein, 1909. S-chaetae as in C. antarcticus. Material: Kerguelen Isl. (sub-Antarctic), leg. J. Travé.
– C. ulrikeae (Najt & Thibaud, 1987), comb. n. S-chaetae sub-equal. Separation of Abd.IV and V as in other species of the genus Cryptopygus. Primarily, it was described as Folsomia ulrikeae (Najt and Thibaud, 1987). Material: holotype, Ecuador. Collections of the Museum national d’Histoire naturelle (Paris, France).
– Cryptopygus sp. 1 (complex ‘antarcticus’). S-chaetae as in C. antarcticus. Characters common with the nominotypic subspecies of C. antarcticus but body more slender. Material: New Zealand (South Island), leg. M. Minor.
– Cryptopygus sp. 2 (complex ‘antarcticus’). S-chaetae as in C. antarcticus. With the characters of C. antarcticus but ms formula of body tergites as 10/000. Material: New Zealand (North Island).
– Cryptopygus sp. 3. S-chaetae as in C. antarcticus. With 6+6 ocelli. Manubrium without anterior chaetae Dens with one anterior chaeta, mucro bidentate. Material: South Africa (Jonkershoek), leg. C. Janion-Scheepers.
– Cryptopygus sp. 4. S-chaetae as in C. antarcticus. With 8+8 ocelli. Dens rather long, with nine anterior and 5 posterior chaetae, mucro bidentate. Common chaetae and macrochaetae on body short. Material: South Africa, (Sutherland), leg. C. Janion-Scheepers.
2. “3+2” pattern with development of dorsal triplet (Figs
– Cryptopygus sp. 5. Dens with 9-10 anterior and 6 posterior chaeta, mucro tridentate. Material: South Africa (Table Mountain), leg. L. Deharveng and A. Bedos.
3. “3+2” pattern with development of lateral duplet (Figs
– Cryptopygus sp.6. With 4+4 or 5+5 ocelli. PAO with strong inner denticles. Dens with ten anterior chaetae, mucro tridentate. Material: South Africa (Little Switzerland), leg. E. Krzemińska.
4. “3+2” pattern with development of all s-chaetae (Figs
– Cryptopygus sp.7. With 4+4 ocelli. Dens with 11-12 anterior and five posterior chaeta, mucro tridentate. Material: South Africa (Sutherland), leg. C. Janion-Scheepers.
S-chaetae patterns of Cryptopygus are probably more diverse than shown above: Cryptopygus yosii Izarra, 1965 (Argentina, after our study of a syntype) shows “3+1+1” pattern in which accp3 is thick and tubular and accp4 is short and moved to the latero-ventral position. More material on less primitive species needs to be studied to complete the generic overview. Nevertheless, s-patterns of Abd.V in Cryptopygus seem to be less divergent than in the larger genus Folsomia (
The dorsal fusion or separation of genital (Abd.V) and pre-genital segment (Abd.IV) is traditionally considered to be of great taxonomic value in the classification of the subfamily Anurophorinaes.l., and the genus Folsomia is defined by the apomorphic condition of this character (fusion). Based on the available literature and our own observations, the s-chaetotaxy of “austral” Cryptopyguss.s. shows principally the same characteristics as in Folsomia, particularly 4,3/2,2,2,3,5 set and arrangement of s-chaetae on Abd.V. The more adaptive characters (furca, ocelli, etc.) vary considerably within both genera. Therefore Abd.IV–V fusion seems to be the only apomorphic character that separates Folsomia from Cryptopygus and the former genus can be easily derived from the latter. This key character can potentially show a high level of homoplasy and Folsomia is probably a polyphyletic or paraphyletic group. In the Northern Hemisphere, the high diversity of Folsomia makes it difficult to find an appropriate ancestor or even ancestors among known taxa. In contrast, at least three “austral” native Folsomia mentioned above show much in common with certain species of Cryptopyguss.s. Thus, all the main characters of F. minorae sp. n. (ocelli, clavate tibiotarsal hairs, outer teeth on claws, dens and mucro) indicate its close relationship to a group of species similar to Cryptopygus antarcticus (
The generic position of both lineages, F. minorae and F. australica-loftyensis, can be modified in the future, depending on the increase of knowledge on the generic groups “Cryptopygus” and “Folsomia”. The genus Folsomia is also very diverse in the Holarctic and consists of several species groups of which several differ in characters of great taxonomical value and may justify the status of new separate genera.
We thank Maria Minor (Massey University, New Zealand) and Arne Fjellberg (Norway) who provided material on F. minorae sp. n. The material belonging to the following persons was also used: Valery Bulavintsev (Moscow), Penelope Greenslade (Ballarat), Morgan Lythe (Melbourne), David Russell (Görlitz) and Wanda M. Weiner (Kraków). We thank the New Zealand Department of Conservation for the collection permit (national authorization # 38116-GEO to M. Minor), the staff of the Snow Farm for help in providing access, Dr. Alastair Robertson (Institute of Agriculture & Environment, Massey University) for help with fieldwork, CapeNature and SanParks (South Africa) and Parks Victoria (Australia) for collection permits. The work in South Africa was supported by the Protea I and II South Africa-France bilateral grants (no. 68652 for L. Deharveng). This work was partly supported by Museum national d’Histoire Naturelle (Paris, France) and RFBR research project № 14-04-01140 (Russia) for MP. Jean-Marc Thibaud (Paris) lent some material of Cryptopygus. This work would not have been possible without creative initiative of Anatoly Babenko (Moscow). We are grateful to Arne Fjellberg, Wanda Weiner and three anonymous reviewers for their comments. The collections in Antarctic Peninsula were supported by Federal Environment Agency and funded by the German Federal Ministry for the Environment, Nature Conservation and Nuclear Safety under the project nr. FKZ 3709 85 157. CJS was supported by Australian Research Council grant DP140102815.