Collembola: Onychiuridae), with a Description of Four New Species from Caves
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SYSTEMATICS Review of North American Species of the Genus Onychiurus (Collembola: Onychiuridae), With a Description of Four New Species From Caves 1 1 2 ROMUALD J. POMORSKI, MICHAŁ FURGOŁ, AND KENNETH CHRISTIANSEN Ann. Entomol. Soc. Am. 102(6): 1037Ð1049 (2009) ABSTRACT The history of the genus Onychiurus is discussed. Four new Onychiurus species are described from North American caves: steinmanni and nathanieli from Colorado, reluctoides from Indiana, and furcisetosus from Virginia. Onychiurus reluctus Christiansen, 1961 is redescribed. A key for all of the known North American species is provided. KEY WORDS Collembola, Onychiurus, new species, cave The genus Onychiurus was created by Gervais in 1841 WrayÕs description is clearly erroneous in some re- with type species Podura ambulans Linnaeus. The spects and inadequate in others such that certain name has had an extremely checkered history (Ellis placement will have to await further material study. and Bellinger 1973) but became widely used by 1900 The junior author, K. C., has several additional species with many species of the genus having been described with inadequate materials to merit description. This under one or another synonym. The subgenus Prota- will be a fertile Þeld for new investigators. All of the phorura was created by Absolon in 1901 and was new species herein described come from caves. If we widely used and elevated to generic status by Stach in accept PomorskiÕs redeÞnition of Onychiurus, then 1954 (Stach 1954). In 1948 and 1949, Bagnall (1948, most the species of the genus can be distinguished 1949) created several new genera from the described from most species of the Onychiurini by having only species of these two genera and in 1954 Stach accepted nine or fewer setae in the distal tibiotarsal whorl. The most of these as valid genera, adding two more. Iron- Nearctic species are distinguished from most of the ically, in his key to the world species, Stach used only thus narrowly deÞned genus members by lacking Onychiurus sensu lato rather than any of these new male ventral organs. We found only seven species of generic names. Gisin (1952, 1960) accepts two Ony- the genus having this feature. Of these species, one, O. chiurus sensu lato taxa as subgenera: Protaphorura and aguzouensis Deharveng, 1978 is distinguished from all Oligaophorura but does not key them out separately. Nearctic species by having four pseudocelli at each In 1996, Weiner revised the genera of old Onychiurus antennal base (Deharveng 1978). A second species, O. sensu lato accepting and redeÞning most of the genera obsiones Cassagnau, 1963, has pseudocelli on the ven- created by Bagnall, Stach, and others and creating ter of the thorax (Cassagnau 1963), unlike any Nearc- several new genera. The now narrowly deÞned genus tic species. O. decemsetosus Yosii, 1966 is unlike any Onychiurus has Ͻ40 species worldwide. It is impossi- other species in having two ϩ two pseudocelli on the ble at the moment to know how many because many Þrst thoracic segment (Yosii 1966). It is possible that of the features used in the new deÞnition of the genus the loss of the male ventral organ is a troglomorphic are unknown in these species. The characteristics of feature because almost all species having this feature these redeÞned genera can be seen in the Onychiurus are either troglophile or troglobiotic. key on the Janssens and Christiansen website at All holotypes are deposited in The Museum of Com- http://www.collembola.org. parative Zoology (MCZ) at Cambridge, Harvard Uni- The genus Onychiurus, sensu Weiner (1996) and versity. The chaetotaxy terminology is that used in Pomorski (1996, 1998) is represented in North Amer- Pomorski (1998). Drawings were made using a camera ica by only one clearly described species O. reluctus lucida. (Bellinger et al. 2008). A second species, O. wilchi Wray 1950 almost certainly belongs in this genus, but Onychiurus reluctus Christiansen, 1961 the type specimens are in such poor condition and (Figs. 1Ð9) 1 Department of Biodiversity and Evolutionary Taxonomy, Zoolog- Redescription. Color white. Length (without an- ical Institute, Wrocław University, Przybyszewskiego 63/77, PL-51- tennae) of adults: holotypeÐ1.94 mm, reproductive 148 Wrocław, Poland. 2 Corresponding author: Department of Biology, Grinnell College, maleÐ1.25 mm, femalesÐ1.15Ð2.0 mm. Body shape cy- Grinnell, IA 50112-0806 (e-mail: [email protected]). lindrical with anal spines. Furca reduced to small area 0013-8746/09/1037Ð1049$04.00/0 ᭧ 2009 Entomological Society of America 1038 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 6 Figs. 1–2. Onychiurus reluctus (Christiansen, 1961): (1) habitus and dorsal chaetotaxy; (2) habitus and ventral chaetotaxy. All Þgures of type specimens. of Þne granulation with 2ϩ2 setae posterior, arranged antennal segment IV in latero-external position, ap- in one row. Granulation more or less uniform, distinct, proximately one-third length from the base. Antennal somewhat coarser on terga, head capsule and around segment III with microsensillum slightly below anten- anal spines. Antennal bases marked? from head. An- nal III sense organ. Thoracic terga II and III with tennae slightly shorter than head. Antennal segment microsensilla located laterally. Antennal organ III with IV with a subapical organ, consists of two pits, from Þve guard setae, two sensory rods, two bent and which the Þrst is deep and usually ampulla like, second smooth sensory clubs and Þve papillae usually irreg- is shallow, with small sensillum. Microsensillum on ular in shape. Postantennal organ consists of 15Ð16 November 2009 POMORSKI ET AL.: REVIEW OF NORTH AMERICAN Onychiurus 1039 Figs. 3–9. Onychiurus reluctus (Christiansen, 1961): (3) antennal III sense organ with papillae and without papillae; (4) top of antennal segment IV; (5) labium, A and C thickened labial papillae; (6) postantennal organ and anterior cephalic pseudocelli; (7) macro-, meso-, and microchaetae; (8) remnant of furca; (9) tibiotarsal chaetotaxy and claw of legs III. vesicles, covered with secondary vesicles. Pseudocel- ferentiated into macro-, meso-, and microsetae. lar formula: dorsal 32/133/33353, ventral 2/000/1112; Macro and mesochaetae minutely forked at the tip subcoxae I of legs I-III with 2-2-2 pseudocelli. Formula (Fig. 7). Body sensilla cylindrical and hidden. Ab- of parapseudocelli ventrally 00/000/111003. Dorsal dominal tergum IV with one medial seta. Subcoxae chaetotaxy (Fig. 1), nearly symmetrical, well dif- with 5Ð5-5 setae. No thoracic ventral setae. Ventral 1040 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 6 tube of males and females with 10ϩ10 setae. Furcula sity, paratypes will be deposited in the collection of vestigal. Claws without teeth. Empodial appendage K. C.). without basal lamella, appendage length Ϸ0.9Ð1.0 Etymology. The species name is derived from two mm inner edge of claw. Tibiotarsi IÐIII with nine Latin words “furca” and “setae”, that mark charac- distal setae. Male ventral organ absent. teristic feature showing very distinctly forked mac- Type Material. HOLOTYPE: Reproductive female ro- and mesochaetae on the dorsum and sides of (mounted on slide) USA, IA, Jackson County, Cave body. 1958, leg. K. Christiansen. Deposited MCZ. PARA- Remarks. The furcate macrochaetae distinguish this TYPES: 4 reproductive females (mounted on two species from all congeners. slides); same data as holotype. Other material: 1 reproductive male and 3 females: USA, WI, Craw- ford County, Star Valley Cave, fungus debris; May VII 1958, leg. K. Christiansen. (Deposited in collec- Onychiurus nathanieli sp. nov. tion of K.C.) (Figs. 20Ð27) Description. Color white. Length (excluding an- tennae) of adults: holotypeÐ1.16 mm, reproductive Onychiurus furcisetosus sp. nov. malesÐ1.12Ð1.16 mm, femalesÐ1.34Ð1.96 mm. Body (Figs. 10Ð19) shape cylindrical with anal spines present. Furca re- duced to small area of Þne granulation with two ϩ two Description. Color white. Length (excluding an- setae posterior, arranged in one row. Granulation of tennae) of adults: holotypeÐ2.15 mm, reproductive body more or less uniform, distinct, somewhat coarser malesÐ2.08Ð2.15 mm, reproductive femalesÐ2.2Ð2.3 on terga, head capsule and around anal spines. An- mm. Furca reduced to small concavity, granulation tennal bases marked from head. Antennae slightly smaller and sparser than on remainder of body, with ϩ shorter than head. Antennal segment IV with a sub- two two setae arranged in one row. Granulation apical organ, consisting of two pits, of which the Þrst more or less uniform, distinct, somewhat coarser on is deep and usually ampulla-like, second is shallow, terga, head capsule, and around anal spines. Antennal with small sensillum. Microsensillum on antennal seg- bases clearly marked from head. Antennae slightly ment IV in latero-external position, approximately shorter than head. Antennal segment IV with a sub- one-half length from the base. Antennal segment III apical organ, consists of two pits, from which the Þrst with microsensillum slightly below antennal III sense is deep and usually ampulla-like, second is shallow, organ. Thoracic terga II and III with microsensilla with small sensillum. Microsensillum on antennal seg- located laterally. Antennal organ III with Þve guard ment IV in latero-external position, approximately setae, two sensory rods, two bent and smooth sensory one-third length from the base. Antennal segment III clubs and Þve papillae of irregular shape (Fig. 22). with microsensillum slightly below antennal III sense organ. Thoracic terga II and III with microsensilla Postantennal organ consists of 15 vesicles, covered located laterally. Antennal organ III with Þve guard with secondary vesicles. Pseudocellar formula: dorsal setae, two sensory rods, two bent and smooth sensory 32/033/33354(3), ventral 2/000/1112; subcoxae of legs clubs and Þve papillae of regular shape (Fig. 15). I-III each with two pseudocelli. Formula of parap- Postantennal organ consists of 16Ð17 vesicles, covered seudocelli ventrally 10/000/101003. Dorsal chaetotaxy with secondary vesicles. Pseudocellar formula: dorsal (Fig.